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The Dictionary of VIROLOGY This page intentionally left blank The Dictionary of VIROLOGY

Fourth Edition

Brian W.J. Mahy

Division of Emerging Infections and Surveillance Services Centers for Disease Control and Prevention Atlanta, GA 30333 USA

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09 10 11 12 10 9 8 7 6 5 4 3 2 1 Preface to the Fourth Edition

The fourth edition of this Dictionary was necessary because of the considerable body of new knowledge concerning viruses which has accumulated since the last edition appeared in 2001. As in previous editions, the Dictionary is confined to viruses affecting vertebrates, from humans to fish, and no attempt has been made to include viruses which infect bacteria, fungi, invertebrates, or plants. Thus this book is mainly aimed at an audience of those interested in human or veterinary virology. Those needing a complete coverage of all viruses can consult the Encyclopedia of Virology (3rd edition, 2008) edited by BWJ Mahy and MHV van Regenmortel (Oxford: Academic Press). W i t hin this field the last 7 years have witnessed the emergence of numerous previously unknown viruses, such as the new human coronavirus causing severe acute respiratory syndrome (SARS). This disease spread rapidly around the world from its origin in China to infect more than 8000 persons, some 800 of whom died. In addition, new technological approaches have uncovered other new viruses such as the human bocavirus and the human metapneumovirus, both of which are associated with respiratory tract infections worldwide. A further reason for this fourth edition is that the ICTV, which is the decid- ing body on virus nomenclature and taxonomy of the International Union of Microbiological Societies, has produced a new eighth Report, under the chair- manship of L Andrew Ball, which includes many changes and refinements, and these are reflected throughout this new edition. Many of the changes are based upon reliable nucleotide sequence analyses of viral genomes. The speed with which this information can be obtained nowadays is evident from the outbreak of SARS which was first isolated in March 2003, but even though the coronavirus genome is the largest of any RNA virus, two complete genome sequences were published by independent laboratories in Canada and US by May of that year. As in previous editions, I have provided many references that will serve to provide easy entry into the literature. In general these have been chosen on the basis of wide coverage of the field, such as review articles, rather than first reports. Space constraints made it impossible to cite all key papers in an area, and omission of a relevant publication does not mean that it was not considered in writing the entry. I wish to thank the many fellow virologists who provided comments or information during the preparation of this edition, especially Tom Barrett, Charles Calisher, Rosa Gualano, James Mills, Stuart Nichol, Colin Parrish, Peter Tattersall, Bill Taylor Marc van Regenmortel, Peter Walker, and Scott Weaver. If any readers wish to offer suggestions for correction or other improvement to this text, I would appreciate receiving them by e-mail to [email protected]. Finally, I wish to express my deepest thanks to my dear wife Penny for the great skill and dedication she was able to bring to editing this new edition.

Brian WJ Mahy Atlanta, Georgia USA This page intentionally left blank A

10924 virus An isolate of Latino virus in lung carcinomatous tissue from a 58- the genus Arenavirus . year-old Caucasian male.

12056 virus An isolate of Paraná virus in A6 cells (CCL 102) Epithelial cell line the genus Arenavirus . initiated by primary cultivation of normal kidneys from an adult male 127 virus See egg drop syndrome 1976- toad. The cells support the replication associated virus . of frog virus 3 but not the Lucke frog herpesvirus. 13p2 virus American oyster reovirus in the genus Aquareovirus . A9 cells (CCL 1.4) A fibroblastic cell line derived from wild-type L929 cells. 1324Cg/79 virus A strain of Puumala virus Sensitive to HAT selection media, in the genus Hantavirus . deficient in adenosine phosphoribosyl transferase (APRT) and hypoxanthine 2060 virus Classified originally as echo- phosphoribosyl transferase (HPRT). virus 28. Now designated a strain of See HAT selection . human rhinovirus subtype 1A in the genus Rhinovirus . A particles See A-type virus particles .

3076 virus An isolate of Mobala virus in A23 virus A subtype in the genus Human the genus Arenavirus . enterovirus B . Originally thought to be coxsackie A23 virus but identical to 3099 virus An isolate of Mobala virus in human echovirus 9. the genus Arenavirus . AA288-77 virus An isolate of Machupo 3739 virus An isolate of Pichinde virus in virus in the genus Arenavirus . Isolated the genus Arenavirus . in Bolivia from the rodent Calomys callosus . 63U-11 virus (63UV) A strain of Marituba virus in the genus Orthobunyavirus . AAV Abbreviation for Adeno-associated virus in the genus Dependovirus . 75V 446 virus (San Juan virus) A strain of Alajuela virus in the genus Abacavir n 6-Cyclopropylamino 2 3 - Orthobunyavirus . dideoxyguanosine. A carbocyclic nucleoside analog of deoxyguanosine, which 75V 2374 virus (V2374V) A strain is phosphorylated to form carbovir of Alajuela virus in the genus triphosphate, an inhibitor of reverse Orthobunyavirus . transcriptase activity. Potentially active against retrovirus infection, but resist- 75V 2621 virus (V2621V) A strain ant mutations have been observed in of Gamboa virus in the genus clinical isolates. Orthobunyavirus . Abadina virus (ABAV) A serotype of 78V 2441 virus (V2441V) A strain Palyam virus in the genus Orbivirus of Alajuela virus in the genus belonging to the Palyam serogroup. Orthobunyavirus . Isolated from Culicoides sp.

A549 cells (CCL 185) Epithelial cell line Abelson murine leukemia virus initiated through the explant culture of (AbMLV) An acutely transforming Abelson murine leukemia virus (AbMLV) 2

strain of Murine leukemia virus in the blood of a 3-year-old boy with biphasic genus Gammaretrovirus isolated from fever and signs of meningitis. Found prednisolone-treated BALB/c mice in Sweden, Finland, Poland, former inoculated with Moloney leukemia Czechoslovakia, Hungary, Austria, virus. It has a short latent period and Bulgaria, and western parts of the produces lymphoid leukemia of B-cell former USSR. type. It can transform 3T3 mouse cells in vitro. Requires a helper virus for absorbance The amount of light absorbed complete virus replication. by a solution or substance at a particu- Risser R et al (1982) Biochim Biophys Acta 651 , lar wavelength. 213 Synonym : optical density. abl gene The oncogene of Abelson absorption Uptake of one substance by murine leukemia virus. The gene prod- another, e.g. removal of antibodies uct, a 160-kDa fusion protein, attaches from a mixture by adding soluble anti- to the cell plasma membrane via a gens, or vice versa . myristic acid residue and has tyrosine- specific protein kinase activity. absorption spectrum Graphical represen- tation of the absorbance of a substance Abney virus A virus, isolated from an at different wavelengths. anal swab of a child with upper respi- Abu Hammad virus (AHV) ratory illness, which became a proto- A strain of Dera Ghazi Khan virus type strain of reovirus type 3. in the genus Nairovirus. Isolated from a tick, Argas Rosen L et al (1960) Am J Hyg 71 , 258 hermanni, in Egypt. Not reported to cause disease in humans. abortive infection Infection in which some or all virus components are syn- Abu Mina virus (ABMV) A strain of thesized but no infective virus is pro- Dera Ghazi Khan virus in the genus duced. Also termed ‘ non-productive Nairovirus . Not reported to cause disinfection.’ Usually occurs because the ease in humans. host cell is non-permissive. May also result from infection with defective Acado virus (ACDV) A strain of viruses; in these cases it may be possi- Corriparta virus in the genus Orbivirus . ble to rescue the virus by co-infection Isolated from Culex antennatus and with a helper virus or by co-cultivation. C univittatus neavi in Ethiopia. Not reported to cause disease in humans. abortive transformation Transformation of cells which is unstable. A few gen- Acará virus (ACAV) A species in the erations after transformation the cells genus Orthobunyavirus . Isolated from revert to normal. sentinel mice, Culex sp mosquitoes, and the rodent Nectomys squamipes in Para, Above Maiden virus A serotype of Great Brazil, and in Panama. Not reported to Island virus in the genus Orbivirus . cause disease in humans.

Abras virus (ABRV) A strain of Patois accessory proteins Virus-specified provirus in the genus Orthobunyavirus . teins that are not required for virus rep- Isolated from Culex adamesi and C lication. Found in the arteriviruses and paracrybda in Ecuador. Not reported to coronaviruses as nonstructural proteins cause disease in humans. of unknown function. Calisher CH et al (1983) Am J Trop Med Hyg 32 , 877 acciptrid herpesvirus 1 (AcHV-1) An unassigned virus in the family Absettarov virus (ABSV) A strain of Herpesviridae , isolated from a nesting Tickborne encephalitis virus (Far Eastern bald eagle, Haliaetus leucocephalus . subtype) in the genus Flavivirus . Synonym : bald eagle herpesvirus. Isolated in 1951 in Leningrad from the Docherty DE et al (1983) Avian Dis 27 , 1162 3 acquired immunodeficiency syndrome (AIDS) acetoxycycloheximide A glutarimide acipenserid herpesvirus 2 (AciHV-2) antibiotic. A potent reversible inhibitor An unassigned member of the family of protein synthesis in animal cells. See Herpesviridae , isolated from the inter- also cycloheximide . nal organs of adult white sturgeon, Acipenser transmontanus . Biochemically acetylcysteine A mucolytic agent used and biologically distinguishable from for adjunct therapy of bronchopulmo- acipenserid herpesvirus 1. nary disorders to reduce the viscosity Synonym : white sturgeon herpesvirus 2. of mucus. Appears to have antiviral effects in HIV patients due to inhibition acquired immunodeficiency syndrome of viral stimulation by reactive oxygen (AIDS) A disease of humans caused intermediates. by human immunodeficiency viruses (HIV) 1 and 2. Globally, more than 42 N-acetylethyleneimine (AEI) An azirid- million people were infected with HIV ine compound which is a potent inac- by the year 2006, and millions had tivator of virus infectivity. Used in already died from the disease. The incu- inactivated vaccine formulation, e.g. bation time from infection to develop- for foot-and-mouth disease virus vac- ment of AIDS appears to range from 6 cines. Has also been used to inactivate to 13 years (median 10 years). AIDS is rabies virus, but yielded a less potent primarily a disease of the immune sys- immunogen than virus inactivated tem so the infection usually results in with β -propiolactone. a wide range of adverse immunologi- Wiktor TJ et al (1972) Appl Microbiol 23 , 914 cal and clinical conditions. The extent of the disease is generally measured by 2,3-bis-(acetylmercaptomethyl)quinoxalin the CD4 lymphocyte count, and as the An antiviral agent. Inhibits polio- count declines to below 200 per micro- virus RNA synthesis in vitro and in vivo . liter there is serious risk of AIDS-related Inhibits human herpesvirus 1 multipli- complex (ARC), a syndrome involving cation in vitro . Does not interfere with opportunistic infections, such as recur- attachment, penetration, or DNA syn- rent bacterial infections, candidiasis, thesis, but interrupts a late stage in pulmonary tuberculosis, Pneumocystis virus assembly and/or maturation. carinii pneumonia, EBV-associated lymphoma, and Kaposi’s sarcoma. Bucchini D and Girard M (1975/1976) The opportunistic infections (i.e. those Intervirology 6 , 285 caused by microorganisms that seldom cause disease in persons with nor- aciclovir (ACV) See acycloguanosine . mal defense mechanisms) and cancers resulting from immune deficiency are acid-stable equine picornaviruses (EqPV) generally the most severe but neurologi- Unassigned species in the family cal problems such as dementia resulting Picornaviridae . from HIV infection of the brain cells can also occur. The disease is almost always acipenserid herpesvirus 1 (AciHV-1) fatal. The virus is generally transmitted An unassigned member of the family through blood and body fluids, usually Herpesviridae, isolated from juvenile through unprotected sexual intercourse, white sturgeon, Acipenser transmonta- but vertical and/or perinatal trans- nus , suffering mortality during rearing mission is also very common. AIDS is in north-west American hatcheries. The prevalent among injecting drug addicts virus replicates in white sturgeon epi- and in patients receiving transfusions of dermal cell cultures, inducing syncytia. blood which was not screened for the Associated with epidermal hyperpla- presence of HIV. It can be partly control- sia and necrosis in the fish. The virus led by antiretroviral drugs such as AZT could be transmitted to juvenile white (azidothymidine) or non-nucleoside sturgeon but not to trout. reverse transcriptase inhibitors such Synonym : white sturgeon herpesvirus 1. as nevirapine combined with protease Hedrick RP et al (1991) Dis Aquat Organ inhibitors (see HAART), but the side 11 , 49 effects of the drugs are not negligible. acquired immunodeficiency syndrome (AIDS) 4

Jasny B (Editor) (1993) Science 260 , 1253 acute adult T-cell leukemia See human Kaplan JE et al (2000) Clin Infect Dis 30 , T-lymphotropic virus 1 . Suppl 1 Levy JA (1993) Microbiol Rev 57 , 183 acute anterior poliomyelitis virus Synonym for poliovirus. acridine orange A fluorescent derivative of acridine which will bind to nucleic acute epidemic gastroenteritis virus of acids in cells or within the virion. humans Synonym for Norwalk virus, the When exposed to ultraviolet light, the type species of the genus Norovirus in dye fluoresces orange if the nucleic the family Caliciviridae. Causes diarrhea acid is single-stranded; green if it is and vomiting in children and adults. double-stranded. See also photody- There are at least seven related cali- namic inactivation . civiruses in the group: Desert Shield, Hawaii, Lordsdale, Mexico, Norwalk, acriflavine A photoreactive dye. See pho- Snow Mountain, and Southampton. todynamic inactivation . Virus particles are 27 nm in diameter, and are ether- and acid-stable. Found acronym (Greek: acro extreme, onoma in the feces originally only by electron name) A special case of sigla, frequently microscopy, but now detected and used in virology. A word created from distinguished using PCR. Antibodies the initial letters of the principal words can be demonstrated in patients. The in a compound term. See CELO virus virus is very difficult to propagate in and human echoviruses as examples. vitro . See also gastroenteritis viruses of humans . acrylamide A chemical which is polym- Estes MK and Hardy ME (1995) In Infections erized using a cross-linking agent of the Gastrointestinal Tract, edited by to give polyacrylamide, one of the MJ Blaser et al . New York: Raven Press, p. most commonly used supports for gel 1009 electrophoresis. Fankhauser RL et al (1998) J Infect Dis 178 , 1571 actidione Synonym for cycloheximide. Talal AH et al (2000) J Med Virol 61 , 117 actinomycin D An antibiotic produced by acute epidemic hemorrhagic conjuncti- the fungi Streptomyces chrysomallus and vitis See Apollo virus . S antibioticus. Inhibits DNA-dependent RNA transcription. Interacts with the acute hemorrhagic conjunctivitis virus guanine residues of helical DNA. Not A strain of Human enterovirus D in the readily reversible by removal of drug genus Enterovirus, designated human from the culture medium. Blocks inter- enterovirus 70. Causes acute hemor- feron production by inhibiting mRNA rhagic conjunctivitis in humans in all synthesis. Most single-stranded RNA parts of the world except the Americas viruses are not significantly affected by and Australia. The prototype strain the drug at concentrations of 1–5 μ g/ml, J670/71 isolated in Japan multiplies which inhibit host cell DNA-dependent optimally at 32–34°C in monkey kid- RNA transcription; influenza viruses ney cell cultures. A low temperature and retroviruses are notable exceptions. should be used for isolation, although Synonyms: dactinomycin; meractino- the virus can be adapted to higher tem- mycin. peratures. Acute hemorrhagic conjunctivitis may also be caused by a strain of activator A protein which binds to DNA Human enterovirus C, known as human upstream of a gene and activates tran- coxsackievirus A24. scription of that gene. Now usually Yin-Murphy M (1984) Prog Med Virol 29 , 23 called a transactivator. acute infection Virus infection which active immunity Immunity induced results in brief symptoms, lasting a by injection of virus or virus subunit few days (such as the common cold), antigens. after which the virus is eliminated 5 Adelaide River virus (ARV)

completely by the immune system. See altered DNA polymerase may arise also: persistent infection . during prolonged treatments. The drug does not affect latent herpesviruses, so acute laryngo-tracheo-bronchitis virus recurrence after treatment is possible Synonym for Human parainfluenza and has been described. virus type 1, a species in the genus Synonyms: Wellcome 248U; aciclovir; Respirovirus . acyclovir; Zovirax. Darby G (1995) Antivir Chem Chemother 5 , acute respiratory distress syndrome Suppl 1, 54 (ARDS) A fulminant lung alveolar Elion GB (1989) Science 244 , 41 and interstitial edema, which develops rapidly, due to increased capillary acyclovir See acycloguanosine 9-(2- permeability. One instance, hantavirus hydroxyethoxymethyl) guanine . pulmonary syndrome, is caused by Sin Nombre virus, but there may be other acylation Introduction of an acyl radical viral or bacterial causes of this disease (RCO) into an organic compound. Acyl syndrome. transferase is an enzyme that transfers an acyl group from acyl-coenzyme A to acyclic nucleoside analogs A series of another compound. During influenza antiviral compounds active against var- virus replication, the three cysteine ious species in the family Herpesviridae . residues at the carboxyterminal por- They include acyclovir, bucyclovir, gan- tion of the hemagglutinin of the virus ciclovir, penciclovir, and 2HM-HBG. are acylated. de Clercq E (1995) Rev Med Virol 5 , 149 1-adamantanamine hydrochloride See acyclic nucleoside phosphonates A series amantadine . of antiviral compounds active against a wide range of DNA viruses. They Adansonian system A polythetic hierar- include HPMPA, HPMPC, PMEA, chical classification system proposed PMPA, and PMPDAP. by Adanson in 1763, which considers many criteria of equal importance to acycloguanosine 9-(2-hydroxyethoxyme- define a species, and is essentially the thyl) guanine A nucleoside analog. basis of the present universal classifica- An antiviral agent with a potent and tion system for viruses in use today. highly specific action against human herpesvirus 1, 2, and 3 both in vitro Adefovir 9-(2-phosphonylmethoxy ethyl) and in animal models of skin, eye, adenine . A nucleoside inhibitor of and brain infections. It is only weakly reverse transcriptase activity, poten- active against human herpesvirus 5. tially active against retrovirus infec- The drug is selectively phosphorylated tion. It is also active against chronic by herpesvirus induced thymidine hepatitis B infection even in the pres- kinase, and once phosphorylated is a ence of virus mutants that are resistant potent inhibitor of herpesvirus-induced to lamivudine. DNA polymerase. In a clinical study, Synonym : PMEA. 24 patients with dendritic corneal epithelial ulcers were treated by minimal Peters MG et al (2004) Gastroenterology wiping debridement, 12 then receiving 126 , 91 the drug topically as eye ointment, the others being given a placebo. Seven Adelaide River virus (ARV) A species of the placebo patients showed recur- in the genus Ephemerovirus . Isolated rence of herpetic corneal lesions within in 1981 from cattle in Tortilla Flat, a week. There was no recurrence in Northern Territory, Australia. Although the patients receiving acycloguanos- known to affect cattle, the virus is not ine. Acute toxicity studies have shown known to cause any serious disease that the drug has a very low toxicity. symptoms, and has been isolated from Acyclovir-resistant mutants with an healthy sentinel cattle. 9-(S)-(2,3-dihydroxypropyl)adenine 6

9-(S)-(2,3-dihydroxypropyl)adenine An with disease. For all species except antiviral agent which inhibits rotavirus AAV-5 replication is dependent upon infection. the presence of a helper adenovirus for Smee DF et al (1982) Antimicrob Agents complete virus production, but infec- Chemother 21 , 66 tious DNA and antigens demonstrable by immunofluorescence are made in adenine arabinoside 9-d -Arabino- the presence of a helper herpes-type furanosyladenine. An analog of virus. A latent infection may be estab- deoxyadenosine, which is active as lished in the absence of a helper virus. an antiviral agent particularly for the Replicate in cells which support adeno- treatment of severe herpes simplex virus replication to a higher titer than virus infections. Synthesized in 1960, the adenovirus whose replication may it was subsequently found as a natu- be depressed. Not genetically related rally occurring nucleoside in culture to adenovirus. Mature virus particles filtrates of Streptomyces antibioticus . contain equivalent numbers of posi- Phosphorylated to the active form, tive or negative strands of DNA, each araATP, by cellular kinases it inhibits approximately 4.7 kb in length, pack- viral DNA synthesis at lower concen- aged into separate virions. The strands trations than are required to inhibit are complementary, and after extraction host cell DNA synthesis, by inhibiting anneal to form double-stranded DNA. the viral DNA polymerase. It neither There are a number of serotypes. The directly inactivates virus nor prevents type species is type 2. Types 1, 2, 3, 4, attachment. In the body it is speedily and 5 are primate adeno-associated converted to the hypoxanthine, with a viruses. There are also bovine, avian, decline to less than 50% of the original canine, ovine, and equine types. antiviral activity. It is active against her- Antibodies can be found in human sera, pesvirus and poxvirus; less so against but none of these viruses is known to be adenovirus and papovavirus. The drug pathogenic. The use of AAV as a vector has no action against RNA viruses. for potential gene therapy in a clinical Although acyclovir is the drug of choice setting is under development, and has for treatment of herpes simplex or been shown to restore vision in a canine varicella-zoster infections, adenine ara- model of childhood blindness. binoside appears useful for treatment Synonym : adeno-satellite virus. of acyclovir-resistant mutant viruses. Acland GM et al (2001) Nature Genet 28 , 92 Synonyms : vidarabine; ara A; vira A. Berns KI and Linden RM (1995) BioEssays 17 , 237 Whitley RJ et al (1980) Drugs 20 , 267 Matsushita et al (1998) Gene Therapy 5 , 938 Tattersall P and Cotmore SF (2005) In adenine arabinoside monophosphate A Topley & Wilson’s Microbiology and Microbial phosphorylated derivative of adenine Infections , vol. 1, Tenth edition, edited by arabinoside with greater solubility than BWJ Mahy and V ter Meulen. London: the parent compound. Hodder Arnold, p. 407 Spruance SL et al (1979) N Engl J Med 300 , 1180 adeno-associated virus vectors Because adeno-associated viruses are wide- Adeno-associated virus 1–5 (AAV1–5) spread in nature and cause no appar- Species in the genus Dependovirus . AAV- ent disease, the possible use of human 2 and AAV-3 were isolated from humans AAV as a gene vector has been actively in association with adenovirus, and explored. The virus is known to inte- AAV-5 is a human isolate that appears grate at a specific site in the human to depend upon herpesvirus for its rep- genome (19q 13.4), and several clinical lication. AAV-6 is now regarded as a trials have been carried out with encour- strain of AAV-1, which is a simian iso- aging results for diseases such as cystic late like AAV-4. The genus also contains fibrosis, Factor IX hemophilia, and other two tentative species, AAV-7 and AAV- disease with a known genetic cause. 8, which were also simian isolates. None Linden RM and Berns KI (2005) In Topley & of these viruses has yet been associated Wilson’s Microbiology and Microbial Infections , 7 adenovirus death protein (ADP)

vol. 2, Tenth edition, edited by BWJ Mahy There are inverted terminal repeti- and V ter Meulen. London: Hodder Arnold, tions of 50–200 base pairs in all viruses p. 1590 sequenced. GC content of the DNA varies between 34% and 64%. Viral adenoid degeneration agent Synonym for maturation takes place in the nucleus. human adenovirus, for which it was the Virus is liberated by cell disruption. original name. Causes degeneration of Adenoviruses may be divided on the human tonsillar tissue grown in culture. basis of their host species and further on the basis of hemagglutination, anti- adenoidal–pharyngeal–conjunctival genic structure, and oncogenicity. They agent Synonym for human adenovirus. express immunoregulatory genes that act against the host defense mecha- adenoma An epithelial cell tumor that is nisms. Usually found in the respiratory usually benign; the cells form gland- tract where they are often associated like structures. with disease. adeno-satellite virus Synonym for Adeno- Benko M and Harrach B (1998) Arch Virol 143/144 associated virus , 829 . Benko M et al (2002) J Virol 76 , 10056 Horwitz M (2001) Virology 279 , 1 adenosine-deaminase ADAR1 An Russell WC (2000) J Gen Virol 81 , 2573 interferon- induced RNA- specific Russell WC (2005) In Topley & Wilson’s cytokine which catalyzes the conver- Microbiology and Microbial Infections , vol. 1, sion of A to I. Tenth Edition, edited by BWJ Mahy and V ter Meulen. London: Hodder Arnold, p. 439 adenosine triphosphatase (ATPase) An enzyme which catalyzes the conver- Adenovirus General name for species sion of ATP to ADP with the release of in the family Adenoviridae . Most ade- Pi. Some viruses (e.g. vaccinia virus) noviruses grow in only one species, possess an ATPase activity. and they are named accordingly, e.g. human adenovirus 2, where 2 refers Adenovir See penciclovir. to the serotype. Adenovirus serotypes are generally defined by neutraliza- Adenoviridae A family of double- tion assays with a cutoff ratio (homolo- stranded DNA viruses with icosahedral gous:heterologous titer) greater than symmetry. There are four recognized 16. Most adenovirus infections are genera: Mastadenovirus, Aviadenovirus, asymptomatic, but they can cause seri- Atadenovirus, and Siadenovirus. The ous diseases of the respiratory, ocular, virion is non-enveloped, 70–90 nm in and gastrointestinal systems. These can diameter, formed of 240 non-vertex be fatal in some patients, and a case capsomeres (hexons) 8–10 nm in diame- of fatal adenovirus type 3 pneumonia ter, and 12 vertex capsomeres (pentons) has been reported in identical immu- that consist of a base and an outward nocompetent adult twin sisters. The projection (fiber) up to 77.5 nm long use of human adenoviruses as poten- with a knob at its end. Members of the tial vectors for gene therapy, includ- Aviadenovirus genus have two fibers at ing selective toxicity to tumor cells, is each vertex. The virion is resistant to under active development. See also mild acid, lipid solvents, and trypsin. Adenoviridae . Buoyant density (CsCl): 1.30–1.37 g/ml. Barker JH et al (2003) Clin Infect Dis 37 , 142 Inside the capsid is the core, consisting of protein and a linear molecule of double-stranded DNA mol. wt. 20–30 1 0 6 adenovirus death protein (ADP) A pro- with an inverted terminal repetition and tein which is expressed in increased no single-stranded breaks. The DNA amounts late in infection, and facili- sequence of human adenovirus 2 has tates the release of progeny virions 35,937 base pairs, but over the whole from adenovirus-infected cells by trig- family sequence lengths from 26,163 to gering host cell lysis. 45,063 base pairs have been reported. Hausmann J et al (2000) Virology 244 , 343 adenovirus-SV40 hybrids 8 adenovirus-SV40 hybrids First reported adventitious viruses Contaminant between human adenovirus 7 and viruses present by chance in a virus SV40. An adenovirus 7 isolate was preparation or vaccine. Animals and found to be contaminated with SV40 cell cultures are often infected with after isolation and passage in primary adventitious viruses, whose presence rhesus monkey kidney cells. Infectious may go unrecognized for a period. SV40 virus was eliminated by passage in the presence of SV40 antiserum but Aedes cells (CCL 125 and 126) Cell lines on injection into newborn hamsters established from the mosquitoes, Aedes the tumor cells produced contained albopictus and A aegypti, able to sup- adenovirus 7 and SV40 T antigens. The port the replication of a number of virus could be neutralized by adeno- arboviruses. virus antiserum, and SV40 genome sequences appeared to be in an adeno- aerosol A gaseous suspension of ultrami- virus particle. The adenovirus SV40 croscopic particles or liquid droplets. hybrid stock virus was named E46 . The viral DNAs in the hybrid virus are aerosol transmission A common form of covalently linked. A number of hybrids transmission of viruses which infect via between SV40 and other adenoviruses the respiratory tract, such as influenza, have since been described. They may rhinoviruses, measles, and mumps be divided into those that produce free viruses. infective SV40 virus and thus contain the complete SV40 genome, and those affinity chromatography Chromato- that do not. Used as experimental tools, graphy using ligands attached to an e.g. to study SV40 T antigen. insoluble support which interacts with Tjian R (1978) Cell 13 , 165 the molecule of interest, retaining it, and allowing unwanted molecules to adhesion molecules Cell surface mole- be washed away. On changing the con- cules that mediate leukocyte binding to ditions the molecule of interest can be the vascular endothelium. They include eluted. Examples are the selection of selectins, integrins, and molecules of antigen by immobilized antibody and the immunoglobulin superfamily. Many the selection of polyadenylated mRNA of these molecules are used as receptors using oligo dT sepharose. on the cell surface for viruses. Afluria A trivalent inactivated influenza vaccine approved in the USA in 2007 adjuvant Substance added to antigens to for intramuscular injection in persons enhance immune response. Salts of alu- over 18 years of age. minum (e.g. hydroxide or phosphate) acceptable for use in humans. Saponin AFP Acute flaccid paralysis, such as may or Freund’s adjuvant used only in be caused by poliovirus infection. experimental animals. African green monkey cytomegalovirus AdoHcy Abbreviation for S -adenosyl- See Cercopithecine herpesvirus 5 . homocysteine. African green monkey EBV-like virus AdoMet Abbreviation for S -adenosyl- See Cercopithecine herpesvirus 14 . methionine. African green monkey HH4-like virus adsorption See attachment . See Cercopithecine herpesvirus 14 . adult respiratory distress syndrome African green monkey kidney cells (ARDS) See acute respiratory distress Continuous line of cells used for syndrome . growth of certain viruses of vertebrates including adenoviruses, SV40, rotavi- adult T-cell leukemia virus See human T ruses, poliovirus, rubella, arboviruses, lymphotropic virus 1 . and paramyxoviruses such as measles 9 African swine fever virus (ASFV)

virus. There are three commonly used Infectivity is ether-resistant but acid- lines: BSC-1, CV-1, and Vero cells. sensitive, being inactivated below pH 6. Horse erythrocytes are agglutinated. African green monkey polyomavirus Virus contains double-stranded RNA in (AGMPyV) A species in the genus 10 segments. Multiplies in eggs in yolk- Polyomavirus . Isolated from a South sac, and in cell cultures of many species. African vervet monkey kidney cell Although originally confined to Africa, culture. Has been adapted to growth the virus was inadvertently introduced in primary rhesus monkey kidney cell into Spain, and is now endemic around cultures. Virion diameter is 40–45 nm. Madrid and other areas to the south. The superhelical, circular double- Synonyms: African horse plague virus; stranded DNA has a mean length 101% perdesiekte virus; pestis equorum of the length of SV40 DNA (5.2 kb). Has virus. the internal capsid antigen common to Mellor PS and Mertens PPC (2008) In all papovaviruses of the SV40 polyoma Encyclopedia of Virology , Third edition, BWJ group. Unrelated to other papova- Mahy and MHV van Regenmortel (eds). viruses by neutralization tests. T antigen Oxford: Academic Press, vol. 1, p. 37 is indistinguishable from the T antigen of SV40. Transforms hamster kidney African monkey cytomegalovirus cells. Natural host probably the chacma Synonym for Cercopithecine herpesvirus 2 . baboon, Papio ursinus . Synonym : B-lymphotropic papovavirus strain K38. African Rift Valley fever virus See Rift Valley fever virus . Valis JD et al (1977) Infect Immun 18 , 247 African swine fever virus (ASFV) Only African horse plague virus Synonym for member of the genus Asfivirus. The African horse sickness virus . genome has double-stranded DNA, about 170–200 kb, depending on the African horse sickness virus (AHSV) isolate. The DNA is linear, covalently A species in the genus Orbivirus with closed-ended with inverted comple- nine serotypes. mentary tandem repeats and is infectious. Causes a fatal disease resembling African horse sickness viruses 1–9 classic swine fever in domestic pigs (AHSV1–9) Serotypes in the genus (Sus scrofa domesticus). There is high Orbivirus . There are nine serotypes fever, cough, and diarrhea. Incubation identified by neutralization tests. There period 7–9 days. Virus replication is a group-specific CF antigen. Only begins in the tonsils but soon becomes equids and dogs are naturally suscepti- generalized: especially involved are ble. Causes disease which may be fatal lymph nodes and spleen. Surviving in horses, mules, and donkeys. Dogs, pigs may have viremia for months. elephants, and zebras are possible res- Natural hosts are domestic and wild ervoir hosts. Camels, cattle, sheep, swine (Sus scrofa), warthogs, bush pigs, goats, and jackals may also be infected. and giant forest hogs. Infection is by In severe cases death occurs from pul- contact and fomites. Premises may be monary edema. In chronic cases there infective for months. Virus has been is cardiac involvement with edema of recovered from argasid ticks, especially the head and neck. Some infections are Ornithodorus moubata in East Africa mild. Viremia often occurs. Transmitted and O erraticus in Spain and Portugal, by nocturnal biting flies of the genus and replication in the ticks demon- Culicoides. Experimentally goats are strated. Virus diameter is 200 nm; enve- slightly susceptible but ferrets and dogs lope is acquired as it buds through the are infected more readily. Mice, rats, and plasma membrane. Survives dry at guinea pigs can be infected i.c. A mouse room temperature for years. Resists brain passage virus vaccine is effective. inactivation by some disinfectants Virion is 75–80 nm in diameter, icosa- but inactivated by 1% formaldehyde hedral and similar to bluetongue virus. in 6 days, 2% sodium hydroxide in African swine fever virus (ASFV) 10

24 days. Chloroform- and ether-resistant. agammaglobulinaemia A condition in Replicates in the chick embryo yolk-sac which there is no immunoglobulin in killing the embryo, and in cell cultures the blood, either familial, such as X- of swine macrophages, such as pig linked agammaglobulinemia, or follow- bone marrow. Hemadsorption of pig ing replacement of lymphoid tissues by erythrocytes is seen after 24 h and CPE lymphoma or leukemia for example. later. After 100 passes the virus loses virulence for pigs but does not give agar A mixture of polysaccharides, some protection from infection with virulent anionic, extracted from red algae, that virus. Antibodies do not provide immu- forms a gel at temperatures below about nity. Originally observed in East, South, 40° C. Used as a support medium when and West Africa, the virus reached supplemented by appropriate buffers/ Portugal and Spain in 1957, France in media ingredients for electrophoresis, 1964, Italy in 1967, and Cuba in 1971. production of microbial cultures, over- Outbreaks occurred in Malta, Sardinia, laying tissue culture cells, etc. and Brazil in 1978, and Haiti in 1979. In 1982 a severe outbreak occurred for agarose A sulfate-free neutral fraction of the first time in West Africa. The dis- agar. Often used in preference to agar ease is believed to have been eradicated as it does not contain inhibitors of virus from Europe (except Sardinia) since growth frequently present in agar, and 1995. In Brazil the disease is mild and as lower temperature gelling products has been difficult to eradicate. Sporadic of agarose are now available. Also used outbreaks have occurred in and been widely in gel electrophoresis as the pore eradicated from Cuba, the Dominican size is more uniform than that of agar. Republic, and Haiti. Probably infection can be spread by waste food from ships agarose gel electrophoresis Technique and aeroplanes. No vaccine is available. used for separating proteins or nucleic Synonyms: warthog disease virus; mal- acids by passage of an electric current adie de Montgomery virus. through the gel. Andres G et al (1997) J Virol 71 , 2331 Dixon LK et al (1993) Arch Virol Suppl 7 , 185 age-dependent polio-encephalitis of Dixon LK et al (1994) J Gen Virol 75 , 1655 mice virus Synonym for Lactate dehy- Dixon LK and Chapman D (2008) In Encyclopedia of Virology, Third edition, drogenase elevating virus . BWJ Mahy and MHV van Regenmortel (eds). Oxford: Academic Press, vol. 1, p. 43 Agenerase An antiviral drug that binds Yáñez RJ et al (1995) Virology 208 , 249 to the active site of the HIV-1 protease, resulting in the formation of imma- AG80-663 virus (AG80V) A species in the ture non-infectious virions. Also called genus Alphavirus , antigenically related Amprenavir. to Venezuelan equine encephalitis virus, but genetically distinct. agglutination test Some viruses will cause clumping of cells due to attach- AG83-1746 virus (AG83-1746) An ment to more than one cell and this isolate of Maguari virus in the can be used as the basis of an assay. genus Orthobunyavirus belonging to the Hemagglutination of red blood cells Bunyamwera serogroup. Isolated from by influenza virus is an example. Cells mosquitoes, Psorophora varinervis . other than red blood cells also exhibit Mitchell CJ et al (1987) Am J Trop Med Hyg the phenomenon. Also used to refer 36 , 107 to the clumping of inert particles, e.g. latex, coated with antibody and mixed AG83-497 virus (AG497V) An isolate of with homologous virus antigens. Melao virus in the genus Orthobunyavirus belonging to the California serogroup. agnoprotein A late protein encoded by Isolated from mosquitoes, Culex sp. SV40 and related viruses (BK and JC Mitchell CJ et al (1987) Am J Trop Med Hyg polyomaviruses) in the leader region of 36 , 107 late mRNA. Function unknown. 11 AKR (endogenous) murine leukemia virus (AKRMLV) agouti type C retrovirus See DPC-1 virus . reported to cause disease in humans. Antigenically indistinguishable from Aguacate virus (AGUV) A tentative spe- Samford virus, and cross-reacts with cies in the genus Phlebovirus related Akabane virus in CFT but not in neu- to the sandfly fever virus group, not tralization or hemagglutination inhibi- assigned to an antigenic complex. tion tests. Aino and Samford appear Isolated from Lutzomyia sp in central to be identical viruses, so the name Panama and Canal Zone. Not reported Samford has been dropped. to cause disease in humans. Levin A et al (2006) Virus Res 120 , 121 Yanase T et al (2003) Virus Res 93 , 63 Água Preta virus Probably a herpes-like virus as determined by thin-section electron microscopy. Isolated from the Akabane virus (AKAV) A species in the bat, Carolloa subrufa, in Utinga Forest genus Orthobunyavirus, belonging to near Belem, Brazil. the Simbu serogroup. Isolated from mosquitoes, but flies of Culicoides spp AHL-1 cells (CCL 195) Cell line derived are the most probable vectors. Found from the lung of a normal, adult, in Guma Prefecture, Honshu, Japan, male Armenian hamster, Cricetulus Queensland, Australia, and Vietnam. migratorius . There is serological evidence to associate infection with epizootic bovine Aichi virus (AiV) The type species of the congenital arthrogryposis and hydra- genus Kobuvirus. A cytopathic, small, nencephaly (A-H syndrome) in Japan round virus with icosahedral surface and Australia. The syndrome has also structure isolated from Pakistani chil- been reported in Israel, Argentina, dren and Japanese travellers return- South Africa, and Zimbabwe, caused by ing home from South-East Asia. Akabane-related viruses of the Simbu Biophysically and biochemically iden- antigenic group. Experimental infection tical to enteroviruses, but with differ- of pregnant sheep and goats causes dis- ent polypeptide bands and having no ease in the fetus. Not reported to cause reaction to the neutralizing antibod- disease in humans. ies or PCR amplification methods for Bryant JE et al (2005) Am J trop Med Hyg 73 , enteroviruses. 470 Yamashita T et al (1998) J Virol 72 , 8408 Parsonson IM et al (1981) Am J Trop Med Hyg 30 , 660 Yamakawa M et al (2006) Virus Res 121 , 84 AIDS Acronym for a cquired i mmuno d e- ficiency s yndrome. Akadon A large genus of South American AIDS-related complex A group of malig- grass mice with 41 currently recog- nancies and infections that occur in nized species. Can serve as hosts for the late stages of infection with human arenaviruses. immunodeficiency virus. See acquired immunodeficiency syndrome . AKATA An EBV-infected cell line derived from a human Burkitt’s lymphoma. AIDS virus See Human immunodeficiency virus types 1 and 2 . AKR (endogenous) murine leukemia virus (AKRMLV) A strain of Aino virus (AINOV) A strain of Shuni Murine leukemia virus in the genus virus in the genus Orthobunyavirus , Gammaretrovirus , belonging to the rep- belonging to the Simbu serogroup. lication competent virus group. An Isolated from mosquitoes, Culex tri- endogenous virus which causes spon- taeniorhynchus , in Nagasaki Prefecture, taneous leukemia in AKR mice. In mice Kyushu, Japan and C brevitarsis in which do not spontaneously develop Queensland, Australia, when it was leukemia earlier, T lymphomas develop named Samford virus. Antibodies after about 9 months. Leukemogenesis are present in cattle and horses. Not in AKR mice is accelerated by infection AKR (endogenous) murine leukemia virus (AKRMLV) 12

of mink cell focus-forming (MCF) virus can be detected in serum or plasma. recombinants. The MCF recombinants Elevations of ALT activity occur as a have a chimeric env gene between the result of liver damage, and are indica- endogenously activated and infect- tive of such damage in hepatitis virus ing virus genes. In AKR mice which infections. develop spontaneous leukemia, the provirus found in the tumor cells is usu- Alastrim virus Synonym for variola ally an MCF recombinant. It appears minor virus. See Variola virus . that the MCF envelope glycoprotein can bind to hematopoietic growth fac- albatrosspox virus A probable species in tor receptors, such as the erythropoietin the genus Avipoxvirus . receptor, and induce transformation. Hesse I et al (1999) Lab Anim Sci 49 , 488 Alcelaphine herpesvirus 1 (ALHV-1) Hung Fan (1994) In The Retroviridae , vol. 3, A species in the genus Rhadinovirus . edited by JA Levy. New York: Plenum Press, Causes a widespread, sporadic, often p. 313 fatal disease of cattle, with fever, acute Rich RF et al (2006) Virology 346 , 287 inflammation of nasal and oral mem- Stoye JP et al (1991) J Virol 65 , 1273 branes, and involvement of pharynx and lungs. There is often keratitis and AKR mink cell focus-inducing virus nervous symptoms. The virus can be A possible subspecies in the genus transmitted experimentally to cattle Gammaretrovirus. A helper independ- and rabbits. There is replication in cell ent virus isolated from AKR thymoma cultures of fetal bovine thyroid, adre- cells. Probably a recombinant between nal, kidney, spleen, and lung. The virus ecotropic AKR mouse leukemia virus will also replicate in Vero cells. and a xenotropic virus. Synonyms: bovine epitheliosis virus; Hartley JW et al (1977) Proc Natl Acad Sci bovine herpesvirus 3; malignant 74 , 789 catarrhal fever virus. Bridgen A (1991) Arch Virol 117 , 183 AKT The oncogene specified by AKT8 Bridgen A and Reid HW (1991) Res Vet Sci virus. The human cellular homologs, 50 , 38 known as c-AKT1 and c-AKT2, encode Rossiter PB (1985) Prog Vet Microbiol Immun protein kinase C-related serine/threo- 1 , 121 nine kinases. Alcelaphine herpesvirus 2 (ALHV-2) AKT8 virus A probable species in the A species in the genus Rhadinovirus . genus Gammaretrovirus. An acute trans- Epidemiological evidence suggests forming strain of mouse leukemia that infection can be transmitted to cat- virus. tle from African antelope of the family Bovidae, subfamily Alcelaphinae, which Alagoas virus See Vesicular stomatitis includes wildebeest, Connochaetes tau- Alagoas virus . rinus and C gnu, hartebeest, Alcephalus spp and topi, Damaliscus spp, which Alajuela virus (ALJV) A species in the may carry the virus as a latent infection. genus Orthobunyavirus in the Gamboa Synonyms : hartebeest malignant serogroup. Isolated from mosquitoes, catarrhal fever virus; Snotsiekte virus. Aedeomyia squamipennis in Alajuela, Costa Rica. Not reported to cause dis- Alenquer virus (ALEV) A serotype of ease in humans. Candiru virus in the genus Phlebovirus in the Candiru antigenic group. Calisher CH et al (1981) Am J Trop Med Hyg 30 , 219 Herrero MV et al (1994) J Med Entomol Aleutian disease virus See Aleutian 31 , 912 mink disease virus . alanine aminotransferase (ALT) A met- Aleutian mink disease virus (AMDV) The abolic enzyme the activity of which type species of the genus Amdovirus . 13 allotropic specificity

Causes an economically important, Porter DD et al (1980) Adv Immun 29 , 261 lethal disease in ranch-raised mink. All Qui J et al (2006) J Virol 80 , 654 types of mink are susceptible, but the Aleutian genotype, so named because Alexander cells A human hepatoma- the bluegray coat color is similar to that derived cell line, PLC/PRF/5, that car- of the Aleutian blue fox, develop more ries the hepatitis B virus genome and severe lesions and die sooner. Ferrets secretes hepatitis B surface antigen. are also susceptible. The virus causes Alexander J et al (1978) Perspect Virol 10 , 103 a persistent infection associated with a progressive disorder of the immune Alfuy virus (ALFV) A serotype of Murray system. The virus can cross the pla- Valley encephalitis virus in the genus centa to infect the fetus, but chronically Flavivirus. Isolated from mosquitoes in infected females produce few live kits. Queensland, Australia. Not reported to Animals infected in utero have a less cause disease, but antibodies to it or to severe disease than those infected after a closely related virus are common in birth. Virus is excreted in the urine, humans in northern Queensland. feces, and saliva, and infection is readily transmitted by contact and handling. alkaline phosphatase An enzyme with an After infection there is rapid replica- optimum pH above 8.0, which removes tion and high virus titers are present in the 5 terminal phosphate from a wide spleen, liver, and lymph nodes within variety of phosphate esters, includ- 10 days. A proportion of non-Aleutian ing linear DNA molecules. Occurs in mink clear the virus and develop no a variety of normal and malignant tis- disease. A chronic infection occurs in sues, and its level in serum may be the majority and high antibody levels high in certain diseases such as hepati- develop resulting in hypergammaglob- tis or bone disease. ulinemia. Viremia persists for months. Virus–antibody complexes are formed Alkhurma virus (ALKV) A tick-borne and deposited in the tissues, produc- flavivirus isolated in Saudi Arabia in ing glomerulonephritis, the usual cause 1995 from the blood of two male butch- of death, as well as arteritis of the cor- ers who died of hemorrhagic fever in onary, hepatic, gastrointestinal, and Dr Suliman Fakeeh Hospital, Jeddah. cerebral vessels. There is a systematic Several other cases were identified in plasmacytosis involving bone marrow, patients potentially exposed to sheep spleen, lymph nodes, liver, and kid- ticks. The genome sequence shows a neys. Ferrets and skunks can be infected close relationship to that of Kyasanur experimentally, but not rabbits, guinea Forest disease virus, from India, pigs, hamsters, rats, or mice. Human which also causes severe hemorrhagic infection is doubtful, although it would disease. be prudent to handle the virus with Synonyms : Fakeeh virus; Jeddah virus. caution. Replication in tissue culture is Bessaud M et al (2005) Virus Res 107 , 57 doubtful. Virion diameter is 23 nm, den- Charrel RN et al (2001) Biochem Biophys Res sity 1.29–1.41 g/ml in CsCl. The genome Commun 287 , 455 consists of a single negative-stranded Zaki AM (1997) Trans R Soc Trop Med Hyg DNA, 4748 nt in length. It is more resist- 91 , 179 ant to heat than most viruses, 60° C for 30 min causing only partial inactivation. Allerton virus Synonym for Bovine her- Resistant to lipid solvents and desoxy- pesvirus 2 . cholate. Passes through a filter of average pore diameter 50 nm. Control of the allotropic specificity The ability of par- disease can be obtained by killing all voviruses to replicate in different host hyperglobulinemic animals. cell types is governed by a small region Synonym : Aleutian disease virus. of the virus capsid called the allotropic Alexandersen S et al (1988) J Virol 62 , 1495 determinant. Best SM et al (2003) J Virol 77 , 5305 Agbandje-McKenna M et al (1998) Structure Parrish CR (1990) Adv Virus Res 38 , 403 6 , 1369 Allpahuayo virus 14

Allpahuayo virus A species in the genus of the second genus is Human herpesvi- Arenavirus. Isolated in 1997 from rus 3 and other species are Bovine her- arboreal rice rats (Oecomys bicolor pesvirus 1, Bubaline herpesvirus 1, Suid and Oecomys paricola) collected at the herpesvirus 1, and Equid herpesvirus 4 . Allpahuayo Biological Station in north The type species of the third genus is east Peru. A New World arenavirus Gallid herpesvirus 2 and the fourth is belonging to the Tacaribe complex, Gallid herpesvirus 1. Psittacid herpesvirus genetically related to Pichinde virus. 1 is the only unassigned species in the Not known to cause disease in humans. subfamily. Moncayo AC et al (2001) Virology 284 , 277 Synonym : herpes simplex virus group. McGeoch DJ et al (2000) J Virol 74 , 10401 Almeirim virus (ALMV) A serotype of Minson AC (2005) in Topley & Wilson’s Changuinola virus in the genus Orbivirus Microbiology and Microbial Infections , vol. belonging to the Changuinola virus 1, edited by BWJ Mahy and V ter Meulen. serogroup. Isolated from Lutzomyia spp. London: Hodder Arnold, p. 506 Roizman B et al (1992) Arch Virol 123 , 425 Almpiwar virus (ALMV) An unas- alpha interferon A class of human inter- signed rhabdovirus. Isolated from a feron produced primarily by leukocytes skink, Ablepharus boutonii virgatus , in following stimulation by viruses. The Northern Queensland, Australia. Not biologically active molecule consists reported to cause disease in humans. of 166 amino acids. Alpha interferon is used as a treatment for hairy cell leuke- alphafetoprotein A serum protein (the mia and AIDS-related Kaposi’s sar- fetal equivalent of albumin) which can coma. It is also moderately effective as be used to monitor the development a treatment for chronic infection with of hepatocellular carcinoma (HCC) hepatitis viruses B and C. Synthetic resulting from infection with hepatitis alpha interferon is now produced in B virus. Normal levels (less than 20 ng/ various forms by recombinant DNA ml) may rise above 1000 ng/ml in the technology for drug use. Encoded in presence of HCC. chromosome 9. At least 20 genes and 9 pseudogenes encoding alpha interferon Alphaherpesvirinae A subfamily of the have been cloned. family Herpesviridae . The nucleotide sequences of the subfamily members Pestka S et al (1987) Ann Rev Biochem 56 , 727 form a distinct lineage within the family. Replicate rapidly, usually with CPE Alphanodavirus A genus in the family in fibroblasts in culture or epithelial Nodaviridae containing five species iso- cells in vivo . Many members cause lated from insects. The type species is vesicular epithelial lesions in their Nodamura virus . Virions are spherical, natural hosts. Latent infection is often 32–33 nm in diameter with icosahedral demonstrable in nerve ganglia. Host symmetry (T 3). Virions are stable range is very variable. DNA mol. wt. at acid pH. The genome consists of 85–110 1 0 6 (120–180 kb). There are two molecules of positive sense single- two unique sequences separated by stranded RNA, 3.1 and 1.4 kb in length. other sequences which repeat in an Nodamura virus can infect both verte- inverted orientation. Genes homolo- brates and invertebrates, and there is gous to human herpesvirus 1 are found serological evidence that it may natu- within the unique short sequence (Us) rally infect pigs and herons. The virus and flanking inverted repeats (Irs and is also transmissible to suckling mice TRs). Four genera are identified so far: by Aedes aegypti mosquitoes, and will Simplexvirus, Varicellovirus, Mardivirus , also replicate in yeast cells. and Iltovirus. The type species of the Johnson KN et al (2001) J Gen Virol 82 , 1855 first is Human herpesvirus 1 , and other Price BD et al (2005) J Virol 79 , 495 species are herpesvirus B (Cercopithecine herpesvirus 1 ), Human herpesvirus 2 and Alpharetrovirus A genus in the subfamily Bovine herpesvirus 2 . The type species Orthoretrovirinae that includes avian 15 alternate reading frames

retroviruses. The type species is Avian Table A1. The 29 species of Alphavirus leukosis virus. Virus particles have a C-type morphology. The genome is Aurá (C) 7.2 kb in length (one monomer), with Barmah Forest (C) four genes: gag, pro, pol, and env . The Bebaru (C) LTR is U3-250, R-20, U5-80 nt in size. Cabassou (Ci) Many members of the genus contain Chikungunya * (C, H, B, Ba) additional transduced oncogenes. The Eastern equine encephalitis * (C, A, Cu, H, R, natural host range is restricted to birds, B, M) Everglades * (C, A, R, M) although some will infect mammalian Fort Morgan (Ci) cells experimentally. The oncogene- Getah (C, A) containing viruses can be divided into Highlands J (C, R, B) two groups: (1) Replication competent Mayaro * (C, H) viruses, e.g. Rous sarcoma virus and (2) Middelburg (C) Replication defective viruses, e.g. avian Mosso das Pedras (R) myeloblastosis virus, avian sarcoma Mucambo * (C, H, R, B, M) virus, and Fujinami sarcoma virus. Ndumu (C) Synonym : ‘ Avian type C retroviruses. ’ O’Nyong-Nyong * (A, H) Pixuna (C, A, R) Payne LN and Purchase HG (1991) In Rio Negro (C) Diseases of Poultry , Ninth edition, edited Ross River * (C, H, B, M) by BW Calnek et al . Ames: Iowa State Salmon pancreas disease (F) University Press, p. 386 Semliki Forest * (C, A, B) Sindbis * (C, A, H, B) Southern elephant seal (L, S) Alphaviridae A misnomer – alphaviruses Tonate * (B, C, Ci) form a genus, Alphavirus, in the family Trocara (C) Togaviridae . Una (C, A) Venezuelan equine encephalitis * (C, A, H, R, Ba, M) Alphavirus A genus in the family Western equine encephalitis * (C, A, H, R, B) Togaviridae , the type species of which is Whataroa (C) Sindbis virus . All species in this genus except salmon pancreas disease virus Isolated from: A, anopheline mosquitoes; B, multiply in mosquitoes or other arthro- birds; Ba, bats; C, culicine mosquitoes; Ci, pods as well as in vertebrates. All are Cimicidae; Cu, Culicoides; F, fish; H, humans; L, lice; M, marsupials; R, rodents; S, serologically related to each other but seals. not to other members of the family. * Can cause disease in humans. The HAI test is best for demonstrating an antigenic relationship between members of the genus and the CF test or neutralization test for differentiating between members. Cause encephalitis belonging to the Changuinola virus on i.c. injection in suckling mice. The serogroup. Isolated from phlebotomine positive-stranded genomes of alphavi- sandflies in the Amazon region of Brazil. ruses are 11–12 kb in size, exclusive of alternate reading frames The produc- the 3 terminal poly (A) tract. There are tion of more than one mRNA from a 29 currently known species, as listed in single coding sequence, exemplified Table A1 . by members of the genus Respirovirus . Calisher CH et al (1980) Intervirology 14 , 229 The P gene of respiroviruses encodes Kielian M (1995) Adv Virus Res 45 , 113 an mRNA that encodes the P protein Strauss JH and Strauss EG (1994) Microbiol (568–603 amino acids long), but a sec- Rev 58 , 491 ond open reading frame near the 5 - end of the mRNA is translated into a ALT See alanine aminotransferase . number of proteins named C, C, ’ Y1, and Y2. Sendai virus encodes all four Altamira virus (ALTV) A serotype of such proteins, but other members of Changuinola virus in the genus Orbivirus the genus may only encode one or two. alternate reading frames 16

All have the same carboxy-terminus, Effective (about 70%) against influenza but differ in the start site in the mRNA, virus A but not against influenza virus which may be an AUG but in at least B, C nor measles. Amantadine-resistant one example is an ACG. They are all influenza virus mutants are readily iso- small (175–215 amino acids long) basic lated, and since 2006 there has been a proteins, and their functions are not great increase in the number of resist- completely understood. ant viruses isolated from 9% to more Latorre P et al (1998) J Virol 72 , 5984 than 90%. The drug does not inhibit viral attachment but acts at an early stage by blocking the function of the aluminium hydroxide (alum ) An adju- influenza M2 protein, a 97 amino acid vant used in many inactivated virus protein containing a transmembrane- vaccines to increase immunogenicity. spanning region of 19 amino acids that Vaccines containing alum lose potency is an important channel for entry of if the vaccine is frozen, however. protons into the virion during uncoating. The drug may cause some mental ALVAC An attenuated vaccine strain of disturbance in patients with a history Canarypox virus under development as of psychiatric disorder. Drowsiness, a potential poxvirus-vector vaccine for slurred speech, lethargy, dizziness, and use in humans. Replicates only mini- insomnia are side effects. The α -methyl mally in mammalian cells, but expresses derivative, rimantadine, reportedly recombinant genes sufficiently to cause produces fewer side effects. See riman- an immune response. tadine hydrochloride . Paoletti E (1996) Proc Natl Acad Sci 93 , 11354 Synonym : Symmetrel. Lamb RA et al (1994) In Cellular Receptors of alveolar macrophages Mononuclear Animal Viruses . Cold Spring Harbor: Cold phagocytes present in the respiratory Spring Harbor Press, p. 303 tract, loosely attached to the walls of pulmonary alveoli. Derived from circu- Amapari virus (AMAV) A species in lating monocytes which mature in situ . the genus Arenavirus , belonging to Respond to virus infection by release the Tacaribe antigenic group. Isolated of chemokines and pro-inflammatory from forest rodents ( Oryzomys capito ; cytokines such as tumor necrosis factor Neacomys guianae ) in the Amapari region (TNF)-alpha and interleukins. of northern Brazil. Produces plaques in Vero cell cultures. Not known to cause Amaas virus Synonym for variola minor disease in humans. virus. See Variola virus . Bowen MD et al (1996) Virology 219 , 285 α -amanitin A cyclic polypeptide. The Pinheiro FP et al (1966) Proc Soc Exp Biol Med 122 , 531 principal toxin found in the poison- ous mushroom, Amanita phalloides . A potent and selective inhibitor of nucle- Ambe virus A species in the genus oplasmic form II DNA-dependent RNA Phlebovirus, isolated from phlebotom- polymerase of eukaryotic cells. It binds ine sandflies in the Amazon region to the RNA polymerase and blocks of Brazil. Antigenically related to the RNA synthesis after initiation, prevent- Phlebotomus virus complex. ing chain elongation. Viruses which Tesh RB et al (1989) Am J Trop Med Hyg require cellular RNA polymerase II 40 , 529 for their replication, e.g. Adenoviridae, Arenaviridae, Orthomyxoviridae , and amber Name given to a triplet codon Retroviridae , are inhibited. (UAG) specifying chain termination. Fiume L and Wieland Th (1970) FEBS Lett 8 , 1 Other chain termination codons are ochre (UAA) and opal (UGA). None amantadine 1-Adamantanamine hydro- of these three trinucleotides will bind chloride. A primary symmetrical amine aminoacyl tRNA. Some cells contain with limited prophylactic activity. suppressor tRNAs which will cause the 17 amino acid

insertion of an amino acid instead of Arenavirus usually called the New terminating the protein chain at the site World Arenaviruses. Sometimes used of a chain termination codon. The name erroneously as a synonym for the ‘ amber ’ was obtained by translation of Tacaribe antigenic group viruses. Four the German name Bernstein, one of the of them are associated with human contributors to the original work. hemorrhagic fever: Guanarito virus in Venezuela, Junín virus in Argentina, amber mutant Virus with a genome Machupo virus in Bolivia, and Sabiá mutation resulting in a chain termina- virus in Brazil. Wild rodents are the tion codon UAG. See amber . natural hosts and transmission from humans to humans is rare. The viruses ambisense viruses Viruses with a single- have also been isolated from mites stranded RNA genome which use an and other ectoparasites, but it is doubt- ambisense expression strategy, with ful whether arthropods actually trans- partial genetic coding in the positive mit them. sense as well as the negative sense. Arenaviruses are an example, but it Johnson KM et al (1967) Prog Med Virol also occurs with some members of the 9 , 105 Bunyaviridae , the Circoviridae , and several plant viruses. American oyster reovirus 13p2V (13p2V) A strain of Aquareovirus A in the genus Bishop DHL et al (1986) Adv Virus Res 31 , 1 Nguyen M and Haenni AL (2003) Virus Res Aquareovirus. A reovirus that appeared 93 , 141 in fish cell cultures of bluegill, Lepomis macrochirus, that had been inoculated Amdovirus A genus in the family Parvo- with extracts of apparently normal viridae containing only one species – oysters, Crassostrea virginica . Double, Aleutian mink disease virus . Shares fea- encapsidated, icosahedral particles, tures with the Bocavirus and Parvovirus 75 nm in diameter, were found. On genera. The genome is single-stranded injection into bluegills, focal hepatic negative sense DNA 4748 nt in length. necrosis developed, and 40% of the fish Permissive replication occurs only in died. The genome consists of 11 seg- Crandell feline kidney cells, though ments of dsRNA distributed among restricted replication occurs in cells bear- three size classes. The virus has five ing Fc receptors, such as macrophages. major structural proteins. Will infect many species of Mustilidae Synonyms : bluegill hepatic necrosis such as badgers, martens, mink, otters, virus; 13P2 virus. skunks, raccoons, and weasels. The vir- Meyers TR and Hirai K (1980) J Gen Virol 46 , ion structure differs from that of bocavi- 249 ruses and parvoviruses, and resembles Winton JR et al (1987) J Gen Virol 68 , 353 more closely that of dependoviruses. Wolf K (1988) Fish Viruses and Fish Viral This is due to the presence of three Diseases. Ithaca: Cornell University Press mounds elevated above the capsid surface around the threefold icosahedral American Type Culture Collection A axis of symmetry. repository containing a large collec- Dworak LJ et al (1997) Arch Virol 142 , 363 tion of cell lines and viruses of certified authenticity. The details of the collection Amdur II cells (CCL 124) Cell line devel- are available at http://www.atcc.org oped from a skin biopsy of a 1-year-old Caucasian boy with methylmaloni- amino acid Basic unit of proteins, con- cacidemia, a genetic disease. taining an amino (—NH2 ) and a car- Mellman WJ (1969) Pediatr Res 3 , 217 boxyl (—COOH) group and a variable side chain which determines the prop- American ground squirrel herpesvirus erties of the individual amino acid. Synonym for sciurid herpesvirus 1. The side chain may be simple (glycine) or a complicated ring structure (tryp- American hemorrhagic fever viruses tophan). There are 20 commonly occur- A group of 17 species in the genus ring amino acids in nature from which amino acid 18

proteins are synthesized during ribos- by affecting the permeability of the omal translation of mRNA. cytoplasmic membrane. aminoacyl-tRNA An amino acid attached amphotropic murine type C virus A to its specific tRNA by covalent link- strain of Murine leukemia virus in the age between the carboxyl group on the genus Gammaretrovirus which infects amino acid and the 2 or 3 hydroxyl and replicates in murine and non- group on the ribose at the 3 end of the murine cells. It thus differs from the tRNA. In this form the amino acid is ecotropic and xenotropic subspecies. said to be ‘ activated. ’ Further differentiation is possible on the basis of antigenicity and interfer- aminoacyl-tRNA synthetases (amino ence. Neither syncytia nor plaques are acid-RNA ligases) Enzymes which induced in XC cells. The virus was iso- bring about the covalent bonding of lated from wild mice. amino acids to their specific tRNAs. Hartley JW and Rowe WP (1976) J Virol Each enzyme is capable of selecting 19 , 19 one of the 20 amino acids and uniting it with its specific tRNA. Once charged amphotropic virus A virus which will with their amino acids, the tRNA mol- replicate in the cells of one or more ecules are ready to provide them to the species in addition to its natural host. protein-synthesizing system. Term usually applied to members of the family Retroviridae . p-amino-benzaldehyde, 3-thiosemicarbazone One of the first antiviral compounds to be developed, which inhibits amplicon 1 . The product nucleic acid replication of vaccinia virus. obtained from a polymerase chain reaction. 2. A cloning-amplifying vector Brownlee KA and Hamre D (1951) J Bacteriol based on repeat units of herpes simplex 61 , 127 virus (HSV) defective genomes and containing both bacterial and HSV origins of aminopeptidase An amino acid cleaving DNA replication and viral genome pack- enzyme which is found on the surface aging and cleavage signals. Can replicate of certain cells where it may act as the in the presence of standard HSV helper receptor for attachment of coronavi- virus. Co-transfection of cells with helper ruses and possibly certain retroviruses. virus DNA and amplicon generates concatameric defective genomes composed aminopterin A folic acid analog that of multiple reiterations of the repeats. inhibits dihydrofolate reductase activ- Foreign DNA sequences can be intro- ity. See HAT selection . duced to form concatameric chimeric defective genomes that are efficiently aminotransferase An enzyme which packaged and can be stably propagated transfers an amino group between in serially passaged virus stocks. amino acids. Alanine aminotransferase (ALT) and aspartate aminotransferase Advani SJ et al (2002) Clin Microbiol Infect (AST) are commonly measured in 8 , 551 serum to provide an indication of hepatocellular damage, which causes ele- amplified reverse transcriptase (Amp- vated levels of these enzyme activities. RT) An ultrasensitive detection method for the presence of the enzyme, reverse ammonium sulfate Salt commonly used transcriptase, which depends upon to precipitate enzymes, proteins, and PCR amplification of the DNA product viruses without denaturation. Used of the enzyme. frequently in the initial stages of protein purification as proteins precipitate Amprenavir An antiretroviral drug at different concentrations of the salt. which binds to the active site of HIV-1 protease, preventing processing of the amphotericin B An antibiotic produced gag and gag-pol polyprotein precursors by Streptomyces nodosus which operates resulting in the formation of immature 19 anatid herpesvirus 1 (AnHV-1)

non-infectious particles. Also known as ana1-ana2 Serotype designation given to Agenerase. fowl adenovirus A derived from the duck, Anas domestica . Amur tiger prion protein The amino acid sequence of the cellular prion pro- Anajatuba virus A probable species in tein of the Amur tiger (Panthera tigris the genus Hantavirus , identified from altaica ) was determined by sampling Fornes ’ colilargo ( Oligoryzomys fornesi ) the blood of 25 Amur tigers in zoos in Brazil. Has been associated with in China. The sequence was found to hantavirus pulmonary syndrome cases be similar to that found in house cats in Maranhao State, Brazil. (Felis domesticus ). analytical ultracentrifuge Instrument Amur virus A strain of Hantaan virus used to sediment macromolecules at in the genus Hantavirus. Found in high centrifugal force with appropriate Apodemus peninsulae rodents from optics so that the rates of sedimenta- Far East Russia, and subsequently in tion and diffusion of macromolecules patients from China. A closely related can be measured with great accuracy. virus known as Soochong virus was Markham R (1962) Adv Virus Res 9 , 241 recently isolated from A peninsulae in Korea, and it may be another isolate of Ananindeua virus (ANUV) A strain Amur virus. of Guama virus in the genus Jiang J et al (2007) J Med Virol 79 , 1792 Orthobunyavirus belonging to the Lokugamage K et al (2004) Virus Res 101 , 127 Guama serogroup. Isolated in Para, Brazil from marsupials, Caluromys amyloid A pathologic extracellular pro- philander (the woolly opossum) and teinaceous substance which is depos- Didelphis marsupialis , several species of ited in the form of insoluble fibrillar mosquitoes (especially culicine) and proteins in various organs and tissues a bird, the mouse-colored antshrike in amyloidosis, affecting vital functions. (Thamnophilus murinus) . Not reported to cause disease in humans. amyotrophic lateral sclerosis (ALS) A devastating motor neuron disease, also Anapu virus A probable species in the known as Lou Gehrig’s disease. First genus Orbivirus , isolated from phlebot- discovered in 1869, involving degen- omine sandflies in the Amazon region eration of upper motor neurons in the of Brazil. Antigenically related to the motor cortex and lower motor neu- Changuinola virus serogroup. rons in the brain stem and spinal cord. Can be familial, but 90% of cases anatid herpesvirus 1 (AnHV-1) An appear to be sporadic. Little is known unassigned species in the family of the etiology, but it was reported that Herpesviridae . Differs from other herpes- enterovirus RNA sequences related to viruses with respect to its structure and echovirus 7 are present in spinal cord maturation. Mature virus particles are neurons of patients with the disease. seen accumulated in long extensions of A similar disease is caused by Vilyuisk the endoplasmic reticulum; their size human encephalomyelitis virus, genome varies between 160 and 380 nm, and sequences of which are similar to those they are embedded in an osmiophilic found in Saffold virus, a newly isolated matrix. In addition, capsids (about member of the genus Cardiovirus . 80 nm in diameter) and developmental stages of the viral nucleoid (about Berger HH et al (2000) Neurology 54 , 20 40 nm in diameter) are encountered in the nuclei of infected cells. The genome AN 20410 virus A strain of Mopeia virus DNA has a high G C content (64.3%). in the genus Arenavirus . A natural infection in domestic ducks, and possibly of mallard, Anas platy- AN 21366 virus A strain of Mopeia virus rhynchos, in the UK. There is nasal and in the genus Arenavirus . ocular discharge, and diarrhea, with anatid herpesvirus 1 (AnHV-1) 20

up to 97% mortality. At post-mortem human transmission of Andes virus examination petechial bleedings in infection has been reported. mucosal membranes and many organs López N et al (1996) Virology 220 , 223 are prevalent. In less acute cases hem- Martinez VP et al (2005) Emerg Infect Dis orrhagic or pseudomembranous phar- 11 , 1848 yngitis, esophagitis and cloacitis are frequently observed. Typical herpes- Anellovirus A genus of non-enveloped, virus particles are observed by electron single-stranded (ss) DNA viruses which microscopy in the nucleus and the are commonly found in humans and cytoplasm of infected cells which are many other vertebrate species exam- of diagnostic relevance. Only ducks, ined, but have not so far been associ- geese, swans, and day-old chicks can ated with any disease. First reported be infected experimentally. The virus from Japan as a virus thought to be can be cultivated on the chorioallantoic associated with transfusion-related hep- membrane (CAM), killing the embryo atitis, and named after the initials of the in 4 days. Can be adapted to replicate patient (TT), subsequent investigation in chick fibroblast cultures, but loses showed that the virus was ubiquitous virulence. Attenuated virus vaccine is in human populations and not associ- effective in control of the disease. ated with hepatitis. The International Synonyms : anserid herpesvirus 1; duck Committee on Taxonomy of Viruses enteritis virus; duck plague herpesvirus. (ICTV) therefore renamed it Torque teno virus (TTV, retaining the initials TT). Gardner R et al (1993) Intervirology 36 , 99 Mallanna SK et al (2006) Virus Res 115 , 192 Virions of TTV are 30–32 nm in diameter, but a related virus less than 30 nm anchorage dependence Growth of nor- in diameter has also been found and mal fibroblasts or epithelial cells in named Torque teno mini virus (TTMV). culture which requires attachment to The genome of anelloviruses consists a glass or plastic surface in order to of circular ssDNA 3.5 to 3.8 kb in length divide. Such cells will not grow in liq- for TTV and 2.8 to 2.9 kb for TTMV. uid or semisolid suspension. This prop- Virions in human blood are bound to erty can be used to distinguish normal IgG forming immune complexes. TTV from virus-transformed cells, which has been found globally wherever it will grow in suspension in semisolid has been looked for. In African children media, such as soft agar. the infection seems to be acquired at an early age, and spread by saliva or feces Freedman VH and Shin S (1974) Cell 3 , 355 is suspected. In most human populations more than 80% infection rates Andasibe virus (ANDV) A tentative speare reported, making it the commonest cies in the genus Orbivirus . The Perinet human virus infection. In addition to virus was isolated in the same area and the human population, TTV and TTMV Andasibe is the Malagasy name of the have been found in non-human pri- locality, so they could be one and the mates such as chimpanzee, macaque, same virus, but Perinet virus is a tenta- tamarin, and douroucouli, and in cats, tive species in the genus Vesiculovirus . dogs, farm animals, and tupaias. Bendinelli M and Maggi F (2005) In Topley & Clerc Y et al (1984) Arch Inst Pasteur Wilson’s Microbiology and Microbial Infections , Madagascar 51 , 135 vol. 2, edited by BWJ Mahy and V ter Meulen. London: Hodder Arnold, p. 1276 Andes virus (ANDV) A species in the Bendinelli M et al (2001) Clin Microbiol Rev genus Hantavirus which caused fatal 14 , 98 human pulmonary disease in south- Prescott LE and Simmonds P (1998) N Engl J western Argentina. Genetically distinct Med 339 , 776 from, though related to, Sin Nombre virus. The presumed rodent reservoir angelfish herpesvirus Herpesvirus-like is Oligoryzomys longicaudatus . In con- particles were observed in sick South trast to other hantaviruses, human to American angelfish, Pterophyllum 21 anterior poliomyelitis virus

altum, imported to Denmark. Virus iso- sodium deoxycholate (DOC) or sodium lation has not been reported. dodecyl sulfate (SDS). Mellergard S and Bloch B (1988) Dis Aquat anisometric Adjective to describe virus Organ 5 , 151 particles that are not isometric, e.g. angelfish reovirus (AFRV) A strain of rod-shaped particles. Aquareovirus A in the genus Aquareo- annealing Synonym for hybridization. virus , associated with head and lateral line erosion disease of angelfish, Anopheles A group viruses There are six Pomecanthus semicirculatis . mosquito-borne viruses in this serologi- Varner PW and Lewis DH (1991) J Aquat cal group of the genus Orthobunyavirus , Anim Health 3 , 198 all isolated only from mosquitoes: angiotensin-converting enzyme 2 A met- Anopheles A -1940 alloprotease in the renin–angiotensin Anopheles A -CoAr 3624 system which causes vasoconstric- Anopheles A -ColAn 57389 tion, and is expressed on the surface Las Maloyas-AG8-24 of pneumocytes and enterocytes. Has Lukuni TRVL 10076 been identified as the main receptor Trombetas for severe acute respiratory syndrome All are considered serotypes of a single (SARS) coronavirus. species, Anopheles A virus . Guo Y et al (2008) Virus Res 133, 4 To KF and Lo AW (2004) J Pathol 203 , 740 Anopheles A virus (ANAV) A species in the genus Orthobunyavirus comprised anguillid herpesvirus 1 (AngHV-1) of six serotypes. Isolated from Anopheles An unassigned virus in the family sp in Colombia, South America. Not Herpesviridae , isolated from Japanese reported to cause disease in humans. and European eels which grows in an eel kidney cell line, has typical herpes- Anopheles B virus (ANBV) A species in virus morphology, and induces syn- the genus Orthobunyavirus . Isolated cytia and intranuclear inclusions. Has only from mosquitoes in South not been associated with morbidity or America. Boraceia virus is a closely mortality in eels. related serotype. Not reported to cause disease in humans. Kobayashi T and Miyazaki T (1997) Fish Pathol 32 , 89 Ueno Y et al (1992) Gyobyo Kenkyu 27 , 7 ansamycins Derivatives of rifamycin.

Anhanga virus (ANHV) A tentative ans1-ans3 Serotype designation given to species in the genus Phlebovirus . Not the fowl adenovirus 1 (CELO) derived serologically related to existing anti- from the goose, Anser domesticus . genic complexes of that genus. Isolated from the sloth, Choloepus didactylus , in anserid herpesvirus 1 Synonym for Castanhal Forest, Brazil. Not reported anatid herpesvirus 1. to cause disease in humans. ansteckende schweinelähmung virus Anhembi virus (AMBV) A strain of Synonym for porcine enterovirus. Wyeomyia virus in the genus Orthobun- yavirus , belonging serologically to the Antequera virus (ANTV) An unassigned Bunyamwera serogroup. Isolated from species of the family Bunyaviridae , sero- the rodent, Proechimys iheringi, and an logically belonging to group 5 with arthropod, Phoniomyia pilicauda, in São Resistencia and Barranqueras viruses. Paulo, Brazil. Not reported to cause Isolated from Culex (Melanoconion) del- disease in humans. pontei from Argentina. anionic detergent A detergent having anterior poliomyelitis virus Synonym a negatively charged surface ion, e.g. for Poliovirus . Antheraea cells, adapted 22

Antheraea cells , adapted (CCL 80) This can belong to several different classes, cell line was derived from the ovarian e.g. IgM, IgG, and IgA (secretory anti- tissues of the moth, Antheraea euca- body). See immunoglobulin . lypti , and constituted the first true line of arthropod cells established in cell antibody-dependent cell-mediated cyto- culture. Because of the difficulty and toxicity (ADCC) Specific killing of a expense in obtaining significant vol- target cell by cytotoxic T lymphocytes umes of lepidopteran hemolymph, (CTLs) or natural killer (NK) cells in the Antheraea cells were adapted to the presence of antibody. CTLs usually hemolymph-free culture medium. recognize a foreign (e.g. virus-induced) The adapted Antheraea cells are able protein and are restricted by MHC pro- to support the growth of a number of teins. Following recognition, the CTL arboviruses. releases molecules such as perforin which cause lysis of the target cell. NK anthroponosis A disease which is spread cells attach to antibody coating the tar- from humans to humans. get cell membrane via Fc receptors, and release toxin causing target cell death. anti-apoptosis Many viruses induce cell NK cells can also lyse uninfected cells killing by apoptosis, but with viruses bearing Fc receptors in the presence of that persist in their host, such as her- antibody to the T-cell receptor. pes viruses, they may actively inhibit apoptosis through a virus encoded antibody-dependent enhancement (ADE) protein or a virus encoded microRNA. Virions complexed to antibody can be In herpesviruses, during latent infec- taken up by cells by binding to the Fc tion the latency-associated transcript receptors on the cell surface, or in the encodes a microRNA which inhibits case of complement cascade by attach- apoptosis, thus contributing to the per- ment to the C3 receptor, circumvent- sistence of HSV in a latent form in sen- ing the requirement for a virus-specific sory neurons. receptor and enhancing viral pathogen- Gupta A et al (2006) Nature 442 , 82 esis. Microglial cells are prime targets Liu M and Vakharia VN (2006) J Virol 80 , for ADE in the central nervous system. 3369 Sarnow P et al (2006) Nat Rev Microbiol 4 , 651 antibody-mediated neutralization The antibiotic Substance used to inhibit the basis of the serum neutralization test, growth of microorganisms including in which antisera prepared against bacteria and fungi. They do not affect different viruses are allowed to react the growth of viruses. Many different after which the infectivity of the virus antibiotics are now available. Their is determined. An important technique principal application in virology is to used for the initial characterization of prevent bacterial and fungal growth in newly isolated viruses prior to genome tissue culture media used for the culti- sequence analysis. vation of cells and growth of viruses. anticodon A group of three consecutive antibody An immunoglobulin molecule bases in a tRNA molecule which recog- produced by lymphocytes following nizes a codon in an mRNA molecule. administration of a foreign protein The bases pair in an antiparallel man- or carbohydrate (antigen) into a ver- ner: A with U and G with C, at least as tebrate host. The induced antibody far as the first two bases in the codon reacts specifically with the adminis- are concerned. The pairing with the tered antigen. In virology, antibodies third base is more complicated as one are frequently used as a means of dis- tRNA can recognize several codons criminating between different viruses provided they differ only in the last using a range of serological techniques place. This is the ‘ wobble ’ hypothesis including virus neutralization, immu- which states that a certain amount of nodiffusion, complement fixation, variation or ‘ wobble ’ is tolerated in the ELISA, or Western blotting. Antibodies third nucleotide of the codon. 23 anti-idiotypes antigen Molecule of carbohydrate or pro- antigenic site The portion of a protein tein which stimulates the production which reacts with the antibody induced of an antibody, with which it reacts in response to the entire antigen. A specifically. complex antigen (e.g. a protein) will contain several antigenic sites. Also antigen–antibody complex A macromo- termed ‘ epitope. ’ lecular complex of antigen and antibody molecules specifically bound antigenic variation Altered antigenic- together. Important in pathogenesis ity resulting from genetic changes in a of immune complex diseases such as virus population which lead to resist- serum sickness or glomerulonephritis. ance to neutralization by antibodies. Occurs most commonly with RNA viruses such as influenza A virus, hep- antigen–antibody reaction The specific atitis C virus, and HIV. interaction between an antigen and an antibody which recognizes a struc- antigenome The complementary posi- tural feature of the antigen and binds tive RNA strand on which is made the to it. This reaction can be measured negative-strand genome of viruses by a variety of serological methods. of the order Mononegavirales such as See complement fixation test , ELISA , parainfluenza virus type 1 murine. immunodiffusion , neutralization , precipitin , radioimmunoprecipitation , and antigen-presenting cells T-lymphocytes Western blotting . are central for inducing immunological responses and the recognition and antigenemia Presence of circulating viral destruction of virus infected cells. antigen in the bloodstream. CD8 T-cells recognize a complex of MHC class I with a virus peptide on antigenic determinant The portion of infected cells. The endogenous antigen- an antigen which is recognized by the presenting pathway involves the deg- active site of an antibody. radation of newly synthesized viral proteins by a complex of proteolytic antigenic drift The appearance of a virus enzymes called the proteosome. Small with slightly changed antigenicity after peptides generated by this process are frequent passage in the natural host. actively transported by peptide trans- This is presumably due to selection of porters (TAP) to the endoplasmic retic- mutants under pressure of the immune ulum where they associate with MHC response. Commonly described in class I heavy chain. Peptides of about 9 influenza virus infections, but also amino acids can fit into the MHC cleft. observed with many other viruses. The light chain associates with the heavy Synonym : immunological drift. chain and the complex is transported to the cell surface for presentation. antigenic modulation Disappearance of membrane proteins from the surface anti-HBc Antibody to hepatitis B core of a cell after combination with specific antigen. antibodies, due to internalization of the complexed molecules. anti-HBe Antibody to hepatitis B e antigen. antigenic shift A sudden and major change in the antigenicity of a virus anti-HBs Antibody to hepatitis B surface resulting from genetic recombina- antigen. tion (gene reassortment). Most likely to occur in viruses with segmented anti-idiotypes Antibodies resulting from genomes, but only reported in influ- immunization with a specific antibody enza virus A to date. Occurred in 1957 (e.g. antiviral antibody). The result- when Asian influenza appeared, and ing anti-idiotype antibody can have a again in 1968 when Hong Kong influ- conformation which mimics the origi- enza appeared. nal antigen, and so can be used itself anti-idiotypes 24

to immunize and may induce neutral- Synonym : herpesvirus aotus 1 and 3. izing antibodies against the original Ebeling A et al (1983) J Virol 47 , 421 virus. Such anti-idiotype antibodies should provide a noninfectious anti- aotine herpesvirus 2 (AoHV-2) A strain genic mass which could form the basis of Bovine herpesvirus 4 (BHV-4) in the of a vaccine. Although protection has genus Rhadinovirus, isolated from a been demonstrated experimentally, no kidney cell culture of an owl monkey, anti-idiotype vaccines have yet been Aotus trivergatus. Does not cross-react developed. serologically with aotine herpesviruses 1 or 3. antimessage Viral RNA which is negative sense and cannot act as mRNA. It Barahona HH et al (1973) J Infect Dis 127 , 171 is transcribed by a viral transcriptase Bublot M et al (1991) J Gen Virol 72 , 715 to a positive-strand which functions as mRNA. The genome of negative-strand AP-1 transcription factor A nuclear tran- viruses and Delta virus is an antimes- scription factor complex which is the sage molecule. proto-oncogene transduced as the viral oncogenes v- fos (Finkel–Biskis–Jenkins antireceptor Virion surface protein which murine sarcoma virus) and v-jun (avian binds specifically to a cell surface sarcoma virus 17). receptor. See hemagglutination . Vogt PK (2001) Oncogene 20 , 2365 antisense oligonucleotides Oligonucle- Ape Aime-Itapua virus A probable spe- otides synthesized to represent the cies in the genus Hantavirus , identified complementary strand to the coding from the rodent Akodon montensis in strand (mRNA strand). Candidates for eastern Paraguay. possible use as inhibitors of virus rep- Chu YK et al (2006) Am J Trop Med Hyg lication since they can hybridize to the 75 , 1127 virus mRNA and prevent its expression. See siRNA . apelin A peptide which binds to an HIV coreceptor, APJ, and inhibits HIV repli- antisense RNA RNA which is comple- cation. See APJ . mentary (opposite sense) to a given mRNA, and may interfere with its Kleinz MJ and Davenport AP (2005) expression. Pharmacol Ther 107 , 198 Simons RW and Kleckner N (1983) Cell 34 , 683 Apeu virus (APEUV) A strain of Caraparu virus in the genus Orthobunyavirus , antiserum The serum from a vertebrate belonging serologically to the C group which has been exposed to an antigen viruses. Isolated from the woolly opos- and which contains antibodies that sum, Caluromys philander , the murine react specifically with the antigen. opossum, Marmosa cinerea , sentinel Cebus monkeys and mice. Also from mosquitoes in Para, Brazil. Has been antiviral agent A chemical compound associated with a few cases of febrile which inhibits virus replication. illness in humans. antiviral chemotherapy Treatment of Nunes MR et al (2005) J Virol 79 , 10561 virus diseases using drugs which inhibit or prevent virus entry, replica- aphidicolin A cyclic compound iso- tion or release. lated from the fungus, Cephalosporum aphidicola, which inhibits cellular DNA aotine herpesvirus 1 and 3 (AoHV-1 ; polymerase alpha and DNA polymer- AoHV-3) Two tentative species in the ases of vaccinia and herpesviruses. genus Cytomegalovirus . Found in owl Bucknall RA (1973) Antimicrob Agents monkeys, Aotus trivergatus . See also Chemother 4 , 294 Herpesviridae . de Filippes FM (1984) J Virol 52 , 474 25 Aquareovirus aphthous fever virus Synonym for Foot- apoptin A 13.6 kDa protein encoded by and-mouth disease virus . chicken anemia virus which induces apoptosis. aphthovirus Synonym for Foot-and-mouth Danen-van Oorschot AA et al (2000) J Virol disease virus . 74 , 7072 Heilman DW et al (2006) J Virol 80 , 7535 Aphthovirus A genus of the family Picornaviridae. Unstable below pH 6.8. apoptosis Mechanism by which many Buoyant density in CsCl: 1.43–1.45 g/ viruses induce cell death; transform- ml. Poly C tracts occur about 360 bases ing viruses encode proteins that inhibit from the 5 terminus of genome RNA. cellular apoptotic pathways. Plays a Three species of genome-linked protein major role in viral infections and in the (VPg) are encoded. All species of virus host response to them. CTLs, NK cells, share more than 50% sequence identity and cytotoxic cytokines all kill virus- over the entire genome. Type species infected target cells through apoptotic Foot-and-mouth disease virus O . pathways. In the infected host, T- lymphocyte apoptosis plays a role in the APJ A seven-transmembrane protein natural history of the T-cell responses which is a coreceptor for HIV found to viral infection. The immune response on nerve cells. A homolog of the angio- is silenced by the physiological elimina- tensin receptor. The natural ligand of tion of lymphocytes by apoptosis, and APJ, termed apelin, specifically inhibits an overzealous elimination can lead to the replication of HIV-1 virus in cells viral persistence. bearing APJ. Basu A et al (2006) Apoptosis 8 , 1391 Cayabyab M et al (2000) J Virol 74 , 11972 Razvi ES and Welsh RM (1995) Adv Virus Fan X et al (2003) Biochemistry 42 , 10163 Res 45 , 1 Shen Y and Shenk TE (1995) Curr Opin Genet Apoi virus (APOIV) A species in the genus Dev 5 , 105 Flavivirus belonging to the Modoc virus group. Isolated from healthy rodents, aptamers High affinity nucleic acid lig- Apodemus speciosus ainu and A argentus ands obtained by affinity and amplifi- hokkaidi , on the foothills of Mount Apoi, cation procedures. If prepared against Hokkaido, Japan. There is one report of a viral target, such as influenza virus infection in a laboratory worker who hemagglutinin, the aptamer will developed encephalitis. inhibit hemagglutinin-mediated membrane fusion and has more than 15-fold apolipoprotein B mRNA editing enzyme higher affinity for binding as compared A normal cell enzyme, also known with a monoclonal antibody. Aptamers as CEM15, which is a member of the can also be used as diagnostic tools, in cellular cytidine deaminase-editing addition to therapeutic applications. family. It is an inhibitor of HIV-1 rep- Gopinath SCB (2007) Arch Virol 152 , 2137 lication, causing hypermutation of the genome. However, the HIV-induced Aquabirnavirus A genus in the family Vif protein which binds to CEM15 Birnaviridae containing virus species induces its polyubiquitination and pro- which only infect fish, mollusks, and teosomal degradation. crustaceans. Isolated from a variety Mangeat B et al (2003) Nature 424 , 99 of aquatic animals, sometimes in the absence of disease. The type species is Apollo virus Synonym for enterovirus Infectious pancreatic necrosis virus , which 70. A name given to the virus isolated causes disease in a variety of salmonid in central western Africa from the first fish as well as Japanese eels. Other spe- group of cases of acute hemorrhagic cies in the genus are Tellina virus 2 and conjunctivitis. Named Apollo disease Yellowtail ascites virus . after the first Apollo–Saturn 11 moon- craft landing which took place about Aquareovirus A genus in the family the same time, in 1969. Reoviridae comprising at least six species Aquareovirus 26

(A–F) infecting fresh as well as seawater 9-β -D -arabinofuranosyladenine See ade- fish and some marine invertebrates such nine arabinoside . as clams and oysters. The genome consists of 11 segments of double-stranded 1-β- D -arabinofuranosylcytosine hydro- RNA (three large, three medium, and chloride See cytarabine hydrochloride . five small) ranging in length from 0.8 to 3.8 kb. On the basis of RNA–RNA 1-β -D -arabinofuranosylthymidine See hybridization, six genogroups are cur- spongothymidine . rently recognized. The type species, Aquareovirus A , includes at least 10 1-β -D -arabinofuranosyluracil See spongo- viruses which affect fish and oysters. uridine . Aquareoviruses have been isolated from wide geographic areas, and at least five arabinosyl adenine See adenine different viruses cause economically arabinoside . important diseases of finfish: the golden shiner, grass carp, smelt, angelfish, and grouper reoviruses. Most of the viruses arabinosyl cytosine See cytarabine replicate well in fish cell lines. hydrochloride . Hetrick FM and Hedrick RP (1993) Ann Rev ara C See cytarabine hydrochloride . Fish Dis , 187 Winton JR et al (1987) J Gen Virol 68 , 353 Araçai virus A probable species in the genus Orbivirus , isolated from phlebot- Aquareoviruses A–F (ARV-A–ARV-F) The omine sandflies in the Amazon region six species in the genus Aquareovirus , of Brazil. Antigenically related to the family Reoviridae, are recognized as Changuinola virus serogroup. distinct based on RNA–RNA hybridization. It is assumed that the members Aragão’s myxoma virus A strain of of a single species are able to exchange Myxoma virus from the South American genetic information by reassort- tapeti, Lepus brasiliensis (the Forest ment during mixed infection. Species rabbit). A includes members isolated from angelfish, Atlantic salmon, Chinook Araguari virus (ARAV) An unas- salmon, Chum salmon, Masou salmon, signed virus isolated from a Philander smelt, striped bass, and American oys- opossum in 1969 at Serra do Navio, ters; Species B includes isolates from Brazil. Virions are enveloped, with an Chinook salmon and Coho salmon; ‘ arenavirus-like ’ morphology. Pathogenic Species C, D, and E have one isolate for certain laboratory vertebrates and each, from golden shiner, Channel cell cultures. Eight RNA species. Three catfish, and turbot, respectively; and major polypeptides (67 kDa, 58 kDa, Species F has isolates from Coho 30 KDa; 1:2:3), two minor polypeptides salmon and Chum salmon. There are (43.5 kDa); 67 kDa and 30 kDa proteins also tentative species in the genus iso- appear to be glycoproteins (i.e. not lated from Chub, grass carp, hard clam, similar to arenaviruses). landlocked salmon, and tench. Zeller HG et al (1989) Arch Virol 108 , 191 Chodosh J et al (2008) in BWJ Mahy and MHV van Regenmortel (eds) Encyclopedia Aransas Bay virus (ABV) An unassigned of Virology , Third edition, Oxford: Academic species in the family Bunyaviridae , Press, vol. 2, p. 491 serologically related to Upolu virus. Isolated from Ornithodorus capensis in AR 86 virus A strain of Sindbis virus . Texas, USA. Not known to cause disease in humans. AR 339 virus A strain of Sindbis virus . Araraquara virus (ARAV) A probable spe- ara A See adenine arabinoside . cies in the genus Hantavirus , identified 27 Arenaviridae

from the rodent Necromys lasiurus in or (3) isolated from a vertebrate or an South America. Has been associated arthropod and shown to be antigeni- with hantavirus pulmonary syndrome. cally related to an established arbovi- Johnson AM et al (1999) J Med Virol 59 , 527 rus. A number of antigenic groups have been designated. An antigenic group is ara T See spongothymidine (Ara T) created when a newly discovered virus can be shown to be serologically related Aratau virus A probable species in the to, but clearly distinguishable from, a genus Orbivirus , isolated from phlebot- previously isolated arbovirus. The orig- omine sandflies in the Amazon region inal groups were A, B, and C but now of Brazil. Antigenically related to the new groups take the name of the first- Changuinola virus serogroup. discovered member of the group. Groups A and B form the genera Alphavirus and ara U See spongouridine (Ara U) Flavivirus, respectively, of the families Togaviridae and Flaviviridae . Other arbovi- Aravan virus (ARAV) A tentative species ruses belong to the families Arenaviridae, in the genus Lyssavirus, isolated from a Bunyaviridae, Reoviridae (genus Orbivirus), lesser mouse-eared bat (Myotis blythi ) and Rhabdoviridae . A few are unclassi- in southern Kyrgysztan in 1991. fied and there is one virus in each of the Kuzmin IV et al (2006) Dev Biol 125 , 273 taxa Iridoviridae, Paramyxoviridae, and Poxviridae . The name ‘ arbovirus ’ is not Arawete virus A probable member of the accepted as a legitimate taxonomic term genus Orbivirus , isolated from phlebot- since it has no relevance to chemistry, omine sandflies in the Amazon region morphology, or mode of viral replication. of Brazil. Antigenically related to the Karabatsos N (1978) Am J Trop Med Hyg 27 , Changuinola virus serogroup. Suppl 372

Arbia virus (ARBV) A serotype of Arbroath virus (ABRV) A serotype of Salehabad virus in the genus Phlebovirus , Great Island virus in the genus Orbivirus , belonging to the Rift Valley fever com- belonging to the Great Island antigenic plex. Isolated from Phlebotomus pernici- complex. Isolated from a pool of ticks, osus from Toscana, Italy. Not known to Ixodes uriae , found on a dead puffin, cause disease in humans. Fratercula articula, in Arbroath, Scotland in 1978. Arboledas virus (ADSV) A tentative species in the genus Phlebovirus belonging ARC See AIDS-related complex . to the sandfly fever antigenic group. Isolated from Lutzomyia spp from Norte arctic squirrel hepatitis virus (ASHV) de Santander, Colombia. Not known to cause disease in humans. A tentative species in the genus Orthohepadnavirus found in arctic Tesh RB et al (1986) Am J Trop Med Hyg 35 , ground squirrels in Alaska. 1310 Testut P et al (1996) J Virol 70 , 4210 Arboviridae Old name (no longer used) for a family encompassing the Flaviviridae ARDS See acute respiratory distress and Togaviridae families. syndrome . arbovirus A term ( arthropod- borne virus ) Arenaviridae (Latin: arenosus sandy) used to describe any virus of ver- A family of RNA viruses 50–300 nm in tebrates which is transmitted by an diameter (usually 110–130 nm), spherical arthropod. For inclusion in the catalog or pleomorphic. They have a dense lipid of arboviruses they must be: (1) iso- bilayer membrane covered with projec- lated from a vertebrate and shown to tions, surrounding a core which con- be infectious to an arthropod, (2) iso- tains ribosome-like particles, 20–25 nm lated from an arthropod and shown to in diameter. Virion proteins include two be pathogenic to a vertebrate, e.g. mice, glycopeptides and two polypeptides. Arenaviridae 28

Genome consists of two species of single- depletes local arginine concentrations, stranded RNA with mol. wt. of 2.3 1 06 and effectively aborts herpes simplex and 1.2 1 0 6 called L and S segments. virus infection. Viral RNA is transcribed by a virion Mistry SK et al (2001) Virus Res 73 , 177 polymerase into complementary RNA which acts as mRNA. Both the L and S Aride virus An unclassified arbovirus, segments are ambisense. The L segment isolated in suckling mice from a pool of encodes a large polymerase protein, L, in ticks, Amblyomma loculosum , collected negative sense, and a small zinc-binding from dead roseate terns on Bird Island, protein, Z, in plus sense. The S segment Seychelles. Not known to cause disease encodes a nucleocapsid, N protein in in humans. negative sense, and a glycoprotein, GPC, in plus sense. Virus is synthesized in Converse JD et al (1976) Arch Virol 50 , 237 the cytoplasm and matures by budding through the cell membrane. Infectivity arildone An aryl-alkyl-diketone (4-[6- sensitive to lipid solvents. There is a sin- (2chloro-4-methoxyphenoxy) hexyl]- gle genus: Arenavirus . The name was at 3,5heptanedione) which inhibits first spelt arenovirus (arenosus sandy) replication of several viruses, includ- but this was altered to avoid confusion ing human poliovirus, at non-cytotoxic with the adenoviruses. concentrations. Probably acts by inhibiting uncoating. Bishop DHL and Auperin DD (1987) Curr Top Microbiol Immun 133 , 5 Diana GD et al (1977) J Med Chem 20 , 750 Salvato MS (Editor) (1993) The Arenaviridae . McSharry JJ et al (1979) Virology 97 , 307 New York: Plenum Press Ariquemes virus A probable species in the Arenavirus The only genus in the fam- genus Phlebovirus, isolated from male ily Arenaviridae. The type species is phlebotomine sandflies in Ariquemes, Lymphocytic choriomeningitis virus . All Rondonia State, in Brazil. Antigenically species are antigenically related but related to the Phlebotomus serogroup. can be divided into two phylogenetic and serological groups: (1) the Old Arkonam virus (ARKV) A serotype World Arenaviruses, e.g. Lassa virus of Ieri virus in the genus Orbivirus . and Lymphocytic choriomeningitis virus Isolated from the mosquitoes Anopheles and (2) New World Arenaviruses, e.g. subpictus, A hyrcanus, and Culex tritaen- the Tacaribe antigenic group viruses. iorhynchus in Tamil Nadu, India. Not Most species have a single rodent or bat reported to cause disease in humans. host in which they cause a persistent infection with viremia and/or viruria. Armstrong virus A strain of Lymphocytic Bowen MD et al (2000) J Virol 74 , 6992 choriomeningitis virus in the genus Salvato MS and Rodas JD (2005) In Topley & Arenavirus . Wilson’s Microbiology and Microbial Infections , vol. 2, Tenth edition, edited by BWJ Mahy Aroa virus (AROAV) A mosquito-borne and V ter Meulen V. London: Hodder species in the Flavivirus genus. Related Arnold, p. 1059 strains are Bussuquara virus, Iguape virus, and Naranjal virus. Argentina virus Synonym for the Indiana Vesicular stomatitis virus 2 A strain of . Arsenic trioxide A drug which causes apoptosis of cells transformed by Argentine hemorrhagic fever virus human T-cell lymphotropic virus type Junín virus Synonym for . I, and acts synergistically in combination with alpha-interferon. Argentine turtle herpesvirus Synonym for chelonid herpesvirus 4. Bazarbachi A et al (1999) Blood 93 , 278 arginase It has been shown that one anti- Arteriviridae A family of enveloped viral mechanism used by macrophages positive-stranded RNA viruses that is the production of arginase, which was established in 1996; contains a 29 aseptic meningitis

single genus Arterivirus. Arteriviridae , isolated from anopheline mosquitoes Coronaviridae , and Roniviridae make up in the Amazon region of Brazil. the order Nidovirales . Snijder EJ and Meulenberg JJM (1998) J Gen Arumowot virus (AMTV) A tentative Virol 79 , 961 species in the genus Phlebovirus not serologically related to other mem- Arterivirus A genus of positive single- bers of that genus. Isolated from the stranded RNA viruses which includes mosquito, Culex antennatus, the mice, Equine arteritis virus , Lactate dehydroge- Thamnomys macmillani and Lemniscomys nase-elevating virus , Porcine respiratory striatus , a shrew of Crocidura sp, the and reproduction syndrome virus , and Kusu rat, Arvicanthis niloticus , and Simian hemorrhagic fever virus . Virions are Kemp’s gerbil, Tatera kempi , in Sudan, 60 nm in diameter having an isometric Nigeria, Ethiopia, and the Central nucleocapsid 35 nm in diameter and an African Republic. Not reported to envelope containing 12–15 nm ring-like cause disease in humans. surface structures. Virion density (CsCl): 1.17–1.20 g/ml. The genome RNA is ARV AIDS-related virus. See Human 13 kb in length, positive-stranded, and immunodeficiency virus type 1 . infectious with a 5 terminal cap and a 3 terminal poly A tract. Leader- ascitic fluid Fluid from the peritoneal containing mRNAs are synthesized cavity of mice. Used in raising mono- during replication forming a ‘ nested- clonal antibodies in large quantities fol- set’ similar to Coronaviridae . Individual lowing injection of hybridoma cells into viruses are not related antigenically and the peritoneal cavity. Can also be used each infects a different single species. in the culture of encephalomyocarditis The primary host cells are macrophages. virus. No arteriviruses infecting humans have been identified. aseptic lymphocytic choriomeningitis Plagemann PGW and Moennig V (1992) Adv See Lymphocytic choriomeningitis Virus Res 41 , 99 virus . Snijder EJ and Meulenberg JJM (1998) J Gen Virol 79 , 961 Snijder EJ et al (2003) J Virol 77 , 97 aseptic meningitis Meningitis in which the raised cell count in the CSF is predominantly lymphocytic, and the pro- arthropod-borne Virus which multiplies tein level is only moderately raised in an arthropod and is transmitted to (80–120 mg/ml). Most commonly its animal or plant host(s) by this route. caused by a viral infection, but the term is not synonymous with viral men- artificial top component Name given to ingitis because the condition may be picornavirus empty capsids produced caused by leptospirosis, syphilis, tuber- by heating virions. culosis, brucellosis, cryptococcosis, infiltration of the meninges with malig- Aruac virus (ARUV) An unassigned ver- nant or granulomatous tissue, cerebral tebrate rhabdovirus. Isolated from the abscess and meningeal infiltration in mosquito, Trichoprosopon theobaldi , in collagen disease, or following the intro- Trinidad. Not reported to cause disease duction of drugs or contrast media. in humans. The common viral causes are enteroviruses, mumps virus, lymphocytic cho- Aruana virus (ARNV) A probable species riomeningitis virus and arboviruses. in the genus Orbivirus , isolated from Enteroviruses of most types have been phlebotomine sandflies in the Amazon isolated from cases of aseptic menin- region of Brazil. Antigenically related gitis. Outbreaks have been associated to Changuinola virus . with echovirus 2, 3, 4, 7, 9, 11, 14, 15, 16, 17, 18, 19, 25, 27, 30, 33, coxsackie Arumateua virus (ARMV) A probable virus A7, A9, B1, B2, B3, B4, B5, B6, and species in the genus Orthobunyavirus , enterovirus 71. Asfarviridae 30

Asfarviridae (Sigla: A frican s wine f ever asinine herpesvirus 2 Synonym for Equid a nd related) A family consisting of one herpesvirus 7 , a species in the genus genus Asfivirus. The family description Rhadinovirus . corresponds to the genus description. asinine herpesvirus 3 Synonym for Equid Asfivirus A genus with only one species, herpesvirus 8 , a species in the genus African swine fever virus , in the family Varicellovirus . Asfarviridae. Viruses consist of a nucleoprotein 70–100 nm in diameter, sur- assembly During virus replication, the rounded by internal lipid layers and an formation of mature virions from com- icosahedral capsid, 200 nm in diameter. ponent nucleic acid and proteins. The DNA genome is linear, covalently closed-ended double-stranded DNA, Assurinis virus A probable species in the 170–190 kb in length, encoding 150–200 genus Orbivirus , isolated from phlebot- proteins. Virus replication occurs in omine sandflies in the Amazon region swine macrophages in vitro and in of Brazil. Antigenically related to the vivo . Virus morphogenesis takes place Changuinola virus serogroup. in virus factories. Domestic and wild swine are the only natural hosts, but Astra virus An isolate of Dhori virus made soft ticks of the genus Ornithodorus are from the tick, Hyalomma plumbeum , and also infected, and probably maintain the the mosquito, Anopheles hyrcanus , in the virus in nature. Virus can be transmit- former USSR. ted in ticks trans-stadially, trans-ovari- ally, and sexually. Warthogs, bushpigs, Butenko AM and Chumakov MP (1971) Aka and swine can be infected by bites from Med Nauk SSR Part 2 , 11 , 1 infected ticks. The disease is endemic in many African countries and in Sardinia. Astroviridae A family containing two genera, Avastrovirus and Mamastrovirus . Dixon LK et al (1994) J Gen Virol 75 , 1655 Virions are spherical, 28–30 nm in diam- Tulman ER and Rock DL (2001) Curr Opin Microbiol 4 , 456 eter, and non-enveloped. A distinctive star is seen on the surface of about 10% Ash river virus A probable species in of the virions, from which the family the genus Hantavirus , identified from name is derived. The genome is posi- the masked shrew (Sorex cinerus ) in tive sense, single-stranded RNA, of Minnesota, USA. length 6.4–7.4 kb. At least eight serotypes of human astroviruses have been Arai S et al (2008) Am J Trop Med Hyg 78 , 348 defined by immuno-electron microscopy and neutralization tests, and Ashy-headed sheldgoose hepadnavirus confirmed as genotypes. Astroviruses (ASHBV) A virus detected by DNA have also been described from a wide cloning of tissues from the ashy-headed variety of mammals and birds. sheldgoose (genus Chloephaga ) and complete sequencing. Related to duck Jiang B et al (1993) Proc Natl Acad Sci 90 , hepatitis B virus, and so an assumed 10539 member of the genus Avihepadnavirus . Noel J et al (1995) J Clin Microbiol 33 , 797 Guo H et al (2005) J Virol 79 , 2729 asymptomatic infections Virus infections which produce no apparent symp- Asian influenza virus The cause of the toms in the host. Also known as silent 1957 influenza pandemic. See Influenza infections. A virus .

Asibi virus The original virulent strain of Atadenovirus A genus in the family Yellow fever virus from which the first Adenoviridae , which would accom- attenuated vaccine strain was derived. modate certain avian, bovine, ovine, and reptilian adenoviruses which asinine herpesvirus 1 Synonym for Equid have a different genome organization herpesvirus 6, a tentative species in the from members of the Mastadenovirus, genus Varicellovirus . Siadenovirus, and Aviadenovirus genera. 31 Atlantic salmon papillomatosis virus

The name reflects the high AT-content, Synonym : herpesvirus ateles 2. which may be up to 65% AT in some Albrecht J-C (2000) J Virol 74 , 1033 regions of the DNA. The type species is Albrecht J-C et al (2004) J Virol 78 , 9814 Ovine adenovirus D, and other species in Melendez LV et al (1972) Nature (New Biol) the genus are Bovine adenovirus D, Duck 235 , 182 adenovirus A , and Possum adenovirus . Duck adenovirus A causes egg drop ateline herpesvirus 3 (AtHV-3) An syndrome in poultry worldwide. There unassigned member of the family are several tentative species in the Herpesviridae . Originally isolated from genus, including adenoviruses from a cell culture of kidney tissue from a reptiles such as the bearded dragon, Guatemalan spider monkey, Ateles geof- chameleon, gecko, and snake. froyi, which developed characteristic Benko M and Harrach B (1998) Arch Virol herpes-type CPE. Four isolates from 143/144 , 829 peripheral lymphocytes of Colombian Benko M et al (2002) J Virol 76 , 10056 spider monkeys, Ateles fusciceps robus- tus , are antigenically slightly differ- Wellehan JF et al (2004) J Virol 78 , 13366 ent. The virus appears to be a natural, horizontally transmitted infection of ataxia of cats virus Synonym for Feline spider monkeys in which it rarely, if panleukopenia virus . ever, causes disease. It is very similar in behavior to cebine herpesvirus 2 caus- Atazanavir A protease inhibitor which ing lymphomatous neoplasms in mar- prevents cells infected with HIV from mosets, owl monkeys, and other species. producing new virus. It is used in com- Marmoset lymphocytes are transformed bination with at least two other drugs by it in vitro and the transformed cells in HIV therapy. have T-cell markers. Synonym : Reyataz Synonym : herpesvirus ateles strain 73. Prober CG and Kimberlin AW (2005) In Deinhardt FW et al (1974) Adv Cancer Res Neonatal and Pediatric Pharmacology , Third 19 , 167 edition, edited by SJ Yaffe and JV Avanda. Falk L et al (1978) Int J Cancer 21 , 652 Philadelphia: Lipincott, Williams & Wilkins, Luetzeler J et al (1979) Arch Virol 60 , 59 p. 475. atherosclerosis Many studies have Ateline herpesvirus 1 (AtHV-1) A spe- attempted to link virus infection with cies in the genus Simplexvirus , sub- vascular disease. There is some experi- family Alphaherpesvirinae, isolated from mental evidence that human herpes- a fatal infection of a 5-month-old virus 5 (human cytomegalovirus) may female spider monkey, Ateles geoffroyi , play a role in atherosclerosis, but this is born in a Californian zoo. The virus by no means conclusive. kills suckling mice and marmosets on inoculation. Libby P (2002) Nature 420 , 868 Synonym : spider monkey herpesvirus. Nieto FJ (2002) Semin Vasc Med 2 , 401 Stassen FR et al (2006) J Clin Virol 35 , 349 Hull RN et al (1972) J Natl Cancer Inst 49 , 225 Atlantic cod nervous necrosis virus Ateline herpesvirus 2 (AtHV-2) A species (ACNNV) A tentative species in the in the genus Rhadinovirus , subfamily genus Betanodavirus . Gammaherpesvirinae, which infects spider monkeys. The virus was isolated Atlantic cod ulcus syndrome virus from kidney cell cultures of a mature Synonym for Viral hemorrhagic septi- male spider monkey, Ateles geoffroyi , cemia virus . from Guatemala. The virus is oncogenic in marmosets and owl monkeys Atlantic halibut nodavirus A tentative (Aotus trivirgatus), which develop species in the genus Betanodavirus . malignant lymphomas with leukemia. Grotmol S et al (2000) Dis Aquat Organ 39 , 79 The virus genome encodes an oncoprotein, Tio, that mediates the transform- Atlantic salmon papillomatosis virus ing phenotype. A probable species in the family Atlantic salmon papillomatosis virus 32

Herpesviridae . Examination of wart- electrostatic forces but independent of like lesions among Atlantic salmon temperature except that collisions are revealed herpesvirus-like particles, more frequent at higher temperatures. 200–250 nm in diameter, as well as Absence of suitable receptors can give non-enveloped 110 nm particles within a cell immunity from infection. The cell nuclei. There has been no reported virion protein which binds to the cell is virus isolation. called the ‘ antireceptor. ’ Synonym : salmon wart disease virus. Lonberg-Holm K and Philipson L (1974) Carlisle JC (1977) J Wildl Dis 13 , 235 Monogr Virol 9 , 148 Schelkunov IS et al (1992) Bull Eur Assoc Fish Pathol 12 , 28 attachment interference See interference . Atlantic salmon paramyxovirus (ASPV) A paramyxovirus first isolated from attachment proteins The protein(s) on a the gills of farmed Atlantic salmon suf- virus which attach to the cell receptor fering from proliferative gill inflam- in the initial stage of infection. mation in 2003. The complete genome sequence revealed a close relationship attack rate In epidemiology, the number to respiroviruses, in the subfamily of cases of a disease incident to a Paramyxovirinae . defined population over a defined time Falk K et al (2008) Virus Res , 133 , 218 period. Also called ‘ disease incidence. ’ Nylund S et al (2008) Virology 373 , 137 attenuated virus strains Mutant strains Atlantic salmon reovirus HBR (HBRV) with low virulence or which are aviru- A strain of Aquareovirus A in the genus lent for their natural host species, and Aquareovirus . in which they can thus be used as a vaccine. Often obtained by passage in Atlantic salmon reovirus TSV (TSRV) cell culture or in a host different from A strain of Aquareovirus A in the genus the one in which they usually cause Aquareovirus . disease.

Atlantic salmon swim-bladder fibrosa- attenuation The process of producing an rcoma virus A probable species in the attenuated virus strain. family Retroviridae . Particles resembling a C-type retrovirus were seen in A-type inclusion body See Cowdry type fibrosarcomas on the swim bladder of A inclusion bodies . Atlantic salmon in a commercial cage culture in Scotland. Histologically, the A-type virus particles A term used origi- tumors were classified as leiomyosa- nally by electron microscopists to des- rcomas. There has been no reported ignate a morphologically defined group virus isolation. of RNA virus particles, often found in tumor cells. They are double-shelled Duncan IB (1978) J Fish Dis 1 , 127 spherical particles, appearing in thin McKnight IJ (1978) Aquaculture 13 , 55 sections as two concentric rings, the outer with a diameter of 65–75 nm and AtT-20 cells (CCL 89) An ACTH-secret- the inner with a diameter of approxi- ing line cloned from earlier cultures mately 50 nm. The inner ring usu- established after alternate passage of ally appears more dense. The center mouse pituitary tumor cells as tumors is electron-lucent but contains some in animals and in cell culture. amorphous material. They are always intracellular and morphologically simi- attachment The first stage of infection lar but there are at least two groups. of a cell by a virus following chance (1) Intracytoplasmic particles within collision of the virus with a suitable the ground substance of the cytoplasm receptor on the cell. It is dependent on where they may form large paranuclear 33 Australian X-disease virus

masses within or close to the Golgi area. Synonyms: HBs Ag; hepatitis-associated They are typically seen in mouse mam- antigen. mary tumor cells but are also present Blumberg BS (1977) Science 197 , 17 in cells of lymphomas in mice with the mammary tumor virus. They are Australian bat lyssavirus (ABLV) A spe- intermediates in the assembly of B- or cies in the genus Lyssavirus , originally D-type virions. (2) Intracisternal A par- isolated from an Australian black fly- ticles (IAP) which appear in uninfected ing fox (Pteropus alecto ) from Ballina, rodent-derived cells, budding from New South Wales, Australia in 1996. the inner surface of the cisternae of the An additional isolate was made from endoplasmic reticulum. They are more a woman bat-carer who died of rabies- variable in size than the first group of like encephalitis in 1996, and further A-type particles and are not considered isolates were made from other spe- to be related to other morphological cies of flying-fox (P poliocephalus and P types such as B- and C-type particles. scapulatus ) and from an insectivorous They are believed to represent expres- bat (yellow-bellied sheath-tailed bat). sion of endogenous proviral genetic There are probably several strains of elements, present in high copy number ABLV circulating in Australia. The virus in rodent cells. is closely related to North American Bernhard W (1960) Cancer Res 20 , 712 and other strains of Rabies virus, and is Dalton AJ (1972) J Natl Cancer Inst 49 , 323 neutralized by antirabies serum. Synonym : Ballina virus. atypical measles A syndrome reported in Foord AJ et al (2006) Aust Vet J 84 , 225 children exposed to wild-type measles Speare R et al (1997) Commun Dis Intell virus who had received inactivated 21 , 117 measles vaccine 2–4 years previously. The symptoms were severe and Australian encephalitis (Murray Valley included an extensive maculopapular encephalitis) Name used for arbovi- rash and interstitial pneumonia. ral encephalitis in Australia. The most Frey HM and Krugman S (1981) Am J Med important cause is Murray Valley Sci 281 , 51 encephalitis virus. Mackenzie JS (1994) Arch Virol 136 , 447 atypical pneumonia of rats See pneumonia virus of rats . Australian infectious bronchitis virus A strain of avian infectious bronchitis atypisches geflügelpest virus Synonym virus originally isolated by Cumming for Newcastle disease virus . in 1962 from poultry in Australia suffering from a kidney disease known as Aujeszky’s disease virus Synonym for uremia. Infectious bronchitis viruses pseudorabies virus (Suid herpesvirus 1 ). isolated in Australia before 1980 induced nephritis with high mortal- Aurá virus (AURAV) A species in the ity, whereas viruses isolated in the genus Alphavirus . Isolated from mos- USA and elsewhere outside Australia quitoes in Brazil. No known associa- showed little evidence of nephropatho- tion with disease. Antibodies found in genicity. Since 1980, most viruses iso- low percentage of humans, rodents, lated from Australia appear to be less marsupials, and horses but not in mon- nephrotropic, and no longer cause high keys, bats, lizards, cows, and sheep. mortality. Causey OR et al (1963) Am J Trop Med Hyg Ignjatovic J and McWaters PG (1991) J Gen 12 , 777 Virol 72 , 2915 Rümenapf T et al (1995) Virology 208 , 621 Klieve AV and Cumming RB (1988) Avian Pathol 17 , 829 Australia antigen An antigen first found in the blood of an Australian aborigi- Australian X-disease virus An histori- nal and later identified as hepatitis B cal name for Murray Valley encephalitis surface antigen. virus . autoantibodies 34 autoantibodies Antibodies made against is Turkey astrovirus . All members of the host proteins which are not foreign. genus infect avian species. There are Found in patients with infectious mono- three recognized species in the genus. nucleosis. See Paul–Bunnell antibody . Chicken astrovirus 1 which includes the strains avian nephritis virus 1 and autoimmunity Production of autoanti- 2, Duck astrovirus, and Turkey astrovi- bodies, e.g. Hashimoto’s disease which rus, which includes the strains turkey is caused by production of antibodies astrovirus 1 and 2. Infection with an to thyroglobulin. avastrovirus causes intestinal disease, but also hepatitis and damage to the autointerference See interference . kidney and thymus. Koci MD and Schultz-Cherry S (2002) Avian autonomous replicating sequence (ARS) Pathol 31 , 213 A specific DNA sequence responsible for initiation of DNA replication, first Aviadenovirus A genus in the family seen in yeast, then in other eukaryotes. Adenoviridae comprised of the viruses ARS elements support independent isolated from birds of many different replication of a plasmid in host yeast species in many countries. They share a cells. Can be used for cloning large common antigen which does not cross- DNA molecules. react with the genus Mastadenovirus Struhl K et al (1974) Proc Natl Acad Sci common antigen. The 12 vertex cap- 76 , 1035 someres each have two filaments of different lengths. Genome mol. autoradiography A method for detecting wt. 30 1 06 . G C content: 54–55%. the location of a radioisotope in tissue, Gallus-adeno-like virus agglutinates rat cell, or molecule. The sample is placed erythrocytes and is the only species to in contact with an X-ray film, whereby do so. Type species fowl adenovirus 1 the emission of β particles from the (chicken embryo lethal orphan (CELO) sample activates the silver halide grains virus). There are at least 12 serologi- in the emulsion and allows them to be cally distinct fowl adenoviruses. Three reduced to metallic silver when the film serologically distinct types have been is developed. Widely used to detect isolated from turkeys and three from virus-induced nucleic acids or proteins geese. Other strains have been isolated after radioisotopic labeling followed by from ducks, pheasants, guinea fowl, separation on a polyacrylamide gel. and goshawks.

Auzduk disease virus A tentative species McFerran JB and Connor TJ (1977) Avian Dis 21 , 585 in the genus Parapoxvirus . Synonym : camel contagious ecthyma virus. avian acute leukemia virus An acutely transforming strain of avian type C Av-3 cells (CCL 21) A line of normal retrovirus. human amnion cells. Avian adeno-associated virus (AAAV) AV 9310135 virus A strain of Whitewater A species in the genus Dependovirus . Arroyo virus in the genus Arenavirus . Antigenically distinct from primate adeno-associated viruses. A defective Avalon virus (AVAV) A strain of Sakhalin parvovirus that accompanies most virus in the genus Nairovirus belonging avian adenovirus infections in the field, to the Sakhalin virus antigenic group. and contaminates most laboratory Isolated from mosquitoes on Great strains. Some herpesviruses can also Island, Witless Bay, Newfoundland, provide helper activity to AAAV. Virus Canada. Not reported to cause disease is easily isolated from feces or intes- in humans. tinal epithelial cells. Virion 18–20 nm in diameter with a density (CsCl) of Avastrovirus A genus in the family 1.43 g/ml. No evidence of pathogenic- Astroviridae , the type species of which ity has been found, but the presence 35 avian erythroblastosis virus

of AAAV appears to reduce the patho- ataxia followed by tremors of the head, genicity of avian adenoviruses for somnolence, and often death. Young 1- to 5-day-old chicks. Humans who ducks, turkeys, and pigeons can be work closely with chickens frequently infected experimentally but not mam- develop antibodies to AAAV, which mals. Has caused outbreaks of disease presumably multiplies with the help of in pigeons in Turkey. Virus is present a human adenovirus. in the excreta and in the eggs which Synonym : quail parvovirus. may contain antibodies protective for Dawson GJ et al (1982) Nature 298 , 580 the chick. Strains vary in virulence. Siegl G et al (1983) Intervirology 23 , 61 Inoculation of eggs may kill the embryo. Replicates in chick embryo cell cultures. avian adenovirus See Aviadenovirus . Synonym : epidemic tremor virus. Calnek BW et al (1991) In Diseases of Poultry , avian arthritis virus A possible species Ninth edition, edited by BW Calnek et al . in the family Poxviridae . Unrelated to Ames: Iowa State University Press, p. 520 fowlpox virus. Ether- and chloroform- Marvil P et al (1999) J Gen Virol 80 , 653 resistant. Replicates in yolk sac of embryonated eggs. Causes arthritis on avian enteric cytopathogenic virus An old injection into the footpads of chicks. term used for fowl adenovirus when it Taylor DL et al (1966) Avian Dis 10 , 462 was thought to be a picornavirus.

Avian carcinoma virus Mill Hill virus avian entero-like virus 2 to 4 (AELV-2 2 (ACMHV-2) A species in the genus to 4) Unassigned viruses in the family Alpharetrovirus . Isolated from a ‘ globular Picornaviridae . tumor in the ovarian region ’ in a white leghorn chicken. Causes endotheliomas, avian enteroviruses Include avian enceph- leukemia, kidney carcinomas, hepa- alomyelitis virus, duck hepatitis virus tocarcinomas in chickens, and hemor- 1, duck hepatitis virus 3, and avian rhagic disease in embryos. Transforms nephritis virus, as well as a number of immature macrophages or fibroblasts in strains isolated from normal birds. vitro . It is defective, lacking full function of the gag, env , and pol genes necessary avian erythroblastosis virus A mix- for replication. Contains two oncogenes: ture of viruses belonging to the genus mil and myc . Alpharetrovirus . Consists of a transform- Synonyms : MH 2 virus; Mill Hill virus ing virus and a leukemia virus which 2. acts as helper for the defective trans- Alexander RW et al (1979) J Natl Cancer Inst forming virus. The helper belongs to 62 , 359 subgroup B of the chicken leukosis sar- Begg AM (1927) Lancet II , 912 coma viruses. The transforming virus Hu SSF and Vogt PK (1979) Virology 92 , 278 is very closely related to the transforming component in avian myeloblastosis. avian diarrhea virus Synonym for infec- Contains two transduced oncogenes: tious enteritis virus. See also blue v-erbA and v-erbB . Expression of comb virus . v-erbB , which is a homolog of the cellular epidermal growth factor receptor, is avian encephalomyelitis-like virus necessary and sufficient for induction (AEV) A tentative species in the genus of erythroblastosis. On i.v. injection Hepatovirus . The complete genome RNA into chickens the virus causes rapid sequence (7032 nt) shows 39% amino erythroblastosis and anemia but on acid identity with hepatitis A virus. i.m. injection sarcomas are produced. Virions 24–32 nm in diameter without Transforms fibroblasts and bone mar- envelope. Density (CsCl): 1.31–1.33 g/ row cells in vitro. Non-producer cells ml. There are at least 15 serotypes. A can be obtained and the defective widespread and silent infection of adult genome rescued by superinfection domestic fowl, but causes severe dis- with a chicken leukosis sarcoma virus. ease in 2- to 3-week-old chicks. There is The anemia and the occasional case of avian erythroblastosis virus 36

lymphatic leukemia produced are coat proteins. The helper thus deter- probably due to the helper virus. mines the subgroup and host-range of Synonym : erythroblastosis virus. the progeny virus. (2) Strains which in Graf T et al (1983) Cell 34 , 7 vivo transform hematopoietic cells and Hayman MJ et al (1979) Virology 92 , 31 induce leukemia on injection into birds Ishizaki R and Shimizu T (1970) Cancer Res and can act as helpers for defective sar- 30 , 2827 coma viruses. (3) Endogenous viruses found in virtually all normal chickens avian H10 influenza A virus An avian and other gallinaceous birds which are subtype, H10N4, of Influenza A virus transmitted vertically, in Mendelian which causes severe respiratory dis- fashion, to their progeny as they exist ease in mink (Mustela vison) both under as proviral DNA sequences integrated field conditions and experimentally. into somatic and germ line cells. The Originally found to be the cause of a sites of these integrated viral genes are severe epidemic of respiratory disease termed endogenous virus (ev ) loci, and in farmed mink in Sweden in 1984. on average each chicken carries five such loci. Related ev loci are found in Englund L and Segerstad CH (1998) Arch red jungle fowl, ring-necked pheasants, Virol 143 , 653 partridge and grouse, but not in guinea fowl, quail, peafowl, ruffed pheasants, avian hepatitis E virus A tentative spe- gallopheasants, or turkeys. If the ev Hepevirus cies in the genus . Associated locus contains a complete endogenous with hepatosplenomegaly in chickens. viral genome, virus may be produced Big liver and spleen disease virus See . spontaneously or after induction, e.g. by mutagens, and belongs to subgroup avian infectious bronchitis virus (IBV) E. Expression from ev loci containing See Infectious bronchitis virus . partial genomic information may be observed by complementation of defec- avian infectious laryngotracheitis virus tive avian leukosis sarcoma viruses by Synonym for Gallid herpesvirus 1 . providing envelope antigens, termed chick helper factor (chf), or by the pres- avian influenza A virus See Influenza A ence of group-specific antigen (gs ). virus . Antisera raised against the gs antigen, p27, are used in the complement avian leukemia virus See Avian leukosis fixation test for avian leukosis virus virus . (COFAL test) to screen flocks for presence of avian leukosis viruses. Avian avian leukosis sarcoma virus See Avian leukosis viruses are transmitted natu- leukosis virus . rally in three ways: (1) horizontally, by direct contact from bird to bird; (2) ver- Avian leukosis virus (ALV) A species in tically when exogenous virus is trans- the genus Alpharetrovirus. There are 10 mitted via the egg; and (3) genetically subspecies or subgroups designated in the germ line when endogenous A–J. Subgroups A–E and J are chicken viruses pass from parents to offspring. viruses; subgroups F–I are isolated from Synonym : chicken leukosis sarcoma other avian species. They are defined virus. by envelope properties such as host- Crittenden LB (1991) Crit Rev Poult Biol 3 , 73 range, sensitivity to viral interference, Payne LN (1992) In The Retroviridae, vol. 1, and antigenicity. There are: (1) strains edited by JA Levy. New York: Plenum Press, producing solid tumors, sarcomas, p. 299 on injection into birds. These sarcoma Payne LN and Purchase HG (1991) In viruses, such as Bryan high titer strain Diseases of Poultry , Ninth edition, edited of Rous sarcoma virus, can transform by BW Calnek et al . Ames: Iowa State University Press, p. 386 cells in culture but most of the strains are defective. They contain the src gene, avian leukosis virus – HPRS103 (ALV-J) but require a leukemia virus to provide A tentative species in the genus the genetic information for the viral Alpharetrovirus . 37 avian nephritis virus 1 to 3 (ANV-1 to 3) avian leukosis virus – RSA (ALV- transforming virus is probably defec- A) A tentative species in the genus tive. Non-producer transformed cells Alpharetrovirus . can be obtained and from these cells leukemogenic virus can be rescued by avian lymphoid leukosis virus See Avian superinfection with chicken leukosis leukosis virus . sarcoma virus of subgroup B, C, or D. MAV of subgroup A does not rescue avian lymphomatosis virus See Avian transforming AMV. leukosis virus . Moscovici C (1975) Curr Top Microbiol Immunol 71 , 79 avian monocytosis virus Synonym for Payne LN (1992) In The Retroviridae, vol. 1, infectious enteritis virus. See also blue edited by JA Levy. New York: Plenum Press, comb virus . p. 299

Avian myeloblastosis virus (AMV) A Avian myelocytomatosis virus 29 replication-defective species in the (AMCV29) A replication-defective spe- genus Alpharetrovirus . The original cies in the genus Alpharetrovirus . Related strain, reported by Ellerman and Bang viruses are CMII and OK10. All carry the in 1908, has been lost. The most widely myc oncogene. It causes myelocytoma- used strain is BAI (Bureau of Animal tosis, renal and liver tumors and occa- Industry) strain A which originated sionally erythroblastosis in chickens. from a 1941 isolate from two birds with Causes cell transformation in vitro and neurolymphomatosis. After several can produce foci of such cells in chick passages from nerve and bone mar- embryo monolayers. The transforming row tissue a virus with leukemogenic virus is defective and carries the trans- activity was obtained. BAI strain A duced oncogene vmyc . Non-producer is a complex of several viruses. Two cells can be obtained from which infec- viruses, AMV-1 and AMV-2, have been tive virus is produced on superinfection isolated from it. They belong, based on with chicken leukosis sarcoma viruses envelope properties, to chicken leu- such as Rous-associated virus or ring- kosis sarcoma virus subgroups A and necked pheasant virus. B, respectively, and on injection into Bister K et al (1977) Virology 82 , 431 day-old chicks cause osteopetrosis and Hu SSF et al (1979) Proc Natl Acad Sci 76 , 1265 kidney tumors, but not leukemia. Payne LN (1992) In The Retroviridae, vol. 1, They have been renamed ‘ myelo- edited by JA Levy. New York: Plenum Press, blastosis-associated virus ’ (MAV-1 p. 299 and MAV-2). It is suggested that the original mixture of viruses be called avian nephritis virus 1 to 3 (ANV-1 to ‘ standard ’ AMV, which contains 3) Unassigned viruses in the family cell-transforming and leukemogenic Picornaviridae . Isolated in chick kid- viruses, as well as the non-transforming ney culture of the rectal contents of MAV-1 and 2. There are stocks of AMV normal 1-week-old chicks in Japan in free of subgroup A virus which are 1976. Virions 28 nm in diameter, ether- referred to as subgroup B AMV. No resistant, containing RNA were found, subgroup A AMV with leukemogenic distinct antigenically and pathogeni- activity is reported. ‘ Standard ’ AMV cally from avian encephalomyelitis causes myeloblastosis, osteopetrosis, virus. Appears to be a widespread lymphoid leukosis, and nephroblas- infection of commercial chicken and toma in chickens. It seems likely that turkey flocks, transmission occurring the different conditions are caused by by direct and indirect contact. Main individual viruses or combinations of target organ for replication is the kid- viruses in the ‘ standard ’ AMV. AMV ney, but there are few clinical signs and can cause transformation of yolk-sac the disease significance is uncertain. or bone-marrow cells in vitro . Contains Imada T (1993) In Virus Infections of the oncogene v-myb incorporated at the Vertebrates, vol. 4, edited by JB McFerran 5 leader sequence of the gag gene, and and MS McNulty. Amsterdam: Elsevier, encodes a DNA-binding protein. The p. 479 avian nephritis virus 1 to 3 (ANV-1 to 3) 38

Shirai J et al (1992) Avian Dis 36 , 369 avian papillomavirus Synonym for fring- Yamaguchi S et al (1979) Avian Dis 23 , 571 illa papillomavirus. avian nephrosis virus Synonym for avian parainfluenza virus 1 (APMV-1) Infectious bursal disease virus . Synonym for Newcastle disease virus .

Avian orthoreovirus (ARV) Species in the avian paramyxovirus 1 (APMV-1) genus Orthoreovirus which do not infect Synonym for Newcastle disease virus . mammalian species and share a common group antigen, distinct from the Avian paramyxovirus 2 (Yucaipa) mammalian reovirus group antigen. (APMV2) A species in the genus The genome consists of 10 segments of Rubulavirus. Isolated in southern double-stranded RNA. Unlike mamma- California, USA from tracheal exudate lian orthoreoviruses, infectivity is not of chickens with severe laryngotrachei- increased by heating in the presence of tis in which the main cause of disease MgCl2 . There are at least 11 distinct sero- was infectious laryngotracheitis virus. types. The viruses all induce syncytia in Yucaipa virus causes only mild disease. cell culture and cause a variety of symp- Similar to Newcastle disease virus but toms in chickens and turkeys following an antigenically distinct species. a viremia which results in widespread Synonym : Yucaipa virus. virus dissemination. The principal site Dinter Z et al (1964) Virology 22 , 297 of replication is the gastrointestinal tract, Ozdemir I et al (1990) Avian Pathol 19 , 395 and the most serious disease symptoms are arthritis (tendosynovitis) caused by Avian paramyxovirus 3 (APMV-3) A spe- infection of the tendons and associated cies in the genus Rubulavirus . Isolated tissues of the hock joint. Other symp- from turkeys in Wisconsin, USA and toms, which vary depending upon the in Ontario, Canada. Isolates have also strain of virus, include hepatitis, gas- been made from psittacines. Similar to troenteritis, myocarditis, and paling, Newcastle disease virus but antigeni- and weight loss due to maladsorption. cally distinct from it, and from Yucaipa The virus is widespread in commercial virus and Bangor virus (Avian para- poultry flocks, but some control can be myxovirus 5). achieved by good management proce- Anderson C et al (1987) Avian Pathol 16 , 691 dures, especially thorough cleansing and disinfection of houses between suc- Avian paramyxovirus 4 (APMV-4) A spe- cessive crops of broilers. To date, the cies in the genus Rubulavirus . Isolated development of an attenuated vaccine from domestic poultry in Hong Kong has not produced reliable results. and USA and from wild ducks in Japan. Shortridge KF and Alexander DJ (1978) Res avian orthoreovirus 176 (ARV-176) A strain Vet Sci 25 , 128 of avian orthoreovirus in the genus Orthoreovirus. Isolated from chickens. Avian paramyxovirus 5 (Kunitachi) (APMV-5) A species in the genus avian orthoreovirus 1733 (ARV-1733) Rubulavirus. Isolated from the lung A strain of avian orthoreovirus in the of a budgerigar, Melopsittacus undula- genus Orthoreovirus. Isolated from tus , which died during an epizootic in chickens. Kunitachi, Tokyo, Japan. Replicates in the amniotic cavity of embryonated avian orthoreovirus S1133 (ARV-S1133) hen’s eggs and in chick-embryo cell A strain of avian orthoreovirus in the cultures. Agglutinates chicken, goose, genus Orthoreovirus. Isolated from duck, guinea pig, and human O eryth- chickens. rocytes at 4ºC and 25ºC, but activity not stable in amniotic fluid. Causes a fatal avian orthoreovirus SK138a (ARV-138) disease on injection into budgerigars. A strain in the genus Orthoreovirus . Synonym : Kunitachi virus. Isolated from chickens. Nerome K et al (1978) J Gen Virol 38 , 293 39 Avian sarcoma virus 17

Avian paramyxovirus 6 (APMV-6) A avian reticuloendotheliosis virus See species in the genus Rubulavirus . Reticuloendotheliosis virus . Isolated from ducks in Hong Kong and Japan. avian retroviruses Most avian retroviruses are members of the genus Nerome K et al (1984) J Virol 50, 649 Alpharetrovirus , but the avian Shortridge KF et al (1980) J Gen Virol 49 , 255 (Reticuloendotheliosis) virus group is in the genus Gammaretrovirus . Avian paramyxovirus 7 (APMV-7) A species in the genus Rubulavirus . Isolated avian rotaviruses Group A, D, and F from doves in Tennessee, USA. Other species in the genus Rotavirus contain members of the species identified in strains that infect birds. The group A Japan and UK. viruses are closely related to mammalian group A strains, but two other Alexander DJ et al (1991) Arch Virol 116 , 267 groups do not cross-react with mammalian rotaviruses; they are: group Avian paramyxovirus 8 (APMV-8) A D (prototype, chicken 132 virus) and species in the genus Rubulavirus , serogroup F (viruses isolated from pheas- logically distinct from other avian par- ants and turkeys in the USA). amyxoviruses. The prototype virus was isolated from a hunter-killed Canadian avian sarcoma and leukosis viruses goose, Branta canadensis , in Delaware, (ASLV) Members of the genus USA in 1976. Alpharetrovirus of the Retroviridae family. Although information on these Alexander DJ et al (1983) Arch Virol 78 , 29 viruses has largely been derived from domestic chickens, a recent sequencing Avian paramyxovirus 9 (APMV-9) A study reported new viruses in 19 spe- species in the genus Rubulavirus , sero- cies of birds in the order Galliformes (e.g. logically distinct from other avian par- chicken, grouse, partridge, ptarmigan, amyxoviruses. The prototype virus was and quail) representing 3 families and isolated from a sick domestic white 14 genera. On this basis it was proposed Pekin duck on Long Island, New York, that ASLVs should be classified into two USA in 1978. subgenera of Alpharetrovirus as follows: Alexander DJ et al (1983) Arch Virol 78 , 29 Subgenus 1: Gallus ASLV, Perdix ASLV, and Phasianidae/Odontophoridae ASLV Avian paramyxoviruses (APMV 1–9) Subgenus 2: Colinus ASLV, Tetraonine Species in the genus Rubulavirus . ASLV, and Bonasa/Phasianus ASLV Includes Newcastle disease virus (avian paramyxovirus 1) and a number of See Avian leukosis virus . viruses isolated from birds which are Dimcheff DE et al (2000) J Virol 74 , 3984 antigenically distinguishable from it and from each other. They include Avian sarcoma virus CT10 (ASV-CT10) A Yucaipa virus, Bangor virus, turkey species in the genus Alpharetrovirus . A parainfluenza virus, budgerigar parain- defective virus which carries the trans- fluenza virus, and Kunitachi virus. duced oncogene v-crk , the product of which binds to several cellular proteins in vitro avian plague virus Synonym for highly and causes cell transformation . pathogenic avian influenza A virus. Infected birds develop fibrosarcomas. Mochizuki N et al (2000) J Biol Chem 275 , avian pneumoencephalitis virus Syno- 12667 nym for Newcastle disease virus . Avian sarcoma virus 17 A replication- defective virus which causes fibrosarco- avian pneumovirus Synonym for Turkey mas in young chickens and transforms rhinotracheitis virus . chick embryo fibroblast cells in culture. avian reovirus (AVREO 1–9) See Avian Contains the v-jun oncogene. orthoreovirus . Cavalieri F et al (1985) Virology 143 , 680 avian type C oncovirus group 40 avian type C oncovirus group See the mucous membranes of the upper avian sarcoma and leukosis viruses respiratory tract, mouth, and esophagus. (ASLV) . The species currently listed under this genus are Fowlpox, Turkeypox, Canarypox, avianized virus Virus adapted to growth Pigeonpox, Quailpox, Sparrowpox, in birds. Starlingpox, Juncopox, Mynahpox , and Psittacinepox viruses . Other probable members of this genus are crowpox, Avibirnavirus A genus of the family peacockpox, and penguinpox viruses. Birnaviridae including only viruses Fowlpox virus (type species of the genus which infect birds. Only one species, Avipoxvirus ) has been investigated most Infectious bursal disease virus , has been extensively. recognized and this is the type species. Synonym : fowlpox subgroup viruses. There are two recognized serotypes: serotype 1 strains are pathogenic in Schnitzlein WM et al (1988) Virus Res 10 , 65 Tripathy DN (1991) In Diseases of Poultry , chickens, causing immunosuppres- Ninth edition, edited by BW Calnek et al . sive disease by destruction of cells in Ames: Iowa State University Press, p. 583 the bursa of Fabricius; and serotype 2 strains which are nonpathogenic. avirulent strain A strain of virus which does not cause disease. See attenuated avidity Intensity of binding of, e.g., an virus strains . antibody molecule to the antigen which induced its formation. Avulavirus A genus of the family Paramyxoviridae consisting of avian spe- Avihepadnavirus A genus of the family cies. The type species is Newcastle disease Hepadnaviridae containing only viruses virus . All species have both hemagglu- which infect birds. The type species is tinin and neuraminidase activities, but Duck hepatitis B virus . The only other do not possess an SH gene. confirmed species in the genus is Heron hepatitis B virus, but Ross’s goose axenic Not contaminated with any for- hepatitis B virus may also be a mem- eign microbes; a pure preparation or ber. There are also five new candidate culture. avihepadnaviruses detected by DNA cloning of avian tissues from the Chloe axonal spread Movement of virus by wigeon, mandarin duck, puna teal, transport through nerve axons seen Orinoco sheldgoose, and ashy-headed with, e.g. rabies virus. sheldgoose. 5-azacytidine A base analog that has Guo H et al (2005) J Virol 79 , 2729 proved to be a mutagen for RNA viruses.

Avipoxvirus A genus of the subfamily 3-azido-3-deoxythymidine See AZT . Chordopoxvirinae, consisting of viruses of birds. Ether-resistant. Species are anti- azidothymidine See AZT . genically related. Infected cells develop type A inclusion bodies which are rich AZT (azidothymidine) A synthetic in lipid. Hemagglutinin is not formed. pyrimidine dideoxynucleoside ana- Mechanical transmission by arthro- log that inhibits replication of retro- pods is common. Type species Fowlpox viruses, including HIV by interfering virus . DNA 260 kb. Avipoxviruses infect with DNA synthesis mediated by the domestic, wild, and pet birds. Infections viral reverse transcriptase. AZT-5 - are characterized by the development of triphosphate interacts preferentially with proliferative lesions ranging from small reverse transcriptase rather than cell nodules to spherical tumor- or wart-like DNA polymerase. The use of this drug masses on the skin of unfeathered areas. by oral administration in vivo is recom- In some cases, proliferative lesions or mended for reducing viral load, restor- diphtheric membranes may develop on ing T-cell function, and prolonging life 41 AZT (azidothymidine)

in persons with clinical AIDS. It is also benefits of AZT therapy need to be bal- recommended for use in HIV-infected anced against these opposing factors. pregnant women in order to reduce Synonyms : 3 -azido-2 ,3 -dideoxythy- the risk of transmission of HIV to the midine; AZT; azidothymidine; retrovir; baby. The drug is toxic to dividing cells zidovudine. such as bone marrow cells, and drug- de Clercq E (1993) Adv Virus Res 42 , 1 resistant HIV mutants arise readily; the de Clercq E (1995) Rev Med Virol 5 , 149 B

B cells Lymphocytes derived from hemo- females, 80% of cases have associated poietic stem cells in the bone marrow arthropathy. B19 virus infects eryth- (or the bursa of Fabricius in birds) rocyte precursors, and if the infected which synthesize immunoglobulin individual has pre-existing hemolytic and play a major role in the adaptive anemia a transient aplastic crisis occurs immune response. which usually lasts for 7 days, before a humoral immune response is mounted. B14–150 cells (CCL 14.1) Heteroploid More serious, chronic anemia occurs in peritoneal cells from the Chinese ham- immunocompromised individuals. In ster, Cricetulus griseus . Initiated from pregnant women, intrauterine B19 infec- the original B14 cell line by selection of tion may cause fetal loss and hydrops cells resistant to bromo-deoxyuridine fetalis . in the culture medium. This was later Anderson LJ and Young NS (1997) Monogr replaced by idoxuridine to which the Virol 20 , 153pp cells are now resistant. Berns KI (1990) Microbiol Rev 54 , 316 Erdman DD et al (1996) J Gen Virol 77 , 2767 B95–8 cells An Epstein–Barr virus transformed marmoset B-lymphocyte cell 734 B virus A possible species in the line. Used for primary measles virus genus Betaretrovirus. Present in MCF-7 isolation. cells, a line derived from a pleural effu- Kobune F et al (1990) J Virol 64 , 700 sion from a patient with disseminated mammary adenocarcinoma. No antigenic relationship with type C retro- B-virus Synonym for Cercopithecine her- viruses, but some cross-reaction with pesvirus 1 (herpesvirus B). mouse mammary tumor virus. B19 virus (B19V) The type species of the B814 virus The first strain of Human coro- genus Erythrovirus, the cause of ery- navirus isolated in organ culture from a thema infectiosum (fifth disease) in chil- patient with a common cold. dren and of aplastic crisis in children and adults with chronic hemolytic ane- Tyrrell DAJ and Bynoe ML (1965) BMJ 1 , mia. The linear single-stranded DNA 1467 genome of B19 virus is 5.5 kb in length and has long (about 300 bases) inverted B-virus of monkeys Synonym for Cercop- terminal repeats at the 3 and 5 ends. ithecine herpesvirus 1 . Both plus and minus strands are packaged with equal efficiency into B19 viri- Babahoya virus (BABV) A sero- ons. Human B19 infection is common type of Patois virus in the genus between 4 and 12 years but is usually Orthobunyavirus belonging to the asymptomatic; serosurveys have shown Patois serogroup. Isolated from mos- that about 80% of the adult popula- quitoes, Culex (Melanoconion) ocossa , in tion have experienced B19 infection. Ecuador. Not known to cause disease Erythema infectiosum is a common in humans. manifestation of childhood infection Calisher C et al (1983) Am J Trop Med Hyg and starts with intense erythema of the 32 , 877 cheeks (slapped-cheek disease), then moves to the trunk and limbs, lasting Babanki virus (BBKV) A serotype of about 2 days. In a few cases ( 10%) Sindbis virus in the genus Alphavirus . arthropathy with joint symptoms is Isolated from the mosquito, Mansonia seen, but when infection occurs in adult africana , in Cameroon. 43 bacteriocins baboon endogenous virus (BaEV) A Antigenically distinct from simian agent probable species in the genus Gammaret- 12 so named polyomavirus papionis-2. rovirus. A vertically transmitted endo- Gardner SD et al (1989) Arch Virol 105 , 223 genous virus which is present in multiple copies in many Old World primate spe- baboon T-cell leukemia virus (BTLV) cies. Remarkable sequence conservation See Primate T-lymphotropic viruses . is maintained between different species of primate. Isolated from a baboon, baboon type C virus See baboon endo- Papio cynocephalus , by co-cultivation genous virus . of the cells with various mammalian cells. Replication was most efficient in baby hamster kidney cells See BHK21 fetal canine thymus cells. Virus can be cells . isolated directly from placental extract. Bacajaí virus A virus in the genus The virus designated M7 has reverse Orbivirus isolated from phlebotom- transcriptase and group-specific protein ine sand flies in the Amazon region immunologically distinct from other of Brazil. Antigenically related to the C-type viruses. It also contains an env gene Changuinola virus serogroup. homologous to that of a Deltaretrovirus . However, there is strong sequence rela- BAC (Bacterial Artificial Chromosome) tionship to RD114 virus, suggesting A DNA construct based on a fertility that the cat endogenous virus may have plasmid (F-plasmid) used for trans- evolved from this baboon endogenous forming and cloning in bacteria. The virus. It has been reported that sera from usual insert size is 150 kb (range from human patients with schizophrenia 100 to 300 kb). displayed a significantly increased incidence of antibodies to BaEV. bacilliform Description of the shape of Synonym : baboon C-type virus. certain virus particles which are cylin- Hu S et al (1977) J Virol 23 , 345 drical with two rounded ends. Lillehoj EP et al (2000) J Neurovirol 6 , 492 Schnitzer TJ (1979) J Gen Virol 42 , 9 bacteriocinogen A plasmid DNA present in van der Kuyl AC et al (1995) J Virol 69 , 5917 certain strains of bacteria, which specifies production of a bacteriocin. Normally baboon herpesvirus Synonym for Cercop- the bacteriocinogen is repressed, and the ithecine herpesvirus 12 . cell carrying it does not produce bacteriocin. The circumstances in which dere- baboon lymphotropic herpesvirus Syno- pression occurs are complex and not nym for Cercopithecine herpesvirus 12 . completely understood.

Baboon orthoreovirus (BRV) A species Hardy KG (1975) Bacteriol Rev 39 , 464 in the genus Orthoreovirus. The only bacteriocins Protein substances of vary- member of subgroup IV. ing complexity released by some types of bacteria which kill bacteria of certain baboon polyomavirus 1 (Ppy-1) A virus strains within the same species. The isolated from baboon cell cultures that producing strain is generally immune was originally called simian agent 12. to the effects of its own bacteriocins. Replicates productively in African When purified, bacteriocins seem to green monkey kidney cells, in contrast fall into two classes: some are simple to human polyomaviruses. Now a spe- proteins or proteins associated with cies called Simian virus 12 in the genus cell wall components; others resem- Polyomavirus. ble bacteriophages or fragments of Synonym : polyomavirus papionis 1. them. Bacteriocin formation is due to Cunningham TP and Pipas JM (1985) J Virol a bacteriocinogen in the cell, which is 54 , 483 normally repressed and behaves like a defective prophage. Bacteriocins Baboon polyomavirus 2 (PPyV-2) A spe- adsorb to specific receptors on the cell cies in the genus Polyomavirus . Isolated wall, and bacterial mutants which lack from baboon kidney cell cultures. these receptors may arise and will be bacteriocins 44

resistant. The potency of bacteriocins is Bahig virus (BAHV) A serotype of Tete exceedingly high, and in several cases virus in the genus Orthobunyavirus , it has been shown that the lethal action belonging to the Tete antigenic virus is mediated without penetration of the group. Isolated from birds in Egypt cell. Some bacteriocins appear to be and Italy. Serological surveys suggest enzymes which cause cell lysis. presence in Cyprus and Israel. Not Kraus D and Peschel A (2006) Curr Top reported to cause disease in humans. Microbiol Immunol 306 , 231 Bakau virus (BAKV) A species in the bacteriophage A virus which replicates genus Orthobunyavirus . With Ketapang, inside a bacterium. Nola, Tanjong Rabok, and Telok Forest viruses forms the Bakau serogroup. baculovirus expression vector A gene Isolated from mosquitoes in Malaysia expression system which utilizes a and Pakistan. Not reported to cause strong promoter found in baculoviruses disease in humans. to obtain high-level expression of foreign genes. The baculovirus vector uses Bakel virus (BAKV) A strain of Qalyub the highly expressed and regulated virus in the genus Nairovirus , isolated polyhedrin promoter modified for the from ticks. Not reported to cause dis- insertion of foreign genes. The baculovi- ease in humans. rus used is usually Autographa californica nuclear polyhedrosis virus (NPV). One Baku virus (BAKUV) A serotype of of the major advantages of this inver- Chenuda virus in the genus Orbivirus . tebrate virus expression vector over Isolated from a tick, Ornithodoros cap- bacterial, yeast, and mammalian expres- ensis. Antibodies found in nestlings of sion systems is the abundant expression a gull, Larus argentatus , on Glinyanyi of proteins coded by the inserted gene. Island in the Caspian Sea. Not reported In addition, recombinant proteins pro- to cause disease in humans. duced in insect cells with baculovirus vectors are biologically active and, for balano-posthitis virus of sheep A pos- the most part, appear to undergo post- sible species in the genus Parapoxvirus translational processing to produce causing venereal infection in sheep in gene products with similar properties America. It can cause ulcerative derma- to the authentic proteins. titis as well as balanitis and ulcerative vulvitis. Similar disease pictures are Bacuri virus A probable species in the reported from Australia, South Africa, genus Orbivirus , isolated from phlebot- and the UK. They may be caused by omine sand flies in the Amazon region a variant of Orf virus but the relation- of Brazil. Antigenically related to the ships of these viruses to other parapox- Changuinola virus serogroup. viruses require study. badger herpesvirus See Mustelid herpes- Synonyms: ulcerative dermatosis of virus 1 . sheep; pizzle rot; foul sheath. Bagaza virus (BAGV) A species in the Trueblood MS (1966) Cornell Vet 56 , 521 genus Flavivirus belonging to the Ntaya virus subgroup. Isolated in BALB/3T3 cells (CCL 163) Heteroploid suckling mice from a pool of Culex fibroblast cell line developed as 3T3 mosquitoes collected from humans cells, but from disaggregated 14- to at Bagaza, Central African Republic. 17-day-old embryos of inbred BALB/c Has also been found in Cameroon and mice. Exhibit low saturation density, Senegal. Not reported to cause disease are extremely sensitive to contact inhi- in humans. bition, grow at high dilution, and are highly susceptible to transformation Bahia Grande virus (BGV) An unassigned by the oncogenic DNA virus SV40 and species in the family Rhabdoviridae iso- mouse sarcoma virus. Do not form lated from Aedes sollicitans . With Muir tumors on injection into weanling irra- Springs and Reed Ranch forms the diated BALB/c mice whereas 3T12-B Bahia Grande serogroup. cells do. Also known as 3T3-B cells. 45 Bangor virus

BALB/3T12-3 cells (CCL 164) This is one of feces from these calves and infants of several cell lines developed from a 32 years later showed the presence of pool of disaggregated 14- to 17-day-old rotavirus-like particles. BALB/c mouse embryos. The cells are Hodes HL (1977) Am J Dis Child 131 , 729 extremely insensitive to contact inhibition, exhibit a high saturation density, Bamble disease are tumorigenic, and are susceptible to This disease was first transformation in tissue culture by the described in 1872 in Norway, and takes oncogenic DNA virus SV40. The cell its name from the village in which it line has been used in studies relating to was prevalent. There is an incubation in vitro properties associated with tum- period of 2–4 days followed by sudden origenicity and contact inhibition. onset, with ‘ stitch-like ’ pain in chest, epigastrium, abdomen, and more rarely, the limb muscles, accompanied BALB/c (Mo) mice A strain of mice by fever, headache, coughing, and hic- derived from a pre-implantation cough. May be caused by coxsackie embryo infected with Moloney leuke- viruses types B1–6, A4, 6, 9, and 10, or mia virus. They have the viral DNA echovirus types 4, 6, and 9. The first N transmitted as a single Mendelian American outbreak was described in gene. Heterozygous animals have one 1888 by Dabney, whose name was offi- copy of the viral DNA in each diploid cially bestowed on the disease in the cell, homozygous individuals have USA, in 1923. The Bornholm outbreak two. Infective virus is produced in their was not described until 1932. tissues and 90% of the mice develop Synonyms: Bornholm disease; Dabney’s thymus-derived lymphomas before grippe or grip; devil’s clutch; devil’s they are 10 months old. grippe or grip; epidemic myalgia; pleu- Jaenisch R (1976) Proc Natl Acad Sci 73 , 1260 rodynia; Taarbaek disease. Miles J (1971) Hist Med 3 , 28 Balbina virus A probable species in the genus Orbivirus , isolated from phlebot- banded krait herpesvirus Synonym for omine sand flies in the Amazon region elapid herpesvirus 1. of Brazil. Antigenically related to the Changuinola virus group. Bandia virus (BDAV) A serotype of Qalyub virus in the genus Nairovirus . bald eagle herpesvirus Synonym for With Bakel, Omo, and Qalyub viruses acciptrid herpesvirus 1. forms the Qalyub serogroup. Isolated from mice and ticks of Ornithodoros Balkan nephropathy virus Virus parti- sp from the Bandia Forest, Senegal. Not cles morphologically resembling coro- reported to cause disease in humans. naviruses seen in sections of kidney tissue from human cases of a slowly bandicoot papillomatosis carcinomato- progressive kidney disease. This dis- sis virus 1 (BPCV1) A virus isolated ease occurs only in the Balkans, mainly from lesions on the western barred Bulgaria, and is rare in Muslims. Virus bandicoot (Perameles bougainville ), an antigen in tissue sections reacts with endangered Australian marsupial. The patients’ serum, but does not react with genome appears to have sequences in pig or bird coronavirus antiserum. common with both papillomaviruses Apostolov K et al (1975) Lancet ii , 1271 and polyomaviruses. Woolford L et al (2007) J Virol 81 , 13280 Ballina virus Synonym for Australian bat lyssavirus . Bangor virus A strain of Avian paramyxovirus 2. Isolated from a finch in Baltimore virus A virus isolated in 1942 Northern Ireland. Antigenically related from four nursery outbreaks of gas- to, but distinct from, Yucaipa virus. A troenteritis. Caused severe diarrhea in similar virus has been isolated from a calves. Diameter 40–80 nm. Examination parrot. Bangor virus 46

Collings DF et al (1975) Res Vet Sci 19 , 219 Banna virus (Indonesia-6969) (BAVIn- McFerran JB et al (1974) Arch Ges Virusforsch 6969) A strain of Banna virus , isolated 46 , 281 from mosquitoes in Indonesia.

Bangoran virus (BGNV) An unassigned Banna virus (Indonesia-7043) (BAVIn- vertebrate rhabdovirus. Isolated from 7043) A strain of Banna virus , isolated the mosquito, Culex perfuscus , and from mosquitoes in Indonesia. the Kurrichane thrush, Turdus liboya- nus , in the Central African Republic. Banna virus LY1 A tentative species in Not reported to cause disease in the genus Seadornavirus . humans. Banna virus LY2 A tentative species in Bangui virus (BGIV) An unassigned the genus Seadornavirus . member of the family Bunyaviridae . Isolated from a man with fever, head- Banna virus LY3 A tentative species in ache, and rash in Bangui, Central the genus Seadornavirus . African Republic. Banna virus M14 A tentative species in Banna virus (BAV) The type species of the genus Seadornavirus . the genus Seadornavirus, related species are Kadipiro virus and Liao ning virus . Banna virus TRT2 A tentative species in Originally isolated in Yunnan Province, the genus Seadornavirus . China from the cerebrospinal fluids and sera of patients with febrile illness and Banna virus TRT5 A tentative species in encephalitis. Isolates have also been the genus Seadornavirus . made from mosquitoes, ticks, and some domestic animals in southern China, and from mosquitoes in Indonesia. Banna virus WX1 A tentative species in The virus appears to be widespread in the genus Seadornavirus . South-East Asia, but determination of the exact relationships between the var- Banna virus WX2 A tentative species in ious isolates will require further genetic the genus Seadornavirus . and antigenic analyses. Banna virus WX3 A tentative species in Brown SE et al (1993) Virology 196 , 363 Xu P et al (1990) Chin J Virol 6 , 27 the genus Seadornavirus . Banna virus WX8 Banna virus (China-HN131) (BAVHN- A tentative species in the genus Seadornavirus . 131V) A tentative species in the genus Seadornavirus , isolated in China. Banzi virus (BANV) A species in the genus Flavivirus , serologically a mem- Banna virus (China-HN191) (BAVHN- ber of the Yellow fever virus group. 191V) A tentative species in the genus Isolated in South Africa from a boy Seadornavirus , isolated in China. with a fever. Mosquito-borne. Natural hosts may be cattle and sheep. Found Banna virus (China-HN295) (BAVHN- in South Africa, Kenya, Tanzania, 295) A tentative species in the genus Zimbabwe, and Mozambique. Does Seadornavirus , isolated in China. not appear to be a common cause of disease in humans. Banna virus (China-HN59) (BAVHN- 59V) A tentative species in the genus Barbarie duck parvovirus (BDPV) A Seadornavirus , isolated in China. strain of Duck parvovirus in the genus Dependovirus . Banna virus (Indonesia-6423) (BAVIn- 6423) A strain of Banna virus , isolated Barfin flounder nervous necrosis virus from mosquitoes in Indonesia. (BFNNV) A species in the genus 47 Batken virus (BKNV)

Betanodavirus. Isolated from juvenile base analog A substance with a structure marine fish in Japan. resembling one of the purine or pyri- Morit K et al (2003) Dis Aquat Org 57 , 19 midine bases found in nucleic acid. Watanabe KI et al (2000) Dis Aquat Org It can be incorporated into a growing 41 , 219 nucleic acid chain by substitution for the proper base, resulting in mutations Barmah Forest virus (BFV) A species in or cessation of growth. It is thus an the genus Alphavirus related genetically antimetabolite. to Ross River virus, but not antigenically related to other alphaviruses. First iso- base pair substitution A point mutation lated in 1974 from mosquitoes, Culex (transition or transversion) in a double- annulirostris , collected in the Barmah stranded nucleic acid. In a transition Forest, Victoria, Australia. Shown to there is replacement of one purine by cause a clinical disease, Barmah Forest another or one pyrimidine by another. disease, indistinguishable from Ross In transversion there is replacement of River virus infection in humans, who a purine by a pyrimidine or vice versa . present with a rash, fever, and malaise; sometimes with arthritis/arthralgia. basement membrane A thin extracellular Marsupials are likely primary hosts, structure that forms a substratum for and the virus has been isolated from orderly growing cells and plays a role a wide variety of mosquito species. in cellular growth and cell differentia- Barmah Forest disease became notifi- tion, tissue regeneration, and filtration able in Australia in 1995. The virus of macromolecules between tissue grows well in the C6/36 mosquito compartments. In polarized cells some cell line. enveloped viruses, e.g. rhabdoviruses, bud preferentially from the basement Lee E et al (1997) Virology 227 , 509 membrane. See polarized epithelial Mackenzie JS (1999) Emerg Infect Dis 5 , 1 cells . Quinn HE et al (2005) J Med Entomol 42 , 882 bass virus See largemouth bass virus . barramundi virus-1 (BaV) An unassigned virus in the family Picornaviridae , iso- bat salivary virus Synonym for Rio Bravo lated in Australia from degenerative virus . areas of the brain and retina of diseased larva of barramundi, Lates calcarifer . Batai virus (BATV) A strain of Highly transmissible from diseased to Bunyamwera virus in the genus healthy fish. Orthobunyavirus , belonging antigeni- Glazebrook JS et al (1990) J Fish Dis 13 , 245 cally to the Bunyamwera virus sero- Munday BL et al (1992) Aquaculture 103 , 197 group. Isolated in India, Malaysia, Thailand, former Czechoslovakia, and Barranqueras virus (BQSV) An unas- Ukraine. Mosquito-borne. Not reported signed virus in the family Bunyaviridae . to cause disease in humans. Isolated from Culex ( Melanoconion ) delpontei in Argentina. Antigenically Batama virus (BMAV) A species in the related to Resistencia virus and genus Orthobunyavirus . Isolated from Antequara virus. a bird, Euplectes afra . Not reported to cause disease in humans. Barur virus (BARV) An unassigned virus in the family Rhabdoviridae . With Batken virus (BKNV) A strain of Dhori Fukuoka, Kern Canyon, and Nkolbisson virus in the genus Thogotovirus . Isolated viruses forms the Kern Canyon sero- from mosquitoes and from the sheep group. Isolated from Rattus rattus and tick, Hyalomma p plumbeum in Kirghizia. from the goat tick, Haemaphysalis inter- Antigenically and genetically related to media, in Karnataka, India. Also found Dhori virus. Not reported to cause dis- in northern Canada. Not reported to ease in humans. cause disease in humans. Frese M et al (1997) J Gen Virol 78 , 2453 Batu cave virus (BCV) 48

Batu cave virus (BCV) A strain of Phnom budgerigars. The virus attacks the cells Penh bat virus in the genus Flavivirus . of the immune system, and those which Isolated from bats in Malaysia. Not produce feather and beak. Affected reported to cause disease in humans. birds gradually lose their feathers and develop beak abnormalities. Because Bauline virus (BAUV) A strain of Great the immune system is attacked, affected Island virus in the genus Orbivirus . birds succumb to a variety of second- Isolated from the tick, Ixodes uriae , on ary bacterial, fungal, parasitic, or viral Great Island, Newfoundland, Canada. infections. Antibodies present in puffins and pet- Bassami MR et al (2001) Virology 279 , 392 rels. Not reported to cause disease in Heath L et al (2006) J Virol 80 , 7219 humans. Niagro FD et al (1998) Arch Virol 143 , 1723 Ritchie B et al (1989) Virology 171 , 83 BAY-57-1293 A thiazolylsulfonamide Todd D et al (1991) Arch Virol 117 , 129 which is an inhibitor of herpes virus helicase primase. Not yet licensed for BeAn 157575 virus (BeAnV-157575) A ten- use in humans. tative species in the genus Vesiculovirus . Kleymann G (2004) Antiviral Chem Chemother Isolated from a bird, Pyriglena leucop- 15 , 135 tera . Not reported to cause disease in humans. Bayou virus (BAYV) A species in the Calisher CH et al (1989) Intervirology 30 , 241 genus Hantavirus . Cause of fatal hantavirus pulmonary syndrome in BeAn 277 virus (GMAV) A probable spe- Louisiana, USA. Rodent host is the rice cies in the genus Bunyavirus , isolated rat, Oryzomys palustris . from mosquitoes. Antigenically related Morzunov SP et al (1995) J Virol 69 , 1980 to Guama virus .

BB cells (CCL 59) Cell line initiated from BeAn 293022 virus A strain of Flexal virus posterior trunk tissue, not including in the genus Arenavirus . fins, of 2-year-old brown bullheads. Susceptible to infectious pancreatic BeAn 47693 virus (BUJV) A strain of necrosis virus and is the only cell line Bujaru virus in the genus Phlebovirus . susceptible to channel catfish virus. BeAn 70563 virus A strain of Amapari B-DNA A right-handed conformation of virus in the genus Arenavirus . the DNA double helix which is stable at high humidity and is considered to be BeAn 8582 virus (CAPV) An isolate of the biologically most important form. Capim virus in the genus Orthobunyavirus , B-DNA is in equilibrium with the left- isolated from mosquitoes. handed, Z-DNA, conformation. The equilibrium depends upon parameters BeAr 328208 virus (BAV) such as cations in the environment A strain Wyeomyia virus and covalent modifications of DNA. of in the genus Orthobunyavirus The B-DNA helix has two grooves . Isolated from mos- Sabethes Sabethoides glaucadae- (one major and one minor) as opposed quitoes, ( ) mon to the Z-DNA helix which has only a . Not reported to cause disease in s ingle groove. humans.

Beak and feather disease virus (BFDV) A Bear Canyon virus A species in the genus species in the genus Circovirus. First iso- Arenavirus isolated from Californian lated from cockatoos in Australia with deer mice ( Peromyscus californicus ). psittacine beak and feather disease. Fulhorst CF et al (2002) Emerg Infect Dis Since found in Europe and America. 7 , 717 Causes chronic and often fatal disease in large psittacine birds includ- Bebaru virus (BEBV) A species in the ing parrots, cockatoos, lorikeets, and genus Alphavirus . Isolated in Malaysia 49 benzo[de]isoquinoline-1,3-diones

from mosquitoes. No known associa- Belterra virus (BELTV) A strain of tion with disease in humans. Rift Valley fever virus in the genus Scherer WF et al (1962) Am J Trop Med Hyg Phlebovirus, belonging to the Rift Valley 11 , 269 fever antigenic complex. Isolated from the rodents, Proechimys longicaudatus beclin A mammalian protein that acts and nudicaudatus in Brazil. An arthro- as a tumor-suppressor by interacting pod vector has not been identified. Not with Bcl-2, an anti-apoptosis protein known to cause disease in humans. that can prolong the life of tumor cells. An autophagy-promoting protein. Benevides virus (BENV) A species in Experimentally, beclin was found to the genus Orthobunyavirus, belong- protect mice against fatal Sindbis virus ing to the Capim serogroup. Isolated encephalitis. from a sentinel mouse and Culex mosquitoes at Belém virus laboratory, Liang XH et al (1998) J Virol 72 , 8586 Brazil. Antibodies have been found in Liang XH et al (1999) Nature 402 , 672 Nectomys squamipes rodents in Brazil. Not known to cause disease in humans. Bedsonia An old name for chlamydia, after Sir Samuel Bedson, who discov- Benfica virus (BENV) A strain of ered them. Bushbush virus in the genus Orthobun- yavirus , belonging to the Capim sero- BeH 2251 virus (CDUV) An isolate of group. Isolated from a sentinel mouse, Candiru virus in the genus Phlebovirus . Nectomys squamipes rodents and Culex sp in Belém, Brazil. Not known to Beilong virus (BeV) A novel paramyxo- cause disease in humans. virus found in a persistently infected human mesangial cell line, and named benign epidermal monkeypox virus for a collaboration between Beijing, Synonym for Tanapox virus . China, and Geelong, Australia which led to the discovery. The virus has typi- benign inoculation lymphoreticulosis cal paramyxovirus morphology, but the virus Synonym for cat-scratch disease genome is the largest so far described virus, now known to be a tick-borne (19,212 nt long). The virus origin and rickettsial agent, Bartonella henselae . possible role in disease are unknown. Li Z et al (2006) Virology 346 , 219 bent DNA DNA molecules bend when continuous dA.dT tracks of at least Belém virus (BLMV) An unassigned three contiguous dAs are repeated at and ungrouped virus in the family an interval of 10.5 bp, equivalent to Bunyaviridae . Isolated from the bird spe- one turn of the DNA helix. This may cies Pyriglena leucoptera and Hylophilax result in retarded mobility during naevia in Brazil. Not reported to cause polyacrylamide gel electrophoresis. disease in humans. Bent DNA may facilitate initiation of DNA replication and transcription. Belgrade virus An isolate of the spe- Has been found in autonomously rep- cies now called Dobrava-Belgrade virus licating sequences (ARSs) of yeast and in the genus Hantavirus belonging to in the DNA genome of the baculovirus, the Hantaan antigenic group, which nuclear polyhedrosis virus. caused a fatal meningo-encephalitis. See Dobrava–Belgrade virus . benzo[de]isoquinoline-1,3-diones Com- pounds reported to inhibit replication Belmont virus (BELV) An unassigned of human herpesvirus 1 and vaccinia and ungrouped species in the fam- virus in chick embryo cultures. Infection ily Bunyaviridae . Isolated from Culex of rabbit skin or eye with vaccinia virus annulirostris in eastern Queensland, could be prevented or reduced in sever- Australia. Antibodies found in cattle, ity by treatment with the drugs. wallabies, and kangaroos. Not known Garcia-Gancedo A et al (1979) Chemotherapy to cause disease in humans. 25 , 83 benzothiadiazines 50 benzothiadiazines Compounds that relatively slowly remaining mainly have been shown to block the synthesis cell-associated with spreading CPE. of virus RNA in hepatitis C virus rep- Cause enlargement of infected cells, licon cell systems. Not yet licensed for hence the common name ‘cytomegalo- cl inical use. virus. ’ Latent infection in the salivary Sarisky RT (2004) J Antimicrob Chemother glands and other tissues is frequent. 54 , 14 Large inclusion bodies (‘ owl eye’ inclusions) containing DNA are often Bermejo virus A strain of Andes virus in present in the nuclei and cytoplasm the genus Hantavirus identified from late in infection. The host range is usu- the rodent Oligoryzomys chacoensis , in ally narrow and they generally repli- South America and from a human case cate best in fibroblasts. DNA mol. wt. 6 of hantavirus pulmonary syndrome in 150 1 0 , 240 kb. G C content 58%. Bolivia. Sequences from either or both termini may be present in an inverted form Padula P et al (2002) Emerg Infect Dis 8 , 437 internally. Three genera are identified: Cytomegalovirus , Muromegalovirus , and Berne virus (BEV) See Equine torovirus . Roseolovirus. The type species of the first is Human herpesvirus 5 , the sec- Berrimah virus (BRMV) A species in the ond Murid herpesvirus 1 , and the third genus Ephemerovirus. Isolated in 1981 Human herpesvirus 6 . from healthy sentinel cattle, Bos tau- Synonym : cytomegalovirus group. rus, in Australia. Antigenically related to bovine ephemeral fever, Kimberley, Betaherpesviruses Cytomegalovirus- Malakal, and Puchong viruses. Not related viruses. See Betaherpesvirinae . known to be pathogenic. Betanodovirus A genus in the family Berry–Dedrick phenomenon An exam- Nodaviridae consisting of viruses iso- ple of non-genetic reactivation. Rabbits lated from juvenile marine fish which infected with a mixture of heat- cause nervous necrosis. The type spe- inactivated myxoma virus and infectious cies is Striped jack nervous necrosis virus . rabbit fibroma virus die of myxomatosis. Virions are spherical, non-enveloped It was originally thought to be an exam- 25–30 nm in diameter, containing a ple of genetic cross-reactivation, but genome consisting of two molecules of now appears to result from use by the positive-sense single-stranded RNA, inactivated virus of the DNA-d ependent both of which lack poly A tails at their RNA polymerase of the active virus to 3 ends. The sizes of the RNAs, both of make its own mRNA. which are required for replication, are Berry GP and Dedrick HM (1936) J Bacteriol 3.1 and 1.4 kb. Replication occurs in 31 , 50 the cytoplasm and involves a third subgenomic RNA species 0.4 kb in length. Bertioga virus (BERV) A species in the The viruses cause significant problems genus Orthobunyavirus . Isolated in 1962 in commercial fish hatcheries. Infected from sentinel mice in Bertioga, State of fish develop a vacuolating encephalo- São Paulo, Brazil. Not reported to cause pathy and retinopathy associated with disease in humans. behavioral abnormalities and high mortality. Viral antigens can be found Bet protein A nonstructural protein that in eggs, larvae, and ovaries of hatch- is induced in cells infected with simian ery-reared and wild spawner fish, sug- foamy virus, and is absolutely required gesting that transmission of the virus is for replication. May play a role in virus both horizontal and vertical. latency. Ball LA (1999) In Encyclopedia of Virology , Second edition, edited by A Granoff and RG Betaherpesvirinae A subfamily of the fam- Webster. London: Academic Press, p. 1026 ily Herpesviridae . Nucleotide sequences of subfamily members form a distinct β -propiolactone An organic solvent lineage within the family. Replicate that can be used to prepare subunit 51 big bone disease virus

preparations of virus antigens for use normal peritoneal cells of an adult female in tests such as complement fixation or hamster with a m ethylcholanthrene- hemagglutination. induced fibrosarcoma. Hierholzer JC et al (1996) In Virology Methods Manual edited by BWJ Mahy and HO Bhanja virus (BHAV) An unassigned Kangro. London: Academic Press, p. 47 member of the family Bunyaviridae serologically related to Forecariah virus Betaretrovirus A genus in the family and Kismayo virus. Isolated from ticks Retroviridae , the type species of which in India, Nigeria, Cameroon, Senegal, is Mouse mammary tumor virus (MMTV). the former Yugoslavia, and Italy. Also Virions exhibit a B-type morphology from cattle, sheep, hedgehog, and with prominent surface spike projec- squirrel. Has been described in associa- tions and an eccentric condensed core. tion with a febrile disease in humans. Capsid assembly occurs within the Gaidamovich SY et al (1979) Intervirology cytoplasm to form A-type particles. The 11 , 288 RNA genome, 10 kb in length, is linked Sang R et al (2006) Emerg Infect Dis 12 , 1074 to the primer tRNALys 3 for MMTV Lys 1,2 and tRNA for other members of BHK21 cells (CCL 10) Heteroploid cells the genus. The long terminal repeat derived from kidneys of 5 unsexed (LTR) is about 1300 bases long, with the day-old Syrian or golden hamsters, structure U3-1200, R-15, U5-120. Both Mesocricetus auratus. Used for polyo- exogenous and endogenous species mavirus transformation, aphthovirus are known. The genome contains the vaccine production, and replication usual retroviral genes gag , pro , pol , and studies of many viruses. env plus (in MMTV only) an additional gene sag, located at the 3 end whose Stoker M and Macpherson I (1964) Nature 203 , 1355 product functions as a superantigen. No members of the genus contain an biased hypermutation oncogene. The known species in the There are several genus are Mouse mammary tumor virus , examples of biased hypermutation, Langur virus , Mason–Pfizer monkey virus , e.g. in retroviruses and hepadnaviruses, Jaagsietke sheep retrovirus , and Squirrel involving G to A mutation. However, monkey retrovirus. Related endogenous the best studied example is that of mea- proviruses have been found in other sles virus in the brain associated with mammalian species. measles inclusion body encephalitis or subacute sclerosing panencephali- Corporale M et al (2006) J Virol 80 , 8030 tis. Measles virus isolated from these Gardner MB et al (1994) In The Retroviridae , conditions is found to have many U vol. 3, edited by JA Levy. New York: Plenum residues substituted by C residues, pre- Press, p. 133 dominantly in the matrix protein gene. In extreme cases more than 50% of the BeWo cells (CCL 98) This is the first U’s are mutated to C’s. human endocrine cell type to be maintained in continuous cultivation and was Baczko K et al (1993) Virology 197 , 188 initiated from a malignant gestational choriocarcinoma of the fetal placenta. bicyclams Macrocyclic polyamines consisting of two linked units of cyclam BF-2 cells (CCL 91) The bluegill fry cell (1,4,8,11-tetra-azacyclotetradecane) line was derived from a trypsinized have been found to be potent selective suspension of pooled caudal portions inhibitors of HIV replication in vitro ; of the trunk of 1-year-old fingerlings. they appear to inhibit the uncoating or Susceptible to lymphocystis disease fusion stages of infection. virus of fish and tadpole edema virus de Clercq E et al (1992) Proc Natl Acad Sci of frogs. 89 , 5285

B14FAF28-G3 cells (CCL 14) This line of big bone disease virus Synonym for Chinese hamster cells was derived from osteopetrosis virus. big liver and spleen disease virus 52 big liver and spleen disease virus A type morbillivirus, and Bluetongue virus of hepatitis E virus which causes hepat- would become Bluetongue orbivirus , for osplenomegaly in chickens. In addition example. This proposal is still at the to human infections, viruses related discussion stage and has not yet been to hepatitis E virus have been found adopted by the ICTV. in pigs and chickens. The virus is cur- van Regenmortel MHV and Mahy BWJ rently termed avian hepatitis E virus (2004) Emerg Infect Dis 10 , 8 and is a tentative species in the genus Hepevirus . bioassay Determination of the infectious Huang FF et al (2002) J Clin Microbiol 40 , titer of a virus by measuring its biologi- 4197 cal activity (e.g. infectivity for its host). Payne CJ et al (1999) Vet Microbiol 68 , 119 biological containment Reducing or Bijou Bridge virus (BBV) A strain of eliminating the risk of viruses or other Tonate virus in the genus Alphavirus . microorganisms escaping from the Isolated from a mixed infection with laboratory. Containment conditions Fort Morgan virus from the Cimicidae usually involve reducing the labora- (Cliff swallow nest ) bug, Oeciacus vicar- tory air pressure so that it is negative ius , in eastern Colorado, USA. Related with respect to the environment, pass- to Cabassou virus , within the VEEV ing exhaust air through high efficiency antigenic complex. Tonate virus was particulate air (HEPA) filters, and steri- previously isolated only from birds in lizing all fluid waste by boiling. See French Guiana. also biosafety. Kinney RM et al (1983) J Gen Virol 64 , 135 Monath TP et al (1980) Am J Trop Med Hyg biological control Pest control agents of 29 , 969 biological origin including parasites, predators and pathogens. Some viruses Biken-1 virus Probably a strain of Seoul have been used as biological control virus in the genus Hantavirus . agents, e.g. myxoma virus and rabbit Isegawa Y et al (1990) Nucleic Acids Res 18 , hemorrhagic disease virus for rabbit 4936 control and baculovirus for insect pest control. Biken CAM virus An attenuated strain of Measles virus used as a vaccine in biosafety Work with viruses infectious Japan. to humans poses a risk of laboratory acquired infection. The currently Bimbo virus (BBOV) An unassigned ver- accepted guidelines for biosafety and tebrate rhabdovirus. Isolated from a appropriate containment levels for var- healthy specimen of the Golden Bishop ious viruses can be found in Biosafety bird, Buplectes afer , in Central African in Microbiological and Biomedical Republic. Not reported to cause disease Laboratories, Fifth Edition (2007), edited in humans. by LC Chosewood and DE Wilson. Published by the US Department of Bimiti virus (BIMV) A species in the Health and Human Services. genus Orthobunyavirus , belonging to the Guama serogroup. Isolated from biotechnology Industrial processes Culex spissipes in Trinidad. Also found requiring the use of biological systems, in Brazil, Guinea, and Surinam. Not including genetic engineering, fermen- reported to cause disease in humans. tation technology, hybridoma technology, and agricultural technology. binomial nomenclature It has been proposed that a binomial system of virus biotin A small water-soluble macromole- nomenclature be adopted in which the cule (vitamin B complex) which is a coen- virus name is followed by the name zyme in carboxylation–d ecarboxylation of the genus to which it belongs, So reactions. Used as a non-radioactive Measles virus would become Measles reporter group for labeling a ntibodies 53 BK polyomavirus (BKPyV)

or nucleic acid probes. It has a very now three genera: Aquabirnavirus , high affinity for avidin (streptavidin) Avibirnavirus , and Entomobirnavirus . which, when coupled with an indicator molecule (enzyme, fluorescent dye), is birth defects See congenital infection . used to detect biotinylated anti-species antibody. Bittner mouse mammary tumor virus A Gould EA et al (1985) J Virol Methods 11 , 41 strain of Mouse mammary tumor virus in the genus Betaretrovirus. The first of biphasic milk fever virus Synonym for the mouse mammary tumor viruses Tick-borne encephalitis virus . isolated from C3H mice. It is a highly oncogenic virus transmitted in the milk. Foster-nursed mice are free of the Birao virus (BIRV) A strain of Bunyamwera virus but may continue to carry a mam- virus in the genus Orthobunyavirus , mary tumor virus of low oncogenicity. belonging to the Bunyamwera sero- The virus is also known as C3H mam- group. Isolated from Anopheles pharoen- mary tumor virus and as MTV-S. See sis and A squamosus in Central African Mouse mammary tumor virus . Republic. Not reported to cause disease Synonym : milk factor. in humans. bird flu Popular name for influenza virus biundulant meningoencephalitis virus A (H5N1). Synonym for Tick-borne encephalitis virus . bird papillomavirus Synonym for fringilla papillomavirus. Bivens Arm virus (BAV) An unassigned animal rhabdovirus in the birdpox virus Synonym for Fowlpox family Rhabdoviridae. Isolated from virus . Culicoides insignis in Florida, USA. Antigenically related to Coastal Plains and Tibrogargan viruses, found only in Birnaviridae A family of viruses roughly Australia. Not known to cause disease spherical in shape with a diameter of in humans. 60 nm, sedimenting at 650S and band- ing in CsCl at 1.33 g/ml; there is no Gibbs EPJ et al (1989) Vet Microbiol 19 , 141 envelope. The icosahedral capsid structure is a single protein layer based on a BK polyomavirus (BKPyV) A species in T 13 lattice, and is composed of four the genus Polyomavirus . Distinct from major polypeptides. The genome com- JC polyomavirus and other members of prises two segments of linear double- the genus. Agglutinates human group stranded RNA, 3092 and 2784 bp in O erythrocytes. Originally isolated from length in infectious pancreatic necrosis the urine of a patient on immunosup- virus, and 3129 and 2795 bp in infec- pressive therapy after renal transplanta- tious bursal disease virus. Both seg- tion. Presence of antibodies in humans ments contain a 94 kDa genome-linked suggests it is a common infection of protein. There are no poly A tracts at humans, probably of the kidneys, usu- the 3 ends of the segments, and there ally silent but activated by immuno- is no evidence for 5 capping of any of suppression. Can be propagated in Vero the viral mRNAs. The family contains cells, human diploid lung fibroblasts viruses infecting fish, chickens, insects, (WI 38) or primary human fetal kidney and rotifers, and contains three gen- cells. Has not been associated with pro- era, Aquabirnavirus , Avibirnavirus , and gressive multifocal leukoencephalo- Entomobirnavirus . pathy (caused by JC virus). Transforms rat and hamster cells in culture and is Dobos P (1995) Annu Rev Fish Dis 5 , 25 oncogenic on injection into newborn Galloux M et al (2007) J Biol Chem 282 , 20774 Pous J et al (2005) J Gen Virol 86 , 2329 hamsters. Has been associated with mild respiratory disease in children. Birnavirus Former name for a genus Synonym : polyomavirus hominis. in the family Birnaviridae . There are Goudsmit J et al (1982) J Med Virol 10 , 91 BK polyomavirus (BKPyV) 54

Imperiale MJ (2000) Virology 267 , 1 routine screening include hepatitis B, Marshall WF et al (1990) J Clin Microbiol 28 , hepatitis C, human immunodeficiency 1613 viruses 1 and 2, and human T-cell lym- Nishimoto Y et al (2006) J Mol Evol 63 , 341 photropic viruses I and II. Unfortunately transmission of other viruses may Black Creek Canal virus (BCCV) A spe- occur, particularly as a consequence of cies in the genus Hantavirus. The cause organ transplantation. For example one of a non-lethal case of hantavirus pul- donor transmitted rabies virus to four monary syndrome in Florida, USA. organ transplant recipients all of whom Host is the cotton rat, Sigmodon hispidus . died about 2 weeks later since the diag- Rollin PE et al (1995) J Med Virol 46 , 36 nosis of rabies in the donor was not suspected. black-footed penguin herpesvirus Synonym for sphenicid herpesvirus 1. Srinivasan A et al (2005) N Engl J Med 352 , 11 black stork herpesvirus Synonym for blood group P antigen A neutral glycoli- ciconiid herpesvirus 1. pid called globoside which is expressed on erythrocytes and erythroid progeni- black-tailed deer adenovirus A probable tors. It is the cell surface receptor for species in the genus Mastadenovirus . human parvovirus B19. Causes a fatal hemorrhagic disease in blacktailed deer, Odocoileus hemionus Bloodland Lake virus (BLLV) A strain columbianus. The virus is closely related of Prospect Hill virus in the genus to Bovine adenovirus B . Hantavirus , isolated from voles, Microtus ochrogaster, in the USA and Canada. Not Lapointe JM et al (1999) Arch Virol 144 , 393 known to be associated with disease in Lehmkuhl HD et al (2001) Arch Virol 146 , 1187 humans.

BL cells Burkitt’s lymphoma cells Blotched snakehead virus A probable species in the genus Aquabirnavirus bleomycin An antibiotic that is use- found in a fish cell line. Genome ful in chemotherapy of patients with sequence studies suggest that it may be Kaposi’s sarcoma. more closely related to the Avibirnavirus genus, however. The structure of the blind passage Transmission of material viral protease is unusual, and may be a from an inoculated animal or cell cul- target for antiviral chemotherapy. ture which shows no evidence of infec- Da Costa B et al (2003) J Virol 77 , 719 tion, to a fresh animal or cell culture. Feldman AR et al (2006) J Mol Biol 358 , 1378 Evidence of infection may appear after John RK and Richards RN (1999) J Gen Virol several such blind passages. 80 , 2061

BLO-11 cells (CCL 198) A fibroblast-like blotting A technique first developed by cell line derived from the abdomi- Sir Edwin Southern in Oxford to trans- nal wall muscle of a 30-day-old male fer nucleic acids or protein to an immo- Blotchy mouse (Moblo/Y) with heredi- bilizing matrix such as nitrocellulose, tary emphysema. nylon filters or diazobenzyloxymethyl paper. See Southern blotting ; Northern blood borne virus infections Many virus blotting ; and Western blotting . infections can be present in blood for short periods when viremia occurs dur- blue comb virus An early name for Turkey ing infection. Other infections remain coronavirus , the cause of turkey blue chronic and persistent in the blood, comb disease, also known as transmis- and blood destined for transfusion is sible enteritis of turkeys. screened by a variety of methods to remove any positive units to eliminate blue ear disease A rapidly fatal disease transfusion–transmission of virus infec- of piglets. See Porcine reproductive and tion. In the USA, viruses eliminated by respiratory syndrome virus . 55 Bobaya virus (BOBV) blue eye Corneal keratitis in dogs, 1–2 size 19.2 kb. There are three large (3.9– weeks after canine adenovirus infection. 2.8 kb), three medium (2.0–1.8 kbp), and four small (1.1–0.8 kb) segments. blue fox parvovirus A tentative species in Virions are icosahedral, 80 nm in dia- the genus Parvovirus . Related to mink meter. Unlike the Orthoreovirus , there is enteritis virus. Isolated from the blue a diffuse outer capsid removed by expo- fox, Alopex lagopus . sure to CsCl. The outer coat may con- Veijalainen PM and Smeds E (1988) Am J Vet tain 92 capsomeres. The inner shell has Res 49 , 1941 a diameter of 54–64 nm and contains 32 large ring-shaped capsomeres. There blue gill virus See American oyster are seven viral proteins. Replication reovirus . occurs on yolk sac inoculation of 6-day- old eggs at 33.5°C. Replication with Blue River virus A strain of Sin Nombre CPE occurs in cell cultures of lamb kid- virus in the genus Hantavirus . Isolated ney, hamster, and chick embryo tissue from Peromyscus leucopus rodents and in BHK21 cells. An egg-attenuated from Indiana and Oklahoma in the vaccine is effective if polyvalent. MidWestern USA. Synonyms: ovine catarrhal fever virus; sore mouth virus. bluegill hepatic necrosis reovirus A Barber TL and Jochim MM (Editors) (1985) strain of Aquareovirus A , related to Bluetongue and Related Orbiviruses . New American oyster reovirus 13p2. York: Alan Liss Maan S et al (2007) J Gen Virol 88 , 621 Bluetongue virus (BTV) The type spe- Mertens PPC et al (2007) J Gen Virol 88 , 2811 cies of the genus Orbivirus . Twenty-five Roy P and Noad R (2006) Curr Top Microbiol Immunol 309 , 87 serotypes can be identified by neutralization tests. There is a group-specific bluetongue viruses 1–25 (BTV-1 to 25) CF antigen. Causes a serious disease of Serotypes of Bluetongue virus . sheep with a mortality of 5–30%. There is fever, edema of the head and neck, blunt end DNA fragments generated by cyanosis, erosions around the mouth, certain restriction endonucleases, e.g. sometimes pulmonary edema, and Hae III, and which are perfectly base- lameness due to involvement of the paired along their entire length, i.e. they hooves and muscle damage. Cattle and do not carry single-stranded regions goats develop a much milder disease. after cleavage with the enzyme. Foot lesions occur in pigs. Wild ruminants are often infected. The infection ‘ BLV-HTLV viruses ’ An old name for is prevalent between 50° North and 30° viruses now included in the genus South of the Equator. Occurs mainly in Deltaretrovirus . Africa, especially in the east and south, but has occurred in Cyprus, Palestine, B lymphocytes See B cells . Turkey, Spain, Portugal, Pakistan, India, Japan, and southern and western USA. B-lymphotropic papovavirus strain Recently the virus has spread to north- K38 Synonym for African green monkey ern Europe including Belgium, the polyomavirus . Netherlands, and the UK. Bluetongue virus is transmitted by nocturnal bit- B-lymphotropic polyomavirus (LpyV) A ing flies of the genus Culicoides , and it strain of African green monkey polyoma- appears that they have survived over virus in the genus Polyomavirus . winter in Europe. Virus replication occurs in the midges but there is no boa herpesvirus An unassigned member evidence of transovarial transmission. of the family Herpesviridae . Synonym Infectivity is ether-resistant. Virions for boid herpesvirus 1. have a sedimentation coefficient of 550S and contain 20% double-stranded Bobaya virus (BOBV) An unassigned and RNA which is in 10 segments, total ungrouped member of the Bunyaviridae Bobaya virus (BOBV) 56

family. Isolated in suckling mice from boot-strapping A method for assessing the pooled brain, liver, and heart of an confidence in phylogenetic analyses. adult African thrush, Turdus libonya- Felsenstein J (1985) J Mol Evol 39 , 783 nus , netted at M’Boko, Central African Republic. Not known to infect humans. Boraceia virus (BORV) A strain of Anopheles B virus Bobia virus (BIAV) A strain of Olifantsvlei in the genus Orthobunyavirus virus in the genus Orthobunyavirus . . Isolated from mos- Anopheles cruzii Phoniomyia With Olifantsvlei, Dabakala, and Oubi quitoes and pilicauda virus forms the Olifantsvlei serogroup. in Casa Grande, State of São Isolated from Culex trigripes at Bobia, Paulo, Brazil. Not reported to cause Central African Republic. Not known disease in humans. Together with to cause disease in humans. Anopheles B virus forms the Anopheles B serogroup. bobwhite quail herpesvirus Synonym Lopes OS and Sachetta LA (1974) Am J for perdicid herpesvirus 1. Epidemiol 100 , 410

Bocas virus A coronavirus, closely Border disease virus (BDV) A species related to or identical with mouse hep- in the genus Pestivirus . Causes a dis- atitis virus. On first isolation thought ease of sheep first described from the to be an arbovirus of the California borders of Wales and England. A con- serogroup. genital condition of newborn lambs Bardos Y et al (1980) Intervirology 13 , 275 characterized by an abnormally hairy birthcoat and a tremor. There is defective myelination of the CNS caused by Bocavirus A genus of the family the virus crossing the placenta to infect Parvoviridae containing three species the fetus. Virus can be isolated from the closely related by their genome DNA CNS of affected lambs. A similar dis- sequence. They are bovine parvovi- ease has been described in Germany, rus, canine minute virus, and human Australia, and New Zealand. The virus bocavirus. replicates in cultures of primary calf and fetal lamb kidney cells. Cell deple- Bohle iridovirus (BIV) A species in the tion is seen and infected cells can be genus Ranavirus, isolated in north- stained with fluorescent-labeled anti- ern Queensland, Australia from the serum. Infected lambs develop antibodies ornate burrowing frog Limnodynastes to hog cholera virus and to bovine viral ornatus in which it causes disease soon diarrhea virus to which border disease after metamorphosis. Experimentally virus is antigenically related. Injection it causes disease in several species of of non-pregnant ewes causes no obvi- native Australian frogs and also in the ous disease, but in pregnant animals fish, barramundi (Lates calcarifer). there is necrotizing carunculitis, abor- Couper BEH et al (2005) Arch Virol 150 , 1797 tion, and border disease in those young Marsh IB et al (2002) Mol Cell Probes 16 , 137 which are not aborted. Synonyms : pestivirus ovis; hairy shaker boid herpesvirus 1 (BoiHV-1) An disease of lambs. A virus related to unassigned member of the family BDV was found in an outbreak of fatal Herpesviridae. Isolated from a young disease in Pyrenean chamois (Rupicapra boa constrictor. pyrenaica pyrenaica ) in Spain in 2002. Hauser B et al (1983) J Comp Pathol 93 , 515 Arnal M et al (2004) J Gen Virol 85 , 3653 Barlow RM et al (1983) J Comp Pathol 93 , 451 Bolivian hemorrhagic fever virus Collett MC et al (1989) J Gen Virol 70 , 253 Synonym for Machupo virus . Meyers G and Thiel H-T (1996) Adv Virus Res 47 , 53 Bollinger bodies Intracytoplasmic acidophilic inclusion bodies found in cells Border disease virus BD31 An isolate of infected with fowlpox virus. Border disease virus . 57 bottlenose dolphin parainfluenza virus

Border disease virus X818 An isolate of Bornaviridae A family within the order Border disease virus . Mononegavirales. Consists of the genus Bornavirus of which Borna disease virus Borna disease virus (BDV) The only spe- is the type and only species. cies in the genus Bornavirus. Named after a town in Saxony where the first Bornavirus The only genus in the family major outbreak in horses occurred. Bornaviridae, containing only one spe- Negative-strand RNA virus which pro- cies, Borna disease virus named after duces persistent infection in a variety Borna, a town in Saxony, where many of experimental animals. Virions are horses died of neurological disease enveloped with spherical morphol- during an epidemic in 1885. Virions ogy 90 nm in diameter and contain a are spherical, with a diameter of about genome of single-stranded RNA, 8.9 kb 90 nm, and sensitive to heat and lipid in length. Replication involves the cell solvents. The genome is a single mol- nucleus. Horses and sheep appear to ecule of linear negative-stranded RNA be the main natural hosts. Causes las- 8.9 kb in length. It codes for at least 6 situde, followed by excitation with ORFs in the order 3 -N-P/X-M-G-L- tonic spasms and later paralysis in 5 . The virus is highly neurotropic and horses. A similar disease is produced causes CNS disease in several non- in sheep, cattle, and probably deer. human vertebrate species, including A reservoir in the bicolored shrew horses. (Crocidura leucodon ) has been suggested from studies in Switzerland. Sporadic Bornholm disease A name assigned in cases of natural infection have been Denmark in 1932 to a local outbreak of reported in donkeys, mules, and lla- a disease more accurately referred to as mas. Guinea pigs, rabbits, rats, and Bamble disease. mice can be infected experimentally. Human infection has been suggested Borrel bodies Minute granules compos- by the reported detection of virus- ing the Bollinger bodies found in fowl- related sequences in peripheral blood pox virus-infected cells. leukocytes of some patients with psychiatric disorders. These results are Borrielota variolae bovis Elementary controversial and require confirmation. bodies associated with cells infected The disease in horses has been detected with smallpox or Vaccinia virus . mainly in Saxony but also in other parts of Germany, the Principality of Goodpasture EW (1933) Science 77 , 119 Liechtenstein, Poland, Rumania, Russia, Syria, and Egypt. A virus causing stag- bos1–bos10 Serotype designation given gers in horses in Nigeria may be the to the mammalian adenovirus iso- same. The virus has been isolated from lated from cattle, Bos taurus . Genus ticks of several genera and from the Mastadenovirus , family Adenoviridae . brains of herons and other wild birds. Transmission by oral and nasal secre- Botambi virus (BOTV) A species in tions is possible. A virus vaccine has the genus Orthobunyavirus . Isolated been used with success. Replicates on from mosquitoes, Culex guiarti, in the the CAM and in cultures of lamb testis Central African Republic. Not reported and monkey kidney cells with CPE. to cause disease in humans.

Gonzalez-Dunia D et al (2000) J Virol 74 , Boteke virus (BTKV) A tentative species 3441 in the genus Vesiculovirus. Isolated from Hilbe M et al (2006) Emerg Infect Dis 12 , 675 the mosquito, Mansonia maculipennis , in Koprowski H and Lipkin WI (1995) Curr Top Central African Republic. Not reported Microbiol Immunol 190 , 134pp Lipkin WI et al (2001) Trends Microbiol 9 , 295 to cause disease in humans. Planz O et al (1999) J Virol 73 , 6251 Staeheli P et al (2000) J Gen Virol 81 , 2123 bottlenose dolphin parainfluenza virus Wolff T et al (2006) J Clin Virol 36 , 309 A novel parainfluenza virus isolated bottlenose dolphin parainfluenza virus 58

from the lung tissue of an Atlantic bot- Associated worldwide with respiratory tlenose dolphin (Tursiops truncatus ). infection and conjunctivitis in cattle. Nollens HN et al (2008) Vet Microbiol Serological evidence suggests a high 128 , 231 incidence of infection. Replication with CPE in bovine kidney cell cultures. Bouboui virus (BOUV) A species in the Agglutination of erythrocytes of several genus Flavivirus , antigenically a member species: mouse, monkey, cattle, horse, of the Yellow fever virus group. Isolated goats, guinea pig and hamster. from mosquitoes and a baboon, Papio Benkö M et al (1988) Intervirology 29 , 346 papio , in the Central African Republic, Mohanty SB (1978) Adv Vet Sci Comp Med Senegal, and Cameroon. Probably 22 , 83 present in Zaire. Not reported to cause disease in humans. Bovine adenovirus A (BAdV-A) Originally called BAdV-1. Boudicca A retrovirus-like long termi- Evans PS et al (1998) Virology 244 , 173 nal repeat retrotransposon present in the genome of the human blood fluke Bovine adenovirus B (BAdV-B) Originally Schistosoma mansoni . Named after the called BAdV-3. Celtic warrior-queen Boudicca who led Reddy PS et al (1999) Virology 253 , 299 a revolt against the Romans in London in ad 61. Bovine adenovirus C (BAdV-C) Originally Copeland CS et al (2003) J Virol 77 , 6153 called BAdV-10. Matiz K et al (1998) Virus Res 55 , 29 bovid herpesvirus 2 Synonym for Bovine herpesvirus 2 . Bovine adenovirus D (BadV-D) Strains include the original serotypes BadV-4, bovid herpesvirus 3 Alcelaphine her- See 5, 8 and Rus. pesvirus 1 . Bovine adenovirus E (BadV-E) Originally bovine AAV Synonym for Bovine adeno- called BadV-6. associated virus . Bovine adenovirus F (BadV-F) Originally Bovine adeno-associated virus (BAAV) called BAdV-7. A species in the genus Dependovirus . Found in association with bovine aden- Bovine astrovirus (BastV) There are two ovirus types 1, 2, and 3. Not known to recognized strains of Bovine astrovirus . be pathogenic. The expression profile of BAAV is similar to that of human AAV5, and also avian adeno-associated virus. bovine astrovirus 1 and 2 (BastV-1 and BastV-2) Strains of Bovine astrovirus in Synonym : bovine AAV. the genus Mamastrovirus similar, mor- Coria MF and Lehmkuhl HD (1978) Am J phologically, to human astrovirus. Can Vet Res 39 , 1904 be grown in primary bovine kidney Luchsinger E and Wellemans G (1971) Arch Ges Virusforsch 35 , 203 cell cultures in the presence of trypsin. Myrup AC et al (1976) Am J Vet Res 37 , 907 Isolated from calves with diarrhea, in Qiu J et al (2006) J Virol 80 , 5482 association with calicivirus, coronavirus, and rotavirus. No clinical signs bovine adenoviruses Originally 10 sero- attributable to bovine astrovirus infec- types of BAdV were described, but tion have been reported. now, on the basis of sequence, only six Aroonprasert D et al (1989) Vet Microbiol species are recognized. Three are in the 19 , 113 genus Mastadenovirus (BadV A, B, and Woods GN et al (1985) J Clin Microbiol 22 , 668 C) and one (BAdV D) is a species in the genus Atadenovirus . Two others (BadV E bovine calicivirus (VESV/Bos-1) A strain and BadV F) are tentative species in the of Vesicular exanthema of swine virus , iso- genus Atadenovirus, family Adenoviridae . lated from cattle. 59 Bovine ephemeral fever virus (BEFV)

Bovine coronavirus (BCoV) A species in and both agglutinate bovine erythro- the genus Coronavirus which causes neo- cytes at 5–8°C. Serotype 2 viruses have natal calf diarrhea. Under natural con- only been isolated from domestic cat- ditions, affects calves 1 day to 3 weeks tle, but serotype 1 viruses have been old. Only infects bovine species. Virus isolated from a wide range of species can be found in feces and diagnosis including domestic cattle, Bos taurus , made by electron microscopy examina- water buffalo, Bubalus bubalis, sheep, tion, reverse passive hemagglutination Ovis aries, goats, Capra hircus, Sika or ELISA. Virus of the same serotype deer, Cervus nippon, African buffalo, has also been isolated from trachea and Syncerus caffer, impala, Aepyceros melam- lungs of calves with respiratory disease, pus, and Australian brushtail possums and on injection into newborn calves by (Trichsurus vulpecula ). Transmission is the nasal route caused both respiratory by oral–fecal spread, and the bovine disease and diarrhea. The virus repli- enteroviruses are very resistant to acid cates in cell cultures of bovine kidney, and alkali so they pass easily through bovine embryonic lung or the human the digestive tract. Most infections are rectal tumor cell line HRT-18. Another subclinical, but occasional cases of abor- disease, winter dysentery of adult cat- tion, still-birth, and infertility in bulls tle, has been associated with the pres- have been ascribed to bovine entero- ence of bovine coronavirus in the feces virus infection. One strain is reported to of affected animals. produce diarrhea in colostrum-deprived Synonym : neonatal calf diarrhea calves. Recent genome sequencing stud- coronavirus. ies show that the bovine enteroviruses McNulty MS et al (1984) Vet Microbiol 9 , 425 are most closely related to the porcine Mebus CA et al (1973) Am J Vet Res 34 , 145 enteroviruses. Park SJ et al (2006) J Clin Microbiol 44 , 3178 Synonym : ecboviruses. Reynolds DJ et al (1985) Arch Virol 85 , 71 Knowles NJ and Barnett ITR (1985) Arch Virol 83 , 141 bovine diarrhea virus See Bovine viral Zell R et al (2006) J Gen Virol 87 , 375 diarrhea virus 1 . Zheng T (2007) Arch Virol 152 , 191 bovine encephalitis herpesvirus Syno- bovine enzootic leukosis virus Synonym nym for Bovine herpesvirus 5 . for Bovine leukemia virus . bovine enteric calicivirus strain NB Bovine ephemeral fever virus (BEFV) A (BEC-NB) An unassigned species in species in the genus Ephemerovirus iso- the family Caliciviridae. Identified in lated in South Africa, Australia, and fecal samples of cattle in the USA and Japan from cattle with fever, respira- in England (Newbury agent), but no tory symptoms, increased salivation, antigenic cross-reaction was found. lacrimation, joint pains, tremors, and Significance and importance in calf stiffness. Cattle and water buffalo are diarrhea not clear. susceptible to clinical disease but other ruminant species may have subclini- Bridger JC et al (1984) Infect Immun 43 , 133 cal infection. The disease is of short duration and is not contagious, and bovine enteric torovirus See Bovine in nature is spread only by an insect torovirus . bite. Virus replicates in mice and in cell cultures such as BHK21. Loses patho- Bovine enterovirus (BEV) Two serotypes genicity for calves on passage in new- of Bovine enterovirus are recognized born mice, hamsters or BHK21 cells. within the genus Enterovirus . Can be isolated in eggs by inoculation of embryo. In Kenya transmitted by bovine enteroviruses 1 and 2 (BEV-1 and Culicoides sp, probably from the reser- BEV-2) Serotypes of Bovine enterovirus voir hosts buffalo and water buck. No in the genus Enterovirus . Picornaviruses evidence of human infection. isolated in cell cultures from bovine tis- Synonyms : dengue fever of cattle virus; sues or excreta. There are two serotypes ephemeral fever virus; bovine epizootic Bovine ephemeral fever virus (BEFV) 60

fever virus; bovine influenza virus; three- infected experimentally develop fever; day stiff-sickness virus; stiffsiekte. in rabbits there is meningo-encepha- Della-Porta AJ and Brown F (1979) J Gen litis with paralysis of the hind legs. Virol 44 , 99 Transmission of the virus is by contact, Gaffar Elamin MA and Spradbrow PB (1978) especially under crowded conditions. J Hyg ( Camb ) 81 , 1 Has been cultivated in bovine embryo Walker PJ (2005) Curr Top Microbiol Immunol cell cultures, with CPE in 1–2 days, but 292 , 57 there is loss of virulence. Replication Walker PJ et al (1994) J Gen Virol 75 , 1889 also occurs in pig, sheep, goat, and horse kidney cell cultures and in human bovine epitheliosis virus Synonym for amnion cultures. No evidence of human Alcelaphine herpesvirus 1 . infection has been recorded. There is transformation of hamster cells in vitro . bovine epizootic fever virus Synonym No growth in eggs. All strains are anti- for Bovine ephemeral fever virus . genically very similar and numerous Bovine foamy virus (BFV) A species in the attenuated, as well as inactivated, virus genus Spumavirus . Isolated from buffy vaccines are available. coat, spleen, lymph nodes, and milk of Synonyms : infectious bovine rhinotra- lymphosarcomatous and normal cat- cheitis virus; infectious pustular vul- tle. Can be propagated in BHK21 cells vovaginitis virus; necrotic rhinitis producing syncytia. Little free virus is virus; red nose virus. produced. Infection appears to be com- Straub OC (1990) In Virus Infections of mon in cattle in the USA and Europe, Ruminants , edited by Z Dinter and B Morein. but there is little evidence of disease. Amsterdam: Elsevier, p. 71 Produces a persistent infection. Different Thiry J et al (2006) Vet Res 37 , 169 from Bovine respiratory syncytial virus . Tikoo SK et al (1995) Adv Virus Res 45 , 191 Dermott E et al (1971) J Gen Virol 12 , 105 Greig AS (1978) Can J Comp Med 43 , 112 Bovine herpesvirus 2 (BoHV-2) A species Kong XH et al (2005) Arch Virol 150 , 1677 in the genus Simplexvirus. Not distinguishable from Allerton virus. Isolated bovine hemadsorbing enteric virus in South Africa from lumpy skin disease Synonym for Bovine parvovirus . and once thought to be the cause of that condition. Causes deep, slowly healing bovine herpes mammillitis virus Syno- ulcers on the teats and udders of milk- nym for Bovine herpesvirus 2 . ing cows, and lesions are produced on Bovine herpesvirus 1 (BoHV-1) A spe- the lips of calves which suckle them. In cies in the genus Varicellovirus . A natu- South Africa the virus has been isolated ral worldwide infection in cattle, but from buffalo, Syncerus caffer, in which it antibodies can also be found in mule produces disease, and may be transmit- deer, Odocoileus hemionus , and ferrets ted by insects or ticks. Infection of day- are susceptible to disease in the USA. old rats, mice, and hamsters may lead Wild ruminants in Africa are probably to stunted growth, rashes, and death. the original host. The virus has also There is replication in calf kidney cell been isolated from soft-shelled ticks, cultures with formation of large syncy- Ornithodorus coriaceus , collected from tia. A viable unattenuated virus given mule deer in western USA, but it is not i.m. gives some protection, but no com- clear whether ticks are a reservoir host. mercial vaccine is available. Shares 14% May cause a silent, mild infection, or of DNA sequences with human herpes- acute disease of the whole respiratory virus 1. tract. Mortality can be as high as 75%. In Synonyms : Allerton virus; bovid her- Europe it has been known to cause con- pesvirus 2; bovine herpes mammillitis junctivitis and, notably, disease of the virus; bovine mammillitis virus; bovine genital tract when lesions appear on the ulcerative mammillitis virus; pseu- external genitalia. There is no evidence dolumpy skin disease virus. of antigenic difference between the res- Wellenberg GJ et al (2002) Vet Microbiol piratory and genital strains. Young goats 88 , 27 61 Bovine leukemia virus (BLV) bovine herpesvirus 3 Renamed Alcelap- c ounterparts in HIV-1. Transmission has hine herpesvirus 1 . not been well studied, but iatrogenic transmission through reuse of syringes Bovine herpesvirus 4 (BoHV-4) A species and needles is likely. in the genus Rhadinovirus. Isolated from Gonda MA (1992) AIDS 6 , 759 cattle in Germany and the USA, and Gonda MA et al (1987) Nature 330 , 388 in the UK and Africa from pulmonary adenomatosis of sheep, but is probably bovine influenza virus Synonym for not the cause of that condition. African Bovine ephemeral fever virus . buffaloes have a high seroprevalence, and may be the natural host species. Bovine leukemia virus (BLV) Type spe- Cell-free transmission to cultures is cies of the genus Deltaretrovirus. Occurs possible, but animals are not infected worldwide, especially in dairy cat- by cultivated virus. Infection is prob- tle. Causes enzootic bovine leukosis, ably by the respiratory route, but most a B-cell lymphoma, in some infected contact experiments failed. May be a animals, but persistent inapparent infec- cause of minor respiratory disease in tion is common and may involve 80% calves which predisposes them to bacte- of an affected herd. Mitogen-stimulated rial disease of the respiratory tract, but cultures of lymphocytes from infected in general the virus appears to be non- cattle produce virus, but in bovine cells pathogenic. Viruses which appear to be only a little virus is produced. However, strains of BoHV-4 have been isolated in bat lung cells, fetal lamb kidney and from cats and owl monkeys, and mis- several other cell lines, virus is con- takenly reported as feline cell-associated tinuously released in abundant quan- herpesvirus and aotine herpesvirus 2, tity. Cattle, sheep, goats, and rabbits respectively. This might reflect contami- injected with the virus become infected, nation of biologicals used in virus isola- produce antibodies to the viral internal tion, such as bovine sera, by BoHV-4. antigen, and some sheep and cattle later Synonym : Movar herpesvirus. develop lymphosarcoma. Human, simian, bovine, ovine, bat, and caprine cells Bartha A et al (1987) Intervirology 28 , 1 Bublot M et al (1990) J Gen Virol 71 , 133 exposed to the virus or bovine leuke- Bublot M et al (1991) J Gen Virol 72 , 715 mic cells form syncytia. Antiserum pre- Dewals B et al (2005) Vet Microbiol 110 , 209 vents syncytia formation. This can be Thiry E et al (1992) Vet Microbiol 33 , 79 used to assay both virus and antiserum. Antibody-positive animals are common Bovine herpesvirus 5 (BoHV-5) A species in herds with high incidence of leuke- in the genus Varicellovirus. Isolated from mia and colostrum provides protection calves with encephalitis in Australia. against infection in the first 5–6 months Causes encephalitis in European breeds of life. Infection is acquired by contact of cattle. with infected animals after the maternal Synonyms: bovine encephalitis herpes- antibody has disappeared. The virus virus; herpesvirus caprae. differs from other mammalian type C Brake F and Studdert MJ (1985) Aust Vet J retroviruses in producing syncytia, in 62 , 331 not having the interspecies mammalian Engels M et al (1986) Virus Res 6 , 57 antigen, and in having a DNA polymer- Thiry J et al (2006) Vet Res 37 , 169 ase with preferential activity in the presence of Mg2 . The RNA genome is Bovine immunodeficiency virus (BIV) A a dimer, 8.4 kb in length (each mono- species in the genus Lentivirus . Isolated mer), with tRNAPro base-paired to in 1972 from a Holstein cow obtained serve as primer RNA for reverse tran- from a herd in Louisiana, USA, with per- scription. Several strains have been sistent lymphocytosis. Morphologically completely sequenced. Only one anti- similar to HIV-1. The RNA genome genic type is known, but there is some is 8.5 kb in length and contains five relationship with human T-cell lympho- accessory genes in addition to gag , pol , tropic virus types 1 and 2. However, and env , namely tat , rev , vif , W , and Y , there is no evidence of human infection which have some sequence similarity to with bovine l eukemia virus. Control is Bovine leukemia virus (BLV) 62

by elimination of seropositive animals mice, and connective tissue tumors from infected herds. (sarcoids) in horses. Cause transfor- Synonyms: bovine enzootic leukosis mation of embryo cultures of mouse, virus; bovine leukosis virus; bovine hamster, and bovine tissues. The major type C oncovirus; enzootic bovine leu- transforming protein of BPV1, E5, is kosis virus. a small hydrophobic protein local- Burny A (1988) Vet Microbiol 17 , 197 ized in the endoplasmic reticulum, Dube S et al (2000) Virology 277 , 379 which activates several cellular protein Mussgay M and Kaaden OR (1978) Curr Top kinases, including growth factor recep- Microbiol Immun 79 , 43 tors. Virus particles (full or empty) agglutinate mouse erythrocytes at 4°C bovine leukosis virus Synonym for between pH 6.8 and 8.4. Elutes read- Bovine leukemia virus . ily at 37°C. Receptors not destroyed by influenza virus neuraminidase. bovine mammillitis virus Synonym for Bovine herpesvirus 2 . Chen EY et al (1982) Nature 299 , 529 Lancaster WD and Olson C (1982) Microbiol bovine norovirus-CH126 A tentative spe- Rev 46 , 191 Roperto S et al (2005) Vet Pathol 42 , 812 cies in the genus Norovirus , isolated Tsirimonaki E et al (2006) Virus Res 115 , 158 from cattle in the Netherlands in 1998. van der Poel WHM et al (2000) Emerg Infect Bovine papular stomatitis virus (BPSV) A Dis 6 , 36 species in the genus Parapoxvirus . Causes a usually benign, non-febrile disease, bovine norovirus-Jena A tentative spe- most often in young cattle. There are cies in the genus Norovirus , isolated crateriform ulcers up to 1 cm in diam- from cattle in Germany in 1980. eter in the mouth. Some strains may Liu BL et al (1999) J Virol 73 , 819 infect sheep and goats. Transmission to humans is reported, causing local skin bovine noroviruses Viruses with a calici- lesions, similar to those caused by orf virus morphology were first reported virus or by pseudocowpox virus. in cattle in the UK in 1978, (Newbury Synonyms : erosive stomatitis virus of agent 1) and subsequently in Germany cattle; papular stomatitis of cattle virus; in 1980. These viruses are genetically pseudo-aphthous stomatitis of cattle distinct, but share a cross-reactive virus; stomatitis papulosa of cattle virus; a ntigenic epitope with human noro- ulcerative stomatitis of cattle virus. viruses. There is some evidence that the Gassman U et al (1985) Arch Virol 83 , 17 bovine noroviruses can infect humans Menna A et al (1979) Arch Virol 59 , 145 in close contact with cattle, such as farm-w orkers and veterinarians. Bovine parainfluenza virus 3 (BPIV-3) A Oliver SL et al (2006) J Clin Microbiol 44 , 992 species in the genus Paramyxovirus first Widdowson et al (2005) J Med Virol 76 , 119 isolated from cattle with shipping fever in the USA. Related to human parainflu- Bovine papillomavirus 1 (BPV-1) A spe- enza virus 3, but can be distinguished cies in the genus Deltapapillomavirus by a variety of tests including reac- with two genotypes, bovine papilloma- tion with monoclonal antibodies and viruses 1 and 2 (BPV-1 and -2), origi- genome sequence analysis. Widespread nally recognized by restriction enzyme endemic infection of cattle populations digest patterns of viral DNA. A natural worldwide. Replicates in a variety of infection of cattle causing papillomas cells in vitro, including calf, goat, buf- with underlying fibroma mainly on the falo, and camel kidney cell cultures. head, neck, legs, back, and abdomen Inoculation of calves causes fever, con- but also in the mouth and esophagus. junctivitis, and rhinitis. The virus is associated with the etiol- Synonym : shipping fever virus. ogy of bladder tumors. Experimental Rydbeck R et al (1987) J Gen Virol 68 , 2153 inoculation of BPV types 1 or 2 pro- Shibuta H et al (1979) Microbiol Immunol duces slowly growing fibrosarcomas 23 , 617 on injection into hamsters and C3H/EB Shioda T et al (1988) Virology 162 , 388 63 bovine rotavirus

Bovine parvovirus (BPV) A species in the in swine embryonic kidney cells, ham- genus Parvovirus, first known as Haden ster lung and kidney cells, monkey Vero (h em ad sorbing en teric) virus. A com- cells, and human embryonic lung and mon infection of cattle; can cause enteri- kidney cells. Inactivated by lipid sol- tis and diarrhea in calves. Has usually vents. Reciprocal antigenic cross-reaction been isolated from fecal specimens from with human respiratory syncytial virus. calves with enteric disease, but can be Synonym : respiratory syncytial virus of isolated from many tissues after injec- bovines. tion into colostrum-deprived calves. Lehmkuhl HD et al (1979) Am J Vet Res The only strains antigenically differ- 40 , 124 ent from the original Haden strain of Stott EJ and Taylor G (1985) Arch Virol 84 , 1 bovine parvovirus have been isolated in Japan. Unrelated antigenically to other bovine respiratory viruses Respiratory parvoviruses. Agglutinates erythrocytes disease is an important problem in the of several species including guinea cattle industry. Viruses appear to be the pig and humans. Replicates in bovine primary cause but stress and weather embryo kidney cell cultures with CPE. play an important role and viral infec- Synonyms: bovine hemadsorbing tion predisposes to bacterial invasion. enteric virus; Haden virus; hemadsorb- The viruses are: ing enteric virus of calves. Cotmore S and Tattersall P (1987) Adv Virus Alcelaphine herpesvirus 1 Res 33 , 91 Bovine adenovirus ( A , B and C) Storz J and Bates RC (1973) J Am Vet med Bovine ephemeral fever virus Assoc 163 , 884 Bovine herpesvirus 1 and 4 Bovine parainfluenza virus 3 Bovine polyomavirus (BPyV) A species Bovine respiratory syncytial virus in the genus Polyomavirus , originally bovine rhinoviruses 1–3 isolated from a stump-tailed macaque Bovine viral diarrhea viruses 1–2 kidney cell culture. After a few pas- Mammalian orthoreoviruses sages the cells showed vacuolation and Mohanty SB (1978) Adv Vet Sci Comp Med signs of degeneration. Subsequently, 22 , 83 an identical virus was isolated from cultured kidney cells of a newborn calf, bovine rhinoviruses 1–3 (BRV-1, -2, and -3) and it became clear that the stump- Serotypes in the genus Rhinovirus , not tailed macaque virus was of bovine yet assigned to a species. Three sero- origin, probably introduced into the types have been described. Serotype 1 culture as a contaminant of fetal calf contains strains Sd-1, 181-V (Germany), serum. Antibodies to the virus have C-07, VC-96, and FS1-43 (USA), RS3X been found in persons having close (UK) and M47 and Chitose (Japan); sero- contact with cattle, e.g. 71% of vet- type 2 contains EC-11 (UK); serotype 3 erinary practitioners tested in the UK. contains H-1 (Japan). Replicate best in Infection has not been associated with calf kidney cells at 33°C and low bicar- disease in humans or cattle. bonate. A widespread infection with low Strains : stump-tailed macaque virus; pathogenicity for calves, causing fever, fetal rhesus kidney virus. nasal discharge, cough, and difficulty in Parry J et al (1983) Arch Virol 78 , 151 breathing. Does not infect other animals. Schuurman R et al (1990) J Gen Virol 71 , 1723 Lupton HN et al (1980) Am J Vet Res 41 , 1029 Yamashita H et al (1985) Arch Virol 83 , 113 Bovine respiratory syncytial virus (BRSV) A species in the genus Pneumovirus . bovine rotavirus A probable species in the Causes a mild to severe respiratory genus Rotavirus. Rotaviruses of all six disease in cattle. Widespread in most species (groups A–F) may infect calves, European countries, North America, causing diarrhea. Circulating antibody Australia, Japan, and North Africa. does not protect against rotavirus infec- Replicates in bovine kidney and lung cell tion, although rotavirus antibody in cultures causing syncytia. Also re plicates the gut from colostrum or milk does bovine rotavirus 64

provide protection. Calf rotavirus infec- Hoet AE and Saif LJ (2004) Anim Health Res tions are highly contagious and occur Rev 5 , 157 worldwide. A similar disease is caused by neonatal calf diarrhea coronavirus. bovine type C oncovirus Synonym for Synonym : calf rotavirus. Bovine leukemia virus; enzootic bovine leukosis virus. Ojeh CK et al (1984) Arch Virol 79 , 161 Saif LJ et al (1994) Curr Top Microbiol Immunol 185 , 339 bovine ulcerative mammillitis virus Synonym for Bovine herpesvirus 2 . bovine spongiform encephalopathy (BSE) Bovine viral diarrhea virus 1 (BVDV-1) A transmissible fatal neurodegenerative The type species of the genus Pestivirus , prion disease of cattle first detected in with four recognized serotypes, NADL, 1986 in the UK. Although the epidemic Osloss, SD-1 and CP7. The genome is was restricted to the UK and Ireland, single positive-stranded RNA, about sporadic cases occurred in, e.g. Canada, 12.3 kb long. Infection appears to be France, Germany, Switzerland, Oman, confined to artiodactyls – cattle, sheep, Portugal, Italy, and the Falkland Islands. pigs, buffaloes, moose, and deer. Both Incubation period is 2–8 years. Can be cytopathogenic and non-cytopathogenic experimentally transmitted to mice, pigs, strains can be isolated. Most infections cats, and bovines. Natural infection is by are inapparent, but a small number of the oral route. Believed to have origi- cattle in the herd develop severe, fre- nated by adaptation of sheep scrapie to quently fatal mucosal disease with cattle around 1981/1982, and was exac- diarrhea, fever, and oral ulceration. erbated by feeding cattle-derived meat Necrotic lesions are found in mucosa, and bone meal to cattle. This was banned hooves, and lymph nodes. If primary in 1988 and the epidemic is declining in infection occurs during pregnancy the UK. See prion diseases . the virus may cross the placenta, Bradley R (1991) Eur J Epidemiol 7 , 532 causing abortion or fetal abnormali- Hope J (1995) Nature 378 , 761 ties. Infection of the fetus with a non- Prusiner S (Editor) (1999) Prion Biology and cytopathic strain before development of Diseases. Cold Spring Harbor: Cold Spring Harbor Laboratory Press immunological competence (110 days ’ gestation) can result in the animal being persistently infected with BDV for life. bovine syncytial virus (BSV) See Bovine It is these animals that maintain the foamy virus . virus in the population, and that may develop severe mucosal disease which Bovine torovirus (BoTV) A species in develops in such persistently infected the genus Torovirus . Originally discov- cattle when they become infected with ered in 1979 in cattle in Breda, Iowa, a second cytopathic strain of BDV. The and named Breda virus, it was associ- disease is often mild and antibodies ated with severe neonatal calf diarrhea. may be present in most members of a Not adapted to growth in cell culture. herd. Antibodies not found in humans Evidence of infection by bovine toro- or horses. A related virus occurs in virus has been found in every country sheep in southern Germany and pigs where serological or virological stud- in Australia. Serial passage in rabies have been done, including Western bits leads to attenuation of virulence Europe, India, South Africa, and New for cattle and this virus may be used Zealand. As with other members of as a vaccine. There are probably at the family Coronaviridae, the genome of least seven antigenically distinguish- Breda virus is a long strand of positive- able types of the virus. It is spherical, sense RNA, 28,475 kb in length. 57 nm in dia meter with a 24 nm wide Synonym : Breda virus. core, and envelope without projections. Draker R et al (2006) Virus Res 115 , 56 Infectivity sensitive to lipid solvents, Horzinek MC (1999) Encyclopedia of Virology , exposure to 56°C and to pH 3 or below. Second edition, edited by A Granoff and Replicates in bovine cell cultures with- RG Webster. London: Academic Press, p. 1798 out CPE. 65 Breu Branco virus

Synonyms : mucosal disease virus; pes- Bowen’s disease A pre-cancerous squa- tivirus diarrhea virus; diarrhea virus of mous cell carcinoma of the nail beds bovines; pestivirus bovis; bovine virus of humans which has been linked to diarrhea virus; bovine diarrhea virus. infection with HPV 16. Baxi M et al (2006) Vet Microbiol 116 , 37 Belak S and Hakhverdyan M (2006) Dtsch Box turtle virus 3 (TV3) A strain of Frog Tierarztl Wochenschr 113 , 129 virus 3 in the genus Ranavirus . Brownlie J and Clarke MC (Editors) (1990) Bovine Virus Diarrhea: Scientific and Technical Bozo virus (BOZOV) A serotype Review . Paris: OIE of Bunyamwera virus in the genus Harkness JW et al (1978) Res Vet Sci 24 , 98 Orthobunyavirus. Isolated from the mosquito, Aedes opok , in Central African bovine viral diarrhea virus 1 CP7 A Republic. Not reported to cause disease strain of bovine viral diarrhea virus 1. in humans. bovine viral diarrhea virus 1 NADL A Saluzzo JF et al (1983) Ann Virol ( Inst Pasteur ) strain of bovine viral diarrhea virus 1. 134E , 221 bovine viral diarrhea virus 1 Osloss A branched DNA assay A nucleic acid strain of bovine viral diarrhea virus 1. detection assay. A type of signal amplification assay performed in microtiter bovine viral diarrhea virus 1 SD-1 A plates. RNA or DNA molecules released strain of bovine viral diarrhea virus 1 from virions are captured using specific (BVDV-1). probes bound to the surface of the plate. Bovine viral diarrhea virus 2 (BVDV-2) Branched DNA molecules are added to A species in the genus Pestivirus . Three the well followed by an alkaline phos- isolates of BVDV, strains 890, New York phatase-linked synthetic probe which ’93 and C413, which differ serologically is detected by chemiluminescence, from BVDV-1. which is proportional to the concentration of target nucleic acid in the original bovine viral diarrhea virus 2 C413 A specimen. strain of bovine viral diarrhea virus 2. branched DNA probes Reagents used for bovine viral diarrhea virus 2 strain signal amplication which consist of a New York ’93 A strain of bovine viral network of simultaneous hybridization diarrhea virus 2. events between multiple sets of oligonucleotide probes (capture probes, extender, bovine viral diarrhea virus 2 strain 890 A and amplifier secondary probes, labelled strain of bovine viral diarrhea virus 2. tertiary probes, etc.) For the detection of HIV, branched DNA assays can detect as bovine visna-like virus Probably a species few as 50 viral genomes per milliliter of of the genus Lentivirus . Isolated from plasma. a cow with persistent lymphocytosis. Replicates in cultures of bovine embry- Brazil 2– 4 virus An isolate of hantavi- onic spleen cells, with the slow forma- rus from Rattus norvegicus in Brazil. tion of multinuclear giant cells similar Antigenically belongs to the Seoul virus to those produced by bovine syncytial serotype. virus. However, the two viruses differ morphologically and antigenically. Breakbone fever virus Synonym for Experimental infection of cows results Dengue virus . in enlarged peripheral lymph nodes and hyperlymphocytosis. The disease Breda virus (BRV) See Bovine torovirus . may last over a year, during which time the virus can be re-isolated. Breteau index A measure of the mosquito Georgiades JA et al (1978) J Gen Virol infestation rate used in arbovirology. 311 , 375 van der Maaten MJ et al (1972) J Natl Cancer Breu Branco virus An unclassified virus Inst 49 , 1649 isolated from Culicine mosquitoes in Breu Branco virus 66

the Amazon region of Brazil. Not asso- bromovinyldeoxyuridine See Brivudin . ciated with disease in humans. bronchovirus syncytialis Synonym for brilliant cresyl blue A photoreactive dye. Human respiratory syncytial virus . See photodynamic inactivation . Brovavir See Sorivudine . Bristol virus An isolate of a Sapporo- like virus (Bristol 98) was found dur- brown bullhead papillomavirus A virus ing surveillance of gastroenteritis in found in papillomas of the brown bull- South West England, 1997–2000. The head, Ictalurus nebulosus . complete genome sequence of the virus showed differences from Sapporo virus brown trout rhabdovirus Rhabdoviruses Manchester, suggesting the existence of have been isolated from brown trout, two distinct genetic groups of Sapporo- Salmo trutta, in Finland, Italy, and like viruses in the UK. Northern Ireland using a variety of fish cell lines. The virus appears to be sero- Robinson S et al (2002) J Med Virol 67 , 282 logically related to pike fry rhabdovirus but not to spring viraemia of carp Brivudin A halogenated nucleoside ana- rhabdovirus. log, bromovinyl deoxyuridine (BVDU) Koski P et al (1992) Bull Eur Assoc Fish Pathol with a similar but less active inhibitory 12 , 177 action against herpesvirus replication. Rowley H et al (2001) Dis Aquat Organ 48 , 7 Descamps J et al (1982) J Virol 43 , 332 Bruconha virus (BRUV) A strain of Broadhaven virus (BRDV) A strain of Caraparu virus in the genus Orthobunya- Great Island virus in the genus Orbivirus , virus belonging to the group C viruses. belonging to the Kemerovo serogroup. Isolated from mosquitoes, Culex (Melanoconion ) sp. Broad-range polymerase chain reac- Calisher CH et al (1982) Proceedings of the tion The use of a variety of primers to International Symposium on Tropical Arboviruses amplify genetic sequences in order to and Hemorrhagic fevers , Belem, Brazil, April implicate a virus or group of viruses 14–18, 1980. Braz Acad Sci, p. 355. that may be involved in the etiology of a suspected viral disease. Priming Brunswick bird plague virus A strain of sites that are conserved among a broad avian influenza A virus. group of viruses are used initially, then more specific primers are used to posi- Brus Laguna virus (BLAV) A virus in the tively identify the species involved. genus Orthobunyavirus , belonging to the Gamboa serogroup. Isolated from bromelain A proteolytic enzyme iso- mosquitoes, Aedeomyia squamipennis . lated from pineapple stem which can Calisher CH et al (1988) Am J Trop Med Hyg be used to remove the peplomers from 39 , 406 the surface of enveloped viruses such as influenza. brushtail possum virus See possum Compans RW et al (1970) Virology 42 , 880 papillomavirus .

5-bromo-2-deoxyuridine A halogenated Bryan strain of Rous sarcoma virus A pyrimidine which can become incorpo- high titer strain, highly oncogenic, and rated into cellular DNA in place of thy- much used in early experimental work. midine. Incorporation may activate the A defective virus which contains the transcription of viral RNA from genes src oncogene and lacks an env gene, integrated into the cellular DNA, such and so requires a helper avian leukosis as those of Retroviridae . virus for replication. The env gene can also be supplied in trans by a helper bromovinylarabinosyl-uracil See Sori- cell line, such as transfected Qt6 cells, vudine . in which the virus grows to high titer. 67 Buenaventura virus (BUEV)

Boerkoel CF et al (1993) Virology 195 , 669 place. Occurs with many enveloped Rubin H (2003) Carcinogenesis 24 , 803 animal viruses. The viral ribonucleo- Sudol M et al (1986) Nucleic Acids Res 14 , protein, with or without a surrounding 2391 membrane protein, associates with an area of the cell membrane which starts BSC-1 (CCL 26) A continuous cell line to form a coat around the virus protein. from kidneys of the African green mon- In electron micrographs it appears as key, Cercopithecus aethiops . Develops a crescentic structure. The cell mem- characteristic CPE when infected with brane closes around the virus and the Simian virus 40 . A valuable tool for viral particle leaves the cell. During the bud- diagnostic studies. ding process, all cellular proteins in the area of membrane destined to become BSL-4 Biosafety level 4 Reserved for the virus coat are replaced by virus- dangerous viral pathogens such as coded proteins. Thus, in the mature Ebola virus, for which no vaccines or virus particle, the envelope lipid is host antiviral treatments exist. cell-derived, but all proteins are virus- coded. The exact mechanism of this BT 4971 virus (PATV) A strain of Patois process is unknown. virus , in the genus Orthobunyavirus . Budgerigar fledgling disease polyoma- B-type inclusion body Inclusion bodies virus (BFPyV) A species in the genus observed in the cytoplasm of vertebrate Polyomavirus . Isolated from budgerigar poxvirus-infected cells, representing embryo fibroblasts inoculated with tis- sites of virus synthesis; characteristic of sues of affected birds. Also grown in all productive poxvirus infections. chick embryo fibroblasts. Genome is double-stranded DNA, 5.1 kb, circular, B-type virus particles A morphologically similar to other polyomaviruses. The defined group of enveloped RNA virus disease is clinically similar to psitta- particles seen typically outside the cells cine beak and feather disease. Avian in mouse mammary carcinoma. They polyoma viruses have also been iso- are always extracellular. They have a lated from buzzard (Buteo buteo ), fal- dense body or core (nucleoid) 40–60 nm con (Falco tinnunculus ), and lovebirds in diameter, contained within a mem- (Agapornis pullaria ) with fatal disease. branous sac or envelope 90–120 nm in The isolated viruses all showed close diameter. The membrane is covered sequence similarity to BFPyV. with protrusions. The nucleoid is usually eccentrically located within the Davis RB et al (1981) Avian Dis 25 , 179 envelope. Occasionally one envelope Johne R and Müller H (1998) Arch Virol 143 , may contain more than one nucleoid. 1501 B-type particles bud through the cell Lehn H and Müller H (1986) Virology 151 , membrane and contain an internal con- 31162 stituent morphologically indistinguishable from an A-type particle, which budgerigar parainfluenza virus A species becomes the core. The prototype is of Avian paramyxovirus 5 in the genus Mouse mammary tumor virus . Avulovirus. Isolated from the lung of caged budgerigars in Kunitachi, Japan Bernard W (1960) Cancer Res 20 , 712 during an outbreak of severe enteritis. Dalton AJ (1972) J Natl Cancer Inst 49 , 323 Similar to Newcastle disease virus but distinct from it and from Yucaipa virus. Bubaline herpesvirus 1 (BuHV-1) A spe- Synonym : Avian paramyxovirus 5 cies in the genus Varicellovirus . A her- (Kunitachi) virus. pesvirus isolated from water buffalo. Brake F and Studdert MJ (1985) Aust Vet J Nerome K et al (1978) J Gen Virol 38 , 293 62 , 331 de Carlo E (2004) Vet Rec 154 , 171 Buenaventura virus (BUEV) A serotype of Punta Toro virus in the genus budding A method of virus release from Phlebovirus. Isolated from Lutzomyia sp the cell in which replication has taken in Colombia. buffalopox virus (BPXV) 68 buffalopox virus (BPXV) A strain of Bujaru virus (BUJV) A species in the genus Vaccinia virus in the genus Ortho- Phlebovirus, belonging to the sandfly poxvirus . Causes occasional outbreaks fever group. With Munguba virus forms of severe disease in India, and has been the Bujaru antigenic complex. Isolated reported in Egyptian water buffalo. from the rodent, Proechimys guyannensis The pocks produced on the CAM are oris, in Para, Brazil. Not known to cause similar to those produced by vaccinia disease in humans. virus, but they do not appear at temperatures above 38°C. Although some Bukalasa bat virus (BBV) A species in strains may differ from vaccinia virus, the genus Flavivirus . Isolated from bats many cases of clinical disease in buffa- of Tadarida sp in Uganda. Serologically loes are caused by viruses that appear a member of the Rio Bravo virus group. to be identical to it, based on sequence No known arthropod vector. analysis of the envelope protein gene. Very similar to Lenny virus. Bulbar poliomyelitis The most serious Synonym : vaccinia subspecies. clinical form of poliomyelitis in which Baxby D and Hill BJ (1971) Arch Ges the medulla oblongata (the bulb) is Virusforsch 35 , 70 involved. This affects the upper spine Singh RK et al (2006) Arch Virol 151 , 1995 and the brainstem and consequently Tantawi HH et al (1977) J Egypt Vet Med may involve paralysis of the respira- Assoc 37 , 15 tory system.

Bufo bufo United Kingdom virus A Bundibugyo ebolavirus A probable spe- strain of Frog virus 3 in the genus cies in the genus Ebolavirus . Caused at Ranavirus. Caused the deaths of many least 16 human deaths in Bundibugyo, frogs and toads in the UK in 1993. a thickly forested area in Uganda, 250 Buggy Creek virus A serotype of Fort miles west of Kampala, in 2007. Morgan virus in the genus Alphavirus . Isolated from the bird nest bug, Bungowannah virus A probable species Oeciacus vicarius . Not associated with in the genus Pestivirus. Isolated dur- disease in humans. ing an outbreak of sudden deaths in 3–4-week-old piglets on a farm in New Calisher CH et al (1988) Am J Trop Med Hyg South Wales, Australia, who developed 38 , 447 myocarditis. Later in the epidemic Bu (IMR-31) cells (CCL 40) A line of near many foetuses were stillborn. The virus diploid fibroblasts from male yearling could be transmitted experimentally to buffalo, Bison bison. Finite life expect- porcine foetuses, and was identified ancy of at least 30 passages. Supports following sequence independent single replication of a large number of viruses. primer amplification (SISPA) as a pestivirus related by phylogenetic analysis Buist bodies A synonym and probably to pestiviruses isolated from giraffe or the proper name for the elementary pronghorn antelope. particles known as Paschen bodies seen Kirkland PD et al (2007) Virus Res 129 , 26 in cells infected with smallpox or vaccinia virus. The credit for their discov- Bunyamwera serogroup A group of more ery is usually given to Paschen (1906), than 20 antigenically related viruses but Buist’s account anticipates this by isolated from mosquitoes that are many years. members of the genus Orthobunyavirus . Buist JB (1886) Proc R Soc Edinb 13 , 603 Bunyamwera supergroup of arbo- Buistia pascheni A synonym proposed viruses Old name for the genus for Paschen bodies in order to com- Orthobunyavirus . memorate the work of both Paschen and Buist. See Buist bodies . Bunyamwera virus (BUNV) A species in Mackie TJ and van Rooyen CE (1937) Edin the genus Orthobunyavirus . A member Med J 44 , 72 of the Bunyamwera serogroup. Isolated 69 Bwamba virus (BWAV)

in Uganda, Nigeria, Cameroon, Kenya, serogroup. Isolated in 1962 from a and Central African Republic where it bird, Sylvia curraca, in Bahig, Egypt. has caused a few cases of fever with a Probably also present in Europe since rash in humans. Mosquito-borne. the bird was viremic on arrival in Egypt on its way south. Not reported Bunyaviridae A large family of arthropod- to cause disease in humans. borne, enveloped, spherical or pleomorphic viruses 80–120 nm in diameter Buritirana virus An unclassified arthropod- with glycoprotein surface projections. borne virus isolated in the Amazon There are three internal ribonucleocap- region of Brazil. sids which are circular helical strands, 2–2.5 nm in diameter, 200–3000 nm long. Burkitt’s lymphoma virus Synonym for The RNA genome is composed of three Human herpesvirus 4 . segments of negative or ambisense single-stranded RNA, termed L (large), burst size In a one-step growth curve, M (medium), and S (small) with lengths the burst size is the average number of of 6.5–9.0, 3.2–6.3, and 0.8–2.0 kb, respec- progeny virus particles produced per tively. There are four major virion pro- infected cell. teins: L (transcriptase), N (nucleocapsid), G1 and G2 (external glycoproteins). The busatin See methisazone . L-RNA encodes the L protein; M RNA encodes the glycoproteins which are Buschke–Lowenstein tumor A rare ano- co-translationally cleaved to form G1 genital form of condyloma acuminata and G2, and in some viruses a nonstruc- which is locally invasive but non- tural protein NSM; the S-RNA encodes metastasizing. These tumors usually the N protein, and in some viruses a contain human papillomavirus 6 or 11. non-structural protein NSS. Virions develop in the cytoplasm and mature by Bushbush virus (BSBV) A species in the budding into smooth-surfaced vesicles genus Orthobunyavirus belonging to the in the Golgi region or nearby, but matu- Capim serogroup. Isolated from mos- ration at the plasma membrane has also quitoes in Trinidad and Belem, Brazil. been observed. There are five genera: Not known to cause disease in humans. Orthobunyavirus , Phlebovirus , Nairovirus , Hantavirus , and Tospovirus . Bussuquara virus (BSQV) A strain of Barrett ADT and Shope RE (2005) In Topley & Aroa virus in the genus Flavivirus . Wilson’s Microbiology and Microbial Infections , Isolated from humans, sentinel howler vol. 2, Tenth edition, edited by BWJ Mahy monkeys, Alouatta sp, sentinel mice and V ter Meulen. London: Hodder Arnold, and Culex spp in Colombia, Panama p. 1025 and Para, Brazil. A case of febrile dis- Bouloy M (1991) Adv Virus Res 40 , 235 ease in humans has been reported. Elliott RM (1990) J Gen Virol 71 , 501 bunyavirus A member of the family Buttonwillow virus (BUTV) A strain Bunyaviridae . of Manzanilla virus in the genus Ortho- bunyavirus, belonging to the Simbu Bunyip Creek virus (BCV) A serotype serogroup. Isolated from rabbits, hares of Palyam virus in the genus Orbivirus . and Culicoides spp in Kern County, Isolated from cattle, Bos taurus , in California also New Mexico and Texas, Grafton, Australia. USA. Probably present in other parts of the USA. Not reported to cause disease buoyant density The density of a par- in humans. ticle expressed in terms of the density of a fluid in which it neither sinks BVDV B ovine v iral d iarrhea v irus. See nor floats. See isopycnic gradient Bovine viral diarrhea virus 1 . centrifugation . Bwamba virus (BWAV) A species in the Burg el Arab virus An unclassified genus Orthobunyavirus. With Pongola a rbovirus belonging to the Matariya virus forms the Bwamba serogroup. Bwamba virus (BWAV) 70

Isolated from nine cases of fever in myeloid leukemias of any inbred mouse humans in Bwamba, Uganda. Also from strain. The BXH-2 mouse strain was Anopheles funestus . Present also in Nigeria obtained by crossing C57BL/6 J with and the Central African Republic. C3H/HeJ mice, two strains with low lymphoma incidence. The integration BXH-2 virus A strain of Murine leukemia site of the BXH-2 provirus is within the virus in the genus Gammaretrovirus pro- Nf1 tumor-suppressor gene. duced by recombinant inbred BXH-2 Bedigian HG et al (1981) J Virol 39 , 632 mice, that have one of the highest spon- Largaespada DA et al (1995) J Virol 69 , 5095 taneous incidences of retrovirus-induced C

C1q complement protein The first com- C6/36 cells A line of insect cells derived ponent of the classical complement from the mosquito, Aedes albopictus . pathway, the first line of defense against Supports the growth of many arboviruses. Human C1q binds directly to viruses, including dengue virus. certain viruses, and this result in activation of the complement system and Barth OM (1992) Mem Inst Oswaldo Cruz 87 , 565 neutralization of the virus by virolysis (damage to the virion envelope). CA protein The capsid protein ( p24) of C2 The second component of complement. HIV which forms the viral core enclosing the genome and enzymes required C211 cells (CCL 123) A cell line derived for infectivity. from a skin biopsy obtained from a 11-year-old Caucasian male with Cri Cabassou virus (CABV) A species in du Chat syndrome. The cell line has the genus Alphavirus , antigenically a partial deletion of the short arm of related to Venezuelan equine encepha- chromosome 4. litis virus. No association with human disease. C3 complement protein The third component of complement which can directly Cacao virus (CACV) A strain of Chilibre activate the alternative pathway, and virus in the genus Phlebovirus , in the can cause opsonization of an antigen. Chilibre antigenic complex of the Reacts fourth in the hemolytic com- sandfly fever group. Isolated from plement activation sequence, and is Lutzomyia sp in Panama. Not reported the most abundant of the complement to cause disease in humans. proteins. Cache Valley virus (CVV) A strain C3a complement protein A biologically of Bunyamwera virus in the genus active fragment of C3, which is an Orthobunyavirus. Isolated in N Dakota, anaphylatoxin. Maryland, Virginia and Indiana, USA, and in Jamaica. Mosquito-borne. C4 complement protein The fourth com- Antibodies are found in humans, mon- ponent of complement which actually keys, horses, cattle, deer, and wild reacts second in the hemolytic comple- rodents. Can cause disease in humans, ment activation sequence. but as laboratories rarely test for it, the true incidence is unknown. C5 complement protein The fifth component of complement. Activated by C5 Blackmore CGM and Grimstad PR (1998) convertase, which splits off a small frag- Am J Trop Med Hyg 59 , 704 ment, C5a, leaving C5b, the remainder Campbell GL et al (2006) Emerg Infect Dis 12 , 854 of the molecule, which forms a complex Inverson JO et al (1979) Can J Microbiol 25 , with C6, C7, C8, and C9 known as the 760 membrane attack complex that has an affinity for cell membranes and medi- Cacipacore virus (CPCV) A species in ates immune cytolysis and hemolysis. the genus Flavivirus , in the Japanese encephalitis group, with no known vec- C5a complement protein A 74 residue tor. Isolated from the blackfaced ant- peptide derived from C5 by tryptic bird, Formicarius analis , in Para, Brazil. cleavage. A chemotactic factor and anaphylatoxin. Batista WC et al (2001) Virus Res 75 , 35 CaCo2 cell line 72

CaCo2 cell line A continuous human a plentiful source of vaccine that was colon adenocarcinoma cell line derived used to eradicate smallpox from many from a 72-year-old Caucasian male. countries, but in tropical countries it Can be used for isolation and growth was unstable at high temperatures there, of human astroviruses. and so in the 1950s Leslie Collier, work- Willcocks MM et al (1990) Arch Virol 113 , 73 ing at the Lister Institute in London, developed a lyophilized vaccine that Caddo Canyon virus (CDCV) An unas- was stable up to 3 months at 37°C. signed, ungrouped member of the Bunyaviridae family. Isolated from the calf rotavirus See bovine rotavirus . tick, Ixodes baergi, in Caddo canyon, Oklahoma, USA. Caliciviridae A family of non-enveloped RNA viruses, 35–40 nm in diameter, caesium chloride density gradient cen- icosahedral with 32 typical surface trifugation Method for separating structures, sometimes described as hollows or cups, hence the name molecules or viruses according to their density. Sedimentation ceases when from Latin: calyx cup. The cap- the molecules reach the position in the sid is composed of 180 protein molecules arranged in dimers to form gradient that is the same as their own buoyant density, termed ‘ the isopycnic 90 capsomeres arranged in a T 3 point. ’ icosahedral lattice. The genome consists of positive single-stranded RNA 6 Caimito virus (CAIV) A tentative spe- of mol. wt. 2.6–2.7 1 0 (7.4–8.3 kb cies in the genus Phlebovirus , antigeni- in length). During replication, a sub- cally unrelated to other members of genomic mRNA (2.4 kb) is synthesized the genus. Isolated from the fan fly, and encodes the major capsid protein. Lutzomyia lephilator , in central Panama. There is one major virion protein of Not reported to cause human disease. mol. wt. 58,000–60,000. A protein of mol. wt. 10–15 1 0 3 which is essential for infectivity, is covalently linked Cajazeira virus (CAJV) An unclassified virus isolated from the viscera of to the 5 end of virion RNA. Replicates bats of undetermined species in the in the cytoplasm. There are four gen- Amazon region of Brazil. era: Lagovirus , Norovirus , Sapovirus , and Vesivirus . Species include Rabbit Calabazo virus (CALV) A species in hemorrhagic disease virus , Norwalk virus , the genus Hantavirus . Isolated from Sapporo virus, and Vesicular exanthema cane mice, Zygodontomys brevicauda , of swine virus. Other viruses in the fam- in Panama. Not so far associated with ily (unassigned) include species which human disease. infect reptiles, walrus, pigs, cattle, chickens, mink, pigs, and even insects. Vincent MJ et al (2000) Virology 277 , 14 With a few exceptions, it has not been possible to cultivate caliciviruses in Calchaqui virus (CQIV) A tentative spe- vitro, and this has hampered detailed cies in the genus Vesiculovirus . Isolated molecular studies of their replication. from mosquitoes, Aedeomyia squamipen- Exceptions are the feline calicivirus and nis , in Argentina. the murine norovirus. et al Am J Trop Med Hyg Calisher CH (1987) Carter MJ et al (1992) Virology 190 , 443 36 , 114 Cubitt WD (1989) Prog Med Virol 36 , 103 Jiang X et al (1991) Science 250 , 1580 calf diarrhea virus See neonatal calf Sosnovtsev SV et al (2006) J Virol 80 , 7816 diarrhea coronavirus , bovine rotavi- Stuart AD and Brown TD (2006) J Virol 80 , rus , and bovine noroviruses . 7500 calf lymph vaccine When vaccination Calicivirus Formerly a genus in both the against smallpox began in the 1880s, it family Picornaviridae and the family was produced on the skin of calves, and Caliciviridae . It has now been retired as was called calf lymph vaccine. This was a genus name. 73 calnexin

California encephalitis group Fourteen Table C1. California serogroup viruses viruses in the genus Orthobunyavirus which are antigenically and genetically California encephalitis virus AG83-497 * (C) related and form the species California California encephalitis virus BFS283 * (C) Inkoo virus * (C) encephalitis virus . See California sero- Jamestown Canyon virus (C) group viruses . Keystone virus (C, A, R) La Crosse virus * (C, H) California encephalitis virus (CEV) A Lumbo virus (C) Melao virus (C) species in the genus Orthobunyavirus San Angelo virus (A) belonging to the California encepha- Serra do Navio virus (C) litis group. Isolated from mosquitoes Snowshoe hare virus (L, C) in California, Utah, New Mexico and South River virus (A) Texas, USA. Has been associated with Tahyna virus * (C, A, H) a few cases of encephalitis in humans. Trivittatus virus (C, R) Antibodies to other group members cross-react so that presence of antibod- All may be regarded as subtypes or strains of ies does not prove presence of a spe- California encephalitis virus. Isolated from: A, anopheline mosquitoes; C, cific virus. culicine mosquitoes; H, humans; L, lemmings, hares; R, rodents. California harbor sealpox virus (SPV) * Can cause disease in humans. An unassigned virus in the family Poxviridae . cotton-top marmoset, Saguttus oedipus , California hare virus (CTFV-Ca) A strain in which it causes a silent infection. So of Colorado tick fever virus in the genus named because of its original isolation Coltivirus . from marmosets (callitrichid monkeys). Synonym : Herpesvirus sanguinis . Melendez LV et al (1970) In Infections and California rabbit fibroma virus See Immunosuppression in Subhuman Primates , Rabbit fibroma virus . edited by H Balner and WIB Beveridge. Copenhagen: Munksgaard, p. 111 California serogroup viruses Fourteen Melendez LV et al (1971) Lab Anim Sci 21 , 1050 serologically related viruses which form the species California encephalitis virus, in the genus Orthobunyavirus callitrichine herpesvirus 2 (CalHV-2) ( Table C1 ). An unassigned member of the family Herpesviridae . Isolated from the salivary Hubalek Z et al (1979) J Gen Virol 42 , 357 glands of white-lipped marmosets, Sanguinus fuscicollis . callitrichid hepatitis A highly fatal disease Synonym : marmoset cytomegalovirus. of captive marmosets and tamarins, Nigida SM et al (1979) Lab Anim Sci 29 , 53 caused by infection with the arenavirus lymphocytic choriomeningitis calnexin A 90 kDa integral calcium-binding virus (LCMV). The zoo animals become protein of the endoplasmic reticulum. infected by contact with LCMV-infected A molecular chaperone which binds to newborn mice (pinkies) provided as N-linked glycoproteins and assists in food for the primates. There is no evi- protein folding. Calnexin acts to retain dence to suggest that spread of the dis- unfolded or unassembled N-linked ease occurs between primates. glycoproteins, such as the influenza Montali RJ et al (1995) Am J Pathol 148 , 1441 virus hemagglutinin in the endoplasmic Stephensen CB et al (1995) J Virol 69 , 1349 reticulum, ensuring that only folded and assembled proteins proceed further callitrichine herpesvirus 1 (CalHV-1) An along the secretory pathway. unassigned member of the subfamily Chen W et al (1995) Proc Natl Acad Sci 92 , Gammaherpesvirinae. Isolated from the 6229 Calomys 74

Calomys A genus of mice living mainly Afonso CL et al (2002) Virology 295 , 1 in cornfields in South America, also Fenner F, Wittek R and Dumbell KR known as the vesper mouse. A host and (Editors) (1989) The Orthopoxviruses . San vector for several arenaviruses. Diego: Academic Press Gubser C and Smith GL (2002) J Gen Virol 83 , 855 Calovo virus Name given to a Czech- Kriz B (1982) J comp Path 92 , 1 oslovakian isolate from mosquitoes of a virus identical to Batai virus, a camostat A serine protease inhibitor. strain of Bunyamwera virus in the genus Orthobunyavirus . Lee MG et al (1996) Arch Virol 141 , 1979 calreticulin A major calcium-binding Camp Ripley virus A probable species protein in the endoplasmic reticulum in the genus Hantavirus , identified which acts together with calnexin as a from the Northern short-tailed shrew molecular chaperone responsible for (Blarina brevicauda ) in Minnesota, USA. quality control and folding of newly Arai S et al (2007) Emerg Inf Dis 13 , 1420 synthesized glycoproteins, such as the influenza virus hemagglutinin or her- Canadian vomiting and wasting disease pes simplex virus glycoprotein gB. of pigs virus Synonym for porcine Brodsky JL (2007) Biochem J 404 , 353 hemagglutinating encephalitis virus.

Camberwell virus A strain of Norwalk Cananéia virus (CNAV) A strain of Bertioga virus in the genus Norovirus , isolated virus in the genus Orthobunyavirus from an outbreak of gastroenteritis in belonging to the Guama serogroup. a hostel for the aged in Camberwell, Isolated from sentinel mice in Brazil. near Melbourne, Victoria, Australia. Calisher CH et al (1983) Am J Trop Med Hyg Genetically related most closely to 32 , 424 Lordsdale virus. Cauchi MR (1996) J Med Virol 49 , 70 Canarypox virus (CNPV) A species in the Marshall JA et al (2005) J Med Virol 75 , 321 genus Avipoxvirus. Similar to fowlpox virus. Causes a fatal infection in canar- camel contagious ecthyma virus ies, with pneumonia and exudate over Synonym for Auzduk disease virus. serous membranes. Sparrows are susceptible, chickens usually not. A vac- Camelpox virus (CMLV) A species in the cine strain is under development as a genus Orthopoxvirus . Causes pustules candidate human vector vaccine. The around the lips and nose of camels, vaccine virus, known as ALVAC, abor- and sometimes keratitis. Usually mild tively infects mammals with limited but may be severe, causing abortions expression of the vectored foreign pro- and up to 25% mortality, especially teins, but sufficient to elicit an immune in young animals. Incubation period response. Results with an ALVAC-HIV-1 4–15 days. Epidemics occur every 3–5 vaccine in infants are encouraging, and years in camels in the Middle East, protection of pigs from challenge with North Africa, Pakistan, and the former Nipah virus using an ALVAC-NIV Southern USSR. May cause lesions on glycoprotein has been reported. the hands of camel drivers. Lesions Evans TG et al (1999) J Infect Dis 180 , 290 produced on the CAM are very similar McFarland EJ et al (2006) AIDS 20 , 1481 to those caused by variola virus but in Taylor J et al (1994) Dev Biol Stand 82 , 131 cell cultures giant cells are produced. Weingartl HM et al (2006) J Virol 80 , 7929 Has a very limited host range. Genome sequence analysis shows that it is most Candiru complex Species in the genus closely related to variola (smallpox) Phlebovirus which were isolated from virus. The attenuated MVA strain of humans with febrile illness in the vaccinia virus can be used as an effec- Amazon region of Brazil, 1954–1994, tive vaccine. and are antigenically related to Candiru Synonym : photo-Shootur virus. virus. 75 canine calicivirus (CaCV)

Candiru virus (CDUV) A species in the canine adenoviruses 1 and 2 (CAdV- genus Phlebovirus . With Alenquer, 1 and 2) Two serotypes of Canine Itaituba, Morumbi, Nique, Oriximina, adenovirus , a species in the genus Serra Norte, and Turuna viruses Mastadenovirus . A natural infection of forms the Candiru antigenic complex. dogs, often silent, but in puppies there Isolated from humans with febrile ill- is often fever, vomiting, and diarrhea ness in Para, Brazil. Not found to repli- with up to 25% mortality. There is cuta- cate in mosquitoes experimentally. neous edema, ascites, hemorrhages into the viscera and hepatitis. Also Canid herpesvirus 1 (CaHV-1) A species a cause of laryngotracheitis (kennel in the genus Varicellovirus. There is cough). In foxes there is acute encepha- only one serotype, which has world- litis and hemorrhage into the brain. wide distribution. A natural infection Spread of infection is from the respira- in dogs, often silent but may cause tory tract and urine. Experimentally, necrotizing rhinitis and pneumonia, dogs and foxes may be infected by frequently fatal in newborn puppies. any route. Bears, coyotes, wolves, and The necropsy findings are predomi- raccoons are susceptible. Virus rep- nantly disseminated focal necroses lication with CPE occurs in cultures and hemorrhages in kidneys, liver, of dog, ferret, raccoon, and pig cells. lungs, spleen, thymus, and brain. May Hemagglutination is reported. A modi- cause tracheobronchitis (kennel cough) fied live vaccine, attenuated by passage in older animals but this condition in pig kidney cell cultures, produces can also be caused by an adenovirus. solid immunity following a single Replicates in primary dog kidney cell dose in dogs of any age. The complete cultures and all canine cell lines that DNA sequence of canine adenovirus have been tested with CPE. No CPE in 1 has been determined. There was lit- human, bovine, or porcine cell cultures. tle identity to human adenoviruses in Has been used as the basis of an anti- the early region genes, more in the late fertility vaccine for foxes. genes. Sequences of the early region Synonyms: canine herpesvirus; canine (E3) genes showed distinct sequences tracheobronchitis virus; kennel cough in canine adenovirus 2 as compared to virus. canine adenovirus 1. Appel MJ (1987) In Virus Infections of Synonyms : canine hepatitis virus; Carnivores , edited by MJ Appel. Amsterdam: canine laryngotracheitis virus; fox Elsevier, p. 3 encephalitis virus; hepatitis infectiosa Ronsse V et al (2005) Theriogenology 64 , 61 canis virus; infectious canine hepatitis Strive T et al (2006) Vaccine 24 , 980 virus; Rubarth’s disease virus.

Caninde virus (CANV) A serotype of Emery JB et al (1978) Am J Vet Res 39 , 1778 Koptopoulos G and Cornwell HJC (1981) Changuinola virus in the genus Orbi virus . Vet Bull ( Lond ) 51 , 135 Isolated from Lutzomyia sp in Para, Linne T (1992) Virus Res 23 , 119 Brazil. Morrison MD et al (1997) J Gen Virol 78 , 873 Schultz RD (2006) Vet Microbiol 117 , 75 canine AAV See Canine adeno-associated virus . canine astrovirus Particles resembling astroviruses have been seen by elec- Canine adeno-associated virus (CAAV) tron microscopy of dog feces by sev- A species in the genus Dependovirus . eral investigators, but the virus has not Antibodies are commonly present been further characterized. in dogs in Japan. No evidence of pathogenicity. canine calicivirus (CaCV) An unassigned Ishihara C and Yanagawa R (1975) Jpn J Vet species in the family Caliciviridae . Res 23 , 95 Isolated from the feces of a dog with diarrhea, and propagated in canine Canine adenovirus (CAdV) A species in kidney cells and in a dolphin cell line. the genus Mastadenovirus comprised of two serotypes. Schaffer FL et al (1985) Arch Virol 84 , 181 Canine coronavirus (CCoV) 76

Canine coronavirus (CCoV) A species in droplets and is highly infectious. The the genus Coronavirus . Causes vomit- virus is antigenically closely related to ing and diarrhea in dogs. Antigenically measles and rinderpest viruses. The related to feline infectious peritonitis roughly spherical virion is 100–250 nm virus and to porcine transmissible gas- in diameter. The helical nucleocapsid troenteritis virus, which can be trans- has a diameter of 15–18 nm. Buoyant mitted to dogs, but only CCoV causes density in CsCl: 1.20g/cm3 . Lacks disease. Replicates in a variety of neuraminidase. Chick and guinea pig canine and feline cell lines. Fecal mate- erythrocytes are agglutinated irregu- rial is the main source of infection, and larly. Can be readily isolated by co- disease can be prevented by avoiding cultivation of lymphocytes from infected contact with infected dogs and their animals with mitogen-stimulated excretions. Usually infection is self- canine lymphocytes. Once isolated, dis- limiting but a more pathogenic variant temper virus replicates in ferret, dog, has been described in Italy (the CB/05 and monkey (Vero) cell cultures and strain). This highly virulent strain can be adapted to eggs. Giant cell and causes a fatal disease in pups, and the cytoplasmic inclusions are produced. In sequences of the envelope, membrane 2000, an epidemic of canine distemper and nucleocapsid proteins of the virus occurred in Caspian seals (Phoca caspica ) are most closely related to those of por- in the Caspian Sea and more than 10,000 cine coronavirus causing transmissible seals were found dead. gastroenteritis of swine. Synonym : distemper virus. Synonym : gastroenteritis virus of dogs. Appel MJ (1987) In Virus Infections of Buonavoglia C et al (2006) Emerg Infect Dis Carnivores . Amsterdam: Elsevier, p. 133 12 , 492 Barrett, T (1999) In Encyclopedia of Virology , Cartwright S and Lucas M (1972) Vet Res 91 , Second edition, edited by A Granoff and 571 RG Webster. London: Academic Press, Decaro N et al (2007) Virus Res 125 , 54 p. 1559 Pratelli A et al (2000) J Virol Methods 84 , 91 Kennedy S et al (2000) Emerg Infect Dis 6 , 637 Tennant BJ et al (1993) Vet Rec 132 , 7 Kuiken T et al (2006) Vet Pathol 43 , 321 Montali RJ et al (1983) J Am Vet Med Assoc canine dermal papillomavirus Synonym 183 , 1163 for canine papillomavirus. canine hepatitis virus Synonym for Canine distemper virus (CDV) A species Canine adenovirus . in the genus Morbillivirus . A natural infection of dogs, foxes, wolves, ferrets, canine herpesvirus Synonym for Canid weasels, raccoons, and mink, with a herpesvirus 1 . worldwide distribution. Outbreaks of canine distemper have occurred in canine laryngotracheitis virus Synonym seals in Lake Baikal, Siberia in 1987 and for Canine adenovirus . the Caspian Sea in 2000. Exotic carnivores kept in zoos, such as red pan- Canine minute virus (CMV) A species in das, tigers, kinkajous, and binturongs, the genus Bocavirus. The dog is the only are also susceptible. The infection is known host. Isolated from dog feces in endemic in dogs and usually attacks the the Walter–Reed canine cell line dur- young, causing fever, nasal and ocular ing the 1960s. Causes CPE in this cell discharge, and sometimes skin erup- line but not in primary canine kidney tions, after an incubation period of 4–5 or thymus cell cultures, nor in cells of days. The fever subsides to rise again, human, simian, porcine, bovine, feline, this time with vomiting, diarrhea, and or murine origin. Agglutinates rhesus often pneumonia. The disease may be monkey erythrocytes at 5°C but not mild, especially in puppies, but signs guinea pig, human, rat, or pig eryth- of CNS involvement, particularly fits, rocytes. Antibodies are found in the may occur. The virus also causes hard- blood of many dogs. Can cause repro- pad, and probably canine rhinotonsil- ductive disease (abortion and infertil- litis. The disease is spread by airborne ity) in dogs, and may be associated 77 Cantagalo virus

with some cases of canine enteritis. These minimal changes allowed CPV to Antigenically distinct from H-1 virus, replicate systemically in dogs and to be latent rat virus and minute virus of shed from the intestine. Within 1 year mice. See also Canine parvovirus . of its recognition, CPV had spread from Synonyms: minute virus of canines; its probable origin in Europe globally MVC. to Australia, New Zealand, Japan, and Parrish CR (1990) Adv Virus Res 38 , 403 the USA. Subsequently, minor variants (0.2% nucleotide sequence changes) Canine oral papillomavirus (COPV) appeared in 1981, 1984, and 1990, and A species in the genus Lambda- in each case these also spread globally. papillomavirus. A natural infection of All types of domestic and wild dogs as dogs worldwide, causing papillomas well as cats are susceptible to infection. on the lips, which spread inside the After oronasal infection the virus rep- mouth. Has been detected in feral coy- licates in the tonsils and lymph nodes. otes. Does not infect mice, hamsters, or By 4–6 days the virus spreads to the guinea pigs experimentally. intestine, causing enteritis by destruction of regenerating intestinal epithelial Chambers VC (1960) Cancer Res 20 , 1083 cells. A diffuse myocarditis is occa- Delius H et al (1994) Virology 204 , 447 sionally seen in puppies. Both inacti- Teifke JP et al (1998) Vet Microbiol 60 , 119 vated and live attenuated vaccines are available. canine papillomavirus 2 A probable species in the family Papillomaviridae . A Carmichael LE (2005) Vet Med B 52 , 303 natural infection of dogs causing skin Ikeda Y et al (2000) Virology 278 , 13 Parrish CR (1990) Virus Res 38 , 403 papilloma. Distinct from canine oral Parrish CR (1994) Semin Virol 5 , 121 papillomavirus. Has been detected in immunocompromised dogs associ- canine respiratory coronavirus A prob- ated with cyclosporine treatment or able species in the genus Coronavirus . X-linked SCID. Isolated from respiratory samples of Synonym: canine dermal papilloma- dogs. Could be cultured in HRT-18 virus. cells. The genome sequence is similar Favrot C et al (2005) Am J Vet Res 66 , 1764 to that of bovine coronavirus, a corona- Goldschmidt MH et al (2006) J Virol 80 , 6621 virus group 2 species. Zaugg N et al (2005) Vet Dermatol 16 , 290 Erles K et al (2007) Virus Res 124 , 78 Canine parvovirus (CPV) A strain of Feline panleukopenia virus in the genus canine rhinotonsillitis A disease closely Parvovirus , closely related to mink related to canine distemper. enteritis virus and raccoon parvovirus, differing less than 2% by sequence canine tracheobronchitis virus Synonym analysis of the single-stranded DNA for Canid herpesvirus 1 . genome (5124 nt in length). Canine parvovirus type 2 (CPV-2) was first iso- Canoal virus A probable species in the lated or described in 1978. Serological genus Orbivirus isolated from phlebot- studies indicated that CPV-2 was a new omine sand flies in the Amazon region virus of dogs, as anti-CPV-2 antibod- of Brazil. Antigenically related to the ies could only be demonstrated in dog Changuinola virus group. Not reported sera collected after mid-1976. Termed to cause disease in humans. CPV-2 to distinguish it from the previously described, but unrelated, canine Caño Delgadito virus (CADV) A spe- minute virus (CMV). The name CPV- cies in the genus Hantavirus . Isolated 2 is no longer used. CPV is over 98% from the cotton rat, Sigmodon alstoni , identical in sequence to feline panleu- in Venezuela in 1995. Disease potential kopenia virus and it appears that the unknown. feline virus acquired a new host range in canines as a result of only two amino Cantagalo virus A strain of Vaccinia virus acid changes in the capsid protein VP2. in the genus Orthopoxvirus , isolated in Cantagalo virus 78

1999 from lesion samples taken during a rodent, Proechmys guyannensis oris , episodes of exanthem affecting dairy opossums, Caluromys philander , and cattle and human milkers in Cantagalo mosquitoes of Culex sp in Para, Brazil. county, state of Rio de Janeiro, Brazil. No association with disease in humans Sequence analysis suggests that it is a reported. derivative of vaccinia virus which has persisted some 20 years after smallpox capped terminus An unusual sequence vaccination ceased in Brazil. of methylated bases at the 5 - Damasco CR et al (2000) Virology 277 , 439 terminus of an mRNA molecule. The general structure is m 7 G5 ppp 5 Np … (7-methylguanine linked to the adja- Cao Bang virus A probable species in the cent nucleotide by a 5 -5 triphosphate genus Hantavirus , identified from the bridge). Such caps are present on many Chinese mole shrew (Anourosorex squa- viral and cellular mRNAs. The cap mipes ) in Vietnam. is involved in translation, but some Song J-W et al (2007) Emerg Infect Dis 13 , viruses, notably members of the fam- 1784 ily Picornaviridae, do not have capped termini on their mRNA molecules and cap The modified 5 end of eukaryotic instead there is a genome-linked pro- and most viral mRNAs. See capped tein (VPg) covalently linked to the 5 terminus . terminus. Viruses which have a virion transcriptase responsible for mRNA cap-binding protein Any of the proteins synthesis may contain a virion enzyme that specifically recognize the capped function which caps the mRNA, or an terminus of mRNA. Promote unwind- endonuclease which cleaves short- ing of the ribosome binding site in reg- capped oligonucleotides from host cell ulation of mRNA translation. mRNA and uses these to prime viral mRNA synthesis. Influenza virus uses Sonenberg N et al (1978) Proc Natl Acad Sci 75 , 4843 such a mechanism, referred to as ‘ cap snatching. ’ cap-snatching A 5 -cap structure is Clemens MJ (1979) Nature 279 , 673 essential for translation of most virus and cellular mRNAs, and many nega- capping See capped terminus . tive strand viruses have an enzyme in the virion which synthesizes a cap capping enzyme A viral enzyme which structure as the mRNA is synthe- adds a capped terminus to mRNA. sized. However two genera of viruses, Present in many viruses which contain Orthomyxovirus and Bunyavirus , acquire a virion transcriptase for mRNA syn- a cap structure on their mRNAs by thesis. Also specified in the genome stealing the 5 - 10–15 nt, including the DNA of viruses such as poxviruses. cap, from host cell mRNA as part of their replication cycle, and this process capravirine A non-nucleoside reverse is called cap-snatching, or cap-stealing. transcriptase inhibitor which is potent See capped terminus . in vitro and under development for clinical use. Cape Wrath virus (CWV) A serotype of Great Island virus in the genus Orbivirus , caprine adenovirus (GAdV) A tentative belonging to the Kemerovo serogroup. species in the genus Mastadenovirus , Isolated from a female tick, Ixodes uriae , also called goat adenovirus. Two types found under rocks at Cape Wrath, can be distinguished serologically. Scotland. Not reported to cause disease When inoculated into 3-week-old kids in humans. respiratory tract disease developed. Lehmkuhl HD and Cutlip RC (1999) Arch Capim virus (CAPV) A species in the Virol 144 , 1611 genus Orthobunyavirus. A member of Lehmkuhl HD et al (1997) Am J Vet Res 58 , the Capim serogroup. Isolated from 608 79 Capripoxvirus

Caprine arthritis encephalitis virus Caprine herpesvirus 1 (CpHV-1) A spe- (CAEV) A species in the genus Lenti- cies in the genus Varicellovirus . Isolated virus belonging to the ovine/caprine from kids, Capra hircus , with a severe lentivirus group. A non-oncogenic ret- generalized infection. Experimental rovirus causing a disease complex first infection of these animals produces described in 1974 that affects goats a severe febrile disease, and in adult of all ages and breeds. In young ani- pregnant goats there is fever and abor- mals, paresis, and paralysis result from tion. Not pathogenic for lambs or infection, but in older animals there is calves. The virus replicates in bovine, chronic persistent arthritis and masti- rabbit, and lamb cell cultures with CPE, tis. Concomitant bacterial and parasitic but not in HeLa, Vero, or chick embryo opportunistic infections are common. cells. It is antigenically related to, but The virus is widely disseminated in distinct from, bovine herpesvirus 1. goat herds in North America, Europe, Synonyms: goat herpesvirus; herpes- and Australia, especially in dairy goats virus caprae. where kids are commonly fed pooled Berrios PE et al (1975) Am J Vet Res 36 , 1755, colostrum and milk from dairy moth- 1763 ers which disseminates the infection. Engels M et al (1987) J Gen Virol 68 , 2019 The CAEV genome is 9.2 kb in length and exists in the virion as a dimer. The caprine respiratory syncytial virus Pos- primer for DNA synthesis is tRNA- sibly a strain of Bovine respiratory syn- Lys1,2 . In addition to gag , pol , and env cytial virus. Comparative studies of genes, the genome encodes three pro- bovine, caprine and human RSV in teins: Q (230 amino acids) which is an ovine kidney cell line showed the analogous to vif of HIV; tat (87 amino caprine and bovine viruses were most acids) which activates viral transcrip- closely related. tion; and rev (33 amino acids) which is Easton AJ et al (2004) Clin Microbiol Rev 17 , 390 involved in transportation of genomic Lehmkuhl HD et al (1980) Arch Virol 65 , 269 and mRNA to the cytoplasm. Goat Trudel M et al (1989) Arch Virol 107 , 141 synovial membrane cell cultures are permissive for replication. Although caprinized virus Virus adapted to goats. a common disease of goats (in most When rinderpest virus is adapted to countries surveyed, 80% of goats are goats it ceases to be virulent for cattle. infected), the virus has also been isolated from sheep with progressive Capripoxvirus A genus of the subfamily pneumonia. Arthritic disease is seen Chordopoxvirinae . Viruses of ungulates. frequently in adult goats, and is pro- The virions (300 270 200 nm) are gressive with gradual swelling of the longer and narrower than vaccinia knee joints. Goat kids are more likely virions. Infectivity is sensitive to ether to develop encephalomyelitis with and trypsin. Species show serologi- posterior paresis, occasionally leading cal cross-reactivity. They produce no to limb paralysis. When affected kids hemagglutinin. Genome DNA is about recover they go on to develop severe 145 kbp in length. Mechanical trans- arthritis as adults. In dairy goats masti- mission by arthropods occurs. Type tis is also common. Control is by sepa- species is Sheeppox virus. Other species rating kids from animals immediately are Goatpox virus and Lumpy skin disease after birth to prevent colostrum trans- virus . Capripox of sheep and goats is mission. No vaccines are available. enzootic in Africa north of the equa- Natural transmission from goats to tor, the Middle East and Turkey, Iran, sheep can occur in mixed flocks. Afghanistan, Pakistan, India, Nepal, and parts of China. There are sporadic Germain K and Valas S (2006) Virus Res 120 , outbreaks in Europe. Control is by 156 slaughter of infected animals. Narayan O and Cork LC (1990) In Virus Synonym : sheeppox subgroup viruses. Infections of Ruminants, edited by Z Dinter and B Morein. Amsterdam: Elsevier, p. 441 Bhanuprakash V et al (2006) Comp Immunol Pisoni G et al (2005) Virology 339 , 147 Microbiol Infect Dis 29 , 27 Ravazzolo AP et al (2006) Virology 350 , 116 Gershon PD et al (1989) J Virol 63 , 4703 capsid 80 capsid A protein shell which surrounds antigenically to the C group viruses. the virus nucleic acid and its associated Mosquito-borne. Isolated from a sen- protein (the nucleoprotein core). The tinel Cebus monkey and mice in Para, capsid usually has icosahedral symme- Brazil. Also isolated from bats. Has try but in some cases is helical. Capsid been associated with a febrile illness and core together form the nucleocap- in humans. Found in Brazil, Panama, sid. See capsomeres . Trinidad, French Guiana, and Surinam. Green J et al (2000) Virus Genes 20 , 227 carbovir Carbocyclic 2 ,3 -didehydro- capsid polypeptide Protein-forming 2 ,3 -dideoxyguanosine. A potent part of the capsid structure of a virus inhibitor of human immunodeficiency particle. virus replication in vitro . Carter SG et al (1990) Antimicrob Agents capsomeres Units from which the capsid Chemother 34 , 1297 is built, visible in the electron microscope, and consisting of groups of iden- carboxymethylcellulose A cellulose tical protein molecules (protomers). In derivative which is used for the sepa- icosahedral capsids, the capsomeres ration of proteins by ion exchange at the 12 corners are called ‘ pentons ’ chromatography. because they have five neighboring capsomeres. All other capsomeres have carcinoembryonic antigen The cell recep- six neighbors and are called ‘ hexons. ’ tor for murine hepatitis virus. Each penton contains five protomers, each hexon three or six. Many animal Cardiovirus A genus of the fam- viruses have their capsomeres arranged ily Picornaviridae . Consists of: (1) in icosahedral symmetry. Encephalomyocarditis virus (strains include Columbia SK virus, mengo virus, and capuchin herpesvirus AL-5 Synonym for Maus Elberfeld virus); and (2) Theilovirus cebine herpesvirus 1. (strains include Theiler’s murine encephalomyelitis virus, Vilyuisk human capuchin herpesvirus AP-18 Synonym encephalomyelitis virus, Saffold virus, for cebine herpesvirus 2. and Theiler-like virus of rats). Differ from enteroviruses in losing infectivity below CAR C oxsackie a denovirus r eceptor. pH 4 instead of pH 3 and being unsta- A cellular receptor shared by group ble at pH 6 in the presence of chloride or B coxsackie virus and adenovirus. A bromide ions. The viruses have a poly C member of the immunoglobulin pro- tract (80–250 bases) about 150 bases from tein superfamily. the 5 terminus of the genome. Martino TA et al (2000) Virology 271 , 99 Cooper PD et al (1978) Intervirology 10 , 165 Paul S and Michiels T (2006) J Gen Virol 87 , 1237 Caraipe virus A probable species in the Tracy S et al (2006) Curr Top Microbiol genus Orthobunyavirus, isolated from Immunol 299 , 193 anopheline mosquitoes in the Amazon region of Brazil. Not associated with Carey Island virus (CIV) A species in the disease in humans. genus Flavivirus . A member of the Rio Bravo virus group. Isolated from bats Carajas virus (CJSV) A species in the in Malaysia. Not reported to cause dis- genus Vesiculovirus . Isolated in Brazil ease in humans. in 1983 from sand flies, Lutzomyia sp. Travassos da Rosa AP et al (1984) Am J Trop Carlow virus A probable species in the Med Hyg 33 , 999 genus Norovirus. Isolated from an outbreak of gastroenteritis in Ireland in Caraparu virus (CARV) A species in 2002. the genus Orthobunyavirus , belonging Kearney K et al (2007) Virol J 4 , 61 81 cat fever virus carnivorepox virus A strain of Cowpox protease inhibitor drugs that has sig- virus which differs from the reference nificantly reduced morbidity and mor- strain (Brighton) by a low ceiling tem- tality associated with HIV-1 infection. perature for pock development on the Palella FJ et al (1998) N Engl J Med 338 , 853 CAM. Caused an epizootic among car- Palella FJ et al (2006) J Acquir Imm Defic Syndr nivores in Moscow Zoo. 43 , 27 Marennikova SS et al (1978) Arch Virol 56 , 7 case–control studies In an epidemiologi- carp pox herpesvirus Synonym for cal investigation to determine factors cyprinid herpesvirus 1. responsible for a disease outbreak, both case–control and cohort studies carp virus A transmissible virus obtained are used. In case–control studies, it is from cases of human multiple sclerosis. necessary first to decide on a case defi- Depresses the number of circulating nition, then persons meeting the case polymorphonuclear neutrophils within definition are compared with controls 16–48 h of inoculation into adult mice. who do not have the disease and are The effect lasts at least 11 months dur- judged comparable in age and other ing which time the mice remain normal. characteristics. Past exposures or risk The virus passed through membranes factors are then examined to determine of 50 nm average pore diameter but if they are associated with the disease. not of 25 nm. Replicates in PAM cells, Schlesselman JJ (1982) Case Control Studies: a line of mouse fibroblasts. The effect Design , Conduct , Analysis. New York: Oxford in mice was neutralized by serum University Press. from patients with multiple sclerosis. However, the mouse test has proved caspase A protease, specific for cysteinyl- difficult to reproduce and is unreliable. aspartate residues, which is involved It is possible that these results are an in cell death and apoptosis. artifact and the virus does not exist. Carp RI et al (1977) Lancet ii , 814 caspase inhibition Several viruses Carp RI et al (1978) Prog Med Virol 24 , 158 encode proteins designed to inhibit caspase activity and so evade apop- carrier cultures A type of persistent infec- tosis. Examples are the CrmA protein tion of cell cultures in which only a encoded by several poxviruses, the small proportion of the cell population E3-14.7K protein of adenoviruses, is infected. These cells release virus and an inhibitor of apoptosis protein and are killed, but the released virus encoded by African swine fever virus. infects a small number of other cells. A Castelo dos Sanhos virus (CDSV) carrier culture can be ‘ cured ’ of virus A probable species in the genus infection by adding antiviral antibody. Hantavirus, isolated from a suspected Many different viruses can cause car- case of hantavirus pulmonary syn- rier infections. Carrier cultures may drome which occurred in 1995 in arise because most of the cells in it Castelo dos Sanhos, Mato Grosso State, are genetically resistant to the infect- Brazil. The presumed rodent host of ing virus, or because of the presence of this virus is unknown. weak antibody or interferon in the cell culture medium. Johnson AM et al (1999) J med Virol 59 , 527 Castleman’s disease A rare B-cell lym- carrier state The condition in which an ani- phoproliferative disorder which has mal is persistently infected with a virus, been associated with human herpes often without showing the signs of the virus 8 infection. disease associated with the virus. The virus can persist even in the presence of cat distemper virus Synonym for Feline the specific neutralizing antibody. panleukopenia virus .

CART regime C ombination a nti -r etro- cat fever virus Synonym for Feline panleu- viral therapy. A regime including three kopenia virus . cat flu virus 82 cat flu virus Synonym for Feline mine sand flies in the Amazon region calicivirus . of Brazil. Not associated with disease in humans. cat leukemia virus See Feline leukemia virus . catfish reovirus See channel catfish reovirus . cat plague virus Synonym for Feline panleukopenia virus . cationic detergent A surface-active agent having positively charged surface ions. cat sarcoma virus See Feline leukemia virus . cattle plague virus Synonym for Rinderpest virus . cat-scratch disease virus A disease once thought to be due to a virus, but actu- Catu virus (CATUV) A species in the genus ally a rickettsial infection, caused by Orthobunyavirus , antigenically related Bartonella henselae . It is a mild disease, to the Guama serogroup. Originally with fever, malaise, and a local lesion isolated from a young man with a which becomes a pustule, and with febrile illness in Para, Brazil; 10 further lymphadenitis in the draining lymph isolations have been reported. Also iso- nodes. A skin test has been used to lated from bats, mosquitoes, sentinel confirm the diagnosis, using pus from monkeys and mice in Brazil, Trinidad, a case as antigen. and French Guiana. Synonyms: benign inoculation lymphoreticulosis virus; non-bacterial regional lymphadenitis virus. Caudovirales An order consisting of three families of tailed bacterial viruses infecting Bacteria and Archaea. They cat type C oncovirus Synonym for Feline are: Myoviridae (long contractile tails), leukemia virus and feline sarcoma virus. Siphoviridae (long non-contractile tails), and Podoviridae (short non-contractile Catacamas virus A probable species tails). in the genus Hantavirus, isolated in Catacamas, in eastern Honduras from Coue’s rice rats (Oryzomys couesi ). Some cauliflower disease of eels virus A virus 20% of 24 rats examined had antibodies (EV2) isolated from a European eel, to the virus. Catacamas virus is geneti- Anguilla anguilla , with stomatopapil- cally most closed related to Bayou virus , lomas. Eels develop lesions with a which is found in marsh rats (Oryzomys cauliflower-like appearance, predomi- palustris ). nantly around the mouth and head and consisting of hyperplastic squamous Milazzo ML et al (2006) Am J Trop Med 75 , cells. An icosahedral virus has been 1003 isolated from the blood of affected eels, but the relationship of this virus to EV2, Catarina virus A probable species in the and of both to the disease is not clear. genus Arenavirus found in the Texas EV2 virus is 90–140 nm in diameter and southern plains woodrat (Neotoma pleomorphic, with surface projections micropus ). Genetically related to 10 nm long. Concentrated preparations Whitewater Arroyo virus, which is agglutinate chick and sheep erythro- associated with the white-throated cytes. Spontaneous elution occurs at woodrat (Neotoma albigula ). room temperature. Replicates in FHM Cajimat MN et al (2007) Am J Trop Med Hyg cells, optimally at 15°C, producing syn- 77 , 732 cytia and irregular masses of rounded cells which eventually lyse. Infectivity catarrhal jaundice virus Synonym for is ether-sensitive and does not survive Hepatitis A virus . much over 3 weeks at 4°C. Replication inhibited by actinomycin D but not Catete virus A possible species in the idoxuridine. See eel virus American . genus Orbivirus, isolated from phleboto- Synonym : EV2 virus. 83 CD

Nagabayashi T and Wolf K (1979) J Virol 30 , injection grow into tumors and contain 358 viral antigen. The virus is not neutral- Pfitzner I and Schubert G (1969) Z Naturforsch ized by caviid herpesvirus 1 antiserum. B 24 , 790 Synonyms: guinea pig herpesvirus; guinea pig cytomegalovirus; guinea caveolae Bottle-shaped invaginations pig salivary gland virus; guinea pig of the plasma membrane of cells. leukemia virus. Some viruses such as SV40 virus and echovirus 1 enter cells via a caveolae- Rhim JS (1977) Virology 82 , 100 mediated endocytic pathway which delivers the virus to an intermedi- caviid herpesvirus 3 (CavHV-3) An unas- ary compartment, the caveosome. The signed virus in the family Herpesviridae . caveosome acts as a sorting organelle Originally isolated from the buffy coat for delivery of the virus to the endo- of healthy guinea pigs and designated plasmic reticulum. GPXV. Produced CPE in guinea pig embryo fibroblast cells. The viral DNA Norkinm LC and Kuksin D (2005) Virol J 2 , could be distinguished from other cav- 38 Pietiainen VM et al (2005) Ann Med 37 , 394 iid herpesvirus DNAs by restriction enzyme analysis. caveosome See caveolae . Synonym : guinea pig herpesvirus 3. Bia FJ et al (1980) J Virol 36 , 245 caviid herpesvirus 1 (CavHV-1) An unassigned virus in the family Herpesviridae . CbaAr426 virus (CAV) An isolate from A silent infection of guinea pigs; inclu- mosquitoes of Bunyamwera virus in the sion bodies can be found in cells of the genus Orthobunyavirus belonging to the salivary gland ducts. The infection can Bunyamwera serogroup. be passed experimentally by peripheral injection of salivary gland extract. CC chemokines A group of chemokines Inoculation i.c. causes fatal meningitis. (also known as beta-chemokines) in Intratracheal inoculation causes pneu- which there are two adjacent cysteine monia. There is replication in primary residues. They have chemoattractant guinea pig fibroblast culture with the and other activating effects on mono- appearance of small foci of enlarged cytes and macrophages. Some also cells in 10 days. Nuclear inclusions stimulate T lymphocytes, eosiniphil appear and on passage CPE may be leukocytes, and basophil leukocytes. seen in 1–2 days. The virus is cell associated. CCL Abbreviation for c ertified c ell l ine, Synonyms : guinea pig herpesvirus 1; registered with the American Type Hsiung–Kaplow herpesvirus. Culture Collection (ATCC). See certi- Fong CKY et al (1979) J Gen Virol 42 , 127 fied cell line . Smith MG (1959) Prog Med Virol 2 , 171 CCR5 A chemokine receptor expressed on caviid herpesvirus 2 (CavHV-2) An the surface of macrophage cells and T- unassigned virus in the subfamily cell lymphocytes which is used as a co- Betaherpesvirinae. Isolated from cultures receptor (with CD4) for HIV, and binds of strain 2 guinea pig cells which devel- cell signaling molecules called beta- oped focal areas of swollen rounded chemokines (e.g. RANTES, MIP-1α , cells 9–13 days after preparation. No and MIP-1β ). This interaction results in virus was obtained from homogenized activation and movement of immune guinea pig tissue extracts. Not reported cells to sites of infection, leading to to cause disease in guinea pigs or other enhanced clearance of pathogens. animals. Replicates in rabbit kidney cell, mink lung cell, and Vero cell cul- CCRSF Complement component recep- tures, but not in monkey, hamster, tor superfamily. human, or fowl CMA cells. Infective virus in rat cells does not kill them all CD Cluster of d ifferentiation. An inter- and transformed cells appear which on nationally agreed system of naming CD 84

differentiation antigens on cell surfaces, vaccine and adapted measles virus based on their reaction with panels of strains, but not wild-type strains. monoclonal antibodies. Several CD Naniche D et al (1993) J Virol 67 , 6025 antigens act as relatively specific receptors for virus attachment, e.g. CD4 is a CD49 The fibronectin receptor. A series receptor for human immunodeficiency of type 1 transmembrane proteins virus ; CD21 is a receptor for Epstein– (CD 49a–f) which are members of the Barr virus; CD46 is a receptor for mea- integrin superfamily. CD 49e is a recep- sles virus; and CD155 is a receptor for tor for vaccine strains of measles virus. poliovirus.

؉ CD56 A transmembrane protein of the Ig CD4 T lymphocytes which contain the superfamily which, together with CD CD4 antigen on their surface. CD4 is 16, is present on the surface of natu- a type 1 transmembrane protein of the ral killer (NK) cells. It is an isoform of Ig superfamily. It has four extracellular N-CAM ( n eural c ell a dhesion m olecule) immunoglobulin-like domains. Found found in the CNS. on about 60% of T lymphocytes in human blood. CD62 A family of type 1 transmembrane proteins on the surface of leukocytes CD4 cells T lymphocytes which are effec- which contain an N-terminal C-type tors of the specific cellular immune lectin domain which can bind to car- response. Class II MHC restricted, the bohydrates. Known as selectins, they CD4 antigen is a receptor for binding mediate the adhesion of leukocytes to HIV-1, and the onset of AIDS is asso- vascular endothelium, including the ciated with marked depletion of CD4 brain capillary endothelium. cells in the blood. Dalgleish AG et al (1984) Nature 312 , 776 CD81 A tetraspanning membrane protein which is widely distributed on a vari- CD8 cells T lymphocytes which are effec- ety of cell types, and has been impli- tors of the specific cellular immune cated as the receptor for hepatitis C response. Class I MHC restricted. virus. Zhang J et al (2004) J Virol 78 , 1448 ؉ CD8 T lymphocytes which contain CD8 on their surface. CD8 is a heterodimer CD150 The signaling lymphocyte acti- consisting of an alpha and a beta chain, vation molecule (SLAM). A CD2-like both type 1 transmembrane proteins of molecule of the Ig superfamily, which the Ig superfamily. Found on thymo- acts as the receptor for the H protein of cytes, and on 15–25% of mature T lym- all strains of measles virus. phocytes, which are CD4 negative. Erlenhoefer C et al (2001) J Virol 75 , 4499 ؉ CD34 Hemopoietic precursor cells cDNA Abbreviation for complementary which contain CD34, a heavily glyco- DNA; formed by reverse transcription sylated protein on their surface. Serves of RNA. as a receptor for simian foamy virus. cDNA cloning Molecular cloning of a CD46 A membrane cofactor protein that double-stranded DNA copy of an RNA is on the surface of most leukocytes molecule. and other nucleated cells, but not on erythrocytes. Binds complement cDNA library A series of clones which components C3b and C4b, prevent- represent all the mRNA sequences ing their deposition and formation of expressed in the cell from which they an attack protein on the cell surface. are derived. Only contains transcribed Downregulation of CD46 therefore DNA sequences (exons), and no non- enhances complement-mediated cell transcribed sequences (introns, spacer lysis. The first measles virus receptor DNA). Obtained by synthesizing a protein to be identified, it binds mostly cDNA copy of the mRNA population, 85 cell fusion

converting this to double-stranded cell culture Growth of dispersed cells DNA, and integrating this DNA into in vitro as single layers (monolayers) the restriction site of a cloning vector. on a glass or plastic surface, or as sus- If an expression vector is used (e.g. pensions, in liquid or soft gel medium. lambda gt11), specific mRNAs may be A culture derived from cells taken identified using antibodies which react directly from the tissue of origin is with particular expressed proteins. called a ‘ primary culture.’ The first subculture of a primary culture may give cebine herpesvirus 1 (CbHV-1) An rise to a cell line. See tissue culture . unassigned species in the family Herpesviridae, isolated from a captive cell cycle Cytological study identifies capuchin monkey, Cebus appella . only two phases in the cell growth Synonyms: capuchin herpesvirus AL-5; cycle: (1) mitosis, during which the owl monkey herpesvirus. chromosomes become visible and Lewis MA et al (1976) Infect Immun 14 , 759 undergo redistribution within the cell which then divides; and (2) interphase , which occupies the majority of the cebine herpesvirus 2 (CbHV-2) An cell growth cycle. Biochemical analy- unassigned species in the family sis has resulted in the subdivision of Herpesviridae , isolated from a captive interphase into three phases charac- Cebus appella capuchin monkey, . terized by their metabolic activity, the Synonyms : capuchin herpesvirus AP-18; most prominent of which is the period owl monkey herpesvirus. of DNA synthesis (S), usually lasting Sabin AB et al (1934) J Exp Med 59 , 115 6–8 h, which occurs in the middle of interphase. The gap between mitosis cell-associated virus Virus particles (M) and the S phase is known as G1, which remain attached to or inside the and the second, between S and M, is host cell after replication. The amount known as G2. Thus the cell cycle occurs of cell-associated compared to released as M–G1–S–G2–M, etc. Most viruses virus varies considerably with the are able to multiply in cells independ- virus and/or the host cell involved. ently of cell division or the stage in the Cytomegaloviruses are a typical exam- cell cycle at which infection occurs. ple of a strongly cell-associated virus. However, the Parvoviridae are an exception and require rapidly dividing cells cell attachment The first step in virus-cell for the establishment of infection. interaction is a specific non-covalent binding between attachment sites on cell death Programmed cell death is a the virion surface and receptors on physiological process that induces the plasma membrane of the host cell. apoptosis, a form of cellular suicide. All vertebrate viruses carry multiple Apoptosis enables unwanted cells such copies of their attachment proteins or as virus-infected cells to be eliminated, sites. For example small picornaviruses and rapid cell death reduces virus carry 60 receptor-binding sites per spread. virion, and influenza virus has several Dix MM et al (2008) Cell 134 , 679 hundred. cell-free translation An in vitro system cell counts The number of viable cells in for translating added viral or cellular a culture can be determined using a mRNAs into proteins. The most com- vital stain, which is excluded by living monly used systems are derived from cell membranes and is incorporated wheat germ, rabbit reticulocytes or only into dead cells. Trypan blue is the ascites tumor cells. most commonly used vital stain for this procedure. cell fusion Formation of multinucleate George VG et al (1996) In Virology Methods giant cells (syncytia) by fusion of cell Manual , edited by BWJ Mahy and HO membranes. Enveloped viruses have Kangro. London: Academic Press, p. 3 a membrane glycoprotein with fusion cell fusion 86

activity. Virus-induced fusion from out- peptides fuse with MHC class II mol- side the cell can be caused by exposure ecules and are presented in association to high multiplicity of virus (e.g. Sendai with the MHC class II molecules on the virus), even when the virus nucleic cell surface. The complex so presented acid has been inactivated. Such fusion is recognized by the T-cell receptor and can be mediated by the fusion protein CD4 molecule on CD4T cells resulting alone, acting at the cell surface. Fusion in clonal expansion, and production of within the cell occurs with several growth factors that induce prolifera- enveloped viruses such as influenza, tion and differentiation of both T and Semliki Forest virus, and vesicular B lymphocytes. Cytotoxic CD8 T lym- stomatitis virus, which are taken up phocytes recognize a complex of MHC into endosomal vesicles by receptor- class I with a foreign peptide on the mediated endocytosis. Fusion between surface of infected cells and destroy the viral and endosomal membranes is them. B lymphocytes are stimulated to triggered by a reduction in pH within produce humoral antibodies to neutral- the endosome. ize free virions, prevent spread of the infection and provide subsequent pro- cell-fusing agent virus (CFAV) A tenta- tection from reinfection. tive species in the genus Flavivirus isolated from Aedes aegypti cells that may cell transformation See transformation . be the present-day survivor of the pri- meval Flavivirus . cellular receptors A wide variety of cell surface molecules are used as recep- cell line A population of cells arising tors by different viruses. These vary from a primary cell culture at the time from glycoproteins such as sialic acid of the first subculture. They are of two or heparan sulfate to specific proteins types, euploid (e.g. diploid) or aneu- on the cell surface such as integrins, or ploid (heteroploid). Euploid cells, e.g. molecular immunoglobulins. normal human fibroblasts, retain their normal karyotype throughout their CELO virus An acronym for c hicken culture lifespan, rarely give rise to con- e mbryo l ethal orphan virus. An avian tinuous cell lines, and stop dividing adenovirus. after about 50 generations. Aneuploid cells, e.g. tumor derived cells, may give rise to continuous cell lines that can central dogma The idea originally probe cultured indefinitely, but have an posed by Crick FHC (1958, Symp Soc abnormal karyotype. A cloned cell line Exp Biol 12 , 137) that genetic informa- is descended from a single cell clone. tion can be perpetuated in the form of A diploid cell line is one where at least nucleic acid, but cannot be retrieved 75% of cells have the same karyotype from the amino acid sequences of pro- as normal cells of the species from teins. Prions appear to challenge this which the line was derived. A hetero- dogma. ploid cell line has less than 75% of cells which are of normal karyotype. Crick FHC (1970) Nature 227 , 561 cell-mediated immunity Immunity Central European encephalitis virus mediated by cells of the T-lymphocyte (CEE) The former name for Tick- lineage, distinguished from humoral borne encephalitis virus. See also Tick- immunity which results from secre- borne encephalitis virus (European tion of immunoglobulins by B lym- subtypes). phocytes. T and B lymphocytes form an integrated system to generate a spe- central nervous system viral infections cific response to virus infection. The Many viruses such as rabies, measles, initial stages of induction of immunity mumps and herpes simplex may infect involve uptake of antigen (virus or the nervous system and cause serious virus proteins) by dendritic cells, where disease. Viruses may enter the nervous proteolysis occurs and antigen-specific system along nerves and through the 87 Cercopithecine herpesvirus 9 (CeHV-9)

blood. Once in the central nervous sys- Genome DNA is 67% G C. Partial tem viruses may cause acute or slow genome sequences of several genes chronic infections. are related to their homologs in her- Johnson RT (2003) J Neurovirol 9 , 140 pes simplex virus. Infectivity is neutralized by antisera to herpesvirus B Central Plata virus A probable species in (Cercopithecine herpesvirus 1 ). the genus Hantavirus , identified from Synonym : SA8 virus. the rodent Oligoryzomys flavescens in Eberle R et al (1993) Arch Virol 130 , 391 Uruguay. Has been associated with Hilliard J et al (1989) Arch Virol 109 , 83 hantavirus pulmonary syndrome. cercopithecine herpesvirus 3 (CeHV-3) centrifugation-enhanced culture Detec- An unassigned species in the family tion of a cytopathic effect in a culture Herpesviridae. Isolated from vervet of cells may require days or weeks. monkey, Cercopithecus aethiops , kidneys. For some viruses this process can be Genome DNA is 51% G C . speeded up by centrifuging the culture. Synonym : SA6 virus. Faisst S (1999) In Encyclopedia of Virology , Malherbe H and Harwin RS (1963) S Afr Second edition, edited by A Granoff and RG Med J 37 , 407 Webster. London: Academic Press, p. 1408 centrophyten Minute basophilic particles cercopithecine herpesvirus 4 (CeHV-4) reported by Lipschütz (1928, Wien Klin An unassigned species in the family Wschr 41, 365) to be present in epider- Herpesviridae . Isolated from a vervet mal cells infected with measles virus. monkey, Cercopithecus aethiops . Synonym : SA15 virus. Cercopithecine herpesvirus 1 (CeHV-1) Malherbe H and Harwin RS (1963) S Afr A species in the genus Simplexvirus . A Med J 37 , 407 natural infection of Asiatic macaque monkeys. 10% of newly caught rhe- Cercopithecine herpesvirus 5 (CeHV-5) A sus monkeys have antibodies, and the species in the genus Cytomegalovirus . virus is frequently present in kidney Isolated from vervet monkey, cell cultures of this animal. Reservoir Cercopithecus aethiops pygerythrus , kid- species include Macaca mulatta , ney and salivary gland cell cultures. M fascicularis , M fuscata , M arctoides , Focal lesions were observed in mon- M cyclopsis, and M radiata . Causes olayer cultures. Giant cells and eosi- vesicular lesions on the tongue and nophilic inclusion bodies were seen. lips, and sometimes of the skin. Synonyms: African green monkey cyto- Infection of humans by monkey bites megalovirus; macaque monkey virus. or other means leads to ascending Black PH et al (1963) Proc Soc Exp Biol Med myelitis or acute encephalitis; almost 112 , 601 all cases are fatal. Mice under 3 weeks old, day-old chicks and guinea pigs can be infected experimentally. Not all Cercopithecine herpesvirus 8 (CeHV-8) A strains are antigenically identical. species in the genus Cytomegalovirus . Synonyms: B-virus of monkeys; herpes- Causes persistent infection of rhesus virus simiae. monkeys, Macaca mulatta . Genome DNA is 52% G C . Holmes GP et al (1995) Clin Infect Dis 20 , 421 Synonym: rhesus monkey cytome- Ostrowski SR et al (1998) Emerg Infect Dis 4 , galovirus. 117 Perelygina L et al (2003) J Virol 77 , 6167 Smith AL et al (1998) J Virol 72 , 9224 Cercopithecine herpesvirus 9 (CeHV-9) Weigler BJ (1992) Clin Infect Dis 14 , 555 A species in the genus Varicellovirus . Causes mild exanthematous disease Cercopithecine herpesvirus 2 (CeHV-2) in captive rhesus monkeys, Macaca A species in the genus Simplexvirus . mulatta . Isolated from vervet monkey, Synonyms : Medical Lake macaque her- Cercopithecus aethiops , kidney cultures. pesvirus; simian varicella herpesvirus, Cercopithecine herpesvirus 9 (CeHV-9) 88

Liverpool vervet herpesvirus, Patas Synonyms: baboon herpesvirus; baboon monkey herpesvirus. lymphotropic herpesvirus; herpesvirus Blakely GA et al (1973) J Infect Dis 127 , 617 papio; papio Epstein–Barr herpesvirus. Clarkson MI et al (1976) Arch Ges Virusforsch Falk L et al (1976) Int J Cancer 18 , 798 22 , 219 Heller M and Kieff ED (1981) J Virol 37 , 698 Gray WL and Gusick NJ (1996) Virology 224 , Meseda CA et al (2000) J Gen Virol 81 , 1801 161 cercopithecine herpesvirus 13 (CeHV-13) cercopithecine herpesvirus 10 (CeHV-10) An unassigned virus in the family An unassigned species in the family Herpesviridae, isolated from captive Herpesviridae. Appears to be a com- macaque monkeys, Macaca cyclopis . mon infection of rhesus monkeys but Synonym : herpesvirus cyclopis. there is no evidence that it causes disease in them. Isolated by co-cultivation Cercopithecine herpesvirus 14 (CeHV-14) A of rhesus blood leukocytes with sim- species in the genus Lymphocryptovirus . ian or human fibroblasts such as Wl- Synonym : African green monkey EBV- 38 or MRC-5 cells. Causes CPE within like virus. 6–8 days, which slowly progresses. On staining with acridine orange, mul- Bocker JF et al (1980) Virology 101 , 291 tiple green inclusions are seen in the nucleus but not in the cytoplasm. More Cercopithecine herpesvirus 15 (CeHV-15) A than half the virus infectivity is cell- species in the genus Lymphocryptovirus . associated. Experimental inoculation Isolated from a malignant lymphoma of mice, hamsters and rabbits caused of a captive rhesus monkey, Macaca no obvious disease. Infection of rhesus mulatta . monkeys resulted in a rise in antibod- Synonym: rhesus EBV-like herpesvirus; ies. No cross-neutralization could be rhesus lymphocryptovirus. demonstrated with a range of other Rangan SRS et al (1986) Int J Cancer 38 , 425 human and simian herpesviruses. There appear to be at least two anti- Cercopithecine herpesvirus 16 (CeHV-16) genic types. A species in the genus Simplexvirus , Synonym : rhesus leukocyte-associated isolated from a baboon. herpesvirus, strain 1. Synonym : Herpesvirus papio 2 Frank AL et al (1973) J Infect Dis 128 , 618, 630 Eberle R et al (1995) Arch Virol 140 , 529

Cercopithecine herpesvirus 12 (CeHV-12) Cercopithecine herpesvirus 17 ( CeHV-17) A species in the genus Lympho- A species in the genus Rhadinovirus , iso- cryptovirus . Isolated from cell lines lated from rhesus monkeys. Genetically established from splenic lymphocytes related to human herpesvirus 8. of a baboon, Papio hamadryas , by co-cul- Synonym: Rhesus monkey rhadino- tivation with X-irradiated marmoset or virus. baboon lymphoblastoid cell cultures. The baboon had been injected with Desrosiers RC et al (1997) J Virol 71 , 9764 blood from another baboon which had been injected with human blood and cercopithecine herpesvirus SA8 Synonym had a lymphoma. The cell lines have B for Cercopithecine herpesvirus 2 . lymphocyte characteristics and carry a virus related to, but not identical with, certified cell line A line of cells certi- Epstein–Barr virus as demonstrated by fied by the American Cell Culture cross-reactivity of viral capsid antigens, Collection Committee. Each cell line is and conservation of many of the EBV assigned an accession number, e.g. CCL latency-associated genes, but only par- 2 HeLa, and derivatives are given a tial (40%) DNA homology. It can trans- decimal notation, e.g. CCL 2.1 HeLa form other nonhuman primate cells 229. Information on cell lines can be in vitro but its relationship to disease obtained from the American Type in the baboon is uncertain. Culture Collection, 10801 University 89 Ch1ES(NBL-8) cells (CCL 73)

Boulevard, Manassas, VA 20110-2209, sea were found to be dying from an USA. infection termed dolphin morbillivirus. Two years previously a closely related Cervid herpesvirus 1 (CvHV-1) A spe- virus had been isolated from porpoises cies in the genus Varicellovirus , iso- (Phocoena phocoena). Serological evi- lated from British red deer with ocular dence of infection with this virus was infection. subsequently found in many other spe- Synonym : red deer herpesvirus. cies in the Atlantic ocean, and the name cetacean morbillivirus was adopted. Deregt D et al (2005) Virus Res 114 , 140 Nettleton PF et al (1986) Vet Bull 118 , 267 Nielsen O et al (2000) J Wildl Dis 36 , 508 Rima BK et al (2005) Virus Genes 30 , 113 van de Bildt MW et al (2005) Arch Virol 150 , Cervid herpesvirus 2 (CvHV-2) A spe- 577 cies in the genus Varicellovirus , isolated from a reindeer, Rangifer tarandus . cetyltrimethylammonium bromide (CTAB) Synonyms: Rangifer tarandus herpes- A solvent that is widely used to dissolve virus; reindeer herpesvirus. DNA samples. Deregt D et al (2005) Virus Res 114 , 140 El-Kommonen C et al (1986) Acta Vet Scand CEV BFS-283 virus A strain of Cali- 27 , 299 fornia encephalitis virus in the genus Orthobunyavirus. Isolated from mosquitoes. cervine adenovirus (OdAdV-1) A tentative species in the genus Atadenovirus . CFT Abbreviation for c omplement f ix- Isolated from black-tailed deer during ation t est. an outbreak of hemorrhagic disease affecting mule deer, ( Odocoileus hemi- CFU Abbreviation for c olony f orming onus), black-tailed deer ( Odocoileus u nit. hemionus columbianus) and Rocky Mountain mule deer (Odocoileus hemi- CG1820 virus (CG1820V) Bashkiria onus hemionus) in central and northern virus, a Russian strain of Puumala virus California in 1993. Fawns were mostly in the genus Hantavirus. Closely related affected, with pulmonary edema and by sequence analysis to the Sotkamo intestinal lumenal hemorrhage causing strain of Puumala virus . high mortality. The disease could be reproduced experimentally in white- Piiparinen H et al (1997) Virus Res 51 , 1 tailed deer (Odocoileus virginianus ). A similar disease was subsequently CGBQ cells (CCL 169) This cell line was reported in captive moose (Alces alces ) derived from the sternum of a 15- to in Toronto, Canada. 17-day-old embryo of the domestic goose, Anser anser . Shilton CM et al (2002) J Zoo Wildl Med 33 , 73 C group viruses Fourteen serologically Woods LW et al (2001) J Wildl Dis 37 , 153 Zakhartchouk A et al (2002) Arch Virol 147 , related viruses ( Table C2 ) in the 841 genus Orthobunyavirus , found only in the Western Hemisphere. All except Nepuyo and Gumbo limbo viruses cetacean calicivirus (VESV/Tur-1) A have been associated with febrile ill- strain of Vesicular exanthema of swine ness in humans. There are three sub- virus in the genus Vesivirus . Isolated groups. from a bottle-nosed dolphin (Tursiops truncatus ) with vesicular skin lesions. Ch1ES(NBL-8) cells (CCL 73) A hetero- Smith AW et al (1983) J Am Vet Assoc 183 , ploid line derived from the trypsinized 1223 esophagus of a male fetus of the goat, Capra hircus. Developed for study of Cetacean morbillivirus (CeMV) A species viruses that affect domestic animals, in the genus Morbillivirus . In 1990 many and has been used for scrapie prion striped dolphins in the Mediterranean studies. Chaco virus (CHOV) 90

Table C2. The 14 C group viruses

(1) Apeu (C, H, M) (2) Bruconha (C) (3) Itaqui (C, H, R, M) Caraparu (C, H, R) Gumbo limbo (C, H, R) Oriboca (C, H, R, M) Madrid (C, H, R) Marituba (C, H, M) Ossa (C, H, R) Murutucu (C, H, R, M) Nepuyo (C, R, Ba) Restan (C, H) Vincés (C) 63U11

Isolated from: Ba, bats; C, culicine mosquitoes; H, humans; M, marsupials; R, rodents.

Chaco virus (CHOV) An unassigned spe- Chagres virus (CHGV) A tentative species in the family Rhabdoviridae . With cies in the genus Phlebovirus , belong- Timbo virus and Sena Madureira virus ing to the sandfly fever group with forms the Timbo serogroup. Isolated no complex assigned. Isolated from a from the lizards, Ameiva ameiva ameiva man with a febrile illness in the Canal and Kentropyx calearatus, in Para, Brazil. Zone, Panama. Also isolated from the Not isolated from arthropods but con- mosquito, Sabethes chloropterus and sidered to be arthropod-transmitted Lutzomyia sp. Causes CPE in primary as it will replicate in experimentally rhesus monkey kidney, human amnion, infected mosquitoes. Kills newborn and mouse embryo cell cultures. On mice which are more sensitive than injection i.c. kills newborn mice. Vero cells in which it replicates with Robeson G et al (1979) J Virol 30 , 339 CPE at 30°C. Monath TP et al (1979) Arch Virol 60 , 1 chamois contagious ecthyma virus chaffinch papillomavirus (ChPV) A strain Tentative species in the genus of Fringilla coelebs papillomavirus, a spe- Parapoxvirus . Causes lesions similar to cies in the genusEtapapillomavirus . Causes those produced by goatpox virus, but papillomas exclusively on the legs of the lesions are primarily around the finches (chaffinch, Fringilla coelebs , and lips and mouth, and infrequently on brambling, Fringilla montifringilla ). the udder, thighs, anus, nostrils, and Usually only one leg is affected, and eyes. Antigenically different from goat- the whole foot may be affected with a pox virus and the two do not cross- highly keratinized papillomatous lesion protect. An important cause of disease in severe cases. Typical papillomavirus in sheep, and a live vaccine is used. As particles, 52 nm in diameter, were first virus remains active in scabs for a long seen in the tumors by electron micros- period the vaccine is used annually. copy, and shown to have 72 capsomeres Occurs in many parts of the world. arranged in a skew (T 7d) lattice as Synonyms : contagious ecthyma virus; for human papillomavirus. Virus DNA chamois papillomavirus. has been characterized and is partially Renshaw HW and Dodd AG (1978) Arch related to bovine papillomavirus, and Virol 56 , 201 more closely to parrot papillomavirus. Mercer A et al (1997) Arch Virol Suppl 13 , 25 The incidence of the tumors, based on data from European bird-ringers, is chamois papillomavirus Synonym for 1–2%, and only chaffinches and bram- chamois contagious ecthyma virus. blings seem to be affected. Synonym : fringilla papillomavirus. Lina PHC et al (1973) J Natl Cancer Inst 50 , 567 Chandipura virus (CHPV) A species in Moreno-Lopez J et al (1984) J Virol 51 , 872 the genus Vesiculovirus . Antigenically Tachezy R et al (2002) BMC Microbiol 10 , 19 related to vesicular stomatitis virus. 91 chelonid herpesvirus 2 (ChHV-2)

Isolated from a man with dengue/ This results from the pairing of the chikungunya-like disease, in Nagpur, bases. Maharashtra, India. Also isolated from sand flies, Phlebotomus sp, and a hedge- Charleville virus (CHVV) An unassigned hog, Atelerix sp, in Nigeria. vertebrate rhabdovirus. Isolated from sand flies of Phlebotomus sp and the liz- Chandiru virus (CDUV) Does not exist. A ard, Gehyra australis , in Charleville and spelling error in the 7th and 8th ICTV Mitchell River, Queensland, Australia. Reports. See Candiru virus . Not reported to cause disease in humans. Chang conjunctiva cells (CCL 20.2) Wong Kilbourne derivative. A clone chelating agent Compounds which bind (1-5c-4) will support replication of divalent cations, so inhibiting their influenza viruses. activity. Can be used to inhibit biological interactions which require divalent Chang liver cells (CCL 13) A heteroploid cations, e.g. deoxyribonuclease activ- cell line established from non-malignant ity, or attachment of some viruses (e.g. human liver. One of the original suc- Foot-and-mouth disease virus) to cells. cessful attempts in serial propagation of human cells from normal tissues chelonid herpesvirus 1 (ChHV-1) An and named after the man who accom- unassigned species in the family plished it. Herpesviridae . Epizootics of gray patch disease were observed in green sea tur- Changuinola virus (CGLV) A species tles, Chelonia mydas, kept in captivity in the genus Orbivirus. There are 12 in the West Indies. Two types of lesion serotypes. Isolated from sand flies were seen: papules and spreading (Phlebotomus sp) and small rodents in gray patches 7–8 weeks after hatching. Panama. Has also been isolated from a Intranuclear inclusions were present in mosquito catcher with a febrile illness. sections of the lesions and herpesvirus- like particles were present in scrapings channel catfish herpesvirus (CCHV) See from the lesions. The disease could be Ictalurid herpesvirus 1 . transmitted between turtles by a cell- free extract. Fibropapillomas of both channel catfish reovirus (CRV) A strain green sea turtles, Chelonia mydas , and of Aquareovirus D in the genus Aqua- loggerhead turtles, Caretta caretta , in reovirus , isolated from catfish, Ictalurus Florida appear to be caused by the punctatus, but not associated with same herpesvirus, found only in asso- disease. ciation with the tumor cells. The virus Chinchar VG et al (1996) Dis Aquat Org 33 , could not be cultivated in chelonian 77 cell lines, which support the replication Hedrick RP et al (1984) J Gen Virol 65 , 152 of other chelonian herpesviruses. Synonyms: green sea turtle herpesvi- chaperones Molecules which assist the rus; gray patch disease agent of green replication and assembly of viruses. sea turtle; gray patch disease of turtles Examples are human heat-shock provirus. teins Hsp40 and Hsp70 which assist Haines H and Kleese WC (1977) Infect the human papillomavirus E1 to bind Immun 15 , 756 to the origin of DNA replication, or Lackovich JK et al (1999) Dis Aquat Org 30 , the lectin chaperones calnexin and cal- 89 reticulin which facilitate folding of the Rebell G et al (1975) Am J Vet Res 36 , 1221 hemagglutinin of influenza virus in the endoplasmic reticulum. chelonid herpesvirus 2 (ChHV-2) An unassigned virus in the family Chargaff’s rule In double-stranded DNA Herpesviridae . Isolated from the Pacific molecules, the proportion of adenine pond turtle. equals that of thymine, and the propor- Synonym : Pacific pond turtle herpes- tion of guanine equals that of cytosine. virus. chelonid herpesvirus 3 (ChHV-3) 92 chelonid herpesvirus 3 (ChHV-3) An chemotherapeutic index The ratio unassigned virus in the family Herpes- between the lowest effective antiviral viridae. Isolated from the painted turtle, concentration and the highest non- Chrysemys picta . toxic concentration. Synonyms: map turtle virus; painted turtle herpesvirus. chemotherapy The use of compounds which will inhibit the growth of infec- chelonid herpesvirus 4 (ChHV-4) An tious agents or tumors without unduly unassigned virus in the family Herpes- affecting normal host cell metabo- viridae. Isolated from Argentine lism, e.g. acyclovir for herpesvirus tortoises, Geochelone chilensis , with infections. necrotizing stomatitis. Red-footed tortoises, Geochelone carbonaria , housed Chenuda virus (CNUV) A species in together with the Argentine tortoises, the genus Orbivirus belonging to the remained healthy and did not develop Chenuda complex of the Kemerovo the disease. serogroup. There are seven serotypes. Synonyms: Argentine turtle herpes- Isolated from ticks, Argas hermanni , and virus; geochelone carbonaria herpes- other species, in Egypt and Africa. Not virus; geochelone chilensis herpesvirus. reported to cause disease in humans. Jacobsen ER et al (1985) J Am Vet Med Assoc 187 , 1227 Chiba virus (ChV) A possible species in the genus Norovirus, it was first iden- chemokine receptors Type III trans- tified as a cause of an oyster associ- membrane proteins of the rhodopsin ated outbreak of gastroenteritis that superfamily (with seven membrane- occurred in Chiba prefecture, Japan in spanning domains). There are different 1987. However, since the first isolation, receptors for alpha-chemokines and multiple genotypes of norovirus have beta-chemokines. been isolated in Chiba prefecture, and the name is no longer used. chemokines A class of cytokines with cell-specific chemo-attractant and other Kobayashi S et al (2000) J Med Virol 62 , 233 activating activity for various cell-types Okada M et al (2005) J Clin Microbiol 43 , 4391 Someya Y et al (2000) Virology 278 , 490 within the immune system. Divided into two types. The alpha-cytokines such as interleukin 8 and macrophage chick embryo fibroblast (CEF) cells A inflammatory protein (MIP) 2 act pri- primary cell culture system pre- marily upon neutrophil leukocytes. pared from chick embryos. Used for The beta-chemokines such as mono- culture of many viruses, especially cyte chemotactic protein (MCP) and orthomyxoviruses. RANTES (r egulation upon a ctivation n ormal T -cell expressed and s ecreted) Chicken anemia virus (CAV) The only act on monocytes and macrophages. species in the genus Gyrovirus , which causes aplastic anemia in chickens. chemokinesis Random migration of cells Distributed worldwide both in com- brought about by a specific chemical mercial and specific pathogen-free substance in their environment. poultry flocks. May also infect turkey and quail. Probably immuno- Lusso P (2000) Virology 273 , 228 suppressive and most serious as a chemotaxis Directed migration of cells dual infection with avian reticulo- in response to a chemical substance endotheliosis virus, avian adenovirus in their environment. Cells may move or infectious bursal disease virus. The toward (positive chemotaxis) or away single-stranded circular DNA genome from (negative chemotaxis) the source has a negative-sense organization, in or highest concentration of the chemi- contrast to other circoviruses, and the cal substance. Important in recruit- virions are larger. ment of leukocytes to sites of injury or Noteborn MH (2004) Vet Microbiol 98 , 89 inflammation. Toro H et al (2000) Avian Dis 44 , 51 93 Chikungunya virus (CHIKV)

Chicken astrovirus 1 (CastV-1) A species chicken rotavirus G/555 (AvRV-G/555) A in the genus Avastrovirus . Antibody strain of Rotavirus G, a tentative species to the virus is widespread in broiler in the genus Rotavirus . flocks. The virus is present in all parts of the small intestine of infected birds, chick helper factor (chf) An endogenous causing mild diarrhea and distension avian leukosis virus, present in certain of the small intestine. chicken cells. It can act as helper for a defective avian sarcoma virus by pro- Baxendale W and Mebatsion T (2004) Avian Pathol 33 , 364 viding genetic information for the viral Koci MD and Schultz-Cherry S (2002) Avian surface glycoproteins of the infectious Pathol 31 , 213 progeny virus. Probably the same as RAV-60. See Avian leukosis virus . chicken embryo lethal orphan virus Hanafusa H et al (1970) Proc Natl Acad Sci (CELO) A strain of Fowl adenovirus 66 , 314 A in the genus Aviadenovirus , closely related to quail adenovirus. Isolated Chick syncytial virus (CSV) A species in originally from embryonated eggs the genus Gammaretrovirus , belonging inoculated with various materials, and to the Avian (Reticuloendotheliosis) in which the embryos died. It became virus group. Causes syncytia formation clear that the source of the virus was in some cell lines, and in young birds the eggs and not the injected material. there is immunosuppression, and other It has been associated with various dis- syndromes such as abnormal feather eases in chickens, turkeys and pheas- development, bursal and thymic atro- ants. Oncogenic in newborn hamsters. phy, and anemia. Less pathogenic than Replicates with typical adenovirus CPE reticuloendotheliosis virus. A repli- in cell cultures. The use of this virus as cation competent virus that does not a gene vector in humans is under active carry an oncogene. Causes reticulo- investigation. endotheliosis, splenomegaly, and occa- Synonym : fowl adenovirus 1. sionally lymphomas in chickens. Aghakhan SM (1974) Vet Bull 41 , 531 Chiocca S et al (1996) J Virol 70 , 2939 Chikungunya virus (CHIKV) A species Stevenson M et al (2006) Gene Ther 13 , 356 in the genus Alphavirus. Causes an epidemic disease in humans charac- chicken leukosis sarcoma virus See terized by joint and back pains which Avian leukosis virus . are so severe that the patient is doubled up; hence the name, a Tanzanian Chicken parvovirus (ChPV) A species in word meaning ‘ that which bends up.’ the genus Parvovirus. Causes runting Incubation period 3–12 days. Onset disease in broiler chickens with intesti- sudden and the disease biphasic. nal lesions and poor weight gain. Following 1–6 days ’ fever the temperature returns to normal for 1–3 Kisary J et al (1985) J Gen Virol 66 , 2259 days before the second period of fever which lasts a few days. In this phase chickenpox virus Not a poxvirus. A a maculopapular pruritic rash on the herpesvirus called varicella virus. trunk and limbs is common. After Name probably derived from Old 6–10 days, recovery is usually com- English gican, meaning itch, as this is a plete although rarely joint pains may characteristic symptom. See Human persist. Hemorrhagic fever-type dis- herpesvirus 3 . ease may occur but shock is uncommon. Transmitted by various Aedes sp chicken rotavirus D/132 (AvRV-D/132) including A furcifer , A taylori , A africa- Reoviridae. A strain of Rotavirus D , a nus , and A aegypti. Mansonia africana species in the genus Rotavirus . may also transmit the virus. No vertebrate host is known, but antibodies are chicken rotavirus F/A4 (AvRV-F/A4) present in monkeys, birds and many Reoviridae. A strain of Rotavirus F , a ten- mammals. Occurs in India, South East tative species in the genus Rotavirus . Asia, eastern, western, central and Chikungunya virus (CHIKV) 94

southern Africa. The virus re-emerged genus Mastadenovirus, related to human to cause massive outbreaks in India adenovirus A. and Malaysia in 2006. Strains from Asia and Africa show some antigenic differ- chimpanzee agent Synonym for Pongine ences. Kills suckling mice on injection herpesvirus 1 (chimpanzee herpesvirus). i.c; guinea pigs and rabbits show no signs of infection. Replicates with CPE chimpanzee coryza agent The origi- in chick embryo, rhesus kidney, and nal name given to the virus now des- HeLa cell cultures. ignated respiratory syncytial virus. Jain M et al (2008) Trop Doct 38 , 70 Isolated from a throat swab from a Chanas AC et al (1979) Arch Virol 59 , 231 chimpanzee with clinical symptoms Sam IC and AbuBaker S (2006) Med J of coryza at the Walter Reed Army Malaysia 61 , 264 Institute of Research in 1955. Saxena SK et al (2006) Euro Surveill 11 , EO60810.2 chimpanzee cytomegalovirus Synonym for Pongine herpesvirus 4 . Chilibre virus (CHIV) A species in the genus Phlebovirus isolated from Chimpanzee foamy virus (CFV) The sand flies. With Cacao virus forms type species of the genus Spumavirus . the Chilibre virus antigenic complex. Causes characteristic ‘ foamy ’ cytopa- Isolated from insects of Lutzomyia sp in thology in cell culture. No diseases Canal Zone, Panama. Not reported to have been associated with infection. cause disease in humans. chimpanzee foamy virus human iso- Chilibre virus VP-118D (CHIV) A strain late (CFV/Hu) Probably a strain of of Chilibre virus . Chimpanzee foamy virus . No natural human infections have been reported. Chiloe wigeon hepadnavirus An avihepadnavirus cloned from captive chimpanzee herpesvirus 1 Synonym for Chiloe wigeon. The genomic sequence Pongine herpesvirus 1 . is closely related to duck hepatitis B virus. Chinese hamster ovary See CHO-K1 cells . Guo H et al (2005) J Virol 79 , 2729 chinook salmon reovirus (DRCRV, Chim virus (CHIMV) An unassigned, GRCV, LBSV, YRC, ICRV) Serotypes ungrouped virus in the family of Aquareovirus A and B in the genus Bunyaviridae. Isolated from ticks in Aquareovirus. Isolated from adult Coho Uzbekistan. Not reported to cause dis- salmon, Oncorhynchus kisutch, and chi- ease in humans. nook salmon, Oncorhynchus tshawytscha , in Oregon, USA. Distinct antigenically chimera An individual composed of two from chum salmon reovirus. genetically distinct types of cells. Synonym : Sacramento River chinook salmon disease virus. chimeric DNA A recombinant DNA molecule containing sequences from more Chipmunk parvovirus A tentative spe- than one organism. cies in the genus Erythrovirus . Isolated in Korea from the Manchurian chip- chimpanzee adenovirus C2 (ChAdV-C2) munk (Tamias sibiricus asiaticus ). The A strain of simian adenovirus, a nucleotide sequence of the genome tentative species in the genus Mas- showed significant homology to parvo- tadenovirus , related to human adenovirus B19. virus C. Yoo BC et al (1999) Virology 253 , 250 chimpanzee adenovirus strain Y34 chlamydia A group of obligate intracellu- (ChAdV-Y34) A strain of simian lar microorganisms, once thought to be adenovirus, a tentative species in the viruses, but now classified as bacteria. 95 Chordopoxvirinae

They are unlike true viruses in several chloroquine An antimalarial 4-amino- important respects: quinoline drug which is a potential (1) they multiply by binary fission; antiviral agent against chikungunya (2) they have cell walls like bacteria, virus. Infection of cells by viruses that with muramic acid-containing enter through endosomes can be inhib- mucopeptides; ited by lysosomotropic agents such as (3) they have a number of enzymes chloroquine. which are metabolically active; (4) they possess both DNA and RNA, CHO-K1 cells (CCC 61) A cell line as do bacteria; derived from Chinese hamster ovary (5) unlike viruses, they possess ribo- tissue. A range of drug-resistant somes; mutants of these cells has been devel- (6) growth can be inhibited by a num- oped, including lines resistant to ber of antibacterial agents, includ- α -amanitin. ing tetra cyclines, chloramphenicol, Kao F and Puck TT (1968) Science 164 , 312 rifampicin, and sulfonamides. Lobban PE and Siminovitch L (1975) Cell 4 , Synonyms : Bedsonia; Miyagawanella; 167 TRIC agent. Moulder JW (1966) Annu Rev Microbiol 20 , Chobar Gorge virus (CGV) A species 107 in the genus Orbivirus. Isolated from ticks of Ornithodoros sp in Nepal. Not chlamydozoa ribasi A name given to the reported to cause disease in humans. elementary bodies seen in association Fomeda virus is a serotype of the with cells infected with variola minor species. virus. Aragao HB (1933) C R Soc Biol Paris 113 , Choclo virus (CHOV) A tentative species 1271 in the genus Hantavirus . Isolated from patients and trapped rodents in 2000 chlamydozoa variolae Synonym for during an outbreak of hantavirus pul- Paschen bodies. See also Buist bodies . monary syndrome in Las Tablas, Los Santos Province, Panama. The rodent chloramphenicol An antibiotic isolated host is the pygmy rice rat, Oligoryzomys from Streptomyces venezuelae which fulvescens . inhibits protein synthesis in prokaryo- Vincent MJ et al (2000) Virology 277 , 14 tes and in mitochondria of eukaryotes. CHOHR genes Genes which permit chloramphenicol acetyl transferase (CAT) a virus to grow in Chinese hamster gene The CAT gene product catalyzes ovary cells. The CP77 gene of cowpox- the acetylation of chloramphenicol. virus encodes a 77 kDa protein that The most commonly used type was allows the virus to grow on CHO cells, obtained from the bacterial transposon, but it is deleted or non-functional in Tn9. The CAT gene is frequently used vaccinia virus, which cannot grow on as a marker in genetic cloning experi- CHO cells. ments to detect gene expression. cholecystokinin A neurotransmitter 1-(p-chlorophenyl)-3-(m-3-isobutylgua- whose expression is greatly reduced in nidinophenyl)urea hydrochloride A the acute phase of infection with Borna guanidine derivative differing signifi- disease virus. cantly from guanidine hydrochloride in its antiviral properties. Has very cholestasis Stopping or suppression of high in vitro activity against rhino- the flow of bile, a severe complication viruses. Relatively non-toxic to labora- of hepatitis virus infection. tory animals. Synonym : ICI 73602. Chordopoxvirinae A subfamily of the Swallow DL et al (1977) Ann NY Acad Sci family Poxviridae , comprising the pox- 284 , 305 viruses of vertebrates. Contains eight Chordopoxvirinae 96

genera: Orthopoxvirus , Parapoxvirus , wrapped around this protein core. The Avipoxvirus , Capripoxvirus , Leporipox- nucleosomes are arranged in supra- vi rus , Suipoxvirus , Molluscipoxvirus , and nucleosomal clusters and packaged into Yatapoxvirus . loops, each containing 30–90 kb of DNA by supercoiling to form the 30 nm chro- chorioallantoic membrane (CAM) A matin fiber. This fiber undergoes fur- highly vascularized extra-embryonic ther coiling and condensation to form membrane formed in birds and reptiles visible chromosomes. by apposition of the allantois to the inner face of the chorion. Many viruses chromatography A method of separating such as orthomyxoviruses and para- and analyzing chemical substances by myxoviruses will grow on the CAM of preferential retention, either as a gas or the 11- to 13-day fertilized egg. liquid, on to a solid medium, such as paper or a gel. chorionic villus sampling A procedure for direct testing of the fetus using a chromoneme Genome of bacteria or needle guided by ultrasound. Can be viruses: term introduced by White- used to test, e.g. for rubella infection. house (1969) to distinguish the ‘ thread- Daffos F et al (1983) Prenat Diagn 3 , 271 like ’ structure of bacterial or viral genetic material from the true chromo- chorioretinitis Inflammation of the some of plant and animal cells. choroid and retina of the eye, which can proceed to blindness. Occurs Whitehouse HLK (1969) Towards an in infections by measles virus caus- Understanding of the Mechanism of Heredity . ing subacute sclerosing panencepha- London: Edward Arnold. litis and in congenital infection by cytomegalovirus. chromosomal translocations One theory concerning the development of choroid plexus An important point of Burkitt’s lymphoma is that in the pres- entry to the brain for blood-borne ence of hyper-endemic malaria, three viruses because it contains a fenes- specific chromosomal translocations trated endothelium and a sparse base- can occur (18:14; 18:22; 12:8) each of ment membrane. which can deregulate the c-myc oncogene on chromosome 8 by bringing it CHP 3 and 4 (WW) cells (CCC 132 and under the influence of an immunoglob- 133) These fibroblast-like cell lines ulin gene promoter on chromosomes were established from skin biopsies 14, 22, or 2. of two young sibling black males, one with classic galactosemia and one with chromosomes Structures formed from asymptomatic galactosemia. chromatids during mitosis. Contain the nucleoprotein of the nucleus in chromatids Thread-like structures for- an organized form in which it can be med from chromatin during the first divided into two equal parts and thus stage of mitosis (between prophase), pass on the genetic information to the after which they are known as daugh- daughter cells. ter chromosomes. chronic adult T-cell leukemia Following chromatin The material which makes infection with T-cell lymphotropic virus up cell chromosomes, consisting of 1 person has an estimated 1% chance equal parts by weight of DNA and of developing adult T-cell leukemia histones (low-molecular-weight basic (ATL). In chronic ATL there is a high proteins), and varying amounts of leukocytes count associated with lym- non-histone chromosomal proteins. phadenopathy and hepatosplenom- Eukaryotic chromatin is built up from egaly. The mean survival time is 24 nucleosomes, each consisting of eight months and a projected 4-year survival histone molecules with 146 bp of DNA time of 5.7%. 97 Circoviridae chronic fatigue syndrome (CFS) A Chuzan virus A synonym for Kasba virus, severely disabling fatigue with self- a strain of Palyam virus in the genus reported impairments in concentration Orbivirus. Causes congenital abnor- and short-term memory, sleep dis- malities in cattle. Phylogenetically it is turbances and musculoskeletal pain. most related to African horsesickness Occurs worldwide. A number of infec- virus. tious agents have been proposed as eti- Yamakawa M et al (1999) J Gen Virol 80 , 937 ologic agents of CFS, including human herpesviruses 4 (EBV), 5 (CMV) and chymotrypsin A protease which hydro- 6 (HHV6); enteroviruses; retroviruses; lyzes the peptide bond on the and Borrelia burgdorferi . None has C-terminal side of valine, isoleucine, proved to be a unique causative agent, phenylalanine, tyrosine, methionine, but it remains possible that such infec- and some alanine residues. tions act as a trigger for the syndrome. Synonyms : post-viral fatigue syndrome; Aler RP and Meadway RJ (1968) Biochem J myalgic encephalitis (ME). 108 , 893 Devanur LD and Kerr JR (2006) J Clin Virol ciconiid herpesvirus 1 (CiHV-1) An unas- 37 , 139 signed virus in the family Herpesviridae . Fukuda K et al (1994) Ann Int Med 121 , 953 Heneine W et al (1994) Clin Infect Dis 18 , Viruses have been isolated from black S121 storks, Ciconia nigra, and white storks, Mawle AC et al (1994) Infect Agents Dis 2 , Ciconia ciconia . The disease significance 333 is not known. Synonym : black stork herpesvirus. chronic progressive pneumonia of sheep Kaleta EF and Kummerfeld N (1983) Avian virus Synonym for Visna/maedi virus . Pathol 12 , 347 chronic wasting disease See wasting disease . cidofovir Hydroxyphosphonylmethoxy- cytosine (HPMPC). A potent inhibi- chronic wasting disease of mule deer tor of the replication of several DNA and elk A transmissible spongiform genome containing viruses, including encephalopathy caused by a prion. orthopoxviruses. Originally thought to be localized to Christensen ND et al (2000) Antiviral Res 48 , the western United States of Colorado 131 and Wyoming, but now detected over a wider geographical range. cinchocaine A local anesthetic which Belay ED et al (2004) Emerg Infect Dis 10 , 977 has been found to lyse chick embryo Manson JC et al (2006) Biochem Soc Trans 34 , fibroblasts. Cells transformed by avian 1155 leukosis sarcoma virus are more sensi- Williams ES (2005) Vet Pathol 42 , 530 tive to lysis than normal cells, perhaps because they attain higher intracellular chub reovirus (CHRV) A tentative spe- levels of the drug. cies in the genus Aquareovirus , found Synonym : dibucaine hydrochloride. in chub, Leucuscus cephalus. Not associ- Rifkin DB and Reich E (1971) Virology 45 , ated with disease. 172 Ahne W and Kolbi O (1987) J Appl Ichthyol 3 , 129 Circinoviridae Suggested but not adopted family name for TT virus and chum salmon reovirus CS (CSRV) A related viruses, which have a circular strain of Aquareovirus A in the genus negative single-stranded DNA genome Aquareovirus , found in normal salmon, about 3.8 kb in length. Now classified Oncorhynchus keta , in Japan. in a floating genus, Anellovirus. See TT Winton JR et al (1987) J Gen Virol 68 , 353 virus . chum salmon reovirus PSR (PSRV) A Circoviridae A family of small non- strain of Aquareovirus F in the genus enveloped viruses, 17–22 nm in diam- Aquareovirus . eter, with a circular single-stranded Circoviridae 98

DNA genome 1.7–2.3 kb in length. defined by the cis–trans test: if x- and y Only one protein has been found in the are two function-abolishing mutations virion. There are two genera, Circovirus and they are on the same cistron, then and Gyrovirus. The name derives from in a mixed infection, x y /x − y − , the circular form of the ca 2 kb genome. the cis diploid, will be functional but x−y /x y− , the trans diploid will not Circovirus A genus in the family function because x and y , being on the Circoviridae, containing six species: Beak same cistron, cannot function alone – and feather disease virus , Canary circo- they are indivisible. See gene . virus , Goose circovirus , Pigeon circovirus , Porcine circovirus 1 , and Porcine circo- citrullinemia cells (CCC 76) A human virus 2 . Tentative species in the genus diploid fibroblast cell line derived from include circoviruses of duck, finch, and skin of a female Caucasian infant with gull. citrullinemia. circular dichroism (CD) spectroscopy A Mellman WJ (1967) Proc Natl Acad Sci 57 , 829 form of spectroscopy based on the differential absorption of left- and right- handed circularly polarized light. c-Jun transcription factor A member of the AP-1 family of transcription fac- circulating recombinant forms (CRFs) tors. Interacts with the genome of JC Isolates of HIV-1 containing RNA from polyomavirus. Present as v-Jun as an more than one subtype. At least 14 oncogene in avian sarcoma virus 17. have so far been isolated. clade A segment of a phylogenetic tree; Robertson DL et al (2000) Science 288 , 55 a monophyletic group sharing a closer common ancestry with one another cirrhosis A serious liver disease charac- than with members of any other clade. terized by diffuse interlacing bands of fibrous tissue dividing the hepatic parenchyma into micronodular or Classical swine fever virus (CSFV) A macronodular areas. May result from species in the genus Pestivirus . A highly chronic infection with viruses such as contagious disease which only affects hepatitis B or hepatitis C. pigs. Causes fever, apathy, vomiting, eye discharge, diarrhea and cutane- Cisternavirus A A name once proposed ous hemorrhages, and is frequently but not adopted for a genus in the fam- fatal. Secondary bacterial infection ily Retroviridae, which would contain often occurs. Strains vary in virulence: the viruses with A-type virus particles. some are mild, others neurotropic and some are poorly neutralized by anti- Dalton AJ et al (1975) J Natl Cancer Inst 55 , serum. Disease first reported in the mid- 941 western USA in about 1810, then in the UK in 1862, and has since spread cis–trans test A genetic complementa- worldwide. The pathological lesion tion test originally used in fine struc- consists of degeneration of small blood ture genetic mapping of Drosophila and vessels causing hemorrhage. There is T4 bacteriophage. Used to determine leukopenia, atrophy of the thymus, whether two mutations affecting the and lymphocytic depletion of periph- same character lie within the same or eral lymphoid tissues. Infection of different cistrons. See cistron . sows 10–50 days pregnant may result Benzer S (1955) Proc Natl Acad Sci 41 , 344 in infection of the fetuses, abortion, or congenital tremor due to cerebellar cistron The unit of genetic function; a hyperplasia. Hyperimmune serum can segment of DNA or RNA which codes provide passive immunity, and there for a single gene product. An mRNA are several vaccines available, attenu- which encodes two or more proteins ated in rabbits or in cell culture, which is called polycistronic. No smaller unit provide active immunity. However, or part of a cistron is functional. It is the European Union has adopted a 99 cleavage

non-vaccination policy, and control classical swine fever virus Brescia A in those countries is by slaughter of strain of Classical swine fever virus . infected herds. Although pigs are the only natural host, experimental hosts classical swine fever virus C strain A include goats, sheep, calves, deer, and strain of Classical swine fever virus . laboratory rabbits, but not wild mice, cottontail rabbits, rats, or sparrows. classification The process of grouping Passage in laboratory rabbits, which biological entities such as viruses, on only show transient fever experimen- the basis of features they have in com- tally, results in attenuation of the virus mon, into a hierarchical series. The for pigs. Inactivated on drying in the formal hierarchy proceeding from the air and following exposure to lipid largest to the smallest group includes solvents. The virus is antigenically kingdom, phylum, class, order, family, related to bovine viral diarrhea virus. genus, and species. In some classifica- The virion is 40–50 nm in dia meter tions the group name cohort is inserted with a nucleocapsid core of 29 nm between class and order, and the group diameter and an envelope 6 nm thick. name tribe between family and genus. The genome is a linear positive single- In the case of viruses, a taxonomic stranded RNA molecule, 12.3 kb in grouping above the level of family has length, and encodes a polyprotein that not been developed, except for three is cleaved post-translationally to form orders (Caudovirales , Mononegavirales , the virion and nonstructural proteins. and Nidovirales ). In addition, the ques- There is no 3 terminal poly A tract. In tion of what constitutes a virus species recent years the virus has been eradi- has been the subject of much heated cated from Australia, Canada, the USA, debate. Scandinavia, and Ireland, but it has van Regenmortel MHV (2007) Infect Genet continued to circulate and cause out- Evol 7 , 133 breaks of disease in Europe, especially van Regenmortel MHV and Mahy BWJ in The Netherlands in 1997 and in the (2004) Emerg Infect Dis 10 , 8 UK in 2000. To maintain disease-free status, early rapid diagnosis is essential clathrin A scaffolding protein composed to eradicate the disease if it appears. of three heavy chains (180 kDa) which Vaccination is also feasible in endemic associates with three light chains areas but precludes the use of serologi- (34–36 kDa) to form a three-legged pro- cal diagnosis of infection, and so there tein complex called a triskelion, which is a need to develop marker vaccines forms a basket-like network around a that can be distinguished from natural coated pit. During receptor-mediated infection. endocytosis, a ligand on the virus Synonyms : European swine fever virus; surface binds to a plasma membrane pestivirus suis; swine fever virus; hog receptor, followed by lateral movement cholera virus. of the ligand–receptor complex toward Greiser-Wilke I and Moennig V (2004) Anim a clathrin-coated pit, which eventu- Health Res Rev 5 , 223 ally invaginates into the cytoplasm, Meyers G and Thiel H-J (1996) Adv Virus Res where a small endocytotic vesicle (the 47 , 53 endosome) is pinched off the pit. Once Moennig V and Plagemann PGW (1992) Adv the ligands have been internalized in Virus Res 41 , 53 an endosome, the receptor molecules van Oirschot JT and Terpstra C (1989) In are returned intact to the plasma Virus Infections of Porcines , edited by MB membrane. Pensaert. Amsterdam: Elsevier, p. 113 Clavelée virus Synonym for Sheeppox classical swine fever virus Alfort/187 A virus . strain of Classical swine fever virus . cleavage The cutting of nucleic acid or classical swine fever virus Alfort- protein, usually enzymatically, at spe- Tübingen A strain of Classical swine cific sites. See restriction endonucle- fever virus . ases and protease . Clinical Laboratory Improvement Act 100

Clinical Laboratory Improvement Act single restriction site for DNA inser- An act passed in the USA in 1992, tion, or a pair of restriction sites defin- which provides strict standards for ing a region that can be removed and quality assurance and control for all replaced with foreign DNA, and a tran- aspects of laboratory diagnosis of scription promoter sequence located infections. upstream of the insertion site. clinical trials Trials to determine the effi- closed reading frame A reading frame cacy and safety of antiviral drugs or which contains terminator codons that vaccines for use in the human popula- prevent its translation into protein. tion. Normally carried out in human subject volunteers. CM2 protein The M2 protein of influenza virus C. Clo Mor virus (CMV) A strain of Sakhalin virus in the genus Nairovirus . Isolated c-myc oncogene The cellular gene, from a tick, Ixodes (Ceratixodes) uriae , located on chromosome 8, encoding found under rocks in Cape Wrath, the MYC protein. The virally trans- Scotland. Not reported to cause disease duced oncogene v-myc was originally in humans. found in avian myelocytomatosis virus MC29. MYC is a nuclear transcription clone (1) A population of identical recom- factor. Amplification of c-myc expres- binant DNA molecules, all carrying the sion is found in many tumors, and in same inserted DNA sequence. (2) A the case of Burkitt’s lymphoma, asso- colony of microorganisms containing ciated with human herpesvirus 4, this a specific DNA fragment inserted into gene amplification is linked to chro- a vector. (3) A cDNA clone is a copy mosomal translocation and consequent of an RNA virus genome obtained by deregulation of c-myc . reverse transcription and placed into a subcloning vector. (4) A population CoAr 1071 virus (CA1071V) A strain of of cells or microorganisms of identical Tacaiuma virus in the genus Ortho- genotype. bunyavirus . clone 1-5c-4 cells See Chang conjunctiva CoAr 3624 virus (CA3624V) A strain of cells . Anopheles A virus in the genus Ortho- bunyavirus . cloned library A collection of cloned DNA sequences representative of the CoAr 3627 virus (CA3627V) A strain of genome under study. Tacaiuma virus in the genus Ortho- bunyavirus . cloning (1) Molecular cloning is the production of many identical copies of a Coari virus A probable species in the gene from a single gene inserted into genus Orbivirus isolated from phlebot- a cloning vector. (2) Cell cloning is the omine sand flies in the Amazon region production of genetically identical of Brazil. Antigenically related to the cells from a single cell. (3) Virus clon- Changuinola virus antigenic group. ing from a mixed population can be Not associated with disease in humans. achieved by repeatedly picking a single plaque and so selecting a single repli- Coastal Plains virus (CPV) An unas- cating species. signed vertebrate rhabdovirus. Isolated from a steer, Bos taurus , in northern cloning vector Any DNA molecule Australia. Not associated with disease capable of autonomous replication in humans. within a host cell, into which other Cybinski DH and Gard GP (1986) Aust J Biol DNA sequences can be inserted and Sci 39 , 225 thus amplified. Can be derived from a bacterial plasmid, a bacteriophage, coat The protective proteins surrounding or an animal virus. There is usually a the virus nucleic acid in mature virions. 101 codon

Includes the capsid, and may consist of Appleton H and Pereira MS (1977) Lancet ii , a single layer as in the Picornaviridae , or 780 several layers as in the Poxviridae . Turton J et al (1990) Epidemiol Infect 105 , 197 coated pit A cell surface depression or co-cultivation A mixed culture of two or invagination that mediates receptor- more different types of cell. If one of dependent endocytosis of a variety of them is permissive for the replication macromolecules, and buds into the cell of a latent virus present in the other to form a coated vesicle. Viruses such cell line, this virus may replicate and as Semliki Forest virus or influenza become detectable, i.e. be ‘rescued. ’ virus exploit this pathway to enter The cells in which replication takes cells. See also clathrin . place are known as ‘ indicator cells.’ The frequency of virus rescue during coated vesicle Intracellular vesicles com- co-cultivation can often be increased by posed of a membrane surrounded by a artificially fusing the cells, with result- protein lattice containing clathrin. ing heterokaryon formation. An exam- During receptor-mediated endocytosis, ple of rescue is the recovery of measles viruses such as influenza bind to a virus from the brain tissue of patients cell surface receptor containing sialic with subacute sclerosing panencephali- acid, and are taken into a coated pit. tis by co-cultivation with Vero cells. A coated vesicle is formed from the coated pit by pinching off into the Codajas virus An unclassified arbovirus cytoplasm with the virus inside. isolated from Culicine mosquitoes in Acidification of the interior of the vesi- the Amazon region of Brazil. Not asso- cle then occurs by a proton pump in the ciated with disease in humans. membrane. When the pH is sufficiently low, the virus and vesicle membranes Code of Safety Practices The appropri- fuse releasing the nucleocapsid into the ate practices for handling viruses in cytoplasm. See also clathrin . the laboratory are described in detail in the US Government HHS publi- Helenius A (1992) Cell 69 , 577 cation number (CDC) 93–8395 enti- Simons K et al (1982) Sci Am 246 , 58 tled Biosafety in Microbiological and Bio-medical Laboratories, Fifth edition, cobra herpesvirus Synonym for elapid LC Chosewood and DF Wilson (eds) herpesvirus 1. (2007).

Cocal virus (COCV) A species in the coding sequence The portion of a gene genus Vesiculovirus. Isolated from mites, that directly specifies the amino acid Gigantolaelaps sp, in east Trinidad. sequence of its protein product. Non- coding sequences include control cockatoo entero-like virus (CELV) regions such as promoters, and poly- An unassigned virus in the family adenylation signals and the intron Picornaviridae . sequences of unknown function found in certain eukaryotic genes. cockle agent A virus resembling a parvovirus found in many outbreaks of codon A group of three consecutive gastroenteritis involving ingestion of bases in a nucleic acid molecule which shellfish or cold foods. See cockle together specify a particular amino virus . acid during translation from mRNA. Since there are four bases, there are 64 cockle virus A probable species in the possible codons, but as only 21 amino genus Erythrovirus. A virus associated acids need to be specified, most are with gastroenteritis following inges- coded for by several alternative codons tion of cockles. Particles 25–26 nm, dis- (degeneracy of the code). However, no tinct from Norwalk virus, later found codon specifies more than one amino to have a single-stranded DNA genome acid. The sequence of bases which with sequence homology to B19 virus. make up a gene is translated as a series codon 102

of codons, beginning with an initiation antigen present in all a vian l eukosis codon AUG. This establishes the read- viruses. See Avian leukosis virus . ing frame within the gene. In some virus nucleic acids there is more than cohort study A type of prospective epi- one reading frame, so that up to three demiological study of a group of peo- different proteins may be specified by ple which aims to gather new data a single sequence of bases. Termination and identify cause–effect relation- of translation is specified by a termina- ships. When successful, it can provide tor codon UAG, UAA, or UGA except incontrovertible proof of a cause–effect in mitochondria where UGA appears relationship. to code for tryptophan. There is also evidence that UGA may code for seleno- cohesive ends The projecting 5 -single- cysteine in certain contexts in eukaryo- stranded ends occurring on certain tic cells (Taylor, 1994). double-stranded nucleic acid mole- Barrell BG et al (1979) Nature 282 , 189 cules which, through sequence comple- Crick FHC (1963) Prog Nucleic Acid Res Mol mentarity, can base-pair and thus form Biol 1 , 163 a circular molecule. Such structures Taylor EW et al (1994) J Med Chem 37 , 2637 occur in the genome DNA of temperate bacteriophages and facilitate integra- codon 129 The human prion protein con- tion into host DNA. tains an amino acid polymorphism for methionine or valine at codon 129, and it appears that those who are Cohn fractionation technique A method homozygous for valine at this position for fractionation of proteins from appear to be more likely to develop plasma by precipitation with cold etha- Creutzfeldt–Jakob disease whether nol. It yields three fractions, enriched spontaneously or following iatrogenic in cryoprecipitates, globulins and albu- infection. In the normal Caucasian pop- min, respectively. A modification of the ulation, 51% have methionine/valine original technique is used to prepare at codon 129, 12% are homozygous for human immunoglobulins for viral valine, and 37% are homozygous for immunoprophylaxis or treatment. methionine. Cunningham-Rundles C (1992) In Encyc- Collinge J et al (1991) Lancet 337 , 1441 lopedia of Immunology, edited by IM Roitt and PJ Delves. London: Academic Press, codon bias The non-uniform distribution p. 598 of codon usages which results in preferred codons being used by viruses or Coho salmon reovirus CSR (CSRV) A organisms to specify a particular amino strain of Aquareovirus B in the genus acid. Aquareovirus . Air GM et al (1976) J Mol Biol 107 , 445 Coho salmon reovirus ELC (ELCV) A cod ulcus virus A virus found in associa- strain of Aquareovirus B in the genus tion with ulcus syndrome of cod, Gadus Aquareovirus . morhua. A tentative member of the family Iridoviridae . Coho salmon reovirus SCS (SCSV) A Jensen NJ et al (1979) Norg Vet Med 31 , 436 strain of Aquareovirus B in the genus Aquareovirus. Isolated from adult Coho Coe virus An old name for coxsackie salmon,Oncorhynchus kisutch, in Oregon, virus A2l, which can cause a ‘common USA. cold ’ -like disease. Winton JR et al (1989) In Viruses of Lower cofactor Additional (non-protein) com- Vertebrates , edited by W Ahne and E Kurstak. ponent required by an enzyme for its Berlin: Springer, p. 257 action. Coho salmon reovirus SSR (SSRV) A COFAL test A co mplement f ixation test strain of Aquareovirus F in the genus for the common group-specific (gs) Aquareovirus . 103 Colorado tick fever virus (CTFV) coital exanthema virus Synonym for colony A cluster of cells derived from equid herpesvirus 3. However, some a single cell by division on a solid cases of coital exanthema have been medium, e.g. agar. attributed to equid herpesvirus 1, which also causes genital infection. Colony B North virus A strain of Great Island virus in the genus Orbivirus . cold-adapted mutants Mutants which colony-forming units (CFU) The number can replicate at temperatures below of colonies formed per unit of volume that at which the wild type gener- or weight of a cell suspension. ally replicates or replicates optimally. Influenza virus can be adapted to rep- colony-stimulating factors Substances licate in eggs at 25°C and then has a which stimulate cell growth. Human reduced efficiency of replication at herpesvirus 4 (EBV) expresses a gene 37°C, making it suitable for use as a (BARF-1) that encodes the receptor for candidate live virus vaccine. colony-stimulating factor 1. ColAn 57389 virus (CA57389V) A Colony virus (COYV) A serotype of Great strain of Anopheles A virus in the Island virus in the genus Orbivirus . genus Orthobunyavirus, isolated from mosquitoes. Colorado tick fever virus (CTFV) The type species of the genus Coltivirus . col factors Bacteriocinogens present in Causes disease in humans 4–5 days coliform bacteria, which carry genes after the bite of an infected tick, for colicin production. Dermacentor andersoni, but other tick species may also serve as vectors. Ticks colicins Bacteriocins produced by Esche- become persistently infected, provid- richia coli . ing an over-wintering mechanism. In humans there is fever, usually saddle- coliphage A bacteriophage whose host back type, leukopenia, headache, limb cell is Escherichia coli . pains, and often abdominal pain and vomiting. Rash is uncommon. There collectins A group of soluble proteins may be encephalitis, especially in chil- found in serum, lung, and nasal secre- dren. Virus has been isolated from tions that can provide a first line of wild rodents but in them infection is defense against infectious agents. inapparent. Hamsters can be infected Include mannose-binding protein experimentally i.p. and on passage the which interacts with influenza virus, virus may kill them. Occurs in north- herpes simplex virus, and human west USA but does not reach the Pacific immunodeficiency virus and can acti- coast. Infectivity is acid-sensitive but vate complement to cause virus neu- ether-resistant. The virion, 80 nm in dia- tralization. Collectins are composed of meter, has two protein shells and con- a collagen stalk and a globular head, tains 12 segments of double-stranded usually present as trimers, and form a RNA. Replicates in eggs when injected primitive antibody-like defense mecha- into the yolk sac, and in cells with nism which targets carbohydrate struc- CPE. Control depends on protection tures on infectious pathogens. from ticks. Natural host probably the ground squirrel. Another coltivirus, Epstein J et al (1996) Curr Opin Immunol 8 , 29 Eyach virus has been isolated from ticks in France and Germany, where Colobus papillomavirus A possible spe- it causes febrile illness and neurologic cies in the genus Alphapapillomavirus , syndromes in humans. An isolate of affecting Colobus monkeys, causing CTFV from a hare in California was venereal warts. closely related to Eyach virus. Reszka et al (1991) Virology 181 , 787 Attoui H et al (2000) Biochem Biophys Res Commun 273 , 1121 colonic carcinoma cell line (CaCo2) See Emmons RW (1988) Ann Rev Microbiol 42 , 49 CACO2 cell line . Klasaco R (2002) Med Clin North Am 86 , 435 Coltivirus 104

Coltivirus A genus in the family cascade reaction sequence in response Reoviridae. Spherical virus particles to injury or infection and induce vari- 80 nm in diameter with two outer cap- ous effects including hemolysis, phago- sid shells and a core which possesses cytosis, opsonization and vasodilation. no projections. Genome consists of 12 Acts at an early stage, between innate segments of double-stranded RNA immunity and adaptive immunity. So ranging from 240 bp to 2.53 kb long; called because it complements the act - total mol. wt. 18 1 0 6. Primarily iso- ion of antibodies. Consists of two arms: lated from Ixodid ticks, but also from the classical complement pathway is humans, deer, and small animals. activated by immune complexes for- There are two serotypes: Colorado med when antibody combines with tick fever (North America) and Eyach antigen, or by certain viruses or virus- (Europe). Several isolates made in infected cells. The alternative pathway China and Indonesia are classified in is activated by bacterial endotoxin, fun- the Seadornavirus genus. gal or plant polysaccharides, antigen- IgA complexes, or virus-infected cells. Columbia SK virus A strain of Encephalo- The components of the system are myocarditis virus . designated by numbers, and in the classical pathway complement factors columbid herpesvirus 1 (CoHV-1) C1–C9 are activated sequentially in a An unassigned species in the fam- cascade reaction resulting in lysis of the ily Herpesviridae . A natural and wide- infected cell. In the alternative path- spread infection of pigeons. Not way factor C3 is cleaved and C5–C9 transmissible to chickens, ducks, geese, are formed without a requirement for or quail, but budgerigars and turtle C1, C2, or C4. Some viruses, notably doves can be infected experimentally. poxviruses and herpesviruses, encode Related viruses have been isolated proteins that contain short consensus from falcons, falconid herpesvirus 1, repeats which are present in many and owls, strigid herpesvirus 1. Only complement control proteins, such as one serotype has been found. Causes CD46 (membrane factor control pro- conjunctivitis, respiratory lesions, and tein) and CD55 (decay accelerating fac- focal necrosis of the liver, but is usually tor). These viral homolog bind to C3b carried by apparently normal birds as and C4b, subverting the action of the a latent infection. Replicates with CPE complement system. in primary chicken embryo fibroblasts. Propagated on the CAM, it produces Nash AA and Dutia BM (2005) In Topley & Wilson’s Microbiology and Microbial Infections , plaques and kills the embryo in 4 days. vol. 1, Tenth edition, edited by BWJ Mahy Synonyms : inclusion disease of pigeons and V ter Meulen. London: Hodder Arnold, virus; pigeon herpesvirus. p. 270 Vindevogel H et al (1985) J Comp Pathol 95 , complement fixation test (CFT) A sero- 105 Marlier D and Vindevogel H (2006) Vet J logical test for detection of serum 172 , 40 antibody which binds complement on reaction with antigen. To measure the Common Cold Research Unit A research binding, the amount of complement station established at Salisbury, UK in remaining in the serum is estimated by 1946 which was responsible for many reaction with antibody directed against important advances through experi- membrane antigens of erythrocytes; ments on human volunteers including the reaction leads to lysis of the eryth- the isolation and cultivation of rhino- rocytes and released hemoglobin can viruses in vitro. Sadly it was closed in be estimated spectrophotometrically. 1990. From the amount of complement in the original serum, the amount fixed common cold virus See human rhino- by the virus–antibody complex can be viruses . estimated. complement A system of at least 18 serum complement-mediated lysis A high den- proteins which interact in a complex sity of membrane-bound antibody is 105 concatemeric DNA

required for complement-mediated defective in different genes assist lysis of virus-infected cells, and so this each other by providing the missing defense mechanism is effective late in functions – the assistance is often the infectious cycle. A more efficient unequal with one virus replicating rap- process is antibody-dependent cell- idly and the other very slowly and (2) mediated cytotoxicity (ADCC), which allelic (intragenic) in which mutants requires only small quantities of mem- defective in the same gene produce a brane-bound antibody, and cell lysis is functional gene product by each pro- mediated by neutrophil leukocytes and viding a part of it; this occurs between natural killer (NK) cells. ts mutants of bacteriophage T4 but has not been unequivocally observed complementary base sequence A nucleic among vertebrate viruses. acid sequence which is able to form a perfect hydrogen-bonded duplex, by complementation group A group of G–C and A–T pairing, with the one to mutant viruses with mutations in the which it is complementary. same gene or cistron. They cannot com- complementary DNA (cDNA) DNA plement each other. There should be as transcribed from single-stranded RNA many complementation groups as cis- using reverse transcriptase. When trons but mutations in some cistrons made double-stranded and cloned in a are not always lethal. vector, it is termed a ‘ cDNA clone. ’ complementation map A genetic map complementary RNA (cRNA) A single- constructed on the basis of comple- stranded RNA molecule that is com- mentation experiments. plementary in base sequence to the single strand from which it was tran- complementation test A test to deter- scribed. Most single-stranded RNA mine whether two virus mutants are viruses (except retroviruses) use com- defective in the same cistron. A mixed plementary RNA as intermediates in infection with the two viruses is car- replication. ried out; complementation is positive if complementary strand A single-stranded the virus yield exceeds the sum of the nucleic acid molecule complementary yields in single infection. If no increase in base sequence to the single strand in yield results from the mixed infec- from which it was transcribed. All tion, the two viruses are said to be in single-stranded RNA and DNA viruses the same complementation group. employ complementary strands as intermediates in their replication. COMUL test Complement fixation test Strands which are complementary to for mu rine l eukemia virus antigens. one another can hybridize to form a double strand. Comvax A combination vaccine containing Hemophilus B conjugate complementation Interaction between (meningococcal protein conjugate) and viral gene products or gene functions hepatitis B (recombinant vaccine). in mixed infected cells which results in increased yield of infective virus of one Con A Abbreviation for con canavalin A . or both parental types. Usually consists in one of the viruses providing a gene concanavalin A A lectin from the jack- product which the other requires but is bean, Canavalia ensiformis. It has high unable to make. The genotypes of the affinity for terminal α - d -mannosyl infecting viruses are not altered and residues and can be used for purifying the progeny virus has the genome of glycoproteins. It agglutinates cells of its parent although it may have certain many types and acts as a potent T-cell structural proteins such as coat pro- mitogen. teins, e.g., specified by the other virus. There are two types of complementa- concatemeric DNA Long DNA mol- tion between mutant viruses: (1) non- ecules made by continual repetition of allelic (intergenic) in which mutants a certain basic DNA chain sequence. concatemeric DNA 106

Can be formed during virus DNA rep- Best JM et al (2005) In Topley & Wilson’s lication involving a circular intermedi- Microbiology and Microbial Infections , vol. ate molecule, and is a precursor from 2, Tenth edition, edited by BWJ Mahy and which mature genome DNA is cut and V ter Meulen. London: Hodder Arnold, p. 959 packaged. congenital varicella syndrome (CVS) Thomas CA et al (1968) Cold Spring Harbor Symp 33 , 417 A severe disease of the fetus resulting from maternal infection with human herpesvirus 3 during the first two tri- concatenates Long molecules made mesters of pregnancy. Clinical mani- by continuously repeating one basic festations range from multisystem molecular unit. involvement resulting in death in the neonatal period to dermatomal skin conditional lethal mutants Mutants scarring, limb hypoplasia, or both as which will not replicate under condi- the only defects. The disease is rare tions in which the wild-type replicates, and the overall risk of zoster in infancy but will replicate under permissive following maternal varicella infection conditions, such as in another cell line, in the second and third trimesters of at a different temperature, or as a com- pregnancy is about 2%. ponent of a mixed infection (see com- 9 plementation da Silva O et al (1990) Pediatr Infect Dis J , ). Most studies of virus 854 gene function using conditional lethal mutants have employed temperature- sensitive (ts ) mutants. conglutinin A protein in the collectin family that acts as a first line of defense against infectious agents. congenital infection Infection occurring before birth. May follow a number of Epstein J et al (1996) Curr Opinion Immunol viral infections and is sometimes lethal. 8 , 29 May produce fetal abnormalities, e.g. rubella virus and human herpes virus Congo virus See Crimean-Congo hemor- 5. Some viruses affect particular organs rhagic fever virus . depending on the stage of fetal development at which infection occurs, conjunctival infection The most common while others which are non-cytocidal conjunctival infections are associated may infect every cell in the embryo with adenovirus, coxsackie virus A24, and persist throughout adult life, e.g. and enterovirus 70, and the conjunc- lymphocytic choriomeningitis virus in tiva may represent a primary portal of mice. entry for these viruses. Yin-Murphy M (1984) Prog Med Virol 29 , 23 congenital rubella syndrome Although rubella is usually a trivial childhood exanthem, if infection occurs in utero Connecticut virus (CNTV) An unassigned during the first 3 months of pregnancy, species in the family Rhabdoviridae , 20% of infected infants are born with belonging to the Sawgrass serogroup. one or more multiple severe congenital Isolated from ticks Ixodes dentatus in abnormalities, including neurosensory Connecticut, USA. deafness, total or partial blindness, congenital heart disease and micro- consensus sequence A short nucleotide cephaly with mental retardation. There sequence in which each position repre- may also be bone translucency, retarda- sents the base most often found when tion of growth, hepatospleno megaly, many actual sequences of a genetic and thrombocytopenic purpura. Vacci- element are compared. For example, nation of girls age 15 months against consensus sequences are found at rubella as part of the MMR vaccine has the exon–intron junction in DNA, at resulted in a dramatic fall in congenital preferred splice junctions in RNA, rubella syndrome cases since the vac- and at ribosomal binding sites involv- cine was licensed in 1969 in the USA ing translation initiation. See Kozak and 1970 in the UK. sequence . 107 cordycepin conservative replication A model for combinations of laboratory practices nucleic acid replication which does not and techniques, safety equipment, and occur in nature. The original or parent laboratory facilities. See Code of Safety double strand is preserved intact and Practices . the progeny molecule has both strands newly synthesized. In nature a semi- continuous cell lines Cells of uniform conservative method takes place. morphology which are capable of indefinite propagation in vitro . They contact infection A disease transmitted originate by transformation of primary by close mechanical contact between cell cultures, frequently of tumor tis- organisms, e.g. herpes. sue. Often aneuploid in chromosome number and on injection into an immu- contact inhibition When normal cells nologically compatible animal may come into contact their movements grow into a tumor. Most do not show cease. Cell division is also inhibited, contact inhibition. but since this is dependent on other factors as well, the term ‘ contact inhi- continuous flow centrifugation Centri- bition’ is used to imply loss of move- fugation in a rotor which has a fluid ment only. See density-dependent seal that allows the continuous flow inhibition . of a sample into and out of the rotor Abercrombie M (1979) Nature 281 , 259 while it is rotating at high speed. Used for large-scale purification of viruses. contagious ecthyma virus Synonym for chamois contagious ecthyma virus. contour length The length of a nucleic acid strand measured by electron contagious pustular dermatitis of horses microscopy. virus Synonym for horsepox virus. Convict Creek 107 virus An isolate of Sin contagious pustular dermatitis of sheep Nombre virus, from a rodent ( Peromyscus virus Synonym for Orf virus . maniculatus) trapped in Convict Creek, on the California–Nevada border, USA. contagious pustular stomatitis of horses virus Synonym for horsepox virus. Li D et al (1995) Virology 206 , 973 contagious pustular stomatitis of sheep Copenhagen vaccinia virus A strain of virus Synonym for Orf virus . Vaccinia virus , the first to be completely sequenced. contagium vivum fluidum A name for Goebel SJ et al (1990) Virology 179 , 247 the cause of tobacco mosaic disease, coined by Beijerinck in 1897 for the copia element Transposable DNA seq- ‘ living infectious agent’ which we now uence in Drosophila chromosome that call tobacco mosaic virus. Although closely resembles a vertebrate retrovi- work by Ivanowski in 1892 had also rus sequence. Consists of a central cod- shown that tobacco mosaic disease was ing region (4.7–7 kb) flanked by 276 bp transmissible by filtered sap, he had direct terminal repeats (LTRs). Occur at interpreted this to mean that a bacte- widely separated chromosomal sites, rial toxin was the causative agent. and code for poly A-containing cyto- Beijerinck, working in Delft, was una- plasmic RNAs. ware of Ivanowski’s earlier work in St. Petersburg. Both can be given credit as Dunsmuir P et al (1980) Cell 21 , 575 founders of the discipline of virology. coproantibodies One year after Beijerinck’s work, the Antibodies that can be first animal virus was shown to be fil- detected in feces, e.g. in rotavirus trable by Loeffler and Frosch, working infection. with foot-and-mouth disease virus on Corthier G and Franz J (1981) Infect Immun the island of Riems. 31 , 833 containment levels Four biosafety con- cordycepin 3 -Deoxyadenosine, an inhib- tainment levels (BSLs) are defined as itor of RNA polyadenylation. core 108 core The central part of the virion proteins are N -glycosylated. The M enclosed by the capsid and consisting protein is N -glycosylated in avian of protein and the viral nucleic acid infectious bronchitis virus (the type genome. The core of reovirus, e.g., can species) and in porcine transmissible be isolated by removing one of the two gastroenteritis virus and turkey corona- capsid protein shells, leaving a single- virus, but is O-glycosylated in mouse shelled particle which is transcription- hepatitis virus and bovine coronavirus. ally active. Replication involves synthesis of a complementary (negative) strand RNA of Corfu virus (CFUV) genome length that acts as a template An unassigned species in the Phlebovirus genus, belong- for synthesis of a 3 -co-terminal nested ing to the sandfly fever virus group. set of 5–7 subgenomic mRNAs which have a capped 5 terminus and are Isolated from Corfu Island, Greece from Phlebotomus major . 3 -polyadenylated. Only the 5 -unique region of each mRNA appears to be Rodhain F et al (1985) Ann Inst Pasteur Virol translationally active. Viruses mature 136E , 161 by budding through the endoplasmic reticulum and Golgi membranes, not at cormorant herpesvirus Synonym for the plasma membrane. Coronaviruses phalacrocoracid herpesvirus 1. infect birds and many mammals, including humans, especially the res- corn snake retrovirus (CSRV) First piratory tract, gastrointestinal organs, observed by electron microscopy of and neurological tissues. a rhabdomyosarcoma from a female Siddell SG et al (2005) In Topley & Wilson’s corn snake, Elaphe guttata . A rattle- Microbiology and Microbial Infections , vol. 1, snake fibroma cell line (C-89) could be Tenth edition, BWJ Mahy and V ter Meulen, infected, and some characterization of London: Hodder Arnold, p. 823 the virus was carried out. Coronavirus A genus in the family Clark HF et al (1979) J Gen Virol 43 , 673 Coronaviridae . May require subdivision when more data are available on the species which cause disease in a wide Coronaviridae A family of single- range of birds and mammals. Type stranded RNA viruses, belonging to the species is avian Infectious bronchitis Order Nidovirales. There are two genera: virus (IBV). There are three groups of (1) Coronavirus , type species Infectious species based on features such as the bronchitis virus; and (2) Torovirus , type number and arrangement of nonessen- species Equine torovirus. The virions are tial genes, and the presence or absence pleomorphic, approximately spheri- of a hemagglutinin-esterase protein in cal, 120–160 nm in diameter, covered the virion. The species are: with petal-like projections (peplomers) 12–24 nm long and arranged in a char- Group 1 acteristic fringe giving the appearance Canine coronavirus of a crown (corona) from which the Feline coronavirus family name is derived. Virus is assem- Human coronavirus 229E bled in the cytoplasm and matures by Porcine epidemic diarrhea virus budding through the endoplasmic Transmissible gastroenteritis virus reticulum. Nucleic acid consists of one Group 2 molecule of infectious single-stranded Bovine coronavirus RNA about 30 kb (coronavirus) or Human coronavirus OC43 20 kb (torovirus) in length. Virion RNA Human enteric coronavirus has a 5 terminal cap and a 3 termi- Murine hepatitis virus nal poly A tract. There are at least five Porcine hemagglutinating encephalomyeli- virus-specific polypeptides. All corona- tis virus viruses have spike (S), membrane Puffinosis coronavirus (M), and nucleocapsid (N) proteins Rat coronavirus and some also have a hemagglutinin- Severe acute respiratory syndrome corona- esterase (HE) protein. The S and HE virus 109 Cowbone Ridge virus (CRV)

Group 3 Cotia virus (CPV) An unassigned virus Infectious bronchitis virus in the family Poxviridae . Isolated from Pheasant coronavirus sentinel mice and mosquitoes in the Turkey coronavirus forest of Cotia, São Paulo, Brazil. Isolated from a man in French Guiana, Only two species agglutinate erythro- but not reported to be a significant cytes: Human coronavirus OC-43 and cause of disease in humans. Replicates Porcine hemagglutinating encephalomyeli- in human embryo lung cells as well as tis virus . in several other cell lines. Shares some IBV is not antigenically related to any antigens with vaccinia virus but not other coronavirus. Rabbit coronavirus neutralized by antisera to vaccinia, is a tentative species in the genus. fowlpox, goatpox, myxoma, or tanapox viruses. Cavanagh D et al (1997) Arch Virol 145 , 629 Gonzales JM et al (2003) Arch Virol 148 , 2207 Esposito JJ et al (1980) J Gen Virol 47 , 37 Snijder EJ et al (2003) J Mol Biol 331 , 991 Ueda Y et al (1978) J Gen Virol 40 , 263 Zhang X et al (2007) Virology 358 , 424 Ueda Y et al (1995) Virology 210 , 67 coronavirus-like (CV-like) superfamily cottontail rabbit herpesvirus Synonym See Nidovirales . for leporid herpesvirus 1.

Corriparta virus (CORV) A species in the Cottontail rabbit papillomavirus (Shope) genus Orbivirus, isolated from culicines (CRPV) Type species of the genus and birds in northern Australia. With Kappapapillomavirus. The 72 capsomeres Acado and Jacareacanga viruses forms are arranged with a left-hand skew lat- the Corriparta serogroup. Not known tice. A natural infection of cottontail to cause disease in humans. rabbits, Sylvilagus floridanus . Domestic Carley JG and Standfast HA (1969) Am J rabbits, Oryctolagus, and several spe- Epidemiol 89 , 583 cies of Lepus can be infected by scarification into the skin. Skin warts appear Coryza virus See human rhinoviruses . and regress but may become malignant, more often in domestic rabbits COS-1 cells A line of African green mon- than in cottontails. Serial propagation key kidney cells established from CV-1 in cell cultures has not been reported. cells which were transformed by Although rabbit erythrocytes adsorb an origin-defective mutant of SV-40 the virus they are not agglutinated. virus, so contain SV-40 large T antigen. Synonym : Shope papillomavirus. Grow as fibroblast-like, adherent cell Bryan WR and Beard JW (1940) J Natl Cancer monolayers. Inst 1 , 607 Hopfl R et al (1995) Arch Dermatol Res 287 , Côte d’Ivoire Ebola virus (CIEBOV) 652 A species in the genus ‘Ebola-like Stevens JG and Wettstein FO (1979) J Virol viruses, ’ isolated from a woman who 30 , 891 became ill with Ebola fever after examining a dead chimpanzee in the Tai for- Councilman-like bodies Collections of est, Côte d’Ivoire, in 1994. Genetically eosinophilic necrotic hyaline cells in distinct from other Ebola virus isolates. the livers of yellow fever patients. covalently closed circular DNA Cotia body Inclusion body seen late in (cccDNA) A form of double-stranded infection with Cotia virus. Eosinophilic DNA in which both strands are cir- when stained with hematoxylin and cular, i.e. do not have free ends. Also eosin, but appears as a reddish-purple known as form I DNA. See supercoiled ring with an inner pale blue area when DNA . stained with Giemsa. Morphologically different from those of cowpox and Cowbone Ridge virus (CRV) A species other poxviruses. in the genus Flavivirus , serologically Ueda Y et al (1978) J Gen Virol 40 , 263 belonging to the Modoc subgroup. Cowbone Ridge virus (CRV) 110

No known arthropod vector. Isolated Hu FQ et al (2004) Virology 204 , 343 from a cotton rat in Florida, USA. Not Pandey R et al (1985) Prog Vet Microbiol reported to cause disease in humans. Immun 1 , 199

Cowdry type A inclusion bodies coxsackie-adenovirus receptor (CAR) Intranuclear acidophilic inclusions It was established by in vitro experi- as seen in human herpesvirus 1 and ments that all six serotypes of human 2 infected cells. Cowdry described a coxsackie B virus bound to the same second type of inclusion (type B), also cell surface receptor, a 46 kDa protein, acidophilic and intranuclear, but the as human adenoviruses 2 and 5. These basophilic chromatin of the nucleus is two viruses are unrelated and struc- not marginated. This type was found turally quite different. However when in poliomyelitis, Borna disease and Rift clinical isolates of coxsackie B virus Valley fever, but the term is no longer were examined it was found that cox- used. sackie viruses B1, B3, and B5 may also bind to the complement cascade regu- Cowdry EV (1934) Arch Pathol 18 , 527 lating protein known as decay accelerating factor. cow papillomavirus See bovine papillomavirus 1 . Bergelson JM et al (1997) J Infect Dis 175 , 697

Cowpox virus (CPXV) A species in the coxsackie viruses See human coxsackie genus Orthopoxvirus. Causes papules, viruses . developing into vesicles on a firm inflamed base. Crusting follows and CP81 cells A lymphoid cell line derived may not clear for several weeks. from an orang-utan with spontaneous Lesions appear on teats and udders of myelomonocytic leukemia from which cows. May infect the hands of milk- an Epstein–Barr-like herpesvirus has ers who may then spread the infec- been isolated. tion among cattle. Transmission Rasheed S et al (1977) Science 198 , 407 from humans to humans is rare. A number of infections of humans have CPAE cells (CCC 209) A diploid endothe- been reported in which there was no lial cell line derived from the main obvious contact with cattle. Cases of stem pulmonary artery of a young cow, transmission to humans by domes- Bos taurus . Tests for BVD indicate that tic cats, and transmission from rats the cell line is positive for the virus. to monkeys have been reported. It is likely that the natural host is a small CPE See cytopathic effect . mammal rather than cattle. A survey in the UK found antibodies to CPXV CpG island A region of genome DNA in bank voles, field voles, wood mice and house mice. Pocks on the CAM with a high G C content and a high are intensely hemorrhagic and smaller frequency of CpG dinucleotides rela- at 48 h than those caused by vaccinia tive to the bulk genome DNA. virus. Pocks not produced above 40°C. Variant strains may produce white CPV See Canine parvovirus . pocks. Lesions produced in rabbit skin are large and indurated, with a purple- CR 326 virus A strain of Hepatitis A virus . black center. Replicates in many cell Isolated from patients in Costa Rica lines, some of which, e.g. RK 13, may by the injection of serum or extracts be more sensitive than CAM. Outside of clotted blood into white mustached Europe, lesions in the cow are usually marmosets, Saguinus mystax. Can be due to vaccinia virus. serially passed in marmosets, causing hepatitis. Human convalescent serum Archard LC et al (1984) J Gen Virol 65 , 875 neutralizes the virus. Crouch AC et al (1995) Epidemiol Infect 115 , 185 Carmine C et al (1973) Proc Soc Exp Biol Med Gubser C et al (2004) J Gen Virol 85 , 105 142 , 276, 1257 111 Crimean-Congo hemorrhagic fever group viruses

Provost PJ et al (1975) Proc Soc Exp Biol Med conducted with extreme care. Disease 148 , 532 can be transmitted to Old World and New World monkeys, cats, hamsters, crane herpesvirus See gruid herpes- guinea pigs, and mice. The disease is virus 1 . caused by a prion, which has a very small size; indicated by extreme resist- crane hepatitis B virus (CHBV) An unu- ance to irradiation. Very heat-resistant, sual hepadnavirus isolated from sera of some infectivity surviving 100°C. Not demoiselle cranes (Anthropoides virgo) inactivated by formalin, alcohol, or and gray crowned cranes ( Balearica ether. Following the epidemic of bovine regulorum) from German zoos. Several spongiform encephalopathy in the UK other crane species were negative. in the 1990s, a related disease with a CHBV has a broad host range and is distinct pathology has been detected in only distantly related to avihepadna- humans, and this has been termed new viruses of other species such as Ross ’ variant (nv) CJD. See prion diseases . goose hepatitis B virus and duck Synonym : transmissible virus–dementia hepatitis B virus. However, the virus virus. infected primary duck hepatocytes almost as efficiently as duck hepatitis Brown P et al (2001) Emerg Infect Dis 7 , 6 Chesebro BW (Editor) (1991) Curr Top B virus. Microbiol Immunol 172 , 288pp Prassolov A et al (2003) J Virol 77 , 1964 Holman RC et al (1995) Neuroepidemiology 14 , 174 Crawley virus Prusiner S (Editor) (1999) Prion Biology and A species in the genus Diseases. Cold Spring Harbor: Cold Spring Orthoreovirus . A strain of Avian orthore- Harbor Laboratory Press ovirus . Isolated from chickens with chronic respiratory disease. Synonym : Fahey–Crawley virus. CRFK cells (CCC 94) This cell line was established from the cortical portion of the kidneys of a normal female CRELM test An immunofluorescence domestic cat (10–12 weeks old) and has assay using a set of reagents designed been extensively used in feline virus to detect the possible presence of the research. most serious African hemorrhagic fever viruses, i.e. C rimean-Congo, R ift Valley fever, E bola, L assa, and M arburg. cricetid herpesvirus 1 (CrHV-1) An unassigned virus in the family Herpesviridae . Isolated from a Syrian golden hamster, Creutzfeldt–Jakob disease (CJD) One of Mesocricetus auratus, with a regional the subacute spongiform encephalopa- enteritis which has been considered a thies; a progressive degeneration of the neoplastic disease. Replicates in ham- CNS in humans, with dementia in the ster embryo fibroblast cell cultures with early stages. There may be myoclonus CPE in 1–16 days. Non-pathogenic on and typical EEG changes. Onset usu- injection into adult hamsters and mice, ally between 50 and 65 years. Occurs but fatal for suckling hamsters. sporadically all over the world with an Synonym : hamster herpesvirus. annual incidence of 1 per million population, but small clusters of cases are Tomita Y and Jonas AM (1968) Am J Vet Res reported. Some cases have familial his- 29 , 445 tory. Has been transmitted in humans by corneal graft when the incubation Cri du Chat cells (CCC 90) This cell line period was 18 months, and by contami- was established from the skin of an nated human pituitary-derived growth adult Caucasian female with Cri du hormone with incubation periods of 10– Chat syndrome. The cells show a dele- 30 years. The mode of natural transmis- tion in number 5 chromosome. sion in humans is not known. There is no evidence of increased risk of devel- Crimean-Congo hemorrhagic fever oping the disease in health-care work- group viruses An antigenic group ers. However, postmortems should be within the genus Nairovirus . There Crimean-Congo hemorrhagic fever group viruses 112

are four closely related viruses in this species of the order Crocodylia . The group: Hazara, Khasan, Kodzha, and complete genome sequence suggests Crimean-Congo hemorrhagic fever virus . that this reptile virus is quite distinct They are tick-borne and have been iso- from other chordopoxviruses. Causes lated mostly from ixodid ticks. variable disease symptoms from a non- fatal dermatitis to a more serious dis- Crimean-Congo hemorrhagic fever virus ease with ophthalmia, rhinitis resulting (CCHFV) A species in the genus in asphyxia, and debilitating illness Nairovirus . Belongs to the Crimean- with high mortality. Congo serogroup viruses. Recognized Afonso CL et al (2006) J Virol 80 , 4978 first when it caused an epidemic of acute severe hemorrhagic fever in Crohn’s disease A chronic granulomatous 1944 and 1945 in the western Crimea. disease of unknown etiology, involving Isolated from patients and from the the gastrointestinal tract, particularly tick, Hyalomma marginatum margina- the terminal ileum. It seems to have tum. A similar disease had been known both genetic as well as environmental for many years in the central Asian causes. It has been suggested to have republics of the former USSR and has a link with measles virus infection, since been observed on the borders including measles vaccination, but no of the Black and Caspian Seas, and in convincing evidence has yet been pub- Bulgaria and the former Yugoslavia. lished in support of this hypothesis. A similar virus isolated independently in Zaire and Uganda in the 1960s, cross-hybridization Hybridization bet- called Congo virus, was subsequently ween complementary nucleic acids found to be identical, and the names from different sources. Percentage were combined. Occurs over a wide cross-hybridization can provide a area from South Africa to the Middle measure of the relatedness of two East and Asia. Wildlife hosts in Africa nucleic acids. include large herbivores. Human infections commonly occur in abattoir cross-neutralization Neutralization of workers having contact with infected infectivity of a virus preparation by sheep or cattle tissues. Mortality is high immune sera against different viruses. (10–50%). Usually this indicates a close antigenic and taxonomic relationship between Crimean-Congo serogroup viruses See the viruses involved. Crimean-Congo hemorrhagic fever group viruses . cross-protection Protection conferred on a host by infection with one strain of a virus which prevents infection by Crixivan An alternative name for a closely related strain (see superin- Indinavir, an anti-HIV drug which fection exclusion ). Mild strains of a inhibits proteolytic cleavage of the HIV virus have been used to protect against polyprotein. One of the side effects is infection with a severe, related virus, deposition of adipose tissue in the cen- and this was the basis for vaccinia pro- tral region, termed ‘ Crix belly. ’ tection against variola infection. The phenomenon is also used to assess crmB gene cytokine response modifier relatedness of virus strains. gene B, found in the genome of several species of the genus Orthopoxvirus . cross-reactivation See reactivation . Pickup DJ (1994) Infect Agents Dis 3 , 116 croup-associated virus Synonym for Human parainfluenza virus types 1 and 2 . cRNA See complementary RNA . crowpox virus (CRPV) A tentative spe- crocodilepox virus (CRV) An unclas- cies in the genus Avipoxvirus . sified member of the subfamily Chordopoxvirinae which infects host CRS See congenital rubella syndrome . 113 culture medium cryo-electron microscopy Electron micro- C-type virus particles A term used origi- scopy where the specimen is studied at nally by electron microscopists to cryogenic temperature (the tempera- describe a morphologically defined ture of liquid nitrogen). This technique group of enveloped RNA virus par- enables the sample to be viewed with ticles, often seen outside the cells in minimal distortion and the fewest leukemic tissues. The avian and mam- possible artifacts, and is superior to malian leukemia–sarcoma viruses are negative-staining. C-type particles as are many endog- Adrian M et al (1984) Nature 308 , 32 enous viruses with no known biological function. They are never seen inside the cytoplasmic matrix, but within cryopreservation Preservation of virus or cytoplasmic vacuoles or at the cell sur- cell samples at low temperatures, usu- face from which they bud. Just after ally between –70°C and –180°C. budding they are described as immature C-type particles. They mature crypt cell hyperplasia A common patho- rapidly, the core seems to collapse and logical finding associated with diseases become more electron dense. They of the gastrointestinal tract such as cali- have a diameter of 90–110nm and the civirus or rotavirus infection. core is centrally located. There is a lipoprotein envelope covered with knobs cryptic infections In cryptic infection, 8 nm in diameter, but devoid of promi- e.g. with parvoviruses, the viral life nent projections. The core appears to cycle does not proceed further than have cubic symmetry and to consist nuclear sequestration of the incoming of an outer layer of ringlike subunits virion, and remains completely silent 6 nm in diameter forming a hexagonal with respect to gene expression. pattern, and an inner membrane 3 nm thick. Within this is a tubular structure cryptogram In virology this was pro- which usually appears as a ring but posed as a cipher used to record cer- may fill the core and may have helical tain basic properties of viruses. Each symmetry. cryptogram consists of four pairs of Bernard W (1960) Cancer Res 20 , 712 symbols with the following meanings: Dalton AJ (1972) J Natl Cancer Inst 49 , 323 1st pair: type of nucleic acid and strandedness; 2nd pair, molecular weight cubic viruses Viruses with icosahedral of nucleic acid in millions and per- (cubic) symmetry of their capsid. centage in infective particle; 3rd pair, outline of particle and shape of nucleocapsid; 4th pair, kind of host infected Cuiaba virus A probable species in the and kind of vector. For example, genus Vesiculovirus , isolated from a human polyomavirus D/2:3.4/13: toad in the Amazon region of Brazil. S/S:V/O. Although cited in the litera- Not associated with disease in humans. ture for a number of years, it was not universally accepted, and has dropped Cuiaba-d’Aguilar virus See d’Aguilar out of use. virus . Gibbs AJ et al (1966) Nature 209 , 450 Culicoides A genus of minute biting flies cryptovirogenic Having the potential (midges), dipteran insects in the family to produce infective virus particles Ceratopogonidae, that feed on various after derepression of the viral genome warm-blooded animals and on mosqui- present within the cell. Analogous to the toes. They transmit a number of virus term ‘ lysogenic ’ used for bacterial cells. infections, most notably a large number of species of the Orbivirus genus. CSIRO Village virus (CVGV) A serotype of Palyam virus in the genus Orbivirus . culture medium A liquid or semi-solid Isolated from Culicoides brevitarsis in mixture which supplies the physical Northern Territory, Australia. conditions and substances necessary culture medium 114

for cell growth or maintenance in vitro . adult male African green monkey, All culture media must provide: Cercopithecus aethiops, in 1964, for use in (1) The correct osmotic pressure. This transformation studies on Rous sarcoma is largely due to the concentration virus . of sodium chloride, but other ions and glucose also contribute. C value The amount of DNA (picograms (2) The correct pH. This is usually per cell) in the haploid genome of a obtained with bicarbonate buffer, eukaryotic cell. The DNA content of with up to 5% carbon dioxide in diploid nuclei is the 2C value. The lack the closed head-space above the of correlation between the amount of medium. DNA per cell and the phenotypic com- (3) The necessary inorganic ions: plexity of an organism is termed ‘ the sodium, potassium, calcium, mag- C value paradox. ’ nesium, iron, carbonate, phosphate and sulfate. CX3CR1 A chemokine receptor that is (4) Carbohydrate, usually glucose. involved in the attachment of respira- (5) Amino acids. About 12 are necessary. tory syncytial virus to host cells. (6) Vitamins and growth factors. These may be provided either in the form Tripp RA et al (2001) Nat Immunol 2 , 732 of pure substances or as undefined products such as yeast or embryo CXCR4 A T-cell receptor that can act as extract. a coreceptor (with CD4) for the attach- (7) Peptides and proteins. Some cells ment and entry of human immunodefi- will grow in completely defined ciency virus. media, but for most tissue cultures a supply of serum and peptides is necessary. Most culture media con- cyanogen bromide A chemical which tain phenol red to give visual indi- reacts with methionine, converting it cation of change of pH. Antibiotics to homoserine lactone, splitting the are virtually always included to peptide chain on the C-terminal side of maintain sterility, although nei- each methionine. It is used in studies of ther phenol red nor antibiotics are the structure and amino acid sequence required for cell metabolism. See of proteins. also tissue culture . cyanophage Virus which replicates in Cupixi virus (CPXV) A species in the blue-green algae. genus Arenavirus, belonging to the New World Arenavirus group, iso- cybrid Result of the fusion of a cell with lated from phlebotomine sand flies a cytoplast. The cytoplast can transmit (Oryzomys sp) in the Amazon region cytoplasmic components which may of Brazil. Antigenically related to the not be under the control of the cell Changuinola virus group. Not associ- genome. For example, intracisternal ated with disease in humans. A-type virus particles can be transmit- curing Conversion of a lysogenic bacte- ted to a cell which does not contain rial culture to a non-lysogenic state. them by fusion of the cell with a cyto- Can occur spontaneously or can be plast which does. The result of the induced, e.g., by heating the culture fusion is a cybrid. briefly or by exposure to irradiation. cyclical variations Many virus diseases Curionopolis virus (CUR) An unas- undergo monthly or yearly cyclical signed animal rhabdovirus isolated variations which are not well under- from Culicoides spp. midges in the stood, e.g. influenza occurs almost Amazon region of Brazil. Not associ- exclusively during the winter months, ated with disease in humans. and hepatitis A virus is associated with epidemics every 5–7 years. CV-1 cells (CCL 70) A heteroploid cell line derived from the kidney of an Dowell SF (2001) Emerg Infect Dis 7 , 369 115 cyprinid rhabdoviruses cyclic AMP A compound derived from infect arthropod species. Attempts to ATP by the action of the enzyme ade- infect vertebrates or vertebrate cell nyl cyclase. It is an important regula- lines have failed. More than 230 cypo- tory molecule in higher eukaryotes. viruses have been described. They are grouped into 16 species based on cyclins Proteins (A and B forms are the electropherotype patterns of their known) whose levels fluctuate during dsRNA genome. the cell cycle, reaching a peak at mitosis then falling to zero. They are thought to be responsible for driving cells into cyprinid herpesvirus 1 (CyHV-1) An the G2 phase, and so into mitosis. unassigned virus in the family Herpes- viridae . Isolated from epithelioma of cycloheximide 3-[2-(3,5-dimethyl-2-oxo- carp. Produces specific CPE in cell cul- cyclohexyl)-2-hydroxyethyl]glutarim- tures of a warm water aquarium fish, ide. A glutarimide antibiotic isolated Lebistes reticulatus . from streptomycin-producing strains of Synonyms : carp pox herpesvirus; epi- Streptomyces griseus . A potent reversible thelioma of carp virus; epithelioma inhibitor of protein synthesis but does papillosum of carp virus; fishpox virus. not affect the maturation of ribosomes. Grutzner L (1956) Zentbl Bakt ParasitKde I Active against a wide range of eukary- Abt Orig 165 , 81 otic cells but does not inhibit prokaryotic systems. Synonym : actidione. cyprinid herpesvirus 2 (CyHV-2) An unassigned virus in the family Herpes- cyclo-octylamine A compound struc- viridae which causes necrosis of turally related to amantadine which hematopoietic tissue and anemia with inhibits influenza virus replication in high mortality in goldfish Carassius cell cultures. auratus . Synonyms : goldfish herpesvirus; cyclophilin A (CypA) Cyclophilins hematopoietic necrosis herpesvirus of are intracellular proteins which bind goldfish. cyclosporin (e.g. cyclosporin A). There Goodwin AE et al (2006) Dis Aquatic Organ are 14 known members of the cyclo- 69 , 137 phin family but only cyclophilin A Groff JM et al (1998) J Vet Diagn Invest 10 , plays an essential role in the replication 375 of HIV-1 by binding to the gag protein of the virus. cyprinid herpesvirus 3 (CyHV-3) Greene WC and Peterlin M (2002) Nat Med An unassigned virus in the family 8 , 673 Herpesviridae which causes massive Luban J (1996) Cell 87 , 1157 mortality of koi and other species of cyclophosphamide An alkylating anti- carp. The DNA genome is exceptionally cancer drug which is an immunosup- large at 295 kbp, and related to the pressant, and has been extensively genomes of CyHV-1 and CyHV-2, and used experimentally to inhibit the more distantly to that of channel cat- immune response, especially through fish virus (IcHV-1). inhibiting B-cell division. Synonyms : koi herpesvirus; carp interstial nephritis gill necrosis virus. cyclosporin A An 11-amino-acid cyclic Ilouze M et al (2006) FEBS Lett 580 , 4473 peptide with potent immunosuppres- Waltzek TB et al (2005) J Gen Virol 86 , 1659 sive activity on both the humoral and cellular systems. Widely used in organ and bone marrow transplantation. cyprinid rhabdoviruses There are several rhabdoviruses that affect carp species, Cymevene Se e ganciclovir . including spring viremia of carp virus, swim-bladder inflammation virus, Cypovirus A genus in the family and pike fry rhabdovirus (grass carp Reoviridae the members of which only rhabdovirus). Cysternaviridae 116

Cysternaviridae A name proposed Cytomegalovirus One of three genera in but not adopted for the family the subfamily Betaherpesvirinae , con- Coronaviridae . taining three species, Human herpesvirus 5 (human cytomegalovirus), Cercopithecine herpesvirus 5 , and cytarabine hydrochloride (ara C) 1- Cercopithecine herpesvirus 8 . Tentative d beta, -arabinofuranosylcytosine species in the genus are aotine herpes- hydrochloride. An antiviral and anti- virus 1 and aotine herpesvirus 3. The leukemic agent which inhibits DNA members of the genus are grouped on synthesis. In the body the drug is the basis of the nucleotide sequence converted to araCTP, when it is able similarity of their genome DNAs. to inhibit both DNA polymerase and nucleoside reductase. Its antiviral spec- cytomegalovirus group Synonym for trum resembles that of idoxuridine. Betaherpesvirinae . Rapidly inactivated in vivo. Was used with some success in the treatment of herpes keratitis and severe generalized cytopathic effect (CPE) Alteration in herpes infection, but now replaced by the microscopic appearance of cells acyclovir as the drug of choice. in culture following infection with a virus. May consist of rounding up, cell detachment, cell fusion, production of cytochalasins A group of mold metabo- inclusion bodies, etc. Neutralization of lites that bind to actin filaments and CPE is widely used in serological iden- so stop cell movement. Virus entry tification of viruses. through clathrin-coated pits is sensitive to cytochalasins. cytoplasmic amphibian viruses Synonym for icosahedral cytoplasmic cytochrome c A basic protein which is used deoxyviruses of amphibians. in the electron microscopy of nucleic acids as it binds to them and renders cytoplasmic filament One of three classes them detectable in the electron micro- of filaments found in eukaryotic cells; scope. See Kleinschmidt procedure . actin-containing microfilaments, tubu- lin-containing microtubules, and intermediate filaments. cytocidal Causing cell death. cytoplast An enucleated eukaryotic cell. cytokines Small proteins (5–20 kDa) Used to study dependence of virus rep- secreted by cells which affect the lication on the cell nucleus. growth, differentiation or activation of other cells. They are important non- Cytorhabdovirus A genus of plant rhabdo- antigen-specific effector molecules viruses. The type species is Lettuce which help to mediate the immune necrotic yellows virus . response. Examples are interferons, lymphokines and tumor necrosis factor. cytoryctes variolae The name originally applied by Guarnieri to the inclusion bodies which now take his name, who cytolysis The lysis of cells. was under the impression that they were protozoa. cytomegalic inclusion disease (CID) Diseases caused by cytomegalovirus cytosine arabinoside hydrochloride See infection are marked by characteristic cytarabine hydrochloride . large refractile inclusion bodies (known as ‘ owl eye ’ inclusions). These are cytosis A general term for pinocytosis found in patients suffering from classi- and phagocytosis. cal congenital CMV infection, and also in AIDS patients suffering from CMV, a cytosol The fluid portion of the cyto- frequent opportunistic infection. plasm outside the organelles. 117 cytovene cytotoxic Harmful to cells. A property of directly lyse target cells and are certain chemicals (drugs), viruses and usually CD8 positive, class I MHC cells (e.g. cytotoxic T cells). The viruses antigen-restricted. may be toxic without replicating, or as a result of replicating in the cell. cytovene See gancyclovir . cytotoxic T lymphocytes (CTLs) A subset of effector T lymphocytes which D

D17 cells (CCL 183) A cell line, derived Orthobunyavirus. Isolated from Culex from an osteosarcoma, metastatic to the guiarti and C ingrami in the Ivory lung of an 11-year-old female poodle. Coast. Riggs J et al (1974) J Gen Virol 25 , 21 Institut Pasteur Dakar (1984) Annual Report , p. 99 D1R/D12L A heterodimer composed of the 97 kD DIR gene product and the Dabney’s grippe or grip Synonym for 33 kD D12L gene product, which func- Bamble disease. tions as the mRNA capping enzyme of poxviruses. dactinomycin Synonym for actinomycin D. D4T 2 -3 -Didehydro-2 -3 -dideoxythymi- dine. A potent and relatively selective d’Aguilar virus (DAGV) A strain of inhibitor of human immunodeficiency CSIRO village virus in the genus virus replication in vitro . Orbivirus, belonging to the Palyam serogroup. Isolated from Culicoides brevitarsis in south-east Queensland, D5 virus A virus isolated from a child Australia. Antibodies are present in with summer diarrheal illness which cattle and sheep. Not reported to cause became the prototype for reovirus sero- disease in humans. type 2. Synonym : Jones virus. Dak AN B 188d virus A strain of Ippy Sabin AB (1959) Science 130 , 1387 virus in the genus Arenavirus .

Da Bie Shan virus A strain of Hantaan Dakar bat virus (DBV) A species in virus isolated from the rodent the genus Flavivirus belonging to the Niviventer confucianus in Thailand. Rio Bravo virus antigenic group. No known arthropod vector. Isolated from Pattamadilok S et al (2006) Am J Trop Med Hyg 75 , 994 insectivorous bats of Scotophilus sp in Senegal, Nigeria, Uganda, and Central African Republic. Not known to cause Da Fano bodies Minute basophilic intra- disease in humans. cytoplasmic inclusion bodies found in cells infected with human herpesvirus DakArk 7292 virus (DAKV-7292) An 1 or 2. unassigned animal virus of the family Rhabdoviridae . Isolated from ticks, Da virus A strain of parainfluenza Amblyomma variegatum . Not associated virus type 5 in the genus Rubulavirus . with disease in humans. Isolated from postmortem blood from a case of infectious hepatitis. Calisher CH et al (1989) Intervirology 30 , 241 Antigenically identical to simian virus SV5 and SA virus. DALDs Disability-adjusted life days. Hsiung GD (1959) Virology 9 , 717 DALYs Disability-adjusted life years. dab lymphocystis disease virus A tentative damselfish neurofibromatosis virus species in the genus Lymphocystivirus . (DNFV) A possible species in the family Retroviridae . Virions 90–100 nm in Dabakala virus (DABV) A strain diameter, density 1.14–1.17 g/cm3 , with of Olifantsvlei virus in the genus Mn2 -dependent reverse transcriptase 119 Deer papillomavirus (DPV)

activity, were found in association with DDC 2 ,3 -Dideoxycytidine. An inhibi- transmissible neurofibromas of bicolor tor of human immunodeficiency virus damselfish, Pomacentrus partitus , from replication. In combination with zido- coral reefs in Florida, USA. vudine, better results were obtained Schmale MC et al (1966) J Gen Virol 77 , 1181 in HIV-AIDS patients than with either drug alone. Danazol A synthetic androgen, an ante- Synonym : Zalcitabine. rior pituitary suppressant, that is used Nowak R (1995) Science 269 , 1666 to relieve some of the symptoms of HTLV I-associated myelopathy/tropi- DDI 2 ,3 -Dideoxyinosine. An inhibitor cal spastic paraparesis. of reverse transcriptase activity which has been approved for therapeutic dandy fever virus Synonym for Dengue treatment of AIDS patients with intol- virus . erance of zidovudine. Synonyms : Videx; didanosine. Dane particle The original name for the complete human hepatitis B virus par- Dearing virus A virus isolate, from the ticle first described by David Dane; a stool of a child, which became a proto- 42 nm double-shelled particle with a type for reovirus serotype 3. 7 nm outer shell and a 27 nm inner core. Sabin AB (1959) Science 130 , 1387 Dane D et al (1970) Lancet i , 695 3-deazaguanine A nucleoside analog dark field microscopy A method of which has been found to inhibit rota- microscope illumination in which the virus replication. specimen is seen with light scattered or Smee DE et al (1982) Antimicrob Agents diffracted by it. Chemother 21 , 66

Daudi cells (CCL 213) A B lymphoblast 3-deazauridine A nucleoside analog cell line, derived from a 16-year-old which has been found to inhibit orbi- black male with Burkitt lymphoma. virus replication. The cells are positive for the EBNA and VCA antigens of human herpesvirus 4. Oshiro LS et al (1978) J Gen Virol 39 , 73 Klein E and Klein G (1968) Cancer Res 28 , decay-accelerating factor (DAF) A widely 1300 distributed membrane-anchored protein (CD 55) involved in regulation of the DBM paper Abbreviation for d iazo b en- complement cascade. It appears to func- zyloxymethyl paper, used to bind RNA tion as an echovirus 7 receptor protein. for Northern blotting experiments. Ward T et al (1994) EMBO J 13 , 5070 DBS-FCL-1 cells (CCL 161) A diploid fibroblast-like cell line, derived from Dede cells (CCL 39) A Chinese hamster enzymatically dispersed lung tissue cell line, established from the lung of a 135-day-old normal male African of an adult female Chinese hamster, green monkey fetus. Cricetulus griseus .

DBS-FCL-2 cells (CCL 162) A fibroblast- deer fibroma virus Synonym for Deer like cell line, derived from enzymatically papillomavirus . dispersed lung tissue of a 141-day-old normal male African green monkey Deer papillomavirus (DPV) A species in fetus. the genus Deltapapillomavirus . A natural infection of deer in USA producing DBS-FRhL-2 cells (CCL 160) A diploid tumors which are histologically fibro- fibroblast-like cell line, derived from papillomas. Not transferable to calves, enzymatically dispersed lung tissue rabbits, guinea pigs, or sheep, but of a 248 g normal male rhesus monkey fibroplastic tumors can be produced in fetus. hamsters. Deer papillomavirus (DPV) 120

Synonym : deer fibroma virus. passed at high multiplicity the propor- Groff DE and Lancaster WD (1985) J Virol tion of defective particles may increase. 56 , 85 See defective interfering (DI) virus Sundberg JP and Nielson SW (1981) Can Vet and von Magnus phenomenon . Some J 22 , 385 defective viruses can replicate in a mixed infection with a helper virus. deer tick virus A virus which infects The helper often provides the coat pro- New England deer ticks, Ixodes dam- teins. See phenotypic mixing . mini . Appears to be a strain of Powassan Huang AS (1977) Bacteriol Rev 41 , 811 virus in the genus Flavivirus . Ebel GD and Kramer LD (2004) Am J Trop defensins A family of small arginine-rich Med Hyg 71 , 268 cationic proteins found in vertebrate Ebel G et al (1999) Emerg Infect Dis 5 , 570 phagocytes which are active against enveloped viruses. They can be divided defective interfering (DI) virus Virus based on structural considerations generated by growth at high multi- (cysteine-spacing) into alpha-defensins plicity of infection which can interfere and beta-defensins. Both types exist in with the replication of normal virus humans, but only beta-defensins are (known as ‘ standard ’ virus) and may found in bovine or chicken cells. modify the outcome of disease. The Daher KA et al (1986) J Virol 60 , 1068 term does not include virus particles Hazrati E et al (2006) J Immunol 177 , 8658 in which the complete genome is miss- Yanagi S et al (2007) Clin Microbiol Infect ing. DI virus has four main properties: 13 , 63 defectiveness (inability to grow in the absence of helper virus), dependence degeneracy of the code See codon . (ability to be complemented by and to replicate in the presence of helper delavirdine A non-nucleoside inhibitor virus), interference (causing a decrease of reverse transcriptase which binds in yield of standard virus), and enrich- directly to the enzyme. Effective for the ment (ability to increase the proportion treatment of HIV. of its own yield). Originally discovered Synonym : Rescriptor with influenza virus (see von Magnus phenomenon ), it now appears that any McCance-Katz EF et al (2006) Clin Infect Dis virus may generate defective interfer- 43 , S224 ing particles if passaged at high multiplicity. In all cases examined, the delayed-type hypersensitivity (DTH) A defective virus is found to lack a part state of heightened immune respon- of the genome nucleic acid, but always siveness following previous exposure retains the genome sequences essential to an antigen. On re-exposure to anti- for its own replication. The missing gen there is a cell-mediated response genome functions are provided by the that takes about 24 h to develop, in helper virus. contrast to immediate-type sensitivity, mediated by antibody, which occurs in Roux L et al (1991) Adv Virus Res 40 , 181 minutes. Strahle L et al (2006) Virology 351 , 101 deletion mutant A mutant generated defective viral genome A virus genome by the loss of one or more nucleotides which lacks adequate function in one from the virus genome. or more genes essential for autonomous virus replication. Delgadito virus See Caño Delgadito virus . defective virus Virus which is unable to replicate because it lacks a complete delta 1 protein An outer capsid protein genome. In all virus preparations there of reovirus. are many defective particles and they may interfere with the replication of delta agent Synonym for Hepatitis delta complete particles. When viruses are virus . 121 dengue hemorrhagic fever (DHF) delta antigen A nuclear antigen, first and catalytic activities resemble those described in 1977, that by 1980 was of viroids and satellite viruses found characterized as hepatitis delta virus. in plants, but are unique and distinct Rizzetto M et al (1980) Proc Natl Acad Sci from all other known animal viruses. 77 , 6124 Dempsey cells (CCL 28) A human skin delta herpesvirus Synonym for cercop- cell line, established from a two-and-a- ithecine herpesvirus 7. half-year-old boy with Klinefelter syndrome, that contains a sex chromosome Deltaretrovirus A genus in the family complement of XXXXY. Retroviridae , the type species of which is Bovine leukemia virus. The viruses demyelination The loss of myelin, the are spread by horizontal infection. The lipid sheath surrounding the neuro- genome is 8.3 kb in length, and contains nal axon. It may be associated with a structural genes (gag , pro, pol , and env ) number of different virus infections. as well as nonstructural genes (tax and Fazakerley JK and Buchmeier MJ (1993) Adv rex ). The tRNA primer is tRNAPro. Virus Res 42 , 249 The genus includes human and nonhuman primate viruses; Primate T-lympho- denaturation of nucleic acid Dissociation tropic virus I, human T-lymphotropic of double-stranded molecules into virus 1, and simian T-lymphotropic single strands, caused by high tem- virus I, Primate T-lymphotropic virus II , perature or extremes of pH. Of protein: human T-lymphotropic virus II and any change in the native conformation Simian T-lymphotropic virus II, and without breaking the primary chemical Primate T-lymphotropic virus III and sim- bonds that join the amino acids; may ian T-lymphotropic virus III. Infection is involve breaking noncovalent bonds associated with B- or T- cell leukemias (e.g. hydrogen bonds) or covalent or lymphomas as well as neurological bonds (e.g. disulfide). diseases, and a long latency. No oncogene-containing members of the genus Dendrid A trade name for idoxuridine have been recognized. eye drops. Δ tas virus A replication-defective foamy dendritic cells A system of cells of stel- virus generated by pregenome splicing late or dendritic morphology which with the removal of the ORF-1 intron. are strongly positive for MHC class II Saib A et al (1995) J Virol 69 , 5261 antigen. They are important accessory cells for the primary immune response. DELTA study An early clinical trial of Originally derived from bone marrow, nucleoside reverse transcriptase inhibi- they are present throughout the body tors which showed that a combination in skin, gut, and lung and in peripheral of drugs was more successful than lymphoid organs. They can take up azidodideoxythymidine (AZT) mono- antigens such as viruses and process therapy in delaying disease, heralding them to peptides for presentation by the concept of combination therapy MHC class II molecules. for HIV. dengue fever of cattle virus Synonym for delta virus Synonym for Hepatitis delta Bovine ephemeral fever virus . virus . dengue hemorrhagic fever (DHF) A Deltavirus A genus containing a single severe hemorrhagic disease first identi- species, Hepatitis delta virus . Virions fied in epidemic form in the Philippines are spherical, 36–43 nm in diameter, in 1954 as the result of infection with with an inner nucleocapsid of 19 nm dengue virus types 3 and 4. A large containing the circular negative sense outbreak occurred in Cuba in 1981, fol- single-stranded RNA genome, about lowed by outbreaks in Mexico (1984), 1.7 kb in length. The genome structure Nicaragua (1985), Puerto Rico (1986), dengue hemorrhagic fever (DHF) 122

El Salvador (1987), Venezuela (1989), dengue virus occurs in HeLa cells with and Rio de Janeiro (1990). The disease is CPE and in suckling mouse brain. The believed to result from sequential infec- virus can be adapted to eggs and other tions with different serotypes of dengue cell types. Adapted experimentally to virus and to have an immunopatho- mice, it causes flaccid paralysis of the logic basis, but only 6% of patients with limbs. Most nonhuman primates have sequential infections actually develop an inapparent infection following exper- the severe form of the disease. See imental inoculation. Currently, control dengue viruses 1–4 . of dengue fever relies upon control of Gubler DJ and Meltzer M (1999) Adv Virus the mosquito vector, and despite much Res 54 , 35 developmental research, no vaccine is Halstead SB (1988) Science 239 , 476 available. See also dengue viruses 1–4 . Kurane I and Ennis FA (1992) Semin Immunol Synonyms: breakbone fever virus; 4 , 121 dandy fever virus; polka fever virus. dengue shock syndrome (DSS) Shock Gubler DJ (2006) Novartis Found Symp 277, 3 is one of the severe manifestations in dengue hemorrhagic fever and, if untreated, can cause up to 50% mortal- dengue viruses 1–4 (DENV-1–4) Four ity in affected patients. serotypes of Dengue virus in the genus Flavivirus which together with Kedougou Dengue virus (DENV) A species in the virus form the dengue virus serogroup. genus Flavivirus. Causes an acute febrile Double diffusion tests reveal a common illness in humans with symptoms rang- antigen and specific antigens. Type 1 ing from clinically inapparent to severe occurs in South-East Asia from India fatal hemorrhagic disease. There is an to Japan and Hawaii, with temporary incubation period of 5–8 days, and spread to Greece, South Africa, and the symptoms last about 10 days with Australia. The virus is not endemic in severe headaches, retro-ocular pain, Japan except for a transient outbreak in back and limb pains, nausea and vom- the 1940s. Type 2 occurs in South-East iting. Often there is a scarletiniform or Asia, central America, and the Caribbean. maculopapular rash. There is no specific Types 3 and 4 occur in Thailand and treatment, but analgesics containing the Philippines. Hemorrhagic fever acetaminophen can be used to relieve with dengue shock syndrome prob- pain. The most severe symptoms, hem- ably results from infection with one orrhagic fever with shock, probably type in persons immune to another. An result from infection with one dengue antigen–antibody reaction occurs in the virus serotype in persons immune to tissues and results in increased vascular another (see dengue viruses 1–4). The permeability and leakage of plasma, but natural hosts for the virus are Aedes the molecular basis of the pathological mosquitoes, humans, and nonhuman process is not well understood. No vac- primates. Aedes aegypti is the principal cines are currently licensed, but consid- vector worldwide, but other important erable progress has been made toward vectors are Aedes albopictus in Asia and the development of a tetravalent live the Americas, Aedes scutellaris in the attenuated vaccine. Control is presently Pacific, and Aedes africanus and Aedes aimed at the principal vector species, luteocephalus in Africa. The virus is only Aedes aegypti . transmitted by the bite of an infected Alvarez M et al (2006) Am J Trop Med Hyg 75 , mosquito vector, and cannot be spread 1113 from person to person. Following infec- Gubler DJ (1998) Clin Microbiol Rev 11 , 480 tion, humans and nonhuman primates Gubler DJ and Meltzer M (1999) Adv Virus usually develop a high level of viremia Res 54 , 35 lasting about 5 days, and if a competent mosquito vector takes a blood meal Denhardt’s solution A solution com- during this viremic phase, it becomes prising 0.02% Ficoll, 0.02% polyvinyl infective after 8–12 days and capable pyrrolidone and 0.02% bovine serum of transmitting the virus. Replication of albumin used in the preincubation of 123 deoxyribonucleic acid (DNA)

nitrocellulose and nylon filters. This which selectively inhibits replication of treatment prevents nonspecific binding herpesviruses. in Northern and Southern blots. Colacino JM and Lopez C (1985) Antimicrob Denhardt DT (1966) Biochem Biophys Res Agents Chemother 28 , 252 Commun 23 , 641 6-deoxyacyclovir An acyclic nucle- dense virus particles The virions of oside analog that is absorbed orally Picornaviridae which band at a density and converted by xanthine oxidase to of 1.44 g/ml compared to 1.34 g/ml at form acyclovir. Has been used to treat which standard particles band. Dense chronic hepatitis B infection as well as and standard virions are probably two herpes zoster. configurations of the virion structure. Synonym : desciclovir. Rowlands DJ et al (1975) J Gen Virol 29 , 223 Peterslund NA et al (1987) J Antimicrob Wiegers KJ et al (1977) J Gen Virol 34 , 465 Chemother 20 , 743 Weller IV et al (1986) J Hepatol 2 , 5119 density-dependent inhibition Inhibition

of cell division in tissue cultures due deoxycholate (C 24 H39 O4 Na) (mol. wt. to the presence of neighboring cells, 414.6), an anionic detergent. although other factors are involved. For example, if cell metabolism has ren- 1-deoxynojirimycin An alpha- dered the medium acid, restoration of glucosidase inhibitor with some anti- an alkaline pH will temporarily restore viral activity against HIV. growth. Addition of serum or other growth factors may also have a similar deoxyribonuclease (DNase) A phos- effect. See contact inhibition . phodiesterase enzyme which degrades DNA. There are two types: exonu- density gradient A gradient of a solute in cleases which require a terminus for a solvent used to support macromole- hydrolysis, and endonucleases which cules during their fractionation. Usually may act broadly or on specific base applied to the separation of macromo- sequences (restriction endonucleases). lecular species by centrifugation in a Often refers to DNase I, an endonucle- supporting column of fluid whose den- ase which digests single-stranded and sity is lowest at the top of the tube, and double-stranded DNA to give oligonu- increases toward the bottom where it is cleotides terminating in a 5 -nucleotide, greatest. This technique is particularly and is dependent upon the divalent useful for the fractionation of virus- cation, Mg2 , for activity. DNase I associated macromolecules. See rate digests single-stranded and double- zonal centrifugation and isopycnic stranded DNA to give oligonucleotides gradient centrifugation . terminating in a 3 -nucleotide.

Densovirinae A subfamily of the fam- deoxyribonucleic acid (DNA) A polymer ily Parvoviridae comprised of parvo- of deoxyribonucleotides which is the viruses infecting invertebrate (arthropod) primary genetic material of all cells. A species. very large molecule: mol. wt. 106 –1010 . Adenine (A), cytosine (C), guanine (G), Densovirus A genus of the subfamily and thymine (T) are the four bases char- Densovirinae , family Parvoviridae , con- acteristic of DNA. A and G are purines taining parvoviruses infecting arthro- (with a double-ring structure), and C pods, which replicate without a helper and T are pyrimidines (with only a sin- virus. The virions contain single- gle ring). However, in certain types of stranded DNA, 6 kb in length, which is viral DNA other bases occur, e.g. in T2, either a positive or a negative strand. T4, and T6 bacteriophages cytosine is Siegl G et al (1985) Intervirology 23 , 61 replaced by 5-hydroxymethylcytosine. The nucleotides in DNA are joined by 2-deoxy-2؅-F-ß-arabinosylfuranosyl- diester links in which one phosphoric –1 5-iodocytosine A nucleoside analog acid molecule forms bonds between the deoxyribonucleic acid (DNA) 124

3 and 5 positions of consecutive nucle- DNA, about 4.7 kb in size, and upon otides to form a chain called a ‘poly- extraction the DNA readily forms deoxyribonucleotide.’ The sequence of double strands. Replication is depend- nucleotides in the chain is the primary ent upon a helper virus, adenovirus structure of DNA. In double-stranded or herpesvirus. In the absence of a DNA, the adenine and thymine are helper virus, the AAV genome can be present in equimolecular amounts, as integrated into cellular DNA to estab- are guanine and cytosine (see Chargaff’s lish a latent infection. Subsequent rule). The pairing between bases of the helper virus infection may activate the two chains is highly specific. Adenine latent AAV. All six isolates of adeno- is always hydrogen-bonded to thym- associated virus share a common anti- ine, and guanine to cytosine. Because of gen. Transplacental transmission has this specific base-pairing, the sequences been observed. Species affecting cattle, of nucleotides along the two chains chicken, dog, horse, humans, monkey, are complementary. The two strands and sheep have been described. are of opposite polarity, the 5 end of Berns KI (1990) Microbiol Rev 54 , 316 one chain being opposite the 3 end of Berns KI and Bohenzky RA (1987) Adv Virus the other, and they exist as a double Res 32 , 485 helix. The two helices are usually right- handed (BDNA) but more rarely DNA Dera Ghazi Khan serogroup viruses A can adopt left-handed helical conforma- group of seven antigenically related tions (ZDNA). In addition to the B and tick-borne viruses belonging to the Z conformations, in certain conditions genus Nairovirus : DNA may form single-stranded loops, hairpins or cruciform structures. These Abu Hammad structural forms may play a role as rec- Abu Mina ognition elements in DNA–protein inter- Dera Ghazi Khan actions such as transcription. The exact JD254 conformation and number of residues Kao Shuan per turn depends on the physical condi- Pathum Thani tions. Both single- and double-stranded Pretoria DNA molecules can exist as linear molecules, or circles, and may be coiled or Dera Ghazi Khan virus has been isolated supercoiled, and all these forms can be from ixodid ticks, the remainder from observed in the DNA genomes of vari- argasid ticks. They have not been iso- ous viruses. Single-stranded DNA mol- lated from other arthropods or from ecules are rare in nature but are found vertebrates. Not reported to cause dis- to constitute the genome of viruses such ease in humans. as Circoviridae , Parvoviridae , and bacteriophage Φ X174. Dera Ghazi Khan virus (DGKV) A species in the genus Nairovirus , belonging Watson JD and Crick FHC (1953) Nature to the Dera Ghazi Khan serogroup. 171 , 737 Isolated from a tick, Hyalomma dromedarii, in Pakistan. Not reported to cause deoxyriboviruses DNA-containing disease in humans. viruses. Dermacentor ticks The principal vector of deoxyuridine triphosphate nucleotido- Colorado tick fever virus is Dermacentor hydrolase (dUTPase) A herpesvirus- andersoni , a tick which occurs above specific enzyme which is nonessential 4000 ft in the Rocky Mountains, USA. for growth of virus in vitro . Powassan virus has also been isolated from this species of tick in South Dependovirus A genus in the subfamily Dakota, USA. Parvovirinae, family Parvoviridae . The type species is Adeno-associated virus 2 dermovaccinia virus Strains of Vaccinia (AAV2). Mature virions contain either virus less virulent than others. Produce positive or negative single-stranded opaque white pocks on the CAM. 125 Dhori virus (DHOV) derriengue Mexican name for rabies in Detroit-539 cells (CCL 84) A cell line the vampire bat. derived from a female Caucasian child with Down’s syndrome. Derzsy’s disease A highly contagious parvovirus disease affecting geese and Detroit-548 cells (CCL 116) A cell line Muscovy ducks first reported in China derived from a skin biopsy of a in 1956, and subsequently in many Caucasian female infant exhibiting other countries. See Goose parvovirus . anomalies associated with D trisomy syndrome. desciclovir See 6-deoxyacyclovir . Detroit-551 cells (CCL 110) A cell line derived from the skin of a Caucasian Desert Shield virus (DSV-395) A sero- female embryo which may be consid- type of Norwalk virus in the genus ered to be normal and may serve as Norovirus. Isolated from an outbreak of a control for the study of cells having gastroenteritis amongst US troops dur- mutant chromosomes. ing the 1990–1991 Persian Gulf War. Lew JF et al (1994) Virology 200 , 319 Detroit-562 cells (CCL 138) A heteroploid cell line derived from metastatic Detroit-6 cells (CCL 3) A heteroploid carcinomatous cells in pleural fluid cell line developed from sternal mar- obtained from an adult Caucasian row taken from a human male adult female who had a primary carcinoma with carcinoma of the lung. Now of the pharynx. discontinued because of HeLa cell contamination. Detroit-573 cells (CCL 117) A cell line derived from a skin biopsy of a Detroit-98 cells (CCL 18) A hetero- deceased newborn Caucasian female ploid cell line developed from sternal with multiple congenital abnormalities. marrow taken from a human male adult with no history of malignancy. devil’s clutch Synonym for Bamble Discontinued because of HeLa cell disease. contamination. devil’s grip or grippe Synonym for Detroit-510 cells (CCL 72) A cell line Bamble disease. derived from skin tissue obtained from α α a 9-month-old Caucasian female with dexamethasone 9 -Fluoro-16 -methyl- galactosemia. prednisolone. A synthetic glucocor- ticoid which, when added to certain lines of mammary tumor cells in cul- Detroit-525 cells (CCL 65) A cell line ture, stimulates the rate of produc- derived from skin tissue obtained from tion of mouse mammary tumor virus. a three-and-a-half-year-old Caucasian Probably acts by increasing cell gene girl who exhibited some of the symp- transcription. toms diagnostic of Turner’s syndrome. Dhori virus (DHOV) A species in the Detroit-529 cells (CCL 66) A cell line, genus Thogotovirus , with a genome derived from skin tissue obtained from containing seven negative-stranded a two-and-a-half-year-old Caucasian RNA segments. Isolated from a tick, girl who exhibited symptoms and signs Hyalomma dromedarii , in India, Egypt, of Down’s syndrome. and the Volga River delta in the former USSR. Antibodies are frequently found Detroit-532 cells (CCL 54) A cell line in sera from camels in India. Not derived from the foreskin tissue reported to cause disease in humans. of a 2-month-old Caucasian infant Fuller FJ et al (1989) In Genetics and Patho- with Down’s syndrome. Used for genicity of Negative Strand Viruses , edited by the isolation and propagation of D Kolakofsky and BWJ Mahy. Amsterdam: cytomegaloviruses. Elsevier, p. 279 Dhori virus (DHOV) 126

Leahy MB et al (1997) Virus Res 50 , 215 dibucaine hydrochloride Synonym for Sokhey J et al (1977) Indian J Med Res 66 , 726 cinchocaine.

DHPG See ganciclovir . Dicentrarchus labrax encephalitis virus A possible species in the genus DI Abbreviation for d eaminase i nhib- Betanodavirus isolated from juvenile sea itor. Also an abbreviation for d efective bass. i nterfering. See interference . dichloroflavan A drug that specifi- diabetes mellitus Various viruses have cally inhibits rhinovirus replication, been associated with the onset of juve- by binding to the virion and blocking nile diabetes, including coxsackie virus un coating. See also: disoxaril . B and mumps virus, both of which may infect the pancreas. Diabetes also didanosine (DDI) A nucleoside inhibi- occurs in a significant number of cases tor of retroviral reverse transcriptase, of congenital rubella syndrome. approved as an alternative drug to Craighead JE (1975) Prog Med Virol 19 , 161 AZT for use in adults, or to be used in combination with a protease inhibitor diagnosis In the course of a virus infec- as part of HAART. It can cause pancre- tion one can detect virus, viral antigen, atitis and peripheral neuropathy. nucleic acids, or antibodies to the virus in order to identify the virus. A good didehydrodideoxyuridine (D4T) A knowledge of viral pathophysiology nucleoside inhibitor of reverse tran- and host response is needed to deter- scriptase. See stavudine . mine which specimens to collect and which assays to perform in order to dideoxy nucleotide method A method make a diagnosis. for sequencing DNA based on the ability of DNA polymerase to extend Specter S and Bendinelli M (2005) In Topley & a primer, annealed to the template to Wilson’s Microbiology and Microbial Infections , be sequenced, until a dideoxy chain- vol. 2, Tenth edition, edited by BWJ Mahy terminating nucleotide is incorpo- and V ter Meulen. London: Hodder Arnold, p. 1532 rated. The DNA template is in single- stranded form by cloning, e.g., in M13. dialysis A process of selective diffu- After annealing the primer, four separate reaction mixtures are used, each sion through a membrane. It is usually α used to separate low-molecular-weight containing one -labeled deoxynucle- solutes which diffuse through the oside triphosphate (dNTP), a mixture membrane from the colloidal and high- of unlabeled dNTPs and one chain- molecular-weight solutes which do not. terminating ddNTP, such that each aliq- uot has a different ddNTP. The Klenow fragment of bacterial DNA polymer- 3,8-diamino-5-ethyl-6-phenylphen- ase 1 is added, and after incubation anthridinium bromide See homidium the newly synthesized DNA product bromide . will be terminated at all the positions where a ddNTP can be inserted. In this diarrhea See gastroenteritis viruses of way, each reaction tube forms a nested humans . set of fragments based on the primer and ending in a ddNTP. The fragments diarrhea virus of bovines Synonym for are separated by electrophoresis using pestivirus diarrhea virus. adjacent tracks for the four mixtures, on a high resolution polyacrylamide- diazobenzyloxymethyl (DBM) paper An urea gel, and after autoradiography activated paper used to bind nucleic the DNA sequence of the original tem- acid covalently for hybridization pro- plate can be read directly as a ladder. cedures. Used for binding RNA in Usually sequences up to 350 nt can be Northern blotting experiments. read on a single gel, and a new primer, Alwine JC et al (1979) Methods Enzymol 68, 220 based on the previous sequence, can 127 diphenylamine reaction

be added until the whole of the DNA dihydrofolate reductase An enzyme sequence has been determined. which catalyzes the reduction of Sanger F et al (1977) Proc Natl Acad Sci 74 , dihydrofolate to tetrahydrofolate, 5463 an essential step in purine synthesis. Specified in the genome of some dideoxyadenosine A dideoxynucleoside herpesviruses. in which the 3 -hydroxy group on the 9-(1,3-dihydroxypropoxy)methylgua- sugar moiety is replaced by a hydro- nine (DHPG) See ganciclovir . gen. Rapidly metabolized in vivo to didanosine (DDI) by deamination. 9-S-2,3-dihydroxypropyl adenine A com- DDI is then converted to dideoxyinos- pound reported to inhibit rotavirus ine monophosphate and ultimately to replication. dideoxyadenosine triphosphate, which is a chain-termination inhibitor of DNA Smee DF et al (1982) Antimicrob Agents Chemother 21 , 66 synthesis, and so inhibits HIV. dikkop Means ‘ thick head ’ and is the dideoxyribose A sugar constituent of local name for edematous or cardiac nucleotides which resembles deox- form of African horse sickness. yribose except that it lacks a hydroxyl group at position 3. Thus nucleotides dilution endpoint (DEP) (1) A property of based on it can be incorporated into a virus in which a series of dilutions is a polynucleotide chain but the lack of made of infected material, usually with the hydroxyl group prevents addition phosphate buffer, and the infectivity of of further nucleotides. each dilution measured. The endpoint is the greatest dilution which retains infectivity. DEPs are usually in the range of diethylaminoethylcellulose (DEAE- cellulose) An ion exchange medium 10 1 –10 7 . (2) The greatest dilution of an used in chromatography. It has a p K of antibody which gives a measurable reac- about 9.5 and is used in the chromato- tion with an antigen in a serological test. graphy of proteins and nucleic acids. 3-[2-(3,5-dimethyl-2-oxocyclohexyl)2- diethylaminoethyl dextran (DEAE) An hydroxyethyl]glutarimide See cyclo- anion exchanger, usually used as a col- heximide . umn. DEAE. Sephadex is also available N-N-dimethylamino-2-propanol p-aceta- and has similar properties. midobenzoate See inosiplex .

2,7-bis-[2-(diethylamino)ethoxy]- dimethyl sulfoxide (DMSO) A very fluoren-9-one hydrochloride See tilor- hygroscopic liquid with a slightly bit- one hydrochloride . ter taste and exceptional solvent properties for both inorganic and organic differential host A host for separating chemicals. Its penetrative properties one virus from a mixture of viruses; it have been used to aid the absorption of is susceptible to one virus but not to topically applied drugs such as idoxu- the other(s). ridine. Widely used experimentally as a solvent for substances administered differentiation The differentiation of spe- to cells in tissue culture. cialized functions particular to a cell type by expression of tissue-specific diphasic milk fever Synonym for Tick- genes. Certain viruses require such borne encephalitis virus . tissue-specific expression in order to infect, e.g. via specific cell receptors, diphenylamine reaction A chemical reac- or to replicate – for some viruses no tion that can be used for the colori- in vitro cell culture system exists that metric determination of DNA, e.g. the provides the functions necessary to amount of DNA in a virus preparation. support their replication (e.g. papilloma- See orcinol reaction . viruses, hepatitis B). Burton K (1956) Biochem J 62 , 315 diploid 128 diploid A genome with two homologous severe cases of congenital cytomegalic genome sets. The only examples so inclusion disease. far known amongst vertebrate viruses belong to the Retroviridae . distemper virus Synonym for Canine distemper virus ; hundestaupe; maladie de diplornavirus Double-stranded RNA jeune age. virus. A name once proposed for the family Reoviridae . Ditchling agent See Ditchling virus . direct immunofluorescence An assay for Ditchling virus A Norwalk-like virus, diagnosis of a virus infection in which a very similar or identical to Wollan single fluorochrome-labelled antibody virus. Observed by electron micros- is used. copy in seven or eight fecal specimens collected during an outbreak of acute discontinuous DNA synthesis A simple epidemic gastroenteritis in a school in system of DNA synthesis employed by Ditchling, Sussex, UK. many viruses during replication which consists of the formation of two repli- Appleton H et al (1977) Lancet 1 , 409 cation forks which synthesize DNA in different directions from the origin. D loop Abbreviation for displacement Since all DNA polymerases synthesize or displaced loop. A structure formed DNA only in a 5 to 3 direction, this in supercoiled DNA which has been requires a leading strand and a lag- incubated, under the appropriate con- ging strand which move in opposite ditions, with a short single-stranded directions. DNA or RNA fragment that is homologous for part of the supercoiled mol- Kornberg A and Baker TA (1992) DNA Replication . New York: WH Freeman & Co. ecule. The displaced strand of the supercoiled molecule, the D loop, is disintegrations per minute (dpm) A then available for manipulation, e.g. direct measurement of radioactivity, site-specific mutagenesis. D-loop for- estimated from the counts per minute mation is an initial step in the replica- measured in a radioactivity counter and tion of many circular double-stranded the quenching of counts in the sample. DNA molecules. See R-loop mapping . 1 μ Ci gives 2.2 1 0 6 dpm; 1 Becquerel is one disintegration per second. DMSO See dimethyl sulfoxide . disoxaril A compound that inhibits rep- DNA See deoxyribonucleic acid . lication of enteroviruses and rhinoviruses by binding to the VP1 capsid DNA-binding protein A protein which protein and preventing uncoating. binds to specific DNA sequences such as Rossman MG and Rueckert RR (1987) promoters or enhancers. DNA-binding Microbiol Sci 4 , 206 proteins have many functions, e.g. maintaining DNA in single-stranded form for dispersion factor A measure of the varia- transcription or replication. Many viruses bility in incubation period (the interval encode DNA-binding proteins. from acquisition of infection to onset of illness). If the observed distribution DNA-dependent DNA polymerases (I, of incubation periods is plotted on a II, and III, alpha, beta, gamma) See logarithmic scale, the dispersion fac- DNA polymerase . tor is the antilogarithm of the standard deviation. DNA-dependent RNA polymerase A transcriptase mediating the transference disseminated intravascular coagula- of the information encoded in the base tion (DIC) A serious blood disorder sequence in DNA to an analogous base that has been associated with filovirus sequence in mRNA. The RNA chain is hemorrhagic fever and also occurs in initiated at a specific site on the DNA, 129 DNA microarray

the ‘ promoter region,’ and transcrip- system such as biotin-digoxigenin is tion is terminated at another specific hybridized to the specimen either in site, the ‘ terminator region. ’ These are solution or else in situ . Although used specific signals in the DNA template for many years, this diagnostic method recognized by the transcription appa- has been replaced by methods based ratus. In animal cells, three distinct on nucleic acid amplification by the DNA-dependent RNA polymerases are polymerase chain reaction, which yields found, known usually as forms I, II, and an amplicon that can be sequenced, III (although sometimes as forms A, B, providing more definitive information and C) which mediate the synthesis about the virus in the specimen. from DNA of ribosomal RNA, mRNA, and tRNA (as well as 5S ribosomal DNA insert A DNA sequence inserted RNA and some families of short dis- into a cloning vector. persed repetitive nucleotide sequences), respectively. Only form II, and to a DNA ligase An enzyme involved in minor extent form III, are inhibited by α DNA synthesis, and in repair of single- -amanitin, which binds to the stranded breaks (nicks) in native enzymes. Actinomycin D inhibits the double-stranded DNA, as well as cir- function of all three enzymes by bind- cularization of DNA. Catalyzes the ing to the DNA template. Some viruses, formation of a phosphodiester bond notably the poxviruses, contain their between the 5 -phosphate end of one own virus-encoded, virion-bound, nucleotide and the 3 -OH group of DNA-dependent RNA polymerase another. Requires ATP or NAD as a which mediates the synthesis of virus- cofactor. Ligases are present in both specific mRNA early in infection. animal and plant cells, including bacte- Synonyms : DNA transcriptase; RNA ria. Cf. DNA polymerase and see semi- transcriptase. conservative replication . DNA endonuclease See deoxyribonu- Lehman IR (1974) Science 186 , 790 clease , endonuclease, and restriction endonucleases . DNA methylase An enzyme that catalyzes specific methylation of DNA DNA exonuclease See deoxyribonucle- bases. Occurs in most organisms, and ase, exonuclease . involves the enzymatic transfer of the methyl group of the S -adenosyl- DNA gyrase A topoisomerase type II methionine to specific bases in DNA. present in bacteria which introduces Functions after incorporation of the negative superhelical twists into base into the polynucleotide strand. In eukaryotic cells, the most common relaxed closed circular DNA molecules. Several DNA phages require the action methylated base is 5 -methyl-cytosine. of this enzyme during replication of their DNA. DNA microarray A technique for analyzing the expression of multiple genes Denhardt DT (1979) Nature 280 , 196 in a cell or tissue sample, frequently to compare a normal sample with a virus- DNA helicase A DNA unwinding infected or diseased (e.g. cancerous) enzyme – a single-stranded DNA- one. Known also as functional genomic dependent nucleoside 5 -triphosphatase analysis. The probe DNAs can be involved in DNA replication. Has cDNAs or oligonucleotides which are ATPase activity and hydrolyzes two mol- arrayed on a solid surface such as glass, ecules of ATP per DNA base pair broken. plastic, or a silicon chip. mRNA from Kuhn B et al (1978) Cold Spring Harbor Symp the specimens to be compared is labeled 43 , 63 with a fluorophore such as Cyanine 5 (red) or Cyanine 3 (green), and after DNA hybridization A diagnostic tech- hybridization the relative intensities are nique in which a DNA molecule labeled visualized in a microarray scanner to radioactively or with a signal-generating identify which genes are upregulated DNA microarray 130

or downregulated as a result of disease is only found in mitochondria and func- or infection. These analyses provide a tions in the replication of mitochondrial great deal of information, but must be DNA. (4) DNA polymerase delta has interpreted with caution. For example, a high molecular weight and contains some probes may cross-hybridize to the associated 3 -exonuclease activity. Many wrong mRNA, yielding false-positives viruses induce new DNA polymerase or false-negatives. Microarray analysis activity upon infection (e.g. adeno- or has been used to study the pathologi- herpesviruses) whilst others, notably cal changes in monkeys infected with parvoviruses, use existing host cell the SARS coronavirus and the effects of enzymes. Retroviridae contain a differ- highly pathogenic influenza viruses in ent type of DNA polymerase which is mouse or monkey models. RNA template-dependent. See reverse transcriptase . De Lang A et al (2007) PLoS Pathog 3 , e112 Fornek JL et al (2007) Adv Virus Res 70 , 81 Wickner SH (1978) Annu Rev Biochem 47 , 1163 Weller SK (2005) In Topley & Wilson’s DNA polymerase One of a number of Microbiology and Microbial Infections, vol. enzymes involved in DNA replication 1, Tenth edition, edited by BWJ Mahy and which catalyze the addition of deox- V ter Meulen. London: Hodder Arnold, p. 147 yribonucleotide units to a DNA chain by forming a phosphodiester bond DNA polymerase δ A cellular enzyme between the 3 -OH group on the grow- that is involved in the DNA synthesis ing end of the DNA molecule (known as of parvoviruses and human papilloma- the primer strand) and the 5 -phosphate virus during their replication. on the incoming deoxyribonucleotide. The reaction requires a template Ni TH et al (1998) J Virol 72 , 2777 (DNA or sometimes RNA), all four α deoxyribonucleoside triphosphates DNA polymerase -primase A cellular and a primer with a free 3 -hydroxyl enzyme that is involved in the syn- group. The primer can be an uncom- thesis of SV40 virus, human papillo- pleted DNA strand, but is more usu- mavirus, and bovine papillomavirus 1 ally a short RNA strand. Synthesis of at during their replication. It consists of least one strand is discontinuous and four subunits, and the primase activ- yields a series of Okazaki fragments. ity synthezises short RNA primer Three types of DNA polymerase have molecules. been described in prokaryotes. DNA Taneja P et al (2007) Biochem J 407 , 313 polymerase I (Kornberg enzyme) has a 5 -3 exonuclease activity as well DNA probes Molecules of DNA whose as polymerase activity and is mainly sequence is specific for a particular involved in repair synthesis of DNA. virus nucleic acid and can be used, e.g. DNA polymerase II has a 3 -5 nucle- by hybridization, to detect the presence ase activity but lacks the 5 -3 nuclease of the virus nucleic acid in a sample, activity. DNA polymerase III has both e.g. a PCR product. 5-3 and 3 -5 exonuclease activities and appears to be the true DNA replicat- DNA provirus A duplex DNA copy of ing enzyme in Escherichia coli; the role a virus nucleic acid (usually a retro- of DNA polymerase II is uncertain. In virus) that is integrated into the host eukaryotic cells, four distinct species of cell genome and is transmitted from DNA polymerase have been described. one cell generation to another without (1) DNA polymerase alpha-primase has causing lysis of the host. a high molecular weight and contains associated DNA primase activity; it is DNA relaxing enzyme A topoisomerase the major enzyme involved in DNA rep- that catalyzes the conversion of super- lication. (2) DNA polymerase beta is a helical DNA to a non-superhelical cov- low molecular weight enzyme, present alently closed form. in cell nuclei, the function of which is unknown. (3) DNA polymerase gamma Wang JC (1971) J Mol Biol 55 , 523 131 Dobrava-Belgrade virus (DOBV)

DNA replication See DNA polymerase . DNA topoisomerase An enzyme that catalyzes the breaking and rejoining DNA replicons Mobile autonomously of DNA strands, allowing the strands replicating DNA molecules. May to pass through one another and alter- include plasmids, insertion sequences, ing the topology of the DNA. Type 1 transposons, or retrotransposons. activity catalyzes the removal of superhelical turns and the linking of single- Syvanen M (1994) Annu Rev Genet 28 , 237 stranded rings of complementary DNA sequences. Type 2 activity catalyzes the DNA–RNA blot hybridization See removal or introduction of superhelical Northern blotting . turns and the linking of double-stranded DNA rings. A type 1 DNA topoisomerase activity is found in poxviruses, and DNA sequencing Determination of is associated in vaccinia virus with a 314 the sequence of nucleotides in DNA amino acid polypeptide that is essential or in RNA which has been reverse- for virus replication and probably plays transcribed into DNA. The DNA is a role in transcription and replication of cloned, treated by base-specific chemi- the virus genome DNA. See also DNA cal reactions and the products sepa- gyrase and DNA relaxing enzyme . rated by size on sequencing gels then read as a sequence of bases: A, C, G, DNA transcriptase Synonym for DNA and T. The Maxam–Gilbert method dependent RNA polymerase. involves chemical cleavages at specific bases. The DNA is end-labeled, e.g. using polynucleotide kinase, and after DNA tumor viruses Many of the known completion of the four specific cleavage DNA-containing viruses have been reactions the products are separated on associated with oncogenesis. a polyacrylamide gel. This method is labor-intensive and more difficult to DNA vaccines A promising strategy for perform than the Sanger method. Here, vaccine development which has so the DNA to be sequenced is cloned in far been disappointing. Virus DNA is phage M13 in single-stranded form inserted into a plasmid which can be and a specific primer is annealed to the injected directly into the host as naked sequence immediately 5 to the cloned DNA, free of proteins. The DNA is taken insert. A DNA polymerase reaction is up into host cells, which may express then carried out using the Klenow frag- the encoded proteins, which enter the ment of bacterial DNA polymerase 1 MHC class I pathway and stimulate a and four reactions, each containing four strong cell-mediated immune response. deoxyribonucleotides (one of which is Although candidate DNA vaccines have labeled by a fluorescent or radioactive shown promising results with influenza, marker) and one of the four dideoxyri- Venezuelan encephalitis, and hepatitis B bonucleotides (A, C, G, or T). During viruses in animal models, they have not DNA synthesis, incorporation of a been brought to the market. dideoxyribonucleotide prevents further Srivastava IK and Liu M (2003) Ann Intern chain elongation, and reaction condi- Med 138 , 550 tions are chosen such that the DNA is terminated at all possible positions. DNase See deoxyribonuclease . The four reaction mixtures each yield a nested set of fragments of increasing Dobrava-Belgrade virus (DOBV) A spe- length, and after separation on adjacent cies in the genus Hantavirus . Two lanes of a polyacrylamide gel the DNA viruses that are closely similar and can sequence (up to 350 nt) can be read be considered a single virus species. directly. Dobrava virus was isolated from the Maxam AM and Gilbert W (1977) Proc Natl yellow-necked fieldmouse, Apodemus Acad Sci 74 , 560 flavicollis, in Slovenia, and Belgrade Sanger F et al (1977) Proc Natl Acad Sci 74 , virus came from a fatal case of hemor- 5463 rhagic fever with renal syndrome in the Dobrava-Belgrade virus (DOBV) 132

former Yugoslavia. Dobrava virus has dolphin rhabdovirus Isolated from a also been found in the striped mouse, white-beaked dolphin, Lagenorhynchus Apodemus agrarius, which appears to albirostris, that had beached on the be a reservoir of the virus in Central Dutch coast. Has rhabdovirus morphol- Europe. ogy and is antigenically distinct from Avsic-Zupane T et al (1995) J Gen Virol 76 , rabies and vesicular stomatitis viruses. 2801 Neutralizing antibodies were found in Nemirov K et al (1999) J Gen Virol 80 , 371 several European marine mammal spe- Sibold C et al (2001) J Med Virol 63 , 158 cies, including other species of dolphin and seal. DoCl1 (S؉L2) cells (CCL 34.1) The DoCl1 Osterhaus ADME et al (1993) Arch Virol cell line is one of four canine lines 133 , 189 established from focus-derived clones. A clonal substrain of MDCK (ATCC Don cells (CCL 16) Diploid cells from a CCL 34) was infected by murine sar- normal 8-month-old adult male Chinese coma virus (MSV) in the absence of the hamster, Cricetulus griseus . associated RD-114 helper virus to yield the parent foci. Provides a non-murine Dorcopsis wallaby herpesvirus A strain cell host system for studies on the of macropodid herpesvirus 2. interaction of RNA tumor viruses with mammalian cells. dot–blot hybridization A single diagnostic procedure in which the sample to be doctor fish virus (DFV) A strain of analyzed (e.g. extracted cellular DNA) Santee-Cooper ranavirus in the genus is blotted directly on to a membrane Ranavirus . (usually nitrocellulose) and hybridized with a series of reference probes (dots) dog papillomavirus See canine papillo- prepared from cloned virus-specific mavirus 2 . DNA. The probes may be radioactively or chemically labeled and, after treat- dolphin morbillivirus (DMV) A mor- ment, a signal is detected where hybrid- billivirus, isolated from a striped dol- ization occurs. Used extensively for phin, Stenella coeruleoalba , stranded on the coast of Spain in 1990 during a screening tissues for papillomaviruses. distemper epidemic among dolphins Wickenden C et al (1985) Lancet i , 65 in the Mediterranean. See Cetacean morbillivirus . double membrane vesicles Vesicles Barrett T et al (1993) Virology 193 , 1010 induced in cells infected by arteri- Bolt G et al (1994) Virus Res 34 , 291 viruses, which are probably derived Bolt G et al (1995) J Gen Virol 76 , 3051 from the endoplasmic reticulum, and are the site of accumulation of dolphin poxvirus (DOV) An unassigned replicase subunits, including the RNA- virus in the family Poxviridae . Identified dependent RNA polymerase and heli- morphologically in lesions observed case enzymes. on the skin of captive and free-ranging Pedersen KW et al (1999) J Virol 73 , 2016 bottle-nosed dolphins, Tursiops truncatus , dusky dolphins, Lagenorhyncus obscurus , double-stranded (ds) nucleic acid Nucleic white-sided dolphins, Lagenorhyncus acid with two antiparallel strands acutus, and Burmeister’s porpoises, hydrogen-bonded together. The pairing Phocoena spinipinnis , in waters off the between the bases of the two strands is coast of Peru. The lesions consist of black specific. In DNA, adenine (a purine) is punctiform stippled patterns termed hydrogen-bonded to thymine (a pyri- ‘ tattoo lesions. ’ Convalescent dolphin midine), and guanine (a purine) to cyto- sera react with the poxvirus present in sine (a pyrimidine). As a consequence, the lesions as observed by immunoelec- the sequences of nucleotides along the tron microscopy, but the virus has not two chains are complementary. Usually been isolated in cell culture. both strands are either DNA or RNA van Bressem MF et al (1993) J Wildl Dis 29, 109 but RNA:DNA duplexes can occur. The 133 duck adenovirus 1 (DAdV-1)

genomes of viruses of several groups driving sickness virus Synonym for are either double-stranded DNA or jaagsiekte virus. double-stranded RNA. See double- stranded viruses . DRM Detergent-resistant membranes obtained during the study of lipid double-stranded viruses RNA-containing membrane rafts by extracting cells with viruses: Reoviridae and Birnaviridae . cold nonionic detergents such as Triton DNA-containing viruses: Adenoviridae , X-100 or NP-40. Also called detergent- Asfarviridae , Herpesviridae , Iridoviridae , insoluble glycolipid-enriched com- Papovaviridae , and Poxviridae . plexes (DIGs). The assembly of many nonicosahedral enveloped viruses Douglas virus (DOUV) A strain involves lipid membrane rafts. of Sathuperi virus in the genus Briggs JAG et al (2003) J Gen Virol 84 , 757 Orthobunyavirus , belonging to the Simbu serogroup. Isolated from a droplet transmission An important bovine animal and culicoid flies in mechanism for transmission of respira- Northern Territory, Australia. tory viruses. Aerosolized droplets are deposited at different levels in the res- Downie body Large acidophilic cytoplas- piratory tract depending on their size: mic inclusion bodies found in cowpox those over 10 μ m are deposited in the virus lesions. nose, those 5–10 μ m in the airways, and those less than 5 μ m in the alveoli. Doxorubicin A cytotoxic anticancer drug derived from Streptococcus peucetius . Used dsRNA adenosine deaminase activity as a therapeutic agent against Kaposi’s See DRADA . sarcoma. Also called Adriamycin. D-type virus particles A term used DPC-1 virus An endogenous retrovirus originally to describe the 30 nm isolated from the agouti, Dasyprocta non-enveloped particles seen in punctata . Not related by nucleic large numbers in cell nuclei from acid hybridization to other known polyomavirus-infected mice and ham- retroviruses. sters. They proved to be polyomavirus Sherwin SA et al (1979) Virology 94 , 409 particles and the term passed out of use. However, in their proposed classi- DPSO 114/74 cells (CCL 194) A fibroblast- fication of Retroviridae Dalton et al sug- like cell line derived from the lung tis- gested a genus oncornavirus D, with sue of a caesarean-derived squirrel Mason–Pfizer monkey virus as the type monkey. species. This has resulted in the Mason– Pfizer virus and morphologically similar DRADA D ouble-stranded R NA a denosine viruses being described as having type d eaminase activity. A cellular enzyme D morphology, and the establishment activity present in brain cells which is of the genus Betaretrovirus, with Mouse capable of altering viral RNA sequences mammary tumor virus as the type spe- by introducing clustered transitions into cies. There are at least five such viruses: double-stranded RNA. Has been associ- Mason–Pfizer monkey virus , Mouse mam- ated with alterations in measles virus mary tumor virus , Squirrel monkey retro- RNA sequences found in subacute scle- virus , Langur virus and Jaagsiekte sheep rosing panencephalitis (SSPE). retrovirus . Billeter MA et al (1994) Ann NY Acad Sci Colcher D et al (1977) Proc Natl Acad Sci 74 , 724 , 367 5739 Dalton AJ et al (1974) Intervirology 4 , 201 dragon grouper nervous necrosis virus Fine D and Schochetman G (1978) Cancer (DGNNV) A tentative species in the Res 38 , 3123 genus Betanodavirus . duck adenovirus 1 (DAdV-1) A strain drift See antigenic drift . of Duck adenovirus A in the genus duck adenovirus 1 (DAdV-1) 134

Atadenovirus causing egg drop inoculation. Distinct antigenically from syndrome (EDS). First described in 1976 astroviruses of chickens or turkeys. as a disease of laying hens and referred Gough RE et al (1985) Avian Pathol 14 , 227 to as EDS 76 virus in many publications. A widespread natural infection of ducks and geese, which show little evi- duck circovirus (DuCV) A tentative spe- dence of disease. In chickens the virus cies in the genus Circovirus . replicates in nasal mucosa and subsequently in lymphoid tissues and in the duck embryo cells (CCL 141) A cell pouch shell gland, resulting in eggs line derived from whole Pekin duck with abnormal thin shells and a gen- embryos. eral loss in egg production. The virus is transmitted vertically in chicken flocks, duck enteritis virus Synonym for anatid and may remain silent until the birds herpesvirus 1. approach peak egg production. Isolated from chickens worldwide, which prob- Duck hepatitis B virus (DHBV) The type ably became infected through a con- species of the genus Avihepadnavirus . A taminated vaccine. Infection also occurs natural infection of ducks first found in some flocks by direct contact with in domestic ducks, Anas domesticus wild or domestic ducks or geese. Control platyrhynchos , in the People’s Republic is by eradication through slaughter of of China in 1979. Found in ducks of antibody-positive birds or by immuni- Anas sp worldwide except Muscovy zation with an oil adjuvant-inactivated ducks, Cairina moschata, which cannot vaccine applied at 3–4 months of age. be infected experimentally. Infection of Sequencing of the genome of DadV domestic ducks is usually acquired con- shows significant homology with bovine genitally leading to a chronic viremia. adenovirus 4–8 and ovine adeno- Geese may also carry the infection. The virus isolate 287, but a genome organi- virus has provided a useful experi- zation different from members of the mental model for molecular biological Mastadenovirus or Aviadenovirus genera. studies of hepadnavirus infection and A new genus, Atadenovirus , was created replication. Candidate DNA vaccines to accommodate the duck, bovine, ovine, are under development. and possum viruses. Seven other viruses, including several reptile viruses, are ten- Schultz U et al (2004) Adv Virus Res 63, 1 tative species in the genus. Miller DS et al (2006) Virology 351 , 159 Synonym : EDS virus. duck hepatitis virus 1 and 3 (DHV-1 Benko M and Harrach B (1998) Arch Virol 143/4 , 829 and 3) Unassigned viruses in the fam- McFerran JB and Adair BM (1977) Avian ily Picornaviridae, often referred to as Pathol 6 , 189 ‘ enteroviruses; ’ there are two serotypes, 1 and 3. Cause a widespread infection duck adenovirus 2 (DAdV-2) A tenta- of ducks with disease in ducklings. tive species in the genus Aviadenovirus . There is hemorrhagic necrosis of the Serologically unrelated to duck adeno- liver and high mortality. Older birds virus 1. Isolated from Muscovy ducks are immune. Virus is present in the in France. excreta. Birds hatched from the eggs of immune birds resist infection. Virus Bouquet JJ et al (1982) Avian Pathol 11 , 301 replicates in hens’ eggs, killing the embryo, but becomes attenuated on Duck astrovirus (DastV-1) A species in passage and may be used as a vaccine the genus Avastrovirus , associated with for ducklings. Multiplies without CPE hepatitis in ducklings, and originally in chick cell cultures, but with CPE called duck hepatitis virus type 2. There in duck cell cultures. Experimentally, is one known serotype (duck astrovirus other birds and mammals are resist- 1) identified by electron microscopy in ant. Type 1 virus was isolated from an liver suspensions. Replicates in embry- outbreak of disease with high mor- onated chicken eggs following amniotic tality in white Pekin ducks on Long 135 Duvenhage virus (DUVV)

Island, NY, USA but is now known to Republic, and Senegal. Has been iso- be distributed worldwide. Type 3 virus lated from humans with a febrile infection is less severe and has only illness. been seen in the USA. On the basis of genome sequence data it has been Duke’s disease A childhood exanthem suggested that duck hepatitis type 1 also called Filatow-Dukes’ disease. should be assigned to a new genus of Designated fourth disease in a series the Picornaviridae . of six skin rashes by Clement Dukes in 1900. Probably caused by an entero- Kim MC et al (2006) J Gen Virol 87 , 3307 virus, although some reports claim that Tseng, C-H et al (2007) Virus Res 123 , 190 it does not exist, or that it is due to infec- Woolcock PR and Fabricant J (1991) In tion with Staphylococcus aureus strains Diseases of Poultry, Ninth edition, edited by BW Calnek. Ames: Iowa State University that make an epidermolytic toxin. Press, p. 597 Duncan’s disease A genetic immunode- duck hepatitis virus 2 See Duck ficiency disorder (X-linked lympho- astrovirus . proliferative syndrome) characterized by defective immune responses to Epstein–Barr virus (EBV). Fulminant duck/Hong Kong/D3/75 virus A prob- infections, mononucleosis or fatal able species in the genus Avulavirus . B-cell malignancies can result from An avian paramyxovirus isolated EBV infection. from chickens, ducks, and geese in Hong Kong. Unrelated antigenically to Bar RS et al (1974) N Engl J Med 290 , 363 Newcastle disease virus and the other avian paramyxoviruses. Duncan syndrome See Duncan’s disease .

Alexander DJ et al (1979) Arch Virol 60 , 105 dunkop Means ‘ thin-head ’ and is the local name for the pulmonary form of duck infectious anemia virus A strain of African horse sickness. avian reticuloendotheliosis virus. duovirus Synonym for rotavirus. Name refers to the double-shelled construc- Duck parvovirus A species in the genus tion of the virus particle. Dependovirus. There are two strains, Barbarie duck parvovirus and Muscovy duplex duck parvovirus. Although they are Having two parts. Often used to not helper-dependent viruses, phy- describe the double strand of nucleic logenetic analysis places both the duck acid. and goose parvoviruses in this genus containing mostly helper-dependent dUTPase An enzyme which hydro- viruses. lyzes dUTP to dUMP, making dUMP available for conversion to dTMP by thymidylate synthetase. A dUTPase is duck plague herpesvirus Synonym for encoded by several herpesviruses, but anatid herpesvirus 1. the gene does not appear to be essential for growth in cell cultures in vitro . duck plague virus See anatid herpesvirus 1 . Preston VG and Fisher FB (1984) Virology 138 , 58 duck spleen necrosis virus A strain of Duvenhage virus (DUVV) A species in avian reticuloendotheliosis virus. the genus Lyssavirus . First isolated in South Africa from the brain of a Dugbe virus (DUGV) A species in the farmer who had been bitten on the lip genus Nairovirus. With Nairobi sheep by a bat and died after a rabies-like ill- disease virus forms the Nairobi sheep ness. Negri bodies were present in the disease serogroup. Isolated from cattle patient’s brain tissue and in the brain and ticks in Nigeria, Central African of experimentally infected mice, but Duvenhage virus (DUVV) 136

rabies antigen was not demonstrable Amengual B et al (1997) J Gen Virol 78 , 2319 in the brain tissue by immunofluo- Meredith CD et al (1971) S Afr med J 45 , 767 rescence. In 1985, a bat biologist in Roine RO et al (1988) Brain 111 , 1505 Finland died following infection with Duvenhage virus, and it became clear dwarf gourami iridovirus (DGIV) A that the virus is widely distributed in strain of Infectious spleen and kidney necro- bats in Europe. It has been isolated fre- sis virus in the genus Megalocytivirus . quently from bats, Eptesicus serotinus , The virus was isolated from an orna- in Europe and from a bat in Zimbabwe, mental fish species, dwarf gourami Nycteris thebaica . Antigenically related (Colisa lalia ), several of which died in to rabies virus and resembles labora- aquarium shops in Australia in 2004. An tory or ‘ fixed ’ strains of the virus. Can outbreak of disease in farmed Murray be distinguished by in vivo neutraliza- cod (Maccullochella peelii peelii ) seems to tion and cross-challenge tests in mice, have been caused by this virus. or less easily by CF and fluorescent Go J et al (2006) Mol Cell Probes 20 , 212 antibody tests. Kills mice after a short Sudthongkong C et al (2002) Dis Aquat incubation period with inflammatory Organ 48 , 163 infiltration of brain parenchyma and small intraneuronal inclusion bodies. Grows readily in BHK21 cells. E

E1A protein The first protein expressed Eastern equine encephalitis virus (EEEV) when adenovirus DNA enters the cell A species in the genus Alphavirus , first nucleus. E1A protein acts as a master isolated from the brain of an affected protein to modify the host cell environ- horse in 1933. Maintained in the ment and to assist in the transcription wild as a harmless infection of birds, of other early genes. small rodents, reptiles, and amphibia. Causes encephalitis in humans, horses, E1 glycoprotein A protein on the surface pigeons, and pheasants. Man and of alphaviruses which lies lateral to the horses are infected by mosquito bites. surface membrane. The E2 glycopro- In horses there is viremia and fever tein projects outwards from the virion followed by involvement of the CNS. to form the spikes. Strains vary in virulence but mortality can be 90%. In humans the disease E protein Abbreviation for envelope is similar, with high mortality, 30–50%. protein. Most survivors show paralytic or mental sequelae. There are about five E1 region A region mapping at the left- human encephalitic cases a year in the hand end of the conventional genome USA with more than 100 inapparent of adenovirus which encodes some 10 infections determined serologically. polypeptides required for cell trans- Outbreaks with high mortality occur formation by adenovirus. Divided into in pheasants, and outbreaks have been two domains: E1A, which is required to reported in penguins maintained in an immortalize cells in culture; and E1B, aquarium, and in dogs. The virus is necessary for tumor formation. found in eastern USA, the Caribbean, Central America and north and east- E1A gene See E1 region . ern South America. In North America the principal vector is the swamp mosquito, Culiseta melanura, and the reser- Eagle’s medium (basal medium) Simple voir hosts are birds. In the Caribbean chemically defined medium; used in Culex taeniopus is the principal vector. growing many vertebrate cell lines. Mice, guinea pigs, goats, chicks, snakes Eagle H (1955) J Exp Med 102 , 595 and turtles can be infected experimentally. In birds the virus infects the vis- early antigens The products of expression cera (liver) rather than the CNS. Virus of genes expressed early in the replica- can be propagated in eggs and a vari- tion cycle, before genome replication. ety of tissue cultures in which it causes CPE. Vaccines are available for horses, early enzymes Enzymes, especially in pheasants and human laboratory work- cells infected with bacteriophage, ers exposed to EEEV. which are synthesized de novo under the direction of the invading virus Farrar MD et al (2005) Vet Diagn Invest 17 , 614 genome. Scott TW and Weaver SC (1989) Adv Virus Res 37 , 277 early genes Viral genes which are Weaver SC (1997) In Viral Pathogenesis , expressed early in the replication cycle edited by N Nathanson et al . Philadelphia: before nucleic acid replication occurs, Lippincott Raven, p. 329 e.g. the T antigen genes of SV40. Early gene products are usually involved in EB-3 cells (CCL 85) A cell line of the replication of the viral nucleic acid. lymphoblast-like cells, derived from See late genes . a 3-year-old black male with Burkitt’s EB-3 cells 138

lymphoma. The cells harbor a herpes- (2) virion length is about 970 nm as like virus particle detected by electron opposed to 790 nm; microscopy. (3) genome length of 18.9 kb as opposed to 19.1 kb; EBNA Epstein– Barr virus-induced nuclear (4) several gene overlaps as opposed a ntigen. A neoantigen complex asso- to a single overlap; ciated with the chromatin of latently (5) glycoprotein expression involves infected lymphoblastoid cells carrying transcriptional editing; the EBV (human herpes virus 4) viral (6) transcription of only the first ORF genome. The EBNA complex consists of of gene 4 yields a soluble small at least six virus-specified polypeptides, glycoprotein; which differ in structure dependent (7) protein profile distinct, however upon the inducing virus strain. The best species specific; characterized of the polypeptides are (8) glycoprotein gene nucleotide known as EBNA-1 and EBNA-2, both sequence difference of 57%. of which seem to elicit humoral antibodies in infected individuals. EBNA-1 Reston ebolavirus may be nonpathogenic is a phosphorylated polypeptide con- for humans, but this has not been for- sisting of 641 amino acids, which can mally tested for obvious reasons. be seen diffusely associated with cell chromosomes at metaphase. EBNA-2 EBTr(NBL-4) cells (CCL 44) A hetero- polypeptide consists of 443 (in EBV- ploid bovine cell strain derived from 2) or 491 (in EBV-1) amino acids and minced whole trachea of a male fetus is also phosphorylated. Although the of Bos taurus . Cells decline beyond the precise function of these proteins is not 55th passage. known, EBNA-1 is a DNA-binding protein which is involved in the initiation of EBV DNA replication, and EBNA-2 EBV Abbreviation for E pstein– B arr virus. seems to function as a trans -activator of See Human herpesvirus 4 . virus gene expression. There are three related genes for EBNA-3 tandemly ecbovirus ( e nteric c ytopathic b ovine located in the EBV genome (EBNA-3 A, o rphan v irus) Synonym for Bovine EBNA-3B, and EBNA-3C), which spec- enterovirus , a species in the genus ify large proteins (900–1000 amino acids Enterovirus. There are two serotypes, each) of presently unknown function. but neither has been associated with EBNA-LP is a nuclear phosphopro- disease in cattle. tein which consists of multiple repeat sequences of 22 and 44 amino-acids. Its echinoviruses A name proposed, but not function is presently unknown. adopted, for the group of syncytial or Epstein MA and Crawford DH (2005) In foamy viruses in the genus Spumavirus. Topley & Wilson’s Microbiology and Microbial The name refers to the presence of long Infections , vol. 1, Tenth edition, edited by projections or spines on the surface of BWJ Mahy and V ter Meulen. London: the virus particle. Hodder Arnold, p. 407 echovirus See human echoviruses 1–7 , 9 , Ebolavirus A genus in the family 11–27 , 29–33 . Filoviridae . The type species is Zaire ebolavirus (ZEBOV). There are three eclipse period The time after infection other species in the genus: Cote d’Ivoire between the disappearance of the ebolavirus , Reston ebolavirus, and Sudan infecting virus and the appearance of ebolavirus. A new virus which caused new intracellular virus infectivity. The an outbreak in Uganda in 2007 will infecting virus loses its infectivity soon probably be named Uganda ebolavirus . after penetration and for some time no Differs from members of the genus infective virus can be demonstrated. Marburgvirus. There are eight distin- See also latent period . guishing features: (1) almost no antigenic cross-reactivity; Doermann AH (1952) J Gen Physiol 35 , 645 139 Edmonston B strain ecmovirus ( e nteric c ytopathic m onkey eczema herpeticum A severe vesiculo- orphan virus) Synonym for simian pustular, umbilicated eruption caused enterovirus. by herpes simplex virus infection superimposed upon a pre-existing ecological niche occupancy The biotic atopic dermatitis. Also called Kaposi’s properties of members of a virus spe- varicelliform eruption. A related dis- cies such as host range, tissue tropism, ease, eczema vaccinatum, results from and vector. Provides the needs that vaccinia virus infection superimposed must be met for a virus to survive. upon a pre-existing atopic dermatitis. ecotropic murine type C virus A subspe- ED50 Abbreviation for median effective cies of mouse type C oncovirus in the dose. genus Gammaretrovirus. Infects and replicates in cultures of mouse and E Derm (NBL-6) cells (CCL 57) A hetero- rat cells only, unlike the amphotropic ploid cell strain initiated from a biopsy murine type C virus which will also of the dermis of an approximately 4- replicate in nonmurine cells. year-old quarter-horse, Equus caballus. Synonym : mouse-tropic strain. Cf. xeno- Has finite life expectancy. tropic murine type C viruses . Bryant ML et al (1978) Virology 88 , 389 Edge Hill virus (EHV) A species in the genus Flavivirus , serologically a mem- ecpovirus ( e nteric c ytopathic p orcine ber of the Yellow fever virus sero- orphan virus) Synonym for porcine group. Isolated from mosquitoes in enterovirus. Queensland, Australia. The major vertebrate hosts are probably marsupials. ecsovirus ( e nteric c ytopathic s wine No known association with disease in orphan virus) Synonym for porcine humans or other animals. enterovirus. edible vaccines Transgenic plants have ecthyma contagiosum of sheep virus been engineered to produce viral Synonym for Orf virus. antigens (e.g. HbsAg) which can be administered orally, but to date the Ectromelia virus (ECTV) A species in the development of an edible vaccine for genus Orthopoxvirus which is a natural humans is at an early stage. pathogen of mice. Serologically related Silin DS et al (2007) Expert Opin Drug Deliv to vaccinia virus. A latent infection, 4 , 323 endemic in many mouse stocks, activated by stress. Injection i.p. into mice Edman degradation A method for causes death from hepatitis. Injection sequencing peptides. The N-terminal into the skin of rabbits, guinea pigs, amino acid is removed by cleavage of and cotton rats produces local lesions the peptide bond with trifluoroace- which can be prevented by immuni- tic acid. If the N-terminal residue has zation with vaccinia virus. Replicates been labeled (e.g. with dansyl chlo- slowly on the CAM, with small, white, ride) it can be identified by its chro- irregularly shaped pocks appearing matographic properties. Otherwise the in 72 h. Pocks are not produced above amino acid composition of the remain- 39°C. Replication occurs in a number of ing peptide is compared with that of cell cultures, but the plaques produced the original peptide and the terminal are smaller and appear more slowly amino acid determined by deduction. than those due to vaccinia, cowpox, or monkeypox viruses. Gray WR (1972) Methods Enzymol 25 , 121 Synonyms : mousepox virus; pseudolymphocytic choriomeningitis virus. Edmonston B strain The first attenuated Buller RM and Palumbo GJ (1991) Microbiol measles vaccine strain, produced by Rev 55 , 80 passage in human kidney cells, human Esteban DJ and Buller RM (2005) J Gen Virol amnion cell, chick chorioallantoic mem- 86 , 2645 brane, and chick embryo fibroblast Edmonston B strain 140

cells. This vaccine was 95% protective eel virus American (EVA) A tentative but produced some cases of mild mea- species in the genus Vesiculovirus . sles (fever and rash). Therefore more Isolated in Europe and Japan from the attenuated strains were developed by blood of American and European eels, further passages of Edmonston B vac- Anguilla rostrata and Anguilla anguilla . cine in chick embryo fibroblast cells Diameter of virus particles 46–54 nm. to produce the Moraten and Schwartz Replicates with CPE in RTG-2 or EHM measles virus vaccine strains. monolayer cell cultures. Virus particles are present in clusters in the cytoplasm. Edmonston Zagreb strain A vaccine Injection into eels has not produced produced by further passage of the any lesions or disease. Edmonston B strain in human diploid Sano T (1976) Fish Pathol 10 , 221 cells. It produces a higher seroconversion rate than the Schwarz strain. eel virus B12 (EEV-B12) A tentative species in the genus Novirhabdovirus. Edmonston virus A strain of Measles Isolated from European eels, Anguilla virus in the genus Morbillivirus. anguilla , in France. Isolated in 1952 on the first day of the Castric J and Chastel C (1980) Ann Virol , rash from the blood of a 13-year-old Paris 13E , 435 boy with measles, David Edmonston. Used extensively as a prototype eel virus C26 (EEV-C26) A tentative spe- strain of measles. The original strain cies in the genus Novirhabdovirus. (Edmonston A) after extensive passage to cause attenuation was called the eel virus European (EVE) A tentative Edmonston B strain, from which most species in the genus Aquabirnavirus . current vaccine strains were derived, Caused nephritis disease outbreaks in except the Japanese AIK-C measles Japanese eels, Anguilla japonica , after vaccine, which was derived from the introduction of European eels, Anguilla Edmonston A strain. anguilla , into Japan and Taiwan in the 1970s. Closely related structurally to EDS Abbreviation for egg drop syn- infectious pancreatic necrosis virus. drome. See duck adenovirus 1 . Castric J et al (1984) Ann Virol , Paris 135E , 35

EDS virus Synonym for egg drop syn- efavirenz A non-nucleoside reverse tran- drome 1976-associated virus. scriptase inhibitor that inhibits HIV replication in vivo. Binds to the reverse EDTA Abbreviation for the chelating transcriptase and appears relatively agent e thylene d iamine- t etraacetic a cid. resistant to mutations. Albright AV et al (2000) AIDS Res Hum EEE virus Abbreviation for Eastern equine Retroviruses 10 , 1527 encephalomyelitis virus. Martinez E et al (2000) Clin Infect Dis 31 , 1266 Ren J et al (2000) Struct Fold Des 8 , 1089 eel iridovirus A possible species in the family Iridoviridae, isolated from effective concentration (EC ) The con- Japanese eel, Anguilla japonica , in 50 centration of an antiviral drug that Japan. reduces the titer of the virus by 50% is measured in cell culture and expressed eel orthomyxovirus-like agent A syn- as the EC50 . cytium-inducing virus isolated from European eels, Anguilla anguilla , with efficiency of plating (EOP) In growing ‘ cauliflower disease ’ (stomatopapil- bacterial cells on glass or plastic plates, loma). The virus has not been shown to the percentage of cells that give rise to cause the stomatopapillomatosis. colonies when a given number of cells Nagabayashi T and Wolf K (1979) J Virol 30 , are plated. In virology, a term intro- 358 duced by EL Ellis and M Delbrück in 141 Elaphe virus

1939 ( J Gen Physiol 22, 365) to quantify Ehrlich–Lettre ascites , strain E cells the relative efficiencies with which dif- (CCL 77) A cell line, of mouse origin, ferent cells could be infected and sup- derived as an explant of a 7-day-old port viral replication. It is obtained by tumor from the parental Ehrlich–Lettre dividing the plaque count by the total ascites carcinoma, which has an unusu- number of virions in the inoculum. For ally high mean chromosome number. animal viruses it ranges from 10 1 to 10 6 or less. EID50:HA ratio The ratio between the infective titer measured in eggs (ID50) EgAn 1825-61 (EGAV) A serotype and the hemagglutinating (HA) titer of of Uukuniemi virus in the genus a virus preparation. Especially applica- Phlebovirus . ble to influenza virus. It gives a measure of the proportion of defective virus egg drop syndrome 1976-associated particles present. virus (DAdV-1) A disease involving depressed and abnormal egg produc- EIF2 Eukaryotic initiation factor 2, a fac- tion in broiler chicken breeder flocks tor in translation of mRNA during has been recognized since 1976 in west- protein synthesis. Binds the initiator ern Europe. A number of serologically methionine-tRNA to the 40S ribosomal indistinguishable viruses were isolated, subunit. one of which, isolate 127, was further studied and found to be an adenovirus eIF4A A eukaryotic translation initiation of ducks. See duck adenovirus 1 . factor which is an RNA helicase. Binds Synonym : EDS 1976 virus. in a 1:1 ratio with eIF4G, and together McFerran JB (1979) Vet Q 1 , 176 with eIF4E, forms the eIF4 complex. Todd D and McNulty MS (1978) J Gen Virol 49 , 63 eIF4E A cap-dependent RNA helicase that is required for the translation of egg inoculation Eleven-day fertile chicken almost all capped messenger RNAs. eggs are used for growth of influenza viruses. The virus is inoculated into eIF4F A eukaryotic translation initiation the allantoic cavity and allowed to factor which consists of a protein com- multiply for 24–36 h by which time plex of eIF4E, eIF4A, and eIF4G. eIF4E it contains a great deal of virus, and binds to the cap structure of the mes- the allantoic fluid is harvested. Other senger RNA, eIF4A, an RNA-depend- viruses, such as pox viruses and some ent ATPase and ATP-dependent RNA retroviruses can be grown on the cho- helicase unwinds the secondary struc- rioallantoic membrane of the 11-day ture of the 5 -untranslated region of the fertile egg, yielding pocks that can be messenger RNA to facilitate ribosome counted to determine the infectivity of binding, and eIF4G serves as a scaffold the virus preparation. for eIF4E and eIF4A to coordinate their functions. EGTA Abbreviation for the chelating Imataka H and Sonenberg (1997) Mol Cell agent e thylene g lycolbis-(aminoethyl- Biol 17 , 6940 ether)-t etraacetic a cid. Li W et al (2001) J Biol Chem 276 , 29111

Egtved virus Synonym for Viral hemor- eIF4G A molecular scaffolding protein rhagic septicemia virus, a species in the involved in the translation complex. genus Novirhabdovirus. Elaphe virus A possible species in the EH IV (Elaine IV) cells (CCL 104) A cell genus Gammaretrovirus . Observed to be line derived, by seeding on monolayer associated with the cells of an embryo- WI 38 cells, from the peripheral blood nal rhabdomyosarcoma in a corn snake, leukocytes of a 20-year-old Caucasian Elaphe guttata. Primary heart and kid- female 2 months after the onset of ney cells and a cell line derived from a infectious mononucleosis. Contains rattlesnake fibroma could be infected Epstein–Barr virus in 0.1% of the cells. with a cell-free extract of the original Elaphe virus 142

tumor. Productive infection could not series of electromagnetic lenses. The be demonstrated in mammalian, chick contrast needed to make structures embryo, piscine, or non-ophidian rep- visible is provided by electron-dense tilian cells. salts of heavy metals such as phospho- Lunger PD and Clark HF (1978) Adv Virus tungstate or uranyl acetate. An impor- Res 23 , 159 tant technique in virus diagnosis and Lunger PD et al (1974) J Natl Cancer Inst 52 , characterization. 1231 Chrystie IL (1996) In Virology Methods Manual , edited by BWJ Mahy and elapid herpesvirus 1 (EpHV-1) An HO Kangro. London: Academic Press, p. 91 unassigned species in the family Stefan S et al (1999) J Clin Virol 13 , 105 Herpesviridae. Herpesvirus-like particles were present in the venom of a cobra, electron spin resonance Magnetic res- Naja naja . Also found in a siamese onance arising from the magnetic cobra and a banded krait. moment of unpaired electrons in a Synonyms: cobra herpesvirus; Indian paramagnetic substance. Used in cobra herpesvirus; banded krait her- determining the structure of organic pesvirus; siamese cobra herpesvirus. compounds. Lunger PD and Clark HF (1978) Adv Virus Res 23 , 159 electropherogram A picture (photograph or autoradiograph) which shows the elapid parvovirus A probable parvovirus distribution of proteins or nucleic isolated from a corn snake, Elaphe gut- acids that have been separated by gel tata , with pneumonia. Replicated in electrophoresis. IgH2 cells at 30°C. Synonym : corn snake parvovirus. electropherotype A characteristic profile of proteins or nucleic acids separated Ahne W and Scheinert P (1989) Zentralbl Vet by electrophoresis and often used to Med (B) 36 , 409 distinguish different virus strains. electrofocusing A method for separat- electrophoresis The separation of ing protein molecules by gel or density charged macromolecules, either in free gradient electrophoresis in which a pH solution or in a liquid in the pores of a gradient had been established. Each matrix, by the application of an electri- protein species moves to a pH approxi- cal potential difference. The movement mating to its isoelectric point. of the macromolecules is due to their charge but the separation may also electron cryomicroscopy (cryo-EM) A be due to their Stokes ’ radius. See gel technique for examining virus structure electrophoresis . which is designed to avoid artefacts that may be introduced by dehydra- electrophoretic mobility The movement tion and staining. High titer virus sus- of proteins or nucleic acids during gel pensions are placed on carbon grids electrophoresis, which is characteristic then rapidly frozen in liquid ethane. for each species depending upon size, This embeds the virions in a thin layer charge, and other factors such as fold- of amorphous ice within the holes ing of the molecule. in the carbon grid, and the electron micrographs can be digitized and 3- electroporation A method for introducing dimensional reconstruction performed. nucleic acids or other large macromol- Crowther RA et al (1994) Cell 77 , 943 ecules into cells by creating transient Dubochet J et al (1988) Q Rev Biophys 21 , 129 pores in the plasma membrane by means of an electrical pulse. electron microscopy A technique used Zimmerman U (1982) Biochim Biophys Acta for visualizing virus particles in which 694 , 227 a beam of electrons is focused onto the specimen and an image formed on a elementary bodies Small, round, staina- phosphorescent viewing screen by a ble, extracellular aggregates of virus, or 143 embryonated egg

viral products, seen in large numbers is detected by an antibody to which an by light microscopy in the vesicle fluid enzyme has been linked. The enzyme or scrapings from skin lesions of small- is detected by the production of color pox, vaccinia, varicella and zoster. on reaction with a substrate. Chernesky MA and Mahoney JB (1996) In elephant loxodontal herpesvirus Syn- Virology Methods Manual , edited by BWJ onym for elephantid herpesvirus 1. Mahy and HO Kangro. London: Academic Press, p. 123 elephant papillomavirus (EPV) A prob- Reen DJ (1994) Methods Mol Biol 32 , 461 able species in the family Herpesviridae . Proliferative cutaneous lesions devel- elk papillomavirus See European elk oped in a herd of captive African ele- papillomavirus . phants. Southern blot analysis of DNA from lesion specimens did not indi- Ellidaey virus (ELLV) A serotype of Great cate papillomavirus-specific genomes. Island virus in the genus Orbivirus , Particles resembling herpesviruses belonging to the Great Island complex were seen by electron microscopy. of the Kemerovo serogroup. Isolated Jacobson ER et al (1986) J Am Vet Med Assoc from a pool of engorged ticks, Ixodes 189 , 1075 uriae , from a puffin at Ellidaey, west Iceland in 1981. Designated ELL-3a. elephantid herpesvirus (ElHV-1) An unassigned virus in the family El Moro Canyon virus (ELMCV) A species Herpesviridae. Routine post-mortem in the genus Hantavirus , isolated from examination of 50 elephants, Loxodonta the harvest mouse, Rheithrodontomys africana, killed in the Kruger National megalotis , in western USA, Mexico, and Park revealed lymphoid nodules in Canada. Not known to cause disease in the lungs of 37 of them (74%). There humans. were from 1 to 6 grayish-white nodules Synonym : harvest mouse virus 1. present, with diameters varying from Hjelle B et al (1995) Crit Rev Clin Lab Sci 32 , 3 to 30 mm. Cowdry type A intranu- 469 clear inclusions were present in epithe- Hjelle B et al (1995) Virology 207 , 452 lial cells of the nodules, and electron microscopy demonstrated particles of elongation factor Protein, forming part of herpesvirus morphology. Recent fatal the ribosomal binding complex, which endothelial disease described in 10 promotes elongation of a polypeptide elephants in North American zoos was chain. caused by endotheliotropic elephant herpesvirus. This virus is apathogenic for the African elephant, but causes El Tifu Negro (black typhus) A local fatal hemorrhagic disease in the Asian name for Machupo virus infection. elephant (Elepha s maximus ). Synonym: elephantid herpesvirus; ele- elution of virus Release of virus parti- phant loxodontal herpesvirus. cles from association with a cell sur- Ehlers B et al (2001) J Gen Virol 82 , 475 face or other solid support such as an McCully RM et al (1971) Onderstepoort J Vet ion-exchange column. The elution of Res 38 , 225 Orthomyxoviridae by the action of neu- Richman LK et al (1999) Science 283 , 1171 raminidase from erythrocytes to which they have been adsorbed constitutes an elephantpox virus Family Poxviridae. A effective purification method. poxvirus isolated from an elephant. Now known to be a species of Cowpox EMBO E uropean M olecular B iology virus. O rganization.

ELISA E nzyme- l inked i mmuno s orbent embryonated egg A fertilized egg, used a ssay. A serological method in which an for growing viruses such as influenza antigen, immobilized on a solid matrix, virus when 11–12 days old. Embu virus (ERV) 144

Embu virus (ERV) Unassigned virus termed aseptic encephalitis. Can be of the family Poxviridae. The virus is caused by a wide variety of viruses, transmitted by mosquitoes to rodents. especially arboviruses and some enteroviruses. EMCV See Encephalomyocarditis virus . encephalitozoon rabei Name given to a emerging infections Infectious diseases supposed protozoan etiological agent which are newly recognized, or have of rabies. The structure is now known increased significantly in incidence as the ‘ Negri body. ’ during the previous 10 years. Manouelian Y and Yiala J (1924) Ann Inst Jones KE et al (2008) Nature 451 , 990 Pasteur 38 , 258 emerging viruses The number of newly Encephalomyocarditis virus (EMCV) recognized viruses has increased dra- Type species of the genus Cardiovirus . matically in the past 20 years. Factors The genome of EMC contains 7840 responsible include: mutation; host fac- bases plus a short poly A tract of about tors such as immunodeficiency caused 35 bases at the 3 end. The 5 non- by deliberate (e.g. cancer chemother- translated region is 833 bases long apy) or acquired (e.g. HIV infection) and contains a poly C tract located immunosuppression; new technologies 150 bases from the 5 terminus. The for virus detection (e.g. PCR); and envi- virus causes a natural infection of wild ronmental changes (e.g. forest clearing) rodents in which it probably rarely which bring humans in closer contact causes disease. Has been isolated from with virus vectors such as arthropods blood and stools of humans, from cap- or rodents. tive lower primates, pigs, squirrels, and other animals. May cause febrile illness Mahy BWJ (2006) Biodiversity 7 , 34 with CNS involvement in humans. Many animals can be infected experi- empty particles Virus particles contain- mentally. Mice and hamsters often die ing no nucleic acid, usually identi- with CNS involvement after infection fied by negative staining and electron by any route. All strains are identical microscopy of virus preparations. when compared by neutralization, CFT or hemagglutination–inhibition (HAI), emtricitabine A synthetic nucleoside but many show differences in biologi- analogue of cytosine, active against the cal behavior. Agglutinates sheep eryth- reverse transcriptase of HIV-1. Once rocytes in the cold. Replicates with incorporated into the nascent viral CPE in cell cultures of chick, mouse, DNA it causes chain termination. humans, monkey, hamster, and cattle. Synonyms : Columbia-SK virus; ME emtriva An alternative name for virus; mouse Elberfield virus; mengo emtricabine. virus. encapsidation signal (€) A sequence of Palmenberg AC (1987) In The Molecular 137 nucleotides near the 5 -terminus Biology of the Positive Strand RNA Viruses , of the pregenome of hepatitis B virus, edited by DJ Rowlands et al. London: Academic Press, p. 1 known as epsilon, which is needed for the specific encapsidation of the endemic A disease which persists in a hepadnaviral RNA pregenome. A given population or locality. protein product of the P gene is also needed as a structural component dur- endemic pneumonia of rats See pneu- ing encapsidation. monia virus of rats . Bartenschlager R et al (1990) J Virol 64 , 5324 Junker-Nipmann M et al (1990) EMBO J 9 , endocytosis The process by which extra- 3389 cellular fluid or particles are taken up by eukaryotic cells. Occurs by invagi- encephalitis Inflammation of the brain. nation and pinching off of the plasma When caused by virus infection it is membrane so that internalized particles 145 endoplasmic reticulum

are initially present in vesicles (endo- factors (e.g. hormones) or extrinsic fac- somes) within the cytoplasm. Uptake tors (e.g. radiation, chemicals). Once of large particles is termed ‘ phago- integrated into the DNA, the virogenes, cytosis,’ and uptake of solutes, fluid both endogenous and exogenous, are and small particles such as viruses subject to the same pressures of selec- is termed ‘ pinocytosis. ’ Endocytosis tion and mutation as other genes. The may be a general phenomenon which genomes of all eukaryotic organisms allows various molecules to enter the studied to date contain endogenous cell. Virus entry, however, is usually retroviruses, and amongst vertebrates receptor-mediated and requires a spe- endogenous retrovirus-specific nucle- cific cell-surface receptor. The virus– otide sequences have been found in all receptor complex accumulates in orders examined to date, including car- coated pits of the plasma membrane, tilaginous and bony fish. Endogenous which pinch off as intracellular resi- viruses can be recovered from cells of dues. For many enveloped viruses, humans and many other species, e.g. escape from the vesicle requires acidi- birds, mice, cats, pigs, baboons, etc. fication within the vesicle, and fusion They often have a limited host range of the virion lipid envelope and the (N- or B-tropic) or may not replicate vesicle membrane occurs as the pH is in the species from which they were lowered. The pH at which this fusion isolated. Such viruses are called xeno- occurs is specific for different viruses. tropic or ‘ S ’ -tropic. See also human endogenous retroviruses . endo-epidemic hemorrhagic fever virus Boyce-Jacino MT et al (1992) J Virol 66 , 4919 Synonym for Junín virus. Coffin JM et al (1997) In Retroviruses . New York: Cold Spring Harbor Laboratory Press, endogenous avian retrovirus The pres- p. 343 ence of reverse-trancriptase activity can Ruis BL et al (1999) J Virol 73 , 5345 be detected by highly sensitive meth- Tristem M (2000) J Virol 74 , 3715 ods (amplified reverse transcriptase, amp-RT) and by this means it was endolymphatic labyrinthitis A compli- found that most vaccines prepared cation of infection with mumps virus, in chick embryo fibroblast cells for which may cause deafness, which is human use contain detectable levels usually unilateral. The worldwide of RT. These include measles, mumps, application of the MMR vaccine has and yellow fever vaccines for use in largely eliminated this condition. humans. The RT activity was found endonuclease A phosphodiesterase that to be associated with viral particles does not require a terminus for hydro- containing RNA from the endogenous lytic activity. Makes internal cleavages avian leukosis virus subgroup E (ALV- in nucleic acids. See deoxyribonucle- E) and the endogenous avian virus ase , nucleases . (EAV). However, testing of MMR and yellow fever vaccine recipients has endonuclease restriction-PCR analy- revealed no evidence of the transmis- sis A technique used for the genetic sion of ALV-E or EAV to the vaccine characterization of influenza viruses. recipients. Oligonucleotide primers are designed Mahy BWJ and Hadler S (1996) Clin Diagn to amplify a product by PCR that con- Virol 5 , 1 tains unique restriction enzyme cleav- Tsang SX et al (1999) J Virol 73 , 5843 age sites. The method can be used to discriminate between two lineages of endogenous retrovirus A retrovirus influenza H3N2 viruses circulating whose genome is stably integrated simultaneously, or to identify drug- into the DNA of normal cells. It is thus resistant influenza virus isolates. genetically transmitted from genera- Ellis JS et al (1997) J Med Virol 51 , 234 tion to generation. The gene sequences Klimov A et al (1995) J Infect Dis 172 , 1352 which can code for virus production are virogenes and are normally repressed, endoplasmic reticulum A membrane but may be activated by intrinsic system in the cell cytoplasm consisting endoplasmic reticulum 146

of lamellae and tubules. The site at have been studied in detail: the which biosynthetic processes such as England strain and the Maastricht membrane assembly, protein secretion, strain, both first described in 1982. and organelle biosynthesis occur in Because of profound genetic sequence eukaryotic cells. The site of maturation differences, these now appear to be dis- of membranes of many viruses. tinct species rather than strains. Beisser PS et al (1998) Virology 246 , 341 endosome Cellular organelle that plays a central role in endocytosis and plasma engulfment Synonym for viropexis. membrane recycling. enhancement An increase in yield of endosymbiotic infection General term virus or cytopathic effect, or both, fol- coined to describe infection of cells lowing mixed infection of cells by by viruses in which replication occurs two unrelated viruses. A term used without CPE. The term was employed when the mechanism of the process is to distinguish this type of persist- not clear. See complementation and ent infection from a carrier culture, interference . in which only a proportion of the cell population is infected at any one time. enhancers Non-transcribed genetic ele- Fernandes MY et al (1964) J Exp Med 120 , ments present in the genomes of many 1099 viruses that regulate the transcription of specific genes. May exhibit cell or endotheliotropic elephant herpesvirus tissue specificity, so promoting or lim- An unclassified herpesvirus which is iting virus infection. apathogenic for the African elephant (Loxodonta africana) but causes fatal enhancer sequence A short cis -acting hemorrhagic disease in the Asian ele- DNA sequence that increases the tran- phant (Elephas maximus). Caused dis- scriptional activity of a cellular or viral ease in elephants in a number of North gene. American zoos in the last ten years. McKnight S and Tijan R (1986) Cell 46 , 795 At least four different virus genotypes appear to have been involved in these Enseada virus (ENSV) An ungrouped zoo outbreaks. It has been proposed virus in the family Bunyaviridae. that these viruses should be classified Isolated from Culex (Mel) taeniopus in in a new genus Proboscivirus in the sub- São Paulo, Brazil. Not associated with family Betaherpesvirinae . See elephantid disease. herpesvirus . entamoeba virus (ENTV) An unassigned endothelium Squamous epithelium lin- virus in the family Rhabdoviridae . ing blood vessels, lymphatics and Synonym : rhabdovirus entameba. blood-filled cavities such as the anterior chamber of the eye. A number of Entebbe bat virus (ENTV) A species viruses infect and multiply in endothe- in the genus Flavivirus , belonging to lial cells such as filoviruses, Hendra the Rio Bravo serogroup. No known virus, and Nipah virus. arthropod vector. Isolated from the salivary glands of a pool of bats of enfurvirtide An inhibitor of HIV-1 rep- Tadarida sp in Uganda. Not reported to lication which prevents the virus from cause disease in humans. fusing with CD4 cells. Consists of a 36 amino-acid synthetic peptide which is Entecavir A cyclopentile guanosine nucl- administered subcutaneously. eoside analogue with selective activity Lalezari JP et al (2003) N Engl J Med 348 , against hepatitis B DNA polymerase. 2175 Can be administered orally. The development of resistant mutants is a England rat cytomegalovirus A probable problem. species in the genus Muromegalovirus . Rodriguez-Frias F et al (2007) J Med Virol 79 , Two strains of rat cytomegalovirus 1671 147 enzyme conjugate enteric viruses Viruses which primarily with little loss of cytoplasm. Used to infect the gastrointestinal tract. study whether virus replication events require nuclear functions. enteritis of mink virus Synonym for Feline panleukopenia virus. envelope The outer coat of large viruses such as influenza virus or paramyxo- Enterovirus A genus in the family viruses. Strictly, the outer lipoprotein Picornaviridae. The type species is coat of viruses which mature by bud- Poliovirus . Other species include Bovine ding through the cell membrane. Such enterovirus , Human enterovirus A to D , viruses are inactivated by lipid solvents and Porcine enterovirus A and B . (e.g. chloroform or ether). Clinical manifestations include men- Synonym : peplos. ingitis, encephalitis, myelitis, myocarditis, and conjunctivitis. Distinguished env gene One of the genes in the genome from rhinovirus by: (1) being resist- of the retroviruses. Codes for the major ant to low pH; (2) having a density glycoprotein found on the surface of of 1.32–1.35 g/ml; and (3) the disease the viral envelope, hence the name. produced in humans or experimental animals. Many species have entero- enviroxime A benzimidazole derivative viruses which tend to be species-specific. that inhibits rhinovirus replication in Transmission is horizontal by fecal con- vitro. tamination. Food and water are often Philpotts RJ et al (1981) Lancet i , 1342 involved; arthropods may play a minor role. The first 67 human enteroviruses enzootic A disease constantly present were divided into 4 species (3 polio- in an animal community in a defined virus, 6 coxsackie B, 24 coxsackie A, geographic region but affecting only and 34 echoviruses) but these are a small number of animals at any one now included in a new classifica- time. Analogous to endemic. tion with only 9 species in the genus. Enteroviruses have also been isolated enzootic abortion of ewes virus Not a from several other species (e.g. horse, virus. A chlamydia organism. sheep, buffalo, and dog). Hyypia T et al (1997) J Gen Virol 78 , 1 enzootic bovine leukosis virus Synonym for Bovine leukemia virus. enterovirus See Human enterovirus A–D . enzootic bronchiectasis of rats See pneu- enteroviruses 68–71 See human entero- monia virus of rats . virus 68–71 . enzootic nasal tumor virus (ENTV) enterovirus 72 The original name for the A probable species in the genus virus now known as Hepatitis A virus . Betaretrovirus. Causes enzootic nasal Entomopoxvirinae A subfamily in the tumors in small ruminants. The enve- family Poxviridae containing the pox- lope protein of the virus acts as an viruses of insects. There are three genera: oncogene, in a similar manner to Alphaentomopoxvirus , Betaentomopoxvirus , Jaagseikte sheep retrovirus. Sheep and Gammaentomopoxvirus , including and goat genomes contain copies of viruses of Coleoptera, Lepidoptera, and closely related endogenous retrovirus Diptera, respectively. sequences. Maeda N and Fan H (2008) Virus Genes 36 , entry site The ribosome attachment site 147 on the virus or cellular mRNA that is Palmarini M et al (2000) J Virol 74 , 8065 available for initial binding of transfer RNA during translation. enzyme conjugate Usually an antibody preparation to which an enzyme is enucleated cells Eukaryotic cells from linked covalently. The enzyme is cho- which the nuclei have been removed sen to produce a color reaction with enzyme conjugate 148

an appropriate substrate; enzymes epidemic diarrhea of infant mice virus commonly used include alkaline phos- (EDIM) A probable species in the phatase and horseradish peroxidase. genus Rotavirus . Mainly attacks mice Used in enzyme-linked immunosorb- 11 to 15-days old, causing diarrhea and ent assay and Western blotting. dirty yellow fur. Not necessarily fatal. Disease is most common in first litters. enzyme-elevating virus Synonym for Replication in cell cultures is in doubt. Lactate dehydrogenase-elevating virus. Infectivity resists ether, chloroform, and pH 3. A stock free of virus can be enzyme-linked immunosorbent assay obtained by Caesarean section and fos- See ELISA tering on disease-free females. Virus morphologically identical and antigenically very similar to human and other enzyme-linked virus-inducible system rotaviruses. (ELVIS) A rapid, specific method for the detection of herpes simplex virus Kraft LM (1966) Natl Cancer Inst Monogr 20 , using BHK-21 cells that have been sta- 55 bly transfomed with a β -galactosidase from E coli attached to a herpes simplex epidemic gastroenteritis virus Synonym virus-inducible promoter. When these for Norwalk virus . cells are infected with either HSV-1 or β HSV-2, the -galactosidase is induced, epidemic hemorrhagic fever virus Syn- then an X-gal colorimetric substrate is onym for Hantaan virus . added and herpes virus-infected cells turn blue, but remain colorless in the presence of other viruses. epidemic hepatitis-associated antigen Synonym for Hepatitis delta virus . Olivo PD (1996) Clin Microbiol Rev 9 , 321 epidemic jaundice virus of humans Syn- EOP See efficiency of plating . onym for Hepatitis A virus . ephemeral fever virus See Bovine ephem- epidemic keratoconjunctivitis A highly eral fever virus . infectious disease with round sub- epithelial corneal opacities in asso- Ephemerovirus A genus in the fam- ciation with keratitis, swelling of the ily Rhabdoviridae. The type species is regional lymph nodes, and headache. Bovine ephemeral fever virus. Other spe- Adenovirus 8 has frequently been iso- cies include Adelaide River virus and lated from case patients. Berrimah virus , and possibly Kimberley Synonyms : Sanders ’ disease; shipyard virus, Malakal virus, and Puchong eye; viral keratitis. virus. Virions are bullet-shaped, 140–200 nm long and 60–80 nm across. epidemic myalgia Synonym for Bamble The virus genome is a 14.9 kb negative- disease. stranded RNA molecule encoding 10 genes in the order 3 -N-P-M-G-Gns-α 1- α 2- β - γ -L-5 . The viruses cause an eco- epidemic tremor virus Synonym for nomically important disease of cattle avian encephalomyelitis virus. and water buffalo in tropical regions of Africa, Australia, the Middle East and epidemiology The study of factors Asia. Mortality is about 1–2%. Infection involved in the spread of diseases in is transmitted by arthropods, both human and animal populations. Culicoides sp and mosquitoes. epidermal growth factor (EGF) A potent epidemic A sudden increase in the inci- mitogen that acts on a wide variety of dence of a disease, affecting a large cell types by binding to its cell surface number of susceptible hosts and receptor and inducing a cascade of cel- spreading over a large area. lular events that lead to cell division. 149 epizootic epidermal growth factor receptor dissociation of the antigen; neotope , (EGFR) The cell surface receptor which which arises from changes in folding of binds EGF, a mitogenic polypeptide the polypeptide chain; metatope , which active on a variety of cells. Has tyrosine is formed from adjacent protein mol- kinase activity. ecules during assembly of structures Hügin AW and Hauser C (1994) J Virol 68 , from subunits; and mimotope , which 8409 is a peptide structure that mimics the shape of an epitope. The structure on epidermodysplasia verruciformis (EV) the antibody which is complementary Widespread and persistent dissemina- to the epitope is termed the paratope. tion of flat warts (verruca plana ), some- In the cell-mediated immune response, times for decades, which can progress T-cells recognize epitopes of proc- to malignancy. Often associated with essed proteins in a complex with HLA HPV types 5 and 8. Found particularly (MHC) molecules. in immunosuppressed persons (e.g. Laver WG et al (1980) Cell 61 , 553 AIDS patients). van Regenmortel MHV (1966) Adv Virus Res Synonym : Lewandowsky-Lutz disease. 12 , 207 episomes Plasmids which can be inte- epitope analysis Analysis of the primary grated into the cell genome, where they structures of proteins may not reveal behave as part of a chromosome. Phage the antigenic determinants that are lambda is an example. May behave as important epitopes for the immune an episome in one cell and a plasmid system. The complete structure of an (replicating independently of the chro- epitope can be determined by X-ray mosome) in another. diffraction of crystals of a complex of a monoclonal antibody Fab fragment epithelial cells Epithelial cells in culture with its antigen. form tight junctions and become polar- van Regenmortel MHV (2008) In ized into apical and basolateral regions. Encyclopedia of Virology , Third edition, BWJ Enveloped viruses bud from one or Mahy and MHV van Regenmortel (eds) other of these surfaces, and usually not Oxford: Academic Press, vol. 1, p. 137 from both. epitope mapping Specific epitopes can Tucker SP and Compans RW (1993) Adv Virus Res 42 , 187 be identified by analysis of the reactivity of neutralization escape mutants with panels of monoclonal antibodies. epithelioma contagiosum virus Synonym These epitopes can then be mapped by for Fowlpox virus . nucleotide sequencing of the escape mutants and physical mapping of the epithelioma of carp virus Synonym for mutational changes. If the three dimen- cyprinid herpesvirus 1. sional structure is known, e.g. the haemagglutinin of influenza virus, these epithelioma papillosum of carp virus changes can be accurately located on Synonym for cyprinid herpesvirus 1. the antigen structure. epitope The structure of an antigen Epivir-HBV Approved to fight HIV which elicits the formation of spe- infections in 1995 this drug works as a cific antibodies, e.g. a grouping of reverse transcriptase inhibitor by block- amino acid sequences on a protein, or ing an enzyme the AIDS virus uses to between adjacent protein subunits; also reproduce. It is now also approved to termed ‘ antigenic site. ’ The term was treat chronic hepatitis B. coined in 1960 by Jerne, who proposed Synonyms : Lamivudine, 3TC. that ‘ an antigen particle carries several epitopes.’ Various types of epitope have epizootic A disease temporarily present been distinguished: cryptotope, which is in an animal community, attacking buried inside the antigen and becomes many animals at the same time, spread- reactive only after denaturation or ing rapidly, and becoming widely epizootic 150

diffused. Analogous to an epidemic in Epsilonretrovirus A genus of exogenous human populations. retroviruses that infect reptiles and fish. They are complex, with genomes epizootic cellulitis virus Synonym for ranging from 11.7 to 12.8 kb. Walleye His Equine arteritis virus. dermal sarcoma virus uses a tRNA primer, but snakehead retrovirus uses Arg Epizootic hemorrhagic disease virus 1–8 tRNA . (EHDV 1–8) A species in the genus Orbivirus belonging to the epizootic Epstein–Barr virus Synonym for Human hemorrhagic disease serogroup. Has herpesvirus 4 . caused fatal epizootics in Virginian white-tailed deer, Odocoileus virgin- equestron Name given to a postulated ianus. Mule deer and other species are regulatory protein produced during not susceptible. After an incubation human poliovirus replication. It would period of 6–8 days there are signs of regulate the suppression of host protein shock, multiple hemorrhages, coma, synthesis and control the production of and death. Virus can be passed i.c. viral RNA and protein. No satisfactory in suckling mice and loses virulence evidence in favor of the existence of for deer. There are eight serotypes of this regulator has been forthcoming. EHDV; Ibaraki virus which affects cattle in Japan is an antigenically related Cooper PD et al (1973) Intervirology 1 , 1 member of the EHDV group. EHDV is found in USA, Canada, and Australia, Equid herpesvirus 1 (EHV-1) A species in where cattle, buffalo, and deer are the genus Varicellovirus , closely related infected by at least five serotypes with- to EHV-4. A natural infection confined out clinical disease. Infection does not to equines. A common cause of acute pass by direct contact and the princi- respiratory disease in horses dur- pal arthropod vectors are Culicoides sp. ing their first 2 years of life. Natural Injection into European deer, sheep, transmission probably by respiratory goats, and cattle leads to a silent infec- route. Usually silent in mares but abortion. Not reported to cause disease in tion may occur, especially in months humans. 8–10 of pregnancy. Should not be con- Hourrigan JL and Klingsporn AL (1975) fused with equine infectious arteritis Aust Vet J 51 , 203 virus which can also cause abortion in mares. Experimental infection during the months 3–9 of gestation may epizootic hematopoietic necrosis virus result in encephalitis due to vascu- (EHNV) A tentative species in the litis. Genital vesicular exanthema or genus Ranavirus , affecting at least 13 pustular vulvovaginitis may be pro- genera of fish including rainbow trout duced. Experimentally, causes abor- and redfin perch. tion in guinea pigs; suckling hamsters infected i.p. develop hepatitis. Has epizootiology The study of factors been adapted to growth on the CAM, involved in the spread of diseases in in the yolk sac and amnion. Replicates animal populations. with CPE in fetal horse cell cultures, Dohoo IR et al (1994) Nature 368 , 284 also in HeLa, human amnion, sheep, Mims CA (1991) Epidemiol Infect 106 , 423 pig, cattle, cat, and chick cell cultures. Live virus given to non-pregnant mares epornotic An epidemic of disease is safe and gives protection. amongst birds. Synonyms : Equid herpesvirus 1; equine abortion virus; equine influenza virus; equine rhinopneumonitis virus; Epsilonpapillomavirus A genus in the mare abortion virus; equine abortion family Papillomaviridae containing herpesvirus. viruses which cause cutaneous papillomas in cattle. The type species is Bovine Crabb BS and Studdert MJ (1995) Adv Virus papillomavirus 5 . Res 45 , 159 151 Equid herpesvirus 7 (EHV-7)

Equid herpesvirus 2 (EHV-2) A species vaccine is available, and alternative in the genus Rhadinovirus. Originally recombinant-derived vaccine candi- isolated from a horse with catarrh, but dates are under investigation. usually causes only mild disease or Synonyms : equine herpesvirus 4; equine silent infection. Is widespread and can rhinopneumonitis virus; respiratory be isolated from infected horses over infection virus. a prolonged period. A slowly replicat- Crabb BS and Studdert MJ (1995) Adv Virus ing cytomegalo-type virus that is not Res 45 , 153 host-cell specific. There are several Studdert MJ (1981) Science 214 , 562 serotypes, and the viruses numbered by Plummer as types 2, 3, and 4 are Equid herpesvirus 5 (EHV-5) A species in probably best regarded as serotypes of the genus Rhadinovirus. Following res- Equid herpesvirus 2 . Serologically dis- piratory tract infection, causes a disease tinct from equid herpesviruses 1 and 3. in horses which resembles infectious Synonym : equine cytomegalovirus. mononucleosis, with malaise, fever, Agius CT and Studdert MJ (1994) Adv Virus and enlarged lymph nodes followed by Res 44 , 357 virus persistence in circulating leuko- Browning GF and Studdert MJ (1989) Arch cytes. The consequent loss of physical Virol 104 , 77 performance is a serious concern to the Telford EA et al (1995) J Mol Biol 249 , 520 racehorse industry. The similarities to Epstein–Barr virus suggest that EHV-5 equid herpesvirus 3 (EHV-3) A tenta- may best be classified with the gam- tive species in the genus Varicellovirus. maherpesviruses, and this awaits fur- Isolated from horses during an out- ther molecular characterization of the break of coital exanthema, an acute, genome. relatively mild disease in which Synonym : equine herpesvirus 5. there are pustular lesions on the external genitalia and vagina. Cross- Agius CT and Studdert MJ (1994) Adv Virus neutralization tests showed it to be Res 44 , 357 distinct from equid herpesviruses 1 Browning GF and Studdert MJ (1989) J Gen Virol 68 , 1441 and 2. Transmission experiments with this virus produced typical equine coi- equid herpesvirus 6 (EHV-6) tal exanthema, except in one animal A tenta- Varicellovirus. which had recently recovered from the tive species in the genus disease. Isolated from vesicular lesions on the Synonyms: coital exanthema virus; muzzle of a foal donkey and the exter- equine herpesvirus 3. nal genitalia and udder of its dam. Restriction analysis of the genome Studdert MJ (1974) Cornell Vet 64 , 117 DNA revealed a relationship to EHV-3, Sullivan DC et al (1984) Virology 132 , 352 but the viruses are antigenically quite distinct. Equid herpesvirus 4 (EHV-4) A species in Synonym : asinine herpesvirus 1. the genus Varicellovirus. A major cause of acute respiratory disease in horses Browning GF et al (1988) Arch Virol 101 , 183 worldwide, most horses being infected Jacob RJ (1988) In Equine Infectious Diseases , during the first 2 years of life. Shown vol. 5, edited by DG Powell. Lexington: University of Kentucky Press, p. 140 in 1981 to be distinct from EHV-1 by restriction endonuclease studies on the virus genome. Horses may become Equid herpesvirus 7 (EHV-7) A species in latently infected, and reactivation with the genus Rhadinovirus. Isolated from virus shedding may then occur to infect the peripheral blood leukocytes of a young foals and so maintain the virus healthy donkey by co-cultivation with indefinitely in a population of horses. equine fetal kidney cells. A betaherpes- Acute disease is associated with fever, virus that is distantly related to EHV-2 anorexia, and profuse nasal discharge. and EHV-5. The clinical and epidemio- In extreme cases the disease may logical significance is not known. become a fatal bronchopneumonia. A Synonym : asinine herpesvirus 2. combined EHV4/EHV-1 inactivated Browning GF et al (1988) Arch Virol 101 , 183 Equid herpesvirus 8 (EHV-8) 152

Equid herpesvirus 8 (EHV-8) A species in culture from the feces of two foals the genus Varicellovirus. Isolated from with diarrhea. The virus was distinct an EHV-1 seropositive donkey that from EAdV-1, and was proposed to be was treated with corticosteroids for 3 EAdV-2. A serological survey of 339 days before nasal swabs were taken equine sera showed that 77% contained and inoculated into equine fetal kidney neutralizing antibody to EAdV-2. cells. The virus is genetically related Studdert MJ and Blackney MH (1982) Am J to EHV-1, and is apparently an alpha- Vet Res 43 , 543 herpesvirus of donkeys which causes acute respiratory disease followed by Equine adenovirus A (EadV-A) A species latent infection in the natural setting. in the genus Mastadenovirus . Synonym : asinine herpesvirus 3. Synonym : equine adenovirus 1. Browning GF et al (1988) Arch Virol 101 , 183 Reubel GH and Studdert MJ (1997) Arch Virol 142 , 1193 Equid herpesvirus 9 (EHV-9) A species in Wilks CR and Studdert MJ (1973) Aust Vet J the genus Varicellovirus. Isolated from a 49 , 456 captive gazelle, but zebras may be the primary host. Immunologically related Equine adenovirus B (EadV-B) A species to equine herpesvirus 1. in the genus Mastadenovirus . Synonym : gazelle herpesvirus. Synonym : equine adenovirus 2. Fukushi H et al (1997) Virology 227 , 34 Equine arteritis virus (EAV) The type species in the genus Arterivirus. Horses equilibrium density gradient cen- are the only susceptible species. trifugation See isopycnic gradient Causes epizootics and is highly con- centrifugation . tagious, infecting mainly young animals via the respiratory tract. Causes equine AAV See Equine adeno-associated fever, conjunctivitis, rhinitis, edema of virus . the legs and trunk, enteritis and colitis. In pregnant mares the fetus may equine abortion herpesvirus Synonym become infected and abortion occurs. for Equid herpesvirus 1 . Bronchopneumonia and pleural effu- sions occur in fatal cases. There is equine abortion virus Synonym for Equid medial necrosis of small arteries and herpesvirus 1 . when the intima is involved, throm- bosis. The virion is 50–70 nm in diam- Equine adeno-associated virus (EAAV) eter with a core 20–30 nm in diameter, A species in the genus Dependovirus. enveloped, inactivated by lipid solvents Obtained from a foal. Disease associa- and low pH. Replicates in horse kidney tion, if any, is presently unknown. cell cultures with CPE. Virus becomes Dutta SK (1975) Am J Vet Res 36 , 247 attenuated on passage in tissue culture and can be used as a vaccine. equine adenovirus 1 (EAdV-1) A Synonyms: epizootic cellulitis virus; strain of Equine adenovirus A in the equine infectious arteritis virus; equine genus Mastadenovirus. Isolated from influenza virus; fievre typhoide du horses with respiratory disease. Often cheval virus; infectious arteritis of causes death from pneumonia in foals. horses virus; pferdestaupe virus; pink Isolates in Germany, Australia and USA eye virus. are antigenically similar. de Vries AAF et al (1997) Semin Virol 8 , 33 Fatemie-Nainie S and Marusyk R (1979) Am Snijder EJ and Meulenberg JJM (1998) J Gen J Vet Res 49 , 521 Virol 79 , 961 Studdert MJ et al (1974) Am J Vet Res 35 , 693 equine coital exanthema virus See equid equine adenovirus 2 (EAdV-2) A strain herpesvirus 3 . of Equine adenovirus B in the genus Mastadenovirus. An adenovirus was equine cytomegalovirus Synonym for isolated in equine fetal kidney cell Equid herpesvirus 2 . 153 Equine papillomavirus 1 (EqPV-1) equine encephalitis virus (EEV) Three remissions. Transmission to other spe- alphaviruses cause encephalitis in cies is reported but not confirmed. Insect equines and in humans: Eastern vectors, probably mechanical, are sus- EE, Western EE, and Venezuelan EE pected but contact infection is possi- viruses. ble as virus is present in milk, semen, saliva, and urine. Control is by slaugh- Equine encephalosis virus (EEV) A spe- ter as there is no vaccine of proved cies in the genus Orbivirus, related to efficacy. The RNA genome is 8.2 kb in African horse sickness virus, and caus- length, and the DNA provirus includes ing a similar disease. LTRs of 321 nucleotides at each end. The virion is 80–120 nm in diameter, equine encephalosis viruses 1–7 (EEV1–7) enveloped, possibly with small pro- Species in the genus Orbivirus , belong- jections. Agglutination of fowl, frog, ing to the equine encephalosis virus and human erythrocytes by serum of group. Isolated from horses in South infected horses is reported. Strains can Africa with fatal peracute illness resem- be distinguished by virus neutraliza- bling African horse sickness. tion but have a common CF antigen. Verwoerd DW (1970) Prog Med Virol 12 , 192 Replicates in cell cultures of embryonic equine tissue without CPE. Interference equine foamy virus (EFV) A tenta- with vesicular stomatitis virus mul- tive species in the genus Spumavirus. tiplication has been used for assay. Isolated from blood samples of horses Replication in horse leukocyte cultures with no pathological signs. Induces is accompanied by CPE. The virus is syncytia formation in many cell lines of being used as a model lentivirus for different species (hamster, rabbit, sim- vaccine development. ian and human). The genome sequence Synonyms : infectious anemia of horses resembles that of other spumaviruses, virus; swamp fever virus. but is clearly distinct. Carpenter S and Alexandersen S (1992) Semin Virol 3 , 157 Tobaly-Tapiero J et al (2000) J Virol 74 , 4064 Cheevers WP and McGuire TC (1988) Adv Virus Res 34 , 189 equine herpesvirus 1 Synonym for Equid Craigo JK et al (2007) Proc Natl Acad Sci 104 , herpesvirus 1 . 15105 Montelaro RC et al (1993) In The Retroviridae , equine herpesvirus 2 Synonym for Equid edited by JA Levy. New York: Plenum Press, herpesvirus 2 . p. 257 equine herpesvirus 3 Synonym for equid equine infectious arteritis virus Syn- herpesvirus 3. onym for Equine arteritis virus . equine herpesvirus 4 Synonym for Equid equine influenza virus A poor term herpesvirus 4 . because it has been used for three different viruses: Equid herpesvirus 1 , equine herpesvirus 5 Synonym for Equid Equine arteritis virus and influenza herpesvirus 5 . virus A equine.

Equine infectious anemia virus (EIAV) A equine morbillivirus (EMV) See Hendra species in the genus Lentivirus causing virus . acute or chronic infections in horses. The disease was first described in Equine papillomavirus 1 (EqPV-1) 1843. Incubation period 12–15 days The type species of the genus or longer. There are acute episodes Zetapapillomavirus. A natural infection with fever, anemia, nasal discharge, of horses causing papillomas on the and subcutaneous edema. Remissions nose and lips, but can be transferred occur which may last for years but to the skin. Causes papillomas experi- the infection is usually fatal. Viremia mentally in horses, but not in other may be present for years, even during species. Equine papillomavirus 1 (EqPV-1) 154

Synonym : horse papillomavirus. skin cells and induces CPE 24–30 h after Cook RH and Olson C (1957) Am J Pathol 27 , infection. The virus contains a large 1087 positive-strand RNA genome (25–30kb) and induces a 3 co-terminal nested set equine parvovirus A possible species in of five mRNAs in infected cells. the genus Parvovirus. Isolated from Synonym : Berne virus. the liver of an aborted equine fetus. Koopmans M and Horzinek MC (1994) Adv Hemagglutinates guinea pig, rhe- Virus Res 43 , 233 sus monkey, and fowl erythrocytes. Snijder EJ et al (1990) J Virol 64 , 331 Antibodies were detected in horse sera. equus caballus papillomavirus 1 A strain Wong FC et al (1985) Can J Comp Med 49 , 50 of equine papillomavirus 1 .

Equine rhinitis A virus (ERAV) A spe- eradication The total elimination of cies in the genus Aphthovirus. Causes a virus species from the world. No upper respiratory tract infections of virus has yet been totally eradicated. horses whereas all other members of Smallpox (variola virus) was declared the genus cause foot-and-mouth dis- by WHO to have been eradicated in ease. Included in the genus because 1980, but the virus is still stored in of similarities in genome organization two repositories in Atlanta, USA and and structure to foot-and-mouth dis- Novosibirsk, Russia. Plans are under- ease virus. way to eradicate poliovirus and mea- Synonym : equine rhinovirus 1. sles virus within the next few years.

Equine rhinitis B virus (ERBV) The only erb A An oncogene present in the genome species in the genus Erbovirus. Causes of avian erythroblastosis virus strains upper respiratory tract disease of ES4 and R. Homologous to the thyroid horses, with a viremia and fecal shed- hormone receptor gene. ding. Infections may be persistent. Subtypes include equine rhinitis B erb B An oncogene present in the genome virus 1 and equine rhinitis B virus 2. of avian erythroblastosis virus strains H, R and ES4. Homologous to the epi- equine rhinopneumonitis virus dermal growth factor receptor gene. Synonym for Equid herpesvirus 4 . Erbovirus A genus in the family equine rhinovirus 2 Former name for Picornaviridae, the type and only spe- equine rhinitis B virus 1. cies of which is equine rhinitis B virus. The genome of equine rhinitis B virus equine rhinovirus 3 Former name for contains a long 3 -untranslated region equine rhinitis B virus 2. of 167 nucleotides.

Equine torovirus (EqTV) The type species ERC-55 A calcium-binding protein of of the genus Torovirus. Isolated in 1972 unknown function which binds the E6 from a horse with diarrhea in Berne, oncoprotein of human papillomavirus. Switzerland. However, the virus was The mouse homologue of ERC-55 is the not incriminated; Salmonella lille , iso- vitamin D receptor-associated factor lated from the same swab, was believed that transduces the growth suppressive to have caused the disease from which effects of vitamin D. the horse died a week later. A similar virus was isolated in 1979 from calves Eret-147 virus (E147V) A serotype with diarrhea in Breda, Iowa, USA. The of Nyando virus in the genus Orthobunya- viruses are different but antigenically virus. Transmitted by mosquitoes. related. Antibodies against toroviruses Isolated from mosquitoes, Eretmapodites are widespread in ungulates, and some sp. Not associated with disease. wild mice and laboratory rabbits have been found seropositive. Equine toro- erinaceid herpesvirus 1 (ErHV-1) virus grows in vitro in embryonic mule An unassigned virus in the family 155 escape mutant

Herpesviridae , isolated from hedgehogs, virus, but erythroblastosis occurs to Erinaceus europeus . some degree with other leukemia Synonym: European hedgehog viruses such as Friend leukemia virus. herpesvirus. Stack MJ et al (1990) Vet Rec 127 , 620 erythroblastosis virus Synonym for avian erythroblastosis virus. erosive stomatitis virus of cattle Synonym for Bovine papular stomatitis erythrocyte A mature red blood cell that virus, although other viruses may cause is no longer involved in hemoglobin the same clinical picture. synthesis. error catastrophe RNA viruses mutate erythrocytic inclusion body syndrome and there is a maximum error thresh- (EIBS) A serious disease of salmonid old compatible with the maintenance fish. Epidemics have occurred among of genetic information. When this populations of cultured Coho salmon. error threshold is exceeded, the virus There is anemia, with characteristic ceases replication because of error inclusion bodies in the erythrocytes, catastrophe. One approach to antiviral and a yellowish liver. Probably caused therapy is to increase the rate of muta- by a togavirus, but this is not certain. genesis. The antiviral drug ribavirin has been shown to be mutagenic, and Nakajima K et al (1998) Fish Pathol 33 , 181 mice treated with the mutagen 5-fluorouracil became resistant to the estab- erythrodermatitis of carp The chronic lishment of a persistent infection with form of infectious dropsy. Not caused lymphocytic choriomeningitis virus. by rhabdovirus carpio. The etiological agent has not yet been identified. Crotty S et al (2001) Proc Natl Acad Sci 98 , 6895 Ruiz-Jarabo CM et al (2003) Virology 308 , 37 erythroleukemia A form of acute myeloid leukemia which involves eryth- error frequency Since RNA-dependent roblast cells. RNA polymerases do not have proofreading activity, their genetic variabil- Erythrovirus A genus in the subfamily ity is much higher for RNA viruses as Parvovirinae, family Parvoviridae. The compared to DNA viruses. The error type species is B19 virus. Populations frequency of RNA-dependent RNA of mature virions contain both posi- polymerases is 10 3 to 10 4 . tive- and negative-sense DNA strands, 5 kb in length, which usually form dou- Erve virus (ERVEV) A strain of ble-stranded DNA upon extraction. Thiafora virus in the genus Nairovirus. The DNA contains long, inverted ter- Transmitted by ticks and isolated in minal repeats of 365 nucleotides, and France and the UK. Not known to transcripts are extensively spliced to cause disease in humans. produce mature mRNAs. Replication Chastel C et al (1985) Bull Soc Pathol Exp 78 , occurs in primary erythrocyte precur- 594 sor cells and primary umbilical cord erythrocytes. The cellular receptor is erythema infectiosum (fifth disease) A erythrocyte P antigen (globoside). childhood disease with an erythema- Anderson LJ and Young NS (1997) Monogr tous rash (slapped cheek syndrome) Virol 20 , 153 and transient polyarthropathy caused by the human parvovirus, B19 virus. escape mutant A mutant virus which is resistant to neutralization by a par- erythroblast A nucleated bone-marrow ticular neutralizing monoclonal anti- cell that gives rise to red blood cells. body. By sequencing the genome of the escape mutant, the amino acid changes erythroblastosis of mice virus Has been responsible for the resistance (escape) used as a synonym for Kirsten leukemia can be determined. esocid herpesvirus 1 (EsHV-1) 156 esocid herpesvirus 1 (EsHV-1) An unas- The cytoplasm contains other mem- signed virus in the family Herpesviridae , brane-bound organelles as well as the isolated from pike, Esox lucius , with nucleus. epidermal hyperplasia. Synonyms : northern pike herpesvirus; European bat lyssavirus Species in the pike epidermal proliferative herpes- genus Lyssavirus. Two serologically virus. distinct species. Originally shown to be pathogenic for humans when a bat Yamamoto T et al (1983) Arch Virol 79 , 255 biologist died from EBLV infection in 1985. The two serotypes are distrib- Essaouira virus (ESSV) A serotype of uted throughout Europe from Spain to Chenuda virus in the genus Orbivirus. Russia. EBL-1 virus has been isolated Isolated from ticks, Ornithodorus mariti- from Eptisicus serotinus, and EBLV- mus , collected from seabird colonies on 2 from Myotis dasycneme and Myotis Essaouris Island, Morocco, in 1979. daubentonii. Shope RE (1999) In Encyclopedia of Virology , Estero Real virus (ERV) A species in the Second edition, edited by A Granoff and genus Orthobunyavirus , serologically RG Webster. London: Academic Press, p. 1442 a member of the Patois serogroup. Tordo N et al (2005) In Topley & Wilson’s Isolated from the tick, Ornithodoros Microbiology and Microbial Infections, vol. 2, tadaridae , from Cuba. Tenth edition, BWJ Mahy and V ter Meulen. London: Hodder Arnold, p. 1919 van der Poel WHM et al (2000) Arch Virol Etapapillomavirus A genus of the fam- 145 , 1919 ily Papillomaviridae containing avian papillomaviruses. The type spe- European bat lyssavirus 1 (EBLV-1) A cies is Fringilla coelebs papillomavirus species in the genus Lyssavirus , anti- (Chaffinch papillomavirus). genically related to Rabies virus. See European bat lyssavirus . ethidium bromide A chemical which intercalates between base pairs of a European bat lyssavirus 2 (EBLV-2) A nucleic acid and can be used to detect species in the genus Lyssavirus , anti- nucleic acid in gels due to its bright flu- genically related to Rabies virus. See orescence in UV light at a wavelength European bat lyssavirus . around 300 nm. European brown hare syndrome virus (EBHSV) A species in the genus ets gene An oncogene present in the Lagovirus. Has occurred in wild and genome of avian myeloblastosis virus farmed brown hares, Lepus europaeus , strain E26. This virus also carries the and varying hares, Lepus timidus , in myb oncogene. many European countries since 1980. Causes a hemorrhagic disease with Eubenangee virus (EUBV) A species in the liver necrosis and up to 90% mortal- genus Orbivirus and the type member ity in adult hares. EBHS is caused by a of the Eubenangee serogroup. Isolated non-enveloped virus similar to rabbit in northern Australia from mosquitoes. hemorrhagic disease virus with which Antibodies found in wallabies and kan- it shares several antigenic determinants garoos. Not reported to cause disease ( Table E1 ). in humans. Chasey D et al (1992) Arch Virol 124 , 363 Gavier-Widen D (1994) Vet Pathol 31 , 327 eukaryotic cells Animal or plant cells, Wirblich C et al (1994) J Virol 68 , 5164 which differ from prokaryotes in having cell nuclei with nuclear envelopes European Collection of Animal Cell and more than one chromosome, as Cultures An international repository well as nuclear divisions (mitosis and for reference cell lines and microbio- meiosis) during which the chromo- logical reagents. Can be reached at somes can be seen by light microscopy. http://www.ecacc.org.uk . 157 exogenous virus

Table E1. Strains of European brown hare deletions of the pol gene and parts of syndrome virus the env and gag genes. European brown hare syndrome virus-BS89 Ruis BL et al (1999) J Virol 73 , 5345 (EBHSV-BS89) European brown hare syndrome virus-FRG evolution A change in the genetic compo- (EBHSV-FRG) sition of a population, involving muta- European brown hare syndrome virus-GD tion and selection. (EBHSV-GD) European brown hare syndrome virus-UK91 evolutionary tree A diagram depict- (EBHSV-UK91) ing the evolutionary relationships of virus nucleic acid or protein sequences. The topology (branching order) and branch lengths are proportional to the European elk papillomavirus (EEPV) calculated evolutionary distances. See The type species of the genus phylogenetic tree . Deltapapillomavirus causing cutaneous fibropapillomas, especially in European exanthem A widespread rash, usually of elk in Sweden. Causes fibrosarcomas in viral origin. The six classical childhood hamsters. exanthems are: (1) Rubeola caused by European encephalitis virus A subtype measles virus, (2) varicella caused by of Tick-borne encephalitis virus in the varicella-zoster virus, (3) German mea- genus Flavivirus. sles caused by rubella virus, (4) scarlet fever caused by Streptococcus group European ground squirrel cytomegalo- A bacteria, (5) Erythema infectiosum virus Synonym for sciurid herpesvirus 1. caused by B19 virus, and (6) Roseola or exanthem subitum caused by human Barahona HH (1975) Lab Anim Sci 25 , 725 herpesvirus 6.

European harvest mouse papilloma- exanthem subitum A common febrile ill- virus A probable species in the family ness of early childhood. Also known Papillomaviridae , found in skin lesions as roseola. The sixth of the traditional including papillomas of European har- exanthems of childhood. After a 3-day vest mice, Micromys minutus. febrile prodrome, a faint pink macu- O’Banion MK et al (1988) J Virol 62 , 226 lopapular rash develops. Caused by human herpesvirus 6. In the USA, European hedgehog herpesvirus approximately 12–30% of children have Synonym for erinaceid herpesvirus 1. clinical manifestation of the disease, but 86% of children are seropositive by European swine fever virus Synonym age 1 year, and almost 100% by 4 years for Classical swine fever virus . of age.

EV virus Abbreviation for enterovirus. excision Enzymatic removal of a nucle- Usually followed by the isolate number, otide or polynucleotide from a nucleic e.g. EV70. acid polymer. EV2 virus Synonym for cauliflower disease of eels virus. excess mortality A concept developed in 1885 by William Farr which is the Everglades virus (EVEV) A species in the number of deaths observed during an genus Alphavirus. Isolated from the cot- epidemic of influenza-like illness in ton mouse, rat, and Culex sp in the excess of the number expected. Everglades National Park, USA. Cases Simonsen L et al (1997) Am J Public Health 87 , of infection, with fever and headache, 1944 have been reported in humans. exogenous virus A retrovirus which is ev/J A family of endogenous retroviruses not present in the germ line DNA of found in all chicken cells. They have the species in which it is found. exon 158 exon A region of the genome nucleic acid Table E2. Strains of Eyach virus of a virus or a cell which is expressed in mature mRNA. The exons of split Eyach virus (France-577) (EYAV-Fr577) genes are interrupted in the DNA by Eyach virus (France-578) (EYAV-Fr578) Eyach virus (Germany) (EYAV-Gr) introns (intervening sequences). The initial transcript of such genes is processed by splicing to remove the introns, yielding functional mRNA. See also to Colorado tick fever virus. Isolated intron and splicing . from ticks, Ixodes ricinus , collected Gilbert W (1978) Nature 271 , 501 near the village of Eyach in the Neckar Valley, Germany, and from I ricinus and I ventalloi in France. Infectivity resists exonuclease A phosphodiesterase which treatment with ether or deoxycholate degrades nucleic acids stepwise either but is sensitive to chloroform at room from the 3 end or from the 5 end. temperature for 30 min. Examples are: exonuclease I, which acts specifically on single-stranded DNA in Chastel C et al (1984) Arch Virol 82 , 161 the 3 to 5 direction; exonuclease III, Rehse-Kupper B et al (1976) Acta Virol, which has several activities including Prague 20 , 339 double-strand-specific 3 to 5 exonu- eye disease caused by viruses clease, and an endonuclease activity Viruses which acts on apurinic sites in DNA; from at least 10 families can cause ocu- and exonuclease V, which acts from lar disease in humans. In most cases both 3 and 5 ends of linear double- infection spreads from the upper respi- or single-stranded DNA, and requires ratory tract or skin. ATP for its activity. (1) Adenoviridae. Human species cause: (a) epidemic keratoconjunctivitis; (b) pharyngoconjunctival fever; and exotic ungulate encephalopathy See (c) acute follicular conjunctivitis. prion diseases . (2) Papovaviridae. Human species causes papillomas on the lids. expressed genes The basic functional (3) Herpesviridae. Human herpesvirus units of DNA, identified by cloning 1 and 2 cause keratoconjunctivi- mRNAs and sequencing individual tis. Varicellavirus: in chickenpox, cDNAs. Also called ESTs or ‘ expressed about 4% of cases have some cor- sequence tags. ’ neal or conjunctival involvement; Martin KJ and Pardee AR (2000) Proc Natl in shingles, vesicles may occur on Acad Sci 97 , 3789 the cornea and result in scarring. In congenital cytomegalovirus infec- expression vector A cloning vector tion the virus can be demonstrated which expresses protein from geneti- in the retina at post-mortem. Ocular cally inserted foreign genes. The vector involvement is a rare complication normally contains a strong promoter of infectious mononucleosis. which is inducible and a strong initia- (4) Poxviridae . Four species may affect tion site for translation. Both eukaryo- the eyes: variola virus (now eradi- tic and prokaryotic vectors can be cated), vaccinia virus, Orf virus, used; their suitability may depend and Molluscum contagiosum virus. upon the need for processing of the (5) Togaviridae. Rubella virus infection protein product (e.g. cleavage or gly- during pregnancy is an important cosylation). Commonly used expres- cause of congenital eye disease. sion vectors for virus genes include (6) Flaviviridae . Dengue and yellow those derived from Escherichia coli , vac- fever viruses often cause severe cinia virus, the baculovirus Autographa deep pain behind the eyes. californica nuclear polyhedrosis virus (7) Orthomyxoviridae . In some influ- (AcMNPV) and yeast. enza epidemics conjunctivitis may be a common complication. Eyach virus (EYAV) A species in the (8) Paramyxoviridae . In measles photo- genus Coltivirus antigenically related phobia is a characteristic symptom. 159 eye disease caused by viruses

Mumps virus may rarely cause (10) Retroviridae . HIV causes the focal blindness due to nerve damage. retinal ischemia (cotton–wool Newcastle disease virus can cause spots) seen in AIDS patients. conjunctivitis. Chodosh J et al (2008) in BWJ Mahy and (9) Picornaviridae . Paralysis of oculo- MHV van Regenmortel (eds) Encyclopedia motor nerves can occur in polio- of Virology, Third edition. Oxford: Academic virus infection. Coxsackie virus Press, vol. 2, p. 491 A24 causes epidemic conjuncti- Pomerantz RJ et al (1987) N Engl J Med 317 , vitis and enterovirus 70 causes 1643 acute hemorrhagic conjunctivitis. Ritterbrand DC and Friedberg DN (1998) Conjunctivitis is common in rhi- Rev Med Virol 8 , 187E novirus infections. F

F factor See F plasmid . by the formation of certain inclusion bodies visible by light microscopy. F plasmid A conjugative bacterial plasmid (94.5 kb) which mediates the FAIDS virus See Feline immunodefi- Escherichia coli mating process. Encodes ciency virus . information leading to the formation of sex pili (F pili) on the cell surface, DNA Fakeeh virus See Alkhurma virus . transfer to cells that do not contain an F plasmid, and prevention of self-mating. falcon inclusion body disease virus F plasmid-containing cells are denoted Synonym for falconid herpesvirus 1. male, recipient cells female. Some bacteriophages, such as MS2 or fd phages, falconid herpesvirus 1 (FaHV-1) An unas- adsorb exclusively to F pili. signed virus in the family Herpesviridae . Synonym : falcon inclusion body disease F protein Fusion protein, encoded in virus. the genome of paramyxoviruses. Synthesized in the infected cell as a Famciclovir (FCV) A diacetyl-6-deoxy precursor (F0) which is activated after derivative of the acyclic nucleoside cleavage by cellular proteases to pro- penciclovir that has better absorption duce the virion disulfide-linked F1 and in humans and acts as a prodrug for F2 subunits (amino F2-S-S-F1 carboxyl). penciclovir following oral administration. Active against herpesviruses such Lamb RA (1993) Virology 197 , 1 as HSV1, HSV2, and VZV. Synonym : Famcyclovir. FA tests Fluorescent antibody tests. See fluorescent immunoassay . Cirelli R et al (1996) Antiviral Res 29 , 141 Field HJ et al (1995) Antimicrob Agents Fab fragment The antigen-binding frag- Chemother 39 , 1114 ment of the immunoglobulin molecule that is obtained following hydrolysis familial prion disease Some 10% of cases with papain. A slightly larger fragment of Creutzfeldt–Jakob disease are inher- (Fab ’ ) is obtained following treatment ited. It is now possible to associate a with pepsin followed by reduction. number of mutations in the prion gene Each of these fragments consists of an with inherited prion diseases. intact light chain attached by a disulfide bond to the Fd fragment of one heavy Farallon virus (FARV) A strain of Hughes chain. If the immunoglobulin is treated virus in the genus Nairovirus . Isolated with pepsin without subsequent reduc- from ticks, Ornithodorus capensis from tion, the fragment obtained is a dimer gull nests on the Farallon Islands. of two Fab ’ fragments, containing both Gould EA et al (1983) J Gen Virol 64 , 739 antigen-binding sites, and is termed the F(ab’ )2 fragment. Far East Russian encephalitis virus Synonym for tick-borne encephalitis virus Facey’s Paddock virus (FPV) A strain (Far Eastern subtype). of Oropouche virus in the genus Orthobunyavirus . farnesylation The addition of three iso- prene units (farnesyl group) to a pro- factories Regions within the cell nucleus tein. The farnesyl group is specifically or cytoplasm where virus assembly can attached to proteins that have the C- be detected by electron microscopy, or terminal motif CAAX, and is believed 161 Feline calicivirus (FCV)

to play a role in membrane attachment. mild or subclinical, though pregnant This modification is essential for virion queens may abort. Causes similar clini- formation of hepatitis delta virus. cal disease in wild felids such as lions Otto JC and Casey PJ (1996) J Biol Chem 271 , (Panther leo ) and cheetahs (Acinonyx 4659 jubatus). Focal lesions are produced in cell cultures from cat kidney, lung, and fas-mediated apoptosis The 81 amino testis. There is no CPE in cultures of acid HIV-1 Vpu protein interferes with bovine, human, or monkey cells. MHC class 1 presentation and regu- Synonyms: feline herpesvirus 1; feline lates Fas-familial insomnia . See prion rhinotracheitis virus. diseases . Driciru M et al (2006) J Wildl Dis 42 , 667 Gaskell R and Willoughby K (1999) Vet fat head minnow cells See FHM cells . Microbiol 69 , 73 van Vuuren M et al (1999) J S Afr Vet Assoc FBJ osteosarcoma virus See Finkel– 70 , 132 Biskis–Jinkins murine sarcoma virus . felid herpesvirus 2 (FeHV-2) Probably Fc fragment The crystallizable fragment a strain of Bovine herpesvirus 4 in the of the immunoglobulin molecule that genus Rhadinovirus . A cell-associated is obtained following treatment with herpesvirus isolated during studies of papain. Contains portions of two heavy feline urologic syndrome. Induced for- chains and no antigen-binding sites. mation of syncytia, and nuclear inclusions which were found to contain Fcγ RIII (CD16) Natural killer cells can herpes virions by electron microscopy. recognize IgG antibody-coated cell sur- Intracellular and extracellular crys- faces via Fcγ RIII (CD16) and kill target tals were formed, some of which were cells. identified as cholesterol. More recent restriction endonuclease and fluores- Fc receptor A cell surface component cent antibody analyses indicate a close which binds to the Fc portion of immu- relationship of the feline cell-associated noglobulin molecules. virus to bovine herpesvirus 4. Fabricant CC and Gillespie JH (1974) Infect Fc2Lu cells (CCL 217) A cell line derived Immun 9 , 460 from normal lung tissue of a near full- Gaskell RM et al (1979) Vet Rec 105 , 243 term male cat fetus. Kruger JM et al (1990) Am J Vet Res 51 , 879

Fc3Tg cells (CCL 176) A cell line, estab- feline agranulocytosis virus Synonym lished from normal embryonal tongue for Feline panleukopenia virus . tissue of a female cat, which is susceptible to a wide variety of known feline Feline astrovirus (FAstV) A species in the viruses. genus Mamastrovirus. Only one serotype has been detected. fecal-oral transmission An important mode of transmission for numerous Lukashov VV and Goudsmit J (2002) J Gen enteroviruses, as well as hepatitis A Virol 83 , 1397 and E viruses. feline astrovirus 1 (FAstV-1) A serotype Felid herpesvirus 1 (FeHV-1) A species of Feline astrovirus . in the genus Varicellovirus. Cats which have recovered from infection with this Feline calicivirus (FCV) A species in the virus may still carry it and infect kit- genus Vesivirus . Several serotypes are tens, in which it causes nasal discharge, described. Most strains have been iso- lacrimation, and fever. Virus replicates lated from the respiratory tract but some in the mucous membranes of the nose, may be associated with the gastroin- larynx and trachea, also in the conjunc- testinal tract. Unlike many other calici- tiva, and can infect the genital tract. In viruses, feline calicivirus can be grown cats older than 6 months, the disease is in feline cell culture. The virus attaches Feline calicivirus (FCV) 162

to (alpha) 2.6-linked sialic acid on the by an endogenous cat type C retrovi- cell surface. Experimental exposure to rus obtained from a continuous cat cell aerosol of virus caused rhinitis, conjunc- line CCC. It does not have the group- tivitis, oral ulceration, and pneumonia. specific antigen of the exogenous cat Infection most often seen in catteries, leukemia virus but does have that of may be mild or severe, even fatal. An RD 114 virus. RD 114-related nucleic attenuated strain is used as a protective acid sequences are found at about one vaccine, given intranasally or i.m. complete copy per cell in the six Felis Synonyms : cat flu virus. species of the family Felidae , although Carter MJ et al (1992) Virology 190 , 443 they are also present in Old World Gore TC et al (2006) Vet Ther 7 , 213 monkeys and apes (see baboon endog- Stuart AD and Brown TD (2007) J Gen Virol enous virus ). It is a xenotropic virus, 88 , 177 which can be induced by treatment of CCC cells with idoxuridine. The second feline calicivirus CFI/68 (FCV-CFI/68) A group, endogenous feline leukemia strain of Feline calicivirus . viruses, is very similar to exogenous feline leukemia viruses, and probably feline calicivirus F9 (FCV-F9) A strain of represents multiple integrations of the Feline calicivirus . exogenous virus. The main difference between the endogenous and exog- feline cell-associated herpesvirus enous viruses is in the U3 region of the (FCAHV) A strain of Bovine herpesvirus LTR. These endogenous viruses have 4 . See also felid herpesvirus 2 . not been isolated, but are detectable because of their recombination with Feline coronavirus (FcoV) A group exogenous viruses. The third group, 1 species in the genus Coronavirus . represented by a newly described Causes feline infectious peritonitis. virus, FcEV ( Felis catus endogenous ret- Antigenically related to porcine trans- rovirus), is a type C virus present at 15– missible gastroenteritis virus but infec- 20 copies per cell, which probably gave tion with transmissible gastroenteritis rise to RD 114 virus by recombination virus does not give immunity to infec- with a baboon endogenous retrovirus, tious peritonitis. Cats of any age, chee- from which it acquired the env gene. tahs, leopards, and other large cats are Sheets RL et al (1993) J Virol 67 , 3118 susceptible. Causes a gradual loss of van der Kuyl AC et al (1999) J Virol 73 , 7994 appetite, wasting, and abdominal distension due to fibrinous peritonitis. Fatal feline enteritis virus Synonym for Feline in a few weeks. There is often pleurisy panleukopenia virus . and necrotic inflammatory lesions in many organs. Infective virus is present in ascitic fluid and organ extracts. There Feline foamy virus (FFV) A species in the is hypogammaglobulinemia and there genus Spumavirus . Has been isolated may be meningo-encephalitis and pano- from normal cats and from cats with phthalmitis. Recombinants, known as various diseases in at least three con- type II strains, have been found natu- tinents. There are two serotypes. Not rally which arose by recombination with known to cause disease. Replicates in canine coronavirus. feline embryo cell cultures. Infected Synonym: Feline infectious peritonitis virus . cells contain infectious proviral DNA of mol. wt. 6 1 06 . Late in the course Herrewegh AA et al (1995) Virology 212 , 622 of infection, the provirus is integrated Horzinek MC and Osterhaus ADME (1979) into the host cell genome. Arch Virol 59 , 1 Synonym : Feline syncytial virus. Kennedy MA (2006) Am J Vet Res 67 , 627 Lai MMC and Cavanagh D (1997) Adv Viru s Chiswell DJ and Pringle CR (1979) J Gen Res 48 , 1 Virol 44 , 145 Flugel R (1991) J AIDS 4 , 739 feline endogenous retrovirus There are at least three groups of endogenous feline herpesvirus 1 Synonym for Felid feline retroviruses. One is represented herpesvirus 1 . 163 Feline leukemia virus (FeLV)

Feline immunodeficiency virus – Oma feline infectious enteritis virus Synonym (FIVO) A species in the genus Lentivirus , for Feline panleukopenia virus . isolated in 1995 from a captive Pallas ’ cat, Otocolobus manul , in the USA. A feline infectious peritonitis virus (FIPV) highly cytopathic virus which shares a Synonym for Feline coronavirus . major core protein antigen with FIV-P, but differs more than 20% in sequence Feline leukemia virus (FeLV) A replication- homology based on the pol gene. competent species in the genus Barr MC et al (1995) J Virol 69 , 7371 Gammaretrovirus . A common infection of cats causing leukemia and/or sarco- Feline immunodeficiency virus (Petuluma) mas. Depresses immune system leading (FIV-P) A species in the genus Lentivirus , to a variety of opportunistic infections. isolated in 1987 from cats with AIDS- Virus replicates in cells of feline, human, like illness (feline AIDS). Probably an canine, and pig origin. The viruses rep- endemic infection of domestic cats licate in feline fibroblast cells in vitro throughout the world. Similar viruses and are classified into subgroup A, B, have been isolated from wild felids (e.g. or C according to their interference lion, puma, Tsushima cat) in Africa, patterns in vitro. Subgroup A viruses Japan, and North America but are not are restricted to growth in feline cells, known to cause disease in their host whereas subgroup B and C viruses will species. The virus is present in saliva of also grow in canine, human, or mink infected cats and transmitted by bites. cells. Replication is not cytopathic and Venereal transmission has not been virus can be propagated for long peri- demonstrated. The virus infects cells of ods in feline fibroblast cells. Persistently the monocyte–macrophage system and infected cats shed virus in saliva, urine 4–8 weeks after infection causes a pri- and feces and young kittens are easily mary mononucleosis-like disease with infected up to 4 months old. A major- fever and leukopenia, usually followed ity of infected cats shed virus for up to 3 by recovery. This primary disease may months before developing neutralizing be more severe in cats that are coinfected antibody, but up to 5% of cats remain with feline leukemia virus. It is fol- persistently infected and shed virus lowed by a gradual decline in CD4 for a few years before disease devel- T-cell counts over a period of months ops. Over 80% of these persistently to years until the immune system fails. infected cats will die within 3–4 years. Antibodies to several virion proteins Commonly, the disease involves a T-cell appear in the first few weeks after infec- lymphosarcoma or thymic atrophy lead- tion and can be used in ELISA, IFA, or ing to immunosuppression and con- Western blot tests to diagnose infec- sequent degenerative diseases. Feline tion. In vitro, the virus grows in CD4 leukemia viruses frequently undergo or CD8 primary T-lymphoblast cells as recombination with host cell genes and well as peritoneal macrophages. As with yield replication-defective recombinant HIV, there appear to be many genetic feline sarcoma viruses in which one of variants which may coexist even in a at least seven cellular oncogenes has single host, and have different growth been transduced by FeLV. Several pro- properties on various cell lines. Five tective vaccines have been developed, main subtypes, A–E, are recognized by including a canarypox virus-vectored sequence analysis of the viral env and recombinant FeLV vaccine. gag genes. Provides a useful model for Feline sarcoma virus isolates and their AIDS studies. respective host oncogenes are: Bendinelli M et al (1995) Clin Microbiol Rev 8 , 87 Gardner–Arnstein GA-FeSV (c-fes ) Elder JH and Phillips TR (1995) Adv Virus Gardner–Rasheed GR-FeSV (c-fgr ) Res 45 , 225 Hardy–Zuckerman HZ1-FeSV (c-fes ) Nishimura Y et al (1998) Virus Res 57 , 101 Hardy–Zuckerman HZ2-FeSV (c-abl ) Hardy–Zuckerman HZ4-FeSV (c-kit ) feline infectious leukocytosis virus Hardy–Zuckerman HZ5-FeSV (c-fms ) Synonym for Feline panleukopenia virus . Noronha–Youngren NY-FeSV (c-ki-ras ) Feline leukemia virus (FeLV) 164

Parodi–Irgens PI-FeSV (c- sis ) Parrish CR (2008) in BWJ Mahy and MHV Snyder–Theilen ST-FeSV (c- fes ) van Regenmortel (eds) Encyclopedia of Susan–McDonough SM-FeSV (c-fms ) Virology , Third edition, Oxford: Academic Theilen–Pedersen TP1-FeSV (c- fgr ) Press, vol. 4, p. 85 Synonyms: cat type C oncovirus; feline sarcoma virus. Feline papillomavirus (FdPV) A species in the genus Lambdapapillomavirus , which Grosenbaugh DA et al (2006) J Am Vet Med causes mucosal and cutaneous lesions Assoc 228 , 726 in cats. Papillomavirus particles can Hardy WD (1993) In The Retroviridae, vol. 2, be seen by electron microscopy in cells edited by JA Levy. New York: Plenum Press, of the stratum granulosum. Persistent p. 109 Hofmann-Lehmann R et al (2006) Vaccine 24 , infection has also been reported. 1087 Carney HC et al (1990) J Vet Diagn Invest Rohn JL et al (1996) Leukemia 10 , 1867 2 , 294 Lozano-Alcaron F et al (1996) J Am Anim Hosp Assoc 32 , 392 Feline panleukopenia virus (FPLV) A species in the genus Parvovirus . A natural feline pneumonitis virus Not a virus. infection of felids (domestic cats, lions, A species of chlamydia ( Chlamydia and tigers), that has now adapted to psittaci ). growth in other species, so strains are recognized from canine, mink, and rac- feline respiratory viruses The clinical coon. In zoos usually the smaller species picture of acute upper respiratory dis- (raccoons, mink, and foxes) are suscep- ease in the domestic cat may be caused tible. Chiefly seen as a severe febrile by a number of different viruses, felid illness, with vomiting and sometimes herpesvirus 1, feline calicivirus and blood-stained diarrhea in young cats, feline panleukopenia virus, and by although older cats may be attacked feline pneumonitis agent, which is a when virus is first introduced to a pre- chlamydia. viously virus-free group. Subclinical and mild cases probably occur and give feline rhinotracheitis virus Synonym for immunity. Infected animals may excrete Felid herpesvirus 1 . virus for a year, and virus contaminat- ing the environment may remain infec- feline sarcoma and leukemia viruses See tious for months. Kittens infected before Feline leukemia virus . 9 days of age may suffer damage to the developing cerebellum and at 3–4 weeks feline spongiform encephalopathy (FSE) show ataxia and tremors. Cats, mink, See prion diseases . and newborn ferrets can be infected experimentally. After an initial leuko- feline viral rhinotracheitis virus Synonym cytosis there is a progressive fall in cir- for Felid herpesvirus 1 . culating lymphocytes and polymorphs, with lethargy and anorexia. Virus repli- Felis catus endogenous retrovirus See cates in kitten kidney cell cultures. CPE feline endogenous retrovirus . may be transient. Virus replicates best in rapidly dividing cells. Some strains Fen1 A 48kD cellular protein that is a have weak hemagglutinins which bind 5 -3 exonuclease and removes RNA to pig erythrocytes at 4°C. An attenu- primers during the replication of SV40 ated virus vaccine gives protection. virus. Synonyms: ataxia of cats virus; cat distemper virus; cat fever virus; cat plague fer-de-lance virus (FDLV) An unassigned virus; enteritis of mink virus; feline virus in the family Paramyxoviridae . agranulocytosis virus; feline enteritis Isolated from the lungs of fer-de-lance virus; feline infectious aleukocytosis tropical American pit vipers, Bothrops virus; feline infectious enteritis virus; atrox , from a snake farm in Switzerland Feline parvovirus ; show fever virus. where there was an outbreak of lethal respiratory disease among the snakes. Parrish CR (1990) Adv Virus Res 38 , 403 Replicates with CPE in a wide variety of 165 fievre typhoide du cheval virus

reptilian, piscine, and mammalian cells FHM cells (CCL 42) A heteroploid cell at 30°C. The CPE varies in different cell line derived from tissue posterior to the types; in some, syncytia are a promi- anus, exclusive of the caudal fin, of nor- nent feature. Infected cells hemadsorb mal adult fat head minnows, Pimephales guinea pig erythrocytes. Replicates in promelas, of both sexes. Supports growth hens ’ eggs at 27°C; at first only on injec- of infectious pancreatic necrosis virus, tion into the amniotic cavity but adapts frog virus 1 and some other fish to the allantoic cavity. Agglutinates viruses. chicken and guinea pig erythrocytes and has neuraminidase activity. No antigenic FHs 74 Int cells (CCL 241) A cell line relationship demonstrable by CF test to established from the small intestine of other paramyxoviruses, except a para- an apparently normal, 3- to 4-month- myxovirus that caused epidemics of lung old human female fetus. disease in caiman lizards, Dracaena gulan- ensis , imported into the USA from Peru Fialuridine A pyrimidine analog (2- in 1999. The complete sequence of FDLV fluoro5-iodoarabinofuranosyluracil) revealed similarity to other paramyxovi- used as a potential therapy for hepati- ruses except for a new gene (called U for tis B infection. Withdrawn because of unknown) encoding a 19.4 kda protein multi-organ toxicity with several fatali- that appears to be unique. On this basis ties during early clinical trials. it has been suggested that the ophid- Synonym : FIAU. ian virus represents a new genus in the Colacino JM (1996) Antiviral Res 29 , 125 Paramyxoviridae with the suggested genus name Ferlavirus . fiber diffraction A method for the analy- Clark HF et al (1979) J Gen Virol 44 , 405 sis of helical structures in viruses for Franke J et al (2001) Virus Res 80 , 67 which the sample has to be prepared Jacobson ER et al (2001) J Vet Diagn Invest with the helices all aligned in the same 13 , 143 direction. This is achieved by drying a Kurath G et al (2004) J Virol 78 , 2045 Lunger PD and Clark HF (1978) Adv Virus gel of purified virus under controlled Res 23 , 159 humidity and temperature in a strong magnetic field, then analyzing the sam- Ferlavirus A genus name suggested for ple by X-ray diffraction. ophidian paramyxoviruses such as Fer de lance virus. fibroblasts Cells derived from connective tissue that secrete fibrillar procollagen, fibronectin, and collagenase. fetal calf serum A frequent constituent of culture media used for growing animal fibroma virus See under host animal spe- cells or tissue cultures. See serum-free cies, e.g. Rabbit fibroma virus . medium . fibronectin There is evidence that fetal rhesus kidney virus Synonym for fibronectin is a cell surface receptor for Bovine polyomavirus . the salmonid rhabdovirus, viral hemorrhagic septicemia virus, and this may α -fetoprotein (AFP) A protein present prove to be a general feature of other in the serum of vertebrate embryos. rhabdoviruses. Presence in the amniotic fluid is diagnostic for spina bifida in the human fiebre amarilla virus Synonym for Yellow fetus. Serum levels of AFP are normally fever virus . less than 20 ng/ml, but may rise above 1000 ng/ml when hepatocellular carci- field mouse herpesvirus Synonym for noma (frequently associated with hep- murid herpesvirus 5. An unassigned atitis B or C virus infection) is present. virus in the family Herpesviridae .

Feulgen stain One of the few staining fievre typhoide du cheval virus Synonym methods specific for DNA. for Equine arteritis virus . fifth disease 166 fifth disease Erythema infectiosum, Peters CJ and LeDuc JW (1999) J Infect Dis which was the fifth disease in a classifi- 179 , S1–S288 cation of six erythematous rash illnesses of children. Results from infection of Filovirus See Filoviridae . young children with the parvovirus B19. See erythema infectiosum . finch circovirus (FiCV) A tentative species in the genus Circovirus . Using filipin A polyene antibiotic which inter- the polymerase chain reaction circo- acts with sterols in liposomes and virus DNA was isolated from bursa of biological membranes, producing alter- Fabricius tissue from a Gouldian finch ations in the lipid bilayer structure. (Chloebia gouldiae ). The DNA sequences Exposure of vesicular stomatitis virus were related to those of canary circo- to the drug led to a change in the per- virus, pigeon circovirus, and beak and meability barrier of the virus envelope. feather disease virus. Majuk Z et al (1977) J Virol 24 , 883 Todd D et al (2007) Avian Pathol 36, 75 Filoviridae A family in the Order finch paramyxovirus See Avian para- Mononegavirales containing two genera, myxovirus 2 . Marburgviru s and Ebolavirus . Viruses of both genera cause severe hemorrhagic fevers in humans and other primates, fingerprinting A procedure for character- with extremely high mortality (up to izing DNA, RNA or proteins by electro- 90%). Marburgvirus is a single species, phoretic or chromatographic analysis of Lake Victoria marburgvirus, whereas four specific fragments, e.g. uniformly or ter- species of the Ebolavirus genus have minally labeled RNA is digested using been recognized: Zaire, Sudan, Reston, various ribonucleases (often T1 or pan- and Côte d’Ivoire and a fifth, Ebola creatic) and the products separated by Uganda, is not yet classified. Except electrophoresis in two dimensions. The for their extreme genome length, the oligoribonucleotides are then detected viruses have a morphology somewhat by autoradiography. similar to that of members of the family Rhabdoviridae. They are filamentous, Finkel–Biskis–Jinkins murine sarcoma with U-shaped, 6-shaped or circular virus (FBJMSV) A species in the genus forms produced in cell culture. The Gammaretrovirus . A naturally occurring length is highly variable and can be as mouse sarcoma virus species (MSV). great as 14000 nm but it is usually about Isolated from a spontaneous osteosa- 800–1000 nm; the diameter is 80 nm. rcoma in the thoracic spine of a CFl The particles are enveloped with spikes mouse. Induces perosteal sarcomas ca 7 nm in length and 10 nm apart. The in mice after a short latent period (3 virus RNA is 19.1 kb in length, negative- weeks). In cell cultures produces foci of sense and is therefore noninfectious. altered cells, different in morphology There are at least seven virus pro- from those produced by other strains teins with mol. wt. of ca 267 (RNA of mouse sarcoma virus. It is accompa- transcriptase–polymerase), 75 (sur- nied by a helper leukemia virus. Unlike face glycoprotein), 78 (nucleoprotein), other strains of MSV, it transforms 32 (matrix), 31 (P protein), 32 (minor mouse embryo cells into autonomously nucleoprotein), and 29 (second matrix replicating cells; it thus produces protein). Both viruses are highly viru- plaques without having to produce lent for humans and several species of infective virus. Produces osteosarco- monkey. Despite numerous field inves- mas on injection into newborn Syrian tigations, the presumed reservoir from hamsters. Carries the transduced fos which filovirus infections originate has (f inkel o steogenic s arcoma) oncogene, not been found. which encodes a nuclear DNA-binding protein. Feldmann H and Klenk H-D (1996) Adv Virus Res 24 , 1 Finkel MP and Biskis BO (1968) Prog Exp Klenk H-D (Editor) (1999) Curr Tops Microbiol Tumor Res 10 , 72 Immunol 235, 225 Levy JA et al (1978) J Virol 26 , 11 167 FL amnion cells fin V707 virus (FINV) A strain of Rio Grande cichlid virus (Rhabdoviridae ) Uukuniemi virus in the genus Phlebovirus . sheatfish iridovirus (Iridoviridae ) Isolated from five male ticks, Ixodes smelt reovirus ( Reoviridae ) uriae , from a murre colony at Vedoy snakehead rhabdovirus ( Rhabdoviridae ) Rost Islands, Lofoten, Norway in 1975. spring viremia of carp virus Synonym : NorV-707. (Rhabdoviridae ) Striped jack nervous necrosis virus Saikku P et al (1980) J Med Entomol 17 , 360 (Nodaviridae ) fishpox virus Synonym for cyprinid her- Tiger puffer nervous necrosis virus pesvirus 1. (Nodaviridae ) viral deformity virus ( Birnaviridae ) fish viruses Viruses that affect fish occur Viral hemorrhagic septicemia virus in many virus families. In addition, a (Rhabdoviridae ) number of viruses have been isolated from fish with no known disease asso- Agents of low virulence: ciation. Here, the main disease-causing Atlantic cod epidermal hyperplasia viruses are listed; further details may (Adenoviridae ) be found under the individual entry bluegill hepatic necrosis reovirus for each virus. (Reoviridae ) fishpox virus ( Herpesviridae ) Agents of moderate to high virulence: golden shiner reovirus ( Reoviridae ) angelfish reovirus ( Reoviridae ) grouper reovirus (Reoviridae ) Barfin flounder nervous necrosis virus Lymphocystis disease virus ( Iridoviridae ) (Nodaviridae ) walleye herpesvirus ( Herpesviridae ) cichlid virus ( Rhabdoviridae ) Walleye dermal sarcoma virus ( Retroviridae ) Dicentrarchus labrax encephalitis virus white sturgeon herpesvirus (Nodaviridae ) (Herpesviridae ) eel virus European (Birnaviridae ) yellowtail ascites virus ( Birnaviridae ) erythrocytic inclusion body syndrome Bernard J and Bremont M (1995) Vet Res (Togaviridae ) 26 , 341 esocid lymphosarcoma virus (Retroviridae ) Herrick FM and Hedrick RP (1993) Annu grass carp reovirus ( Reoviridae ) Rev Fish Dis , 187 herpesvirus salmonis ( Herpesviridae ) Leong JC (2008) in BWJ Mahy and MHV van Hirame rhabdovirus ( Rhabdoviridae ) Regenmortel (eds) Encyclopedia of Virology , Ictalurid herpesvirus 1 (channel catfish Third edition, Oxford: Academic Press, vol. 3, p. 227 virus) (Herpesviridae ) Nakajima K et al (1998) Fish Pathol 33 , 181 Infectious hematopoietic necrosis virus Perez SI and Rodriguez S (1997) (Rhabdoviridae ) Microbiologia 13 , 149 Infectious pancreatic necrosis virus Walker PJ et al (2005) Disease in Asian (Birnaviridae ) Aquaculture V . Quezon City: Asian Fisheries infectious salmon anemia virus Society (Orthomyxoviridae ) Wolf K (1988) Fish Viruses and Fish Viral Japanese eel iridovirus ( Iridoviridae ) Diseases . Cornell: Cornell University Press. Japanese flounder nervous necrosis virus fitness of viruses A measure of the rela- (Nodaviridae ) tive replication capacity of a virus in a Lates calcarifer encephalitis virus given environment. (Nodaviridae ) Oncorhynchus masou virus Ball SC et al (2003) J Virol 77 , 1021 (Herpesviridae ) Carillo C et al (1998) J Gen Virol 79 , 1699 perch iridovirus (Iridoviridae ) FIV-Oma Feline immunodeficiency perch rhabdovirus disease See virus – Oma (Rhabdoviridae ) . pike fry rhabdovirus disease fixed virus Attenuated Rabies virus . (Rhabdoviridae ) red sea bream iridovirus (Iridoviridae ) FL amnion cells (CCL 62) A heteroploid Redspotted grouper nervous necrosis virus cell line derived from normal human (Nodaviridae ) amnion cells. Flanders virus (FLAV) 168

Flanders virus (FLAV) An unassigned length. Structural proteins are encoded species in the family Rhabdoviridae. With at the 5 end, and nonstructural pro- Hart Park, Kamese, Mosqueiro, and teins at the 3 end. Except for a few Mossuril viruses forms the Hart Park tick-borne flaviviruses, there is no poly serogroup. Isolated from mosquitoes, A tract at the 3 end of the genome. The Culiseta melanura and Culex pipiens, and biological properties of different viruses from an ovenbird, Seiurus aurocapillus , in the family vary widely. in New York State, USA. Serologically Flavivirus A genus in the family similar viruses have been isolated Flaviviridae , consisting of about 55 anti- from mosquitoes in Texas, California, genically related viruses. Some multiply Washington, a number of SE states of in mosquitoes and some in ticks with USA, and Canada. Replicates in and transovarial transmission, but others kills newborn mice on i.c. injection but have no known arthropod host. Some multiplies poorly if at all in embryo- replicate in eggs producing pocks on the nated eggs and cell cultures. Not CAM and in mouse embryo and HeLa known to be pathogenic in the wild. cell cultures, but CPE is not invariably Murphy FA et al (1966) Virology 30 , 314 seen. Produce encephalitis on i.c. injection in mice; most are pathogenic on Flaviviridae A family consisting of i.c. injection in rhesus and cynomolgus Flavivirus Pestivirus three genera: , , monkeys. The type species is Yellow fever Hepacivirus and . Virions are spherical, virus. They can be subdivided according 40–60 nm in diameter, with a lipid enve- to their principal vectors ( Table F1 ). lope. The genome is a single molecule of linear positive-sense, single-stranded flavivirus febricis Synonym for Yellow RNA, ranging from about 9 to 13 kb in fever virus .

Table F1. The flaviviruses

(1) The mosquito-borne species are : Kadam (I) Aroa Kyasanur Forest disease * (I, Ag, H, R, B, Ba) Bagaza (H, R) Langat (I) Banzi * (C, H) Louping-ill * (I, H, R, B) Bouboui (C, A) Meaban (I, Ag) Cacipacore (C) Omsk hemorrhagic fever * (I, H, R) Dengue (C, H) Powassan * (I, H, R) Edge Hill (C, A) Royal Farm (Ag) Ilhéus * (C, H, B) Saumarez Reef (I) Israel turkey meningoencephalomyelitis (C) Tick-borne encephalitis (I) Japanese encephalitis * (C, A, H, B, Ba) Tyuleniy (I) Jugra (C, Ba) Kokobera (C) (3) The species never isolated from wild-caught Koutango (C) arthropods: Murray Valley encephalitis * (C, H) Apoi * (R) Ntaya (C) Bukalasa bat (Ba) Saboya (C) Carey Island (Ba) Sepik * (C) Cell fusing agent St Louis encephalitis * (C, A, H, B, Ba) Cowbone Ridge (R) Tembusu (C, A) Dakar bat (Ba) Uganda S (C, B) Entebbe bat (Ba) Usutu (C, A, H, R) Jutiapa (R) Wesselsbron * (C, A, H, R) Modoc (R) West Nile * (C, A, I, Ag, H, R, B, Ba) Montana myotis leukoencephalitis (Ba) Yaounde (C) Phnom Penh bat (Ba) Yellow fever * (C, H, M) Rio Bravo * (Ba) Zika * (C, H) Sal Vieja (R) San Perlita (R) (2) The tick-borne species are : Tamana Bat (Ba) Gadget’s Gully (I) Yokose (Ba)

Isolated from: A, anopheline mosquitoes; Ag, argasid ticks; B, birds; Ba, bats; C, culicine mosquitoes; H, man; I, ixodid ticks; M, marsupials; R, rodents. * Can cause disease in humans. 169 Flury HEP virus

Flexal virus (FLEV) A species in the genus Hukkanen V et al (2000) J Clin Microbiol 38 , Arenavirus , a member of the Tacaribe 3214 complex. Isolated from Oryzomys Jackwood DJ (2004) Anim Health Res Rev rodents in Brazil in 1975. Not associated 5 , 313 Ni Q et al (2006) Methods 40 , 279 with disease in humans. fluorescent immunoassay A useful flounder lymphocystis disease virus approach to virus diagnosis, especially (FLDV) See Lymphocystis disease when virus-infected cells are avail- virus 1 . able. Specific antibodies (e.g. monoclonal antibodies) are conjugated with flounder virus See Lymphocystis disease fluorochrome dyes (e.g. fluorescein virus 1 . isothiocyanate) and used to detect viral antigens. Virus-infected cells are seen flower cells Characteristic lym- by microscopic examination. A similar phocytes with lobulated nuclei seen approach can be used to detect specific in a peripheral blood film of a patient antibodies in the host. with HTLV1-induced adult T-cell leukemia/lymphoma. fluorocarbon extraction A procedure in antigen purification to free antigens of floxuridine See fluorodeoxyuridine . serum antibodies, host cell components such as lipoproteins and microbial flumadine See rimantadine contaminants. hydrochloride . fluorodeoxyuridine A pyrimidine ana- fluorescein isothiocyanate (FITC) A flu- log and a reversible inhibitor of DNA orescent compound used for labeling synthesis. Phosphorylated by cellular proteins or nucleic acids. It is excited thymidine kinase and thus becomes an by light of wavelength in the range analog of thymidylic acid. Brings about 450–490 nm and emits in the range chromosome breaks. 520–560 nm. Synonym : floxuridine. fluorescent antibody An antibody which fluorography A technique to detect is labeled with a fluorescent dye, e.g. 3 H-labeled molecules in chromato- fluorescein isothiocyanate. It can then be grams and polyacrylamide gels. The used in conjunction with a fluorescence scintillator 2,5-diphenyloxazole (PPO) microscope to detect viral antigens. is introduced into the chromatogram or gel which is then exposed to photo- fluorescence resonance energy transfer graphic film. (FRET) A technique for quantifying Laskey RA and Mills AD (1975) Eur J nucleic acid or protein in samples. For Biochem 56 , 335 measuring the amount of DNA or RNA in a sample, real-time PCR is used to amplify the nucleic acid together fluoroimmunoassays See fluorescent with two fluorescent probes (fluoroimmunoassay . phores) with different fluorescent dyes α α that generate a signal only when the 9 -fluoro-16 -methylprednisolone See probes bind next to each other on the dexamethasone . amplicon. Has been adapted to many methods for virus detection, such as 5-fluorouracil See base analog . hepatitis B DNA in serum, herpes simplex viruses in CSF, identification of Flury HEP virus A fixed strain of Rabies mutations in infectious bursal disease virus derived from Flury LEP virus by virus dsRNA, and measuring kinase further passage in chick embryos. HEP enzyme activities in living cells. stands for h igh e gg p assage, i.e. more than 180 times. It is avirulent for adult Garson JA et al (2005) J Virol Methods laboratory mammals but causes death 126 , 207 in suckling mice. Used as a vaccine. Flury HEP virus 170

Can be propagated in human diploid focus-forming units (ffu) Units of quan- cell lines, chick embryo fibroblasts and tification for the focus-forming assay. BHK21 cells. folate receptor It has been reported that Flury LEP virus A fixed strain of Rabies folate receptor- α is a cofactor for cellu- virus . Isolated in 1939 from a girl named lar entry by the filoviruses, Ebola and Flury who contracted the disease in Marburg viruses. Georgia, USA. Brain tissue was injected Chan SY et al (2001) Cell 106 , 117 into day-old chicks, subsequently passaged in chick brain and later in eggs. follicular dendritic cells Stromal cells in LEP stands for l ow egg passage, i.e. less the spleen which act as a reservoir for than 80 times. Has been used as live prions. vaccine for dogs but is insufficiently attenuated for use in cats and cattle. FoLu cells (CCL 168) A cell line derived Can be propagated in human diploid from the normal lung tissue of an adult cell lines and in BHK21 cells. female gray fox. Susceptible to vesicular stomatitis virus, herpes simplex and FMDV See Foot-and-mouth disease vaccinia viruses. viruses . Fomede virus (FV) A strain of Chobar foamy viruses Members of the genus Gorge virus in the genus Orbivirus . Spumavirus which cause a foamy Isolated from the bat, Nycteris nana . Not appearance of the cells in which they known to cause disease in humans. replicate. Often found in primary tis- Digoutte JP (1981) Institut Pasteur Dakar sue cultures, especially following Annual Report prolonged passage. Usually cause per- Zeller HG et al (1989) Arch Virol 109 , 253 sistent infections in their natural host. There are simian and hamster species fomites Inanimate objects which may similar to the syncytial viruses of cat- serve to transmit virus infection such tle, cats, and humans. Isolated from as blankets, clothing, combs, and writ- chimpanzees and orangutans. No ing materials. confirmed association with disease in humans. ‘ Human ’ foamy virus is the Fomiversen An antiviral drug (phospho- result of rare zoonotic transmission rothioate 21-mer oligonucleotide) tar- from nonhuman primates, and is now geted to the cytomegalovirus IE2 gene. called chimpanzee foamy virus. The Used by intravitreal injection for treat- replication pathway of foamy viruses is ment of cytomegalovirus retinitis in distinct from other retroviruses, and AIDS patients. involves nuclear localization of the Gag protein. Foot-and-mouth disease viruses (FMDVA); FMDV-ASIA1; FMDV-C; FMDV-O; Lecellier CH and Saib A (2000) Virology FMDV-SAT1; FMDV-SAT2; FMDV- 271 , 1 Linial ML (2000) Trends Microbiol 8 , 284 SAT3) Seven serological types, regarded McClure MO et al (1994) J Virol 68 , 71 as a single species in the genus Aphthovirus which causes foot-and- FOCMA F eline o ncovirus-associated c ell mouth disease in cloven-hoofed ani- m embrane a ntigen. Originally it was mals. Infection is endemic in continental thought that anti-FOCMA antibody Europe, Asia, Africa, and South America, protected cats from tumor develop- but Australasia, Japan, USA, and Canada ment, but this remains controversial. are free of it. Cattle are the most commonly infected species, but pigs, sheep, focus-forming assay An assay for non- goats, deer, elephants, and hedgehogs cytolytic transforming viruses, e.g. Rous may also be infected. In cattle the dis- sarcoma virus, based on the morphol- ease is not usually fatal but causes loss ogy of the transformed cells in tissue of condition. There is fever and vesicular culture. The sites of growth of these eruption in mouth, nose, hooves, and modified cells show up as foci. udder. There may be myocardial damage. 171 Four Corners virus

In pigs lameness is the most prominent used at a lower concentration (0.015% sign; sheep and goats are less severely formaldehyde) for virus inactivation affected. The disease is extremely con- during the preparation of antigens for tagious. Guinea pigs and suckling mice immunization. Causes less protein can be infected experimentally. Virus denaturation than glutaraldehyde. multiplies in bovine, porcine, ovine, and mouse embryo cell cultures. Calf thyroid Formivirsen An antisense oligonucle- cell cultures are also often used. There are otide which is complementary to the many subtypes, and more than 40 dis- immediate early region mRNA of tinct antigenic strains are recognized. The human cytomegalovirus, a frequent virus is stable at pH 7.4–7.6 but is inacti- and serious complication of AIDS. vated below pH 6. Control is by slaughter Temsamani J et al (1997) Expert Opin Investig in nonendemic regions and by vaccina- Drugs 6 , 1157 tion elsewhere. The vaccine is commonly grown in BHK21 cells, and inactivated Fort Morgan virus (FMV) A species in the using acetylethyleneimine (AEI). The genus Alphavirus , antigenically related serotype and strain composition of the to Western equine encephalitis virus. vaccines are tailored for local require- Transmitted by swallowbugs to nesting ments. In 2000–2001, a severe pandemic cliff swallows, Petrochelidon pyrrhonota of the disease caused by a strain of sero- and house sparrows, Passer domesti- type O occurred in Korea, Japan, Russia, cus . Not known to be pathogenic for Mongolia, South Africa, the United humans. Kingdom, Republic of Ireland, France, Scott TA et al (1984) Am J Trop Med Hyg 33, 981 and the Netherlands, causing huge economic losses to these countries. In 2007 Fort Sherman virus (FSV) A serotype a small outbreak of disease in the UK of Bunyamwera virus in the genus was caused by an escape of virus from Orthobunyavirus. Causes febrile illness the International Reference Laboratory in humans with symptoms including for Foot and Mouth Disease in Pirbright, fever, malaise, muscle aches, and sore Surrey, UK but was contained within the throat. County. Synonyms : aphthous fever virus; aphtho Mangiafico JA et al (1988) Am J Trop Med Hyg virus; hoof and mouth disease; FMDV; 39 , 593 virus aftosa; le virus de la fièvre apht- euse; maul- und klauenseuchevirus. Fortovase A soft gel formulation of the retrovirus protease inhibitor saquina- Fry E et al (1990) Semin Virol 1 , 439 vir, which has increased bioavailability Knowles NJ et al (2005) Emerg Infect Dis 11 , compared to the parent drug. 1887 Mahy BWJ (2005) Curr Top Microbiol Immunol foscarnet Trisodium phosphonoformate. 288 , 1 A phosphonate analog with activity against herpes and hepatitis B viruses. Forecariah virus (FORV) An unas- Approved for intravenous therapy of signed virus in the family Bunyaviridae , cytomegalovirus-associated retinitis in serogroup 1. Isolated from the tick, immunosuppressed patients. See triso- Boophilus geigyi , in the Republic of dium phosphonoformate . Guinea. Antigenically related to Bhanja virus and Kismayo virus. Foscavir See foscarne t. Institut Pasteur, Dakar (1985) Annual Report , p. 111 Foula virus (FOUV) A serotype of Great Island virus in the genus Orbivirus . formalin A clear, colorless aqueous solu- Isolated from a pool of 10 unfed female tion containing 40% formaldehyde. A ticks, Ixodes uriae , collected from a sea- 10% solution of formalin in phosphate bird colony on Foula, Shetland Islands, buffer (4% formaldehyde) is commonly UK in 1980. used as a fixative for pathological specimens, and as a disinfectant. Also Four Corners virus See Sin Nombre virus . Fourier transformation 172

Fourier transformation An image FOS cells A cell line of fetal ovine syno- processing tool used in the analysis of vial cells that can be used to cultivate the 3D structure of viruses and their small ruminant lentiviruses such as components. maedi-visna virus and caprine arthritis encephalitis virus. Fourth disease See Duke’s disease , exanthem . fowl adenovirus 1 (CELO, 112, Phelps) (FadV-1) A serotype of fowl Fowl adenovirus A–E (FAdV-A–E) Five adenovirus A. species in the genus Aviadenovirus . Differentiated by DNA restriction fowl adenovirus 2 (GAL-1, 685, enzyme fragmentation and sequence SR48) (FadV-2) A serotype of fowl comparisons. Differ from duck adeno- adenovirus D. viruses in having two fibers at each vertex on the virion, whereas the duck fowl adenovirus 3 (SR49, 75) (FadV-3) A virus has only one. Fowl adenovirus serotype of fowl adenovirus D. A agglutinates rat erythrocytes only, whereas duck adenovirus agglutinates fowl adenovirus 4 (KR-5, J-2) (FadV-4) A chicken, turkey, and duck erythrocytes. serotype of fowl adenovirus C. Using neutralization tests, 11 serotypes can be distinguished and are wide- fowl adenovirus 5 (340, TR22) (FadV-5) spread in fowl populations. Originally A serotype of fowl adenovirus B. isolated from an often fatal outbreak of respiratory disease of quail, Colinus fowl adenovirus 6 (CR119, 168) A sero- virginianus, in USA. Chickens, tur- type of fowl adenovirus E. keys, ducks, quail, geese, guinea fowl, pheasants, pigeons, bantams, budgeri- fowl adenovirus 7 (YR36, X-11) (FadV-7) gars, and mallards are also susceptible A serotype of fowl adenovirus E. to infection. Inoculation into embryonated chicken eggs causes death of the fowl adenovirus 8a (TR59) (FadV-8a) A embryo, with necrotic foci in the liver. serotype of fowl adenovirus E. In chicken cell cultures typical CPE is produced. In the older literature strains fowl adenovirus 8b (764) (FadV-8b) A of each serotype were numbered (1–11) serotype of fowl adenovirus E. and are distributed as follows: fowl adenovirus 9 (A2-A) (FadV-9) A FAdV-A Fowl adenovirus 1 (also called serotype of fowl adenovirus D. chicken embryo lethal orphan virus, CELOV). fowl adenovirus 10 (CFA20) A serotype of fowl adenovirus C. FAdV-B Fowl adenovirus 5. fowl adenovirus 11 (380) (FadV-11) A FAdV-C Fowl adenoviruses 4 and 10. serotype of fowl adenovirus D.

FAdV-D Fowl adenoviruses 2, 3, 9, fowl calicivirus (FCV) An unassigned and 11. virus in the family Caliciviridae that has not been well characterized. FAdV-E Fowl adenoviruses 6, 7, 8a, and 8b. fowl diphtheria virus Synonym for Synonym : quail bronchitis virus; Fowlpox virus . chicken embryo lethal orphan virus. fowl leukemia virus See Avian leukosis Gelderblom H and Maiche-Lauppe I (1982) virus . Arch Virol 72 , 289 McFerran JB and Adair BM (1977) Avian Pathol 6 , 189 fowl paralysis virus Synonym for Gallid Monreal G (1992) Poult Sci Rev 4 , 1 herpesvirus 2 . 173 Friend murine leukemia virus (FrMLV) fowlpest A term used to describe infec- stretch of viral mRNA which is long tion with Newcastle disease virus but enough to code for little more than one occasionally applied erroneously also of them. The mRNA is translated from to highly pathogenic influenza A avian different starting points: A→ B and (fowl plague) virus infection. C→D. The triplet code is read in a dif- Barry RD et al (1964) Vet Rec 76 , 1316 ferent frame in each case. A stretch of viral genome can similarly code for different mRNAs. Frameshift mutations fowl plague (classical) A severe epidemic may arise from insertion or deletion of disease of poultry with up to 100% one or more nucleotides (except three mortality caused by highly pathogenic or multiples of three), and may result strains of avian influenza virus. See in termination or alteration of the pro- influenza virus A avian . tein products of a gene. Some viruses, e.g. coronaviruses and retroviruses, fowlpox subgroup viruses Synonym for employ a frameshift during replication avipoxvirus. in order to express certain genes.

Fowlpox virus (FWPV) The type species Fraser Point virus A serotype of Hughes of the genus Avipoxvirus containing a virus in the genus Nairovirus . number of strains varying in host range but otherwise not clearly separable. freeze-drying See lyophilization . Virions have a high lipid content. The genome DNA is 300 kb in length, struc- freeze-fracture A method for preparing turally similar to other poxvirus DNAs, samples for electron microscopy. They with terminal hairpin loops linking the are frozen rapidly at very low tem- two strands. The use of fowlpox virus perature and then the brittle material as a recombinant DNA vector is under is fractured. The exposed surfaces may development. Replicates on CAM probe etched to reveal further details. ducing pocks and in cell cultures of chick embryo with CPE. The disease Freund’s adjuvant A mixture of mineral caused in fowls has a course of 3–4 oil and emulsifier (and, in the complete weeks. There are proliferative lesions adjuvant, killed mycobacteria) with followed by scabbing on the skin, espe- which an antigen is emulsified before cially of the head, sometimes of feet intramuscular or subcutaneous injec- and vent. Involvement of the trachea tion. The antigen is released slowly into is called ‘fowl diphtheria.’ There may the blood stream, often leading to the be eye lesions. Many species are sus- production of a higher antibody titer. ceptible. Transmitted by contact and by mosquitoes which may remain infected Friend murine leukemia virus (FrMLV) for at least 210 days. Control is by a A strain of Murine leukemia virus in vaccine attenuated in eggs. the genus Gammaretrovirus , originally Synonyms: birdpox virus; epithelioma obtained from the leukemic spleen of contagiosum virus; fowl diphtheria a 14-month-old Swiss mouse which virus; poxvirus avium. had been injected at birth with a cell- Boyle DB and Heine HG (1993) Immunol Cell free extract of Ehrlich ascites carcinoma Biol 71 , 391 cells. Causes a disease not observed to Tripathy DN (1991) In Diseases of Poultry , occur spontaneously. Characterized by edited by BW Calnek. Ames: Iowa University gross splenomegaly. Adult mice are sus- Press, p. 583 ceptible and die of the disease in about 3–5 weeks, or longer in a few cases. A fox encephalitis virus Synonym for number of strains have been isolated: Canine adenovirus . some cause hemorrhage into the spleen and death in 3 weeks, others cause foci FPLV See Feline panleukopenia virus . of tumor cells in the spleen and a less acutely fatal disease. The virus prepara- frameshift A process by which two or tions available appear to be a mixture of more proteins can be specified by a two viruses: (1) a lymphoid leukemia Friend murine leukemia virus (FrMLV) 174

virus which does not produce the eryth- Isolated from papillomas excised from roleukemia typical of Friend virus and the leg of a chaffinch, Fringilla coelebs . (2) a replication-defective RNA tumor Virion diameter: 52 nm; sedimentation virus responsible for the leukemogen- coefficient: 300 S; and density in CsCl: esis, designated spleen focus-forming 1.34 g/ml. Composed of 72 morpho- virus (SFFV), which does not contain logical units arranged in a skew T 7 d an oncogene, the active gene product icosahedral lattice. The circular double- of which is a deleted Env protein called stranded genome measured 2.6 μ m. gp55. Presence of the virus in non- Protein composition similar to human producer cells may be detected by papillomavirus. Papillomas have only reverse transcriptase activity. The been found in two species of Fringilla : genome of SFFV consists of two sets of the chaffinch and the brambling, F. nucleic acid sequences: one homologous montifringilla . to part of the helper leukemia virus and Synonyms : avian papillomavirus; bird the other specific to SFFV, but related to papillomavirus; chaffinch papilloma- sequences in xenotropic mouse leukemia virus. viruses. Rat passage or passage at end- Osterhaus ADME et al (1977) Intervirology point dilution results in the separation 8 , 351 of the lymphoid leukemia virus. Many strains in use contain lactate frog adenovirus (FrAdV) The type spe- dehydrogenase-elevating virus as a cies of the genus Siadenovirus . Found contaminant. in a culture of the turtle cell line TH-1 Ikawa Y (1997) Leukemia 11 , S3, 152 inoculated with cells from a granu- Kabat D (1989) Curr Top Microbiol Immunol loma-bearing kidney of a leopard frog, 148 , 1 Rana pipiens . Replicated with CPE in Lee CR et al (2003) Anticancer Res 23 , 2159 turtle cells but not in cells from other Moreau-Gachelin F (2006) Haematologica 91 , reptiles, amphibia, fish, or mammals. 1644 Optimal temperature for virus replica- Steeves RA (1975) J Natl Cancer Inst 54 , 289 Troxler DH et al (1978) J Exp Med 148 , 639 tion is 30°C. Does not have mastadenovirus group antigen: not neutralized Friend spleen focus-forming virus (SFFV) by aviadenovirus antiserum. Does not A defective isolate of Friend murine cause tumors on injection into neph- leukemia virus which encodes a trun- rogenic ridge of Rana pipiens tadpole. cated envelope protein that can bind to Agglutinates rat erythrocytes at 37°C. and activate the erythropoietin recep- Clark HF et al (1973) Virology 51 , 392 tor (EpoR) in the absence of its natural ligand. This results in inactivation of frog herpesvirus 4 Synonym for ranid erythroid signal transduction pathways herpesvirus 2. and proliferation and differentiation of erythroid precursor cells in the absence Frog virus 3 (FV-3) The type species of of erythropoietin resulting in leukemia. the Ranavirus genus, family Iridoviridae . Ruscetti SK (1999) Int J Biochem Cell Biol 31 , A member of the group of icosahe- 1089 dral cytoplasmic deoxyviruses which infect amphibia. Enveloped virion has Frijoles virus (FRIV) A species in the a diameter of 165–200 nm. The icosahe- genus Phlebovirus ; with Joa virus forms dral particle is 130–145 nm in diameter. the Frijoles complex. Isolated from Replicates in both the nucleus and the phlebotomine insects of Lutzomyia sp in cytoplasm. At optimal temperature the Canal Zone, Panama. Not reported of 23–26°C, multiplies in cells of all to cause disease in humans. vertebrate classes in vitro . Envelope is acquired on budding through plasma Frijoles virus VP-161A (FRIV) An isolate membrane. Genome consists of one of Frijoles virus . molecule of double-stranded DNA (105,903 bp). G C ratio 55%. Buoyant Fringilla coelebs papillomavirus The type density (CsCl): 1.287 g/ml. Lethal to species of the genus Etapapillomavirus . tadpoles of several Rana and Bufo sp. 175 fusion of cells

In several toad species the virus pro- Fukuoka virus (FUKAV) An unassigned duces hemorrhagic necrosis of kidneys species in the family Rhabdoviridae and stomach. Replicates in adult newts belonging to the Kern Canyon sero- but not in frogs. In infected Xenopus group. Isolated in cultures of HmLu-1 spp, adaptive cellular immunity has cells derived from baby hamster lung been described. On infection of cell cul- and from blood samples of four sentinel tures, cellular nucleic acid synthesis is calves having a fever and leukopenia. inhibited and viruses, such as vaccinia Noda M et al (1992) Vet Microbiol 32 , 267 virus, SV40 and several RNA viruses, that were in the process of replication fulminant hepatitis About 0.2% of patients in the cell when superinfection by FV-3 with hepatitis B develop symptoms of occurred have their replication halted, acute hepatic failure within a few days while FV-3 continues to replicate. In and go into hepatic coma. These cases mice FV-3 does not replicate but causes of fulminant hepatitis are often fatal, acute degenerative hepatitis owing to and frequently involve coinfection with this blocking of RNA synthesis. hepatitis delta virus. Survivors recover Murti K et al (1985) Adv Virus Res 30 , 1 completely and develop immunity to Robert J et al (2005) Virology 332 , 667 hepatitis B virus. Tan WG et al (2004) Virology 323 , 70 Fuzeon See enfurvirtide . frog virus 4 Synonym for ranid fusin A human membrane protein herpesvirus 2. involved in the penetration of the HIV- receptor complex into the cell. FS (Family Study) virus A virus isolated Feng Y et al (1996) Science 272 , 872 from an outbreak of gastroenteritis during the course of the Family Study fusion of cells The formation of multinu- in Cleveland, USA. Could be passed in cleate giant cells known as polykaryo- humans, and infection gave immunity cytes or syncytia. Can be caused by to infection by Marcy virus. The strain a variety of agents including some has since been lost and can no longer viruses, notably Paramyxoviridae . There be identified. are two types of virus-induced fusion. Hodges RG et al (1956) Am J Hyg 64 , 349 (1) Fusion from without . Not dependent on virus replication or on the synthesis of new proteins. Occurs not more F3T See trifluorothymidine . than 1–3 h after exposure to high multi- plicities of most of the large enveloped FT-cells (CCL 41) A heteroploid cell line RNA viruses or certain DNA viruses derived from cells obtained by primary such as human herpesviruses 1 and cultivation of normal tongue from an 2 and vaccinia virus, even when they adult female bullfrog, Rana catesbeiana . have been inactivated by UV light or β -propiolactone. May also be caused by FUdR See fluorodeoxyuridine . viral hemolysin, since treatment which will destroy this enzyme activity with- Fujinami sarcoma virus (FuSV) A species out affecting viral infectivity will also in the genus Alpharetrovirus . Obtained prevent the cell-fusing action. (2) Fusion from a transplantable fowl tumor in from within. Begins several hours after 1914. A defective virus, with deletions infection and depends on synthesis of in all three virion genes (env , gag , and viral proteins, especially the F protein pol). Carries the oncogene fps , homolo- of paramyxoviruses. Production of new gous to the mammalian feline sarcoma infectious virus is not necessary. Often virus oncogene, fes . Both encode non- most marked after infection at low receptor tyrosine kinases. The tumor virus multiplicity. The mechanism may induced in chickens can be trans- be the same as fusion from without. planted without difficulty into ducks. Fusion from within can be prevented late in infection by antiviral antibody. Fujinami A and Inamoto K (1914) Z Krebsforsch 14 , 94 Falconer MM et al (1995) J Virol 69 , 5582 fusion protein 176 fusion protein See F protein . into the cell. N-tropic viruses infect NIH 3T3 cells, B-tropic viruses infect fuzzy sets A term used to indicate the BALB/c cells, and NB-tropic viruses difficulty of defining species of viruses. infect both types of cell. The Fv1 gene Biological species have generally been is derived from an endogenous retro- defined as groups of individuals capa- virus unrelated to murine leukemia ble of interbreeding, but this cannot be virus. The viral determinants for N- applied to viruses. So the definition of and B-tropism are present in the cap- a virus species adopted by the ICTV in sid protein at amino acid position 110, 1991 which allows some flexibility is: where an arginine specifies N-tropism ‘A virus species is a polythetic class of and a glutamate B-tropism. viruses that constitute a replicating lineage and occupy a particular ecological FV3 virus See Frog virus 3 . niche. ’ van Regenmortel MHV (2005) In Topley & fyn An oncogene, related to src and Wilson ’ s Microbiology and Microbial Infections , belonging to the non-receptor tyrosine vol. 1, Tenth edition, edited by BWJ Mahy and kinase family which is expressed in V ter Meulen. London: Hodder Arnold, p. 24 brain cells. It was originally cloned from an SV40-transformed human fibroblast Fv1 gene A murine gene that restricts library, and has not been found in any infection by N-tropic or B-tropic murine naturally occurring retrovirus. leukemia viruses at a stage after entry G

G protein Name used for the major glyc- virus, particularly for entry into brain oprotein of a number of viruses, espe- cells. cially negative-strand RNA viruses. Harouse JM et al (1991) Science 253 , 320

GA391 virus A strain of Lassa virus in Galibi virus An unclassified virus iso- the genus Arenavirus , isolated from lated from culicine mosquitoes in the Mastomys spp rodents in West Africa. Amazon region of Brazil. Not known to cause disease in humans. Gabek Forest virus (GFV) A tentative species in the genus Phlebovirus , Gallid herpesvirus 1 (GaHV-1) The type belonging to the sandfly fever virus species of the genus Iltovirus. A wide- group. Isolated from the spiny mouse, spread natural infection of fowls and Acomys cahirinus ( albigena). The vector pheasants, causing hemorrhagic trachei- species has not yet been determined. tis with gasping and coughing. Mortality up to 70%. A less virulent strain in Gadget’s Gully virus (GGYV) A tick- Australia and USA causes only coughing borne species in the genus Flavivirus , and sneezing. Ducks, pigeons, and tur- belonging to the mammalian tick-borne keys are occasionally infected. No infec- virus group. Isolated from ticks, Ixodes tions of mammals reported. Transmission (Ceratixodes ) uriae on Macquarie Island, via the respiratory route; some recovered Australia. Not known to cause disease birds may excrete virus for long periods. in humans. An egg- or cell-culture passage strain can be used as a live vaccine, administered gag gene ( group-specific antigen gene) orally in the drinking water. One of the genes in the genome of retro- Synonyms: phasianid herpesvirus 1; viruses. Codes for the precursor of avian infectious laryngotracheitis virus; all the major internal antigens of the infectious laryngotracheitis virus. virus. Gallid herpesvirus 2 (GaHV-2) The type gag–pol gene The adjacent gag and pol species of the genus Mardivirus . A genes of retroviruses are expressed natural infection of fowls, pheasants, from the full-length unspliced mRNA turkeys, quails, ducks, swans, geese, molecules, but only the gag gene is pigeons, and budgerigars, causing pro- expressed unless a frameshift occurs gressive paralysis, usually in birds aged into the pol open reading frame. This 2–8 months. The neuropathogenic form happens at a frequency of about 1 per of the disease was first described by 100 translation events, so the amount Jozsef Marek in 1907, hence the name of pol precursor represents only 1% of Marek’s disease virus, now reserved as the gag precursor that is synthesized. the basis for the genus name. Causes a neoplastic lymphoproliferative disease gag protein The group-specific antigen of of domestic chickens characterized by retroviruses. mononuclear cell infiltration and the development of lymphomas, princi- gal virus Synonym for gallus adeno-like pally in the peripheral nerves and vis- virus, an Aviadenovirus . ceral organs. It has been recognized as a major cause of economic loss to the galactosyl ceramide A cell surface mole- poultry industry. It is the first common cule that has been proposed as an alter- naturally occurring neoplastic dis- native to CD4 as a receptor for HIV-1 ease in any species to be controlled by Gallid herpesvirus 2 (GaHV-2) 178

vaccination, and is the first example of horizontal spread does not occur. The an oncogenic herpesvirus. There are FC-126 strain is commonly used as obvious similarities between MDV, the the vaccine for chickens, and heterolo- Epstein–Barr herpesvirus of humans gous antigens have been introduced and oncogenic herpesviruses of various to extend the usefulness of the vaccine animal species. A member of the sub- (e.g. NDV, IBDV). family Alphaherpesvirinae of the fam- Synonym: Marek’s disease herpesvirus 2. ily Herpesviridae. The genome DNA is Alfonso CL et al (2001) J Virol 75 , 971 178 kb in length encoding 103 proteins. Meleagrid herpesvirus 1 is an antigeni- GALV See Gibbon ape leukemia virus . cally related but non-pathogenic herpesvirus of turkeys (HVT) belonging Gamboa virus (GAMV) A species in the to the same genus, which can trans- genus Orthobunyavirus. With Maru activate latent Marek’s disease virus 10962, Pueblo Viejo and 75V-2621 genes in quail fibroblast cells. Marek’s viruses forms the Gamboa serogroup. disease virus is cell-associated, but free Isolated from the mosquito, Aedeomyia virus is released from feather follicles squamipennis, in Panama. A similar and infection is spread by dust and virus has been isolated in Surinam oral secretions. It can be cultivated in from mosquitoes of Aedes sp. Not chicken kidney cell cultures with pro- reported to cause disease in humans. duction of plaques, but the virus remains cell-associated. Control is by vaccination gammaglobulin A fraction of serum proof chicks at hatching with live herpes- teins with antibody activity. virus of turkeys (gallid herpesvirus 3) which establishes a permanent infec- Gammaherpesvirinae A subfamily of the tion that prevents lymphoma for- family Herpesviridae . The viruses repli- mation. Since the introduction and cate in lymphoblastoid cells, and some widespread use of live vaccines, an species will also replicate in epithelioid acute form of the disease has appeared and fibroblastic cells, causing cell lysis. in Europe and America, characterized They are specific for either B or T lym- by enlargement and lymphocytic infil- phocytes. In lymphocytes replication is tration of the liver and other viscera, often incomplete with persistence of the rather than the nervous system. viral genome but with minimal expres- Synonyms: fowl paralysis virus; Marek’s sion. Even when replication has caused disease herpesvirus 1; neurolymphom- cell death, little or no complete virus atosis of fowls virus; phasianid herpes- may be produced. Latent virus is fre- virus 2. quently demonstrable in lymphoid tis- Baigent SJ et al (2006) Vet Immunol Immuno- sues. Several species are associated with pathol 112 , 78 tumor formation. Host range narrow, Calnek BW et al (1998) Avian Dis 42 , 124 usually limited to species in the same Lee L et al (2000) Proc Natl Acad Sci 97 , 6091 order as the natural host. There are two Payne LN (1999) In Encyclopedia of Virology , genera: Lymphocryptovirus (type species Second edition, edited by A Granoff and Human herpesvirus 4) and Rhadinovirus RG Webster. London: Academic Press, p. 945 (type species Saimiriine herpesvirus 2). Yamaguchi T et al (2000) J Virol 74 , 10176 Synonym : lymphoproliferative herpesvirus group. Gallid herpesvirus 3 (GaHV-3) A species in the genus Mardivirus . A cell-associated gamma interferon A class of interferon virus antigenically related to gallid (known as ‘ immune interferon ’ ) that herpesvirus 2, and isolated from tur- is produced by lymphoid cells in keys. Non-pathogenic in chickens. Does response to mitogens and by sensitized not cause neoplastic transformation. T lymphocytes in response to specific Replicates in avian cell cultures but, antigen. It is a lymphokine that plays a with the exception of hamsters, not in protective role in response to infection mammalian tissues. It protects fowls by upregulating cytokines such as inter- against Marek’s disease. The virus is not leukin 2 in T cells and tumor necrosis released from the feather follicles so factor (TNF) in monocytes. There is 179 Ganjam virus

evidence that it plays an important role species isolated from mammals, reptiles, in control of pathogenesis in response and birds: mammalian virus group, rep- to HIV-1 infection in chimpanzees. tilian virus group, and avian (reticulo- Guidotti LG and Chisari FV (2000) Virology endotheliosis) virus group, respectively. 273 , 221 There are type-specific or subgroup- Rodriguez AR et al (2007) J Gen Virol 88 , 641 specific glycoprotein antigens associated with the viral envelope and group- Gammapapillomavirus A genus in the specific (gs) polypeptide antigens associ- family Papillomaviridae , containing ated with the virion core: gs-1 antigen is viruses causing papillomas which are shared by viruses from one species and histologically distinguishable by intra- is species-specific, but does not cross- cytoplasmic, species specific, inclusion react with type B viruses; and gs-3 antibodies. The genome lacks the E5 open gen is shared by all mammalian type reading frame. Type species is Human C oncoviruses. Members of the genera papillomavirus 4 . Alpharetrovirus and Betaretrovirus do not have such interspecies gs-3 antigens. Gammaretrovirus A genus in the family Coffin JM (1992) In The Retroviridae, vol. 1, Retroviridae. The type species is Murine edited by JA Levy. New York: Plenum Press, leukemia virus (MLV). Virions have a C- p. 19 type morphology and a centrally located condensed core. Capsid assembly occurs γδ T cells A class of T cells found at the at the inner surface of the membrane sites of virus infection, but their exact coincident with the budding process. role is unclear. The diploid genome is about 8.3 kb in Born WK et al (1991) Curr Opin Immunol size (one monomer) with an LTR about 3 , 455 600 bases long. There are four genes in the order gag-pro-pol-env . Both exogenous ganciclovir (GCV) 9-(1,3-dihydroxy- and endogenous viruses are found 2-propoxy) methylguanine A derivative in many mammalian species, and rep- of acycloguanosine that is phosphor- tilian and avian (reticuloendotheliosis) ylated by an enzyme (product of the types are also known. There are also CMV UL97 gene) encoded by human many replication-defective viruses that cytomegalovirus (HHV-5) and is a potent have acquired distinct cell-derived onco- inhibitor of cytomegalovirus replication. genes belonging to the genus. The genus Used for treatment of retinitis and other embraces the leukemia- and sarcoma- complications of cytomegalovirus dis- producing viruses and a number of ease in immunosuppressed patients. related viruses which are probably non- Boucher A et al (2006) Transplant Proc 38 , 3506 oncogenic. Viruses spontaneously shed Crumpacker CS (1996) N Engl J Med 335 , 721 by infected cells are called ‘ exogenous viruses’ but in addition there are ‘ endo- Gan Gan virus (GGV) An unassigned genous viruses ’ whose genetic material virus in the family Bunyaviridae , sero- is integrated into the cell genome and logically a member of the group 3 is vertically transmitted. The nucleic virus group. Isolated from mosquitoes, acid sequences of endogenous viruses Aedes vigilax, in Nelson Bay, Australia. hybridize with DNA from normal tissue Associated with an acute epidemic poly- cells. The degree of hybridization with arthritic-like illness in humans. different species of host can indicate the evolutionary origins of the virus. Ganjam group viruses A term no longer These endogenous viruses may become in use. The two viruses that it contained: activated in vivo or in vitro , spontane- Ganjam and Dugbe, are now placed in ously or by various chemical or physi- the Nairobi sheep disease serogroup cal agents, and produce virus particles of the Nairovirus genus, and Ganjam is which may be ecotropic or xenotropic now regarded as a strain of Nairobi sheep or amphotropic. The role of endogenous disease virus . viruses in the production of spontaneous tumors is not yet clear. The genus Ganjam virus A strain of Nairobi sheep is divided into three groups containing disease virus in the genus Nairovirus Ganjam virus 180

isolated from ticks of Haemaphysalis sp Gammaretrovirus, Transforms mouse in India. cells in vitro and rapidly induces sarcomas in mice, rats and hamsters. See Garba virus (GARV) An unassigned murine sarcoma viruses . vertebrate rhabdovirus. Isolated from Gazdar AF et al (1971) Nat New Biol 234 , 69 birds in Central African Republic. Not reported to cause disease in humans. gazelle herpesvirus Synonym for Equid herpesvirus 9. Gardner–Arnstein feline sarcoma virus (GAFeSV) A species in the genus GB agents See GB viruses . Gammaretrovirus . See Feline leukemia virus . GB viruses Unassigned viruses in the family Flaviviridae , isolated from a Garissa virus (GAV) A strain of Ngari surgeon (George Barker) with acute virus in the genus Orthobunyavirus , iso- non A-non B hepatitis, by injection of lated from some human cases of hem- his serum into marmosets (tamarins) orrhagic fever in Kenya and Somalia of Saguinus sp which then developed during an outbreak of Rift Valley fever hepatitis. Could be serially passed in 1998. in marmosets. Immunologically and Flick R and Bouloy M (2005) Curr Mol Med structurally distinct from the hepatitis 5 , 827 A, B, C, and E viruses. Two GB viruses (GBV-A and GBV-B) were isolated gasping disease virus Synonym for avian recently by representational difference infectious bronchitis virus. Not to be analysis and molecular cloning from confused with infectious laryngotra- the serum of an infected marmoset, and cheitis virus (gallid herpesvirus 1). have been shown to be new members of the Flaviviridae associated with GB gastroenteritis of dogs virus Synonym agent hepatitis. A third virus, GB virus for Canine coronavirus . C, was identified in human sera by reverse-transcription and polymerase gastroenteritis viruses of humans Many chain reaction using consensus primers viruses, such as species of Enterovirus , based on the GBV-A, GBV-B, and hepa- can be isolated from human feces but titis C helicase gene, and appears to be they are rarely the cause of gastroenteri- identical to hepatitis G virus-1. For this tis. Many other viruses can be seen in reason the virus is usually called GB feces with the electron microscope (small virus C/Hepatitis G virus. Although round-structured virus – SRSV – parti- GBV-C/HGV has not been shown to cles) but these are difficult to cultivate cause hepatitis it seems to be able to in vitro. The application of PCR technol- infect liver as well as spleen cells with- ogy has enabled sequence characteriza- out causing obvious disease symptoms. tion of many of these viruses that cause It is widely distributed in the human gastroenteritis, including Norwalk virus population worldwide. (acute epidemic gastroenteritis virus of Birkenmeyer LG et al (1998) J Med Virol humans); groups A, B, and C human 56 , 44 rotaviruses; human adenovirus types 40 Deinhardt F et al (1967) J Exp Med 125 , 673 and 41; human astrovirus; human corona- Mushahwar IK (2000) J Med Virol 62 , 399 virus; human caliciviruses; picobirnavi- Souza IE et al (2006) J Clin Microbiol 44 , 3105 ruses; sapoviruses; and toroviruses. Dingle KE et al (1995) J Gen Virol 76 , 2349 GB virus A An unassigned species in the Glass RI (1995) In Infections of the Gastro- family Flaviviridae. One of two flavi- intestinal Tract, edited by MJ Blaser et al . viruses isolated from the serum of mar- New York: Raven Press, p. 1055 mosets infected with GB virus. It has Lin BL et al (1995) Arch Virol 140 , 1345 been identified in at least six species of New World monkeys since its discov- Gazdar mouse sarcoma virus A strain ery. The virus is transmissible via blood, of mouse sarcoma virus, in the genus but the natural mode of transmission is 181 Gecko adenovirus

not known. Does not seem to cause hep- longer survival times in the face of HIV atitis in the host species or other sus- infection. The molecular basis for this ceptible species. The positive-stranded phenomenon is not yet clear. RNA genome is related to the hepaci- Fogeda M et al (2000) J Virol 74 , 7936 virus genome but lacks a complete capsid Hadlock KG and Foung SKH (1997) Trans protein gene, and differs in the organi- Med Rev 12 , 94 zation of the 3 -non-coding region. Yirrell DL et al (2007) Int J STD AIDS 18 , 244 Charrel RN et al (1999) J Gen Virol 80 , 2329 Erker JC et al (1998) J Gen Virol 79 , 41 GB virus C troglodytes (GBV-Ctro) A virus genome amplified from the GB virus B (GBV-B) A tentative species serum of a chimpanzee (Pan troglo- in the genus Hepacivirus . One of two dytes ). Probably a chimpanzee virus flaviviruses isolated from the serum of related to the GB viruses, with the marmosets infected with GB virus. It is highest homology to GBV-C/HGV. The transmissible via blood and causes hep- significance of this virus in nature has atitis in several species of New World yet to be determined. monkeys. It has not been detected in Birkenmeyer LG et al (1998) J Med Virol humans. Closely resembles hepatitis 56 , 44 C virus by sequence homology and organization. GBB virus An isolate of Seoul virus in the genus Hantavirus obtained from labora- GB virus C (GBV-C) An unassigned spe- tory rats in England. cies in the family Flaviviridae . A positive single-stranded heterogeneous RNA GB virus A-like agents (GBV-A-like- virus (genome 9.4 kb in length) of human agents) Unassigned species in the fam- and chimpanzee origin. GBV-C was dis- ily Flaviviridae, found in New World covered by reverse transcription and monkeys. Their significance in natural PCR of human plasma using consensus host populations is not yet known. See primers based on the genomes of GBV-A, GB virus A . GBV-B, and hepatitis C viruses. GBV-C is (most closely related to the GBV-A group G ؉ C content The total guanine (G of viruses. Soon after it was discovered, and cytosine (C) content of a double- hepatitis G virus-1 was discovered inde- stranded DNA is usually expressed as a pendently by cloning and expression percentage of the total content of bases. of RNA amplified from human serum, The triple hydrogen bond between G but the genome sequences of the two and C is more stable than the double isolates were virtually identical, so they hydrogen bond between adenine (A) frequently go under the clumsy name and thymine (T). Thus the G C con- of GB virus C/hepatitis G virus. The tent affects the physical properties of viruses do not appear to cause hepatitis, the DNA: both the melting point and even though they have been shown to the density of the molecule are propor- infect the liver, and are widely distrib- tional to the G C content. Most mam- uted geographically, but much needs malian cell DNAs have a G C content to be learned about their natural his- of about 44%; some viral DNAs are tory and biology in humans. They can similar but some are as high as 70%. be transmitted experimentally to chimpanzees, without causing hepatitis. The GD VII virus A strain of mouse poliovirus. viruses appear to be genetically heterogeneous, and distinct variants have been reported to differ in tropism for different Gecko adenovirus A tentative species organs. Some strains appear to be lym- in the genus Atadenovirus . Detected photropic, not hepatotropic, viruses. by PCR amplification in the leopard A number of recent studies in patients gecko, fat-tail gecko, Tokay gecko, and co-infected with GBV-C and HIV have the bearded dragon. No evidence of suggested that GBV-C infection is asso- pathogenesis in the host species. ciated with an improved prognosis and Wellehan JF et al (2004) J Virol 78 , 13366 gecko lung-1 (GL1) cells 182 gecko lung-1 (GL1) cells (CCL 111) A cell genetic basis. In order to conduct gene line derived from the lung of an adult therapy it is necessary to know the wild male Tokay gecko, Gekko gekko; sus- type (normal) sequence of the gene, ceptible to a number of mammalian and a means of delivering it to the cell viruses at 36°C. so that it may be expressed. Viruses are the vector of choice for this delivery gel electrophoresis Electrophoresis of as they may enter specific human cells macromolecules in a matrix of poly- and express their genes in a regulated acrylamide, agarose or similar gel. manner for an extended period of time. Although a number of clinical trials gel filtration A type of column chroma- have presented unforeseen problems, tography which separates molecules there remains considerable optimism on the basis of size. The higher molecu- that virus vectors will prove able to cure lar weight molecules pass through the some of these debilitating diseases. column first, the smaller molecules entering pores in the gel making up Linden RM and Berns KI (2005) In Topley & Wilson’s Microbiology and Microbial Infections , the column and thus being retarded. vol. 2, Tenth edition, edited by BWJ Mahy Usually the gels are made from sugar and V ter Meulen, London: Hodder Arnold, polymers that are flexible (Sephadex) p. 1590 or rigid (Sepharose). The pore sizes are determined by the degree of cross- genetic code The sequence of nucleotides linking of the gel. that specifies the amino acid sequences of proteins and the start and stop sig- GeLu cells (CCL 100) A fibroblast-like nals for protein synthesis. Each of the cell line derived from the normal lung 20 amino acids is specified by one or tissue of a 403-day-old Mongolian ger- more codons, each consisting of three bil, Meriones unguiculatus . nucleotides. The code is degenerate, since most amino acids can be speci- gene A sequence of nucleotides along a fied by more than one codon. The sig- nucleic acid molecule which can deter- nals for initiation of protein synthesis mine the composition of one polypep- (start codons) are AUG (methionine) in tide. A protein of 500 amino acids, each eukaryotes, and either AUG or GUG in one represented by one codon of three bacteria. The major signals for termina- nucleotides in the mRNA, thus requires tion of protein synthesis (stop codons) 1500 nt of genetic information. Many are UAA (ochre), UAG (amber), and eukaryotic and some viral DNA genes UGA (opal). Although the code is ‘ uni- are discontinuous and the sequence of versal,’ a number of deviations are nucleotides is interrupted by introns, known, particularly in mitochondrial which are removed after transcription DNA where UGA, e.g. specifies tryp- by splicing, to form the mature mRNA. tophan rather than acting as a stop gene cloning The isolation and amplifi- codon. It has also been suggested that cation of the nucleic acid sequence of UGA may specify insertion of a 21st a gene. This usually involves insertion amino acid, selenocysteine, during into a suitable plasmid vector that rep- expression of certain virus genomes in licates in bacterial or yeast cells, gener- eukaryotic cells. ating multiple copies of the sequence. genetic marker A mutation in a gene gene enhancers Short cis -acting nucle- which allows its genotypic or pheno- otide sequences that increase transcrip- typic identification. tional activity of genes. genetic reactivation See reactivation . gene expression The multistep process by which the protein product of a gene genetic reassortment The exchange of is synthesized. gene segments between viruses that have a segmented genome (e.g. arena- gene therapy It is now recognized that a viruses, bunyaviruses, influenza viruses, large number of human illnesses have a and reoviruses). See recombinants . 183 genomic masking genetic recombination See recombination . genital herpes simplex virus Commonly human herpesvirus 2 (herpes simplex genetic transmission Involves passage of virus 2), although human herpesvirus a viral genome from one host genera- 1 may also infect the genital tract. tion to the next, either integrated into the cellular DNA as a DNA provirus, genome The genetic information in a or in some other close association with cell or virus. In viruses it may consist the genome of the gamete. Often used of DNA or RNA, but not both. DNA with imprecise meaning. See congeni- viral genomes consist of a single mol- tal infection . ecule which may be single-stranded or double-stranded, circular or linear, containing 1.7 kb (circovirus) to 440 kb genetic variation Alteration in genome (iridovirus). One kilobyte of DNA structure by mutation. Occurs much equals 1.02 1 0 6 pg or 618,000 Da. more frequently with RNA than with The base composition of viral double- DNA, in part because of limited copy- stranded DNA is more variable than ing fidelity of RNA replicases and retro- that of cellular DNA; the G C content transcriptases, as well as the lack of may range between 35% and 74% in dif- RNA proofreading enzymes in the host ferent viruses. RNA viral genomes may cell. Consequently RNA viruses exist be linear or circular, double- or single- as a population of variants known as a stranded, containing 0.25 kb (viroids) to quasispecies. This allows rapid adapta- more than 30 kb (coronaviruses). They tion in response to changes in the host usually consist of a single molecule, but (e.g. presence of neutralizing antibod- may be fragmented into several pieces, ies or an antiviral drug) by selection of as in orthomyxoviruses or reoviruses. genetic variants fitted for replication in The genome structures of the principal the altered environment. virus families which infect vertebrates Drake JW and Holland JJ (1999) Proc Natl are given in Table G1. Acad Sci 96 , 13910 Salzano FM (2000) Proc Natl Acad Sci 97 , genomic masking See phenotypic 5317 mixing .

Table G1. Genome structures of the principal virus families which infect vertebrates

DNA/RNA Virus family Nucleic acid

dsDNA Adenoviridae dsDNA, linear with inverted terminal repeats Hepadnaviridae Partially dsDNA and ssDNA circle Herpesviridae dsDNA, linear with terminal repeats and inverted internal repeats of the termini Iridoviridae dsDNA, linear, circularly permuted and terminally redundant Papillomaviridae dsDNA, closed circular Polyomaviridae dsDNA, closed circular Poxviridae dsDNA, linear with the strands covalently closed at their ends ssDNA Circoviridae ssDNA, circular Parvoviridae ssDNA, linear with repetitions at the ends which facilitate loop formation dsRNA Birnaviridae dsRNA, linear in two segments Reoviridae dsRNA, linear in 10–12 segments ssRNA Arenaviridae ssRNA, linear ambisense strand in 2 segments Astroviridae ssRNA, linear positive strand Bunyaviridae ssRNA linear, negative or ambisense strand in 3 segments genotype 184

Table G1. (continued )

DNA/RNA Virus family Nucleic acid Caliciviridae ssRNA, linear positive strand Coronaviridae ssRNA, linear positive strand Filoviridae ssRNA linear, unsegmented negative strand Flaviviridae ssRNA, linear positive strand Nodavirida ssRNA, linear positive strand Orthomyxoviridae ssRNA, linear negative strand in 7 or 8 segments Paramyxoviridae ssRNA, linear unsegmented negative strand Picornaviridae ssRNA, linear positive strand Retroviridae ssRNA, linear positive strand diploid consisting of 2 identical haploid molecules, each terminally redundant, and linked near their 5 ends Rhabdoviridae ssRNA, linear unsegmented negative strand Togaviridae ssRNA, linear positive strand

ds, Double-stranded; ss, single-stranded. genotype The genetic information con- germ line transmission of viruses See ger- tained in an organism or virus. Many minal cells and genetic transmission . factors determine the phenotype resulting from a particular genotype. Virus Gerstmann–Sträussler–Scheinker syn- genotypes are commonly studied by drome See prion diseases . partial or complete nucleotide sequence analysis and comparison between Getah virus (GETV) A species in the related genotypes is used to infer evo- genus Alphavirus. Isolated in Malaysia, lutionary relationships (phylogenetic Japan, Campuchia, and Australia from analysis). mosquitoes, and from swine in Japan. No known association with disease. geochelone carbonaria herpesvirus Causes viremia in chicks. Antibodies Synonym for chelonid herpesvirus 4. found in pigs, horses, men, and birds. geochelone chilensis herpesvirus Yoshinaka Y et al (1979) Microbiol Immun Synonym for chelonid herpesvirus 4. 23 , 95

German measles virus Synonym for GF cell line (CCL 58) A permanent cell Rubella virus . line established from a marine teleost, the blue-striped grunt, Haemulon sciu- germinal cells Cells that produce hap- rus. Grows in Eagle’s basal medium loid gametes by meiosis, e.g. oocytes with 0.196 M sodium chloride and both or spermatocytes. Some viruses such human and calf serum. Incubation as retroviruses may infect germinal at 20°C. Used to isolate enteric fish cells and so pass vertically from one viruses. generation to the next. See genetic transmission . GG167 Original name for Zanamivir (influenza neuraminidase inhibitor). Germiston virus (GERV) A strain See Zanamivir . of Bunyamwera virus in the genus Orthobunyavirus , belonging to the GH1 cells (CCL 82) A somatotrophin- Bunyamwera serogroup. Isolated from secreting clone of MtT/W5, established culicine mosquitoes, humans, and from a rat pituitary gland tumor of a rodents in South Africa, Zimbabwe, Wistar–Furth rat. Uganda, and Mozambique, where it causes fever in humans. Mosquito- GH3 cells (CCL 82.1) A somatotrophin- borne. Natural hosts are sheep, goats, and prolactin-secreting clone of MtT/ and cattle. W5, established from a pituitary tumor 185 gloves and socks syndrome

of a 7-month-old female Wistar–Furth with GALV. No GALV isolates have rat. Not a direct derivative of the GH1 been made from wild gibbon apes. clone. O’Hara B et al (1990) Cell Growth Differ 1 , 119 Oliveira NM et al (2006) J Virol 80 , 3104 Gianotti-Crosti syndrome A self-limited Reitz MS et al (1979) Virology 93 , 48 childhood skin disease (papular acro- dermatitis) associated with hepatitis gibbon ape lymphosarcoma virus B virus or parvovirus B19 infection Synonym for the complex of simian and usually accompanied by lymph- sarcoma virus and its helper, the sim- adenopathy and anicteric hepatitis. Has ian sarcoma-associated virus. also been reported in association with Epstein–Barr virus infection. Lesions gibbon ape type C oncovirus See Gibbon usually persist for about 3 weeks and do ape leukemia virus . not recur. Prevalent in Italy and Japan. Gill-associated virus The type species of Caputo R et al (1992) J Am Acad Dermatol the genus Okavirus . The natural host is 26 , 207 the black tiger prawn (Penaeus mono- Lowe L et al (1989) J Am Acad Dermatol don ). The geographic range is restricted 20 , 336 to Asia and Australia. Causes high mortality with virus in most tissues giant bacteriophages Aberrant forms of but is particularly concentrated in the certain enterobacteria phages in which ‘ Oka ’ or lymphoid organ. the heads are up to 44 times longer than normal. These forms usually con- gingivostomatitis The primary symptom tain greater than unit length of DNA associated with oral herpes simplex and are infectious. virus infection, usually associated with a sore throat and painful ulcers in the giant cell pneumonia virus Synonym mouth which resolve in a few weeks. for Human herpesvirus 5 (human Meanwhile the virus establishes a life- cytomegalovirus). long latent infection in neurons of the trigemonal ganglia, and periodically Gibbon ape leukemia virus (GALV) A reactivates in the oral cavity with shed- replication competent species in the ding in the saliva. genus Gammaretrovirus . Seen by electron microscopy in a disseminated Girard Point virus An isolate of Hantaan T-lymphoblastic lymphosarcoma in a virus. captive gibbon ape, Hylobates lar . The tumor cells in culture grew as free- Girardi heart cells (CCL 27) A hetero- floating cells, releasing virus parti- ploid cell line established from the cles, which are non-transforming in right atrial appendage of a 41-year-old vitro but in gibbons induce leukemia. human male in 1956. The cells grew as Antigenically related to the simian sar- fibroblasts at first, but after about 18 coma virus. Several strains have been weeks they became epithelial-like. isolated and their origins are indicated by suffixes, thus: H for Hall’s Island glandular fever virus Synonym for near Bermuda; SF for San Francisco; Br Human herpesvirus 4 . for brain extract from gibbons injected with extract of human brain from gliotoxin A common fungal metabolite kuru patients; and SEATO for Seato which inhibits viral RNA-dependent Laboratory, Bangkok. All are closely RNA synthesis without effect on DNA- related by protein serology and nucleic dependent RNA synthesis. Prevents acid homology. The GALVs as a group human poliovirus replication in HeLa are related to murine leukemia viruses, cells. especially the endogenous virus of the Miller PA et al (1968) Science 159 , 431 Asian mouse, Mus caroli . Koala retrovirus, an endogenous retrovirus of gloves and socks syndrome A petechial koalas, shares 78% nucleotide identity or papular–purpuric rash on the gloves and socks syndrome 186

hands and feet of children and adults. goat adenovirus 1 (GAdV-1) A serotype Following infection there is acute onset of ovine adenovirus D, a species in the of fever, exanthem, edema, and ery- genus Atadenovirus . thema of the hands and feet that has most frequently been associated with goat adenovirus 2 (GadV-2) A serotype parvovirus B19 infection. It has also of goat adenovirus, a tentative species been described in relation to cytomega- in the genus Mastadenovirus . lovirus infection. Alfadley A et al (2003) J Am Acad Dermatol goat capripoxvirus See Goatpox virus . 48 , 941 Carrascosa JM et al (1995) Dermatology 191, 269 goat herpesvirus (CpHV-1) Synonym for Halasz CLG et al (1992) J Am Acad Dermatol 27 , 835 Caprine herpesvirus 1 . glugea lyssae Name given to a supposed goat papillomatosis A natural infection protozoan etiological agent of rabies. of some herds, but not common; the The structure is now known as the papillomas may become malignant. ‘ Negri body. ’ The virus has not yet been isolated. Levaditi C et al (1924) C R Acad Sci Paris Moulton JE (1954) North Am Vet 35 , 29 178 , 256 Goatpox virus (GTPV) A species in the glycoproteins Proteins with covalently genus Capripoxvirus. Clinically causes attached sugars, bound either through a disease similar to sheeppox, but the the OH group of serine or threonine disease in goats takes a milder course. (O -glycosylation) or through the amide Causes focal epidermal lesions which NH2 of asparagine (N -glycosylated). proceed through papule, vesicle, and Most virus proteins that bud through pustule stages to scab formation. the surface membrane are glycosylated. Lesions are usually on the udder, teats, Although N-glycosylation is the most scrotum, inside of the thighs and less common, O -glycosylation is found with frequently around the eyes and mouth, some viruses, notably coronaviruses. whereas in contagious ecthyma, the The sugar residues found on virus glyco- lips and mouth are primarily involved. proteins include fucose, galactosamine, Antigenically distinct from contagious galactose, glucosamine, mannose, and ecthyma virus and there is no cross- sialic acid. protection. Replicates on the CAM producing opaque pocks and in cell cultures glycosylation The addition of sugars to of lamb and kid kidney tissue with CPE. proteins. Transmissible to sheep and allegedly to calves and rabbits. Experimental vac- glycyrrhizic acid An antiviral agent cines have been used which protect both derived from the roots of the liquorice sheep and goats against capripoxvirus plant, Glycyrrhiza glabra. The ammo- infection, but a commercial vaccine is nium salt completely inhibited growth not presently available. Occurs in many and CPE of vaccinia, human herpes- parts of the world. virus 1, Newcastle disease, and vesicu- Kitching P et al (1985) J Gen Virol 67 , 139 lar stomatitis viruses when grown in Renshaw HW and Dodd AG (1978) Arch cultures of HEp-2 cells. There was no Virol 56 , 201 effect on poliovirus type 1. In addition to the inhibitory action, there is goatpox virus G20-LKV A strain of irreversible inactivation of human her- Goatpoxvirus . pesvirus 1, although not of the other viruses tested. The mode of action of goatpox virus Pellor A strain of glycyrrhizic acid is not understood, Goatpoxvirus . but it is not thought to be mediated through damage to host cells. golden ide reovirus (GIRV) A tenta- Pompei R et al (1979) Nature 281 , 689 tive species in the genus Aquareovirus . 187 Goose parvovirus (GPV)

Isolated from golden ide, Leuciscus idus Gomoka virus (GOMV) A serotype of Ieri melanotus . virus in the genus Orbivirus . Isolated Attoui H et al (2002) J Gen Virol 83 , 1941 from mosquitoes and birds in Central Neukirch M et al (1999) Dis Aquat Org 35, 159 African Republic. Not reported to cause disease in humans. golden pheasant leukosis virus A strain of subgroup G avian leukosis virus. Goodpasture’s syndrome A syndrome An endogenous virus found in normal with glomerulonephritis associated golden pheasant, Chrysolaphus pictus , with pulmonary hemorrhage and cir- cells giving group G host range specifi- culating antibodies against basement city to virus particles in which it pro- membrane antigens which has been vides the information for the envelope. observed in some cases of influenza A virus infection. Hanafusa T et al (1976) Proc Natl Acad Sci USA 73 , 1333 Wilson CB and Smith RC (1972) Ann Intern Tal J et al (1977) J Virol 21 , 497 Med 76 , 91 golden shiner reovirus (GSV) A Goose adenovirus (GoAdV) A species in strain of Aquareovirus C in the genus the genus Aviadenovirus. Three sero- Aquareovirus, which naturally infects types have been reported. commercially propagated golden shiners (bait minnow), Notemigonus cryso- goose adenoviruses 1 to 3 (GoAdV-1 to leucas. The virus is of low virulence 3) Serotypes of Goose adenovirus in the and disease is seen only in hot weather genus Aviadenovirus . Three serotypes when fish swim near the surface and isolated from liver and gut of young show a hemorrhagic dorsal surface goslings. Unrelated to fowl, turkey or and petechial hemorrhages on the ven- duck adenoviruses. Role in disease tral surface. The virus grows and can uncertain, though adenovirus-like par- be isolated in FHM cells at 25–30°C, ticles have been seen in the nuclei of and identified by serum neutraliza- liver cells in birds with hepatitis. tion tests. Golden shiners can also be Zsak L and Kisary J (1984) Avian Pathol experimentally infected with other 13 , 253 aquareoviruses, such as American oyster reovirus (aquareovirus A) or chum Goose circovirus (GoCV ) A species in the salmon reovirus (aquareovirus F). genus Circovirus . Infections are asso- Brady YJ and Plumb J (1991) J Wildl Dis 27 , ciated with growth retardation and 463 developmental problems in farmed Winton JR et al (1987) J Gen Virol 68 , 353 geese. Does not grow in cell culture. An assay for the virus was developed goldfish herpesvirus Synonym for using a Semliki Forest virus expression cyprinid herpesvirus 2, an unassigned vector to produce GoCV capsid protein virus in the family Herpesviridae . in BHK cells and to detect specific antiviral antibody in goose sera by indirect goldfish virus 1 A virus isolated from immunofluorescence using anti-duck normal healthy goldfish (Carassius fluorescent duck immunoglobulin con- auratus ). Unfortunately the isolate has jugate. The genome DNA is 1821 nt in now been lost. length. Berry ES et al (1983) J Fish Dis 6 , 501 Chen CL et al (2003) Avian Pathol 32 , 165 Scott AN et al (2006) Avian Pathol 35 , 495 Golgi apparatus A cytoplasmic organelle Todd D et al (2001) Virology 286 , 354 composed of membrane-bound vesicles in which glycosylation and pack- goose hepatitis virus See Goose aging of secreted proteins takes place. parvovirus . Part of a complex known as the Golgi– endoplasmic reticulum–lysosome sys- Goose parvovirus (GPV) A species in the tem (GERL) in which the sorting of pro- genus Parvovirus. Causes severe and epi- teins into separate pathways takes place. demic liver disease (Derzsy’s disease) Goose parvovirus (GPV) 188

in goslings and young Muscovy ducks, Gorotire virus A probable species in Cairina moschata. Older birds may the genus Orbivirus. Isolated from show acute or chronic hemorrhagic phlebotomine sand flies in the Amazon disease. Spread of the disease occurs region of Brazil. Not known to cause by horizontal transmission, but verti- disease in humans. cal transmission of the virus has also been demonstrated. Injection of 2- to gosling hepatitis virus See Goose 3-day-old goslings i.m. causes hemor- parvovirus . rhagic liver disease and pericarditis after a latent period of 5–7 days, and Gossas virus (GOSV) An unclassified death in 5–10 days. Widespread in North species in the family Rhabdoviridae . America, Asia, and Europe. Replicates Isolated from a bat of Tadarida sp in on inoculation into allantoic cavity of Senegal. Not reported to cause disease goose or Muscovy duck eggs. In goose in humans. embryo fibroblast cell cultures replicates with CPE. Characterization of the DNA GPC-16 cells (CCL 242) A cell line estab- by sequencing and cross-hybridization lished from a chemically induced suggests a close relationship to human adenocarcinoma of the colon from an dependoviruses (AAV). outbred guinea pig. Synonyms : Derzsy’s disease of geese; goose plague; goose hepatitis; goose G protein Abbreviation for glycoprotein. influenza; infectious myocarditis of goslings virus. Graffi leukemia virus A strain of Murine leukemia virus obtained by injecting Brown KE et al (1995) Virology 210 , 283 extracts of several transplantable mouse Gough RE and Spackman D (1982) Avian Pathol 11 , 503 tumors into newborn Agnes Bluhm Zadori Z et al (1995) Virology 212 , 562 strain mice. Causes a chloroleukemia. The cell type may be granulocytic, lymphocytic, or histiocytic depending on Gordil virus (GORV) A tentative species the mouse strain, age, and hormonal in the genus Phlebovirus not assigned status. to any antigenic complex. Isolated from striped grass mouse and gerbil in Graffi A et al (1955) Wien Klin Wochenschr Central African Republic. Not reported 105 , 61 to cause disease in humans. Grand Arbaud virus (GAV) A serotype or strain of Uukuniemi virus in the gorilla herpesvirus Synonym for Pongine genus Phlebovirus. Isolated from a tick, herpesvirus 3 . Argas reflexus , in southern France. Not reported to cause disease in humans. gorilla immunodeficiency virus During a study of simian immunodeficiency granulocyte-macrophage colony- viruses in Africa, wild gorillas in west- stimulating factor (GM-CSF) A central Africa were found to harbor cytokine which upregulates the repli- HIV-1 group O-like viruses. cation of HIV in primary macrophages van Heuverswyn F et al (2006) Nature 444 , and in monocyte and myeloid cell lines. 164 grass carp reovirus (GCRV) A tentative species in the genus Aquareovirus . gorilla rhadinoherpesvirus 1 (GorRHV1) Causes an acute hemorrhagic disease During a study of chimpanzees and in ‘ fingerling ’ and ‘ yearling ’ grass carp. gorillas from Cameroon and Gabon, Isolated from a moribund grass carp, a herpesvirus genetically similar to Ctenopharyngodon idella , in FHM cells in Kaposi’s sarcoma herpesvirus was which it produces CPE. Optimal tem- detected by PCR amplification of perature for virus replication is 16–23°C. peripheral blood mononuclear cells Infectivity not neutralized by antiserum from a gorilla ( Gorilla gorilla ). to rhabdovirus carpio or swim-blad- Lacoste V et al (2000) Nature 407 , 151 der inflammation virus. Common carp, 189 Grimsey virus (GSYV)

Cyprinus carpio , appear to be resistant Cooper JE et al (1982) Lab Anim 16 , 12 to the virus, but grass carp are easily Raynaud A and Adrian M (1976) C R Hebd infected from the water they are in, and Seances Acad Sci 283 , 845 die in 8–9 days with hemorrhagic inflammation in scale bases and other tissues. green monkey virus Synonym for Marburg virus . grass carp rhabdovirus Synonym for pike fry rhabdovirus. green sea turtle herpesvirus Synonym for chelonid herpesvirus 1. Gray Lodge virus (GLOV) A tentative species in the genus Vesiculovirus . green turtle herpesvirus (GTHV) A Isolated from mosquitoes, Culex tarsa- virus associated with an emerging lis , in California, USA. Not known to neoplastic disease of turtles in Florida cause disease in humans. and Hawaii. Fibropapillomatosis is a debilitating, frequently fatal disease of gray patch disease agent of green sea marine green turtles, Chelonia mydas , turtle Synonym for chelonid herpes- characterized by the presence of epi- virus 1. thelial fibropapillomas and internal fibromas containing herpesvirus-like grease or grease-heel virus A pox- DNA sequences. Although a virus has virus infection of horses. Often consi- not so far been isolated, DNA sequence dered to be the same as horsepox but analysis of fibropapilloma tissue sug- this is not certain. Vesicles appear on gests that there may be at least three flexor surfaces of lower parts of legs, closely related herpesviruses associ- later becoming pustules and crusts. ated with the disease, depending on Recovery usually occurs after 3 weeks. the species and geographical location. In addition to green turtles, loggerhead Greasy grouper nervous necrosis turtles and olive ridley turtles may also virus A tentative species in the genus be infected. When cell-free filtrates of Betanodavirus . cultured cells derived from the fibropapilloma tissue were inoculated into Great Island virus (GIV) A species in the cells derived from healthy green tur- genus Orbivirus. Isolated from ticks, tle lung, tumor-like cell aggregates Ixodes uriae, and sea birds on Great were formed, and electron microscopy Island, Newfoundland, Canada. showed that these contained a small (50 nm diameter) naked virus. The role Great Saltee Island virus (GSIV) A sero- of this virus in disease and its relation, type of Great Island virus in the genus if any, to the herpesvirus sequences, Orbivirus . remain to be determined. Great Saltee virus (GRSV) A serotype of Lu Y et al (2000) Arch Virol 145 , 1885 Hughes virus in the genus Nairovirus . Lu Y et al (2000) J Virol Methods 86 , 25 Quackenbush SL et al (1998) Virology 246, 392 green iguana herpesvirus Synonym for iguanid herpesvirus 1. gray kangaroopox virus (KXV) An unassigned virus in the family Poxviridae . green lizard herpesvirus Synonym for lacertid herpesvirus 1. gray lung disease virus Probably a mycoplasma. See pneumonia virus of rats . green lizard papillomavirus A possible Andrewes CH and Glover RE (1945) Br J Exp virus in the family Papillomaviridae . Pathol 26 , 379 Seen by electron microscopy in skin lesions which were most prevalent in gray patch disease of turtles virus the genital areas in the green lizard, Synonym for chelonid herpesvirus 1. Lacerta viridis . Successful experimental transmission of the disease with virus Grimsey virus (GSYV) A serotype of extracts was not accomplished. Great Island virus in the genus Orbivirus . Gross leukemia virus 190

Gross leukemia virus The first strain of Does not cause disease on injection Murine leukemia virus isolated. Obtained into adult grunts, Haemulon sciurus , or from a spontaneous lymphoma of AKR newborn mice. Ether sensitive. Oval mice. Also known as Gross passage particles, 120–140 nm diameter, with A virus. Causes lymphoid leukemia dense core. on injection into newborn mice, usu- Clem LW et al (1965) Ann NY Acad Sci 126 , 343 ally after a latent period of at least 2–3 months. The virulent passage A virus is transmitted in the milk. grunt fin cells (GF) See GF cell line . Gross L (1951) Proc Soc Exp Biol Med 76 , 27 GSV See golden shiner reovirus . ground squirrel cytomegalovirus Syno- nym for sciurid herpesvirus 1. GU71U344 virus (GU344V) A serotype of Bushbush virus in the genus Ground squirrel hepatitis virus (GSHV) A Orthobunyavirus . Isolated from a senti- species in the genus Orthohepadnavirus nel hamster. Not known to cause dis- isolated from wild Beechey ground ease in humans. squirrels, Spermophilus beecheyi , caught near Palo Alto in the San Francisco GU71U350 (GU350V) A serotype of peninsula, USA. The virus has not Guajará virus , a species in the genus been found elsewhere in California or Orthobunyavirus . the rest of the USA. Will infect woodchucks in laboratory experiments, but Guajará virus (GJAV) A species in the not other rodents or species tested. genus Orthobunyavirus . Isolated from Marion PL et al (1983) Proc Natl Acad Sci 83 , sentinel mice in Amapa and Para, 4543 Brazil. Not reported to cause disease in humans. ground squirrel herpesvirus Synonym for sciurid herpesvirus 2. Guama serogroup viruses A group of serologically related viruses in the group-specific antigen An antigen com- genus Orthobunyavirus. Isolated only in mon to a group of viruses. See type- the Western Hemisphere. specific antigen . Guama virus (GMAV) A species in the grouper sleepy disease iridovirus genus Orthobunyavirus belonging to the (GSDIV) A strain of Infectious spleen Guama serogroup. Isolated from sen- and kidney necrosis virus in the genus tinel monkeys and mice, rodents and Megalocytovirus. Isolated from spleens bats. Mosquito-borne. Found in Brazil, of sick grouper, Plectropomus maculatus , Trinidad, Surinam, French Guiana, and in Singapore. Grows in BF-2 cells. Panama. Can cause a febrile illness with arthralgia in humans. Chew-Lim M et al (1992) J Aquat Anim Health 4 , 222 Mahardika K et al (2004) Dis Aquat Org Guanarito virus (GUAV) A species in the 59 , 1 genus Arenavirus belonging to the South American hemorrhagic fever virus gruid herpesvirus 1 (GrHV-1) An unas- group. The cause of Venezuelan hemor- signed virus in the family Herpesviridae . rhagic fever. Causes a hemorrhagic fever Isolated from a crane in Germany. in humans with a mortality rate from confirmed cases of 36%. The rodent vec- Bürtscher H and Grunberg W (1979) Zentralbl Vet Med B 26 , 561 tor is Zygodontomys brevicauda . Synonym : Venezuelan hemorrhagic grunt fin agent Isolated from a culture fever virus. of GF cell line showing spontane- Tesh RB et al (1993) Am J Trop Med Hyg 49, 227 ous CPE. Replicates in primary grunt Tesh RB et al (1994) Am J Trop Med Hyg 50, 452 fin cells and fish cell lines such as CF, Cajimat MN and Fulhorst CF (2004) Virus but not in KB, HeLa, or chick embryo. Res 102 , 199 191 guinea pig leukemia virus guanidine hydrochloride Selectively Guaroa virus (GROV) A species in inhibits replication of small RNA the genus Orthobunyavirus . Isolated viruses in tissue culture. Antiviral activ- in Colombia, Brazil and Panama. ity is directed against genome-coded Mosquito-borne, probably by Anopheles viral RNA polymerase. Picornaviruses sp. Causes a febrile illness in humans. become completely resistant after only a Antibodies frequently found in humans few passages in the presence of the drug. in the Amazon region. Siegl G and Eggers HJ (1982) J Gen Virol 61 , 111 Guillain–Barré syndrome An acute inflammatory demyelinating poly- guanidinium thiocyanate A powerful radiculoneuropathy. Guillain, Barré, protein denaturant which is used to and Strohl described two cases of paraly- inhibit ribonuclease activity during sis, muscular tenderness, areflexia, and RNA extraction from cells. slight sensory disturbance. Recovery Chomczynski P and Sacchi N (1987) Anal was complete. CSF protein was raised Biochem 162 , 156 but cell count was normal. Landry had described a similar clinical picture in 4-guanidino-2,4-dideoxy-2,3-dehydro- 1856. Cases occur sporadically with an Nacetylneuraminic acid A neuramini- incidence of about 1.6 per 100,000 pop- dase (sialidase) inhibitor which inhibits ulation per year. Cause is probably an the growth in tissue culture of all nine immunological reaction started by an neuraminidase subtypes of avian influ- antigen, which in some cases is a viral enza A viruses. A selective agent for the protein. There was a clear association treatment (and prevention) of influenza of the syndrome with widespread use virus infections. Active against both of inactivated influenza vaccine in the influenza A and influenza B viruses. USA in 1976–1977. Delivered by inhalation for human use. A related drug which can be delivered Hughes RAC (1978) Br J Hosp Med 20 , 688 Schonberger LB et al (1979) Am J Epidemiol orally is oseltamivir. 110 , 105 Synonyms : 4-guanidino-Neu5Ac2en; GG167; zanamivir; Relenza, guinea pig adenovirus (GPAdV) A tenta- Gubareva LV et al (2000) Lancet 355 , 827 tive species in the genus Mastadenovirus . Hayden FG et al (2000) N Engl J Med 343 , Not isolated, but sequence amplifica- 1282 tion of the hexon gene indicates that it is von Itzstein M et al (1993) Nature 363 , 418 not closely related to any other known adenovirus. guanyloribonuclease See ribonuclease T1 . guinea pig adenovirus 1 (GPAdV-1) A guanylyltransferase An enzyme which strain of guinea pig adenovirus. adds guanosine 5 monophosphate from guanosine 5 triphosphate to the guinea pig cytomegalovirus Synonym 5 terminus of nascent RNA chains. for caviid herpesvirus 2. Present in virions of Reoviridae , and involved in formation of the 5 termi- guinea pig endogenous virus Synonym nal cap structure on mRNA. for Guinea pig type C oncovirus .

Guaratuba virus (GTBV) A serotype guinea pig herpesvirus 1 Synonym for of Bertioga virus in the genus Orthobun- caviid herpesvirus 1. yavirus. Isolated from sentinel mice and hamsters, birds and mosquitoes in São guinea pig herpesvirus 3 Synonym for Paulo, Brazil. Not reported to cause caviid herpesvirus 3. disease in humans. guinea pig leukemia virus The L2C Guarnieri bodies Intracytoplasmic aci- transplantable leukemia is a B-cell dophilic inclusion bodies found in cells leukemia transmissible by cell suspen- infected with certain poxviruses, vari- sion, which arose spontaneously in ola, vaccinia, and cowpox. 1954 in an old female strain 2 guinea guinea pig leukemia virus 192

pig. The leukemic cells are always guineafowl transmissible enteritis virus female, even when passed in a male (GTEV) An unassigned virus in the guinea pig. Intracellular and extracel- family Picornaviridae . lular virus-like particles are seen in the leukemic tissue. The particles have dif- Gull circovirus (GuCV) A tentative spe- ferent morphologies and their relation- cies in the genus Circovirus. Using the ship to each other, and to guinea pig polymerase chain reaction, circovirus- type C oncovirus which can be induced specific DNA was isolated from bursa by 5-bromodeoxyuridine treatment, of Fabricius tissues from a herring gull is not clear. Caviid herpesvirus 2 has (Larus argentatus ). been isolated from the leukemic cells Todd D et al (2007) Avian Path 36 , 75 by cocultivation with mink lung cells. Rhim JS and Green I (1977) Proc Fed Am Soc Gumbo Limbo virus (GLV) A strain of Exp Biol 36 , 2247 Marituba virus in the genus Orthobunya- virus belonging to the group C virus guinea pig poxvirus Possibly a species in group. Isolated from mosquitoes in the family Poxviridae. Electron micro- Florida, USA. Not known to cause dis- scopy revealed poxvirus-like particles ease in humans. and inclusion bodies in cell cultures prepared from the thigh tissues of Gumboro disease virus Synonym for approximately 8-month-old guinea Infectious bursal disease virus . Named pigs. The animals from which the tis- after the locality, Gumboro, Delaware, sue was taken were suffering from a where the first outbreaks were spontaneous fibrovascular prolifera- observed in the USA in 1962. tion in the muscles, which had brought about an increase in the normal volume Guppy virus 6 (GV6) A strain of Santee- of the thigh by as much as six-fold. Cooper ranavirus in the genus Ranavirus . Hampton EG et al (1968) J Gen Virol 2 , 205 Gurupi virus (GURV) A serotype guinea pig retrovirus Synonym for of Changuinola virus in the genus Guinea pig type C oncovirus . Orbivirus . Isolated from sand flies, Lutzomyia sp in Para, Brazil. Not guinea pig salivary gland virus Synonym known to cause disease in humans. for caviid herpesvirus 2. Gweru virus A strain of Palyam virus Guinea pig type C oncovirus (GPCOV) A in the genus Orbivirus . Isolated from species in the genus Gammaretrovirus . an aborted cattle fetus in Gweru. An endogenous virus, the formation Zimbabwe. of which can be induced in cell cul- Whistler T and Swanepoel R (1988) J Gen tures from any strain of guinea pig by Virol 69 , 2221 treatment with 5-bromodeoxyuridine. Maximal virus release occurs 2–4 days Gyrovirus A genus in the family after treatment and falls off rapidly, Circoviridae , the type and only spe- being negligible again by 7 days. No cies of which is Chicken anemia cell line has been found to support con- virus. Distinguished from the genus tinued virus production. Although this Circovirus on the basis of its nega- endogenous virus is easily induced, no tive sense genome organization. The exogenous virus has been obtained. virion diameter is 19–26 nm, larger Synonyms: guinea pig endogenous than members of the circovirus genus. virus; guinea pig retrovirus. The genome is a closed circular single- Davis AR and Nayak DP (1977) J Virol stranded DNA about 2300 nt long. 23 , 263 H

H-1 virus (H-1PV) A species in the genus H32580 virus (H32580V) An isolate of Parvovirus. Belongs serologically to Tacaiuma virus , a species in the genus rodent parvovirus group 2. Isolated Orthobunyavirus. Isolated in Cotia from a transplantable human tumor, County, São Paulo State, Brazil from Hep-1, hence the designation. Very sim- the blood and spinal fluid of a patient ilar in biological properties to Kilham with low grade fever and strabismus. rat virus but serologically distinct. Agglutinates preferentially guinea pig, HAART H ighly A ctive A nti r etroviral hamster, human, and rat erythrocytes, Therapy. A strategy to use several in that order. Antibodies not often antiretroviral drugs in combination to present in human serum but virus can suppress HIV replication. Drugs which replicate on injection into humans. The are protease inhibitors (e.g. saquinavir, genome DNA is 5176 bases in length, ritonavir, indinavir, or nelfinavir) are with palindromic sequences at the 5 given two or three times daily together and 3 ends. The strain deposited in the with reverse transcriptase inhibitors American Type Culture Collection was (e.g. ddC, delavirdine, didanosine, isolated from the embryo of a pregnant lamivudine, nevirapine, or zidovu- woman with metastatic carcinoma of dine). New drugs under development the breast. The natural host is probably targeted at virus-cell binding, virus the rat. entry or DNA integration promise to Rhode SL and Paradiso PR (1983) J Virol 45 , enhance opportunities for HAART. 173 de Clercq E (2000) Rev Med Virol 10 , 255 Griffiths PD (1999) Rev Med Virol 9 , 1 H1–H51 The original serotype designa- Telenti A and Rizzardi GP (2000) Rev Med tions of the mammalian adenoviruses Virol 10 , 385 isolated from humans, Homo sapiens . Now grouped into six species, Human HADEN virus Sigla from h em ad- sorbing adenovirus A–F . en teric virus of calves. Synonym for Bovine parvovirus .

H1N1 The first designate influenza A Haematopoietic necrosis herpesvirus of virus subtype which was replaced in goldfish Synonym for Cyprinid herpes- 1957 by the Asian influenza (H2N2), virus 2 . but reappeared in 1977 and now cir- culates each winter in human popu- hairy cell leukemia A disorder character- lations. It is a component of trivalent ized by pancytopenia, splenomegaly, influenza vaccines. and abnormal mononuclear cells, with cytoplasmic projections (hairy cells), H-2 virus Does not exist. infiltrating the bone marrow, spleen, and peripheral blood. An immortalized H-3 virus (H-3PV) A strain of Kilham rat T-cell line (MoT) was established in 1976 virus in the genus Parvovirus . A rodent from the spleen of a hairy cell leuke- parvovirus of serological group 1. mia patient, and from this line human Isolated from a human tumor Hep- T lymphotropic virus 2 (HTLV-2) was 3, which had been transplanted for isolated. There is no good evidence that several years in conditioned rats. HTLV-2 is involved in the etiology of Serologically different from H-1 virus. hairy cell leukemia, however. Agglutinates guinea pig and rat erythrocytes equally. HaK cells (CCL 15) A heteroploid cell Synonym : OLV virus. line derived from the kidneys of two HaK cells 194

normal young adult Syrian or golden in Canada in 1957. The disease is most hamsters, Mesocricetus auratus . common in summer and autumn, and children aged 1–5 years are usu- Halibut nervous necrosis virus A tenta- ally affected. Incubation period is 3–6 tive species in the genus Betanodavirus . days. The patient may feel unwell for a day before red papules and vesicles Hallé measles virus A strain of Measles appear in the mouth and ulcerate. The virus isolated in 1971 from a patient skin lesion is a maculopapular rash with subacute sclerosing panencepha- progressing to vesicles on the feet and litis (SSPE). hands, extending to other parts of the body. At least four types of coxsackie Hamming distance The number of A virus can cause the disease: A16 positions at which two nucleotide most often, but in some cases A5, A9, sequences of defined length differ. and A10; and two types of coxsackie B virus: 2 and 5. Enterovirus 71 is also a hamster enteritis A common disease of major cause of the disease, especially laboratory hamsters with high mor- cases complicated by CNS involve- tality. Characterized by diarrhea, ment. The disease occurs worldwide, dehydration, and weight loss. Cause but most frequently in Asia. In 1997 doubtful, but both viruses and bacteria there was a severe outbreak with 34 have been isolated from cases. Animals deaths in young children in Malaysia. under stress are very prone to this A similar outbreak occurred in Taiwan disease. in 1998, with more than 1.4 million Frisk CS and Wagner JE (1977) Lab Anim 11 , cases and 78 deaths in young children, 79 and another outbreak in Singapore and Malaysia in 2000 when echovirus hamster herpesvirus Synonym for cri- 7 was identified in many of the severe cetid herpesvirus 1. cases. Although enterovirus 71 was the most frequently identified virus in Hamster oral papillomavirus The type association with the severe cases, there species of the genus Pipapillomavirus . are suspicions that another virus may Causes mucosal lesions in Syrian be a cofactor in pathogenesis, and there hamsters. are reports of a subgenus B adenovirus and of echovirus 7 isolated from chil- hamster osteolytic viruses Kilham rat dren who died during the epidemic. virus and related rodent parvoviruses. Cardosa MJ et al (1999) Lancet 354 , 987 Chan LG et al (2000) Clin Infect Dis 31 , 678 Hamster polyomavirus (HaPyV) A spe- Hosoya M et al (2006) Pediatr Infect Dis J 25 , cies in the genus Polyomavirus . Has 691 been isolated from the spleen and kid- Hosoya M et al (2007) J Clin Microbiol 45 , 112 neys of normal European hamsters, Cricetus cricetus . Causes fatal meningi- Hantaan virus (HTNV) A species in tis on i.c. injection into newborn mice the genus Hantavirus. Causes hem- or golden hamsters, Mesocricetus aura- orrhagic fever with renal syndrome tus . The genome DNA is 5366 bp in (HFRS) in humans. First recognized length, with about 50% homology in among soldiers in the Hantaan river the predicted amino acid sequence to region of Korea in 1951, when the dis- Murine polyomavirus . ease became known as Korean hemor- Synonyms : latent hamster virus; ham- rhagic fever. A similar hemorrhagic ster papovavirus. fever is seen in Manchuria, Russia, Scandinavia, and eastern Europe. It is Delmas V et al (1985) EMBO J 4 , 1279 Hannoun C et al (1974) Ann Microbiol 125A , a severe disease with damage to the 215 kidneys, shock, and oliguria. Shown to be transmitted to humans following hand-foot-and-mouth disease First recog- contact with striped field mice of the nized as a clinical entity and its associa- species Apodemus agrarius, the natural tion with coxsackie A virus established host for Hantaan virus. This rodent is 195 Hantavirus

widely distributed over Eastern China, renal syndrome virus; hemorrhagic Korea, and Manchuria. HFRS is char- nephrosonephritis virus; Korean hem- acterized by sudden onset of high orrhagic fever virus; nephropathia epi- fever, headache, myalgia, and severe demica virus. malaise. The fever lasts for up to 7 Kim YK et al (2007) J Infect 54 , 381 days. Hemorrhagic symptoms develop Schmaljohn CS et al (1985) Science 227 , 1041 in about a third of infections, usually Sun Y et al (2007) Antimicrob Agents Chem- after fever has subsided when a hypo- other 51 , 84 tensive phase develops with associ- Wang H et al (2000) Virology 278 , 332 ated thrombocytopenia, proteinuria, and petechial hemorrhages. Death may Hantaan 76-118 virus An isolate of occur from hypovolemic shock during Hantaan virus in the genus Hantavirus . this phase, or more likely during the next phase of oliguria as blood pressure returns to normal. Recovery is Hantavirus A genus of the family slow and convalescence may take sev- Orthobunyaviridae , consisting of at least eral weeks. Overall the mortality from 20 morphologically and serologically HFRS is estimated as 2–5%; there are related viruses, which occur world- about 100,000 cases annually in east- wide. Distinguished from other genera ern China, mostly in rural districts. As in the family as they are not transmit- with most other hantaviruses, there is ted by arthropods, but primarily infect no person-to-person transmission, and rodents, from which humans may humans appear to be a dead-end host. become infected but remain ‘ dead-end ’ Infectious virus is present in saliva, hosts. Each hantavirus species appears urine, and feces of infected rodents to have a single rodent species or sub- for several weeks after they become species as its natural host, which deter- infected, and human infection occurs mines its geographical distribution. following inhalation of aerosolized Most virus species can cause serious urine or feces if not by direct contact. diseases in humans following infection, The broad spectrum antiviral drug rib- but person-to-person transmission has avirin inhibits production of infectious not been observed except for Andes Hantaan virus through interaction with virus in South America. Hantaviruses the RNA polymerase of the virus. have never been found in arthropods. Synonyms: epidemic hemorrhagic The type species is Hantaan virus . Other fever virus; hemorrhagic fever with species are listed in Table H1 .

Table H1. Hantaviruses and their hosts

Hantavirus Disease Known or suspected host Location

Subfamily Murinae -associated viruses (Mice and Rats) Amur HFRS Apodemus peninsulae Asia (China, Russia) Da Bie Shan NR Niviventer confucianus Asia (China) Dobrava HFRS Apodemus flavicollis Europe (Slovenia) Hantaan HFRS Apodemus agrarius Asia (Korea) Saaremaa HFRS Apodemus agrarius Europe (Estonia) Seoul HFRS Rattus rattus, R norvegicus Worldwide (Korea) Thailand NR Bandicota indica Thailand Subfamily Arvicolinae -associated viruses (Voles and Lemmings) Bloodland Lake NR Microtus ochrogaster North America (USA) Hokkaido NR Clethrionomys rufocanus Asia (Japan) Isla Vista NR Microtus californicus North America (USA) Khabarovsk NR Microtus fortis Asia (Far East Russia) Prospect Hill NR Microtus pennsylvanicus North America (USA)

(continued) Hantavirus 196

Table H1. (continued)

Hantavirus Disease Known or suspected host Location

Puumala HFRS Clethrionomys glareolus Europe (Finland) Topografov NR Lemmus sibericus Russia (Siberia) Tula NR Microtus arvalis Europe (Russia) Vladivostok NR Microtus fortis Asia (Far East Russia) Subfamily Sigmodontinae -associated viruses (New World sigmodontine rodents) Anajatuba HPS Oligoryzomys fornesi South America (Brazil) Andes HPS Oligoryzomys longicaudatus South America (Argentina) Ape Aime-Itapua NR Akodon montensis South America (Paraguay) Araraquara HPS Necromys lasiuris South America (Brazil) Bayou HPS Oryzomys palustris North America (USA) Bermejo HPS Oligoryzomys chacoensis South America (Argentina) Black Creek Canal HPS Sigmodon hispidus North America (USA) Blue River NR Peromyscus leucopus North America (USA) Calabazo NR Zygodontomis brevicauda Central America (Panama) Cano Delgadito NR Sigmodon alstoni South America (Venezuela) Castelo dos Sonhos HPS Unknown South America (Brazil) Catacamas NR Oryzomys couesi Central America (Honduras) Central Plata HPS Oligoryzomys flavescens South America (Uruguay) Choclo HPS Oligoryzomys fulvescens Central America (Panama) El Moro Canyon NR Reithrodontomys megalotis North America (USA) HU39694 HPS Unknown South America (Argentina) Itapua NR Oligoryzomys nigripes South America (Paraguay) Juquitiba HPS Oligoryzomys nigripes South America (Brazil) Laguna Negra HPS Calomys laucha South America (Paraguay) Lechiguanas HPS Oligoryzomys flavescens South America (Argentina) Limestone Canyon NR Peromyscus boylii North America (USA) Maciel NR Necromys benefactus South America (Argentina) Maporal NR Oligoryzomys fulvescens South America (Venezuela) Monongahela HPS Peromyscus maniculatus North America (USA) Muleshoe NR Sigmodon hispidus North America (USA) New York HPS Peromyscus leucopus North America (USA) Oran NR Oligoryzomys longicaudatus South America (Argentina) Pergamino NR Akodon azarae South America (Argentina) Rio Mamore NR Oligoryzomys microtis South America (Bolivia) Rio Segundo NR Reithrodontomys mexicanus Central America (Costa Rica) Sangassou NR Hylomyscus alleni Africa (Equatorial Guinea) Sin Nombre HPS Peromyscus maniculatus North America (USA) Insectivore-associated viruses (Shrews) Ash river NR Sorex cinereus North America (USA) Camp Ripley NR Blarina brevicauda North America (USA) Jemez Springs NR Sorex monticolus North America (USA) Seewis NR Sorex araneus Europe (Switzerland) Tanganya NR Crocidura theresae Africa (Equatorial Guinea) Thottapalyam NR Suncus murinus Asia (India)

Disease: HFRS, Hemorrhagic fever with renal syndrome; HPS, Hantavirus pulmonary syndrome; NR, None recorded.

Plyusnin A et al (1996) J Gen Virol 77 , 2677 recognized in 1993. Viruses caus- Schmaljohn C and Hjelle B (1997) Emerg ing this disease have rodent hosts of Infect Dis 3 , 95 the subfamily Sigmodontinae , such as Peromyscus leucopus, the deer mouse. hantavirus pulmonary syndrome A Human infection may occur following cardiopulmonary syndrome caused contact with their droppings (feces or by hantaviruses in the Americas. First urine), including aerosols from them. 197 Haruna virus

There is a sudden onset of febrile ill- and various host oncogenes. HZFeSV ness in infected persons followed by strain 1 carries the transduced fes onco- an often fatal respiratory disease syn- gene, and strain 2 the abl oncogene, drome. The fatality rate in infected both of which encode tyrosine kinase. individuals is close to 40%. HZFeSV strain 4 carries the transduced Duchin JS et al (1994) N Engl J Med 330 , 949 kit oncogene and strain 5 the fms oncogene. Both encode a phosphotyrosine Hanuman langur retrovirus An exog- kinase receptor. See Feline leukemia enous simian retrovirus isolated virus . from the common Hanuman langur Hardy WD (1993) In The Retroviridae, vol. 2, (Semnopithecus entellus ) in Lucknow edited by JA Levy. New York: Plenum Press, and Jodhpur, India. Named simian p. 109 retrovirus 6 by the original discoverers, the virus is clearly related to Mason– Hare fibroma virus (FIBV) A species in Pfizer monkey virus, and appears to be the genus Leporipoxvirus . Serologically a strain of Mason–Pfizer monkey virus in related to myxoma virus. Causes fibro- the genus Betaretrovirus . mas in hares in southern France and Nandl JS et al (2003) Virology 311 , 192 northern Italy. Transmissible to rabbits. May be identical to fibrosarcoma virus Hanzalova virus (HANV) A strain of tick- of hares described by von Dungern and borne encephalitis virus in the genus Coca in 1903. Flavivirus. Isolated from the brain of Fenner F (1965) Aust J Exp Biol Med 43 , 143 a woman with meningo-encephali- von Dungern E and Coca AF (1903) Z tis in Beroun, former Czechoslovakia. ImmunForsch 2 , 391 Also isolated from Ixodes ricinus (the first tick-borne encephalitis strain hare virus See European brown hare syn- isolated in former Czechoslovakia). drome virus . Antigenically very similar or identical to Hypr and Absettarov strains of tick- Harlingen virus (HARV) An unassigned borne encephalitis virus. animal rhabdovirus, isolated from culicine mosquitoes. harbor seal herpesvirus Synonym for Phocid herpesvirus 1 . HART See hybrid arrested translation . harbor seals picorna-like virus (SPLV) An unassigned virus in the family hartebeest malignant catarrhal fever Picornaviridae. The complete genome virus Synonym for Alcelaphine herpes- sequence was determined of a novel virus 2 . virus obtained from 7.4% of 108 ringed seals (Phoca hispida ) caught off the coast Hart Park virus (HPV) An unassigned of California. The disease significance species in the family Rhabdoviridae . of this virus is unknown. With Flanders, Kamese, Mosqueiro, Kapoor A et al (2008) J Virol 82 , 311 and Mossuril viruses forms the Hart Park serogroup. Isolated from mosqui- hard pad A disease of dogs caused by toes in California, USA. Serologically canine distemper virus in which there similar viruses have often been iso- is tenderness and keratinization of the lated in many parts of the USA and skin of the feet. Later nervous involve- Canada. Multiplies in newborn mice ment and death usually occur. on i.c. injection but poorly or not at all in embryonated eggs or cell cultures. Hardy–Zuckerman feline sarcoma virus Kills newborn mice on i.c. injection but (HZFeSV) A replication-defective spe- not known to be pathogenic under nat- cies in the genus Gammaretrovirus . ural conditions. Obtained from multicentric fibrosarcomas in cats, and the result of recombi- Haruna virus Antigenically identical to nation between feline leukemia virus Getah virus. Isolated in Japan. harvest mouse virus 1 198 harvest mouse virus 1 Synonym for El serious disease and is a serious threat Moro Canyon virus . to laboratory workers and to the community, but no effective prophylaxis harvest mouse virus 2 Synonym for Rio or treatment is available. These four Segundo virus . hazard groups require that work with these viruses be carried out at Biosafety Harvey murine sarcoma virus (HaMSV) level 1, Biosafety level 2, Containment A species in the genus Gammaretrovirus . biosafety level 3, or maximum contain- The first strain of mouse sarcoma ment level 4, respectively. See biosafety virus isolated. The virus is replication and biological containment . defective, requiring a helper virus for multiplication, and acutely transform- HB virus (HBPV) A species in the genus ing. Obtained from a rat injected with Parvovirus . Isolated from a cystadeno- murine leukemia (Moloney) virus. carcinoma of the ovary of a 12-year-old A recombinant between the murine girl, one human embryo and two pla- leukemia virus and the cellular onco- centas. Serologically unlike either H-1 gene Ha-ras , which encodes the gua- virus or Kilham rat virus. Agglutinates nine triphosphate binding oncoprotein guinea pig, hamster, and rat erythro- p21ras. cytes, but not human cells. Harvey JJ (1964) Nature 204 , 1104 Toolan HW (1964) Proc Am Assoc Cancer Res 5 , 64 HAT selection Growth of cells in medium Toolan HW (1968) Int Rev Exp Pathol 6 , 135 containing hypoxanthine, aminopterin and thymidine. The presence of amin- HBcAg Hepatitis B core (nucleocapsid) opterin prevents de novo synthesis of antigen. purines or pyrimidines, and only cells having functional thymidine kinase HBeAg Hepatitis B e (truncated, soluble and hypoxanthine–guanine phosphori- core) antigen. bosyl transferase enzymes will survive and form colonies. HBsAg Hepatitis B surface (envelope) antigen. Hawaii virus (HV) A strain of Norwalk virus in the genus Norovirus observed HBV Hepatitis B virus . by electron microscopy in the feces from a family outbreak of gastroenteri- Hbx protein A protein specified by mam- tis in 1971. Antigenically different from malian (but not avian) hepadnaviruses, Norwalk virus. the function of which is unknown. Thornhill TS et al (1977) J Infect Dis 135 , 20 It consists of 154 amino acids in the human virus, 141 amino acids in Hazara virus (HAZV) A strain of woodchuck and 138 in ground squirrel Crimean-Congo hemorrhagic fever virus hepadnaviruses. Thought to play a role in the genus Nairovirus. Isolated from a in oncogenesis. tick, Ixodes redikorzevi, in Pakistan. Not reported to cause disease in humans. HCT-8 cells (CCL 244) A cell line derived by the trypsin dissociation of an ileo- hazard groups A classification system for cecal adenocarcinoma from a 67-year- viruses which varies in different coun- old male. tries but usually contains four components: (1) Unlikely to cause human HCT-15 cells (CCL 225) A colorectal disease. (2) May cause human disease adenocarcinoma cell line which is epi- and is a hazard to laboratory workers, thelial-like in morphology and is tum- but is unlikely to spread to the com- origenic in nude mice. munity. (3) Can cause severe disease and is a hazard to laboratory workers, HCT-116 cells (CCL 247) A cell but an effective vaccine prophylaxis line derived from a male patient or treatment is available. (4) Causes with colonic carcinoma, which is 199 heat-tethered virus

tumorigenic in athymic nude mice giv- precursor. After completion of one ing rise to epitheleoid tumors. ‘ headful, ’ the remaining DNA is cut and the filling of a second head begins. HCV Hepatitis C virus . HEF Hemagglutinin esterase fusion pro- HCV clade 1 (HCV-1) A cluster of geneti- tein. A peplomer protein found on cally related viruses in the species the surface of some viruses, notably Hepatitis C virus . influenza C viruses, some coronaviruses and toroviruses, HEF binds to HCV clade 2 (HCV-J6) A cluster of genet- 9-O -acetyl- N -acetyl-neuraminic acid, ically related viruses in the species and has a receptor-destroying activ- Hepatitis C virus . ity which is a neuraminate-O -acetyl esterase. The protein also has fusion HCV clade 3 (HCV-NZL1) A cluster of activity. There is homology between genetically related viruses in the spe- the HEF proteins of coronaviruses and cies Hepatitis C virus . influenza C virus. de Groot RJ (2006) Glycoconj J 23 , 59 HCV clade 4 (HCV-ED43) A cluster of Pekosz A and Lamb RA (1999) J Virol 73 , genetically related viruses in the spe- 8808 cies Hepatitis C virus . heart disease caused by viruses At least HCV clade 5 (HCV-EVH1480) A clus- 18 viruses have been associated with ter of genetically related viruses in the heart disease in humans but the sig- species Hepatitis C virus . nificance of most of them is doubtful. Rubella virus is an important cause of HCV clade 6 (HCV-EUHK2) A cluster of congenital abnormalities and coxsackie genetically related viruses in the spe- viruses are a cause of myocarditis. cies Hepatitis C virus . See DM and Tilles JC (1991) Rev Infect Dis 13 , 951 HD virus A possible species in the genus Polyomavirus . Isolated from a particular heat shock proteins Proteins that are line of Vero cells in which it does not synthesized by cells de novo or at an produce CPE. It appears to be identical increased rate following heat shock to stump-tailed macaque virus, a strain (shiftup in temperature). Seem to proof Bovine polyoma virus . tect against thermal damage, but the exact mechanism is unknown. Found Howley PM et al (1979) J Virol 30 , 400 in both prokaryotic and eukaryotic cells. Some of these proteins function hDLG A membrane-associated pro- normally as molecular chaperones, and tein which is the human homolog of certain viruses exploit these proper- Drosophila disks large TSG. Binds APC ties to assist their own replication and and negatively regulates cell-cycle pro- assembly. For example, heat shock progression. Is a target for the E6 onco- tein 70 binds transiently to the polio- protein of human papillomaviruses in virus capsid precursor, facilitating causing cell transformation. assembly, and BiP protein binds briefly to newly synthesized integral mem- HDV Hepatitis delta virus . brane proteins such as viral glycoproteins, and assists their maturation. HEAD H igh-throughput E xtraction, A mplification and D etection of RNA. Hartl FU (1996) Nature 381 , 571 Legler TJ et al (1999) J Clin Virol 13 , 95 heat-tethered virus Virus whose replication is temporarily interrupted by an headful packaging Mechanism for DNA increase in temperature. The virus is packaging which occurs during rep- not inactivated and on return to the lication in phage T4 and some other permissive temperature the replication viruses. Empty head structures are process recommences. Seen with cer- filled with DNA from a concatemeric tain Poxviridae . Heine–Medin disease virus 200

Heine–Medin disease virus Synonym Orthomyxoviridae , Paramyxoviridae , and for Poliovirus . Rhabdoviridae .

HEL 229 cells (CCL 137) A diploid helical viruses Viruses whose morphol- fibroblast-like cell line, derived from ogy displays helical symmetry. the embryonic lung tissue of a black male, is one of the few such cell lines helicase primase A herpes simplex virus to contain the A type electrophoretic enzyme that promotes ATP-dependent variant of glucose-6-phosphate dehy- unwinding of the DNA duplex during drogenase. replication. It is a target for anti-herpes viral drugs in development such as the HeLa 229 cells (CCL 2.1) A cell line deriv- thiazolylsulfonamide, BAY-57-1293. ative of the parent HeLa line, which Kleymann G (2004) Antivir Chem Chemother differs chiefly in its relative insuscepti- 15 , 135 bility to polioviruses. helper T cells Thymus-derived lympho- HeLa cells (CCL 2) The first aneuploid cytes, usually class II-MHC-restricted, epithelial-like cell line to be derived whose presence (help) is required for from human tissue and maintained the production of normal levels of anti- continuously by serial cell culture. body by B lymphocytes and also for Derived from cervical adenocarcinoma normal development of cell-mediated of a black female, He nrietta La x. immunity. HeLa S3 cells (CCL 2.2) A cell line that is a clonal derivative of the parent HeLa helper virus A virus which, in a mixed line, is especially hardy and has a plat- infection with a defective virus, pro- ing efficiency of 100%. It is readily vides some factor without which the adaptable to growth in suspension cul- defective virus cannot replicate. See ture for biochemical studies of viruses phenotypic mixing . and cells. Helpin See Brivudin helenine A fermentation product of the fungus, Penicillium funiculosum , and a hemadsorbing enteric virus of calves potent interferon inducer. This activ- Synonym for Bovine parvovirus . ity is due to the presence of a double- stranded RNA viral genome. Electron hemadsorption Adsorption of erythro- microscopic studies have revealed cytes to the surface of virus-infected numerous particles of typical virus cells. Orthomyxoviridae , Paramyxoviridae , morphology some 20–30 nm in diame- and Togaviridae , which bud from the ter, and similar to those seen in statolon cell surface, confer this property on the but serologically distinct from them. cell. When erythrocytes are added to the culture medium, adsorption to the Kleinschmidt WJ et al (1968) Nature 220 , 167 Shope RE (1966) J Exp Med 124 , 15 infected cells makes it possible to identify them in cultures which may show helical symmetry A form of symmetry no other indication of infection. in which many RNA virus capsids are constructed. Each capsomere on the hemadsorption inhibition A serologic helix consists of a single polypeptide test in which hemadsorption is inhib- molecule and establishes bonds with ited by the interaction of the serum two capsomeres on each of the adja- antibodies with the surface of the cent turns, giving stability to the cap- hemadsorbing virus. sid. The overall length of the helix is determined by the length of the RNA hemadsorption virus type 1 An old name molecule. In all animal viruses with for Human parainfluenza virus 3 . helical symmetry the nucleocapsid is folded and packed within a lipo- hemadsorption virus type 2 An old name protein envelope, e.g. Bunyaviridae , for Human parainfluenza virus 1 . 201 hematopoietic necrosis herpesvirus of goldfish hemagglutinin A protein which binds to hemagglutination inhibition (HI) A receptors on red blood cells and causes test for the presence of antibodies, them to agglutinate. An extremely e.g. in serum, that inhibit the agglu- important property of some viruses tination of red blood cells in vitro . such as influenza, since it forms the A wide variety of viruses will cause basis of the hemagglutination (HA) agglutination of certain erythrocytes, test for virus quantitation, and the including adenoviruses, arboviruses, hemagglutination inhibition (HAI) test some enteroviruses, influenza virus, to measure serum antibody levels. In parainfluenza viruses, mumps virus, addition to viruses that have a hemag- measles virus, rubella virus and reovi- glutinin on their surface, some such as ruses. Antibodies that react with either orthopoxviruses induce a hemagglu- the virus or the antigen will prevent tinin in infected cells during replica- hemagglutination in a standard test. tion. For influenza viruses, chicken red This simple test requires making dilu- blood cells are commonly used for HA tions of the patient’s serum, mixing or HAI tests, but for other hemagglu- them with a fixed amount of the virus tinating viruses it may be necessary to hemagglutinin (HA), then adding an substitute cells of other species such as erythrocyte suspension (usually 1%). goose (rubella), guinea pig (parainflu- An extremely important test for influenza), or rhesus monkey (adenovirus). enza diagnosis and subtyping. See also hemagglutination . Chernesky MA (1996) In Virology Methods Manual , edited by BWJ Mahy and HO hemagglutinin esterase fusion protein Kangro. London: Academic Press, p. 114 See HEF. hemagglutination tests See hemaggluti- hemagglutinin/neuraminidase (HN) A nation ; hemagglutination inhibition . glycoprotein on the surface of some paramyxoviruses which has both hemagglutinin–neuraminidase hemagglutinin and neuraminidase In ortho- activities. Parainfluenza and rubula- myxoviruses, the hemagglutinin and viruses have this surface protein, but in neuraminidase activities are on dif- morbilliviruses the equivalent protein ferent glycoprotein peplomers on the has only hemagglutination and no neu- virion surface, but in respiroviruses raminidase activity. both activities are located on a single peplomer called HN. hemagglutinating encephalomyelitis virus of pigs Synonym for Porcine hematocrit A measure of the percentage hemagglutinating encephalomyelitis virus . of erythrocytes in the blood (packed cell volume). A raised hematocrit is hemagglutinating virus of Japan found in severe cases of Lassa fever Synonym for Sendai virus (murine virus infection. parainfluenza virus type 1). hematogenous route of entry Spread of hemagglutination Clumping of red blood virus, especially to the central nervous cells, usually present as a 1% suspen- system, through blood. This neces- sion, by viruses which contain a red sitates virus crossing the blood–brain blood cell attachment protein (hemag- barrier, and may occur by passage of glutinin) on their surface. A large infected cells with viruses such as mea- number of animal viruses hemaggluti- sles or HIV, or directly through viremia nate a wide variety of red blood cells, with some arboviruses. An alternative each virus favoring certain cells from route, favored by some viruses such certain animals. Used as a quick, quan- as polio, rabies, or herpesviruses, is titative assay for certain viruses, espe- through peripheral nerves. cially influenza virus. Some viruses, such as orthopoxviruses, induce a hematopoietic necrosis herpesvirus of hemagglutinin in infected cells but it is goldfish Synonym for cyprinid herpes- not present on the virion surface. virus 2. hematopoietic stem cells 202 hematopoietic stem cells Large cells hemorrhagic nephroso-nephritis virus found in the bone marrow or other Synonym for Hantaan virus . hematopoietic tissues such as the spleen which are the progenitors of hemorrhagic septicemia virus of fish cells of the immune system as well as Synonym for Viral hemorrhagic septi- the vascular system. They differentiate cemia virus . under the influence of colony stimulating factors. Henderson–Paterson bodies An old name for the molluscum body pro- Hemophilus influenzae A bacterium that duced in epidermal cells of patients is frequently associated with secondary infected with Molluscum contagiosum infection during influenza virus infec- virus . An inclusion body. tion. Can be treated with antibiotics. Hendra virus (HeV) The type species hemorrhagic conjunctivitis virus See of the genus Henipavirus , related to acute hemorrhagic conjunctivitis Nipah virus but distinct from other virus . paramyxoviruses by genetic analysis and extended host range. Formerly hemorrhagic encephalopathy of rats called equine morbillivirus, but now virus A strain of Kilham rat virus in the known to be a virus which primarily genus Parvovirus which is highly path- infects large fruit bats, Pteropus sp, and ogenic on injection i.c. into newborn is not a morbillivirus. Caused a serious rats. Isolated by i.c. injection of new- outbreak of acute respiratory disease born rats with brain and spinal cord in the Hendra stables, near Brisbane, extracts from Lewis rats which became Queensland, Australia in 1994 in paralyzed after treatment with cyclo- which 14 horses and their trainer died. phosphamide. Causes hemorrhage and A stablehand was also infected and necrosis in the spinal cord. Injection hospitalized, but survived. In 1995 a into newborn hamsters causes an acute second human death, this time from fatal infection. Adult rats, newborn encephalitis, occurred in Mackay, mice, rhesus, and cynomolgus mon- Queensland in a man who had helped keys are insusceptible. Agglutinates in a postmortem examination of two guinea pig erythrocytes, but not horses 1 year earlier. Experimentally, chicken, sheep, rhesus monkey, or Hendra virus has produced disease human cells. in cats, guinea pigs, and horses, but Synonym : Her virus. not in mice, rabbits, chickens, or dogs. Cole GA et al (1970) Am J Epidemiol 91 , 339 The incubation period in horses is Nathanson N et al (1970) Am J Epidemiol 91 , 8–11 days. There is depression, loss of 328 appetite, fever, labored respiration, followed by substantial nasal discharge, hemorrhagic enteritis of turkeys virus and death within 2 days after onset of See turkey adenovirus 3 . symptoms. Lungs show hemorrhagic as well as pneumonia-like lesions. hemorrhagic fever viruses Synonym Sampling of 2000 horses in Queensland for viral hemorrhagic fever viruses revealed no evidence of infection of humans. Includes members of the except in those related to the Hendra Arenaviridae , Bunyaviridae , Flaviviridae , stables. Antibodies can be found in all and Filoviridae . four species of fruit-eating bats (flying foxes) of the genus Pteropus found hemorrhagic fever with renal syndrome in Queensland, and infectious virus virus Synonym for Hantaan virus . was isolated from one pregnant fruit bat. In January 1999 a horse in Cairns, hemorrhagic nephritis A disease asso- Queensland that had died of unknown ciated with reactivation of human causes was found on postmortem to parvovirus BKV, especially in the pres- have been infected with Hendra virus. ence of immunosuppression caused by In 2007 there were two fatal cases HIV-AIDS. due to Hendra virus in horses, but no 203 Hepadnaviridae

associated human cases. Sequence Simmonds P and Mutimer D (2005) In analysis of the negative single-stranded Topley & Wilson’s Microbiology and Microbial RNA genome of Hendra virus reveals Infections , vol. 2, Tenth edition, edited by an exceptionally large molecule BWJ Mahy and V ter Meulen. London: Hodder Arnold, p. 1189 (18,234 nt), which has only limited homology with all other paramyxoviruses except Nipah virus. Hepadnaviridae A family consisting of DNA viruses which infect humans Field HE et al (2007) Aust Vet J 85 , 268 Gould A (1996) Virus Res 43 , 17 and a variety of other species includ- Hanna JN et al (2006) Med J Aust 185 , 562 ing woodchucks, squirrels, herons Hyatt AD and Selleck PW (1996) Virus Res and ducks. The mammalian and 43 , 1 avian species are in different genera, Lin-Fa Wang et al (2000) J Virol 74 , 9972 Orthohepadnavirus and Avihepadnavirus , Thornley M (2005) Aust Vet J 83 , 2 because of differences in genome structure and pathogenesis. The virus Henipavirus A genus in the family infecting humans is composed of a Paramyxoviridae containing two spe- spherical, enveloped particle (the Dane cies, Hendra virus and Nipah virus that particle), 42 nm in diameter, with no are indigenous to fruit bats (Pteropus evident surface projections, but small spp) and are associated with high mor- spherical particles, 22 nm in diameter tality in infected humans or animals. (HBsAg particles), are also present in The genome is 3000 nt longer than that the plasma of carriers. The virus par- of other paramyxoviruses. The attach- ticle consists of a 27 nm icosahedral ment protein (G) lacks hemagglutinat- nucleocapsid (the core particle) con- ing or neuraminidase activities. taining one major polypeptide species (mol. wt. 20 K) surrounded by a deter- Harcourt BH et al (2001) Virology 287 , 192 gent-sensitive envelope. The lipid- containing envelope contains the Hepacivirus A genus in the family surface (S) antigen (HBsAg, Australia Flaviviridae which includes Hepatitis antigen) against which virus neutral- C and related viruses. Transmission izing antibodies are directed. The between humans occurs primarily envelope is composed of three major through exposure to blood or blood proteins: S (p24, gp27); M (p33, gp36); products carrying the virus. There is and L (p39, gp42). The 22 nm surface no known invertebrate vector. Virions antigen (HBsAg) particles consist are spherical, 50 nm in diameter, and largely of S proteins (226 amino acids), contain a molecule of linear positive- which can assemble in the absence sense single-stranded RNA about 9.6 kb of cores. The M proteins (281 amino long. The 5 -NCR is about 340 nt long acids) are composed of p24 with an and contains an internal ribosomal additional 55 amino acids at the N-ter- entry site. The virion proteins include minus containing the pre-S2 domain. a nucleocapsid protein C (p19) and The L proteins (400 amino acids) con- two envelope proteins, E1 (gp31) and tain an additional 119 amino acids at E2 (gp70). Two nonstructural proteins, the N-terminus, containing the pre-S1 NS2 and NS3, are autocatalytically domain. HBsAg cross-reacts between derived from a single precursor mol- human, woodchuck, and ground squir- ecule by a Zn-dependent proteinase rel hepadnaviruses, but not with avian activity that is not found in the other hepadnaviruses. The virion core con- genera of the Flaviviridae . The genome tains a single polypeptide (p22) which consists of a single large open reading has both the core antigen (HBcAg) and, frame (ORF) which encodes a poly- in truncated form (p16), the soluble protein of about 3000 amino acids. The e antigen (HBeAg) specificities. The gene order is 5 -C-E1-E2-p7-NS2-NS3- core also contains enzyme activities NS4A-NS4BNS5A-NS5B-3 . At present involved in replication: DNA polymer- Hepatitis C virus is the only species in ase, reverse transcriptase and protein the genus, but several distinct genomic kinase. The genome is a single mol- clades are recognized. ecule of non-covalently closed, circular Hepadnaviridae 204

DNA (3.0–3.3 kb in different viruses) heparin A highly sulfated gly- which is partially single-stranded and cosaminoglycan that has anticoagu- partially double-stranded. The long lant activity; it inhibits the action of strand is termed negative and is com- thrombin by activating antithrombin plementary to viral mRNA. It encodes III and interfering with the blood- four open reading frames, the core, clotting cascade. polymerase, envelope and X gene. The X gene encodes a transactivator pro- hepatic steatosis A condition known as tein and is absent from avian hepadna- fatty liver which can be caused by hep- viruses. The positive strand varies from atitis C virus infection. 1.7 to 2.8 kb in different molecules. The negative strand is not a closed circle hepatitis-associated antigen See but has a nick at a unique site, which Australia antigen . differs between mammalian and avian hepadnaviruses; there is a polypeptide Hepatitis A virus (HAV) Type species of covalently attached to its 5 end which the genus Hepatovirus . Virion diameter acts as the primer for DNA synthe- 27–29 nm, density in CsC1: 1.32–1.34 g/ sis. The positive strand is primed by ml, sedimenting at 160S in sucrose. a short 19-base RNA molecule at its 5 The genome RNA is positive-sense, end. Replication involves the genera- 7.48 kb in length, with a VPg protein covalently attached at the 5 end, and tion of a closed-circular DNA molecule within the cell nucleus, and synthesis of poly A at the 3 end. There is a single a 3.4 kb plus-strand RNA that serves as open reading frame encoding a poly- mRNA and also as template for synthe- protein of 2235 amino acids. The virus sis of negative-strand DNA. The exact replicates in various primate cell cul- mechanisms of cell entry and exit are tures after adaptation, usually without not currently known. Hepadnaviruses cytopathic effects. Causes enterically are highly host-specific, and suitable in transmitted ‘ short incubation ’ hepati- vitro cell culture systems to study these tis (less than 6 weeks) in humans. The events have not been found for mam- virus is present in the feces during the malian viruses. Fetal duck liver hepato- prodromal phase of the disease but cytes support growth of avian viruses. usually disappears about the time jaundice appears. Chronic carriers of the Gerlich WH and Kann M (2005) In Topley & virus are not seen and the virus does Wilson’s Microbiology and Microbial Infections , not cause progressive liver disease. vol. 2, Tenth edition, edited by BWJ Mahy and V ter Meulen. London: Hodder Arnold, Epidemics of hepatitis A are usually p. 1226 due to water- or food-borne infection. Mason WS and Seeger C (Editors) (1991) Antibodies may be demonstrated by Curr Top Microbiol Immun 168 , 206 pp CFT, immune adherence, hemaggluti- Nassal M (1996) Curr Top Microbiol Immunol nation, and radioimmunoassay. They 214 , 297 appear soon after the onset of jaundice and persist. They increase in fre- HEPA filter H igh e fficiency p articulate a ir quency with age and reach a peak at filter, used in biological safety cabinets age 50 years. Chimpanzees, owl mon- of Class II level and higher to provide keys, and marmosets are susceptible clean input air for product (e.g. cell to experimental infection and the virus culture) protection as well as filtered can be demonstrated in their hepato- exhaust air for environmental protec- cytes. Simian virus strains related to tion. In some cases used in series for but genetically distinct from the human added protection. virus have been isolated from African heparan sulfate A glycosaminoglycan green and cynomolgus monkeys. An constituent of membrane-associated excellent formalin-inactivated vaccine proteoglycans. Found on cells of the is now licensed for use in people at risk lungs, arterial walls and many other for hepatitis A infection. cell surfaces. Related to heparin and Synonyms : catarrhal jaundice virus; epi- containing the same disaccharide demic jaundice virus of humans. repeating units, but is less sulfated. Lemon SM (1992) Rev Med Virol 2 , 73 205 Hepatitis C Virus (HCV)

Robertson B (2005) In Topley & Wilson’s persists for years and is a valuable Microbiology and Microbial Infections , vol. marker of previous infection. HBeAg 2, Tenth edition, edited by BWJ Mahy and appears during the incubation period V ter Meulen, London: Hodder Arnold, p. but disappears more rapidly than 1160 Werzberger A et al (1992) N Engl J Med 327 , HBsAg. Persistence beyond 3–4 weeks 453 may herald a chronic infection, which carries risk of hepatic cirrhosis or hepa- Hepatitis B virus (HBV) Type species of tocellular carcinoma. Not all chronic the genus Orthohepadnavirus . Causes carriers of hepatitis B virus develop ‘ long incubation ’ hepatitis (more than chronic liver disease. A few infections 60 days). Classically, infection results result in acute hepatic failure probably from the inoculation of serum from a due to an antigen–antibody reaction. carrier during blood transfusion, vac- Immune serum globulin (ISG) and cination, tattooing or ear-piercing with hyperimmune globulin (HIG) are effec- inadequately sterilized instruments. tive in providing passive immunity. A However, non-parenteral routes are plasma-derived vaccine consisting of also important and many cases result HBsAg is effective and widely used, from domestic and sexual contact, especially in developing countries. This especially homosexual practices. The has been replaced in most other coun- complete virus is an enveloped particle tries by genetically engineered vaccines 42 nm in diameter, but small spherical derived from HBsAg produced in yeast particles 22 nm in diameter and tubu- (Recombivax, Engerovax), which pro- lar forms of the same diameter but up vide solid immunity after three injec- to 100 nm long, are also present in the tions. A variety of nonhuman primates plasma of carriers. The viral genome is can be infected and on several occa- 3.2 kb long, and serves in infected cells sions zoo primates, especially chim- as the template for a 3.4 kb RNA species panzees, have been found to be chronic (pre-genomic RNA), with redundant carriers. The source of their infection is ends and short sequence repetitions, not clear but may be in the wild since that is packaged into core particles. some human cases have been asso- The pre-genomic RNA is then reverse- ciated with contact with wild, non- transcribed to form negative-strand human primates, and hepadnaviruses DNA, which in turn serves as a tem- have now been found in several spe- plate for positive-strand DNA synthe- cies including chimpanzees, gibbons, sis. Virus may be detected in the plasma orangutans, and woolly monkeys. by electron microscopy or by various Synonym : serum hepatitis virus. tests for one of three viral antigens: (1) Baumert TF et al (2007) World J Gastroenterol surface antigen (HBsAg), present on 13 , 82 the surface of all three types of virus Ganem D and Varmus HE (1987) Annu Rev particle; (2) core antigen (HBcAg), Biochem 56 , 651 which is exposed when the outer Hu X et al (2000) Proc Natl Acad Sci 97 , 1661 membrane of the complete virus is dis- MacDonald DM et al (2000) J Virol 74 , 4253 rupted with a detergent; and (3) e anti- Takahashi K et al (2000) Virology 267 , 58 gen (HBeAg), present in the core of the complete particles. Presence of e anti- ‘ Hepatitis C-like viruses ’ Originally a gen in serum suggests high infectivity. genus in the family Flaviviridae which HBsAg is present in the blood 4 weeks includes Hepatitis C and related viruses. prior to the development of symp- In 1996 it was changed to Hepacivirus . toms and usually disappears 6 weeks after the onset of symptoms. If present Hepatitis C Virus (HCV) The only species beyond 13 weeks it is likely that the in the genus Hepacivirus and the major patient will become a chronic carrier cause of hepatitis non-A non-B disease. and may develop chronic liver disease. The virion has not been visualized but Anti-HBs does not usually appear until it is believed, from physicochemical convalescence. HBcAg does not appear studies of the infectious agent, to be in the blood. Anti-HBc appears dur- spherical, enveloped and about 50 nm ing the disease and, unlike anti-HBs, in diameter. In 1988 the genome of Hepatitis C Virus (HCV) 206

hepatitis C virus was molecularly thrice weekly) plus ribavirin (Virazole) cloned from infected chimpanzee tis- which is effective in 50% of cases dur- sues, and later completely sequenced. ing prolonged treatment, although It is a linear, single-stranded RNA of relapses usually occur when the drug positive polarity with a 5 untranslated is withdrawn. region of about 340 nt followed by Synonyms: NANB hepatitis virus; non- a large open reading frame of about A non-B hepatitis virus. 9400 nt and a short 3 untranslated Alter MJ (1997) Hepatology 26 , 62S region of 50 nt, with no 3 -terminal Bartenschlager R et al (2003) Antiviral Res 60 , poly A. The genome organization is 91 similar to that of flaviviruses and pesti- Blight KJ et al (2000) Science 290 , 1972 viruses, and encodes a large protein Lohmann V et al (1999) Science 285 , 110 (3000 amino acids) that is cleaved into Major ME and Feinstone SM (1997) seven proteins: three structural and Hepatology 25 , 1527 four nonstructural. The gene order is 5 -C-E1-E2-p7-NS2NS3-NS4A,NS4B- hepatitis D virus Synonym for Hepatitis NS5A-NS5B-3. The C (core) protein delta virus . is highly basic and not glycosylated. E1 and E2 are glycosylated membrane Hepatitis delta virus (HDV) A satellite proteins. In vitro replication has not so virus, the only species in the genus far been observed, so little is known Deltavirus. First recognized when a concerning the nonstructural proteins. novel antigen (delta antigen) was The entire genome, except for the 5 observed in the nuclei of hepatocytes untranslated region, is highly variable of some patients with chronic hepati- and viruses sequenced from different tis B virus (HBV) infection. Virions are regions of the world form at least six spherical, about 34 nm in diameter, with genotypes and at least 30 subtypes. The no surface projections. Transmission of virus is difficult to grow in cell culture, HDV is dependent upon HBV, since but transfection of a hepatoma cell line it uses HBsAg as its own virion coat. is possible, and adaptive mutations in The HDV genome is a small single- the virus genome allow efficient repli- stranded circular RNA comprised of cation in hepatoma cell lines. No neu- 1675 nt with about 70% base-pairing so tralizing antibodies have been found, that the RNA forms a largely double- and experimentally infected primates stranded, rod-shaped structure. A sin- are not immune to super-infection. In gle conserved open reading frame in humans the incubation period to dis- the negative sense encodes the hepati- ease following infection is 6–8 weeks, tis delta antigen, which consists of two but about 75% of infections are subclin- protein species: one contains 195 amino ical and give rise to chronic persistent acids (24 kDa) and the other is identi- infection. Of these, some 50% develop cal except for an additional 19 amino chronic liver disease with persist- acids at the C-terminus (27 kDa). Most ently elevated liver enzymes in serum. sera contain equal amounts of the two About half of the patients with chronic species of antigen, which appears to liver disease go on to develop chronic function during replication through its active hepatitis and 5–20% of these will nuclear localization and RNA-binding progress to cirrhosis of the liver by five properties. Once inside the nucleus, years from onset. There is also a clear replication is carried out by the host link between HCV infection and the cell RNA polymerase and is independ- development of hepatocellular carci- ent of HBV virus. HDV has only been noma, since up to 75% of such patients isolated from humans and is widely have anti-HCV serum antibodies. The distributed geographically, with some development of anti-HCV screening 5% of HBsAg carriers infected world- tests has greatly reduced the incidence wide. Following infection, there is an of HCV infection through transfu- incubation period of 3–7 weeks before sions. There is no vaccine and the only fatigue, anorexia, and nausea occur, available drug for chronic infection followed by jaundice and other symp- is interferon alpha (2 million units toms of hepatitis. A high proportion 207 Hepatitis G virus (HGV-1)

(60–70%) of patients with chronic HDV signal peptide. Between ORF1 and infection develop cirrhosis, and up to ORF2 the genome encodes a small 20% develop fulminant hepatitis. No protein (123 amino acids) of unknown specific treatment is available, but since function in ORF3, which overlaps ORF1 HDV is dependent upon HBV infec- by one base. Screening tests for anti- tion, vaccination against HBV provides bodies are being developed which use protection. peptide or recombinant-expressed anti- Synonyms : delta agent; delta virus; epi- gens. The use of these prototype assays demic hepatitis-associated antigen. suggests that anti-HEV serum antibody Lai MMC (1995) Annu Rev Biochem 64 , 259 prevalence in the USA population is Taylor JM (2005) In Topley & Wilson’s less than 1%, but it appears to be higher Microbiology and Microbial Infections , vol. in some endemic areas. Antibodies 2, Tenth edition, edited by BWJ Mahy and are also found in a high percentage of V ter Meulen. London: Hodder Arnold, wild rats, Rattus norvegicus , in the USA, p. 1269 and the virus appears to be endemic in swine, and in wild Sika deer Cervus Hepevirus A floating genus of positive- nippon, in Japan. Transmission from a sense single-stranded RNA viruses pet pig to its owner has been reported, which cause outbreaks and sporadic but it is still not clear whether hepatitis cases of enterically transmitted hepa- E is a zoonotic disease. However, it has titis in humans. The type species is been established that thorough cooking Hepatitis E virus . The genome is 7.2 kb of pork will inactivate the virus. in length with a 5 m7 G cap and a 3 Berke T et al (1997) J Med Virol 52 , 419 poly(A) tail. There is one tentative Feagins AR et al (2007) J Gen Virol 88 , 912 species in the genus, avian hepatitis E Panda SK et al (2007) Rev Med Virol 17 , 151 virus. Purdy MA et al (1993) Semin Virol 4 , 319 Reyes GR and Baroudy BM (1991) Adv Virus Hepatitis E virus (HEV) The type species Res 40 , 57 of the genus Hepevirus , which causes acute enterically transmitted epidem- hepatitis F virus A novel agent claimed ics of hepatitis, especially in young to to be responsible for sporadic non- middle-aged persons in Asia, Africa, and A non-B hepatitis in humans. The South America. Large epidemics with infectious material came from French many thousands of cases have occurred patients, so the F stands for ‘ French ori- in India, Burma, and Kyrgystan. The gin.’ The genome consists of a double- virus was molecularly cloned, by the stranded DNA of approximately 20 kb. SISPA technique, from virus-enriched The particles are infectious in mon- bile obtained from experimentally keys, conferring immunity. Epidemics infected cynomolgus monkeys, but so have occurred in England, Italy, France, far has not been reliably grown in cell USA, and India. These claims need culture. The genome is a 7.5 kb single independent confirmation and to date positive-stranded RNA molecule with a this has not happened. 3 poly A tail and a capped 5 terminus Deka N et al (1994) J Virol 68 , 7810 (in contrast to most caliciviruses, which have a VPg covalently attached at the Hepatitis G virus (HGV-1) A strain of GB 5 terminus). The nonstructural proteins virus C, an unassigned species in the are encoded toward the 5 end, and family Flaviviridae , identified by molec- structural proteins toward the 3 end. ular cloning, expression and immuno- Three separate open reading frames reactivity in the plasma of a patient are used. ORF1 encodes up to six non- with chronic hepatitis. The genome is structural proteins, which include a 9792 nt long, positive-sense, contain- methyltransferase, cysteine protease, ing a single, long open reading frame helicase, and RNA replicase. There is encoding a precursor polyprotein of a 37-base untranslated region between 2873 amino acids. The genome organi- ORF1 and ORF2 which is synthesized zation and sequence place the virus as a subgenomic RNA and encodes within the Flaviviridae, distantly related the structural protein(s) preceded by a to hepatitis C virus and GB viruses Hepatitis G virus (HGV-1) 208

A and B. It is so closely related to GB and gastroenteritis, with fever, jaun- virus C, which was discovered earlier, dice, abdominal pain and occasional that it is often referred to as GB virus- diarrhea. Persistent infection does not C/Hepatitis G virus. The virus appears occur in vivo , and the viruses are not to have a global distribution and per- associated with chronic hepatitis. There sists in serum of hepatitis patients for is an excellent vaccine for prevention of at least 7 years. Although there is evi- human hepatitis A virus infection. See dence that the virus is hepatotropic, in Hepatitis A virus . the majority of cases it does not cause hepatitis. Hep-G2 cell line A human hepatocyte Hadlock KG et al (1997) Transfus Med Rev 12 , carcinoma cell line established from a 94 25-year-old male Caucasian. Linnen J et al (1996) Science 271 , 505 Mushahwar IK (2000) J Med Virol 62 , 399 Hep-2 cells (CCL 23) A heteroploid cell Simons JN et al (1995) Nat Med 1 , 564 line derived from tumors produced in irradiated and cortisone-treated wean- hepatitis GB viruses See GB viruses . ling rats after injection with epidermoid carcinoma tissue from the larynx hepatitis infectiosa canis virus Synonym of a 56-year-old human male. for Canine adenovirus . HEPT 1[(2-Hydroxyethoxy)methyl]-6 hepatitis non-A non-B virus In the mid- (phenylthio)thymine. Reported to be a 1970s, when diagnostic tests were specific inhibitor of human immunode- established for hepatitis A and hepa- ficiency virus type 1 replication. titis B viruses, it became clear that much transfusion-related hepatitis was Baba M et al (1989) Biochem Biophys Res Commun 165 , 1375 not caused by either virus. The term ‘ non-A non-B ’ was used together with Her virus hemorrhagic encephalopa- a diagnosis of exclusion to describe See thy of rats virus the agent(s) responsible for this disease. See Hepatitis C , E , F , G , and GB viruses . Heron hepatitis B virus (HHBV) A species in the genus Avihepadnavirus . hepatocytes Epithelial liver cells. Discovered in 1988 in sera of gray herons, Ardea cinerea , from Germany. hepato-encephalomyelitis virus Proto- Shares sequence homology with duck type strain of reovirus type 3, isolated in hepatitis B virus (overall about 78%) Sydney, Australia in 1953 from a child but differs in that a highly conserved with bronchopneumonia, alopecia, and open reading frame is present upstream conjunctivitis. of C in a position analogous to the X gene of orthohepadnaviruses, and the S Stanley NF et al (1953) Aust J Exp Biol Med Sci 31 , 147 protein and not the L protein possesses a potential myristylation site. The virus Hepatovirus A genus in the family does not infect ducks, but in the gray Picornaviridae comprised of one spe- heron it infects the liver and produces cies: Hepatitis A virus , with two strains, a viremia. In one survey about 30% of human hepatitis A virus and simian herons in Germany were infected, but hepatitis A virus. There is one tentative little is known of the worldwide inci- species in the genus, avian encephalo- dence and distribution. myelitis-like virus. Virions are stable, Netter HJ et al (1997) J Gen Virol 78 , 1707 resistant to acid pH and elevated tem- Triyatni M et al (2001) J Gen Virol 82 , 373 peratures. There are a large number of strains of hepatitis A virus which dif- herpangina A short febrile illness with fer by less than 20% in genome RNA sore throat, chiefly affecting young sequence homology, but there is little children in the summer in which there sequence similarity with other picorna- are small papules or vesicles around viruses. The viruses cause hepatitis the fauces, which soon break down 209 Herpesviridae

into shallow ulcers. Dysphagia, fever, surface projections. The capsomeres are vomiting, and prostration may occur. hexagonal in cross-section and have a Classically caused by human coxsackie hollow running down half their length. A viruses, particularly types 1–6, 8, 10, The genome consists of linear double- and 22. Sporadic cases have been asso- stranded DNA 124–235 kb in length, ciated with human coxsackie viruses depending on the species, with both A7, A9, B1–5 and human echoviruses terminal reiterations and internal repe- 6, 9, 16, and 17. titions. G C content 32–75%. There are more than 20 structural polypeptides, herpes ateles 2 Synonym for Ateline her- mol. wt. 12,000–220,000. Lipid content pesvirus 2 . is variable and located in the envelope. Carbohydrate is covalently linked to herpes febrilis Synonym for Human her- envelope proteins. After attachment pesvirus 1 or 2 . and penetration the genome reaches the nucleus where the viral DNA is herpes gladiatorum, herpes rugbeiorum transcribed. mRNA passes to the cyto- Synonyms for scrum-poxvirus. plasm for translation. Virus-specific enzymes and other factors involved in herpes simiae virus Synonym for herpes- replication and DNA metabolism are virus B. induced. Viral DNA is replicated in the nucleus and immature nucleocap- herpes simplex virus group Synonym for sids are formed which bud through the the genus Simplexvirus . inner lamella of the nuclear membrane, acquiring an envelope and becoming herpes simplex viruses 1 and 2 There are infective. Virus particles accumulate two antigenic types: herpes simplex between the inner and outer lamellae type 1 is a synonym for Human herpes- of the nuclear membrane and in the virus 1 ; herpes simplex type 2 is a syno- cysternae of the endoplasmic reticulum nym for Human herpesvirus 2 . from where they are released to the cell surface. Margination of chromatin and herpes venatorum Synonym for intranuclear inclusion bodies are char- scrumpox. acteristic of herpesvirus-infected cells. Production of infectious progeny virus Herpesvirales A proposed taxon for an is accompanied by destruction of the Order containing all known herpesvi- infected cell. All herpesviruses studied ruses in three families, the Herpesviridae , so far are also able to remain latent in the proposed Alloherpesviridae, and the their host cells. In this case, the genome proposed Malacoherpesviridae . becomes closed-circular and only a small subset of viral genes is expressed. Herpesviridae A diverse family of DNA There are many species and they are viruses with characteristic morphology. found in most eukaryotic hosts which There are three recognized subfamilies: have been examined in detail, from Alphaherpesvirinae , Betaherpesvirinae , humans to oysters. Some species are and Gammaherpesvirinae . Classification cell-associated and free infective units is based formally on genetic content. are scarce. A few are oncogenic. Many The virion is 100–200 nm in diameter. cause mild or silent infections in their Buoyant density (CsCl): 1.20–1.29 g/ natural hosts but severe disease in oth- ml. Consists of four structural compo- ers. They replicate with CPE in cell cul- nents: (1) the core, a protein fibrillar tures and some species produce pocks spool on which the DNA is wrapped; on the CAM. (2) the capsid, 100–110 nm in diameter, composed of 12 pentameric and Davison AJ and Clements JB (2005) In Topley & Wilson’s Microbiology and Microbial 150 hexameric capsomeres arranged Infections , vol. 2, Tenth edition, edited by with icosahedral symmetry; (3) the BWJ Mahy and V ter Meulen. London: tegument, an amorphous asymmetri- Hodder Arnold, p. 488 cal layer between the capsid; and (4) Homa FL and Brown JC (1997) Rev Med Virol the envelope, a bilayer membrane with 7 , 107 Herpesviridae 210

Montague MG and Hutchison CA (2000) herpesvirus saimiri Synonym for Proc Natl Acad Sci 97 , 5334 Saimiriine herepesvirus 2 . Roizman B et al (1992) Arch Virol 123 , 425 herpesvirus salmonis Synonym for sal- herpesvirus aotus 1 Synonym for Aotine monid herpesvirus 1. herpesvirus 1 . herpesvirus sanguinus Synonym for cal- herpesvirus aotus 3 Synonym for Aotine litrichine herpesvirus 1. herpesvirus 3 . herpesvirus scophthalmus Synonym for herpesvirus ateles Synonym for Ateline pleuronectid herpesvirus. herpesvirus 2 . herpesvirus simiae Synonym for herpes- herpesvirus ateles strain 73 Synonym for virus B. ateline herpesvirus 3. herpesvirus simian agent 8 (SA 8) herpesvirus B See Cercopithecine herpes- Synonym for Cercopithecine herpesvirus 1 . virus 2 . herpesvirus caprae Synonym for Bovine herpesvirus suis Synonym for pseudo- herpesvirus 5 . rabies virus. herpesvirus cuniculi Synonym for lep- herpesvirus sylvilagus Synonym for leporid herpesvirus 2. orid herpesvirus 1. herpesvirus cyclopsis Synonym for cer- herpesvirus T (tamarinus) Synonym for copithecine herpesvirus 13. Saimiriine herpesvirus 1 . herpesvirus gorilla Synonym for Pongine herpesvirus varicellae Synonym for herpesvirus 3 . Human herpesvirus 3 . herpesvirus hominis Synonym for herpes zoster An acute self-limited infec- Human herpesvirus 1 and/or 2 . tious disease (known also as shingles or zona) caused by reactivation of latent herpesvirus M Synonym for Saimiriine VZV (human herpesvirus 3) infection. herpesvirus 1 . It occurs frequently in persons who are severely immunocompromised, e.g. herpesvirus marmota Synonym for mar- as a result of HIV infection, and may modid herpesvirus 1. be fatal, and in up to 20% of normal immunocompetent persons over 45 herpesvirus ovis Synonym for Ovine years of age. Symptoms include vesic- herpesvirus 2 . ular lesions similar to those of varicella which usually follow an anatomical herpesvirus pan Synonym for Pongine route around the body along the dorsal herpesvirus 1 . route ganglia as the virus spreads from cells along the neurons to epithelial herpesvirus papio Synonym for cells of the skin. The condition may be Cercopithecine herpesvirus 12 . painful, and although the lesions disappear in a few weeks a chronic debili- herpesvirus papio 2 Synonym for Cerco- tating pain (post-herpetic neuralgia) pithecine herpesvirus 16 . often begins about a month after the onset of skin lesions and may persist herpesvirus platyrrinhae type Synonym for months. for Saimiriine herpesvirus 1 . Herpetoviridae A proposed but not herpesvirus pottos Synonym for lorisine adopted name for the family now herpesvirus 1. known as Herpesviridae . 211 high voltage electrophoresis

Herpid Trade name for a 5% solution of heteroploid Having a chromosome idoxuridine in dimethyl sulfoxide. number that is neither the haploid nor the diploid number normal for the Hershey and Chase experiment An species. important experiment carried out in 1951 using bacteriophage T2 that heterozygosis Having one or more pairs demonstrated the importance of virus of dissimilar alleles. nucleic acid rather than protein for transmission of infection. HEV See Hepatitis E virus . Hershey AD and Chase M (1952) J Gen Physiol 36 , 39 hexamer A group of six protein subunits which form a capsomere on the trian- Hershey Medical Center virus A her- gular faces of capsids with icosahedral pesvirus found in human cells trans- symmetry. See also hexon . formed by cytomegalovirus, and subsequently shown to be bovine her- hexon One of a group of six protein units pesvirus 1. Presumably a contaminant on the triangular faces of an icosahe- from the fetal calf serum used in the dral capsomere. culture medium. HFL1 cells (CCL 153) A human fetal Geder L et al (1978) J Virol 27 , 713 lung fibroblast cell line derived from the lung tissue of a 16- to 18-week-old heteroduplex A double-stranded DNA human fetus. molecule in which the strands do not have completely complemen- HG 261 cells (CCL 122) A cell line derived tary base sequences. Regions of non- from the skin biopsy of a 6-year-old complementarity can often be identi- Caucasian male with Fanconi’s anemia fied in electron micrographs of the that exhibits a higher percentage of DNA, and used to map their position transformed colonies following SV40 (heteroduplex mapping). virus infection than normal diploid cell cultures. heteroduplex mobility assay A genotyping assay in which electrophoretic HHBV Abbreviation for Heron hepatitis B mobility is measured following duplex virus . formation with reference amplicons. hierarchical system, viral classification Hestekin CN and Barron AE (2006) Electrophoresis 27 , 3805 Virus classification is based on five taxa, ranked as order, family, subfamily, heterogenotes Bacteria with parts of two genus, and species. Below species level prophages. The first prophage is called are terms, used by working virologists, the ‘ endogenote ’ and the second the such as strains, variants, or genotypes, ‘ exogenote. ’ which may be helpful but have no offi- cial standing within the ICTV. heterogenous nuclear ribonucleoprotein (hnRNP) complex proteins RNA bind- high 5 cells Adherent insect cells derived ing proteins involved in the assembly from egg cell homogenates of lepidop- of the ribonucleosome and in mRNA tera, Trichoplusia ni. Used for propaga- biogenesis in normal cells, and are tion of baculoviruses and expression of thought to play a role in the replication high-level recombinant proteins. of certain RNA viruses (e.g. coronaviruses, flaviviruses). high-risk human papillomaviruses (HR- HPVs) See Human papillomavirus . heterokaryon Hybrid cell formed by fusion of two cells of different species. high voltage electrophoresis Electro- See homokaryon . phoresis at potential differences of more than 1000 V. Used in nucleic acid heterologous interference See inter- sequencing and in paper electrophore- ference . sis of nucleotides. Highlands J virus (HJV) 212

Highlands J virus (HJV) A species in Millon’s reaction is positive for histones the genus Alphavirus. Isolated from but negative for protamines. Found in rodents, birds, and mosquitoes in USA. cell nuclei in varying amounts in close Disease has been seen in turkeys, and association with DNA. There are five horses infected with the virus. No main classes, differing in their relative known association with disease in content of lysine and arginine. humans. HIV Human immunodeficiency virus highly active antiretroviral therapy type 1 or 2 (HAART) The use of a combination of three or more drugs to treat patients Hivid (zalcitabine, ddC) 2 ,3 Dide- infected with HIV. Usually include oxycytidine, a nucleoside analog used both nucleic acid replication and pro- in the treatment of HIV infection. tease inhibitors. Their use has greatly HIX virus improved AIDS therapy, and can A murine non-transforming now be obtained in a single pill. See type C retrovirus with properties of HAART . both an eco- and a xenotropic virus. Fischinger PJS et al (1975) Proc Natl Acad Sci high pressure (performance) liquid 72 , 5150 chromatography (HPLC) A method for separating peptides, oligonucleotides, HK-PEG-1 (formerly PEG1-6) cells etc. with high resolution. (CCL 189) A hybridoma cell line, produced by fusing P3 63Ag8 myeloma Hinze virus Synonym for leporid herpes- cells with spleen cells from BALB/c virus 2. mice that had been immunized with influenza virus; secretes a monoclonal Hippotragine herpesvirus 1 (HiHV-1) A antibody (IgG3) that reacts with the species in the genus Rhadinovirus , iso- virus. lated from a roan antelope, Hippotragus HKU1 virus (HcoV-HKU1) A novel equinus . human coronavirus first isolated in Reid HW and Bridgen A (1991) Vet Microbiol Hong Kong university from a man 28 , 269 just returned from Shenzhen, China. Associated with upper respiratory tract Hirame rhabdovirus (HIRRV) A species infection and community-acquired in the genus Novirhabdovirus, first iso- pneumonia. Genome sequence analysis lated on Honshu Island, Japan in 1984, shows that it is a group 2 coronavirus. from flounder, Paralychthys olivaceus , with hemorrhagic disease. Rainbow Lau SK et al (2006) J Clin Microbiol 44 , 2063 Woo PC et al (2005) J Virol 79 , 884 trout fry can be infected experimentally. The virus is serologically unre- HL 23V virus A mammalian type C lated to most other fish rhabdoviruses. retrovirus from cultured human leuke- Kimura T et al (1986) Dis Aquat Org 1 , 209 mic cells (acute myelogenous). It is a mixture of one virus indistinguishable Hirt supernatant The supernatant from from the woolly monkey type C virus a virus-infected cell culture lyzed (SSAV-1) and another identical to an with sodium dodecyl sulfate (SDS). A endogenous virus of the baboon, Papio method of separating viral from cellu- cynocephalus, (BALV-M7). They are not lar DNA based on the preferential pre- endogenous human viruses, but are cipitation of undegraded cellular DNA probably horizontally transmitted. in the presence of SDS and sodium Similar isolates have been made from chloride final concentration 1 M. other leukemic patients. Hirt B (1967) J Mol Biol 26 , 365 Reitz MS et al (1976) Proc Natl Acad Sci 73 , 2113 histones Basic proteins rather more complex than protamines. They contain HL-60 cells (CCL 240) A promyelocytic tyrosine but little or no tryptophan. cell line, derived from the peripheral 213 horsepox virus

blood leukocytes of a 36-year-old homologous antiserum A serum contain- Caucasian female with acute promy- ing antibodies raised against a specific elocytic leukemia; produces subcutane- antigen and which will react with that ous myeloid tumors in nude mice. antigen.

HLA See human leukocyte antigens . homologous interference See interference . HLF-a cells (CCL 199) A fibroblast-like cell line derived from the periph- homology The degree of relatedness eral lung tissue of a 54-year-old black between the nucleotide sequences female with epidermoid carcinoma of of two nucleic acid molecules or the the lung. amino acid sequences of two protein molecules. Hybridization experiments HMPC See cidofovir . can produce useful information but, for critical analyses, sequence data are HMV-1 See El Moro Canyon virus . needed.

HMV-2 See Rio Segundo virus . Hong Kong virus Synonym for Influenza A virus isolated in Hong Kong in 1968, HN59; HN131; HN191; HN295 viruses which replaced the Asian strain influ- Chinese isolates of Banna virus, tenta- enza A (H2N2) and caused a global tive species in the genus Seadornavirus , pandemic of influenza A (H3N2) infec- family Reoviridae . tion. As of 2007, this is still the main influenza virus subtype circulating hog cholera virus (HCV) See Classical globally. swine fever virus . horizontal transmission Transmission HoJo virus (HOJOV) A strain of Hantaan between animals of any age after birth, virus in the genus Hantavirus , recov- usually excluding via the maternal ered from the blood of a patient with milk. Korean hemorrhagic fever. Antic D et al (1992) Virus Res 24 , 35 ‘ horse colds ’ An old name for equine influenza. homidium bromide 3,8-Diamino-5-ethyl- 6-phenyl-phenanthridinium bromide. horse papillomavirus Synonym for A trypanocide used in veterinary equine papillomavirus. medicine. Has two useful virological properties: (1) it binds to DNA and horsepox virus A possible species in the (2) it fluoresces in ultraviolet light. family Poxviridae . Once a common dis- Nucleic acids which, e.g., have been ease of horses the disease is now rare separated by gel electrophoresis can in the Western world. The complete thus be ‘ stained ’ by it and visualized. genome sequence of a virus isolated Circular forms of DNA such as in mito- in 1976 in Mongolia shows many simi- chondria bind the drug most strongly. larities to vaccinia virus, but with addi- Suppresses acute infection by retro- tional potentially ancestral sequences viruses, blocking integration of the that are absent in other Orthopoxviruses . viral genome. Also blocks induction of Causes papular lesions on lips, buc- mouse type C oncovirus by idoxyuri- cal mucosa and sometimes in the nose dine, and virus production by chroni- with fever and drooling of saliva. Only cally infected cells. Specific mode of a few deaths. Course 10–14 days or 3–4 action not known. weeks in severe cases. May be trans- Avery RJ and Levy JA (1979) Virology 95 , 277 missible to humans; finger lesions have been reported in those working homokaryon Hybrid cell formed by with horses. Not now present in the fusion of two cells of the same species. UK or the USA. Vaccinia virus will also See also heterokaryon . infect horses and in the early days of horsepox virus 214

vaccination Jenner himself suggested cytopathic virus by recombination with that grease of horses might be the ori- host cell transcripts. gin of variola virus in cows, and in Meyers G and Thiel H-J (1996) Adv Virus Res 1817 replaced ‘ vaccination ’ by ‘ equina- 47 , 53 tion ’ and supplied horse lymph to the National Vaccine Establishment. See hot spot A site within a gene at which also grease or grease-heel virus and mutations occur with unusually high Uasin Gishu disease virus . frequency. Synonyms: contagious pustular dermatitis of horses virus; contagious pustu- HR80-39 virus A strain of Seoul virus in lar stomatitis of horses virus. the genus Hantavirus, isolated from rats. Tulman ER et al (2006) J Virol 80 , 9244 Hs888Lu cells (CCL 211) A fibroblast-like horseradish peroxidase An enzyme cell line derived from the normal lung derived from the horseradish plant, tissue of a 20-year-old Caucasian male Armoracia rusticana . Used in ELISA with osteosarcoma metastatic to the tests to give the color reaction lung. with its substrate, e.g. brown with 3 ,3 -dianisidine. HSDM1C1 cells (CCL 148) A cell line derived from the parental HSDM1 host cell A cell in which a virus is line that secretes and synthesizes large replicating. quantities of prostaglandin E2 (PGE2). host initiation factor cleavage A mecha- Hsiung Kaplow herpesvirus Synonym nism by which certain viruses (mem- for caviid herpesvirus 1. bers of the Picornaviridae and the Pestivirus and Hepacivirus genera of HT-1080 cells (CCL 121) A cell line estab- the Flaviviridae ) initiate protein synthe- lished from a fibrosarcoma arising adja- sis without involving a 5 -cap struc- cent to the acetabulum of a 35-year-old ture. These viruses initiate translation Caucasian male. by direct association of the initiation complex with a highly conserved RNA HT virus A strain of H1 virus in the sequence within the 5 -untranslated genus Parvovirus. A rodent parvovirus region known as the internal ribosome of serological group 2. Antigenically entry site (IRES). This gives a trans- related to but not identical with H1 lational advantage to the virus over virus. Shows equal avidity for guinea host mRNAs, and in addition many pig and hamster erythrocytes, but does picornaviruses (aphthoviruses, entero- not react with those of humans or rats. viruses and rhinoviruses) also encode proteases that specifically cleave the HTDV Human teratocarcinoma-derived host initiation factor eIF4G which virus. See human endogenous prevents cap-dependent translation in retroviruses . the cell. HTLV-1 and HTLV-2 See human T-lymphotropic viruses type 1 and host range A listing of species of hosts type 2 . which are susceptible to a given virus (or other pathogen). HTLV III See human T-cell lymphotropic virus type III . host RNA polymerase II See DNA- dependent RNA polymerase . HTLV-associated adult T-cell leukemia (ATL) An aggressive malignancy host transcript recombination There is which develops in about 2% of persons evidence that some viruses such as the infected with HTLV-1. Characterized by pestivirus bovine viral diarrhea virus lymphadenopathy, hepatosplenomegaly, may evolve from a non-cytopathic to a hypercalcemia, and lytic bone lesions. 215 human adenoviruses 1-52 (HAdV-1 to 52)

HTLV-associated arthropathy (HAA) human adeno-associated virus See pri- An inflammatory disease affecting the mate adeno-associated viruses . joints due to infection with HTLV-1 virus. human adenoviruses 1-52 (HAdV-1 to 52) Serotypes which form seven species HTLV-associated myelopathy/tropical in the genus Mastadenovirus . There are spastic paraparesis (HAM/TSP) A 52 serotypes which can be divided into slowly progressive degenerative dis- three groups on the basis of agglutina- ease which develops in 2–3% of per- tion of rhesus monkey and rat eryth- sons infected with HTLV-1 virus. rocytes. Divided into seven species Affects the spinal cord, resulting in (A–G) on the basis of DNA genome weakness and spasticity in the lower homology. This results in the inclusion limbs along with sphincter and sensory of several monkey adenoviruses and dysfunction. one bovine adenovirus as strains or serotypes within the six human species HTLV-associated uveitis Inflammatory which contain the following serotypes eye disease consequent upon infection and associated symptoms: with HTLV-1 virus. A: Human serotypes 12, 18, and 31. Highly oncogenic in animals, cryp- Hu39694 A tentative species in the genus tic enteric infection in humans. Hantavirus . B: Human serotypes 3, 7, 11, 14, 16, 21, Levis S et al (1998) J Infect Dis 177 , 529 34, 35, 50, and simian adenovirus 21. Weakly oncogenic in animals, Huacho virus (HUAV) A serotype of respiratory and kidney infections Chenuda virus in the genus Orbivirus . in humans. Isolated from the tick, Ornithodoros C: Human serotypes 1, 2, 5, 6, and amblus , in Peru. Not reported to cause bovine adenovirus 9. Respiratory disease in humans. disease, latent in adenoids. D: Human serotypes 8, 9, 10, 13, 15, Brown SE et al (1989) Acta Virol 33 , 221 17, 19, 20, 22, 23, 24, 25, 26, 27, 28, 29, 30, 32, 33, 36, 37, 38, 39, 42, Hughes group viruses A group of tick- 43, 44, 45, 46, 47, 48, 49, and 51. borne viruses serologically related to Keratoconjunctivitis. Hughes virus in the genus Nairovirus . E: Human serotype 4, and simian All isolated from argasid ticks found adenoviruses 22, 23, 24, and 25. in areas frequented by sea birds. Their Conjunctivitis and respiratory exact taxonomic status is unknown: disease. Farallon F: Human serotypes 40 and 41, and Fraser Point simian adenovirus 19. Infantile Great Saltee diarrhea. Hughes virus G: Human serotype 52. Gastroenteritis. Puffin Island Punta Salinas Replication occurs in most primary and Raza continuous human cell cultures. There Sapphire II is an early CPE in heavily infected cul- Soldado tures due to penton antigen, and a late Zirqa CPE associated with virus replication. Some animal cells support produc- Hughes virus has also been isolated tive or abortive infection. With some from the blood of a sooty tern, Sterna types, multiplication is promoted by fuscata . concomitant infection with SV40. In mixed infections hybrid particles may Hughes virus (HUGV) A species in the be formed. Infection in humans may be genus Nairovirus . Isolated from sea silent, or respiratory disease of vary- birds in the Atlantic and Pacific and ing severity produced. Transmission in carried by Ornithodoros ticks. Not humans is mainly airborne but virus is reported to cause disease in humans. often present in stools and urine. The human adenoviruses 1-52 (HAdV-1 to 52) 216

strains present in feces may be difficult Lee TW and Kurtz JB (1994) Epidemiol Infect to cultivate in vitro although present in 112 , 187 large numbers. Types 43–49 were all Noel JS et al (1995) J Clin Microbiol 33 , 797 isolated from AIDS patients. A vaccine has been used with success, but the human astrovirus 1–8 (HAstV-1 to 8) oncogenicity of the viruses indicates Serotypes of human astrovirus in the caution in its use. genus Astrovirus . Synonyms: adenoid degeneration agent; adenoidal–pharyngeal–conjunctival human bocavirus A previously unrec- agent. ognized human parvovirus related by sequence to canine and bovine Jones MS et al (2007) J Virol 81 , 5978 parvoviruses in the genus Bocavirus . et al Virology 207 Kidd AH (1995) , 32 et al Russell W (2005) In Topley & Wilson’s Discovered by Allander in 2005 Microbiology and Microbial Infections , vol. 2, during examination of human respi- Tenth edition, edited by BWJ Mahy and V ratory secretions by treatment with ter Meulen. London: Hodder Arnold, p. 439 DNAse followed by restriction enzyme digestion and sequence-independent Human adenovirus A–G (HAdV-A single primer amplification of the frag- to G) Seven species in the genus ments. The virus has since been found Mastadenovirus . in many countries worldwide, in association with lower respiratory tract Human astrovirus (HAstV) The type disease and wheezing in children, but species of the genus Mamastrovirus . its significance as a human pathogen Astroviruses were first described dur- is still under investigation, since it has ing investigation of an outbreak of also been found in association with diarrhea in infants. Virions were origi- gastroenteritis. nally detected by electron microscopy Allander T et al (2005) Proc Natl Acad Sci 102 , as having a diameter of 28–30 nm, with 12891 no envelope and the appearance of a Allander T et al (2007) Clin Infect Dis 44 , 904 5- or 6-pointed star on the surface of about 10% of the virions. This feature human caliciviruses (HuCV) Members of suggested the name astrovirus. Virions the genera Norovirus and Sapovirus . A contain a single molecule of infectious, group of viruses originally called ‘ small positive-sense single-stranded RNA, round-structured viruses ’ (SRSV), 6.8–7.9 kb in length. The eight human including Norwalk virus (the proto- serotypes identified by immunofluores- type virus), strains of which are Desert cence and immunoelectron microscopy Shield, Hawaii, Lordsdale, Mexico, are serologically distinct from animal Snow Mountain, and Southampton. strains and share at least one common These viruses have a diameter of 27 nm epitope recognized by monoclonal and lack typical calicivirus appearance antibody. Serotype 1 is by far the most but genome sequence analysis shows common, accounting for 72% of isola- that they form a group within the tions. Grouping of strains by sequence genus Calicivirus. The Norwalk virus analysis corresponds to the serotype genome is positive single-stranded groups. Infection occurs worldwide, RNA, 7642 nt in length, containing three primarily in infants, but outbreaks open reading frames and a 3 -terminal in elderly and immunocompromised poly A tail. Cause periodic outbreaks patients have been described. By age of water- and food-borne gastroenteri- 10 years, there is 75% antibody preva- tis worldwide. Illness is usually mild lence. A monoclonal antibody-based and self-limited with symptoms lasting ELISA test has been developed for only 12–24 h after an incubation period detecting astrovirus in fecal specimens. of 48 h. However, persistent infection Carter MJ and Willcocks MM (2005) In has been documented with virus shed- Topley & Wilson’s Microbiology and Microbial ding for weeks to months following Infections , vol. 2, Tenth edition, edited by resolution of symptoms. Another group BWJ Mahy and V ter Meulen. London: of similar viruses are members of the Hodder Arnold, p. 888 Sapovirus genus, type species Sapporo 217 human coxsackie virus A7 (CAV7)

virus , including the strains Houston 86, Lai MM and Cavanagh D (1997) Adv Virus Houston 90, London, Manchester, and Res 38 , 1 Parkville. Sapoviruses have mainly Siddell SG et al (2005) In Topley & Wilson’s been associated with pediatric gastro- Microbiology and Microbial Infections , vol. 2, Tenth edition, edited by BWJ Mahy and V ter enteritis cases, but outbreaks and ill- Meulen. London: Hodder Arnold, p. 823 ness in older persons can occur. There Ziebuhr J and Siddell SG (1999) J Virol 73 , are no vaccines or antiviral therapies 177 available for the control of norovirus or sapovirus gastroenteritis. Treatment is Human coronavirus SARS See Severe aimed at preventing dehydration either acute respiratory syndrome corona- orally or through intravenous adminis- virus . tration of fluids. Berke T et al (1997) J Med Virol 52 , 419 human coxsackie viruses Species in Kapikian AZ (2000) J Infect Dis 181 , S295 the genus Enterovirus . Named after a Liu BL et al (1995) Arch Virol 140 , 1345 small town in New York State from O’Ryan ML et al (2000) J Infect Dis 182 , 1519 where the first virus was isolated. They Zheng DP et al (2006) Virology 346 , 12 are divided on biological characters, especially histopathological changes Human coronaviruses 229E, OC43, and induced following infection of suckling human enteric coronavirus (HCV- mice, into two groups: A and B. They 229E, HCV-OC43, HECoV) Species are responsible for a wide range of in the genus Coronavirus. Cause acute clinical manifestations, such as colds, respiratory disease (common colds) skin diseases, eye infections, and CNS and gastrointestinal disease (HACoV) involvement. In suckling mice, group A in humans mainly from January to viruses produce a flaccid paralysis due March. 229E virus was isolated in 1966 to an acute necrotic myositis; group B in human embryonic kidney cells from viruses produce a spastic paralysis due a medical student with a cold. OC43 to encephalitis. Serologically there are virus was isolated in 1967 in organ 23 viruses in group A, A1–22 and A24 culture from a patient with respira- (A23 is identical to echovirus 9), and 6 tory infection. Not always easy to iso- in group B. Newly recognized entero- late; human tracheal organ cultures are viruses (from type 68) are no longer probably the best method for primary assigned to the coxsackie virus or echo- isolations. The corona-like virus parti- virus species but are given the next cles seen in feces and associated with enterovirus number. Type A7 aggluti- diarrhea are difficult to isolate even in nates erythrocytes from fowls whose organ cultures. Strains have a common erythrocytes are agglutinated by vac- CF antigen but differences in antigenic cinia virus. Types A21 and B3 agglu- structure can be demonstrated by neu- tinate human O erythrocytes. Type B tralization tests and they belong to dif- viruses replicate readily in primary ferent coronavirus antigenic groups. monkey kidney cell cultures with CPE. OC43 is antigenically related to mouse Only a few A viruses will replicate in hepatitis virus and 229E is related to tissue culture. All coxsackie viruses coronaviruses of pigs (TGEV) and cats are pathogenic for suckling mice. Both (FECV). Some strains of OC43 virus types A and B are frequent, often silent, agglutinate human and monkey eryth- human infections, although A viruses rocytes at 4°C, and chicken, rat, and may cause herpangina and aseptic mouse erythrocytes at room tempera- meningitis, and B viruses epidemic ture or 37°C. Neuraminic acid recep- pleurodynia or myalgia (Bamble dis- tors are not involved. Can be adapted ease), orchitis, aseptic meningitis and to replicate in suckling mice and will myocarditis. kill them in 2–3 days following i.c. injection. Propagation in vitro is diffi- Hyypiä T and Stanway G (1993) Adv Virus Res 42 , 343 cult, but strains of 229E can be adapted Hyypiä T et al (1993) Virus Res 27 , 71 to growth in human diploid fibroblast cell lines. human coxsackie virus A7 (CAV7) A Gerna G et al (2006) J Med Virol 78 , 938 strain of Human enterovirus A . Causes human coxsackie virus A7 (CAV7) 218

paralysis on injection into monkeys, mice, others in cell cultures and cotton rats, and newborn mice. Has adapted to mice. been associated with outbreaks of aseptic meningitis with paralysis in human coxsackie virus A24 (CAV24) humans. Sometimes called poliomyeli- A strain of Human enterovirus C . tis virus type IV. Associated with several large out- Grist NR and Roberts GBS (1966) Arch breaks of hemorrhagic conjunctivitis in Gesamte Virusforsch 19 , 454 South-East Asia, and smaller outbreaks of conjunctivitis in other parts of the human coxsackie virus A9 (CAV9) world. Human enterovirus 70 has been A strain of Human enterovirus B . associated with similar outbreaks of Resembles the echoviruses in being conjunctivitis. inhibited by 2-(a-hydroxybenzyl) benzimidazole. Has caused many cases human coxsackie virus B1–6 (CBV1–6) of aseptic meningitis sometimes with Species in the genus Human entero- an exanthem. Has some antigenic virus B . Associated with a number of relationship to coxsackie A23-echo 9. human diseases including pleurodynia Replicates in rhesus monkey kidney (Bamble or Bornholm disease), menin- cell cultures with CPE and causes myo- gitis, respiratory illness, and especially carditis in mice. cardiopathy. The virus may affect the myocardium, endocardium, or peri- Chang KH et al (1992) J Gen Virol 73 , 621 cardium or all three, and evidence of infection is usually seen at autopsy. human coxsackie virus A10 (CAV10) Experimentally, cardiac disease can be A strain of Human enterovirus A . induced in monkeys with CBV 4, and Associated with a distinct syndrome, virulence is associated with a single lymphonodular pharyngitis. site in the 5 non-translated region of Steignian AJ et al (1962) J Pediatr 61 , 331 the genome. The possible association of coxsackie B viruses with type 1 insulin- human coxsackie virus A14 (CAV14) A dependent juvenile diabetes remains probable serotype or strain of Human conjectural, although a similar disease enterovirus A . More neurotropic than has been induced experimentally in other coxsackie A viruses on injection mice. into monkeys. Tracy S, Chapman NM and Mahy BWJ (Editors) (1997) Curr Top Microbiol Immun human coxsackie virus A16 (CAV16) A 223 , 311 pp strain of Human enterovirus A . The com- Tu Z et al (1995) J Virol 69 , 4607 monest cause of hand-foot-and mouth disease in humans. Shares an antigen human cytomegalovirus Synonym for with enterovirus 71 but there is no Human herpesvirus 5 . cross-neutralization; however, the complete sequence of CAV16 shows homol- human echoviruses 1–7, 9, 11–27, 29–33 ogy to HEV71. The first coxsackie virus (EV-1 to-7, 9, 11–27, 29–33) ( e nteric to be associated with hand-foot-and- c ytopathic h uman o rphan virus). mouth disease; CAV5 and CAV10 have Serotypes of human enterovirus B in also been associated with this disease. the genus Enterovirus. There are 33 Poyry T et al (1994) Virology 202 , 982 numbered serotypes, but serotype 8 was found to be identical to echo- human coxsackie virus A21 (CAV21) virus 1, serotypes 10 and 28 are now A strain of Human enterovirus C . reclassified as reovirus type 1 and Serologically identical to Coe virus, rhinovirus type 1 A, respectively, and associated with infantile diarrhea and serotypes 22 and 23 are reclassified common colds. as human parechoviruses 1 and 2. Frequently isolated from fecal speci- human coxsackie virus A23 (CAV23) mens in primary monkey kidney cell Identical to Human echovirus 9 . Some cultures but some strains replicate bet- strains may be isolated in suckling ter in human amnion cells. Originally 219 human echovirus 18 (EV18)

thought to be non-pathogenic, human echovirus 8 (EV8) Does not exist. they may rarely cause aseptic menin- A strain of human enterovirus B iso- gitis, encephalitis, respiratory disease, lated from patients with respiratory exanthem, gastrointestinal symptoms, and intestinal symptoms. Subsequently pericarditis, and myocarditis. Do not shown to be identical to Human echo- produce disease in suckling mice. virus 1. Newly recognized enteroviruses, from Rosen L et al (1958) Am J Hyg 67 , 300 type 68, are no longer assigned a coxsackie or echovirus number. Many strains of types 3, 6, 7, 10, 11, 12, 13, 19, human echovirus 9 (EV9) A serotype of 20, 21, 29, 30, and 33 agglutinate human human enterovirus B. Caused wide- O erythrocytes at 4°C. A strain of type spread epidemic aseptic meningitis 9 has been adapted to suckling mice in Europe in 1955 and 1956, and in and causes a disease similar to that America in 1957. There was often a produced by a coxsackie A virus. Focal maculopapular rash. Unlike the pro- lesions can occasionally be produced totype strain, many isolates can be by some strains on inoculation into the adapted to produce coxsackie A-type brain or spinal cord of monkeys. Types disease in newborn mice. Antigenically 4, 6, and 9 have been associated with identical to coxsackie A23 virus. outbreaks of aseptic meningitis. Types 11, 22, and 25 have been associated human echovirus 10 Does not exist. with respiratory illness. Type 18 was Reclassified as a strain of reovirus recovered from infants with diarrhea. type 1. Type l6 caused the ‘Boston Exanthem’ fever with a maculopapular rash. Data human echovirus 11 (EV11) A strain of on the history of various strains is Human enterovirus B . When it causes available in Strains of Human Viruses. disease it is usually in infants or young Synonym : echoviruses. children. Meningeal symptoms are Majer M and Plotkin SA (1972) Strains of most common, but fever, respiratory or Human Viruses . Basel: S Karger. gastrointestinal symptoms may occur. Sometimes associated with Bamble dis- human echovirus 4 (EV4) A strain of ease. Related by sequence homology to Human enterovirus B. Has caused out- human coxsackievirus A9. breaks of aseptic meningitis with gas- Synonym : U virus. trointestinal symptoms in about 70% of cases. Dahlund L et al (1995) Virus Res 35 , 215 Nagington J et al (1968) Lancet ii , 725 Chin TDY et al (1957) Am J Hyg 66 , 76 Johnsson T (1957) Lancet i , 590 human echovirus 16 (EV16) A strain human echovirus 6 (EV6) A strain of of Human enterovirus B. Caused the Human enterovirus B . Causes outbreaks of ‘ Boston Exanthem ’ : an outbreak of aseptic meningitis in children and adults. fever, aseptic meningitis, and a maculo- Gastrointestinal symptoms are uncom- papular rash which did not appear mon. Localized muscle weaknesses until the fever was over. and maculopapular rashes have been Neva FA and Enders IF (1954) J Immun 72 , observed in some outbreaks. Resembles 307 coxsackie virus in causing Bamble disease and carditis in humans, and in being pathogenic for suckling mice. human echovirus 18 (EV18) A strain of Human enterovirus B . Viruses isolated Kibrick S et al (1957) Ann NY Acad Sci 67 , 311 from two outbreaks of gastroenteritis and designated N5 and D-3 proved to human echovirus 7 A strain of Human be echo 18. Probably a cause of gastro- enterovirus B. Sequence of this virus has enteritis in infants. Infection often asso- been reported to occur in spinal cord of ciated with skin rashes. patients with ALS. Eichenwald HF (1958) J Am Med Assoc 166 , Berger MM et al (2000) Neurology 54 , 20 1563 human echovirus 19 (EV19) 220 human echovirus 19 (EV19) A strain of Illinois, USA from a nasal washing Human enterovirus B. Isolated originally from a young man with a mild respira- from an infant with diarrhea and later tory infection. from the CSF of an adult male patient. Has been associated with cases of human echovirus 34 A strain of Human mild respiratory disease and diarrhea coxsackie virus A24 in the genus mainly in children. Human enterovirus C . Cramblett HG et al (1962) Arch Int Med 110 , 574 human embryo kidney cells (HEK) Three common cell lines are available: human echovirus 20 (EV20) A strain of A 704 cells are derived from a human Human enterovirus B. Isolated from the kidney adenocarcinoma from a 78- stools of children with fever, coryza, year-old male; CAKI 2 cells are derived and diarrhea. In adult volunteers from a human kidney carcinoma of infection caused mainly constitutional a 69-year-old male; ACHN cells are symptoms, but some had ‘colds ’ or derived from a human renal adeno- gastrointestinal symptoms. carcinoma of a 22-year-old male. Synonym : JVI virus. Buckland FE et al (1961) BMJ i , 397 human embryo lung cells Non- transformed diploid cells with a finite human echovirus 21 (EV21) A strain life span, e.g. HEL 299 cells. of Human enterovirus B. Isolated in Hayflick L and Moorhead PS (1961) Exp Cell Massachusetts, USA from a child with Res 25 , 585 meningitis. Kibrick S (1964) Prog Med Virol 6 , 27 human endogenous retroviruses (HERV) Endogenous retroviruses can human echoviruses 22 and 23 (EV22 and be detected in the human genome by a 23) These viruses have been associated variety of procedures such as hybridi- with outbreaks of respiratory disease in zation or gene sequencing. They have a young children. They have some of the general structure similar to other retro- characteristics of coxsackie A viruses viruses including long terminal repeat which they resemble in their reaction (LTR) regions and sequences homolo- to 2-α -hydroxybenzyl benzimidazole gous to the gag , pol, and env genes of and the nuclear changes produced in infectious retroviruses. They can be infected cell cultures. Sequence anal- classified on the identity of the tRNA ysis suggests that they are distinct from homologous to the putative minus- prototype echoviruses and they have strand primer binding site (18 bp) been reclassified as human parecho- located immediately downstream of viruses 1 and 2 in a new genus, the 5 - LTR. On this basis, HERV-E Parechovirus . (glutamic acid), HERV-R (arginine), HERV-I (isoleucine), HERV-H (his- Stanway G et al (1994) J Virol 68 , 8232 tidine), HERV-P (proline), HERV-K (lysine), and HERV-W (tryptophan) human echovirus 25 (EV25) A strain of were distinguished. A further six new Human enterovirus B. Has been associ- HERV were detected using the human ated with rashes in infants. Infection genome project mapping database of human volunteers has caused fever, and partially characterized (all were pharyngitis, and cervical adenitis. defective, with partial deletions of the Kasel IA et al (1965) Proc Soc Exp Biol Med genome): HERV-f and HERV-f (type 118 , 381 B), primed by phenylalanine; HERV- R (type B), primed by arginine but human echovirus 28 Does not exist. Now different from the previously known reclassified as human rhinovirus strain HERV-R; HERV-S, primed by serine; 1 A. Isolated in a primary rhesus mon- and HERV-Z69907 and HERVH49C23, key cell culture at the Naval Medical where the 5 -LTR sequence could not Research Unit No. 4, Great Lakes, be detected. Others probably exist 221 human enteroviruses 76, 89, 90, 91 (HEV76, 89, 90, 91)

but have not yet been detected. Most enteroviruses. In some it produces no HERVs contain numerous mutations CPE but in others, such as RK 13 (rab- creating stop-codons and other inter- bit cells) and BK 1 (bovine cells), it pro- ruptions and so cannot be expressed. duces virus and CPE. Prototype strain However, HERV-K contains very few is AHC(J670/71). mutations and is expressed in PBMCs Melnick JL et al (1974) Intervirology 4 , 369 in the presence of HIV-1, and in some Yoshii T et al (1977) J Gen Virol 36 , 377 tumor cell-derived lines including GH cells (human teratocarcinoma cells). human enterovirus 71 (HEV71) A strain Particles of this endogenous human of Human enterovirus A in the genus retrovirus (termed HTDV for h uman Enterovirus . Isolated in 1970 from the t erato-carcinoma d erived v irus) have brain of a fatal case of encephalitis in been seen by thin-section electron California, USA. Related strains have microscopy of these cells. On average, been isolated from stools of sporadic 50 copies of HERV-K are present in the cases of meningitis and encephali- haploid human genome. tis in California and from outbreaks Blond JL et al (1999) J Virol 73 , 1175 in Australia and Sweden. Associated Tristem M (2000) J Virol 74 , 3715 with an epidemic of meningitis among Weiss RA (2006) Retrovirology 3 , 67 children in Bulgaria in 1975, in which Wilkinson DA et al (1994) In The Retroviridae , 21% of cases had paralysis. Has been vol. 3, edited by JA Levy. New York: Plenum isolated from cases of hand-foot-and- Press, p. 465 mouth disease in Japan, Malaysia and Taiwan. Prototype strain Br Cr. human enterovirus A–D (HEV-A to D) Shares an antigen with coxsackie A 16, Four species in the genus Enterovirus , but there is no cross-neutralization, containing 67 strains. although the sequences share considerable homology. human enterovirus 68 (HEV68) A strain of Human enterovirus D in the genus Hagiwara A et al (1978) Microbiol Immun 22 , 81 Enterovirus . Isolated from patients Hsiung GD and Wang JR (2000) J Microbiol with pneumonia and bronchiolitis in Immunol Infect 33 , 1 California, USA. Melnick JL et al (1979) Intervirology 12 , 297 Schieble JH et al (1967) Am J Epidemiol 85 , 297 human enterovirus 73 (HEV73) Name proposed for a group of enteroviruses human enterovirus 69 (HEV69) A strain from California and Oman which could of Human enterovirus B in the genus not be typed with standard enteroviral Enterovirus . Isolated in 1959 from a rec- typing antisera, but were similar by tal swab of a healthy 4-year-old child in sequence analysis and form a distinct Toluca, Mexico. Few strains have since cluster within the genus Enterovirus . been detected. Not reported to cause Oberste MS et al (2001) J Gen Virol 82 , 409 disease in humans. Prototype strain Toluca-1. human enterovirus 74 (HEV74) A strain Melnick JL et al (1974) Intervirology 4 , 369 of Human enterovirus B in the genus Enterovirus . human enterovirus 70 (HEV70) A strain Oberste MS et al (2004) J Gen Virol 85 , 3205 of Human enterovirus D in the genus Enterovirus. Isolated in 1971 from epi- human enterovirus 75 (HEV75) A strain demics of acute hemorrhagic conjuncti- of Human enterovirus B in the genus vitis in Japan, Singapore and Morocco. Enterovirus . These outbreaks were part of a pandemic involving millions of humans in human enteroviruses 76, 89, 90, 91 Africa, South-East Asia, Japan, India, (HEV76, 89, 90, 91) Strains of Human and England during 1969–1971. The enterovirus A in the genus Enterovirus , virus can adsorb in vitro to cells from isolated from patients in France and a wider range of species than most Bangladesh. human enteroviruses 76, 89, 90, 91 (HEV76, 89, 90, 91) 222

Oberste MS et al (2005) J Gen Virol 86 , 445 Human herpesvirus 1 (HHV-1) Type species of the genus Simplexvirus in human enterovirus 77 (HEV77) A strain the subfamily Alphaherpesvirinae . The of Human enterovirus B in the genus genome DNA has been completely Enterovirus , recovered from a refugee sequenced for the 17syn strain, and child from Kosovo. consists of about 150 kb with a G C Bailly J-L et al (2004) Virus Res 99 , 147 of 67%. The DNA is infectious and has two components, L and S, each human enterovirus 78 (HEV78) A strain of which is bracketed by internal of Human enterovirus B in the genus repeats. Primary infection is common Enterovirus . in young children, often subclinical, but occasionally with acute stomatitis. Norder H et al (2003) J Gen Virol 84 , 827 The virus can pass along nerves and become latent in ganglia from whence human enterovirus 94 (HEV94) A strain it can be reactivated by nonspecific of Human enterovirus D in the genus stimuli (fever, sunlight, menstrua- Enterovirus . tion) to cause lesions, often around Isolated from sewage in Egypt and the mouth. Rarely, the virus may cause a patient in the Democratic Republic acute hepatitis, kerato-conjunctivitis, or of the Congo. The complete coding meningo-encephalitis. Vaccination has sequence of the clinical isolate had 70% not been successful but treatment of sequence similarity to EV70. kerato-conjunctivitis and skin lesions Smura TP et al (2007) J Gen Virol 88 , 849 with locally applied acyclovir ointment is beneficial. In cases of encephalitis, Human foamy virus (HFV) Not a neonatal herpes, or disseminated infec- human virus. A species in the genus tion, intravenous acyclovir is used. See Spumavirus , now called Simian foamy Human herpesvirus 2 . virus. Isolated from a human nasopha- Synonyms: herpes febrilis; herpes sim- ryngeal carcinoma explant culture. Can plex virus 1; herpesvirus hominis; be propagated in a variety of human Human herpesvirus 1 . and animal cell cultures but not in chick embryo cell culture. PCR tests Feldman LT (1994) Semin Virol 5 , 207 McGeoch DJ et al (2000) J Virol 74 , 10401 failed to detect any viral sequences Minson AC (2005) In Topley & Wilson’s in samples of 223 patients, or foamy Microbiology and Microbial Infections , vol. 2, virus-specific antibodies in 2688 sera Tenth edition, edited by BWJ Mahy and V ter from suspected high-risk persons. Meulen, London: Hodder Arnold, p. 506 Previous reports may have resulted from contamination with simian foamy Human herpesvirus 2 (HHV-2) A spe- virus. Persistent zoonotic infection of cies in the genus Simplexvirus . Genome a human with simian foamy virus has DNA is related to HSV1 with about been reported. In a study of 187 work- 85% homology in the open reading ers exposed to nonhuman primates frames. Differs from human herpes- at research facilities or zoos, 10 (5.3%) virus 1 in that it is usually, though tested positive for simian foamy virus not always, transmitted venereally. but none tested positive for other Infection is therefore uncommon before simian retroviruses (simian immuno- the age of puberty. The virus usu- deficiency virus, simian type D retrovirus ally causes genital lesions but can also or simian T-cell lymphotropic virus). be responsible for any of the lesions Synonyms : human spumavirus; human characteristic of human herpesvirus 1. syncytial virus. There is a high level of antigenic simi- Callahan ME et al (1999) J Virol 73 , 9619 larity between viruses 1 and 2 although Schweizer M et al (1995) AIDS Res Human each has antigens specific to itself. Retro 11 , 161 Glycoprotein G-specific assays provide Switzer WM et al (2004) J Virol 78 , 2780 the best differentiation. Acyclovir is an Switzer WM et al (2005) Nature 434 , 376 effective treatment. Synonyms: herpes febrilis; herpes sim- human hepatitis A virus (HHAV) See plex type 2; herpesvirus hominis; Hepatitis A virus . Human herpesvirus 2 . 223 Human herpesvirus 5 (HHV-5)

Herbst-Kravovetz M and Pyles R (2005) 172 kb in length with a G C of 60%. Herpes 13 , 37 First isolated from Burkitt tumors of Stanberry LR (1993) Rev Med Virol 3 , 37 African children. A very widespread Wald A et al (2006) J Infect Dis 194 , 42 human infection, mainly of children, in whom it rarely causes disease but Human herpesvirus 3 (HHV-3) Type spe- produces a high level of immunity. cies of the genus Varicellovirus , in the However, primary infection of young subfamily Alphaherpesvirinae . The adults may result in infectious mono- genome DNA has been completely nucleosis, a febrile condition with sequenced for the Dumas strain, and is enlargement of the lymph nodes and 125 kb in length, with a G C of 46%. often a sore throat. The Paul–Bunnell It consists of an L and an S component test is positive. The virus is the prob- bounded by repeats. The S compo- able cause of Burkitt tumors and carci- nent can be inverted to form two iso- noma of the post-nasal space, but as the mers, and both are present in packaged infection is universal and these tumors genome DNA. The cause of common only occur in local areas, some acces- human infection. Causes chickenpox on sory factors must be involved. After primary infection, usually in childhood. infection, the virus remains in the body Incubation period 1–16 days, rarely up as a latent infection of B lymphocytes. to 21 days. It then remains latent, and It can be propagated in human cell may reactivate many years later to cause cultures, but is cell-associated and is herpes zoster, a painful local condition difficult to purify. A number of virus- with skin lesions, usually in adults. The coded antigens have been identified in eyes may be involved. May follow expo- infected tissues, and antibodies to these sure to infection but most commonly are found in humans. They include appears as a reactivation of latent infec- the capsid antigen (VCA), antibodies tion. Encephalitis is a rare complication to which are a good indication of pri- of chickenpox. Fetal malformations mary infection, and a nuclear antigen have been reported to follow maternal (EBNA) which appears 4–6 weeks after infection. All strains are antigenically disease onset. similar. Either acyclovir or famciclovir Synonyms : Burkitt’s lymphoma virus; are effective in treatment. Convalescent Ebb virus; Epstein–Barr virus; glandular serum has no therapeutic use but an fever virus; infectious mononucleosis attenuated vaccine developed in Japan virus; human (gamma) herpesvirus 4. (the Oka strain) is now licensed in many countries for use in children. Animals Epstein MA and Crawford DH (2005) In Topley & Wilson’s Microbiology and Microbial appear to be resistant to infection. The Infections , vol.1, Tenth edition, edited by virus can be cultivated in HeLa and BWJ Mahy and V ter Meulen, London: various monkey tissue cells, with CPE Hodder Arnold, p. 559 in 2–7 days. It remains cell-bound and Lee SP (1994) Semin Virol 5 , 281 difficult to separate, and is usually pas- Middleton T et al (1991) Adv Virus Res 40 , 19 saged with cellular material. Sugden B (1994) Semin Virol 5 , 197 Synonyms: chickenpox virus; herpesvirus varicellae; varicella-zoster virus. Human herpesvirus 5 (HHV-5) Type species of the genus Cytomegalovirus , sub- Arvin AM (1996) Clin Microbiol Revs 9 , 361 family Betaherpesvirinae. The genome Chee MS et al (1990) Curr Top Microbiol Immunol 154 , 125 DNA has been completely sequenced Hay J and Ruyechan WT (1994) Semin Virol for the AD169 strain, and is 230 kb in 5 , 241 length with a G C of 57%. A world- Loparev VN et al (2007) J Infec Dis 195 , 502 wide infection, common in humans. Simpson D and Lyseng-Williamson KA Usually a chronic silent infection of (2006) Drugs 66 , 2397 peripheral blood monocytes and cells of the salivary glands, but primary Human herpesvirus 4 (HHV-4) Type spe- infection during pregnancy may lead cies of the genus Lymphocryptovirus , to infection of the fetus with varying subfamily Gammaherpesvirinae . The degrees of brain damage. Infection genome DNA has been completely in utero or neonatal infection may sequenced for the B95-8 strain, and is cause severe and often fatal hepatitis, Human herpesvirus 5 (HHV-5) 224

splenomegaly and anemia. Patients commonly infects neuronal cells in with AIDS or on immunosuppres- the brain and has been suggested as a sive therapy and, occasionally, healthy possible causative agent in multiple people may develop fatal pneumonia, sclerosis. There are two closely related hepatitis, or a form of glandular fever. subspecies, HHV-6A and HHV-6B, that Virus can often be isolated from the differ in growth properties, antigenic- urine of patients, and sometimes from ity, nucleotide sequence and epidemio- that of healthy children. An active vac- logical characteristics. HHV-6A is the cine (Towne 125 strain) has been made, virus associated with multiple sclero- is safe and will probably come into use. sis. HHV-6B is strongly associated with The virus is much less susceptible than roseola infantum and is the virus that other human herpesviruses to anti- has been found in the CNS. The virus viral drugs such as acyclovir, but the may persist by integration in a host derivative ganciclovir, administered chromosome, and could be detected in parenterally, is effective and is licensed blood in high copy number ( 6 log(10) to treat retinitis in the immunosup- copies/ml) in about 0.8% of blood pressed, especially patients with AIDS. donors in the UK in one study. There is replication in primary human Akhyani N et al (2000) J Infect Dis 182 , 1321 fibroblast cultures with a slowly devel- Gompels UA et al (1995) Virology 209 , 29 oping CPE and formation of inclusion Leong HN et al (2007) J Med Virol 79 , 45 bodies. Hamster and human cells are Miyagawa HH and Yamanishi, K (1999) transformed in vitro . The virus is cell- Curr Opin Infect Dis 12 , 251 associated. No hemagglutinins. Sen- Pellett PE et al (1992) Adv Virus Res 41 , 1 sitive to freezing and thawing and to Zerr DM (2006) Herpes 13 , 20 prolonged storage at –70°C. Synonyms : giant cell pneumonia virus; Human herpesvirus 7 (HHV-7) A spe- human cytomegalovirus; inclusion cies in the genus Roseolovirus . First body disease virus; visceral disease described in 1990 as an infection in virus. activated human T cells. A ubiquitous virus which is shed in the saliva of 75% Britt W (2005) In Topley & Wilson’s of adults. Seroprevalence appears to be Microbiology and Microbial Infections , vol. 1, at least 90% in the normal population Tenth edition, edited by BWJ Mahy and V with infection beginning by 6 months ter Meulen. London: Hodder Arnold, p. 520 of age and widely prevalent by age 3 Chee MS et al (1990) Curr Top Microbiol Immunol 154 , 125 years. More closely related antigeni- Sinclair J and Sissons JGR (1994) Semin Virol cally to HHV-6A and HHV-6B than to 5 , 249 other human herpesviruses. It appears that the two viruses may interact during reactivation from latency. Remains Human herpesvirus 6 (HHV-6) Type species of the genus Roseolovirus in the as a latent infection in CD4( ) T lym- subfamily Betaherpesvirinae. Genome is phocytes. Significance and possible role 160 kb in length, with a base composi- in human disease is presently unclear. tion of 43% G C, and contains 119 The virus replicates in skin lesions of open reading frames. HHV-6 is one lichen planus, but not in psoriasis, but of the most widespread of the herpes- whether it is involved in the pathogen- viruses, and seroprevalence rates exceed esis of lichen planus is unknown. 90% by 2 years of age. Etiologically Black JB and Pellett PE (1993) Rev Med Virol associated with exanthem subitum 3 , 217 (roseola infantum), a common, usu- De Vries HJ et al (2006) Br J Dermatol 154 , ally benign, childhood disease, rarely 361 associated with encephalopathy. There Hall CB et al (2006) J Infect Dis 193 , 1063 Mega WAG et al (1998) Virology 244 , 119 is abrupt onset of fever lasting 3 days Miyake F et al (2006) J Med Virol 78 , 112 then ceasing as a rash appears. HHV-6 Ward KN (2005) Curr Opin Infect Dis 18 , 247 has also been implicated in some cases Wyatt LS et al (1991) J Virol 65 , 6260 of heterophile-negative mononucleosis, lymphadenopathy and extended Human herpesvirus 8 (HHV-8) A species low-grade fever and malaise. The virus in the genus Rhadinovirus . Identified 225 Human immunodeficiency virus type 1 (HIV-1)

by sequence representational differ- DNA provirus requires cellular tran- ence analysis in tissues of patients scriptional activators such as TNFα and with Kaposi’s sarcoma. DNA and NFκB. Several virus-specified proteins monoclonal antibody studies indicate are involved in transcription, including that the virus latently infects cells of Tat, which binds to an element within Kaposi’s sarcoma (KS), multicentric the LTR region of the genome known Castleman’s disease, and the rare pri- as TAR. Expression of different genes mary effusion lymphoma, and is caus- is determined by differential splicing ally associated with these tumors. of the full length genome mRNA tran- Synonym: Kaposi’s sarcoma herpes- script. Splicing is in turn regulated virus. by the Rev protein, which binds to a region of the mRNA termed the Rev Cesarman E and Mesri EA (2007) Curr Top Microbiol Immunol 312 , 263 responsive element (RRE), and inhibits Dezube BJ et al (2006) J Cell Physiol 209 , 659 splicing of the RNA. Early transcripts Dupin N et al (1999) Proc Natl Acad Sci 96 , of the genome are multiply spliced, but 4546 as the amount of Rev accumulates, sin- Ganem DC (1997) Cell 91 , 157 gle spliced and unspliced transcripts McGeoch DJ and Davison AJ (1999) Semin accumulate, and this results in capsid Cancer Biol 9 , 201 protein synthesis. The major groups of Moore P et al (1996) J Virol 70 , 549 proteins that are synthesized in HIV- Sharp TV and Boshoff C (2000) IUBMB Life infected cells are the gag , pol , and env 49 , 97 gene products. Gag proteins include the matrix (p18), capsid (p24) and Human immunodeficiency virus type nucleocapsid (p15) proteins of the vir- 1 (HIV-1) A species in the genus ion; pol proteins include the protease Lentivirus , which is the primary etio- (p10), reverse transcriptase (p51/66), logic agent of AIDS, a fatal disease first and integrase (p34) proteins; and env recognized in 1981 that results from proteins include the surface envelope gradual destruction of the helper T-cell glycoprotein (gp120) which binds to population in infected individuals. The host cell receptors, and a transmem- prototype strain is LAV LA1. Mature brane protein (gp41). In addition, a virions are spherical, 100–120 nm in number of minor proteins are synthe- diameter, with a dense cone-shaped sized from spliced mRNA, including core, comprised of p24 capsid pro- a myristylated membrane-associated teins, surrounded by an envelope nonstructural protein termed Nef that with spike projections. The genome is is not required for replication in cell diploid, consisting of two molecules cultures but plays a role in HIV patho- per virion of positive single-stranded genesis in vivo . RNA 9800 nt in length, linked near HIV-1 infects only humans or chim- their 5 -ends by non-covalent bonds. panzees, although chimpanzees do A host-cell-derived lysine tRNA is not develop disease or evidence of the primer for reverse transcription of immunodeficiency. In humans, the the genome RNA by the action of the virus spreads by sexual transmis- virion reverse transcriptase (p51/66) sion, blood transfusion, perinatal into linear double-stranded DNA, transmission, and intravenous drug which migrates into the nucleus and injection using unsterilized needles. becomes integrated into the host cell There is considerable variation in the genome through the action of a viral clinical outcome of HIV-1 infection, genome-encoded integrase (p34). The which depends upon genetic varia- integrated DNA provirus contains long tion of the human genome. However terminal repeat (LTR) elements at each most infected individuals develop end, as well as three structural genes a humoral immune response to the (gag , pol , and env ) needed for replica- gp120 Env and p24 Gag proteins, tion. Additional genes involved in reg- and the latter is frequently used as a ulation of synthesis and processing of diagnostic indicator of infection. The the virus include vif , vpr , vpu , tat , rev , USA Centers for Disease Control and and nef. Transcription of the integrated Prevention has classified HIV infection Human immunodeficiency virus type 1 (HIV-1) 226

into four stages. In stage 1, acute infec- and disease severity can be correlated tion, there is usually a febrile mono- with the titer of HIV in the periph- nucleosis or influenza-like syndrome eral blood (viral load), but the factors 3–4 weeks after exposure, which lasts which influence this are not yet clearly 2–3 weeks before clinical recovery understood. Drugs which inhibit viral occurs. Stage 2, asymptomatic infec- replication and so decrease viral load tion, usually lasts 7–9 years, but can be (such as zidovudine, AZT and the pro- shorter. During this period, the health tease inhibitors, Indinavir, Nelfinavir, of the affected individual can be moni- Ritonavir, and Saquinavir) prolong tored by determining the numbers of survival of HIV-infected individuals CD4 cells in peripheral blood. Many (see HAART ). Several genomic clades patients progress to stage 3 (persistent, of HIV-1 are recognized (Table H2). generalized lymphadenopathy) dur- Doms RW (2000) Virology 276 , 229 ing otherwise asymptomatic infection. Fellay, J et al (2007) Sciencexpress 19 July Finally, stage 4 disease (symptomatic Hahn BH et al (2000) Science 287 , 607 HIV infection) usually develops after Kolson DL et al (1998) Adv Virus Res 50 , 1 several years. Symptoms may include Korber B et al (2000) Science 288 , 1789 weight loss, diarrhea, encephalopathy, Popik W and Pitha PM (2000) Virology 276 , 1 and vision impairment, as well as a Simon F et al (1998) Nature Med 4 , 1032 profound decrease in numbers of CD4 Schubert U and McClure M (2005) In helper T cells to less than 200 cells per Topley & Wilson’s Microbiology and Microbial microliter of blood. Other manifesta- Infections , vol. 2, Tenth edition, edited by BWJ Mahy and V ter Meulen, London: tions are thrombocytopenic purpura, Hodder Arnold, p. 1322 anemia, dermatitis, and peripheral neuropathies. A number of opportun- Human immunodeficiency virus type istic infections and conditions, includ- 2 (HIV-2) A species in the genus ing Pneumocystis carinii pneumonia, Lentivirus, related to HIV-1 about Kaposi’s sarcoma, mucosal candidia- 40% by RNA sequence homology; the sis, cytomegalovirus infection, and genome structure is similar except herpes simplex ulcers, may develop that the vpu gene is replaced by a vpx at this stage. Once the diagnosis of gene. The virus was identified serolog- AIDS is confirmed, survival is usually ically in West Africa and subsequently less than 1 year. Progression to AIDS isolated and found to be more closely

Table H2. Examples of genomic clades of HIV

HIV virus type Clade

Human immunodeficiency virus type 1 90CR056 (HIV-1.90CR056) Clade H Human immunodeficiency virus type 1 93BR020 (HIV-1.93BR020) Clade F Human immunodeficiency virus type 1 ANT70 (HIV-1.ANT70) Clade O Human immunodeficiency virus type 1 ARV-2/SF-2 (HIV-1.ARV-2/SF-2) Clade B Human immunodeficiency virus type 1 BRU (LAI) (HIV-1.BRU(LAI)) Clade B Human immunodeficiency virus type 1 ELI (HIV-1.ELI) Clade D Human immunodeficiency virus type 1 ETH2220 (HIV-1.ETH2220) Clade C Human immunodeficiency virus type 1 HXB2 (HIV-1.HXB2) Clade B Human immunodeficiency virus type 1 MN (HIV-1.MN) Clade B Human immunodeficiency virus type 1 NDK (HIV-1.NDK) Clade D Human immunodeficiency virus type 1 RF (HIV-1.RF) Clade B Human immunodeficiency virus type 1 U455 (HIV-1.U455) Clade A Human immunodeficiency virus type 2 BEN (HIV-2.BEN) Clade A Human immunodeficiency virus type 2 D205 (HIV-2.D205) Clade B Human immunodeficiency virus type 2 EHOA (HIV-2.EHOA) Clade B Human immunodeficiency virus type 2 ISY (HIV-2.ISY) Clade A Human immunodeficiency virus type 2 ROD (HIV-2.ROD) Clade A Human immunodeficiency virus type 2 ST (HIV-2.ST) Clade A Human immunodeficiency virus type 2 UC1 (HIV-2.UC1) Clade B 227 Human papillomavirus (HPV-1 to 96)

related to Simian immunodeficiency certain sites on the genome, resulting in virus (SIV) than to HIV-1. The descrip- frameshift mutations, especially in the tive pathogenesis of HIV-2 infection is SH protein. A live-attenuated virus is similar to HIV-1, involving immuno- under development to prevent severe deficiency and neurological diseases, disease from infection in infancy. In although HIV-2 appears to be less addition, since monoclonal antibodies virulent than HIV-1. Several genomic to the F protein have been used thera- clades of HIV-2 virus are recognized peutically against RSV infection, such (see Table H2 ). an approach is under development for use against HMPV infection. human leukocyte antigens (HLA) Major Biacchesi S et al (2007) J Virol 81 , 6057 histocompatibility antigens of the Galiano M et al (2006) J Med Virol 78 , 631 human species. Occur as alloantigens on Kahn JS (2006) Clin Microbiol Rev 19 , 546 the cell surface in two classes. Humans Manoha C et al (2007) J Clin Virol 38 , 221 inherit and express different combina- Miller SA et al (2007) J Virol 81 , 141 tions of class I and class II alleles, and O’Gorman C et al (2006) Eur J Clin Microbiol Infect Dis 25 , 190 the combination of class I and class II Sloots TP et al (2006) Emerg Infect Dis 12 , allotypes expressed by a person is called 1263 the HLA type. The genes that encode Ulbrandt ND et al (2006) J Virol 80 , 7799 HLA class I and class I alloantigens Warris A and de Groot R (2006) Adv Exp Med are closely linked on the short arm of Biol 582 , 251 human chromosome 6 in a region called Wolf DG et al (2006) Pediatr Infect Dis J 25 , the HLA complex, encompassing about 320 4 million bp of DNA. See major histocompatibility complex . human microvascular endothelial cell line (HMEC-1) A cell line that has Marsh SS et al (2000) The HLA Facts Book . proved useful for the propagation of London: Academic Press filoviruses, such as Marburg and Ebola virus. human metapneumovirus (HMPV) A Schnittler HJ et al (1993) J Clin Invest 91 , 1301 species in the genus Metapneumovirus . First described in 2001, and now rec- human monkeypox See Monkeypox ognized as an important cause of res- virus . piratory disease worldwide, especially in infants. Serological surveys suggest Human papillomavirus (HPV-1 to 96) A that this virus has been present in the species in the family Papillomaviridae . human population for at least the last There are 96 genotypes currently recog- 50 years. In large surveys the reported nized, each differing by more than 10% percentage of children with respiratory in genome sequence from other types disease associated with HMPV was (see below). They are classified into five 15% in Austria, 7% in Australia, 13% in genera, Alphapapillomavirus (type spe- Israel, and 10% in Sweden. The disease cies HPV 32), Betapapillomavirus (type is similar to that caused by RSV. The species HPV 5), Gammapapillomavirus virus is found in all age groups, and in (type species HPV 4), Mupapillomavirus high risk (e.g. immunocompromised) (type species HPV 1), Nupapillomavirus adults, has been reported to cause (type species HPV 41). Cause papillo- death. The sequence diversity between mas of the epidermis in humans: skin avian and human isolates is less than warts (verrucae of various types), geni- between certain avian isolates, though tal warts (condylomata acuminata), the viruses are host-specific. However and laryngeal papillomas. Malignant turkey poults can be infected by HMPV change is reported in genital and inoculated oculonasally. Two main laryngeal papillomas. The exact role genotypes of HMPV are recognized, A of the virus in carcinoma of the cer- and B, and each can be subdivided into vix and other tumors remains to be four lineages A1, A2, B1, and B2. It has elucidated. Virus is readily extracted been reported that during passage in from skin warts but less easily demon- vitro frequent point mutations occur at strated in genital and laryngeal lesions. Human papillomavirus (HPV-1 to 96) 228

Accidental or experimental inoculation Stanley M (2007) Brit J Cancer 96 , 1320 of virus into the skin of humans results Pollack AE et al (2007) Bull World Health in the development of warts. Does not Organ 85 , 57 replicate or produce papillomas on zur Hausen H (2002) Nat Rev Cancer 2 , 342 injection into animals. Virus replica- human papovavirus This term includes tion in cell culture is extremely difficult human papillomavirus and human to achieve and has only been possible polyomavirus. in ‘ raft ’ cultures of human epithelial cells. When DNA from virus extracted Human parainfluenza virus type 1 from individual warts is cloned and (HPIV1) A species in the genus sequenced, more than 90 different HPV Respirovirus . Antibodies are common types are revealed. Classification as a in humans, and in monkeys who have new genotype requires more than 10% been in contact with humans. Causes dissimilarity in the combined nucleo- acute laryngotracheitis (croup) in tide sequences of the E6, E7 and L1 young children and occasionally mild genes when compared with those of upper respiratory tract infections in any previously known type. Viruses older children and adults. There may with greater than 90% but less than be more than one type of the virus. It 100% sequence homology are classified is best isolated in primary human or as subtypes. Using this classification, monkey kidney cell cultures or diploid association of specific types with dis- human cell lines, such as NC1H292 eases can be observed as shown in Table cells. Replication is poor in eggs. CPE H3 . A vaccine based on alum adjuvan- may be slight at first but becomes ted non-infectious virus-like particles, more marked on passage. Syncytia are containing the L1 protein expressed formed which float off leaving holes in in yeast, has been licensed for use in the cell sheet. Hemadsorption is bet- prepubertal girls in many countries to ter with chick or guinea pig red cells reduce the incidence of cervical can- rather than human cells, but may not cer in women. The vaccine (Gardasil) be demonstrable until day 10 on pri- is quadrivalent, containing HPV mary isolation. Hamsters and guinea types 6, 11, 16 and 18. The vaccines pigs can be infected with the virus but induce high levels of neutralizing anti- no pathological changes are produced. body for at least 5 years. Synonyms: acute laryngo-tracheo-bron- Synonym : human wart virus. chitis virus; croup-associated virus; Chan S-Y et al (1995) J Virol 69 , 3074 hemadsorption virus 2; influenza virus Delius H and Hofmann B (1994) Curr Top D; newborn pneumonitis virus. Microbiol Immun 186 , 13 Leggatt GR and Frazer IH (2007) Curr Opin Human parainfluenza virus type 2 Immunol 19 , 232 (HPIV2) A species in the genus

Table H3. Specific types of human papillomaviruses and associated diseases

HPV types HPV-associated diseases

1, 2, 3, 4, 7, 10, 26–29, 41, 48, 60, 63, 65, 75–78 Common warts (benign) 5, 8, 9, 12, 14, 15, 17, 19–25, 36–38, 46, 47, 49, 50 Epidermodysplasia verucciformis (EV), malignant 6, 11, 42–44, 54–61, 64, 66–68, 70, 74, 79, 84 Anogenital condylomata (rarely malignant) 30, 31, 33, 35, 45, 52, 56, 58, 69, 83 Anogenital condylomata (intermediately malignant) 16, 18, 31, 33, 35, 39, 45, 51, 52, 56, 58 Anogenital condylomata (highly malignant) 6, 11 Laryngeal papilloma 16, 33 Tonsillar carcinoma 13, 32 Oral focal epithelial hyperplasia 229 human polioviruses (PV-1 to 3)

Rubulavirus. Causes croup in children Ljungan virus, found in rodents in under 5 years of age. Induces syncytia Scandinavia, shares many molecular formation in cell cultures. Epidemics features of the parechoviruses. occur in the fall, usually every other Niklasson B et al (1999) Virology 255 , 86 year, in susceptible infants. Stanway G et al (2000) Rev Med Virol 10 , 57 Synonyms: acute laryngo-tracheobronchitis virus; croup-associated virus. human parvovirus For many years the only recognized human parvovirus Human parainfluenza virus type 3 was B19 virus, the cause of erythema (HPIV3) A species in the genus infectiosum in young children and of Respirovirus . Isolation is most efficient aplastic crisis in children and adults in primary human or monkey kidney with chronic hemolytic anemia. Then cell cultures but the virus will replicate in 2005 Allander et al applied a new in cell lines such as HeLa, Hep-2 and method for virus detection in human NC1-H292. CPE on isolation may be samples which involved Dnase treat- minimal and hemadsorption is used. ment followed by sequence inde- On passage the cell sheet is disrupted, pendent single primer amplification the cells becoming long and narrow. (SISPA). This method detected a new Causes pharyngitis, bronchiolitis and human parvovirus which was related pneumonia in young children, espe- by sequence to the bovine and canine cially in nursery schools. Uncommon parvoviruses, in the Bocavirus genus, cause of ‘ colds ’ in adults. Inoculation and so was named human bocavirus. into young hamsters causes inapparent This virus appears to be widespread infection or small lung lesions. geographically, although it’s asso- Synonym : hemadsorption virus 1. ciation with disease remains unclear. Also in 2005, a San Francisco study of Human parainfluenza virus type 4 25 patients with acute viral infection (HPIV4a and HPIV-4b) Species in syndrome following high risk behav- the genus Rubulavirus. More difficult ior for HIV-1 transmission, using the to isolate than HPIV-1 to -3. Isolation SISPA technique identified three novel best in primary monkey kidney cell viruses, two related to TT virus, and cultures but hemadsorption may a new species of human parvovirus not be demonstrable for 3–4 weeks. named PARV4, which is unrelated by Agglutinates guinea pig and rhesus sequence to any other known mamma- monkey cells better than human cells. lian parvoviruses. PARV4 was found in CPE poor in early passages. Common samples from patients born after 1958, infection of young children causing but in older subjects (born between mild upper respiratory disease but 1949 and 1956) a variant genotype, the virus is not easily isolated. No evi- PARV5, was found. To date, PARV4 dence of virus replication in eggs. Two infection has been detected mainly in antigenic types, a and b, recognized injecting drug users and hemophiliacs, by hemagglutination inhibition tests. suggesting a parenteral route for trans- Not pathogenic for laboratory animals mission of this virus. See B19 virus, but produce high antibody levels in Human bocavirus, PARV4 virus . guinea pigs. Fryer JF et al (2007) Transfusion 47 , 1054 Jones MS et al (2005) J Virol 79 , 8230 Human parechovirus (HPeV) Type spe- Manning A et al (2007) J Infect Dis 195 , 1345 cies of the genus Parechovirus , family Picornaviridae . There are three recog- human polioviruses (PV-1 to 3) Serotypes nized serotypes, human parechovirus 1 of Poliovirus in the genus Enterovirus . (formerly called echovirus 22), human There are three serological types: 1, 2 parechovirus 2 (formerly called echo- and 3. The genome of the Mahoney virus 23), and human parechovirus 3. strain of type 1 is 7433 nt in length, Parechoviruses multiply in the respira- polyadenylated at the 3 end with a tory and gastrointestinal tracts either genome-linked protein (VPg) at the 5 asymptomatic or causing diarrhea, fre- end. There is a long 5 non-translated quently with respiratory complications. region (NTR) of 743 nt, including eight human polioviruses (PV-1 to 3) 230

AUG codons before the initiating AUG in the Americas since 1991, and the at position 743. Secondary structure Western Hemisphere has been declared within the NTR apparently obstructs free of polio by WHO. A major pro- ribosome entry before the initiation gram of global vaccination with labora- codon, and two internal ribosome tory support is being led by WHO with entry site (IRES) elements have been the goal of poliomyelitis eradication. identified that promote correct initia- Since outbreaks have occurred in which tion. Specific mutations in the 5 NTR components of oral polio vaccine have attenuate poliovirus neurovirulence, recombined with other enteroviruses, it especially a single C to U substitution is likely that in the final stages of eradi- at position 472. The cell receptor for cation only inactivated poliovirus vac- poliovirus is a member of the immuno- cine will be used. globulin gene superfamily with Synonyms: acute anterior poliomyeli- three Ig-like domains, mol. wt. 67 Da. tis virus; anterior poliomyelitis virus; The presence of the functional recep- Heine–Medin disease virus; infantile tor molecule presumably dictates tis- paralysis virus. sue tropism. Polioviruses infect human John TJ (1993) Rev Med Virol 3 , 149 or primate nasopharyngeal cells, Thompson KM et al (2006) Risk Anal 26 , 1571 Peyer’s patches in the intestinal tract Wimmer E et al (1993) Annu Rev Genet 27 , and motor neurons in the spinal cord, 353 but most other organs are refractory to infection. The virus is easily isolated human polyomaviruses To date five from feces in primary monkey kidney genetically distinct human polyoma- cell cultures or in HeLa cells. Produces viruses have been described in addi- a rapid CPE. A common infection tion to simian virus 40, which was of the human intestinal tract which accidentally injected into humans is usually silent, but disease occurs with some early lots of inactivated when the CNS is invaded with dam- poliovirus vaccine but is not gener- age to the anterior horn cells and lower ally believed to be a human virus. JC motor neuron paralysis. A similar dis- polyomavirus is the cause of progres- ease may be induced experimentally sive multifocal leukoencephalpathy, in chimpanzees and cynomolgus mon- a rare disease in immunosuppressed keys. Suckling mice are not susceptible, patients, from which the virus was but transgenic mice which have been originally isolated, BK polyomavirus transfected with the human poliovirus was originally isolated from the urine receptor gene can be infected. After of a patient on immunosuppressive 1961, following the widespread use of therapy following kidney transplan- vaccine, it was necessary to character- tation, and appears to be a common ize virus isolates as virulent or attenu- infection of humans which is usually ated, vaccine-derived or wild. The silent. Recently, by applying the SISPA monkey neurovirulence test is the only technique to human respiratory sam- one capable of assessing virulence, and ples a third human polyomavirus was transgenic mice may provide an alter- described and named KI polyoma- native, cheaper model for vaccine test- virus. This is genetically distinct from ing. Sequence-based polymerase chain the other polyomaviruses in the late reaction tests can distinguish vaccine region of the genome. Another human from wild strains. Two types of vac- polyomavirus has been reported from cine are available: formalin-inactivated Australia and also found in the USA (Salk) and live-attenuated (Sabin). in patients with respiratory infections The widespread use of Sabin vaccine, and named WU polyomavirus. Then which is given orally as a trivalent mix- in 2008 a fifth human polyomavirus, ture of viruses attenuated by passage in MC, was found in the Merkel cell car- cell culture (type 1, strain LSc1; type 2, cinoma. See BK polyomavirus and JC strain P2712; type 3, strain Leon 12a1b), polyomavirus . has eliminated poliomyelitis from Allander T et al (2007) J Virol 81 , 4130 most developed countries. No cases of Feng H et al (2008) Science 319 , 1096 poliomyelitis have been confirmed Gayner AM et al (2007) PLoS Pathol 3 , e64 231 Human rhinovirus B (HRV-B) human reovirus-like agent Synonym for Collins PL et al (1996) In Fields Virology , human rotavirus. Third edition, edited by BN Fields et al . Philadelphia: Lippincott-Raven, p. 1313 Human respiratory syncytial virus Lazzaro T et al (2007) J Paediatr Child Health 43 (HRSV) , 29 The type species in the genus Nuitjen MJ et al (2007) Pharmacoeconomics Pneumovirus . A common human respi- 25 , 55 ratory virus first isolated from a chim- Tripp RA (2005) In Topley & Wilson’s panzee. An important cause of lower Microbiology and Microbial Infections , vol. 1, respiratory tract infection in infants Tenth edition, edited by BWJ Mahy and V which may be more severe if they have ter Meulen (eds), London: Hodder Arnold, a low level of antibody, because this p. 783 may react with viral antigen in the Wu H et al (2007) J Mol Biol 368 , 652 tissues. The antibody is often mater- human respiratory syncytial virus A2 A nal. For this reason immunization may strain of HRSV. have unfavorable consequences and no protective vaccine has been devel- human respiratory syncytial virus B1 A oped. Reinfection may occur repeatedly, strain of HRSV. although serious disease is associated with the first or second infection. Causes human respiratory syncytial virus S2 A ‘ colds ’ in captive chimpanzees. Causes strain of HRSV. no symptoms in ferrets and can be serially passaged in them. Freezing of Human rhinovirus A (HRV-A) A spe- specimens for virus isolation reduces cies in the genus Rhinovirus containing the chance of success. Diagnosis is by 76 serotypes that all use ICAM-1 as a immunofluorescence using exfoliated receptor and 10 serotypes that use low cells, or by ELISA test of respiratory density lipoprotein as a receptor. The secretions. Replicates in human cell lines cause of most human ‘colds ’ and some such as HeLa and Hep-2 and less read- other respiratory tract infections. Do ily in monkey kidney cell cultures. Small not hemagglutinate. Strains which rep- syncytia appear and within 1–4 days the licate in rhesus monkey kidney cell cul- whole cell sheet is involved. No demon- tures are called M strains; the majority, strable hemadsorption. No replication in which multiply only in human cells, are eggs. Virion 90–120 nm in diameter and called H strains. Some strains can only variable in size. Matures at the cell sur- be isolated in organ cultures of respira- face by budding; in polarized epithelial tory epithelium. Optimal conditions for cells budding is from the apical surface. propagation are sodium bicarbonate There is a helical nucleocapsid 12–15 nm concentration of not more than 0.35 g/ in diameter containing negative-sense l, temperature 33°C, slow rotation of single-stranded RNA, 15222 nt in length, culture, islands of cells rather than con- encoding 10 genes. The virion RNA gene fluent sheet and minimal concentration order is 3 -NS1-NS2-N-P-MSH-G-F-M2- of serum compatible with maintenance L-5. The 3 end has a leader sequence of cells. Best cells for isolation are pri- of 44 nt and the 5 end a trailer sequence mary human kidney or lung or diploid of 155 nt. Inactivated by lipid solvents. human cell lines. CPE may not be seen Envelope covered with projections. for 10 days or only on passage. No vac- There is a single serotype, although neu- cine available but experiments suggest tralization tests reveal some antigenic immunity to an individual strain can differences between strains but no cross- be produced. A drug (Pleconaril) is reaction with other human respiratory under development. Chimpanzees can viruses. Two therapeutic monoclonal be infected but do not show symptoms antibodies which target the virus F proof a ‘ cold ’ . tein, palivizumab, and motavizumab, Synonyms: common cold virus; coryza are undergoing clinical trials for use in virus; ERC group viruses; murivirus; young children. respirovirus; Salisbury virus. Synonyms: bronchovirus syncytialis; chimpanzee coryza agent; respiratory Human rhinovirus B (HRV-B) 25 sero- syncytial virus of humans. types which are distinct from Human Human rhinoviruses B (HRV-B) 232

rhinovirus A. They were originally Blacklow NR and Greenberg H (1991) N human rhinoviruses 3, 14, and 72. Engl J Med 325 , 252 human rhinoviruses There are more than human syncytial virus Synonym for 150 serotypes of rhinoviruses that have Human foamy virus . not yet been assigned to a species. human T-cell leukemia virus Synonym human rotavirus The commonest cause for human T-cell lymphotropic virus of gastroenteritis in infants and less type 1. often in older children and adults. In infants diarrhea is often preceded human T-cell lymphotropic virus type by respiratory symptoms. Worldwide III The name given by workers at the nearly 1 million children die each year National Cancer Institute, USA to early from rotavirus diarrhea. Isolations isolates of the virus causing AIDS. from patients are commonest from Subsequently found to be identical November to April. Newborn rhe- to the French isolate, LAV LAI, and sus monkeys, calves and pigs can be renamed Human immunodeficiency infected experimentally. Antibodies virus type 1 (HIV-1). to calf rotavirus can be used to detect human T-lymphotropic virus type human virus. It may also be detected in 1 (HTLV-1) A strain of Primate fecal extracts by direct electron micro- T-lymphotropic virus I in the genus Delta- scopy, or by centrifugation on to mon- retrovirus. An exogenous retrovirus olayers of primary human embryo isolated in 1980 from a case of adult fibroblasts or LLC-MK2 cells followed T-cell leukemia (ATL), a disease which by overnight incubation and staining is endemic in Japan, the Caribbean, with labeled antiserum. Maternal anti- Melanesia, South America, Iran, and bodies probably provide some protec- parts of Africa. There is strong sero- tion from disease and may permit silent epidemiological evidence that HTLV-I infection. Infection in maternity hospi- is the cause of ATL. In 1985, HTLV-1 tal nurseries tends to cause mild disease was also found to be associated with a but in children over 6 months of age slowly progressive myelopathy known infection can cause severe symptoms. in the Caribbean as tropical spastic The first vaccine to be developed was paraparesis and in Japan as HTLV-1- a tetravalent rhesus-human reassortant associated myelopathy. This disease is vaccine (RRV-TV) which was licensed now termed HTLV-1-associated mye- in 1998 for use in the USA for children lopathy/tropical spastic paraparesis under 6 months of age, but was with- (HAM/TSP). It is believed to be immu- drawn within a year of use because of nologically mediated and is more fre- a rare but serious adverse event, intus- quently found in women than in men. susception, that occurred in about 1 in The latent period between infection 10,000 infants vaccinated. Since then, and disease is years to decades for two new vaccines have been approved ATL, but usually 2–4 years for HAM/ for use in infants – Rotateq, (licensed in TSP. The HTLV-I genome is 9032 nt the USA in 2006 and now in 46 coun- long and contains gag , pro , pol , and env tries worldwide) which is a bovine genes as well as long terminal repeats rotavirus-based pentavalent vaccine, (LTRs). The genome also contains a pX and Rotarix, (licensed in Mexico in 2004 sequence between the env gene and and now in 80 countries worldwide) the 3 LTR that includes two overlap- which is an attenuated human rotavirus ping regulatory genes: tax , the prod- vaccine. Both appear to be safe and to uct of which (p40 tax ) transactivates provide protective immunity to infants proviral transcription; and rex , the in large studies conducted worldwide product of which (p27rex ) modulates and have shown no adverse effects. See RNA processing. Two other proteins of Rotavirus, Rotaviruses A and B . unknown function, also encoded in this Synonyms : human reovirus-like agent; region of the genome, have been iden- infantile gastroenteritis virus. tified in infected cells. Cell-free HTLV-2 Bellamy AR and Both GW (1990) Adv Virus particles have extremely low infectivity Res 38 , 1 in vitro, but co-cultivation of virus- 233 Humpty Doo virus (HDOOV)

producing cells can be used to transmit HTLV-2 infection have only been found the virus to a variety of cells including in intravenous drug users in North human T and B lymphocytes, fibro- America, Italy, and Spain, and in the blasts, and epithelial cells, and cells from Pygmies in Central Africa. other species such as hamster, monkey, Hall WW et al (1994) Semin Virol 5 , 165 rat, and rabbit, but not mice. In infected Lewis MJ et al (2000) Virology 271 , 142 humans, only CD4 T cells appear Novoa P et al (2007) J Med Virol 79 , 182 to be infected. Most human transmis- Wolfe ND et al (2005) Proc Natl Acad Sci USA sion in the past was through blood 102 , 7994 transfusions. With the development of ELISA tests for the presence of p21e human T-lymphotropic virus type 3 antibodies, blood screening has been (HTLV-3) A new unclassified virus introduced in most developed coun- identified in two primate hunters in tries and HTLV-positive blood units Central Africa. The genome sequence are not used. Transmission may also shows that it is closely related to sim- occur through sexual contact and from ian T-lymphotropic virus 3 from mother to child during breast-feeding. Central Africa, and shares about 62% HTLV-1 proviral DNA has been identity with the genome sequences of detected in an Andean mummy about HTLV-1 and HTLV-2, and 87–92% iden- 1500 years old and provides evidence tity with STLV-3. Molecular dating puts of prehistoric migration from Asia to the origin of HTLV-3 as 36,000–50,000 South America. years ago. There is presently no information on the significance of the virus Kaplan JE and Khabbaz RF (1993) Rev Med in human disease. Virol 3 , 137 Li H-C et al (1999) Nat Med 5 , 1428 Calattini S et al (2006) J Virol 80 , 9876 Matsuoka M and Jeang KT (2007) Nat Rev Switzer WM et al (2006) J Virol 80 , 7427 Cancer 7 , 270 Salemi M et al (1998) Virology 246 , 277 Human torovirus (HuTV) A species in Verdock, K et al (2007) Lancet Infect Dis 7 , 266 the genus Torovirus . First detected Vidal AU et al (1994) J Gen Virol 75 , 3655 by electron microscopy of particles detected in human stool specimens. human T-lymphotropic virus type Morphologically resemble Breda virus. 2 (HTLV-2) A strain of Primate In a Canadian study over a 5-year pe- T-lymphotropic virus 2 in the genus riod of infants with necrotizing entero- Deltaretrovirus . Originally isolated colitis 48% had evidence of torovirus from an immortalized T-cell line (MoT) in stool cultures compared to 17% of established from the spleen of a patient controls. with a T-cell variant of hairy-cell leuke- Duckmanton L et al (1997) Virology 239 , 158 mia. The genome is 8952 bases long, Lodha A et al (2005) Acta Paediatr 94 , 1085 with a similar organization but only 60% homology to that of HTLV-1. No human wart virus Synonym for Human clear association with any human dis- papillomavirus . ease has been found to date, and the humoral immune response The devel- virus does not appear to be etiologically opment of antibodies in the ‘humors ’ , linked to hairy-cell leukemia. There plasma, lymph and tissue fluids, result- are two subtypes, known as HTLV2a ing in humoral immunity. and HTLV-2b, based upon nucleotide sequences, of the env gene, but further humoral immunity Immunity conferred subdivision has been found using LTR by antibodies in extracellular fluids sequences, which are more diverse. including the serum and lymph. High incidence of HTLV-2 (up to 10% of the population) has been found in Humpty Doo virus (HDOOV) An unas- certain native American Indian popu- signed vertebrate rhabdovirus. Isolated lations in which the infection appears from the mosquito, Culicoides marksi , in to be endemic. In Brazil, there is evi- Beatrice Hill, Northern Australia. Not dence that multiple subtypes circulate associated with disease in humans. in regions of high endemicity. Outside Standfast HA et al (1984) Aust J Biol Sci 37 , of these endemic foci, high rates of 351 Huncho virus (HUAV) 234

Huncho virus (HUAV) Does not exist. (gel or filter hybridization). The rate of The name appears in the 7th and 8th hybridization increases with salt concen- ICTV Reports due to a misprint. It tration, or with temperature up to just should read Huacho virus (HUAV) qv. below the melting temperature (Tm). hundestaupe Synonym for Canine distem- hybridoma A hybrid cell line produced per virus . from the fusion of a normal lymphocyte with a myeloma cell. After selection hundskrankheit (German: ‘ dog dis- and cloning a hybridoma cell line will ease’ ) Synonym for phlebotomus fever. produce a monoclonal antibody. So called because the conjunctivae become so markedly infected that they hydrocephalus of pike virus See pike fry resemble the eyes of a bloodhound. See rhabdovirus . phlebotomus fever viruses . hydrophobia virus Synonym for Rabies HV-114 virus (HV-114) A strain of virus . Hantaan virus in the genus Hantavirus . hydrops fetalis A fetal abnormality hybrid arrested translation (HART) A with gross edema of the entire body method for identifying the proteins and severe anemia. It can develop as encoded by a cloned DNA sequence. a result of B19 virus infection of the The mRNA preparation is hybridized mother during pregnancy. with the cloned DNA and only mRNA species homologous to the DNA will hydroxyapatite A calcium hydroxide – anneal to it. Comparison of in vitro calcium phosphate complex that binds translation products of annealed with double-stranded DNA but not single- unannealed mRNAs will identify the stranded DNA, and can be used to sep- protein, the synthesis of which is inhib- arate them. ited by hybrid formation. See hybrid released translation . α -[2-hydroxy-2-(3,5-dimethyl-2-oxo Paterson BM (1977) Proc Natl Acad Sci 74 , cyclohexyl)ethyl]glutarimide See 4370 cycloheximide . hybrid released (selected) translation 9-(2-hydroxyethoxymethyl)guanine See A method used to identify proteins acycloguanosine . encoded by a cloned DNA. A preparation of mRNA is hybridized to the cloned DNA immobilized on a solid matrix such hydroxyphosphonylmethoxycytosine as nitrocellulose. The mRNA homolo- (HPMPC) An antiviral compound, also gous to the DNA is retained on the filter known as cidofovir, which appears to and can then be removed by melting the be active against several species of her- RNA–DNA duplex. The purified RNA pesviruses and poxviruses. is then translated in vitro and the protein product(s) identified, often by gel 3β -hydroxysteroid hydrogenase An electrophoresis. enzyme encoded by vaccinia virus which converts pregnenolone to pro- Goldberg ML et al (1979) Meth Enzymol 68 , 206 gesterone. hybridization The formation of double- hyperchromic effect Increase in absorb- stranded nucleic acid molecules from ance of light of wavelength 260 nm by single-stranded polynucleotides with DNA at the melting point or transition complementary base sequences. The temperature. process may be carried out in the laboratory with both the single strands in solu- hyperimmune gammaglobulin Polyclo- tion (liquid hybridization), or with one nal antisera of high titer can be pro- strand immobilized on a solid support duced in animals for use in diagnostic 235 Hypr virus (HYPRV)

assays by repeated injection of virus or Hypr virus (HYPRV) A strain of tick- viral antigen. borne encephalitis virus in the genus Flavivirus . Isolated from a boy with hyperimmune serum Serum from an ani- encephalitis in Brno, Moravia. Anti- mal which has received two or more genically very similar or identical to injections of a foreign antigen for the Hanzalova virus, in the serogroup purpose of producing a reagent for use tick-borne encephalitis virus (Central in serology. European subtype). Frequent human infections in Hungary, Poland, the hypermutation Enhanced frequency former Yugoslavia, Austria, Bulgaria, mutation can be mediated by an error- Sweden, and Finland. Distinguished by prone polymerase. slow growth rate, narrow host range and high degree of cell association. hypersensitivity A state of altered reac- Can often be isolated from the kidney, tivity in which the body reacts with an and with particular frequency from the exaggerated immune response to a for- salivary glands. It is likely that viruses eign substance. Hypersensitivity reac- of this group might be found in all tions can be immediate or delayed. mammalian species were an adequate search to be made. They have already hypertrophy Increase in the size of an been demonstrated in humans, mouse, organ or tissue, e.g. mumps virus infec- ground squirrels, guinea pigs, various tion of the lymph nodes. primates, rats, horses, and pigs. hypovolemia An abnormally decreased volume of circulating fluid, especially plasma, in the body. I

I10 cells (CCL 83) A cell strain, initiated ICNV International Committee on from a mouse BALB/c Leydig cell tes- Nomenclature of Viruses. Established ticular tumor, which secretes proges- at the International Congress of terone and its derivative. Microbiology in Moscow in 1966. In 1973 it became the International Iaco virus (IACOV) A strain of Bunyamwera Committee on Taxonomy of Viruses virus in the genus Orthobunyavirus . (ICTV) which assumed the broader aim Isolated from mosquitoes, Wyeomyia sp. of developing a system of virus classi- Not associated with disease in humans. fication and nomenclature that would become a universally accepted taxon- Iatrogenic Creutzfeldt–Jakob disease omy of viruses. More than 250 cases of iatrogenic CJD have been reported, mostly from Icoaraci virus (ICOV) A serotype of administration of human pituitary hor- Rift Valley fever virus in the genus mone or dura mater grafts, but also Phlebovirus. Isolated from the rodent, from contaminated instruments during Proechimys guyannensis oris, in Para, neurosurgical procedures, and corneal Brazil, and sentinel mice in São Paulo, transplants. Brazil. Does not multiply in mosquitoes on salivary gland inoculation. Vector is Ibaraki virus (IBAV) A serotype of probably sandflies of Phlebotomus sp. Epizootic hemorrhagic disease virus in the genus Orbivirus . Isolated from cattle in icosahedral cytoplasmic deoxyribovirus Ibaraki Prefecture, Japan. Resembles Synonym for Iridovirus . bluetongue virus, from which it is antigenically distinct, in causing a disease icosahedral cytoplasmic deoxyribovi- (often severe) in cattle in Japan, but with ruses of amphibians Several viruses little pathogenicity for sheep. Probably in the genus Ranavirus , isolated from transmitted by arthropods. Replicates in frogs, newts, salamanders, and toads. bovine cell cultures with CPE and in the Recognized species include Ambystoma yolk sac of embryonated eggs. Not path- tigrinum virus , Bohle iridovirus , Frog ogenic for guinea pigs or rabbits. Not virus 3, and Santee-Cooper ranavirus , and to be confused with bovine ephemeral there are a number of tentative species. fever virus which causes a somewhat Frog viruses do not cause disease in similar disease. Found in Japan, Bali, their natural adult host, Rana pipiens , and Taiwan. A live attenuated Ibaraki but are lethal for frog embryos and lar- virus vaccine is produced in Japan. vae and Fowler toads. Grow in piscine, Synonym : Keishi virus. amphibian, avian, and mammalian Inaba Y (1975) Aust Vet J 51 , 178 cells at 12–32°C. The genome is a single Ito Y et al (1973) Arch Gesamte Virusforsch 40, 29 linear double-stranded DNA of about 170 kb which is highly methylated, IBDV See Infectious bursal disease virus . terminally redundant, and circularly permuted. Replication of DNA occurs IBV See Infectious bronchitis virus . in both the nucleus and the cytoplasm, but virus assembles in the cytoplasm. ICAM-1 See intercellular adhesion Synonym : cytoplasmic amphibian molecule-1 . viruses. Chinchar VG and Hyatt AD (2008) in BWJ ICI 73602 See 1-(p-chlorophenyl)-3- Mahy and MHV van Regenmortel (eds) (m-3isobutylguanidinophenyl)urea Encyclopedia of Virology, Third edition, hydrochloride . Oxford: Academic Press, vol. 3, p. 167 237 Ictalurid herpesvirus 1 (IcHV-1)

Kelly DC and Robertson JS (1973) J Gen Virol icosahedron A solid with 20 triangu- 20 , Suppl 17 lar faces and 12 vertices. In a regular Murti KG et al (1985) Adv Virus Res 30 , 1 icosahedron the faces are equilateral triangles and there are axes of two- fold, threefold, and fivefold rotational icosahedral symmetry One of the two symmetry. types of symmetry found in viral capsids, the other being helical symmetry. ICR 134 cells (CCL 128) A cell line estab- Crystallographic considerations pre- lished from the tissue of stage 17 gyno- scribe that the identical units forming genetic haploid embryos of the grass the capsid of an isometric particle must frog, Rana pipiens , and cloned at the be arranged with cubic symmetry. Of eleventh passage. the possible forms that this may take, icosahedral symmetry provides the ICR 2 A cells (CCL 145) A haploid frog facility to make a range of viral capsids cell line established from androgenetic with different numbers of structural haploid embryos of the grass frog, Rana units. An icosahedron has 20 triangular pipiens . faces and 12 vertices. The simplest has 60 identical structural units in regu- Ictalurivirus An unassigned genus in the lar relation to each other, three to a tri- family Herpesviridae. The only species angular face. To make a large virus in in the genus is Ictalurid herpesvirus 1 this simple form from 60 units would (channel catfish virus). require a large protein, which raises difficulties with genome coding capac- Ictalurid herpesvirus 1 (IcHV-1) The type ity, and an alternative is to use a larger species of the genus Ictalurivirus (which number of small units (i.e. more than is unassigned to a subfamily) in the fam- 60). This inevitably means that the units ily Herpesviridae. Isolated from young cannot all have identical relationships to channel catfish, Ictalurus punctatus , in each other. Those not surrounding a ver- which it causes a severe hemorrhagic tex form groups of six called ‘ hexons, ’ disease which may have a mortal- and those at each vertex are in groups of ity rate in excess of 95%. The first fish five called ‘ pentons. ’ Only certain mul- herpesvirus to be isolated, it has been tiples of 60 units are possible, and the extensively studied. Experimentally, the numbers which make up different viral virus will infect blue catfish, Ictalurus capsid structures are defined by the tri- furcatus , but other species of fish are not angulation number, T . There are always affected. The virus replicates in ictalu- 2 2 60T units, where T h hk k (h and rid and clariid fish cell lines, between k are integers having no common fac- 10°C and 33°C, and about 50% of the tors). Examples are T 3 (caliciviruses), progeny is released into the culture T 4 (alphaviruses), T 13 (rota- medium. Morphologically, IcHV-1 is viruses and orbiviruses) T 16 (herpes- a typical herpesvirus. The complete viruses), and T 2 5 (adenoviruses). DNA sequence has been determined The structural units form into morpho- as 134,226 bp, with G C 56%. The pre- logical units on the virus surface. In dicted proteins are quite different from general, the number of morphological those of mammalian and avian herpes- units (capsomeres) which can be visu- viruses. The virus may become latent alized on the surface of an icosahedral in adult fish, and is perpetuated by this virion is 10T 2 (e.g. 162 for herpes- means in channel catfish populations. viruses, 252 for adenoviruses). There are no effective treatments for the disease. No related species have yet Caspar DLD and Klug A (1962) Cold Spring been recognized. Harb Symp Quant Biol 27 , 1 Synonym : channel catfish virus. Crick FHC and Watson JD (1956) Nature 177 , 473 Chinchar VG et al (1996) Dis Aquat Organ Johnson JE and Speir JA (2008) in BWJ 33 , 77 Mahy and MHV van Regenmortel (eds) Davison AJ (1992) Virology 186 , 9 Encyclopedia of Virology , Third edition, Davison AJ (2008) in BWJ Mahy and MHV Oxford: Academic Press, vol. 5, p. 393 van Regenmortel (eds) Encyclopedia of Ictalurid herpesvirus 1 (IcHV-1) 238

Virology , Third edition, Oxford: Academic success as eye drops in the treatment Press, vol. 2, p. 205 of herpes simplex keratitis, particularly dendritic ulcers of the cornea, but is ICTV International Committee on Tax- now largely supplanted by acyclovir. onomy of Viruses. A committee of the Synonyms : dendrid; herpid; idoxyurid- Virology Division of the International ine; Idurin; staysail. Union of Microbiological Societies (IUMS) which decides upon the clas- IDU See idoxuridine . sification and nomenclature of viruses affecting all species of animals, plants, fungi, bacteria, and archea. Since Idurin See idoxuridine . their work began in 1968, the ICTV has issued eight Reports, the most IE genes Immediate-early genes, expressed recent of which was presented to the e.g. during herpesvirus replication. International Congress of Virology held in San Francisco, USA in July 2005. The Ieri virus (IERIV) A species in the genus ICTV is composed of an 18-member Orbivirus . There are three recognized Executive Committee supported by serotypes, Ieri virus , Gomoka virus, and numerous subcommittees and study Arkonam virus. Isolated from mosqui- groups. Deliberations of these groups toes in Trinidad and Brazil. Not known are available online at http://talk. to cause disease in humans. ictvonline.org. IFA test Indirect fluorescent antibody Fauquet CM, Mayo MA, Maniloff J, test, used to detect the presence of virus Desselberger U, and Ball LA (Editors) (2005) antigens in clinical specimens. The Virus Taxonomy , Eighth Report of the ICTV. London: Academic Press specimen is initially reacted with a primary antibody that is directed against ID50 The 50% infective dose. The dose a specific virus antigen. Binding of that on average will infect 50% of the the primary antibody is then detected individuals to which it is adminis- by adding a secondary antibody– tered. They may be human volunteers, fluorochrome conjugate directed against experimental animals, tissue cultures, the primary antibody, and after washing or eggs. When eggs are used the term the presence of bound antibody in the EID50 is often used. specimen can be visualized by fluorescence microscopy. The technique idiotype The structural features of the is generally more sensitive than labe- variable regions of a particular anti- ling the primary antibody itself (direct body from a single individual. Anti- fluorescent antibody technique) since idiotype antibodies combine with it multiplies the number of fluoro- these structures, and may resemble chromes per antigen stained resulting the epitope to which the first antibody in brighter, clearer staining. reacts. See also isotype . Schutzbank TE and McGuire R (2000) In idoxuridine (IDU) 5-Iodo-2 -deoxyuridine. Clinical Virology Manual, edited by S Specter, An antiviral agent and an analog of RL Hodinka and SA Young. Washington: ASM Press, p. 69 thymidine. Its action is due to its incorporation into DNA, when it becomes active against DNA viruses such as Ife virus (IFEV) A tentative species in the human herpesvirus types 1, 2, and 3, genus Orbivirus. Isolated from a bat, vaccinia and suid herpesvirus 1. The Eidolon helvum , and from mosquitoes. drug is probably inactive until enzy- α matic conversion into the nucleotide IFN- See interferons . that is a competitive inhibitor for the incorporation of thymidine nucleotides IFN-β See interferons . into DNA. This occurs during both cellular and viral DNA synthesis. Because IFN-γ See interferons . of its toxicity it is used mainly as a topical application. Has been used with IgA See immunoglobulin . 239 immortalization

Igbo Ora virus A virus in the genus Ilesha virus (ILEV) A serotype of Alphavirus . Isolated from human sera Bunyamwera virus in the genus in Central Africa in 1967 and Nigeria in Orthobunyavirus , antigenically related 1966 and 1969. Caused an epidemic of to Bunyamwera virus. Isolated in fever, body pains, and rash in the Ivory Nigeria, Uganda, Cameroon, Central Coast in 1984, when it was isolated African Republic, and Senegal from the from humans and mosquitoes. During mosquito, Anopheles gambiae . an epidemic of O’nyong-nyong fever in Uganda in 1996, both the O’nyong- Ilhéus virus (ILHV) A species in the genus nyong virus and Igbo Ora virus genomes Flavivirus belonging to the Ntaya virus were sequenced, and it was concluded group. Main natural host not known that Igbo Ora virus is a strain of O’nyong- but antibodies are found in humans, nyong virus. horses, and birds. The virus has been Lanciotti RS et al (1998) Virology 252 , 258 isolated from at least eight genera of Moore DL et al (1975) Ann Trop Med Parasitol mosquitoes and from febrile patients, 69 , 49 sentinel monkeys, and once from a bat in the Amazon region of Brazil. In IgE See immunoglobulin . addition to Brazil, occurs in Colombia, Central America, and the Caribbean. IgG See immunoglobulin . Usually causes an inapparent infection in humans but a few cases of encepha- IgH-2 cells (CCL 108) A cell line derived litis are recorded. Mice injected i.c. from the heart of a normal, immature develop encephalitis. Viremia occurs male iguana,Iguana iguana . Supports the in several species but not in chickens replication of herpes simplex, pseudo- and pigeons. There is replication on the rabies, vaccinia, and iguana viruses at CAM but most virus is in the embryo. 36°C. In chick embryo, BHK21 and Vero cells, virus replicates with CPE. IgM See immunoglobulin . Southam CM et al (1951) Am J Trop Med Hyg 31 , 724 iguana virus Synonym for iguanid herpesvirus 1. Iltovirus A genus in the subfamily Alphaherpesvirinae containing a single iguanid herpesvirus 1 (IgHV-1) A possi- species, Gallid herpesvirus 1 (Infectious ble species in the family Herpesviridae . laryngotracheitis virus). Isolated from a spontaneously degenerating cell culture of tissue from a green IM-9 cells (CCL 159) An immunoglobulin- iguana, Iguana iguana. On injection into secreting cell line, established from a iguanas it caused no consistent disease bone marrow sample from a female pattern observable on necropsy, but 7 patient with multiple myeloma. of 12 animals injected died. Has narrow host range. Replicates in iguana Imiquimod 1-(2-methylpropyl)-1H -imidazo cell cultures with CPE at certain (4.5-c )quinolin-4-amine. An antiviral temperatures. agent active against human papilloma- Synonyms: iguana virus; green iguana viruses. May act by inducing interferon herpesvirus. alpha. Clark HF and Karzon DT (1972) Infect Miller R et al (1995) Int Antivir News 3 , 111 Immun 5 , 559 immobilized DNA (or RNA) Term used to Iguape virus (IGUV) A serotype of Aroa describe nucleic acid attached to mem- virus in the genus Flavivirus . Isolated brane filters or activated paper for the from sentinel mice exposed in a for- purpose of DNA–RNA hybridization. ested area of Iguape, Brazil, in 1979. Antibodies were detected in 25 species immortalization A change produced by of birds belonging to 16 families. virus infection of a cell culture, which Coimbra TLM et al (1993) Intervirology 36 , 144 results in continued growth of the cells immortalization 240

beyond the time at which it would production of specific antibody pro- have been expected to stop. Similar vides protection against reinfection or identical to transformation, but the but there are instances where antibody term is used to emphasize the fact that alone is insufficient. there is no evidence of neoplastic trans- Nash AA and Dutia B (2005) In Topley & formation. Used especially to describe Wilson ’ s Microbiology and Microbial Infections , the continued growth of human cells vol. 1, Tenth edition, edited by BWJ Mahy after infection with Epstein–Barr virus. and V ter Meulen. London: Hodder Arnold, p. 270 immune complex diseases Diseases that result from deposition of virus antigen– immune therapy The effect of licensed antibody complexes in tissues such as vaccination of chronic carriers of hepa- the kidney or blood vessels. Usually titis B virus has only a small effect on occur when an excess of antigen cir- the disease. However the use of a hep- culates as complexes with antibody. atitis B DNA vaccine has proved more Examples are glomerulonephritis and successful. polyarteritis nodosa in chronic hepati- Roy MJ et al (2000) Vaccine 19 , 764 tis B carriers, and the immune complex diseases characteristic of Aleutian mink immunity disease or lymphocytic choriomeningi- The condition of a living tis virus infection. organism whereby it resists and over- comes an infection or disease (pro- Lai KN et al (1991) N Engl J Med 324 , 1457 tective immunity). Active immunity occurs in response to stimulation with immune evasion In order to survive for antigen (e.g. vaccine) during infec- prolonged periods viruses must hide tion. Passive immunity is due to anti- from or evade the immune response of body or primed lymphocytes derived the host. Certain sites such as the nerv- from another immune individual (e.g. ous system are immunologically privi- maternal immunity). leged, and the virus is protected from the immune system while replicating immunization Rendering an organism there. Herpesviruses are one example, immune to a specific disease pathogen. but measles or rabies viruses can also Usually performed by injecting prepa- survive for many years in the human rations into the organism which will CNS. Some complex viruses such as induce antibodies against the causal cytomegalovirus or variola virus have agent of the disease. Vaccines may developed specific ways to inhibit the contain killed virus, e.g. Salk vaccine human immune response to accom- against poliovirus, or live attenuated plish immune evasion. virus e.g. Sabin (live oral poliovirus) Fazakerley JK and Buchmeier MJ (1993) Adv vaccine, which can be given orally. This Virus Res 42 , 249 has helped enormously in the campaign Nash A et al (1996) Semin Virol 7 , 125 to eradicate poliomyelitis. Vaccines based on plants such as banana or immune modulators Substances such potato are under development for oral as interferon which are used to treat administration, which could revolu- chronic hepatitis B or hepatitis C. tionize immunization practices. Interferon may be conjugated with polyethylene glycol (PEG) to improve immunoassays Assays for virus infection its pharmacological properties (PEG- based upon the presence of antibodies interferon). in serum samples. immune response The response of the immunocompromised patients Patients immune system to an antigen, e.g. for- with deficiencies of the normal immune eign proteins or carbohydrates. It can response may suffer from debilitating be a humoral (antibody), cell-mediated manifestations of diseases caused by (T-cell) response, or immunological herpesviruses, but in some infections tolerance. In most virus diseases, the such as hepatitis B, the disease may 241 immunoglobulin class

be unusually mild. The latter condi- immunogenic Capable of inducing tion may result if the immune response humoral or cell-mediated immunity. itself contributes to the disease. immunoglobulin A set of proteins pro- immunocytochemistry The identifica- duced in the immune response. Several tion of the location of antigens in cells classes of immunoglobulins have the by the use of antibodies to which a same basic structure of two identi- reporter molecule, e.g. ferritin, gold, or cal light (L) polypeptide chains and a fluorescent dye, is attached. two heavy (H) chains linked together by non-covalent forces and disulfide immunodeficiency viruses See specific bonds. There are five classes distin- virus, e.g. Human immunodeficiency guished on the basis of five different viruses or Simian immunodeficiency types of H chain: IgG, IgA, IgM, IgD, viruses . and IgE; the heavy chains are termed γ, α, μ, δ, and ε, respectively. The con- immunodiffusion A serological proce- centration of IgD and IgE in animal dure in which antigens and antibod- serum is very low. IgG is the most ies in solution are permitted to diffuse common, comprising about 75% of toward each other through a gel matrix. all immunoglobulins; it has a mol. wt. The interaction between the antigen of 150 1 03 and will fix complement. and antibody is manifested by a ‘ pre- IgM has a mol. wt. of 900 1 0 3 and cipitin’ line produced by the precipita- also fixes complement. IgA has a mol. tion of the antigen–antibody complex. wt. of 160 1 0 3, does not fix complement, and is the major immunoglobulin immunoelectron microscopy Techniques of the external secretions (intestinal in which virus and specific antiserum fluids, saliva, bronchial secretions, etc.). are mixed before examination by elec- Following virus infection, antibodies tron microscopy. The antibody agglu- appear in 5–7 days. The first class to tinates the virus particles into small appear is IgM, which is usually of low clumps which are easier to find. The affinity. Within 2 weeks IgG becomes technique can also be used to test for the the dominant class in serum. IgA pro- presence of antibodies to a known virus, vides the main defense at mucosal sur- or to identify a virus using a range of faces where it is found as a dimer linked sera. See also immunogold labeling . to a secretory piece (a 60-kDa polypep- Almeida JD and Waterson AP (1969) Adv tide synthesized by epithelial cells in Virus Res 15 , 307 the secretory tissues) which has a strong affinity for mucus and protects the IgA immunoelectrophoresis A technique in against destruction by enzymes in the which an antigen mixture is first sepa- digestive tract. rated into its components by electrophoresis in a supporting medium (e.g. immunoglobulin class Classes of agar gel), then allowed to react with immunoglobulin are determined by antiserum so that immunoprecipita- the amino acid sequences of their tion lines are allowed to develop. It is a heavy chains. Most mammals have powerful analytical method for resolv- five classes: IgM, IgG, IgA, IgD, and ing complex mixtures of antigens. IgE, IgA is the major immunoglobulin of external secretions (intestinal flu- immunofluorescence (IF) A technique in ids, saliva, bronchial secretions, etc.). which antigen or antibody are conju- IgD is in low concentration in serum, gated to fluorescent dyes for detecting but in high concentration as mem- the corresponding antigen or antibody brane immunoglobulin on the surface in cells and thin sections by fluores- of B lymphocytes. IgE is in low con- cence microscopy. centration in serum, and is the main immunoglobulin associated with imme- immunogen A substance which induces diate hypersensitivity. IgG is the major the production of specific antibodies in immunoglobulin in human serum. IgM a suitable animal. is a high molecular weight immuno- immunoglobulin class 242

globulin, which is the first antibody immunoperoxidase assay The linking of class to appear in response to virus horseradish peroxidase to antibodies. infection, and activates complement. The antibodies are then used in tests such as ELISA or Western blotting and immunoglobulin prophylaxis Originally in cytological studies; the presence of used to protect travelers against hepa- the peroxidase is detected by reaction titis A virus infection, but now largely with a substrate that gives a color. replaced by specific immunization with hepatitis A vaccine. immunoprecipitation The precipitation of antigen–antibody complexes that immunoglobulin subclass Subdivision forms the basis of several serological within each immunoglobulin class is tests. When this occurs in solution the based upon structural and antigenic test is called a ‘ precipitin ’ test; when the differences in their heavy chains. There reactants diffuse toward each other in a are four subclasses of IgG and two sub- gel it is known as ‘ immunodiffusion. ’ classes of IgA. immunosuppression Suppression of the immunogold labeling The linking of immune response, e.g. by drugs, infec- colloidal gold molecules to antibod- tion, irradiation, or antilymphocyte ies. The gold molecules can then be serum. Deliberate immunosuppression detected by electron microscopy of is needed, e.g. to prevent rejection fol- samples containing antigen which have lowing transplant surgery, and this been treated with the labeled antibody. can lead to reactivation of latent virus Beesley JE and Betts MP (1984) Proc Roy infections such as herpesviruses. Microscop Soc 19 , 36 immunotherapy The use of antibodies immunohistochemistry A technique for to ameliorate or prevent disease symp- detecting specific antigens during his- toms. In the case of virus-induced tological examination by first treating tumors, antibodies against viral the pathological specimen with a spe- antigens may prevent tumor growth. cific antibody linked to a dye such as naphtha red. importins Cellular proteins which medi- immunological drift Synonym for anti- ate bidirectional transport of macro- genic drift. molecules between the nucleus and the cytoplasm through nuclear pore com- immunologically privileged sites The plexes. Importin- and importin- are brain and the kidney are considered cellular proteins which bind to a sig- immunologically privileged sites. nal on the nucleocapsid of hepatitis B Virus may persist in the brain because virus and allow it to pass through the the blood–brain barrier limits the traf- nuclear pore. ficking of lymphocytes through the Pante N and Kann M (2002) Mol Biol Cell brain, and also since neurons express 13 , 425 few MHC class I molecules, so they are poor targets for cytotoxic T lympho- IMR-32 cells (CCL 127) A cell line estab- cytes. Several viruses persist in the lished from an abdominal mass occur- kidney, such as polyomaviruses JC and ring in a 13-month-old Caucasian male. BK, and the arenavirus lymphocytic choriomeningitis virus, but the mechanism for this persistence is not clear. IMR-33 cells (CCL 146) A continuous cell line, derived from the fibroma of a ger- immunopathology Damage to tissues bil, Meriones unguiculatus . by the immune system. In some virus infections, such as hepatitis B infection, IMR-90 cells (CCL 186) A human dip- the disease has a large immunopatho- loid fibroblast cell line, derived from logical component, making treatment the lungs of a 16-week-old female difficult. fetus. 243 Indian muntjac cells in situ hybridization Use of specific of nuclear inclusion bodies in enlarged nucleic acid probes to localize viral infected cells. nucleic acids in cells or tissues. inclusion body hepatitis (IBH) of chickens inactivated poliomyelitis vaccine The A disease caused by several differ- first anti-poliomyelitis vaccine to be ent species in the genus Aviadenovirus . produced was a formalin-inactivated Causes a common and often fatal dis- preparation made by Jonas Salk in 1960 ease of chickens. At least 10 serotypes of administered by injection. However, fowl adenovirus have been associated some early vaccine batches made by with natural outbreaks. There are liver Cutter Laboratories in California con- lesions, intramuscular hemorrhages, tained aggregates of virus which sur- and aplastic anemia. Yolk-sac inocula- vived the formalin treatment, and tion of chick embryos causes death. led to the inoculation of an estimated Plaques are produced on the CAM. 200,000 persons with live virus in what Birds with antibodies and eggs from was termed the Cutter-Incident. 70,000 such birds are resistant to infection. persons became ill and 200 were per- Fadley AM and Winterfield RW (1975) Am J manently paralyzed as a result. Since Vet Res 36 , 532 the 1980s this problem has been over- McCracken RM et al (1976) Avian Pathol come, and the Salk vaccine has been 5 , 325 used extensively in poliovirus control. inclusion body hepatitis (IBH) of psit- Nathanson N and Langmuir AD (1963) Am tacine birds A usually fatal disease of J Hyg 78 , 16 budgerigars and parrots with gross liver lesions first described by Pacheco inactivation Loss of infectivity of a virus. and Bier in 1930 following an epiz- Can result from exposure to certain ootic in parakeets, and also known as chemicals, specific antibodies or adverse Pacheco’s disease. Often appears in physical conditions such as heat or irra- captive birds following movement or diation. See also neutralization . delays in transit. The disease is caused by psittacid herpesvirus 1. inapparent infection An infection which Pacheco G and Bier O (1930) CR Seances Soc does not give recognizable symptoms. Biol Paris 105 , 109 incidence rate The number of cases of inclusion body protein A term some- disease in a specified period of time times used for the main constituent of divided by the population at risk. an inclusion body. Usually applied to acute diseases of short duration. inclusion body rhinitis virus Synonym for suid herpesvirus 2. inclusion body An area of abnormal inclusion disease of pigeons virus staining in a virus-infected cell. Visible Synonym for columbid herpesvirus 1. by light microscopy and may be single or multiple, large or small, round or incomplete virus Virus lacking some part irregular, intranuclear or intracytoplas- necessary for its replication. Synonym mic, acidophilic or basophilic. Often for defective virus. composed of viral nucleic acid or proteins, but in some infections formed incubation period The interval between of cellular material. Of limited use in infection and the development of the diagnosis of certain infections, e.g. disease. Negri bodies in the brain cells of animals suspected of having rabies. Indian cobra herpesvirus Synonym for elapid herpesvirus 1. inclusion body disease virus Synonym for Human herpesvirus 5 . Cytomegalovirus Indian muntjac cells (CCL 157) This cell induces ‘ owl-like’ inclusions, consisting line, derived from a skin biopsy of an Indian muntjac cells 244

adult male muntjac, Muntiacus munt- gasping; rales are heard. The disease jak , has the lowest mammalian diploid lasts 6–18 days and the mortality is up chromosome number 7. to 90%. In laying birds there is a drop in egg production and eggs are defec- Indiana virus Synonym for Vesicular sto- tive. Pheasants may be infected. Mild matitis Indiana virus . endemic infection may result in poor egg production and predispose to bacte- indicator cells See co-cultivation . rial respiratory disease. Avian nephrosis and visceral gout may be caused by the indinavir A licensed antiretroviral drug virus, possibly by certain strains (see which is an inhibitor of the HIV pro- Australian infectious bronchitis virus ). tease. Also known as L-735,524. The virus is very infectious and spreads by the respiratory route. Birds are infec- indirect fluorescent antibody test See tious for up to 35 days after recovery IFA test . but there is no evidence of a carrier state. Virus can be isolated from eggs induction Activation of a latent virus and semen of experimentally infected infection. The activation can occur birds. Both live attenuated and inacti- spontaneously (e.g. herpes simplex vated vaccines are used, but offer poor virus latent in dorsal ganglia and acti- cross protection against other serotypes. vated to give ‘ cold sores ’ ), by changing Antibodies have been found in humans growing conditions of infected primary who handle poultry. Replicates on the cells, or by a variety of exogenous CAM but produces no definite pocks. In stimuli. chick embryo cell cultures there is replication with syncytium formation and infantile gastroenteritis virus Synonym cell necrosis. The viral RNA is a single for human rotavirus. molecule of single-stranded RNA of mol. wt. 9 1 0 6 (27.6 kb). Replication infantile paralysis virus Synonym for involves synthesis of a nested set of 3 Poliovirus . co-terminal subgenomic mRNAs that are 5 capped and 3 polyadenylated. infectious anemia of horses virus Only the 5 unique regions of these Synonym for Equine infectious anemia mRNAs are translated. There are at least virus ; swamp fever virus. five viral polypeptides: spike, membrane, nucleocapsid, hemagglutinin- infectious arteritis of horses virus esterase, and small membrane protein. Synonym for Equine arteritis virus . Synonyms : gasping disease virus; avian infectious bronchitis virus. infectious bovine rhinotracheitis virus Synonym for Bovine herpesvirus 1 . Cavanagh D (2007) Vet Res 38 , 281 Gough RE et al (1992) Vet Rec 130 , 493 Infectious bronchitis virus (IBV) The type species of the genus Coronavirus . infectious bulbar paralysis virus The cause of a common, contagious, Synonym for pseudorabies virus. acute respiratory disease of chickens, but other avian species are suscepti- Infectious bursal disease virus (IBDV) The ble. Neutralization tests using chick type species of the genus Avibirnavirus . embryos indicate multiple variant The virion has a single capsid structure antigenic types. All strains show some of 32 capsomeres. The genome consists antigenic relationships but are unre- of two segments of double-stranded lated to other coronaviruses. Beaudette RNA 3128 and 2795 bp in length, with strain (IBV-42) is serologically similar to a 5 genome-linked protein (VPg) on Massachusetts strain, although on egg each segment and no 3 poly A. There passage it has become lethal for chick is no envelope. Diameter is 60 nm, den- embryos but has lost infectivity for older sity about 1.32 g/ml. Capsid composed birds. Chicks up to 4 weeks old are most of four major polypeptides. Replicates susceptible. They show depression and in chicken embryo fibroblasts rather 245 Infectious hematopoietic necrosis virus (IHNV)

than in epithelioid cells. Resistant to infectious center assay A technique for ether, chloroform, pH 2 and incubation determining the proportion of cells in a at 56°C for 5 h; difficult to remove from cell suspension that are able to release chicken houses. Causes an acute, highly infectious virus. A suspension of cells contagious lymphoproliferative condi- infected with a virus is layered on a tion of chickens; 2- to 15-week-old birds cell monolayer culture, and covered are most commonly affected. The bursa with a layer of solid medium to pre- of Fabricius is involved, so that the vent the virus released from spreading immune system does not develop prop- too widely. Cells which release virus erly, resulting in an inability to resist form plaques (infectious centers) in the other infections. There is nephrosis. monolayer. Transmission is probably via food and water. Egg- or mouse-adapted attenu- infectious dose It is possible to define ated virus has been used as a vaccine the infectious dose of a virus prepara- with success. Two serotypes of IBDV tion by endpoint dilution and titration have been identified by cross-neutrali- in a susceptible animal species, but this zation. Serotype 1 strains are pathogenic is seldom possible in human subjects. in chickens whereas serotype 2 strains The measure used in human studies is are nonpathogenic. The 3D structure of the secondary attack rate (SAR) which the virus has been determined. describes numerically the communica- Synonyms: avian nephrosis virus; bility of the virus. Gumboro disease virus; infectious bur- sitis virus ( Table I1 ). infectious dropsy of carp virus Synonym for spring viremia of carp virus. But Bottcher B et al (1997) J Virol 71 , 325 the chronic form of infectious dropsy, Kilbenge FSB et al (1988) J Gen Virol 69 , 1757 Mahardika GNK and Becht H (1995) Arch erythro-dermatitis, is caused by a dif- Virol 140 , 765 ferent agent: nonmotile bacteria of Morgan MM et al (1988) Virology 163 , 240 Aeromonas sp. Fijan NN (1972) Symp Zool Soc Lond 30 , 39 infectious canine hepatitis virus Synonym for canine adenovirus 1. infectious enteritis A disease of poultry with diarrhea, hepatic necrosis, infectious catarrh of rats See pneumonia and monocytosis occurring mainly in virus of rats . young turkeys and pullets. See Turkey coronavirus . Synonyms : avian diarrhea virus; avian monocytosis virus; pullet disease virus. Table I1. Strains of IBV infectious equine arteritis virus See Infectious bursal disease virus 002–73 Equine arteritis virus . Infectious bursal disease virus 23/82 Infectious bursal disease virus 52/70 Infectious hematopoietic necrosis virus Infectious bursal disease virus Australian (IHNV) The type species of the genus 002–73 Novirhabdovirus. Causes necrosis of Infectious bursal disease virus Cu-1 Infectious bursal disease virus Edgar hematopoietic tissues of spleen and Infectious bursal disease virus Farragher anterior kidney in trout and salmon in Infectious bursal disease virus GPF-1E North America, Europe, Korea, Taiwan, Infectious bursal disease virus KS Japan, and mainland China. Occurs as Infectious bursal disease virus OH epidemic disease in fish hatcheries. The Infectious bursal disease virus OKYM fish do not feed and often have a dark attenuated red subdermal lesion dorsally located Infectious bursal disease virus OKYM at the back of the head. Replicates in Infectious bursal disease virus P2 fat head minnow cells or other fish Infectious bursal disease virus PBG-98 cell lines with CPE. Studies in rainbow Infectious bursal disease virus QC-2 Infectious bursal disease virus STC trout in fish hatcheries in Idaho show Infectious bursal disease virus UK661 that there is wide genetic diversity amongst IHNV populations. Infectious hematopoietic necrosis virus (IHNV) 246

Synonyms : chinook salmon virus; Aquabirnavirus. The cause of an acute, Oregon sockeye disease virus. contagious and highly lethal disease Bjorklund HV et al (1996) Virus Res 42 , 65 of a variety of salmonid fish including Kim CH et al (1999) J Virol 73 , 843 young rainbow trout in North America, Troyer RM et al (2000) J Gen Virol 81 , 2823 Europe, and Japan. Adult trout and Winton JR (1991) Annu Rev Fish Dis 1 , 83 salmon do not exhibit signs of infection but may become lifelong carriers. infectious hepatitis virus An old name Infected young fish swim erratically for Hepatitis A virus . and eventually die. There is necrosis of the pancreatic acinar and islet tissue. infectious immune complexes The cir- Experimentally, the virus causes pancre- culation of infectious virus in the pres- atic lesions in mice. Replicates in vari- ence of neutralizing antibody has been ous fish cell cultures with CPE but not seen with a number of viruses, such in mammalian cells. Virus has a single as lactate dehydrogenase-elevating capsid structure, is 60 nm in diameter, virus, lymphocytic choriomeningitis with 92 capsomeres and two pieces of virus, and Aleutian mink disease virus. double-stranded RNA, 3092 and 2784 bp These viruses all target macrophages in length, with a 5 genome-linked pro- as their host cell, and it is possible that tein (VPg) on each segment and no 3 the immune complexes are bound to Fc poly A. Ether-resistant but acid-labile. receptors, and internalized in vacuoles in which the complex dissociates. Dobos P and Roberts TE (1983) Can J Microbiol 29 , 1377 Duncan R et al (1991) Virology 181 , 541 infectious jaundice Old name for Hepatitis A virus . infectious particle A virus particle that infectious labial dermatitis virus contains the complete viral genome Synonym for Orf virus . and is capable of infecting a susceptible cell. infectious laryngotracheitis virus Synonym for Gallid herpesvirus 1 . infectious porcine encephalomyelitis virus Synonym for porcine enterovirus. infectious mononucleosis virus Synonym for Human herpesvirus 4 . infectious porcine poliomyelitis virus Synonym for porcine enterovirus. infectious myocarditis of gosling virus See Goose parvovirus . infectious pustular vulvovaginitis virus Synonym for Bovine herpesvirus 1 . infectious nucleic acid Nucleic acid which is able to infect cells and initiate Infectious salmon anemia virus (ISAV) the production of complete virus par- The type species of the genus Isavirus ticles. Nucleic acid removed from the in the family Orthomyxoviridae . Isolated virus particle is not protected from inac- mainly from Atlantic salmon (Salmo tivation by tissue nucleases. It does not salar ) in salmon farming areas on the depend on cell receptors for attachment Atlantic coasts of North America and to, and ability to infect, cells and so may Norther Europe. Placed on the number be able to infect species which the intact one list of the most dangerous fish virus cannot. For example, human viruses by the European Union, the dis- poliovirus nucleic acid injected i.c. in a ease is characterized by severe anemia, mouse can initiate one cycle of infection. ascites, and hemorrhagic liver necrosis, Complete virus is produced but cannot infect further cells. Infectious nucleic acid is readily isolated from viruses Table I2. Strains of IPN virus which do not require virion-contained enzymes to initiate replication. Infectious pancreatic necrosis virus DRT Infectious pancreatic necrosis virus Jasper Infectious pancreatic necrosis virus Infectious pancreatic necrosis virus N1 Infectious pancreatic necrosis virus Sp (IPNV) The type species of the genus 247 Influenzavirus A

with a mortality rate of 15–100%. An of this genus all have eight genome enveloped virus, diameter 130–140 nm, segments. Hemagglutination and the with a single-stranded RNA genome neuraminidase receptor-destroying with negative polarity and eight seg- enzyme are different glycoproteins. The ments. Total size of the genome is conserved end sequences of the viral 14.5 kb. The virus replicates optimally at RNAs of the influenzaviruses A are 15°C and does not replicate above 25°C. 5 -AGUAGAAACAAGG … and 3 -UCG Hemagglutinates piscine but not mam- (U/C)UUUCGUCC … The exact order malian or avian erythrocytes, and has a of electrophoretic migration of the RNA receptor-destroying enzyme that is an segments varies with strain and electro- acetylesterase. The virus is now widely phoretic conditions. Although there are spread geographically in the Americas eight genome segments, three (segments and Europe, and there is good evidence 2, 7, and 8) encode more than one pro- that shipping is an important means of tein by using alternative reading frames, its transmission. so that eleven proteins are produced. On Murray AG et al (2002) Emerg Infect Dis 8 , 1 the basis of the gene sequences, for influenza A virus the segments 1–3 encoded Infectious spleen and kidney necrosis PB2, PB1, and PA proteins are estimated 3 3 virus (ISKNV) The type species of to have mol. wt. 87 1 0 , 96 1 0 , 3 the genus Megalocytivirus in the fam- and 85 1 0 , respectively. Segment 2 ily Iridoviridae. Isolated from manda- encodes PB1, and in an alternate reading 3 rin fish and red sea bream. Infection frame PB1-F2. mol. wt. 10.5 1 0 . The is characterized by the formation of segment 4 encoded (unglycosylated) 3 inclusion body-bearing cells, which HA is 63 1 0 (glycosylated HA1 is 3 3 appear in the spleen, hematopoietic 48 1 0 , HA2 is 29 1 0 ). The seg- 3 tissue, gills, and digestive tract. Many ment 5 encoded NP is 56 1 0 . The 3 species of fish are susceptible includ- segment 6 encoded NA is 60 1 0 . ing mandarin fish (Siniperca chuatsi ), The segment 7 encoded M1 and M2 pro- 3 3 red sea bream (Pagrus major ) grouper teins are 28 1 0 and 11 1 0 , respec- (Epinephelus spp ), yellowtail (Seriola tively. The segment 8 encoded NS1 and 3 3 quinqueradiata), striped beakperch NS2 (NEP) are 27 1 0 and 14 1 0 , (Oplegnathus Fasciatus), red drum respectively. Antigenic variation occur- (Sciaenops ocellata) and African lamp- ring within the HA and NA antigens eye ( Aplocheilichthys Normani ). The of influenzavirus A has been analyzed virus can cause significant mortality, in detail. Sixteen subtypes of HA and especially in aquaculture facilities in nine subtypes of NA are recognized for SE Asia. The double-stranded DNA influenzaviruses A, with minimal sero- genome of ISKNV has been completely logical cross-reaction between subtypes. sequenced. Additional variation occurs within subtypes. By convention, new isolates are He JG et al (2001) Virology 291 , 126 designated by their serotype/host/ spe- infectious units Rarely, single virus parti- cies/site of origin/strain designation/ cles or, in most cases, groups of virions and year of origin and (HA [H] and which plaque or titrate as units. The NA [N] subtype), e.g. A/tern/South number of virus particles is often very Africa/1/61 (H5N3). In humans, contin- much larger (up to 106 times) than the ual evolution of new strains occurs and number of infectious units. older strains apparently disappear from circulation. Antibody to HA neutralizes infectivity A measure of the ability of a infectivity. If NA antibody is present virus to replicate within its host’s cells. during multicycle replication it inhib- Usually determined by endpoint dilu- its virus release and thus reduces virus tion, or plaque titration, and expressed yield. Antibody to the amino terminus as infectious units. of M2 reduces virus yield in tissue culture. Epidemics of respiratory disease Influenzavirus A A genus of the family in humans during the twentieth century Orthomyxoviridae containing one spe- have been caused by influenzaviruses A cies, Influenza A virus . The members having the antigenic composition H1N1, Influenzavirus A 248

H2N2, and H3N2. Limited outbreaks of appears to be critical in determining respiratory disease in humans caused the host range and organ tropisms of by antigenically novel viruses occurred influenza viruses. In addition, interac- in 1976 in Fort Dix, New Jersey, USA tions between gene products determine when classical swine H1N1 viruses the outcome of infection. Interspecies infected military recruits, in 1997 in transmission apparently occurs in some Hong Kong when H5N1 viruses caused instances without genetic reassortment outbreaks in poultry and contempora- (e.g. H1N1 virus from swine to humans neous illnesses and deaths in humans, and vice versa, H3N2 virus from humans and in 1998 and 1999 when H9N2 to swine, and the recent transmission of viruses present in poultry caused illness H5N1 and H9N2 viruses from poultry in humans in China. Influenzaviruses to humans). In other cases interspecies A of subtype H7N7 and H3N8 (previ- transmission may involve RNA segment ously designated equine 1 and equine reassortment in hosts infected with 2 viruses) cause outbreaks of respira- more than one strain of virus each with tory disease in horses. Influenzaviruses distinct host ranges, or epidemic prop- A (H1N1) and (H3N2) have been iso- erties (e.g. 1968 isolates of H3N2 viruses lated frequently from swine. The H1N1 apparently were derived by reassort- viruses isolated from swine in recent ment between a human H2N2 virus years appear to be of three general cat- and a virus containing an H3 hemag- egories: those closely related to classi- glutinin; seal H7N7 virus probably was cal ‘ swine influenza’ and which cause derived by reassortment of two or more occasional human cases; those first avian influenzaviruses; and reassortment recognized in avian specimens, but of human H1N1 and H3N2 viruses in which have caused outbreaks isolated 1978 and 1989 led to human infections from epidemics in humans since 1977. by viruses with H1N1 or H1N2 surface H3N2 viruses from swine appear to con- proteins and four to six other genes of tain HA and NA genes closely related H3N2 origin). Laboratory animals that to those from human epidemic strains. may be infected with influenzaviruses Influenzaviruses A (H7N7 and H4N5) A include ferrets, mice, hamsters, and have caused outbreaks in seals, with guinea pigs as well as some small pri- virus spread to non-respiratory tissues mates such as squirrel monkeys. in this host. In two separate cases, H7N7 viruses were isolated from conjunctival Influenza A virus (FLUAV) A species infections of a laboratory worker and a in the genus Influenzavirus A . Type farm worker in 1980 and 1996, respec- species A/PR/8/34 (H1N1). All spe- tively. Pacific Ocean whales have report- cies of the genus share a common ribo- edly been infected with type A (H1N1) nucleoprotein antigen, the NP protein. It virus. Other influenza subtypes have is demonstrated by CF test or immuno- also been isolated from lungs of Atlantic diffusion and is found as part of Ocean whales in North America. the nucleocapsid or as soluble antigen. FLUAV (H10N4) has caused outbreaks The genome (mol. wt. 4.5 1 06 , 13.6 kb) in mink. All subtypes of HA and NA, is comprised of eight segments of lin- in many different combinations, have ear negative-stranded RNA which been identified in isolates from avian range in length from approximately species, particularly wild aquatic birds, 900 to 2500 nt. Conserved sequences chickens, turkeys, and ducks. Pathology are present at the 5 and 3 termini in avian species varies from inapparent (13 and 12 nt, respectively). The seg- infection (often involving replication ments encode 11 proteins: 1, PB2; 2, in, and probable transmission via , the PB1 and PB1-F2; 3, PA; 4, HA; 5, NP; intestinal tract) to virulent infections 6, NA; 7, M1 and M2; and 8, NS1 and (observed with subtypes H5 and H7) NS2 (nuclear export protein, NEP). The with spread to many tissues and high internal virion RNA transcriptase com- mortality rates. The structure of the HA plex includes the PB1, PB2, PA, and protein, in particular the specificity of its NP proteins. This is surrounded by the receptor-binding site and its cleavability matrix or M protein that underlies the by naturally occurring tissue protease(s) outer lipid bilayer envelope in which 249 influenza virus A equine

are inserted the HA (hemagglutinin), lion chickens were slaughtered in Hong NA (neuraminidase), and M2 proteins. Kong. The virus reappeared in 2003 The HA and NA are both species- and in China and since 2004 has spread to type-specific antigens, as demonstrated infect poultry worldwide. There have for HA by hemagglutination inhibition also been more than 200 deaths in per- (HI) or neutralization, and for NA by sons having close contact with infected enzyme-inhibition tests. In avian influ- birds; a fatality rate of some 60% of enza viruses, 15 HA subtypes and 9 NA those infected. A majority of these cases subtypes can be demonstrated, some of have occurred in Indonesia with 124 which occur in various combinations in confirmed cases in February 2008, 101 of influenza viruses from humans, horses, them fatal. Many countries are planning or pigs. To date, only three HA subtypes for a possible pandemic of influenza if (H1–H3) and two NA subtypes (N1 and the H5N1 virus acquires the ability to N2) have been found in human influ- spread directly from person to person. enza A viruses. Gene reassortment is This has occurred to a limited extent in frequent between influenza A viruses northern Sumatra, where a cluster of in mixed infections, and may result in eight cases occurred within an extended dramatic changes in antigenicity termed family in 2007, which fortunately died ‘ antigenic shift, ’ which may precipitate out. Infection of chickens and turkeys pandemic disease. In addition, all strains with virulent avian strains, usually undergo antigenic drift. Transmission is called fowl plague and of subtype H5 by airborne virus or direct contact caus- or H7, is a serious and commercially ing epidemic and sporadic respiratory important disease which varies from disease. Natural hosts are humans, pigs, a mild respiratory to a rapidly fatal horses, birds and occasionally nonhu- pneumonic disease with CNS involve- man primates, dogs, and cattle. Ferrets ment. Stress may increase susceptibil- are experimentally susceptible to mam- ity. Transmission is probably airborne. malian strains. Less susceptible are A vaccine can protect birds but may not mice, hamsters, and guinea pigs. Most be economically practical. Mice, ferrets, influenza strains replicate in eggs and and other mammals have been infected primary cultures of monkey, human, experimentally usually by i.c. inocula- and chick cells. Laboratory strains may tion. Can be propagated in eggs and in be adapted to grow in cell lines. primary fowl or monkey kidney cells. Most strains produce plaques in chick Banks J et al (1998) Arch Virol 143 , 781 Cox NJ and Subbarao K (1999) Lancet embryo fibroblasts. 354 , 1277 Synonym : avian plague virus; fowl plague virus. influenza virus A avian Strains of Peiris JS et al (2007) Clin Microbiol Rev 20 , Influenzavirus A whose natural hosts are 243 birds: fowl, quail, ducks, turkey, pheas- Webster RG and Kawaoka Y (1988) Crit Rev ant, etc. They have also been isolated Poult Biol 1 , 211 from aquatic mammals such as seals and dolphins. Isolates fall into one of influenza virus A equine Strains of 16 antigenic groups based on hemag- Influenzavirus A whose natural host glutination inhibition, and nine based is the horse. Two distinct antigenic on neuraminidase. Natural infection subtypes of HA have been found in is widespread in birds, particularly equines: H3 and H7. Antibodies are water fowl in which infection is water- also found in human sera. Causes res- borne, and usually silent and intestinal. piratory illness in horses and may be They may excrete the virus over pro- fatal in young animals. Causes inap- longed periods. Many strains of virus parent infection in ferrets and can be are present and may provide a source adapted to produce pneumonia in of new mammalian strains. In 1997 an mice. Replicates in eggs, primary cul- unusual avian virus, H5N1, which was tures of bovine, human, monkey, and highly pathogenic for birds, caused chick cells. Agglutinates horse, pig, 18 human cases of influenza with six calf, rhesus, fowl, human, and guinea deaths in Hong Kong. More than a mil- pig erythrocytes. influenza virus A equine 250

Synonym : equine influenza virus. globally. Inactivated influenza virus Webster RG and Guo Y (1991) Nature 351 , vaccines consequently include H1N1, 527 H3N2 and influenza virus B components (trivalent), with annual changes influenza virus A (H5N1) A highly viru- in one or more of these components to lent strain of avian influenza which in accommodate minor changes due to 1997 passed directly to a human child antigenic drift. Influenza virus is usu- in Hong Kong who died of influenza. ally isolated in the amniotic cavity of Later in the year a further 17 persons embryonated eggs or in primary mon- became infected with the virus, and key kidney cell cultures. Virus multipli- 6 of the 18 cases died. The virus was cation is observed by hemadsorption or present in poultry, and was only trans- hemagglutination of human or guinea mitted to humans by close contact with pig erythrocytes. Ferrets can be infected the birds, and as a result all chickens in experimentally and after adaptation, Hong Kong were killed, which ended pneumonia can be produced in mice the potential epidemic. However, H5N1 and hamsters. In monkeys, horses, influenza virus reappeared 6 years dogs, sheep, guinea pigs, and rats infec- later in China in 2003 and has spread tion is usually inapparent. O-phase to many countries worldwide, caus- (o riginal) or recently isolated strains ing illness and death in persons having replicate better in the amniotic than in close contact with infected birds. The the allantoic cavity, and agglutinate country hardest hit with the disease is human and guinea pig erythrocytes Indonesia, where 129 human cases had better than fowl cells. After adaptation been confirmed by 22 February 2008, to the allantoic cavity they are termed 105 of whom had died. Worldwide D-phase ( d erived) and then they agglu- there had been 366 cases, with 232 tinate fowl cells equally well. This phe- deaths. The potential for an influ- nomenon is termed O/D variation. enza H5N1 pandemic depends on the Cox NJ et al (2005) In Topley & Wilson ’ s potential for the virus to spread from Microbiology and Microbial Infections , vol. 1, human to human, which so far has not Tenth edition, edited by BWJ Mahy and V happened. ter Meulen. London: Hodder Arnold, p. 737 Wilson IA and Cox NJ (1990) Annu Rev influenza virus A hominis Strains of Immunol 8 , 737 Influenza virus A whose natural host is humans. The virus was first isolated influenza virus A porcine Strains of in 1933. Causes sporadic and epidemic Influenzavirus A whose natural host respiratory disease. The HA and NA is the pig. Antibodies reacting with antigens of the virus change slowly swine virus hemagglutinin are present (antigenic drift), but periodically a in many human sera, especially from radical change occurs (antigenic shift) older people, and it has been suggested due to gene reassortment with replace- that the pandemic of 1918–1919 in ment of the HA and/or NA gene by a humans was caused by a virus which new subtype. The human population spread from humans to pigs. Sporadic has only slight immunity to the new fatal influenza cases in pig farmers and strain, resulting in a pandemic. To date, other persons in close contact with pigs three HA subtypes (H1–3) and two continue to be reported. Causes pneu- NA subtypes (N1 and N2) have been monia in pigs especially when associ- found in epidemic human influenza ated with Haemophilus suis. Ferrets and viruses. Viruses isolated from 1933 to mice are also susceptible to infection. 1957 were H1N1 (swine-like); in 1957 Transmission is by airborne droplets. they were replaced by H2N2 viruses Human and avian influenza viruses (Asian); in 1968 these were replaced by may also infect pigs, and epidemics of H3N2 (Hong Kong); and H3N2 strains H1N1, H3N2, H1N2, and H2N3 sub- continue to circulate globally in the type viruses in swine are common in human population. However in 1977, several parts of the world. H1N1 (swine-like) viruses reappeared and have also continued to circulate Synonym : swine influenza virus. 251 initiation factor

Webby RJ et al (2007) Curr Top Microbiol antigenic variants of the genus, and Immunol 315 , 67 antibody to HEF protein neutralizes Wells DL et al (1991) J Am Med Assoc 265 , 478 infectivity. Differs from influenzavirus A in the virion having a lower den- influenza virus A/PR/8/34(H1N1) (FLUA) sity (1.17–1.19 g/ml). Isolated in the Type species of the genus Influenzavirus amniotic cavity of embryonated eggs. A, isolated from a case of influenza in Replicates in canine kidney cell line Puerto Rico in 1934. (MDCK), best in presence of trypsin Influenzavirus B (FLUBV) A genus of the (20 g/ml). Infections usually occur in family Orthomyxoviridae consisting of children. A similar virus is also reported one species: Influenza virus B , the mem- to infect swine in China. bers of which infect only humans. The Herrler G and Klenk H-D (1991) Adv Virus virus is defined by the possession of Res 40 , 213 ribonucleoprotein antigen B. Human Nakada S et al (1984) Virus Res 1 , 433 strains in this genus are described according to a convention which sets influenza virus D Not an influenza virus. out information in the following order: Synonym for Human parainfluenza virus (1) antigen B; (2) geographical region; (3) type 1 . isolation number; (4) year of isolation. Example: B/Yamagata/16/88. All Ingwavuma virus (INGV) A strain strains share a common NP and M of Manzanilla virus in the genus protein. There is no M2 protein, but an Orthobunyavirus. Isolated from pigs in analogous protein (NB) is encoded by Taiwan and Thailand, and from birds RNA segment 6. In contrast to influ- in South Africa, Nigeria and the Central enzaviruses A, no distinct antigenic African Republic. Also found in India subtypes are recognized for influen- and Cyprus. Mosquito-borne. Not zavirus B, and antigenic drift occurs reported to cause disease in humans. more slowly. There is no antigenic shift. INH-95551 virus A strain of Guanarito Influenzaviruses B can readily reassort virus in the family Arenaviridae . gene segments between each other, but no reassortment has been found with Inhangapi virus A possible species in the influenzaviruses A. The only natural genus Vesiculovirus . Isolated from phle- host is humans. Causes sporadic and botomine flies in the Amazon region of epidemic respiratory disease, usually Brazil. Not known to cause disease in milder than that caused by influenza A. humans. Ferrets can be infected experimentally causing mild disease or none at all. Inini virus (INIV) A strain of Manzanilla Strains have been adapted to produce virus in the genus Orthobunyavirus . pneumonia in mice. The virus is eas- Isolated from a bird, Pteroglossus ily propagated in eggs and in monkey aracari , in French Guiana. kidney cell cultures. Laboratory strains initiation The start of synthesis of grow with CPE in human amnion, pig, a polypeptide chain. The point at calf, and ferret cells. which translation of mRNA begins Influenzavirus C (FLUCV) A genus in is indicated by the presence of the the family Orthomyxoviridae, the mem- initiation codon, AUG. In eukaryo- bers of which naturally infect humans tic cells AUG binds methionyl-tRNA causing mild, sporadic respiratory and in prokaryotic cells it binds disease. The type species is influenza N -formyl methionyl-tRNA. C/California/78. Its genome has only initiation codon , start codon See seven segments. A single envelope pro- initiation . tein, the hemagglutinin-esterase-fusion (HEF) protein is present on the virion initiation factor One of the several pro- surface, but there is no neuraminidase. tein factors that function in the initia- All strains of the genus share a com- tion of protein synthesis. There are 10 mon ribonucleoprotein antigen, the NP or more in eukaryotic cells, designated protein. The HEF protein distinguishes eIF1, 2, 3, etc. Inkoo virus (INKV) 252

Inkoo virus (INKV) A strain of insertion sequence consist of inverted California encephalitis virus in the genus repeats. It is thought that IS elements Orthobunyavirus . Isolated from mosqui- may have been involved in the origin toes in Finland up to 70°N. Antibodies and evolution of true viruses. found in humans, cattle, hares, foxes, and other wild animals. A few cases of insertional inactivation Inactivation of febrile illness with rising antibodies are a gene by insertion of nucleotides into reported in humans. the coding sequence. innate immune response The first insertional mutagenesis Introduction of response to infection which occurs one or more nucleotides into DNA or before the lymphocyte-mediated adap- RNA to alter gene expression. tive immune response. Involves innate integrase (IN) A virus-induced enzyme defense mechanisms such as natural which cuts and joins DNA molecules at killer cells which induce apoptosis of specific sites, originally described for virus-infected cells. temperate bacteriophages. Involved in Synonym: nonadaptive immune the insertion of viral DNA into chromo- response. somal (host) DNA. In retroviruses, the Inner Farne virus (INFV) A serotype integrase is a product of the pol gene. of Great Island virus in the genus Katz RA and Skalka AM (1994) Annu Rev Orbivirus . Biochem 63 , 133 inoculation Introduction of virus into a integrated viral genome A viral genome host to initiate replication and/or to which is incorporated into the cellular stimulate an immune response. DNA and is replicated with it. inosine The ribonucleoside of integration The process of insertion of hypoxanthine. viral DNA into the host genome. It usually involves a virus-coded enzyme, inosiplex An antiviral drug. The 3:1 the integrase. The viral DNA is then molar complex of N-N- dimethylamino- replicated by the host nucleic acid rep- 2-propanol p -acetamidobenzoate and lication mechanism. inosine. A controlled double-blind study in human volunteers challenged with integrins A family of structurally related rhinovirus suggested that the drug heterodimeric receptors that mostly exerts significant effects when used bind to an RGD sequence of adhesive therapeutically. The clinical effective- proteins. Function as cell surface recep- ness of the drug may be due to its tors for many viruses, including the activity as an immunopotentiator. adenoviruses, papillomaviruses, filo- Synonym : isoprinosine. viruses, morbilliviruses. Ginsburg T and Glasky AJ (1977) Ann NY Wickham TJ et al (1993) Cell 73 , 309 Acad Sci 284 , 128 integrin α 4 β 7 A receptor for prions. Inoue–Melnick virus Synonym for suba- α β cute myelo-optico-neuropathy virus. integrin 5 1 A receptor for measles virus, and also for Ebola virus. insert A segment of foreign DNA cloned integrin α 6 β 4 A proposed receptor for into a bacterial plasmid or other gene human papillomaviruses. vector. integument Structural component of insertion sequences (IS elements) Small some virions situated between the cap- transposable genetic elements first sid and the envelope. detected in bacteria, ranging in size from 0.7 to 1.8 kb. Can insert into several intercalation The insertion of planar mol- sites in the host genome and can cause ecules such as acridine dyes between transposition of the gene segments the adjacent base pairs of double- which they flank. The termini of the stranded DNA or double-stranded 253 interferons

RNA. Intercalating agents can inhibit bacteria, mycoplasma, and rickettsiae; the replication and transcription of to bacterial endotoxins, helenine, stato- DNA and cause a frameshift mutation. lon, phytohemagglutinin, and nucleic acids. They are a new class of proteins intercellular adhesion molecule-1 which are not ordinarily present in the (ICAM1) A cell surface protein that uninduced cell. Once released, interfer- serves as the receptor for human rhino- ons act upon other cells to reduce their viruses. The crystallographic structure susceptibility to a wide range of virus of the rhinovirus–ICAM-1 complex has infections. Interferons are grouped into been analyzed. six structurally distinct types: α (166 amino acids), β (166 amino acids), ω Olson NH et al (1993) Proc Natl Acad Sci γ 90 , 507 (172 amino acids) (146 amino acids), κ (180 amino acids), and the recently interference Prevention of the replication described interferon-λ . Interferons- of one virus by another, the result of a α and - β are also referred to as Type 1 number of different mechanisms. Thus, interferons, and interferon γ as Type II virus attachment to cell receptors can interferon. Interferon-α , produced by be prevented by prior exposure to any leukocytes, is a monomer with some virus which alters or destroys them. An 22 different subtypes; interferon-β , example of this is the interference of produced by fibroblasts, functions as UV-irradiated Newcastle disease virus a dimer; and interferon- (also called with the replication of an infective ‘ immune ’ interferon), produced by preparation of the same virus added lymphocytes, functions as a tetramer later. This is also an example of homol- molecule. Interferons-α and -β have an ogous interference, in which a virus antiviral action. After release from an interferes with its own replication. infected cell they bind to a receptor on Interference which is strictly homolo- other cells which results in the induc- gous cannot be due to interferon which tion of interferon-regulated proteins is active against a range of viruses. which inhibit virus replication directly Auto-interference is said to occur where or indirectly. Interferon- binds to a a large dose of virus produces a smaller different receptor and is an immuno- yield than a small dose, or fails to kill regulatory protein (mitogen), enhancing an animal whereas a small dose will. macrophage production and activating This may be due to interferon or some B cells and NK cells. With few excep- other inhibitor in the inoculum, but is tions, interferon action is limited to cells generally due to the presence of non- of the animal species in which the inter- infective (defective) particles which ferons were produced, or to those of block intracellular replication of infec- species closely related to the producer. tive ones. Heterologous interference is Interferons do not prevent virus pene- observed between different virus spe- tration of the cell and have no effect on cies and is most often due to interferon extracellular virus. Interferon inducers production. It can also be due to attach- have been administered intranasally ment interference, or to some blocking to human volunteers experimentally of virus replication which is poorly infected with rhinovirus or influenza understood and probably varies from virus but only minimal amounts of one case to another. Examples of heter- nasal interferon were induced and ologous interference are frog virus 3 or the overall results were unimpressive. adenovirus with vaccinia virus. However, rhinovirus infection has been prevented by intranasal instillation of interferons A group of related host-specific human leukocyte interferon. A variety nonviral proteins of differing mol. wt. of genetically engineered interferons 150,00–300,00, liberated by cells fol- are now available for therapeutic use. lowing exposure to a variety of induc- Guidotti LG and Chisari FV (2000) Virology ing agents. Viruses are most potent in 273 , 221 this respect, but interferons can also be Sen GC and Ransohoff RM (1993) Adv Virus induced by the exposure of cells to other Res 42 , 57 microorganisms including protozoa, Staeheli P (1990) Adv Virus Res 38 , 147 interferon- α (IFN- α ) 254 interferon-α (IFN-α ) A non-glycosylated STAT pathway. Signal transduction protein of 166 amino acids produced occurs by the activation of a family of by lymphoid cells and macrophages. tyrosine kinases (Janus kinases (JAK)) Induces an antiviral state. that are activated when the interferon receptors are occupied. The activated α α interferon -2a A recombinant inter- JAKs phosphorylate a family of tran- feron consisting of 165 amino acids scription factors (signal transduction with lysine at position 23 and histidine activators of transcription (STAT)) at position 34. which migrate from the cytoplasm α α to the nucleus. Here they form active interferon -2b A recombinant inter- complexes which bind to ISRE present feron consisting of 165 amino acids in the promoter sites upstream of inter- with arginine at position 23 and histi- feron-inducible genes, activating their dine at position 34. transcription. interferon α -2c A recombinant α inter- Aaronson DS and Horvath CM (2002) feron consisting of 165 amino acids Science 296 , 1653 with arginine at positions 23 and 34. interferon-ω (IFN- ω ) An N -glycosylated interferon-β (IFN-β ) An N -glycosylated protein of 172 amino acids produced protein of 166 amino acids produced by leukocytes and trophoblasts. by fibroblast and epithelial cells. intergenic complementation See interferon-γ (IFN-γ ) An N -glycosylated complementation . protein of 146 amino acids produced by T cells, macrophages and NK cells. intergenic regions Non-coding sequences between the genes of a virus genome. interferon-γ -inducible protein (IP-10) A These may be quite short, but in some chemokine which is specific for acti- viruses, e.g. the Mononegavirales, there vated T lymphocytes, upregulating may be several hundred nucleotides T-cell cytokine synthesis. between the coding sequences. Their function is not well understood. interferon-γ (IFN-γ ) receptor Orthopox- virus-infected cells secrete an IFN-γ interjacent RNA 26S Single-stranded receptor which acts as an IFN- γ decoy RNA found in cells infected with certain binding protein which sequesters Togaviridae, such as Semliki Forest virus. IFN-γ and prevents it binding to cell It has the same polarity as the viral RNA receptors. but shares only 1/3 of its base sequences from the 3 end; it is thus termed sub- interferon-κ (IFN-κ ) A protein of 180 genomic RNA. Interjacent RNA codes amino acids produced by keratinocytes for all the viral coat proteins. and dendritic cells. interleukins A variety of substances pro- interferon-λ (IFN-λ ) A recently described duced by leukocytes which function group of glycosylated interferons during inflammatory responses. (lambda1/2/3) produced in most tissues that seem to activate innate interleukin-1 (IL-1) Includes two proteins, immune responses at the site of viral IL-1α and IL-1 β of 152 amino acids infection. secreted by macrophages. Produced by separate genes in the same cluster. interferon regulatory factor-3 (IRF-3) A Involved in the activation of both B- cellular protein which is activated by and T lymphocytes in response to anti- virus infection to induce expression of gens. A key cytokine expressed during interferon-β. inflammation. interferon-specific response elements interleukin-1β (IL-1 β ) A cytokine induced (ISRE) Binding of interferons to their following infection with herpes- receptors initiates signals that are trans- viruses, HIV-1, adenovirus and chronic mitted to the cell nucleus via the JAK/ hepatitis B. 255 interleukin-16 (IL-16) interleukin-2 A lymphokine produced phagocytes, and activated B cells. A naturally by helper T cells follow- gyokine synthesis inhibitory factor ing T-cell recognition of class II MHC which inhibits the production of IFN-γ (associated) antigen, and by mitogens. and secretion of IL-1, IL-6, and TNF-α Activates growth and proliferation of by macrophages. An EBV open reading T lymphocytes. Structurally similar to frame, BCRF1, shows extensive homol- interferon-γ. ogy with IL-10, and viral IL-10 expression may enable infected cells to escape interleukin 3 A hemopoietic growth fac- from immune surveillance. tor derived from T cells that stimulates proliferation and differentiation of Moore PS et al (1996) Science 274 , 1739 bone marrow pluripotential stem cells. interleukin-11 (IL-11) A hemopoietic interleukin-4 (IL-4) A 20-kDa glycopro- growth factor with growth and prolif- tein, B-cell growth and differentiation erative effects on stem cells. Made by factor secreted by Th2 cells which plays fibroblasts. an essential role in humoral immunity. It activates B cells very early in growth interleukin-12 (IL-12 ) A heterodimeric and allows B cells activated by antigen cytokine made by macrophages which to move into the G1 phase of the cell activates NK cells and T cells. One cycle. of its chains has high homology with IL-6 and the other with the IL-6 receptor. interleukin-5 (IL-5) A Th2 cell-derived It causes proliferation and enhances cytokine that induces activation and the cytotoxicity of both NK cells and differentiation of eosinophil leukocytes cytotoxic T lymphocytes. It polarizes and differentiation of B cells to become the immune response toward Th1 cells antibody-secreting cells. by enhancing IFN-γ production and inducing macrophage activation. It is interleukin-6 (IL-6) A cytokine important thus a protective cytokine against virus in inflammatory reaction. Produced by infections. T and B cells, macrophages, fibroblasts, and endothelial cells. Induces B cells to interleukin-13 (IL-13) An IL-4-like differentiate into antibody-producing cytokine, secreted by activated T cells, cells, Both RNA and DNA viruses which activates B-cell growth early in induce IL-6 expression, and virally- the cell cycle. induced IL-6 participates in inflammatory reactions together with IL-1 and α interleukin-14 (IL-14) A B-cell growth TNF- . factor that induces proliferation of acti- interleukin-7 (IL-7) A protein of 177 vated B cells. amino acids secreted by thymic stroma interleukin-15 (IL-15) A cytokine pro- cells that acts as a T-cell growth and duced by many cell types which is an differentiation factor and a macrophage activator of T-cell proliferation and activation factor. effector function. IFN-α / β mediated interleukin-8 (IL-8 ) An α-chemokine induction of IL-15 that promotes sur- made by macrophages and endothelial vival of activated NK cells and their cells following inflammatory stimula- accumulation. tion. Activates neutrophil functions such as adhesion and microbiocidal interleukin-16 (IL-16) A cytokine activity. released by a variety of cells, including lymphocytes and some epithelial interleukin-9 (IL-9) A cytokine derived cells, that is a chemoattractant for cells from activated T cells that supports expressing the CD4 molecule including growth and proliferation of T-cells and monocytes, eosinophils, and dendritic bone-marrow derived mast cells. cells. CD4 is the cell signaling receptor for mature IL-16. Contributes to CD4- interleukin-10 (IL-10 ) A cytokine pro- cell recruitment and activation at sites duced by T lymphocytes, mononuclear of inflammation. interleukin-16 (IL-16) 256

Cruikshank WW et al (2000) J Leukoc Biol interleukin-23 (IL-23) A heterodimeric 67 , 757 cytokine, one subunit of which (p40) is shared with IL-12. In conjunction with interleukin-17 (IL-17 ) The founding IL-6 and TGF-β , IL-23 stimulates naive member of a group of six cytokines, CD4 T cells to differentiate into a novel called IL-17A to IL-17F. All have a simi- subset of cells called Th17 cells. These lar protein structure which differs from cells produce IL-17, a proinflammatory all other known cytokines. Secreted by cytokine enhances T-cell priming. activated T cells. Involved in inducing and mediating proinflammatory interleukin-24 (IL-24) A cytokine related responses in many of the tissues of the to IL-10 that signals through two hetero- body. Herpesvirus saimiri gene 13 is a dimeric receptors, IL-20R1/IL-20R2, homolog of IL-17. and IL-22R1/IL-20R2. Controls cell Moseley TA et al (2003) Cytokine Growth survival and proliferation by inducing Factor Rev 14 , 155 rapid activation of STAT-1 and STAT-3. Released by activated monocytes, interleukin-18 (IL-18) A pro-inflammatory macrophages, and Th2 cells, and acts cytokine that is required, together on skin, lung, and reproductive tissues. with IL-12, for induction of IFN-γ and the generation of an efficient cellular interleukin 25 (IL-25) A cytokine belong- response against viral infection. Several ing to the IL-17 family (also known as poxviruses, including molluscum conta- IL-17E) secreted by Th2 cells and mast giosum virus, encode an IL-18 binding cells. An important molecule control- protein which downregulates IFN-γ pro- ling immunity in the gut. duction and NK cell responses. The viral binding protein has no homology with interleukin 26 (IL-26) A protein of 171 normal cell membrane IL-18 receptors. amino acids related to IL-10. Expressed in herpes virus-transformed T cells. interleukin-19 (IL-19)A cytokine expressed Signals through a receptor complex of by resting monocytes and B cells which IL-20 R1 and IL-10 R2. Induces rapid is upregulated in monocytes following phosphorylation of STAT-1 and STAT- stimulation with granulocyte-macrophage 3, which enhances IL-8 and IL-10 secre- colony stimulating factor (GM-CSF) or tion and expression of CD54 on the lipopolysaccharide. Has a helical crystal surface of epithelial cells. structure. interleukin 27 (IL-27) A heterodimeric Chang C et al (2003) J Biol Chem 278 , 3308 cytokine belonging to the IL-12 family. Composed of two subunits: interleukin-20 (IL-20) A cytokine pro- EBV-induced gene 13 and IL27-p28. duced by activated keratinocytes and Functions in the regulation of the activ- monocytes which belongs to the IL-10 ity of B- and T-lymphocytes. family. Causes proliferation of keratinocytes during inflammation, espe- interleukin 28 (IL-28) A cytokine in two cially of the skin. isoforms, IL-28A and IL-28B, which belongs to the interferon type III family interleukin-21 (IL-21) A cytokine that has of cytokines. Similar to IL-29. Important potent regulatory effects on NK cells in immune defense against viruses. and cytotoxic T cells that can destroy virally infected cells. interleukin 29 (IL-29) A protein of the helical cytokine family which is an interleukin-22 (IL-22) A cytokine with interferon type III. homology to IL-10, derived from T cells. Activates transcription fac- interleukin 30 (IL-30 ) A 28-kDa protein tors STAT-1 and STAT-3 in several which forms one chain of IL-27. hepatoma cell lines. Signals through the interferon receptor related proteins interleukin 31 A cytokine with a four- CRF2-4 and the IL-22 receptor. helix bundle structure produced by Th2 257 intraneuronal spread

cells. Signals via a receptor composed of the Union occur every 3 years. An of IL-31 receptor A and oncostatin important committee of the Virology M. Plays a role in inflammation of Division is the International Committee the skin. on Taxonomy of Viruses. Stacey R et al (2004) Nat Immunol 5 , 752 intestine 407 cells (CCL 6) A cell line derived from the jejunum and ileum of interleukin 32 A cytokine that can a 2-month-old Caucasian embryo. induce monocytes and macrophages to α secrete TNF- in addition to IL-8 and intracellular adhesion molecule 1 MIP2/CXCL-2. (ICAM-1) The receptor for rhinoviruses which belongs to the immunoglobulin interleukin 33 (IL-33) A cytokine belong- superfamily of proteins. ing to the IL-1 family. Induces helper T cells to produce type 2 cytokines. intercellular bridges In some virus infections the entry of viruses into cells can interleukin 34 (IL-34) A cytokine discov- be short-circuited, so that a transient ered in May 2008 which increases the intercellular bridge is formed. These viability of primary monocytes without bridges permit the viral genome to pass affecting other cell types. Structurally from cell to cell without having to sur- unrelated to any other cytokine. vive in the extracellular environment and so avoiding neutralizing antibody. interleukin 35 (IL-35) An anti-inflammatory This probably occurs in subacute scle- cytokine which suppresses the immune rosing panencephalitis virus (SSPE) response through the expansion of regu- where the measles virion spreads grad- latory T cells and suppression of Th17 ually from neuron to neuron despite cell development. high antibody levels in the extracellu- Niedbala W et al (2007) Eur J Immunol 37 , lar fluid of the brain parenchyma. 3021 intracellular enveloped virus During the intermolecular recombination Recombi- replication of poxviruses mature envel- nation due to the reassortment of spe- oped virus particles are formed inside cies of nucleic acid between viruses the cell, then acquires two additional whose genomes are segmented, e.g. seg- membranes by wrapping in Golgi- ments of RNA of reoviruses, influenza derived cisternae then fusing with the virus. Also termed genetic reassort- plasma membrane to become cell-asso- ment. See phenotypic mixing virus . ciated enveloped virus. The structure and biogenesis of poxviruses thus differs markedly from helical or icosahe- internal promoter A promoter located dral viruses. upstream from the first promoter on the virus genome. intracisternal R-type particles Endo- genous retroviruses detected by elec- internal ribosomal entry site (IRES) A tron microscopy in BHK cells. sequence within the noncoding region of positive-strand RNA viruses, espe- intramolecular recombination See cially picornaviruses, which specifies recombinants . ribosomal binding and initiation of protein synthesis. intramuscular immunoglobulin (IMIG) A generally effective technique for pre- International Union of Microbiological exposure prophylaxis against hepatitis Societies (IUMS) The major gov- A. The availability of a killed hepatitis A erning body of microbiological sci- virus vaccine has reduced the need for ences, supported by more than 100 IMIG especially in developed countries. societies worldwide. Affiliated to the International Union of Science. It con- intraneuronal spread The movement sists of three Divisions, of Bacteriology, of viruses such as herpes simplex or Mycology, and Virology. Meetings rabies through neurons. intrapartum infection 258 intrapartum infection Infection occur- reverse orientation at the ends of the ring during childbirth, or during genome. Found in herpesviruses, pox- delivery. Can occur with HIV-infected viruses, and transposons. mothers, with significant infant mortality, but the transmission can be sig- Invirase Synonym for Saquinavir. nificantly reduced by administration of 5-iodo-2 ’ -deoxyuridine See idoxuridine . antiretroviral therapy to mothers and their HIV-exposed infants. In addition, Iopaka virus An unclassified arbovirus intrapartum infection may occur with isolated in the Amazon region of herpes simplex 2 and hepatitis B virus Brazil. infections. Iotapapillomavirus A genus of the fam- intrathecal antibodies Virus persistence ily Papillomaviridae containing rodent in the CNS is usually accompanied by papillomaviruses causing cutaneous long-lasting intrathecal antibody syn- lesions in the host. The E5 open read- thesis with specificity for viral proteins. ing frame is absent from the genome. Selection and recruitment of these anti- The type species is Mastomys natalensis bodies to the CNS can be detected in papillomavirus . the CSF as a restricted oligoclonal pat- Ipixaia virus A possible species in the tern that can be used as a diagnostic genus Orbivirus, isolated from phle- marker. botomine sandflies in the Amazon Felgenhauer K and Reiber H (1992) Clin region of Brazil. Antigenically related Invest 70 , 28 to Changuinola virus. Not known to cause disease in humans. intrauterine transmission Transmission of a viral infection within the uterus. Ippy virus (IPPYV) A species in the genus May occur with rubella virus, human Arenavirus. Isolated in suckling mice parvovirus, human herpesviruses such from the pooled liver, spleen, and brain as cytomegalovirus and varicella-zoster of an adult rat of Arvicanthis sp trapped virus, and hepatitis B and C viruses, in Ippy, Central African Republic. Not all of which may have serious conse- known to cause disease in humans. quences for the unborn infant. IPV Inactivated poliovirus vaccine (Salk Enders G (2005) In Topley & Wilson ’ s Micro- vaccine). biology and Microbial Infections , vol. 2, Tenth edition, edited by BWJ Mahy and V ter Iridoviridae A family of large DNA- Meulen. London: Hodder Arnold, p. 1443 containing viruses, 120–200 nm in diam- Haun L et al (2007) Minerva Ginecol 59 , 159 eter, having icosahedral symmetry. Virions contain many proteins, sev- intravenous immunoglobulin (IVIG) A eral enzymes, lipid and a single struc- treatment used to prevent virus infec- tural unit membrane associated with tions in patients with primary immu- the viral core. The genome is a single nodeficiency disorders associated with molecule of double-stranded DNA, low serum IgG levels. 140–300 kb in length, G C content 28–54%, which is circularly permuted intrinsic interference See interference . with direct terminal repeats. In vertebrate iridoviruses but not insect irido- intron A region of the genome nucleic viruses the DNA is highly methylated. acid of a virus or a cell which, follow- Replication occurs in the cytoplasm. ing transcription, is lost from mature There are four genera: Iridovirus and mRNA during processing – an inter- Chloriridovirus (both of insects); Ranavirus vening sequence. See also exon and (frogs); and Lymphocystivirus (fish). splicing . Synonyms: icosahedral cytoplasmic Gilbert W (1978) Nature 271 , 501 deoxyribovirus; polyhedral cytoplasmic deoxyribovirus. inverted terminal repeats Short regions Essani K and Granoff A (1989) Intervirology of identical sequence that occur in 30 , 187 259 isoforms

Iridovirus A genus of the family in gerbils but not in domestic animals. Iridoviridae which consists of species The ecology of the virus may thus be infecting arthropods. The type species different from that of other serotypes is Invertebrate iridescent virus 6 . and involve chiefly humans, gerbils, and sandflies. Not known to cause dis- Iriri virus An unclassified arbovirus iso- ease in humans. lated from phlebotomine sandflies in the Amazon region of Brazil. Tesh R et al (1977) Am J Trop Med Hyg 26 , 299 ISG20 A human interferon-induced gene Irituia virus (IRIV) A strain of Changuinola which codes for a 3 -5 exonuclease virus in the genus Orbivirus . Isolated specific for single-stranded RNA, and from rodents of Oryzomys sp in Para, interferes with vesicular stomatitis Brazil. Not reported to cause disease in virus, influenza virus, and encephalo- humans. myocarditis virus expression, but Irkut virus A tentative species in the had no effect on a DNA virus, human genus Lyssavirus . Isolated in Irkutsk, adenovirus. East Siberia, from the brain of a greater Espert L et al (2003) J Biol Chem 278 , 16151 tube-nosed bat ( Murina leucogaster ) captured in September 2002. Caused Ishak score A numerical system devel- fatal encephalitis in mouse brain. The oped to evaluate chronic hepatitis nucleotide sequences of the nucleo- caused by hepatitis viruses B and C. capsid, phosphoprotein, and glycopro- Provides semi-quantitative grading tein genes showed that Irkut virus is of necroinflammation and staging of a new genotype within the Old World fibrosis based on a liver biopsy. bat lyssaviruses. Ishak KG (1994) Mod Pathol 7 , 690 Arch Med Res 38 Kuzmin IV et al (2005) Virus Res 111 , 28 Lefkowitch JH (2007) , 34 Isiolo virus A strain of Sheeppox virus . Iruana virus A strain of Changuinola virus in the genus Orbivirus. Isolated from Isla Vista virus (ILV) A species in the phlebotomine sandflies in the Amazon genus Hantavirus, isolated from voles, region of Brazil. Not known to cause Microtus californicus , in California, USA. disease in humans. ILV, which is closely related to Prospect Hill virus, has not been associated with Isavirus A genus within the disease in humans or rodents. Orthomyxoviridae containing a single species, Infectious salmon anemia virus . Song W et al (1995) J Gen Virol 76 , 3195 Morphologically similar to influenza isoelectric focusing (electrofocusing) A viruses, with surface projections of separation technique in which mixtures 10 nm. Virions have both hemaggluti- of proteins and/or viruses are resolved nating and receptor destroying activi- into their components by subjecting ties, the latter being an acetyl esterase them to an electric field in a supporting activity. The genome has eight linear gel or stabilized solution in which a pH negative-stranded RNA segments with gradient is established. The proteins or a total size of 13.5 kb. viruses migrate to the positions in the Isentress Trade name for raltegravir. gel which have a pH equivalent to their isoelectric points. Isfahan virus (ISFV) A species in the isoelectric point The pH value of a solu- genus Vesiculovirus . Antigenically tion in which a given macromolecule related to vesicular stomatitis virus. (usually a protein or virus) does not Isolated from the sandfly, Phlebotomus move in an electric field. At this pH the papatasi, in Iran. Animal pathogenicity, net surface charge is zero. growth rate, morphology, CPE, and plaque morphology are similar to those isoforms Different chemical forms of a of other serotypes. Neutralizing anti- substance with the same activity, such bodies are common in humans in sev- as isoenzymes. In Borna disease virus eral regions of Iran. They are also found two isoforms of the nucleoprotein exist isoforms 260

as either 40 kDa or 38 kDa. Whereas p40 USSR. Antibodies are found in humans uses the entire ORF, p38 initiates at the but the virus is not reported to cause second AUG codon and lacks 13 amino disease. acids at the amino terminus. Synonym : Keterah virus. isolate A virus propagated in pure culture Itaboca virus A possible species in the as far as is known. May later prove to genus Orbivirus, isolated from phle- be a mixture. Often used incorrectly to botomine sandflies in the Amazon describe what would be better known region of Brazil. Antigenically related as a recognizate . to Changuinola virus. Not known to cause disease in humans. isometric particle Particles with identical linear dimensions, distinct from the Itacaiunas virus An unclassified arbo- rod-shaped and bullet-shaped virus virus isolated from a pool of Culicoides particles and viruses enclosed by irreg- midges in the Amazon region of ular capsules. They appear spherical; Brazil. Not known to cause disease in however, their capsids are constructed humans. with icosahedral symmetry. Itaituba virus (ITAV) A strain of Candiru Isoprinosine Trade name for inosiplex. virus in the genus Phlebovirus . Isolated from an opossum, Didelphis marsupialis isopycnic gradient centrifugation A form in Brazil. Not associated with disease of density gradient centrifugation in in humans. which the supporting gradient column includes the entire range of densities Itakura virus An isolate of Sagiyama of the particles to be tested. In this case, virus, a serotype of Ross River virus sedimentation of an individual particle in the genus Alphavirus. Isolated in will cease when it reaches that point Japan. in the gradient matching its own density, i.e. the point of buoyant density. Separation of particles into zones by this Itaporanga virus (ITPV) A tentative spe- technique depends only upon their den- cies in the genus Phlebovirus . Isolated sity differences and is independent of from mosquitoes of Culex sp. Natural time. See also rate zonal centrifugation . hosts inhabit the forest canopy, e.g. bats and birds. Forest floor animals are not isotype An antigenic determinant shared infected. Found in São Paulo, Amapa by all individuals of a given species but and Para, Brazil, and in Trinidad and absent in individuals of other species. French Guiana. Not reported to cause disease in humans. Israel turkey meningo-encephalitis virus (ITV) A species in the genus Flavivirus . Itapua virus A probable species in the Isolated from domestic turkeys in genus Hantavirus , identified from the Israel. Causes a progressive paralysis black-footed colilargo (Oligoryzomys with meningo-encephalitis leading to nigripes) in eastern Paraguay. Not asso- 10–12% mortality. An attenuated vac- ciated with human disease. cine is commercially available, and in enzootic areas such as Israel is given by Itaqui virus (ITQV) A strain of Oriboca i.m. injection at 8 weeks. Not associated virus in the genus Orthobunyavirus and a with human disease. member of the group C viruses. Isolated from sentinel cebus monkeys and mice, Malkinson M (1993) In Virus Infections of Birds , edited by JB McFerran and MS forest rodents and marsupials in Para, McNulty. Amsterdam: Elsevier Science Pub- Brazil. Mosquito-borne. Causes CPE lishers, p. 239 in HeLa cell cultures. Not reported to cause disease in humans. Issyk–Kul virus (IKV) An unassigned virus in the family Bunyaviridae . Isolated Iteravirus A genus of invertebrate parvo- from bats and ticks in Kirghiz, former viruses in the subfamily Densovirinae . 261 IUMS

Itimirim virus (ITIV) A strain of Bertioga IUBS International Union of Biological virus in the genus Orthobunyavirus . Sciences. Isolated from a rodent, Oryzomys sp. IUdR See idoxuridine . Itupiranga virus (ITUV) A tentative species in the genus Orbivirus . Isolated IUMS See International Union of from mosquitoes in the Amazon region Microbiological Societies . of Brazil. Not known to cause disease in humans. J

J virus A probable species in the genus transmitted by a cell-free filtrate or Paramyxovirus. Isolated by kidney cell tumor extracts containing reverse tran- culture from moribund wild mice, scriptase activity associated with par- Mus musculus, with hemorrhagic lung ticles with a buoyant density typical of lesions. Replicates with CPE in a variety retroviruses. The viral nucleic acid is dis- of cell cultures including MRC5, Hep- tinct from that of Visna/maedi , which is a 2, BHK21, and Vero but not Hela cells. lentivirus of sheep. It now appears that Does not hemagglutinate. Antibodies the envelope glycoprotein of JSRV (Env) present in wild mice, rats, pigs, bovines, functions as a dominant oncoprotein and humans. On infection via the nose both in vitro and in vivo . It has been pos- or s.c. injection, rats and mice became sible to eradicate the disease from sheep viremic and lethargic, produced anti- flocks by motherless rearing of lambs; bodies and developed hemorrhagic over a 3-year period the lambs were interstitial pneumonia. Appears to taken away from their mothers at birth, be a natural respiratory pathogen of deprived of maternal colostrum, and wild mice in Northern Queensland, hand reared away from other sheep. Australia. Leroux C et al (2007) Vet Res 38 , 211 Jun MH et al (1977) Aust J Exp Biol Med Sci Varela M et al (2008) Virology 371 , 206 55 , 645 Jacareacanga virus (JACV) A serotype of J 111 cells (CCL 24) A heteroploid cell line Corriparta virus in the genus Orbivirus . derived from peripheral blood of a 25- Isolated from the mosquito, Culex ( Mel ) year-old human female with monocytic sp, in Flexal area, Brazil. leukemia. Jacunda virus A possible species in the Jaagsiekte sheep retrovirus (JSRV) A spe- genus Phlebovirus, isolated from a cies in the genus Betaretrovirus. Infects rodent Mioprocta acouchy in the Amazon mainly domestic sheep worldwide region of Brazil. except for Australasia. Causes a contagious, slow progressive lung disease Janus acivated kinase (JAK) An enzyme which pathologically appears as an involved in activation of the interferon adenomatosis. There are multiple pul- response. monary adenocarcinomas which sometimes metastasize. There are similarities JAK/STAT pathway During interferon to human bronchoalveolar cell carcino- induction, JAK kinases participate in mas. The disease develops slowly and the tyrosine phosphorylation of tran- is most common in the third year of scriptional factors called STAT (s ignal life. Animal signs are respiratory dis- t ransducers and a ctivators of t ranscrip- tress, loss of appetite and coughing. tion). Phosphorylated STAT proteins Lambs less than 6-months old may die are transported to the nucleus where in a few days, but in older sheep the dis- they interact with interferon-respon- ease is chronic, lasting months to years. sive elements to activate the interferon Secondary infections, such as Pasteurella , response. are frequent complications. In the termi- Aaronson DS and Horvath CM (2002) nal stages there is copious secretion of Science 296 , 1653 tracheobronchial fluid. The virus has not Levy DE (1995) Semin Virol 6 , 81 been cultivated in cell culture; it can be transmitted to goats but is not endemic Jamanxi virus (JAMV) A strain of in that species. The disease can be Changuinola virus in the genus 263 JC polyomavirus (JCPyV)

Orbivirus . Isolated from Lutzomyia sp and China an attenuated vaccine is in Para, Brazil. Not associated with dis- now licensed and is very effective. The ease in humans. attenuated vaccine in Japan is used only for pigs. Jamestown Canyon virus (JCV) A sero- Synonyms: Japanese B virus; Russian type of California encephalitis virus in autumn encephalitis virus. the genus Orthobunyavirus . Isolated Barrett ADT (2008) in BWJ Mahy and MHV from mosquitoes, horse, and deer-flies van Regenmortel (eds) Encyclopedia of in Wisconsin and Colorado, USA and Virology, Third edition, Oxford: Academic in Canada. Serological studies suggest Press, vol. 3, p. 182 the white-tailed deer, Odocoileus virginianus, is an important host in Maryland, Japanese flounder nervous necrosis virus USA. Shown to cause widespread (JFNNV) A tentative species in the infection and occasional encephalitis in genus Betanodavirus, isolated from humans. juvenile fish in a commercial hatchery in Japan. Jandiá virus A possible species in the genus Orbivirus, isolated from sand- Japanese hemagglutinating virus See flies in the Amazon region of Brazil. Sendai virus . Not associated with disease in humans. Jari virus (JARIV) A serotype of Changuinola virus in the genus Japanaut virus (JAPV) A tentative spe- Orbivirus . Isolated from the sloth, cies in the genus Orbivirus . Isolated Choloepus didactylus , in Para, Brazil. from culicine mosquitoes and a bat in the Sepik district of New Guinea. Not Jatobal virus A possible species in the reported to cause disease in humans. genus Orthobunyavirus, isolated from the coati (Nasua nasua ) in the Amazon Japanese B virus Synonym for Japanese region of Brazil. Not associated with encephalitis virus . disease in humans. It is a reassortant virus containing one segment (S) Japanese eel herpesvirus Synonym for derived from Oropouche virus . anguillid herpesvirus 1. Jatuarana virus A possible species in the Japanese encephalitis virus (JEV) A spe- genus Orbivirus , isolated from phlebot- cies in the genus Flavivirus , a member omine sandflies in the Amazon region of the Japanese encephalitis serogroup. of Brazil. Antigenically related to Mainly an infection of birds spread by Changuinola virus . culicine mosquitoes, especially Culex tritaeniorhynchus, but infection of bats JC polyomavirus (JCPyV) A species in the may be a factor in winter survival. In genus Polyomavirus . Antigenically dis- humans it causes mild febrile illness tinguishable from BK virus and SV40 but a few cases show signs of encepha- virus. Agglutinates human group O litis. There may be paresis and sequelae erythrocytes. Originally isolated from are common. The case fatality rate is 5– the brain of a patient with progres- 40% in different outbreaks. Encephalitis sive multifocal leukoencephalopathy occurs in horses. Domestic pigs are (PML) by inoculation of homogenized often infected and have a viremia. Virus brain tissue into primary human fetal found in Asia from Siberia to Malaysia glial cell cultures. Has been clearly and southern India. Replicates in eggs, identified as the cause of PML, as well the best method of infection being into as renal disease and upper respira- the yolk sac. Replication also occurs in tory tract infections. Can be propa- cell cultures of chick and various mam- gated in primary human fetal glial malian tissues, as well as mosquito cell cultures which show a CPE in 3–4 cells. Causes encephalitis on i.c. injec- days. Presence of antibodies suggests tion of mice, hamsters, and monkeys. it is a common human infection with Inactivated vaccine is used in Japan, roughly 60% seroconversion by age Korea, Taiwan, and China. In Japan 12 years. The virus persists following JC polyomavirus (JCPyV) 264

primary infection as an infection of the the tumor tissue of a rat sarcoma initi- kidney, with excretion into the urine ated in 1907. from which it can readily be isolated. Immunosuppression due to AIDS or Jerry Slough virus A strain in the genus drug treatments later in life cause reac- Orthobunyavirus , belonging to the tivation and disease, including PML. California serogroup. Isolated from Highly oncogenic in newborn ham- the mosquito, Culiseta inornata, in Kern sters, especially on i.c. injection. The County, California, USA. Antigenically use of PCR to genotype JCV DNA has very closely related to Jamestown led to a genotyping method which can Canyon virus. Not known to cause dis- be used to trace human migrations. ease in humans. Chang D et al (1999) Arch Virol 144 , 1081 Major EO et al (1992) Clin Microbiol Rev 5 , 49 Jeryl Lynn B vaccine A strain of attenu- Sugimoto C et al (1997) Proc Natl Acad Sci ated mumps virus vaccine, obtained 94 , 9191 after multiple passages of the original virus, obtained from Jeryl Lynn, daugh- JC virus See JC polyomavirus . ter of vaccinologist Maurice Hillemann. Licensed in 1967. JD254 virus (DGKV) A serotype of Hilleman MR (1999) Immunol Rev 170 , 7 Dera Ghazi Khan virus in the genus Nairovirus . Isolated from ticks. JH virus A cytopathogenic agent isolated in the USA at the Johns Hopkins Jedda virus The original name for a spe- University from adults and children cies in the genus Flavivirus , which with mild respiratory disease, origi- caused a fatal hemorrhagic fever in nally classified as echovirus 28, then 1995 in Jedda, Saudi Arabia, now called reclassified as human rhinovirus 1 A. Alkhurma virus . JHM strain A strain of murine hepatitis Jembrana disease virus (JDV) A probable virus in the genus Coronavirus . A neu- species in the genus Lentivirus . A bovine rotropic variant. JHM indicates associ- lentivirus first recognized in the Bali ation with Professor J Howard Mueller district of Indonesia in 1964 where it of Harvard. causes an acute disease in Bali cattle, Bos Nagashima K et al (1978) Adv Exp Med Biol javanicus , leading to death in 1–2 weeks. 100 , 395 It multiplies to very high titers in vivo as Robb JA and Bond CW (1979) Virology 94 , 352 it has a Tat-1 protein which is a potent Wege H et al (1979) J Gen Virol 42 , 37 transactivator, the sequence of which is conserved in several different strains. Jijoye cells (CCL 87) A cell line derived Chen H et al (2000) J Virol 74 , 2703 from ascitic fluid of an African black Setiyaningsih S et al (2008) Virus Res 132 , 220 boy with Burkitt’s lymphoma of the liver. The cells contain an unidentified Jemez Springs virus A probable species herpes-like virus. in the genus Hantavirus, identified from the dusky shrew (Sorex monticolus ) in JKT-6423; JKT-6969; JKT-7041; JKT-7075 New Mexico and Colorado, USA. viruses Indonesian isolates of probable members of the genus Coltivirus , family Arai S et al (2008) Am J Trop Med Hyg 78 , 348 Reoviridae .

Jena virus (JV) A tentative species in Joa virus (JOAV) A serotype of Frijoles the genus Norovirus . A noncultivat- virus in the genus Phlebovirus . Not able bovine enteric calicivirus associ- reported to cause disease in humans. ated with diarrhea in calves and first described in Jena, Germany. Joest–Degen bodies Acidophilic intra- Liu BL et al (1999) J Virol 73 , 819 nuclear inclusion bodies found in the brain cells of horses infected with Jensen sarcoma cell line (CCL 45) This Borna disease virus. Pathognomonic cell line was established in 1958 from for the disease. 265 Juquitiba virus

Johnston Atoll virus (JAV) An unas- is a receptor for the sigma 1 protein of signed arbovirus. With Quaranfil virus reovirus. forms the Quaranfil serogroup. Isolated Barton ES et al (2001) Cell 104 , 441 from a tick, Ornithodoros capensis , from the nest of a common noddy tern on Junín virus (JUNV) A species in the Sand Island, Johnston Atoll in the genus Arenavirus belonging to the central Pacific. Also isolated in South Tacaribe serogroup. One of the South Island, New Zealand and Queensland, American hemorrhagic fever viruses. Australia. Not known to cause disease Causes Argentine hemorrhagic fever, in humans. Virions are enveloped with a disease of corn-harvesters in Buenos three major polypeptides. Aires, Cordoba, and Santa Fe prov- Austin FJ (1978) Am J Trop Med Hyg 27 , 1045 inces. Characterized by fever, leukope- Zeller HG et al (1989) Arch Virol 108 , 191 nia, exanthema, and renal involvement. Mortality rate probably 3–15% but Joinjakaka virus (JOIV) An unassigned most cases recover without sequelae. vertebrate rhabdovirus. Isolated from Transmission from humans to humans a pool of mixed culicines in the Sepik rarely, if ever, occurs. Isolated from wild district of New Guinea. Not reported rodents and transmission to humans is to cause disease in humans. probably via contamination with rodent urine and feces. The vector rodent spe- Jones virus An isolate (prototype strain) cies is Calomys musculinus . Experimental of reovirus type 2 from a child with infection of guinea pigs causes hem- diarrhea. orrhagic disease with depression of the immune response. A disease very Josiah virus A strain of Lassa virus . like that in humans is produced in the New World primate, Callithrix jacchus . Juan Diaz virus (JDV) A strain of Thymectomy protects mice against the Bushbush virus in the genus Orthobunya- lethal effects of infection. Causes chronic virus. Isolated from a sentinel mouse in carrier state in wild rodents and persist- Panama. Not reported to cause disease ent infection in cell cultures. Replication in humans. in wide range of cell cultures usually without CPE, but Vero cells are best for Jugra virus (JUGV) A species in the genus isolation, usually with CPE, although Flavivirus , related to yellow fever virus. some cells survive to form a chronically Isolated from mosquitoes and a bat in infected culture. In polarized cells, Junín peninsular Malaysia. Not reported to virus entry and release occurs through cause disease in humans. the apical plasma membrane. Synonyms : Argentine hemorrhagic JUN A member of the AP-1 family of fever virus; endo-epidemic hemor- transcription factors. Binds retinoblas- rhagic fever virus; O’Higgins disease toma protein. Originally detected as a virus. viral oncogene transduced by avian leukosis virus and associated with sarco- Childs JE and Peters CJ (1993) In The mas. Human papillomavirus E7 binds Arenaviridae , edited by M Salvato. New to JUN. York: Plenum Press, p. 331 Cordo SM et al (2005) J Gen Virol 86 , 1475 Romanowski V (1993) In The Arenaviridae , Juncopox virus (JNPV) A species in the edited by M Salvato. New York: Plenum genus Avipoxvirus , isolated from a Press, p. 51 North American sparrow, Junco hyema- lis. Closely related to fowlpox virus by Juquitiba virus A probable species in DNA restriction analysis. the genus Hantavirus, identified by Beaver DL and Cheatham WJ (1963) Am J sequence analysis of human tissues of Pathol 42 , 23 fatal cases of hantavirus pulmonary Schnitzlein WH et al (1988) Virus Res 10 , 65 syndrome in Brazil in 1994. The virus has not yet been isolated. The rodent junctional adhesion molecule 1 (JAM1) vector is the black-footed pigmy rice An integral tight junction protein, which rat, Oligoryzomys nigripes. Jurona virus (JURV) 266

Jurona virus (JURV) A tentative species in Changuinola virus. Not associated with the genus Vesiculovirus . Not assigned to disease in humans. an antigenic group. Isolated from mosquitoes in Para, Brazil. Not reported to Jutiapa virus (JUTV) A species in the genus cause disease in humans. Flavivirus , a member of the Modoc antigenic subgroup. Isolated from the cotton Juruaça virus An unclassified arbovirus rat, Sigmodon hispidus, in Guatemala. Not recovered from the viscera of an unde- reported to cause disease in humans. termined bat in the Amazon region of Juvenile laryngeal papillomatosis (JLP) Brazil. A rare disease of children above the age of 7, caused by HPV types 6 and Jutaí virus A possible species in the 11, which may result from perinatal genus Orbivirus , isolated from phlebot- infection from an infected mother. omine sandflies in the Amazon region of Brazil. Antigenically related to JV1 virus Synonym for echovirus 20. K

K virus (KV) Kilham polyomavirus, a Kaeng Khoi virus (KKV) A species in the strain of Murine pneumotropic virus in genus Orthobunyavirus. Not assigned to the genus Polyomavirus. A natural and an antigenic group. Isolated from bats, probably silent infection of wild mice. rats and bedbugs (Cimicidae ) caught in Causes fatal pneumonia and some- caves in Thailand, and from dead bats times liver lesions on injection into (Chaerephon plicata ) in Cambodia. Not mice less than 10 days old. In mouse reported to cause disease in humans. lung cell cultures, foci of transformed Osborne JC et al (2003) J gen Virol 84 , 2685 cells appear which produce tumors on injection into newborn or X-irradiated Kaffir-pox virus Synonym for Variola virus. mice. Agglutinates sheep erythrocytes at room temperature or 37°C. Kaikalur virus (KAIV) A strain of Kilham L and Murphy HW (1953) Proc Soc Shuni virus, a species in the genus exp Biol Med 82, 133 Orthobunyavirus, belonging to the Law M-F et al (1979) J Virol 30, 90 Simbu serogroup. Isolated in suckling mice and from a pool of mosquitoes, K9 virus A strain of Human herpesvirus 5, Culex tritaeniorhynchus, collected at in the genus Cytomegalovirus, isolated Kaikalur, Andra Pradesh, India. Not from a cell line derived from a tumor known to infect humans. biopsy specimen from a patient with Kaposi’s sarcoma. Similar to Mj strain Rodrigues FM et al (1977) Indian J med Res 66, 719 in producing a slow and incomplete CPE in human embryo lung cells. Kairi virus (KRIV) A species in the genus Glaser R et al (1977) J Natl Cancer Inst 59, 55 Orthobunyavirus, belonging to the Bunyamwera serogroup. Isolated from K27 virus (K27V) A serotype of Puumala mosquitoes in Trinidad, Brazil and virus in the genus Hantavirus, isolated Colombia, and from a febrile horse in from a human case in Russia. The vec- Argentina. Not reported to cause dis- tor is probably a species of rodent of ease in humans. the genus Clethrionomys. Dunn EF et al (1994) J gen Virol 75, 597 Kachemak Bay virus (KBV) A serotype Kaisodi serogroup viruses Three tick- of Sakhalin virus in the genus Nairovirus. borne viruses: Isolated from ticks, Ixodes signatus. Ritter DG and Feltz ET (1974) Can J Microbiol Kaisodi virus 20, 1359 Lanjan virus Silverwater virus Kadam virus (KADV) A species in the Morphologically like Orthobunyavirus genus Flavivirus. Isolated from a tick, but not serologically related to mem- Rhipicephalus pravus, in Uganda. Not bers of that genus. reported to cause disease in humans. Kaisodi virus (KSOV) An unassigned Kadipiro virus (KDV) A species in the virus in the family Bunyaviridae, related genus Seadornavirus. Isolated from to Lanjan virus and Silverwater virus. Culex mosquitoes. A member of the Kaisodi serogroup. Attoui H et al (2000) J gen Virol 81, 1507 Isolated from ticks and a ground thrush in the Shimoga district of Mysore, Kadipiro virus (Java-7075) (KDV-Ja7075) India. Not reported to cause disease in A strain of Kadipiro virus. humans. Kala Iris virus (KIRV) 268

Kala Iris virus (KIRV) A serotype of Kao Shuan virus (KSV) A strain in the Chenuda virus in the genus Orbivirus, genus Nairovirus, belonging to the Dera belonging to the Chenuda antigenic Ghazi Khan serogroup. Isolated from complex. a tick, Argas robertsi, in Taiwan. Not reported to cause disease in humans. Kaletra A protease inhibitor used in HIV therapy which is a combination of Kaplan leukemia virus A strain of Murine Lopinavir and Ritinovir. Used together leukemia virus isolated from tissues of with a nucleoside inhibitor for AIDS a mouse in which leukemia had been therapy. induced by exposure to X-irradiation. Kaposi’s sarcoma Kamese virus (KAMV) An unassigned spe- A skin tumor first cies in the family Rhabdoviridae. Member described in 1872 by Moriz Kaposi of the Hart Park serogroup. Isolated which frequently develops in young from Culex annulirostris in Uganda and males infected with HIV early in the Central African Republic. Not reported symptomatic phase of AIDS, and to cause disease in humans. presents in a slowly progressive invasive form. There is increasing evidence that human herpesvirus 8 (HHV-8) is Kamiiso-8Cr-95 virus A strain of Puumala involved in the etiology of Kaposi’s virus in the genus Hantavirus. Isolated sarcoma. The virus DNA is detectable from mice, Clethrionomus rufocanus. in all epidemiological forms of Kaposi’s sarcoma. See Human herpesvirus 8 . Kammavanpettai virus (KMPV) A tentative species in the genus Orbivirus. Gallo RC (1998) Science 282, 1837 Isolated from a Brahminy myna bird, Kaposi’s sarcoma-associated herpesvirus Sturnus pagoderum, in Vellore, Tamil Human herpesvirus 8 Nadu, India. See . Kararao virus A possible species in the kanapox A strain of Canarypox virus, used genus Orbivirus, isolated in the Amazon as a bird vaccine, from which ALVAC region of Brazil. Antigenically related was derived. to Changuinola virus. Not associated with disease in humans. kangaroo blindness syndrome A disease of kangaroos which become blind due Karelian fever Synonym for Ockelbo to localized virus infection of the ret- virus infection. Also called Pogosta fever. ina and optic tracts, and stumble into bushes and other objects when dis- Karimabad virus (KARV) A serotype of turbed. Isolation of a strain of Wallal Sandfly fever Naples virus in the genus virus (CSIRO 95/223) from eye tissue Phlebovirus. Isolated from females of suggests that it may be the etiological Phlebotomus sp in Iran and Pakistan. agent, but another mosquito-transmit- Not reported to cause disease in ted orbivirus, Warrego virus, has also humans. been isolated from some of the affected Robeson G et al (1979) J Virol 30, 339 kangaroos. The disease could be reproduced experimentally by infection with Karshi virus (KSIV) A serotype of Royal Wallal virus, which appears to be the Farm virus in the genus Flavivirus, major cause of kangaroo blindness. belonging to the mammalian tick- Synonym: choroid blindness syndrome. borne virus group. Isolated from ticks, Hooper PT et al (1999) Aust vet J 77, 529 Ornithodoros papillipes, in the Karshi Reddacliff L et al (1999) Aust vet J 77, 522 desert, Uzbekistan. Agglutinates goose erythrocytes and causes paralysis in Kannamangalam virus (KANV) An unas- 2- to 3-week-old mice. Not reported to signed vertebrate rhabdovirus. Isolated cause disease in humans. from a house crow, Corvus splendeus, in Vellore, Tamil Nadu, India. Not reported karyopherins Cellular proteins of karyo- to cause disease in humans. pherin alpha and beta families which 269 Keishi virus

play a central role in nucleocytoplas- Kazan virus A strain of Puumala virus, mic transport. Also known as import- isolated from the rodent, Clethrionomys ins, qv. glareolus, in Russia. Chook YM and Blobel G (2001) Curr Opin Lundkvist et al (1995) J virol Meth 52, 75 Struct Biol 11 , 703 KB cells (CCL 17) A heteroploid cell line Kasba virus (Chuzan virus) (KASV) A derived from an epidermoid carcinoma serotype of Palyam virus in the genus in the mouth of an adult human male. Orbivirus. Isolated from Culex vishnui in North Arcot District, Tamil Nadu, K-Balb (K-234) cell line (CCL 163.3) A India. Not reported to cause disease in subline of CCL 163 derived after trans- humans. formation with the Kirsten strain of Ohashi S et al (2004) J clin Microbiol 42, 4610 murine sarcoma virus (K-MSV).

Kasokero virus (KASV) An unassigned KBSH virus Name given to an early virus in the family Bunyaviridae, with isolate of porcine parvovirus from a Yogue virus comprises the Kasokero human cell line. Parvoviruses have fre- serogroup. Isolated from a fruit-eating quently been isolated from cell lines, bat, Rousettus aegyptiacus. and this may be the result of using infected pig trypsin. KBSH virus was Zeller HG et al (1989) Arch Virol 108, 211 isolated from a KB cell subline originating from a Hamburg laboratory. Kata virus Synonym for Peste des petits ruminants virus. Hallauer C et al (1971) Arch Virol 35, 80 k-cyclin A protein encoded by human Kawasaki disease A mucocutaneous herpesvirus 8 which is a homologue of lymph node syndrome that has fea- a cellular protein, cyclin D. tures of a virus infection but so far no causative agent has been discovered. Kedong virus A strain of Sheeppox virus. It is the most common cause of pediatric ischemic heart disease in the world, Kedougou virus (KEDV) A mosquito- with significantly higher mortality borne species in the genus Flavivirus, in among males. Appeared in Japan in the the Dengue virus group. Isolated from 1950s and now causes about 120 cases a pool of mosquitoes, Aedes minutus, per 100 000 population, a sixfold higher collected on human bait. Antibodies incidence than in the USA. Can be con- present in human sera but no evidence fused with Stevens–Johnson syndrome. of pathogenicity for humans. Known to occur in Senegal and the Central Kawakami–Theilen strain of feline African Republic. Recent genome leukemia virus Isolated from a case sequence analysis confirms the relation of spontaneous lymphosarcoma in a to dengue virus. Persian cat. Produces leukemia in kittens if injected when they are 1–2 days Kuno G and Chang GJ (2007) Arch Virol 152 , old but is not pathogenic in cats more 687 than 5 months old. The development Robin Y et al (1978) Ann Microbiol, Paris 129A, 239 of antibodies prevents viremia and the induction of leukemia. “ Keep-up, catch-up, follow-up ” sched- Salerno RA et al (1979) Proc Soc exp Biol Med ule A schedule for measles elimination 160, 18 developed by PAHO. It combines routine vaccination (keep-up) with mass Kawino virus (KaV) An unassigned vaccination campaigns (catch-up) and 4- entero-like virus isolated from Kawino year follow-up campaigns (follow-up). village, Kenya from mosquitoes, Mansonia uniformis, in 1973. No evidence Keishi virus A strain of Ibaraki virus. of infection or disease in vertebrates. Omori T (1970) Natl Inst Anim Hlth Quart 10 Pudney M et al (1978) J gen Virol 40, 433 (suppl), 45 Kemerovo virus (KEMV) 270

Kemerovo virus (KEMV) A serotype of Isolated from ticks, Argas pusillus, and Great Island virus in the genus Orbivirus. a bat, Scotophilus temmenckii, in western Isolated from female ticks, Ixodes Malaysia. Not reported to cause dis- persulcatus, and two humans with a ease in humans. febrile illness in western Siberia and Synonym: Issyk-kul virus. also from a bird, Phoenicurus phoenicurus, in Egypt. Antibodies are found in Ketol-enol tautomerism The bases which humans, cattle, horses, small mammals make up DNA or RNA can exist in two and birds in Siberia. forms, the keto- (lactame-) form and the enol-(lactime-) form. The equilib- Kenai virus (KENV) A serotype of Great rium between these two forms is far Island virus in the genus Orbivirus. towards the keto-form, with an equilibrium constant of 10 4 . Thus there is kennel cough See Canine adenovirus, one chance in 10 4 that a nucleotide is also Canid herpesvirus 1 . in the enol form when an RNA or DNA polymerase passes to place the com- keratin intermediate filament A protein plementary base. Normally guanines cytoskeletal network in keratinocytes. pair with cytidines but not with thymi- The human papillomavirus protein E4 dines. But a thymidine in the enol form binds to this structure and, in the case can pair with a keto-guanine, a keto- of HPV16, causes its collapse. A can pair with enol-C, a keto-C with keratinocytes Skin cells of the keratinized enol-A, and a keto-T with enol-G. As a layer of the epidermis. result the polymerase will add a wrong base whenever the template base is Kerecid Trade name for idoxuridine eye in the enol-form. To correct for this, drops. DNA-dependent polymerases have a proof-reading activity that corrects Kern Canyon group A group of four sero- these errors in base-pairing, resulting logically related unassigned rhabdo- in error frequencies in DNA synthe- viruses. They are: Barur, Fukuoka, Kern sis of only about 10 7 per copied base. Canyon and Nkolbisson viruses. The error rate for RNA polymerases is much higher. Kern Canyon virus (KCV) An unassigned virus in the family Rhabdoviridae, Keuraliba virus (KEUV) An unassigned belonging to the Kern Canyon sero- species in the family Rhabdoviridae, group. Isolated from a bat in California, serologically related to Le Dantec virus. USA. Not reported to cause disease in Isolated from gerbils and a rodent in humans. Senegal. Not reported to cause disease in humans. Kernig’s sign A sign of meningitis seen in a number of viral diseases such as Keystone virus (KEYV) A serotype of Japanese encephalitis. The patient can California encephalitis virus in the genus easily and completely extend the leg Orthobunyavirus. Isolated from mosqui- when in the dorsal decubitus position toes of Aedes sp in the Tampa Bay area but not when in the sitting posture or of Florida; also in Texas, Louisiana, when lying with the thigh flexed upon Mississippi, Georgia, N Carolina and the abdomen. Virginia, USA. The principal vector is Aedes atlanticus. The natural mamma- Ketapang virus (KETV) A serotype of lian host has not been identified, but Bakau virus in the genus Orthobunyavirus. studies in Florida, Texas and Virginia Isolated from mosquitoes in Malaysia. suggest that cottontail rabbits, grey Antibodies are found in humans but the squirrels and cotton rats contribute virus is not associated with disease. to the maintenance of the virus. Not Ketarax Trade name for levamisole reported to cause disease in humans. hydrochloride. See levamisole . Fine PEM and Le Duc JW (1978) Am J trop Med Hyg 27, 322 Keterah virus (KTRV) An unassigned Watts DM et al (1979) Am J trop Med Hyg virus in the family Bunyaviridae. 28, 344 271 Kimberley virus (KIMV)

Khabarovsk virus (KHAB) A species in which includes the H-3, X-14 and HER the genus Hantavirus. Isolated from strains. A natural and latent infection a vole, Microtus fortis, in far-east- of rats. Causes an acute fatal enteritis ern Russia. Serologically and geneti- on i.p. injection into newborn hamsters cally distinct from other hantaviruses. of Mastomys sp. A smaller dose given Disease potential unknown. to slightly older hamsters results in Hörling J et al (1996) J gen Virol 77, 687 stunted growth with abnormal development of teeth and skull bones. On i.c. injection in newborn hamsters Kharagysh virus (KHAV) A serotype of there may be cerebellar hypoplasia Great Island virus in the genus Orbivirus. and ataxia. Infection in pregnant hamsters and rats may cause congenital Khasan virus (KHAV) A serotype of abnormalities. Replicates in rat but Crimean-Congo hemorrhagic fever virus not mouse cell cultures with CPE. Nairovirus. in the genus Isolated from Agglutinates guinea pig, hamster, Haemaphysalis longicornis, ticks, in the human and rat erythrocytes at 23–24°C. Primorie region of Russia. Pathogenic Does not spontaneously elute. Infected when injected i.c. into suckling and 2- cells hemadsorb. week-old mice. Replicates in primary Synonyms: rat virus; latent rat virus; cultures of chick, duck and green mon- R virus. key cells without CPE. No hemagglutinin. Size 90–110nm and morphology Kilham L and Oliver LJ (1959) Virology 7, 428 typical of family Bunyaviridae. Salzman LA and Fabisch P (1978) J gen Virol 39, 571 Lvov DK et al (1978) Acta Virol 22, 249 Tijssen P (editor) (1990) Handbook of Parvo- viruses. Boca Raton: CRC Press Khujand virus A tentative species in the genus Lyssavirus. Isolated from bats in killed vaccine Vaccine comprising virus southern Kyrgyzstan. There have been which has been inactivated, usually by a few human rabies cases in the region, treatment with a chemical such as for- but not so far ascribed to Khujand malin, betapropiolactone, or aziridine, virus infection. e.g. foot-and-mouth disease vaccine, Kuzmin IV et al (2006) Dev Biol (Basel) 125, 273 Salk vaccine.

KI-83-262 virus A serotype of Seoul virus killer cells See natural killer (NK) cells . in the genus Hantavirus. Killer cell immunoglobulin-like recep- KI-85-1 virus A serotype of Seoul virus in tors NK cells represent about 5–15% the genus Hantavirus. of the lymphocytes in the blood, and about 1% of lymphocytes in the spleen. KI-88-15 virus A serotype of Seoul virus in The majority of blood NK cells express the genus Hantavirus. CD56 in combination with various molecules of the killer cell immunoglobulin receptor family. They also express high Kikuchi’s disease A rare, self-limiting, levels of perforin, which cause lysis of necrotizing lymphadenitis of unknown the target cell. etiology that mainly affects young women. Suspected to have a viral Cooper MA et al (2001) Trends Immunol etiology. 22 , 633 Aguiar JI et al (2000) Brazilian J inf Dis 4, 208 Sato Y et al (1999) J Neurol Sci 163, 187 kilobase (kb) A measure of the size of a nucleic acid molecule. One kilo- Kilham polyomavirus (KpyV) A strain of base 1000 nucleotides. Animal virus Murine pneumotropic virus in the genus DNAs range in size from less than 2 Polyomavirus. kilobases (Circoviruses) up to several hundred (Poxviruses). Kilham rat virus (KRV) A species in the genus Parvovirus. Belongs serologi- Kimberley virus (KIMV) A tentative cally to rodent parvovirus group 1, species in the genus Ephemerovirus. Kimberley virus (KIMV) 272

Isolated from cattle, Bos taurus, in Kismayo virus (KISV) An unassigned Australia. virus in the family Bunyaviridae, Calisher CH et al (1989) Intervirology 30, 241 belonging to serogroup 1. Related to Bhanja and Forecariah viruses. Isolated kinase Enzyme that catalyzes phosphor- in Somalia from a tick, Rhipicephalus ylation (e.g. transfer of the phosphoryl pulchellus, which was removed from a group of ATP to another compound). jackal. See polynucleotide kinase and protein Hubalek Z and Holouzka J (1985) Arch Virol kinase . 84, 175

Kindia virus (KINV) A serotype of Klamath virus (KLAV) A tentative spe- Palyam virus in the genus Orbivirus. cies in the genus Vesiculovirus. Isolated Isolated from the tick, Amblyomma vari- from a meadow mouse, Microtus monta- egatum, in the Guinea Republic. nus, in Klamath County, Oregon, USA. Boiro I et al (1986) Bull Soc Pathol Exot Fil Replicates in BHK21 cells with CPE. 79, 187 Day-old mice die 7 days after i.c. injection. Resembles rabies in cytopathology. kinkajou herpesvirus Synonym for lori- Antigenically related to Mount Elgon sine herpesvirus 1. bat virus. Not reported to cause disease in humans. Kirk virus A strain in the genus Parvo- virus. Serologically belongs to rodent Calisher CH et al (1989) Intervirology 30, 241 parvovirus group 1. Isolated from a Murphy FA et al (1972) Arch ges Virusforsch 37, 323 line of Detroit 6 cells which had been inoculated with plasma from an individual who had ingested MS-I infec- Kleinschmidt procedure A technique for tious hepatitis serum. preparing monomolecular films of DNA or RNA for electron microscopy. Nucleic Mirkovic RR et al (1971) Proc Soc exp Biol acids are coated with a basic protein, Med 138 , 626 e.g. cytochrome c, and spread on a dena- Kirsten leukemia virus A strain of tured protein monolayer at an air–water Murine leukemia virus in the genus interface. The nucleic acid molecules Gammaretrovirus, recovered from C3Hf/ are shadowed with a heavy metal, then Gs mice which had been injected with viewed in the electron microscope. a cell-free extract of thymic lymphoma tissue. Newborn mice injected with the Klenow fragment The larger of the two virus develop splenomegaly, excessive fragments of Escherichia coli DNA proliferation of red cell precursors and polymerase I formed after limited pro- a severe, rapidly fatal anemia. teolytic cleavage. It retains the DNA Synonym: erythroblastosis of mice virus. polymerase and the 3 ’ –5 ’ exonuclease activities, but lacks the 5 ’ to 3 ’ exo- Kirsten WH et al (1967) J Natl Cancer Inst nuclease activity of the intact enzyme. 38, 117

Kirsten murine sarcoma virus (KiMSV) knockout A general term for the elimi- A species in the genus Gammaretrovirus nation of a gene (a null allele) from an belonging to the Mammalian virus group, organism. Experimental animals, such isolated from a W/Fu rat injected when as mice, from which a particular gene newborn with Kirsten leukemia virus. has been eliminated (knocked-out), can It appears to be a recombinant between be used to test the functions of related the leukemia virus and an endogenous gene products. rat type C oncovirus genetic sequence, and carries the Ki-ras oncogene, which Knodell score A system for describing encodes a guanine-triphosphate-binding the degree of liver damage in cases of plasma membrane protein. chronic active hepatitis in cases of hep- Kirsten WH and Mayer LA (1967) J Natl atitis B or hepatitis C. See also Ishak Cancer Inst 39, 311 score . 273 Korean hemorrhagic fever virus koala retrovirus (KORV) An endogenous Kodzha virus (CCHFV) A strain of virus detected by electron microscopy Crimean-Congo hemorrhagic fever virus in of PBL cultures and lymphoid tissue of the genus Nairovirus. koalas, Phascolarctos cinereus. The complete proviral DNA sequence showed Kodzha virus AP92 A strain of Crimean- relationship to Gibbon ape leukemia virus. Congo hemorrhagic fever virus in the Hanger JJ (2000) J Virol 74, 4264 genus Nairovirus.

Kobuvirus A genus in the family Picor- Kodzha virus C68031 A strain of Crimean- naviridae containing two species, Aichi Congo hemorrhagic fever virus in the virus (the type species) and Bovine genus Nairovirus. kobuvirus. Unlike other picornaviruses, kobuviruses show icosahedral surface Kokobera virus (KOKV) A species in structure under the electron micro- the genus Flavivirus, belonging to scope. Virions are stable at pH 3.5 . the Japanese encephalitis serogroup. Isolated from mosquitoes in Queens- Koch’s postulates Criteria for determin- land, Australia and in New Guinea. ing whether a particular microorgan- Occasional human infections have been ism is the etiological agent of a disease. reported in patients with acute polyar- They are: ticular disease. (1) The microbe is regularly found in Boughton CR et al (1986) Med J Aust lesions of the disease. 145, 90 (2) It can be grown in pure culture in vitro. Kolongo virus (KOLV) An unassigned (3) When such a pure culture is inocu- vertebrate rhabdovirus. Isolated from lated into experimental animals, a birds, Euplectes afra, in Central African similar typical disease results. Republic. Antigenically related to (4) The microbe can be re-isolated from Mokola virus. Not reported to cause the experimentally induced disease disease in humans. in animals. These postulates were modified for Koolpinyah virus (KOOLV) An unas- application to virus diseases by Rivers signed vertebrate rhabdovirus closely (1937) to read: related to, but distinct from, the rabies- (1) A specific virus must be found related Kotonkan virus. Isolated from associated with the disease with a the blood of bovines near Darwin, degree of regularity. Australia. (2) The virus must be shown to occur in the sick individual not as an Gard GP et al (1992) Intervirology 34, 142 incidental or accidental finding but as the cause of the disease under Koongol virus (KOOV) A species in the investigation. genus Orthobunyavirus. With Wongal Since that time, the ability to grow and MRM31 viruses forms the Koongol viruses in cell culture and the develop- serogroup. Isolated from mosquitoes ment of sequence-based identification in Queensland, Australia and New of viral pathogens has suggested new Guinea. Not reported to cause disease postulates to link a virus as the etiolog- in humans. Antibodies are common in ical agent of a disease (Fredericks and cattle in Queensland. Relman (1996)). Fredericks DN and Relman DA (1996) Clin Koplik’s spots Diagnostic signs seen Microbiol Rev 9, 18 in the prodromal stage of measles Rivers TM (1937) J Bacteriol 33, 1 virus infection. Small irregular bright red spots on the buccal and lingual kodoko virus A novel arenavirus related mucosa, with a minute bluish white to but distinct from lymphocytic cho- speck in the center of each. riomeningitis virus, obtained from an African rodent (Mus nannomys ). Korean hemorrhagic fever virus Synonym Lecompte E et al (2007) Virology 364 , 178 for Hantaan virus. Kotonkon virus (KOTV) 274

Kotonkon virus (KOTV) An unassigned subtype) in the genus Flavivirus, belong- species in the family Rhabdoviridae, ing to the tick-borne encephalitis virus isolated from Culicoides sp in Nigeria. complex (European subtype). Isolated There is serological evidence of infec- from the tick, Ixodes ricinus, squirrel, tion in humans and domestic ani- Sciurus vulgaris, field vole, Microtus mals. Probably the cause of an acute agrestis, hare, Lepus timidus, thrushes, febrile illness similar to ephemeral Turdus sp and a bunting, Emberiza cit- fever in Nigeria. Antigenically related rinella, in Finland. Causes a febrile ill- to Obodhiang virus and Rochambeau ness with encephalitis in humans (5–20 virus. cases annually in Finland). Bauer SP and Murphy FA (1975) Infect Wahlberg P et al (2006) Scand J Infect Dis Immun 12, 1157 38, 1057 Kemp GE et al (1973) Am J Epidemiol 98, 43 Kuzmin IV et al (2006) J gen Virol 87 , 2323 Kunitachi virus Synonym for Avian paramyxovirus 5. Koutango virus (KOUV) A species in the genus Flavivirus, belonging to Kunjin virus (KUNV) A serotype of the Japanese encephalitis serogroup. West Nile virus in the genus Flavivirus, Isolated from Kemp’s gerbil, Tatera belonging to the Japanese encephalitis kempi, and rodents of Mastomys and serogroup. Isolated from mosquitoes, Lemnyscomys sp in Senegal and Central Culex annulirostris, in Queensland, African Republic. Not reported to cause Australia, and in Borneo and Sarawak. disease in humans. Can cause fever with a rash and encephalitis in humans, and has been Kowanyama virus (KOWV) An unas- isolated from one case of infection in a signed virus of the family Bunyaviridae. laboratory worker. Isolated from Anopheles sp in the Mitchell River area, north Queensland, Kununurra virus A probable species in Australia. Antibodies found in domes- the family Rhabdoviridae. Isolated in tic fowls, horses and kangaroos. Not suckling mice from a pool of female reported to cause disease in humans, mosquitoes, Aedeomyia catasticta, col- although antibodies are found in lected with chicken-baited traps at aborigines. Kununurra in Western Australia. Not Doherty RL et al (1968) Trans R Soc trop Med reported to infect humans. Hyg 62, 430 Johansen CA et al (2007) Virus Genes 35, 147 Liehne CG et al (1976) Aust J exp Biol Med Kozak rule For optimal translation of an Sci 81, 499 mRNA, positions –3 and 4 relative to the first nucleotide of the initiation Kupffer cells Specialized macrophages codon must be G or A. of the liver sinusoids, responsible for Kozak M. (1986) Adv Virus Res 31 , 229 removing particulate matter from the circulating blood, such as old erythro- Kozak sequence A base sequence near the cytes and viruses. Ingested virions may 5’ end of untranslated mRNA which is be degraded, may transit macrophages required for optimal recognition of the to underlying parenchyma without initiation codon (AUG) by eukaryotic replicating, or may replicate in the cell ribosomes. The sequence is macrophages with or without spread GCCA/GCCAUGG. of the infection to underlying paren- chymal cells. Kozak M (1987) J mol Biol 196, 947 Kuru A disease caused by a prion, one of Kumba virus A strain of Semliki Forest the transmissible spongiform encepha- virus isolated in Cameroon. lopathy agents. A subacute progressive degeneration of the brain in humans. Kumlinge virus (KUMV) A Finnish strain The natural disease was restricted to a of Tickborne encephalitis virus (European small area in the highlands of Papua 275 Kyzylagach virus (KYZV)

New Guinea, centered round the Fore into the intestine and at other sites. No people. It appears to have been caused CNS involvement. A number of dead by ritual cannibalism of the dead. With langurs and bonnet macaques were the discontinuance of this practice the found during the epidemic and the disease has essentially disappeared. disease may be disseminated by move- See also prion diseases . ment of monkeys and birds. Antibodies Gajdusek DC (1977) Science 197, 943 are present in small forest mammals. The virus is widely distributed in India, Kwatta virus (KWAV) A tentative spe- but human infections occur only in cies in the genus Vesiculovirus. Isolated Mysore. Mice develop encephalitis on from Culex sp in Surinam. Not reported injection by various routes. They may to cause disease in humans. fail to develop antibodies and remain chronically sick for long periods. Calisher CH et al (1989) Intervirology 30, 241 Suckling hamsters are also susceptible but other rodents are resistant. Rhesus Kyasanur Forest disease virus (KFDV) and bonnet monkeys develop viremia A tick-borne species in the genus on i.c. or i.p. injection but show no dis- Flavivirus, member of the Mammalian ease. No vaccine is yet available. tick-borne virus group. The tick vector is Haemaphysalis spingera. In an Bhat HR et al (1979) Ind J Med Res 69, 697 epidemic among forest workers in Mysore State, India, in 1957, symptoms Kyzylagach virus (KYZV) A serotype of included headache, fever, back and Sindbis virus in the genus Alphavirus, limb pains, prostration, conjunctivitis, isolated in Russia from mosquitoes, diarrhea, vomiting and hemorrhages Culex modestus. L

L cell virus Cultures of L cells (a line of laboratory strains Viruses which have mouse cells) were reported to release been propagated in the laboratory in virus particles. These particles resem- vivo or in vitro . They may be different bled C-type viruses. On injection into in many properties from clinical or newborn mice and hamsters no tumors field isolates, known as wild strains. were induced. Infection could not be See vaccine virus markers . transmitted to mouse or rat embryo cultures. A subline of L cells, A9, which lacertid herpesvirus 1 (LaHV-1) An unas- is 8-azaguanine-resistant, also releases signed virus in the family Herpesviridae , virus particles, but they induce mor- identified in the green lizard, Lacerta phological changes (foci) in mouse viridis. Lizards captured in Italy and embryo fibroblast cultures. Focus for- taken to France developed papilloma- mation was more efficient on N-type tous skin lesions. A herpesvirus was than B-type mouse cells. identified by electron microscopy in Botis S et al (1976) J Virol 20 , 690 the lesions from one animal. Synonym : green lizard herpesvirus. L99 virus A strain of Seoul virus in the Raynaud A and Adrian M (1970) CR Acad genus Hantavirus. Isolated from the rat Sci Ser D 283 , 845 Rattus losea . La Crosse virus (LACV) A strain of L-132 cells (CCL 5) A heteroploid cell line California encephalitis virus in the genus derived from normal human embry- Orthobunyavirus. First isolated from the onic lung. Reported to be a sensitive brain tissue of a 4-year-old girl with cell line for the primary isolation of fatal meningoencephalitis in La Crosse, enteroviruses. Wisconsin, USA in 1960. Between 1960 and 1970, 509 cases of human infection laboratory acquired infections Before were reported, mainly in Wisconsin the development of biosafety cabinets and Minnesota. Large serosurveys it was not uncommon for occupational in Midwestern US states indicated virus infections to occur in labora- that there may be 1000 infections per tory workers, especially if spread by reported case, so very few result in the aerosol route. The introduction of overt disease. Classical encephali- guidelines for safe handling of infec- tis occurs, but some 10% of infected tious agents in the early 1980s has been children develop epilepsy, and a few followed in the 1990s by the introduc- deaths have occurred. Antibodies are tion of mandatory safety practices that, found in small forest mammals, and if properly executed, reduce the haz- virus can be isolated from mosqui- ards of working in a virus laboratory to toes, especially Aedes triseriatus , a a minimum. woodland insect that usually feeds Kiley MP and Lloyd G (1998) In Virology , on squirrels and chipmunks. In addi- vol. 1 of Topley & Wilson’s Microbiology and tion, discarded tyres that hold rain- Microbial Infections , Ninth edition, edited by water provide a breeding ground for the BWJ Mahy and L Collier. London: Arnold, mosquitoes and bring the disease to p. 933 urban areas. The range of Aedes triseria- LC Chosewood and DE Wilson (Editors) tus in the USA covers the area east of (2007) Biosafety in Microbiological and Biomedical Laboratories, Fifth edition. the Mississippi river. There are three Washington: US Government Printing Office genetically distinct subgroups: A, B, and C. laboratory safety See biosafety . Bennett RS et al (2007) Virol J 4 , 41 277 lagomorph herpesvirus 1

Calisher CH and Thompson WH (editors) nucleotides derived from the 5 end of (1983) California Serogroup Viruses . New the genome and a 3 non-coding seg- York: Alan Liss ment which is polyadenylated. Two El Said LHE et al (1979) Am J Trop Med Hyg subtypes or strains have been recog- 28 , 364 Huang C et al (1997) Virus Res 48 , 143 nized by complete genome sequence Janssen RS et al (1986) J Virol 59 , 1 analysis: LDV-P and LDV-C which have only 80% nucleotide sequence identity. Productive cytocidal replication of lactate dehydrogenase virus A synonym LDV only occurs in a subpopulation of for Lactate dehydrogenase-elevating macrophages which are continuously virus . renewed in lymphoid tissues, liver, and testis to support the persistent infection. Lactate dehydrogenase-elevating virus It is the continuous destruction of this (LDV) A species in the genus Arterivirus . subpopulation that results in decreased Infects only species of Mus . Has been clearance leading to elevated levels isolated from wild and laboratory mice, of lactate dehydrogenase and other Mus musculus, in Europe, USA, and enzymes in the blood plasma. In certain Australia. Mus caroli can be infected mouse strains, e.g. C58 and AKR, LDV experimentally. The virus productively can cytocidally infect anterior horn neu- infects macrophages and causes a life- rons via interaction with endogenous long infection with permanent viremia murine retroviruses to cause paralysis but no disease unless the mouse is known as ‘ age-dependent poliomy- immunocompromised. Infection is rec- elitis. ’ This disease only occurs in mice ognized by abnormally high levels of with some degree of immunodeficiency, plasma lactate dehydrogenase, and both e.g. as a result of aging. elevated enzyme levels and infectious Synonyms : age-dependent polio- LDV titers (104 –106 ID50/ml) persist encephalitis of mice virus; enzyme- for the life of the mouse. Certain other elevating virus; lactic dehydrogenase plasma enzyme levels are also raised in virus; Riley virus. infected mice due to a failure to clear Palmer GA et al (1995) Virology 209 , 637 them from the circulation, apparently Plagemann PGW et al (1999) Curr Top Virol as a result of impaired function of the 1 , 27 reticuloendothelial system. The infec- Rowson KEK and Mahy BWJ (1985) J Gen tion is usually silent, with no obvious Virol 66 , 2297 pathological changes. Antibodies are produced but infectious virus–antibody lagenavirus A name proposed for viruses complexes continue to circulate. There which appear bottle-shaped on elec- is a minor degree of splenomegaly and tron microscopy. The only one so far certain immunopathological changes. reported is lactate dehydrogenase- Virus is found in all the body tis- elevating virus, but the name was not sues, and is excreted in urine, saliva adopted. and feces. Transmission between mice Almeida JD and Mims CA (1974) Microbios does not occur readily and probably 10 , 175 results mainly from blood transfer during fighting. Fetuses and young can lagging-strand During the synthesis of be infected by a mother who becomes DNA, replication proceeds by the for- infected during pregnancy or lacta- mation of two replication forks which tion. Virions are enveloped, 55 nm in synthesize DNA in different directions diameter, containing a spherical core from the origin. Continuous DNA syn- 25–35 nm in diameter. The nucleocapsid thesis occurs on one strand (leading protein (12–14 kDa) is associated with strand synthesis) and discontinuous positive single-stranded RNA, 14.1 kb synthesis on the other strand (known in length, encoding nine open reading as the lagging strand). frames. Genome expression involves formation of a 3 co-terminal nested lagomorph herpesvirus 1 Synonym for set of subgenomic mRNAs, each carry- leporid herpesvirus 1 (cottontail rabbit ing a non-coding 5 leader of 156–212 herpesvirus). Lagos bat virus (LBV) 278

Lagos bat virus (LBV) A species in the healthy monkeys ( Cercopithecus aethi- genus Lyssavirus . Antigenically related ops ) imported from Uganda. There to, but distinguishable from, rabies were small outbreaks of the disease virus. Isolated from a Nigerian fruit in Zimbabwe in 1975, and in Kenya in bat, Eidolon helvum , in 1956 on Lagos 1980 and 1987, and a large outbreak Island, Nigeria and in 1974 was also with over 100 cases in the Democratic found in bats, Micropteropus pussilis , Republic of the Congo in 1998. Then in the Central African Republic. Not the largest outbreak so far recorded known to cause disease in humans. occurred in northern Angola, West Pathogenic for adult mice, dogs and Africa from October 2004 to July 2005 rhesus monkeys on i.c. injection, but with 252 cases and 227 deaths. Genome adult mice are not affected by i.p. injec- nucleotide sequence analysis showed tion. Guinea pigs, rabbits, and mon- that the virus which emerged in Angola keys, Cercocebus torquatus , are not killed was similar to the viruses causing out- by peripheral inoculation. breaks in East Africa. Bourhy H et al (1993) Virology 194 , 70 There are seven recognized subspecies Markotter W et al (2008) Virus Res 135 , 10 of Lake Victoria marburgvirus: (1) Lake Victoria marburgvirus- Lagovirus A genus in the family Musoke (Kenya, 1980) Caliciviridae comprised of viruses of (2) Lake Victoria marburgvirus-Ozolin rabbits and hares. (Zimbabwe, 1975) (3) Lake Victoria marburgvirus-Popp Laguna Negra virus (LNV) A species in (West Germany, 1967) the genus Hantavirus . Isolated from (4) Lake Victoria marburgvirus- the rodent Calomys laucha in the Chaco Ratayczak (West Germany, 1967) region of Paraguay. Associated with an (5) Lake Victoria marburgvirus-Ravn outbreak of hantavirus pulmonary syn- (Kenya, 1987) drome involving 17 confirmed cases, (6) Lake Victoria marburgvirus-Voege two of whom died. (West Germany, 1967) (7) Lake Victoria marburgvirus-Ang Johnson AM et al (1997) Virology 238 , 115 1379v (Angola, 2005) La Joya virus (LJV) A tentative species in Towner JS et al (2006) J Virol 80 , 6497 the genus Vesiculovirus . Isolated from mosquitoes, Culex dunni, in Panama. Lambdapapillomavirus A genus in Not reported to cause human disease. the family Papillomaviridae contain- Calisher CH et al (1989) Intervirology 30 , 241 ing viruses which infect cats and dogs. The type species is Canine oral Lake Clarendon virus (LCV) A tentative papillomavirus . species in the genus Orbivirus . Isolated from ticks, Argas robertsi, in south-east Lamivudine 2 -deoxy-3 -thiacytidine. A Queensland, Australia. Not known to nucleoside analog inhibitor of reverse cause disease in humans. transcriptase which is also effective George TD et al (1984) Aust J Biol Sci 37 , 85 against hepatitis B virus infection. Synonym : 3TC. Lake Victoria cormorant herpesvirus Dusheiko G (1998) Rev Med Virol 8 , 153 Synonym for phalacrocoracid herpesvirus 1. Landjia virus (LJAV) An unassigned ver- Lake Victoria marburgvirus (MARV) The tebrate rhabdovirus. Isolated from a type species of the genus Marburgvirus . bird, Riparia paludicola, in the Central The virus was originally isolated in African Republic. Not reported to Marburg, Germany and Voege, former cause disease in humans. Yugoslavia in 1967 from patients with severe hemorrhagic fever who landlocked salmon reovirus (LSRV) had become infected by contact with A tentative species in the genus blood and tissues from apparently Aquareovirus . Isolated from landlocked 279 Las Maloyas virus (LMV)

salmon, Oncorhynchus masou , in Lapine parvovirus (LPV) A species in Taiwan. Replicates in BF-2, BB, and the genus Parvovirus , isolated from CCO cells. rabbit feces. Replicates and produces CPE in rabbit kidney cell cultures. Hsu YL et al (1989) Fish Pathol 24 , 37 Agglutinates human group O erythrocytes at 4°C. Stable at pH 3, resistant to Langat virus (LGTV) A species in the chloroform and to heating at 60°C for genus Flavivirus, in the Mammalian 30 min. No cross-reaction with latent tick-borne virus group. Isolated from rat virus in HI tests. In a survey in the a pool of ticks, Ixodes granulatus , in USA, 75% of rabbits were seroposi- Malaysia. Antibodies found in forest tive for parvovirus infection, and virus ground rats. Has low pathogenicity for could be recovered from the kidneys of mice and monkeys, but is very virulent neonatal rabbits. Found in the serum of in severe combined immunodeficient most commercially available rabbits in (SCID) mice. Baby mouse brain cell the USA. cultures show loss of contact inhibi- Synonym : rabbit parvovirus. tion, and the cells pile up. Cultivation in chick embryo fibroblasts causes Gregg DA and House CA (1989) Vet Rec 125 , attenuation and this virus protects 603 mice against homologous and heterol- Matsunaga Y et al (1977) Infect Immun 18 , 495 ogous tick-borne flavivirus. In humans Metcalf JB et al (1989) Am J Vet Res 50 , 1048 it has caused an antibody response but no disease, except in two leukemia patients infected experimentally who lapinized virus Virus adapted to rabbits. developed encephalitis. When rinderpest virus is so adapted it ceases to be virulent for cattle. McLean A (2000) Rev Med Virol 10 , 207 Pletnev AG et al (2000) Virology 274 , 26 largemouth bass virus (LMBV) A strain of Sante-Cooper ranavirus in the genus Lang virus A strain of reovirus type Ranavirus . Has caused recent outbreaks 1. Isolated from a stool sample of a of disease in lakes and reservoirs in healthy child in Cincinnati, OH in Texas and several other Southern 1954. United States.

Langur virus (PO-1-Lu) (LNGV) A spe- Zilberg D et al (2000) Dis Aquat Org 39 , 143 cies in the genus Betaretrovirus . An endogenous virus isolated from a lan- large T antigen A 90-kDa protein induced gur, Presbytis obscuris, by co-cultivation in cells infected with SV40 virus which of lung cells with bat or human cells. is involved in transcription, replication, Related to Mason–Pfizer monkey virus and cell transformation. but distinguishable from it. laryngeal papillomatosis A juvenile dis- Colcher D et al (1978) Virology 88 , 384 ease with growth of benign squamous cell papillomas in the larynx and tra- Lanjan virus (LJNV) An unassigned chea which can lead to narrowing of the virus in the family Bunyaviridae . airway, requiring excisions. The papil- Morphologically like Bunyavirus but lomas are recurrent and usually require serologically unrelated to members multiple excisions. Caused in childhood of that genus. Belongs to the Kaisodi by human papillomavirus, usually ano- serogroup (group 2) together with genital types such as HPV6 and HPV11, Silverwater virus. Isolated from the probably acquired from mothers suffer- tick, Dermacentor auratus , in Malaysia. ing from condyloma acuminatum. Not reported to cause disease in humans. Las Maloyas virus (LMV) A serotype of Anopheles A virus in the genus La-Piedad-Michoacan-Mexico virus Orthobunyavirus . Isolated from mosqui- (LPMV) Synonym for Porcine rubula- toes, Anopheles albitarsis. No evidence virus ; pig’s blue eye disease. of disease in humans. Lassa fever 280

Lassa fever A severe hemorrhagic fever late proteins Virus proteins produced in induced in patients infected with Lassa infected cells after the replication of the virus. viral genome has commenced. They are mainly structural proteins of the virus particle. Lassa virus (LASV) A species in the genus Arenavirus. Causes severe latency See latent infection . human hemorrhagic fever in West Africa (Nigeria, Liberia, Guinea, and latency-associated transcripts (LATs) Sierra Leone). Onset is insidious: 4 During herpesvirus latency, virus gene days of malaise, headache, fever, fol- expression is almost completely abol- lowed by severe limb and back pains, ished except for a set of LATs, synthe- diarrhea, vomiting, and severe pros- sized from a single promoter in the tration greater than expected from the repeat sequences flanking the unique degree of fever. Sore throat with white long region of the viral genome. The patches overlaid by red membrane. LATs do not appear to be translated Low blood pressure. Temperature set- into protein and their function during tles in 2 weeks. An estimated 300,000 latency is not known. cases occur annually in West Africa, causing up to 5000 deaths. In hospital- Ho DY (1992) Prog Med Virol 39 , 76 ized patients, the mortality is 15–20%. Millhouse S and Wigdahl B (2000) J Infectivity survives 1 h at 56°C. The Neurovirol 6 , 6 field rat, Mastomys natalensis , the natural host, comes into villages in the latent hamster virus See Hamster winter when most cases occur. Lassa polyomavirus . fever virus can establish a persistent latent infection An infection in which tolerant infection in this species, with infectious virus is not demonstra- copious shedding of virus in urine. ble until activated. The classic exam- Human contact with the rats or their ple is that of human herpesvirus 1 excreta may result in infection. Close infection of the dorsal root ganglia contact is probably required for case- between episodes of ‘ cold sores. ’ Only to-case transmission in humans, but a small region of the virus genome periodically iatrogenic cases or noso- is expressed during latency yielding comial outbreaks occur due to poor ‘ latency-associated transcripts ’ (LATs), clinical practices. Both ribavirin and and no infectious virus can be found. human convalescent serum have been The role of the LATs is unclear, as they found to be useful in treatment. Mice are not essential to maintain latency. can be infected and produce antibod- In response to certain stimuli, such as ies, but develop no signs except that immunosuppression, UV light, hor- some adults on stimulation may have monal changes or stress, reactivation tonic convulsions. May be isolated in may occur with production of infec- guinea pigs. Liver infection diagnosed tious virus and the reappearance of by fluorescent antibody. Liver biopsy ‘ cold sores. ’ The exact mechanism by may show virus on electron micro- which reactivation occurs is unknown. scopy. Replication in Vero cells can be All herpesviruses appear to be capable detected by fluorescent antibody. Three of establishing latent infections in the strains, Josiah, Nigeria, and AV, have nervous system of their hosts. The term been completely sequenced. ‘ latency ’ is sometimes more widely Bowen MD et al (2000) J Virol 74 , 6992 applied to indicate the presence of Childs JE and Peters CJ (1993) In The integrated viral DNA (of, e.g., adeno- Arenaviridae , edited by MS Salvato. New associated viruses, bacteriophages, or York: Plenum Press, p. 331 retroviruses) in cells, but this is clearly Gunther S (2000) Emerg Infect Dis 6 , 466 a different phenomenon from true latency, which appears to be unique to late genes Genes that are normally the Herpesviridae . Recently evidence has expressed late in infection, after repli- been published suggesting that her- cation of the genome has commenced. pesvirus latency may confer symbiotic 281 Leanyer virus (LEAV)

protection from bacterial infection by rodent, Calomys callosus, in Bolivia and upregulating innate immunity against Brazil. Not reported to cause disease in subsequent infection. humans. Barton ES et al (2007) Nature 447 , 326 Krause PR et al (1995) J Exp Med 181 , 297 LAV See Lymphadenopathy-associated virus and Human immunodeficiency latent nuclear antigen (LANA-1) An virus 1 . antigen induced in cells infected with human herpesvirus 8 which is involved LC16m8 strain A strain of vaccinia virus, in the maintenance of the episomal derived from the Lister strain, which HHV-8 genome in latently infected induces less adverse reactions during cells. Analogous to the Epstein–Barr smallpox vaccination following pri- virus nuclear antigen (EBNA) nuclear mary vaccination in humans. antigen 1, which mediates maintenance of the EBV-episome by binding to the LC-540 cell line (CCL 43) A steroid- origin of replication. secreting cell line from a transplantable Leydig cell tumor which arose latent period In experimental infection, spontaneously in the testes of an adult the time between the disappearance male rat. of the infecting virus and the appearance of new virus in the surrounding LCDV-1 Synonym for Lymphocystis disease medium. See also eclipse period . virus 1 (flounder virus). latent rat virus See Kilham rat virus . LCDV-2 Synonym for lymphocystis disease virus 2 (Dab lymphocystis disease virus). latent viral infections Infections in which the virus genome can persist in a non- LCMV Synonym for Lymphocytic chorio- replicating mode, either integrated into meningitis virus . the host cell DNA (e.g. retroviruses) or as an episome (e.g. herpesviruses). leader sequence A sequence of nucleotides at the 5 end of a mRNA molecule which lateral body A structure seen by electron precedes the AUG initiation codon. The microscopy of vertebrate poxvirus par- leader is an untranslated segment of ticles. The core of the particles appears mRNA that varies in length in different to be biconcave with two lateral bodies viruses. A leader sequence is also found in the concavities. The function of latin the virion RNA of negative strand eral bodies is not known. viruses, such as rhabdoviruses.

Lates calcarifer encephalitis virus (LcEV) leading-strand See lagging-strand . A tentative species in the genus Betanodavirus , causing behavioral leaky mutants Mutants which do not abnormalities and encephalitis in juve- conform completely to the mutant nile barramundi in Australia. characteristic: they have residual activ- Munday BL et al (1994) Aust Vet J 71 , 384 ity under the non-permissive condition. For example, a temperature-sensitive latex agglutination A serological test in mutant which has some activity at the which antibody or antigen is adsorbed restrictive temperature. on to polystyrene latex beads which are then incubated with the other reactant. Leaky virus A possible species in the Positive reactions show as aggregates genus Hantavirus . Isolated from mice, of latex particles that can be detected Mus musculus, in North America. Not readily by the naked eye. known to cause disease in humans. Puthavathan P et al (1993) Virus Res 30 , 161 Latino virus (LATV) A species in the genus Arenavirus . Belongs to the Leanyer virus (LEAV) A tentative species Tacaribe serogroup. Isolated from a in the genus Orthobunyavirus , isolated Leanyer virus (LEAV) 282

from mosquitoes in Northern Australia Lelystad virus Synonym for Porcine respi- in 1974. Not known to cause disease in ratory and reproductive syndrome virus . humans. Stuckley KG and Wright PJ (1983) Aust J Exp Lenny virus A virus isolated from a case Biol Med Sci 61 , 193 of disseminated vaccinia in a severely undernourished patient (Lenny Akpan) in Nigeria. Resembles the Wyeth strain Lebombo virus (LEBV) A species in the of vaccinia virus, producing mixed Orbivirus genus , the only member of pocks on the CAM in 48 h, and replicat- the Lebombo virus group. Isolated ing in rabbit skin; it does not produce Thryonomys swinderi- from a rodent, pocks on the CAM above 38.5°C. anus, and from mosquitoes, Mansonia africana , in Nigeria and Aedes circum- Burke ATC and Dumbell KR (1972) Bull luteolus in South Africa. The virus was World Health Organ 46 , 621 also isolated from a child in Ibadan, Nigeria in 1968. Lentivirinae (Latin: lentus slow) Name for a proposed subfamily of the fam- Brown SE et al (1991) J Gen Virol 72 , 1065 ily Retroviridae; no longer used. See Moore DL et al (1975) Ann Trop Med Parasitol Lentivirus . 69 , 49 Lentivirus A genus in the family Lebombo virus 1 (LEBV-1) The only Retroviridae distinguished from other serotype of Lebombo virus . members of the family by differences in morphology and genetic complex- Lechiguanas virus A strain of Andes virus ity. Although lentivirus assembly and in the genus Hantavirus, isolated from budding resembles that of type C retro- Oligoryzomys flavescens. Has caused viruses, the mature virions of lenti- human cases of hantavirus pulmonary virus (ca 100 nm in diameter) differ in syndrome. that they have a characteristic bar- or cone-shaped nucleoid when visualized Le Dantec virus (LeDV) An unassigned under the electron microscope. All lenti- species in the family Rhabdoviridae . With viruses contain several genes in addi- Keuraliba virus forms serogroup 4, the tion to the gag , pol , and env genes that Le Dantec group. Isolated in Senegal encode the structural and enzymatic from a girl with liver and spleen proteins of retroviruses. These differ in enlargement. Confined to West Africa different lentiviruses. The type species save for rare exceptions, e.g. a dock HIV-1 has six additional genes termed worker in Wales suffered a severe febrile vif , vpu , vpr , tat , rev , and nef , whose illness after contracting Le Dantec virus products are involved in replication. infection following an insect bite while In the HIV-2 and SIV genomes, the vpu unloading cargo from West Africa. gene is absent but a new gene, vpx , is present. Lentiviruses from other animal species contain varying numbers Lednice virus (LEDV) A strain of Turlock of such additional genes. Five sero- virus in the genus Orthobunyavirus . groups of lentiviruses have been recog- Isolated from mosquitoes, Culex modes- nized which reflect the host of origin, tus , in Moravia. as given in Table L1 . Lee virus (LEEV) A strain of Hantaan Barker E et al (1995) In The Retroviridae , vol. virus in the genus Hantavirus . Isolated 4, edited by JA Levy. New York: Plenum Press, p. 1 in cell culture from the blood of a Coffin JM (1995) Science 267 , 483 patient with Korean hemorrhagic fever Cullen B (1998) Cell 93 , 685 in 1981, after passage in Apodemus Miller RJ et al (2000) J Virol 74 , 7187 agrarius . Lee HW et al (1981) Am J Trop Med Hyg 30 , lentogenic strains A term used to 1106 describe mild or avirulent virus strains, Schmaljohn CS et al (1988) J Gen Virol 69 , especially of avian paramyxoviruses. 1949 Strains of Newcastle disease virus, e.g., 283 leukemia viruses

Table L1. The five serogroups of lentiviruses Leporipoxvirus A genus in the subfamily Chordopoxvirinae containing viruses of Serogroups Species rabbits, hares and squirrels. Infectivity is ether-sensitive. DNA cross- Bovine lentivirus Bovine immunodeficiency virus hybridization occurs between species. Equine lentivirus Equine infectious anemia Species show serological cross-reac- virus tivity. Hemagglutinin not produced. Feline lentivirus Feline immunodeficiency Mechan ical transmission by arthropods virus is common. Type species is Myxoma Puma lentivirus virus; other species are Hare fibroma Ovine/caprine Caprine arthritis virus , Rabbit fibroma virus , and Squirrel lentivirus encephalitis viru s fibroma virus . Visna/maedi virus Synonym : myxoma subgroup viruses. Primate lentivirus Human immunodeficiency virus 1 Fenner F (1994) In Virus Infections of Rodents Human immunodeficiency and Lagomorphs , edited by ADME Osterhaus. virus 2 Amsterdam: Elsevier Sciences, p. 51 Simian immunodeficiency McFadden G (2008) in BWJ Mahy and virus MHV van Regenmortel (eds) Encyclopedia of Virology , Third edition, Oxford: Academic Press, vol. 3, p. 225

have been described as lentogenic (low lethal intestinal disease of infant mice virulence), mesogenic (medium viru- virus A strain of Murine hepatitis virus lence), or velogenic (high virulence). in the genus Coronavirus which causes a severe disease in infant mice. They leporid herpesvirus 1 (LeHV-1) A tenta- do not suckle, they lose weight, become tive species in the genus Rhadinovirus . lethargic and die after a short period An indigenous virus of the cottontail of cyanosis. Older animals may have rabbit, Sylvilagus floridanus, in which it diarrhea. The disease can be produced causes a lymphoproliferative disease in in day-old mice by feeding virus. the young in 6–8 weeks. Antigenically distinct from epidemic Synonyms: cottontail rabbit herpes- diarrhea of infant mice virus. virus; lagomorph herpesvirus 1; rabbit herpesvirus. Hierholzer JC et al (1979) Infect Immun 24 , 508 Hinze HC (1971) Infect Immun 3 , 350 Kraft LM (1962) Science 137 , 282 leporid herpesvirus 2 (LeHV-2) A tenta- leucine zipper An amino acid sequence tive species in the genus Rhadinovirus . of 30 residues with leucine at every A widespread silent infection of seventh position. Found in DNA- domestic rabbits, Oryctolagus sp; ‘ blind ’ binding proteins that interact with the passage of the virus in rabbits leads to CAAT box in the region of the leucine increased virulence and the ability to zipper and enhance transcription. produce pericarditis and encephalitis. Other species are not susceptible. There Landschutz HW et al (1988) Science 240 , 1759 is replication in rabbit cell cultures, but leucopenia An abnormally low count of the virus has not yet been grown in circulating leukocytes, often seen as a eggs. result of virus infection. Synonyms: herpesvirus cuniculi; Hinze virus; virus III of rabbits. leukemia viruses Members of the genera Alpharetrovirus , Gammaretrovirus , leporid herpesvirus 3 (LeHV-3) A tenta- or Deltaretrovirus. Isolated from many tive species in the genus Rhadinovirus . species of birds and mammals, the Genome sequencing studies indicate most extensively investigated being that this virus is a distinct species from those from fowl, mice, and cats. Do leporid herpesvirus 1. not transform cells in culture but will Synonym : herpesvirus sylvilagus. replicate in them. They cause leukemia Medveczky MM et al (1989) J Virol 63 , 1010 of various types, depending upon the leukemia viruses 284

strain of virus and the strain of host leukovirus An old name for a group of animal, but usually have to be injected RNA tumor viruses now included in into newborn animals to induce the the family Retroviridae . disease. The latent period before leukemia develops may be several levamisole l -2, 3, 5, 6,-Tetrahydro-6- months. They act as helpers for sar- phenylimidazo (2,l-b )-thiazole. An coma viruses, coding for the envelope anthelmintic drug which boosts cell- of the sarcoma virus, i.e. phenotypic mediated immunity. Thus, while not mixing. Usually vertically transmitted, a directly antiviral agent, it may alter but can be passed between animals in the course of virus infection. Has been close contact, especially in cats. There used in recurrent herpetic lesions of are intraspecies group-specific anti- the skin, and in children with frequent gens and interspecies antigens. There respiratory disease. Common side is no cross-reaction between the avian effects are anorexia, diarrhea, irritabil- and mammalian gs antigens. They can ity, fatigue, nausea, and skin rashes. be grouped by their virus envelope Synonym : Ketarax. antigens and by their tropism for cells of a particular genotype: for example, L’Hoest monkey retrovirus A possible NIH Swiss (N tropic) and BALB/c (B strain of Simian immunodeficiency virus tropic). Some passaged viruses are NB in the genus Lentivirus. Originally iso- tropic. Avian virus differs from mouse lated from a wild-caught monkey in and cat viruses in having prominent a North American zoo, then found by surface spikes. Strains vary in leuke- serological analysis in 57% of wild- mogenic potential, and infection with caught L’Hoest monkeys, Cercopithecus viruses of low virulence may protect l ’ hoesti , from the Democratic Republic against strains of high virulence. of the Congo. Maeda N et al (2008) Rev Med Virol 18 , in Beer BE et al (2000) J Virol 74 , 3892 press . Liao ning virus A species in the genus leukocyte-associated herpesvirus Syno- Seadornavirus, similar to Banna virus nym for cercopithecine herpesvirus 10, but isolated from a mosquito (Aedes an unassigned member of the family dorsalis) in China. Replicates well in Herpesviridae . mammalian cells. Induced fatal hemor- Frank AL et al (1973) J Infect Dis 128 , 618, 630 rhagic disease in experimental mice. Attoui H et al (2006) J Gen Virol 87 , 199 leukocytes Strictly, all white blood cells and their precursors of both the mye- ligase An enzyme that catalyzes the join- loid and lymphoid series. Often used ing (ligation) of two different DNA or especially for granulocytes (polymor- RNA molecules, or two ends of the phonuclear leukocytes) as distinct from same molecule, by a phosphodiester lymphocytes. bond. leukopenia An abnormally decreased number of circulating white cells (leu- ligase chain reaction (LCR) A DNA kocytes) in the blood. May be associ- probe amplification method. ated with acute virus infection. Birkenmeyer LG and Mushahwar IK (1991) J Virol Methods 35 , 117 leukoplakia Painless white patches up to 3 cm diameter on the mucosal or epi- Limestone Canyon virus A probable thelial surfaces. Oral hairy leukoplakia species in the genus Hantavirus , found seen in AIDS patients occurs on buccal in brush mice (Peromyscus boylii ) in mucosa and squamous epithelial cells, Southern Arizona, USA. Differs geneti- which may contain actively replicating cally from isolates from Peromyscus EBV. maniculatus such as Sin Nombre virus. Not known to cause human disease. Greenspan JS et al (1985) N Engl J Med 313 , 1564 Sanchez AJ et al (2001) Virology 286 , 345 285 Ljungan virus (LV) line probe assay A method of determining lipovirus Obtained during attempts to the genotype of a virus by hybridizing isolate and propagate the causal agent the virus genome with oligonucleotides of human infectious hepatitis. Could immobilized on to nitrocellulose. In be propagated, and caused changes general, nucleic acid sequence determi- in cells with which it was grown, but nation is more reliable. proved to be an ameboid cell and not a virus. Similar to a Hartmannellid ameba. linker insertion mutagenesis The insertion of a linker molecule at restriction Dunnebacke TH and William RC (1967) Proc sites within an open reading frame. A Natl Acad Sci 57 , 1363 linker of 3 bp (or multiples of 3) will maintain the reading frame beyond Lipovnik virus (LIPV) A serotype of the insertion, or chain termination can Great Island virus in the genus Orbivirus , be produced in all reading frames by belonging to the Kemerovo serogroup. insertion of other linkers. Isolated from the tick, Ixodes ricinus , in the former Czechoslovakia. Not known to cause disease in humans, but anti- lion lentivirus (FIV-Ple) A possible spe- bodies have been found in 18% of the cies in the genus Lentivirus . A survey inhabitants of Lipovnik. of more than 400 free-ranging African and Asian lions, Panthera leo , revealed Lipschütz bodies Intranuclear inclusion high seroprevalence of antibodies to bodies found in cells infected with feline immunodeficiency virus (84% in human herpesvirus 1, 2, or 3. the Serengeti and 91% in the Kruger National Park). Lion lentivirus was iso- live attenuated virus lated by infection of lion lymphocytes Virus which has in vitro , but the virus also causes a lytic been attenuated to low virulence as infection in domestic cat T-lymphoma compared to wild-type virus, and is cells. No evidence of immunodefi- used as a vaccine e.g. the Sabin polio- ciency or other pathological effects of virus vaccine strains, yellow fever vac- the infection has been found so far. cine 17D, measles Edmonston strain, etc. Often accomplished by passage of Brown EW et al (1994) J Virol 68 , 5953 the wild-type virus in an alternative van deWoude S et al (1997) Virology 233 , 185 host cell. In most cases the basis of the attenuation is not known. lipid rafts Sphingomyelin- and cholesterol-enriched microdomains in the live virus vaccines Vaccines containing cellular membrane. They provide plat- virus which replicates in the recipient forms for the assembly and budding of host but is of reduced virulence as com- viruses such as influenza virus. pared to the original wild-type virus. Usually empirically derived by serial passage in cultured cells. Immunity is lipoprotein A conjugated water-soluble commonly stronger and longer-lasting protein containing a tightly bound than that following killed virus vac- lipid or group of lipids. Found in blood cines, although reversion to wild-type plasma, cell membranes, etc. virulence is sometimes a problem even with licensed live vaccines (e.g. Sabin liposome An artificially prepared lipid polio vaccine). vesicle used to introduce biological molecules (virus particles or nucleic Liverpool vervet herpesvirus Synonym acid) into cells. Liposomes may be for Cercopithecine herpesvirus 9 . uni- or multilamellar and of differing net surface charge depending on Ljungan virus (LV) A species in the the method of production and their genus Parechovirus . Isolated from bank composition. voles, Clethrionomys glareolus, near the Poste G and Papahadjopoulos D (1976) Ljungan river in Medelpad County, Methods Cell Biol 14 , 23 Sweden. Causes mild CPE without cell Ljungan virus (LV) 286

lysis in Vero or GMK cells. Particles the presence of the amino acid analog are 27 nm diameter. Some sequence canavanine. similarity is seen in the 5 non-coding Uhlenhopp EL et al (1974) J Mol Biol 89 , 689 region with mengo virus, and in other regions of the genome with echovirus Lone Star virus (LSV) 22. Causes diabetes and myocarditis in An unassigned Bunyaviridae rodents, and stress appears to be a key virus in the family . factor for the development of symp- Isolated from the Lone Star tick, Amblyomma americanum tomatic disorders in rodents. No firm . Antibodies link with disease in humans. have been found in raccoons. Found in Kentucky, USA. Not known to cause Blixt M et al (2007) Gen Comp Endocrinol disease in humans. 154 , 41 Ekstrom J-O et al (2007) Virus Res 130 ,129 Niklasson B et al (2003) Ann NY Acad Sci long terminal repeats (LTRs) Identical 1005 , 170 sequences of several hundred nucleotides which occur as direct repeats Llano Seco virus (LLSV) A serotype of at the ends of linear proviral double- Umatilla virus in the genus Orbivirus . stranded DNA of retroviruses or trans- Isolated from mosquitoes, Culex tarsa- posable genetic elements, and contain lis , in California, USA. Not known to promoters for initiation of transcrip- cause disease in humans. tion. They are formed by reverse transcription of the RNA genome of LLC-MK2 cells (CCL 7) A heteroploid retroviruses and are composed of cell line derived from a pooled suspen- sequences copied from the unique 3 sion of cells from kidneys of six adult (U3) and 5 (U5) ends. Consequently, rhesus monkeys, Macaca mulatta . each end of the provirus contains an LTR that consists of U3-R-U5, R being LLC-RK1 cells (CCL 106) An aneuploid a DNA copy of a sequence found near epithelial-like cell line derived from both ends of the genome RNA. pooled kidneys of several New Zealand white rabbits, Oryctolagus cuniculus , of lookback studies Retrospective studies undetermined sex. to determine the presence of a virus infection in a population. Applied par- LLC WRC 256 cells (CCL 38) A hetero- ticularly to hepatitis C, which could ploid cell line derived from a Walker not be diagnosed until 1990, when tests rat, Rattus norvegicus, carcinoma main- were developed. Lookback studies tained in adult Harlan–Wistar rats. were then applied to find seropositive lobular hepatitis A type of hepatitis seen persons infected earlier for counseling as a result of virus infection, character- and treatment for hepatitis C infection. ized by lymphocytic infiltration within sinusoids surrounding the hepatocytes. Lopinavir A highly active antiretroviral drug which interferes with process- local infection The infection of a few cells ing of viral polyprotein precursors. of a host which is prevented by the host When used in combination, ritonavir response from spreading systemically. enhances the concentration of lopinavir achieved by oral administration. Lokern virus (LOKV) A serotype of Bunyamwera virus in the genus Lordsdale virus (LDV) A strain of Orthobunyavirus . Isolated from the hare, Norwalk virus in the genus Norovirus . Lepus californicus , rabbit, Sylvilagus auduboni, mosquitoes, Culex tarsalis and Dingle KE et al (1995) J Gen Virol 76 , 2349 Culicoides variipennis , in Kern County, California, USA. Not reported to cause lorisine herpesvirus 1 (LoHV-1) An disease in humans. unassigned species in the family Herpesviridae. Isolated from kinka- lollipops Abnormal particles of T-even jou, (Potos flavus ), skin and kidney cell bacteriophage produced in bacteria in culture showing spontaneous CPE. 287 Lumbovirus (LUMV)

Replicates in a narrow range of cell louping ill virus Turkish subtype A cultures: kinkajou, owl monkey, and strain of Louping ill virus found in Vero cells with CPE. Produces A-type Bulgaria and Turkey. intranuclear inclusion bodies. Probably non-pathogenic for kinkajou, rabbit, LP virus A strain of Lassa virus . and owl monkey. Synonyms: kinkajou herpesvirus; kinka- LPMV Abbreviation for L a- P iedad jou kidney virus; herpesvirus pottos. M ichoacan virus. See Porcine rubula- Barahona HH et al (1973) Lab Anim Sci 23 , virus . 830 L-R cells Leukosis virus-negative Rous Louping ill virus (LIV) A species in the cells. A name suggested for cells trans- genus Flavivirus . A member of the mam- formed by Rous sarcoma virus which malian tickborne virus group. There are not producing infective virus par- are four recognized subtypes: British, ticles. They were formerly known as Irish, Spanish, and Turkish. Causes non-producer cells but by electron an acute meningo-encephalomyelitis microscopy can be seen to be produc- disease in sheep and, less often, in cating virus particles indistinguishable tle, deer, and man. The disease has two from the infective ones. The term is not phases: in the first there is fever and in common use. viremia; and in the second, several Hanafusa H and Hanafusa T (1968) Virology days later, in-coordination of move- 34 , 630 ment followed by paralysis and often death. The second phase may be absent LuIII virus (LUIIIV) A species in the and probably many infections are subgenus Parvovirus . Isolated from a line clinical. Laboratory workers and men of human lung cells Lu 106, originating in contact with sheep may be infected, from Stockholm, Sweden. The natural showing signs of severe meningitis host is unknown. Can be maintained and some encephalitis. Rodents, deer, in HeLa cells. The Douglas chim- shrews, and red grouse may be natu- panzee cell line is highly susceptible. rally infected without clinical disease. Agglutinates erythrocytes of the same Horses may also be infected experi- species as H-1 and latent rat virus, but mentally. Injection i.c. in mice causes differs from these viruses in replicat- encephalitis. Pigs can be infected but ing in human cells and not in rodent not guinea pigs or rabbits. Ixodes ricinus cells. On passage in newborn hamsters is the vector and the sole natural reser- becomes adapted and causes massive voir of infection. A formalinized vac- intestinal hemorrhage. In pregnant cine is available for sheep and humans. hamsters it crosses the placenta, infects Synonym: ovine encephalomyelitis the fetus and causes abortion. virus. Soike KF et al (1976) Arch Virol 51 , 235 Heinz FX and Kunz C (1982) J Gen Virol 62 , 271 Lucké frog herpesvirus Synonym for Laurenson MK et al (2007) Epidemiol Infect ranid herpesvirus 1. 135 , 963 McGuire K et al (1998) J Gen Virol 79 , 981 Lukuni virus (LUKV) A serotype of Anopheles A virus in the genus louping ill virus British subtype A strain Orthobunyavirus . Isolated from mosqui- of Louping ill virus found in Scotland, toes in Trinidad and in Belem, Brazil. and in north and south-west England. Not associated with disease in humans. louping ill virus Irish subtype A strain Lumbovirus (LUMV) A serotype of of Louping ill virus found in Ireland. California encephalitis virus in the genus Orthobunyavirus. Originally isolated louping ill virus Spanish subtype A from the mosquito, Aedes pembaensis , strain of Louping ill virus found in in Mozambique. Antigenically indis- Spain. tinguishable from Tahyna virus but Lumbovirus (LUMV) 288

distinct from other members of the who was also in a group at high risk California serogroup. for AIDS. The prototype virus was Ozden S and Hannoun C (1978) Virology 81 , named LAV LAI, after the initials of 210 the patient. Barré-Sinoussi F et al (1983) Science 220 , 868 Lumpy skin disease virus (LSDV) A species in the genus Capripoxvirus , which lymphoblastoid cell lines Cell lines usu- causes a severe disease characterized ally derived from culture of periph- by high fever with multiple skin nod- eral blood leukocytes. Usually grow ules and lesions in viscera of cattle in in suspension and do not attach to Africa. Virus is present in saliva for at glass or plastic. Frequently contain the least 11 days and in semen of bulls for Epstein–Barr virus (EBV) genome in 22 days. The disease is spread by biting latent form. Can be superinfected by flies (which probably act as mechani- EBV and a variety of other viruses. cal short range vectors) and is most prevalent in the central and southern Lymphocryptovirus A genus in the regions. Virus attenuated by egg pas- family Herpesviridae , subfamily sage and tissue culture is used as a vac- Gammaherpesvirinae. Human herpesvirus cine for cattle, but is highly virulent for 4 (Epstein–Barr virus) is the type spe- sheep and goats. An attenuated sheep- cies and Cercopithecine herpesviruses 12 , pox virus vaccine which also protects 14 , and 15 , Pongine herpesviruses 1 , 2 cattle against lumpy skin disease has and 3 are other members. The viruses been described. A feature of the disease all have a distinctive genome struc- is that epidemics recur after an inter- ture. In EBV, the virion DNA of about val of 5 or 6 years in an unvaccinated 180 kb is linear and bounded by termi- cattle population. The virus has been nal repeats of about 500 bp, which fuse isolated from flies, Biomyia fasciata and after infection of lymphocytes to form Stomoxys calcitrans, caught on infected a circular DNA episome. Internally animals, and these are possible vectors the genome contains short (12 kb) and of transmission. long (134 kb) unique sequence regions Synonyms: Neethling virus; exanthema separated by multiple copies of a direct nodularis bovis; Knopvelsiekte. repeat. The genome contains about 90 Cheneau Y et al (1999) Rev Sci Tech 18 , 122 genes, 10 of which express proteins in Gershon PD et al (1989) J Virol 63 , 4703 latently infected cells. Kitching RP (2003) Dev Biol 114 , 161 Sandberg ML et al (2000) J Virol 74 , 9755 Weiss KA (1968) Virol Monogr 3 , 131pp Sugden B (1994) Semin Virol 5 , 197 Woods JA (1988) Trop Anim Health Prod 20 , 11 Lymphocystis disease virus 1 (LCDV-1) Lundy virus (LUNV) A serotype of Great The type species in the genus Island virus in the genus Orbivirus , in Lymphocystivirus which causes a benign the Kemerovo serogroup. superficial cellular hypertrophy in a wide range of marine and freshwater Lungers virus Synonym for Ovine pulmo- fish including flounder and plaice. nary adenocarcinoma virus . Particle size ranges from 198 to 227 nm. Infectivity is ether-sensitive. The Lwoff-Horne-Tournier scheme An early genome of about 150 kb contains 195 virus classification scheme based potential open reading frames. Causes mainly upon morphology. acute to chronic (5 days to 9 months) benign hypertrophic lesions on the lymphadenopathy-associated virus gills and skin, which contain greatly (LAV) The name given by French enlarged cells that eventually degen- workers at the Pasteur Institut to the erate with release of infectious virus. first-reported isolate of human immuno– Transmission is horizontal by con- deficiency virus (HIV). This virus was tact with other fish. Causes a massive recovered from a person with lym- enlargement of cells (up to 100,000- phadenopathy (enlarged lymph nodes) fold). The virus genome DNA (about 289 Lymphocytic choriomeningitis virus (LCMV)

200 kb long) is circularly permuted, containing electron-dense granules terminally redundant and highly meth- appearing at the time of budding from ylated. A one-step growth cycle of the the cell surface. Nucleic acid single- virus at 25°C takes about 4 weeks. stranded RNA separable into two seg- Synonym: flounder lymphocystis dis- ments: L, 7.2 kb and S, 3.4 kb in length. ease virus; flounder virus. The 3 terminal sequences are similar Tidona CA and Darai G (1997) Virology 230 , 207 (19–30 nt) between the two RNA seg- Tidona CA and Darai G (1997) Arch Virol ments. Coding sequences are ambisense. Suppl 13 , 49 The L segment encodes the L protein (2210 amino acids) in negative sense and Lymphocystis disease virus 2 (LCDV-2) two small proteins X (95 amino acids) A tentative species in the genus and Z (90 amino acids) in positive sense. Lymphocystivirus, varying in size from The S segment encodes the NP protein 130 to 300 nm. Lymphocystis is a com- (558 amino acids) in negative sense and mon, chronic, but rarely fatal dis- the GP protein (498 amino acids) in pos- ease affecting dab (Limanda limanda ) itive sense. In addition, virions encap- and other teleost fish. The infection is sidate variable amounts of full-length found worldwide in numerous spe- complementary RNAs and subgenomic cies of fish, both marine and fresh- mRNAs as well as cellular RNAs such water. Tumor-like masses appear on as ribosomal RNAs. The virus repli- the skin and fins, persist for long peri- cates on the CAM but no lesions are ods, but ultimately regress. They are produced. Also replicates in chick, caused by massive increases (100,000- mouse, cattle, and monkey cell cultures, fold) in the size of infected cells, which but CPE may only be seen on adapta- contain large cytoplasmic inclusions. tion. In mouse cell cultures a persistent, Outbreaks in the wild occur mainly non-cytolytic infection is established. in the summer. Parasites may help to Replication is inhibited by actinomy- spread the disease, but the large lym- cin D because host cell mRNAs are phocystis cells burst in water and may required as primers for RNA transcrip- release infective virus. Transmission is tion. Probably an inapparent infection in possible by applying infectious lesion naturally infected house mice, but has material to abraded skin, but may be been isolated from humans, monkeys, difficult between species. The bluegill, dogs, field mice of Apodemus sp, ham- Lepomia macrochirus , is a useful experi- sters, and guinea pigs. When a colony mental subject. of laboratory mice becomes infected, Synonym: dab lymphocystis disease disease appears in young mice infected virus. in utero but the infection soon becomes Weissenberg R (1965) Ann NY Acad Sci 126 , latent. In guinea pigs there is a gener- 362 alized disease, often fatal, with patchy lymphocyte A spherical, non-phago- pneumonia. In humans infection may cytic leukocyte white blood cell with a be inapparent or an influenza-like fever large nucleus which is involved in the but lymphocytic choriomeningitis of specific immune response. Divided by varying severity may occur. An increas- ontogeny and function into B lympho- ingly recognized cause of congenital cytes (humoral immunity) and T disease, which may occur in pregnant lymphocytes (cell-mediated immunity). women who have contact with pet mice Found in blood, lymph and lymphoid and so become infected with LCMV. tissues. When activated by antigen the Mice infected i.c. develop tremors and cells enlarge and proliferate and are have tonic convulsions during which called ‘ lymphoblasts. ’ they may die. Lymphocytic choriomeningitis virus Barton LL and Mets MB (2001) Clin Infect Dis 33 , 370 (LCMV) The type species of the genus Buchmeier MJ and Zajuc AJ (1999) In Arenavirus. In sections virions are ple- Persistent Viral Infections , edited by R Ahmed omorphic, 110–130 nm in diameter, and ISY Chen. New York: John Wiley, p. 575 consisting of a bilayered envelope Salvato MS (Editor) (1993) The Arenaviridae . with thin projections and an interior New York: Plenum Press lymphoid leukemia viruses 290 lymphoid leukemia viruses A name used may exceed 20%. Can be transmitted for mouse leukemia viruses, usually of by injecting the cell-free serum of dis- low leukemogenic potency, which may eased birds into young birds, but has be separated from Friend or Rauscher not been isolated in vitro . Cross-nucleic leukemia viruses. They cause immuno- acid hybridization tests and sequence depression and may also act as a helper analysis distinguish LPDV from avian virus. sarcoma-leukemia viruses or avian reticuloendotheliosis viruses. lymphokines Heterogeneous group of Chajut A et al (1992) Gene 122 , 349 cytokines secreted by T lymphocytes Davison I and Borenstein B (1999) Acta Virol which function as regulators of the 43 , 136 immune response. Examples are interleukin 2 and interferon γ. lymphoproliferative herpesvirus group Synonym for Gammaherpesvirinae . lymphoproliferative assays Lymphocyte lymphotoxins proliferation in response to an anti- Cytotoxic products of T-cells, particularly tumor necrosis genic stimulus can be used to measure β the recall response to the antigen and factor . provides an in vitro correlate of the lymphotropic papovavirus A strain of delayed-type hypersensitivity response. African green monkey polyomavirus in Only persons exposed to a given virus the genus Polyomavirus . Isolated from a by natural infection or vaccination will B lymphoblastic cell line derived from show a lymphoproliferative response an African green monkey. Grows only to it when their lymphocytes are tested in dividing B lymphocytes. in the presence of antigen-presenting cells in culture. lymphotropism Affinity for replication in Mawle AC (1996) In Virology Methods lymphocytes. Shown by some viruses Manual , edited by BWJ Mahy and HO and prions. Kangro. London: Academic Press, p. 147 lyophilization Rapid freezing of a mate- lymphoproliferative disease (LD) A rial at low temperature accompanied progressive disease caused by EBV by rapid dehydration by sublimation (HHV4) infection in immunocompro- in a high vacuum. A method used to mised individuals such as transplant preserve biological specimens or to recipients and those infected with HIV. concentrate macromolecules with little In most cases the proliferating cells are or no loss of activity. of B-cell origin, but about 12% are of T- Synonym : freeze-drying. cell origin. If untreated, the disease can evolve into immunoblastic lymphoma lysis from without Lysis due to adsorp- or multiple myeloma. tion of a large number of virus particles to the host cell surface. Nalesnik M (1998) Springer Semin Immun o- pathol 20 , 325 lysogenic Having a prophage integrated into the bacterial genome. The prophage lymphoproliferative disease of turkeys may become activated spontaneously or virus An unassigned species in the under the influence of certain stimuli, family Retroviridae . Present in the blood when it will replicate phage particles of turkeys with lymphoproliferative and destroy the cell, releasing infective disease, an acute infection occurring bacteriophage. Lysogenic bacteria may only in turkeys. No other natural host have certain properties determined by is known, but chickens can be infected the prophage, such as diphtheria toxin experimentally. There is splenom- production or antigens. egaly, hepatomegaly, and infiltration of other organs and nerves with pleo- lyssa virus Synonym for Rabies virus . morphic mononuclear cells. The disease is of economic importance as in Lyssavirus A genus of the family birds aged 10–12 weeks the mortality Rhabdoviridae . Morphologically similar 291 lytic viruses

to species in the genus Vesiculovirus Rupprecht CE et al (1994) Curr Top Microbiol but antigenically distinct. The viral Immunol 187 , 352pp envelope may be formed within the Tordo N et al (2005) In Topley & Wilson’s cytoplasm or by budding from the Microbiology and Microbial Infections , vol. 2, Tenth edition, edited by BWJ Mahy and plasma membrane. Virions agglutinate V ter Meulen, London: Hodder Arnold, goose erythrocytes. There is serologi- p. 1102 cal cross-reaction between species, but Vazquez-Moron S et al (2006) J Virol Methods the viruses can be grouped into four 135 , 281 serotypes or seven genotypes. Type species is Rabies virus . Other species lytic cycle A term usually applied to the are Australian bat lyssavirus , Duvenhage productive replicative cycle in virus virus , European bat lyssavirus 1 , European infections (particularly phage) which bat lyssavirus 2 , Lagos bat virus and ends with the production of progeny Mokola virus . Rochambeau virus, virus particles and their release from Aravan virus, Khujand virus, Irkut the host, most usually by lysis of the virus, and West Caucasian bat virus host cells. are all tentative species in the genus. The arthropod isolates, Kotonkon and Obodhiang viruses, formerly assigned lytic viruses Viruses whose replica- to the genus, have features in common tion causes destruction (lysis) of the with members of the Ephemerovirus host cell. genus, and are presently unassigned viruses within the family Rhabdoviridae . Jallet C et al (1999) J Virol 73 , 225 M

M virus Synonym for Saimiriine herpes- Calomys callosus. Experimental infec- virus 1 (marmoset herpesvirus). tion of guinea pigs causes subclinical infection but in marmosets, Saguinus M14 virus Chinese isolate of Banna geoffroyi , the infection is fatal. virus, a tentative species in the genus Synonym : Bolivian hemorrhagic fever Seadornavirus . virus.

M459 virus (BWAV) A serotype of Maciel virus A strain of Andes virus iso- Bwamba virus in the genus lated in Argentina from the rodent Orthobunyavirus . Bolomys obscurus .

M67U5 virus (MNTV) A serotype of macromolecule A molecule having a Minatitlan virus in the genus molecular weight in the range of more Orthobunyavirus . than 1000 kD to many millions.

MA104 cells An epithelial cell line estab- macrophage A mononuclear cell of the lished from an African green monkey reticuloendothelial system that func- fetal kidney. tions in phagocytosis. Relatively long-lived. There are different types Maastricht strain A strain of rat cyto- depending on the tissue of origin, e.g. megalovirus (Murid herpesvirus 2) in the peritoneal and alveolar macrophages, genus Muromegalovirus . Because of major Kupffer cells of the liver, histiocytes sequence differences from the England (tissue macrophages), and osteoclasts. strain, it has been proposed that these Play an important first defense role strains represent distinct species. against foreign substances, including Beisser PS et al (1998) Virology 246 , 341 bacteria, viruses, and protozoa; releasing chemokines that stimulate other Macauâ virus (MCAV) A serotype cells of the immune system; and play- of Wyeomyia virus in the genus ing a role in antigen presentation. Orthobunyavirus . Isolated from mosquitoes, Sabethes soperi, and once from a macrophage activating factor A general rodent, Proechimys guyannensis, in Brazil. term for any cytokine that activates Not associated with disease in humans. macrophages.

Machupo virus (MACV) A species in macrophage chemotactic proteins A the genus Arenavirus belonging to the set of at least three infection-induced Tacaribe serogroup. One of the South chemokines which attract inflamma- American hemorrhagic fever viruses. tory mononuclear cells. They can act First recognized in 1959 as the cause of to promote the migration of mono- Bolivian hemorrhagic fever, which is cytes, lymphocytes, eosinophils, and severe and fatal in some 25% of cases. basophils across the blood–brain bar- Hemorrhages occur, with rash, myal- rier into the brain. gia, CNS involvement, and conjuncti- Persidsky Y (1999) J Neurovirol 5 , 579 val inflammation. Sporadic outbreaks occur in the Beni region of Bolivia. macrophage colony-stimulating fac- The most notable of these affected 700 tor ( M-CSF) A homodimeric cytokine people in one town and there was 18% made by lymphocytes which stimu- mortality. Transmission from humans lates committed stem cells of the bone to humans is unusual but does occur. marrow to differentiate toward the pro- The only known reservoir is a rodent, duction of mononuclear phagocytes. 293 Mahogany hammock virus (MHV) macrophage inflammatory protein (MIP) Madin–Darby canine kidney (MDCK) A chemokine which is one of four mem- cells An epithelial-like cell line estab- bers of the MIP-1 CC chemokine sub- lished from the kidney of a normal family. They are CCL3 (MIP1-α ), CCL4 cocker spaniel in 1958. Widely used (MIP-1β ), CCL9/10 (MIP-1δ ), and for virus growth studies, including CCL15 (MIP-1γ ). primary isolation of human influenza viruses. See MDBK (NBL-1) cells . Macropodid herpesvirus 1 (MaHV-1) A species in the genus Simplexvirus , sub- mad itch virus Synonym for pseudora- family Alphaherpesvirinae . Isolated from bies virus. a culture of kidney cells of a parma wallaby, Macropus parma. The animal Madrid virus (MADV) A species in the was one of a number with a fatal gen- genus Orthobunyavirus , belonging to eralized disease taken from Kawan the group C serogroup. Isolated from Island in Auckland Bay, New Zealand. a man with a febrile illness in Panama. The culture developed foci of CPE Also isolated from sentinel mice in the which extended rapidly. Experimental same area, the mosquito, Culex vomer- infection of Parma wallabies causes ifer, and the spiny rat, Proechimys semi- a severe generalized disease with spinosus . Antibodies found in 3 of 96 lesions in lungs and liver. Antibodies humans but no further cases of disease are found in a wide range of macro- reported. pods (kangaroos and wallabies) from different parts of Australia. No CPE in MADT (morphological alteration and bovine, mouse, or hamster cells. The disintegration test) An electron micro- DNA is distinct from other herpes- scopy-based assay for testing for chem- viruses, G C 53%. ical inactivation of hepatitis B virus. Synonym : Parma wallaby herpesvirus. Prince DL et al (1993) J Clin Microbiol 31 , Johnson MA and Whalley JM (1987) Arch 3296 Virol 96 , 153 Johnson MA et al (1985) Arch Virol 85 , 313 maedi-visna virus (Icelandic: maedi dyspnea) See Visna/maedi virus . Macropodid herpesvirus 2 (MaHV-2) A species in the genus Simplexvirus , subfamily Alphaherpesvirinae . Isolated magnetic resonance imaging (MRI) A from cultured cells of the dorcopsis noninvasive method using nuclear wallaby, Dorcopsis muelleri luctuosa . magnetic resonance to render images The genome DNA is 135 kb in length, of the inside of the body. It has been G C 49.5%. used to detect fetal abnormalities in Synonym: dorcopsis wallaby herpes- pregnant women exposed to human virus. cytomegalovirus. Malinger G et al (2002) Ultrasound Obstet et al Arch Virol 85 Johnson MA (1985) , 313 Gynecol 20 , 317 Johnson MA and Whalley JM (1987) Arch Virol 96 , 153 Maguari virus (MAGV) A serotype mad cow disease A popular name for of Bunyamwera virus in the genus the prion disease, bovine spongiform Orthobunyavirus. Isolated from senti- encephalopathy (BSE). nel mice and mosquitoes in Brazil and from a horse in Guyana. Present also in Trinidad, Colombia, and Argentina. Madin–Darby bovine kidney (MDBK) Antibodies found in cattle, sheep, and cells The first permanent cell line of a birds. Not reported to cause disease in large domestic animal, derived from humans. the kidney of a normal adult steer in 1957. Mahogany hammock virus (MHV) Madin SH and Darby NB (1958) Proc Soc A serotype of Guama virus in the Exp Biol Med 98 , 574 genus Orthobunyavirus. Isolated from Mahogany hammock virus (MHV) 294

mosquitoes and a cotton rat in Florida. and rat (Rt1). See also human leuko- Not known to cause disease in humans. cyte antigens . Brutkiewicz RR and Welsh RM (1995) J Virol Mahoney strain A strain of human polio- 69 , 3967 virus 1 isolated in 1941 in Michigan Doherty PC (2002) Immunol Rev 185 , 39 from a pool of stool specimens of three Mahoney children aged 10, 8, and 3 Makonde virus Synonym for Uganda S years. virus in the genus Flavivirus .

Maiden virus (MDNV) A serotype of Malabar grouper nervous necrosis Great Island virus in the genus Orbi- virus A tentative species in the genus virus . Betanodavirus .

MAIDS See murine AIDS . Malacky/Ma32/94 virus A strain of Tula virus in the genus Hantavirus . Main Drain virus (MDV) A species in the maladie de jeune age Synonym for genus Orthobunyavirus , belonging to Canine distemper virus . the Bunyamwera serogroup. Isolated from the hare, Lepus californicus , and Malakal virus (MALV) A tentative Culicoides variipennis . Antibodies found species in the genus Ephemerovirus . in cattle, sheep, and wild animals. The Isolated from the mosquito, Mansonia virus is found in Kern and Mendocino uniformis, in the Sudan. Not reported to counties, California, USA. Not known cause disease in humans. to cause disease in humans. Calisher CH et al (1989) Intervirology 30 , 241 major histocompatibility complex (MHC) A genetic locus in vertebrates mal de los rastrojos (corn pickers ’ dis- consisting of numerous histocompat- ease) A local name for Junín virus ibility genes controlling cell surface infection. immune response determinants and components of the complement sys- malignant aphtha virus A severe form of tem. There are three MHC class I pro- Orf virus infection. teins found at the plasma membrane of most cell types, and at least six MHC malignant catarrhal fever of European class II proteins found on the surface cattle See Alcelaphine herpesvirus 1 . of antigen-presenting cells (T cells, B cells, and macrophages). They occur malignant catarrhal fever virus Synonym in many alternative allelic forms and, for Alcelaphine herpesvirus 1 . depending on their amino acid structure, bind particular peptides, e.g. malignant rabbit fibroma virus A strain processed viral antigens, at their distal of Rabbit fibroma virus in the genus tip to form a peptide–MHC complex. Leporipoxvirus . This complex is recognized by T-cell receptors and their co-receptors CD4 Malpais Spring virus (MSPV) A tenta- for MHC class II (helper T cells) or tive species in the genus Vesiculovirus . CD8 for MHC class I (cytotoxic T cells). Isolated from mosquitoes, Aedes camp- Positive selection of developing T cells estris , in New Mexico, USA. in the thymus by ‘self ’ MHC molecules results in mature T cells that only rec- Mamastrovirus A genus in the family ognize the foreign peptides if they are Astroviridae containing only viruses presented by a ‘ self ’ MHC protein. This which infect mammalian species. is termed ‘ MHC restriction. ’ In differ- Cause gastroenteritis. The type species ent vertebrates different abbreviations of the genus is Human astrovirus . are used to describe the MHC system, e.g. chicken (B), dog (DLA), guinea pig Mammalian orthoreovirus (MRV) The (GPLA), human (HLA), mouse (H2), type species of the genus Orthoreovirus . 295 Mammalian virus group

There are four serotypes: serotype 1 orthoreovirus in the genus Orthoreovirus . (strain Lang); serotype 2 (strain D5/ The type strain is Dearing. Isolated Jones); serotype 3 (strain Dearing); and from a wide range of mammalian serotype 4 (strain Ndelle). Cause benign species including humans, mice, cat- infections in humans worldwide, only tle, and quokkas (short-tailed walla- rarely associated with mild respiratory bies). Antibodies have been detected or enteric symptoms. In experimental in sera from a wide range of species. mice, MRV can cause diarrhea, runting, Experimental infection of mice causes oily hair syndrome, hepatitis, myocar- jaundice, alopecia, conjunctivitis, and ditis, myositis, pneumonitis, encepha- ‘ oily hair ’ associated with steatorrhea. litis, and neurological symptoms. This The mice appear to have been dipped mouse model has been widely used to in oil, and if newborn mice are infected investigate the molecular basis of viral their growth is stunted and they pathogenesis even though reovirus is become runts. There is no clear associa- not a significant human pathogen. tion with disease in humans and infections are usually symptomless. mammalian orthoreoviruses Members of the Orthoreovirus genus, family mammalian orthoreovirus 4 - Ndelle Reoviridae . (MRV-4) A serotype of Mammalian orthoreovirus in the genus Orthoreovirus . mammalian orthoreovirus 1-Lang The type strain is Ndelle. Isolated from (MRV1) A serotype of Mammalian Mus musculoides in Cameroun and from orthoreovirus in the genus Orthoreovirus . ticks and mosquitoes in Senegal. The type strain is Lang. Recovered from a number of mammalian spe- mammalian rotaviruses Rotaviruses cies including humans, dogs, cattle, of groups A, B, C, and E in the genus macaque monkeys, and cercopithe- Rotavirus . cine monkeys. Antibodies have been detected in sera from a wide range of mammalian type B oncovirus group See mammalian species. Experimental Betaretrovirus . infection of mice, hamsters, ferrets, and rats causes obstructive hydrocepha- ‘ Mammalian type B retroviruses ’ A lus; and in pregnant mice fetal resorp- former genus in the family Retroviridae , tion, intrauterine death, malformation, now named Betaretrovirus . or neonatal death occurs. In humans infection is usually symptomless or ‘ Mammalian type C retroviruses ’ A associated with mild upper respiratory former genus of the family Retroviridae , disease (common cold). Rarely, reo- now named Gammaretrovirus . virus type 1 has been isolated from fatal encephalitis cases in children. Mammalian virus group A subgroup of the genus Gammaretrovirus contain- mammalian orthoreovirus 2-D5/Jones ing five replication-competent viruses (MRV-2) A serotype of Mammalian and eight replication-defective sarcoma- orthoreovirus in the genus Orthoreovirus . inducing viruses ( Table M1 ). The The type strain is D5/Jones. Recovered genome is single-stranded RNA, 8.3 kb from a number of mammalian spe- in length, and the tRNA primer is tRN- cies including humans, mice, cattle, APro. The LTR is about 600 bases in macaque monkeys, and chimpanzees. length (U3-500, R-60, U5-75). There Antibodies have been detected in sera are two cross-reacting antigens: those from a wide range of mammalian spe- shared by virus from one host species cies. Type 2 reovirus strains show con- (gs-1 species-specific antigens) and siderably more antigenic variation than those shared by different virus spe- types 1 or 3. Little evidence of associa- cies from different mammalian hosts. tion with disease in humans. There are two interspecies antigens, a and b, both present in the mouse virus, mammalian orthoreovirus 3-Dearing but in the feline, porcine and woolly (MRV-3) A serotype of Mammalian monkey viruses there is only b. The Mammalian virus group 296

Table M1. Mammalian virus group transcriptase or other viral proteins, and by nucleic acid hybridization. They Replication-competent Replication-defective may be ecotropic (replicating only in viruses viruses the species of origin), xenotropic (un- Feline leukemia virus Finkel–Biskis–Jinkins able to replicate in the species of origin), murine sarcoma or amphitropic (can replicate in the spe- virus cies of origin and in other species). The Gibbon ape leukemia Gardner–Arnstein mammalian type C viruses of different virus feline sarcoma virus species appear to differ in their mode of Guinea pig type C Hardy–Zuckerman transmission and in disease production. oncovirus feline sarcoma virus Murine leukemia virus Harvey murine mammary tumor viruses Viruses in ( MLV ) sarcoma virus the genus Betaretrovirus. The only Abelson MLV Kirsten murine well-characterized member is Mouse sarcoma virus AKR (endogenous) Moloney murine mammary tumor virus but related endo- MLV sarcoma virus genous viruses have been detected in a Friend MLV number of primate and rodent species. Murine leukemia Snyder–Theilen feline virus sarcoma virus Manawa virus (MWAV) A serotype Moloney MLV Woolly monkey of Uukuniemi virus in the genus sarcoma virus Phlebovirus . Isolated from ticks, Argas Porcine type C oncovirus abdussalami and Rhipicephalus sp, in western Pakistan. Not reported to cause disease in humans.

sarcoma virus genomes contain addi- Manchester virus A strain of Sapporo tional genetic information (oncogene) virus in the genus Sapovirus . Isolated which, when integrated into a host cell, from a boy aged 6 months. Complete causes transformation to a neoplastic sequencing of the genome (7266 nt) cell, but they may lack the informa- suggested a virus more closely related tion to make their coat proteins and are to animal caliciviruses than to enteric then defective, being unable to produce small round structured viruses such as infective virus particles without the help Norwalk. of the leukemogenic viruses. The helper Liu BL et al (1995) Arch Virol 140 , 1345 virus provides the coat proteins and thus determines the host range of the Mandarin duck hepadnavirus A novel progeny sarcoma virus. Not all type C hepadnavirus cloned from a captive viruses are oncogenic since they may mandarin duck. The genome nucleo- lack the oncogene. The genetic informa- tide sequence is similar to that of Ross tion of some type C viruses, the viro- goose hepatitis B virus. gene, which may or may not contain Guo H et al (2005) J Virol 79 , 2729 the oncogene, is present in all the cells of many species of animal. These viro- Mandarin fish rhabdovirus A rhabdo- genes are replicated with the cell genetic virus isolated from mandarin fish, material. The virogene on activation Siniperca chuatsi, cultivated in China. can produce a virus which may or may The complete genome of 11,545 nt not be oncogenic. The role of these was sequenced and found to be viruses in the development of sponta- most closely related to viruses in the neous tumors is not clear. Laboratory Vesiculovirus genus. strains of C-type oncoviruses replicate and produce much free virus: they Tao J-J et al (2008) Virus Res 132 , 86 are ‘ exogenous. ’ In contrast, the integrated C-type oncoviruses only appear Manitoba virus (MNTBV) An unassigned as virus particles on activation and are vertebrate rhabdovirus. Isolated from termed ‘ endogenous. ’ When produced the mosquito, Culex tarsalis , in Manitoba, they may be detected by electron micro- Canada. Asymptomatic in mice. scopy, by the presence of their reverse Artsob H et al (1991) Can J Microbiol 37 , 329 297 Marboran mannose-binding protein One of a group complex (PIC). It mediates the plasma of proteins (collectins) that can act as membrane targeting of the Gag poly- part of the innate immune response to protein and lines the inner shell of the interact with viruses such as HIV, HSV, mature virus particle. or influenza to activate the classical complement pathway and neutralize map turtle virus Synonym for chelonid virus infectivity. herpesvirus 3. Malhotra R and Sim RB (1995) Trends Microbiol 3 , 240 Mapuera virus (MPRV) A species in the genus Rubulavirus. Isolated from the M-antibody capture radioimmunoassay salivary glands of a fruit bat, Sturnira (MACRIA) An assay for IgM antibody lilium , captured in the tropical rain- developed for use in rubella virus forest of Brazil in 1979. diagnosis. Henderson GW et al (1995) J Gen Virol 76 , Mortimer PP et al (1981) J Hyg (Camb ) 86 , 139 2509

Manzanilla virus (MANV) A species in Maraba virus (MARAV) A species in the the genus Orthobunyavirus, related to genus Vesiculovirus . Isolated from the the Simbu serogroup. Isolated from the sand fly, Lutzomyia sp. howler monkey, Alouatta siniculus insu- Travassos da Rosa APA et al (1984) Am J Trop laris , in Trinidad. Not associated with Med Hyg 33 , 999 disease in humans. Marajo virus An unclassified virus iso- Maporal virus (MAPV) A probable lated from Culicine mosquitoes in the member of the genus Hantavirus , iso- Amazon region of Brazil. Not known lated from two fulveus pigmy rice to cause disease in humans. rats (Oligoryzomys fulvescens ) captured in western Venezuela. When inocu- marble bone disease virus lated into the Syrian golden hamster Synonym for (Mesocricetus auratus) it caused symp- osteopetrosis virus. toms similar to those seen in hantavirus pulmonary syndrome in humans. marble spleen disease of pheasants A disease of pen-raised ring-necked Fulhorst CF et al (2004) Virus Res 104 , 139 pheasants which occurs in Europe and North America, caused by Pheasant Mapputta virus (MAPV) An unas- adenovirus 1, a species in the genus signed virus in the family Bunyaviridae . Siadenovirus . Young adult birds die Serologically related to Gan Gan, after an inapparent or short illness due Maprik, and Trubanaman viruses. to pulmonary edema. The spleen is Isolated from Anopheles meraukensis enlarged with extensive necrosis and in Queensland, Australia. Antibodies amyloidosis. The main route of infec- present in humans, cattle, horses, pigs, tion appears to be fecal–oral. A virus kangaroos, and rats. Not reported to with the morphology of an adenovirus cause disease in humans. can be seen in the diseased tissue and Boughton CR et al (1990) Aust NZ J Med 20, 51 extracted, but has not been replicated in cell culture. Related antigenically Maprik virus (MPKV) An unassigned to turkey adenovirus strain 3, which virus in the family Bunyaviridae . causes turkey hemorrhagic enteritis. Serologically related to Gan Gan, Iltis JP and Daniels SB (1977) Infect Immun Mapputta, and Trubanaman viruses. 16 , 701 Isolated from mosquitoes in the Sepik Kunze LS et al (1996) Avian Dis 40 , 306 District of New Guinea. Not reported to cause disease in humans. marble spleen disease virus Synonym for pheasant adenovirus. MA protein The matrix protein of HIV, which forms part of the preintegration Marboran Trade name for methisazone. Marburger Affenkrankheit virus 298

Marburger Affenkrankheit virus Table M2. Strains of Lake Victoria marburg Synonym for Marburg virus . virus

Marburgvirus (MBGV) A genus in the Marburg Ang 1379v (Angola 2005) Marburg Ravn (Kenya 1987) family Filoviridae , containing a sin- Marburg Musoke (Kenya 1980) gle species, Lake Victoria marburgvirus . Marburg Ozolin (Zimbabwe 1975) Causes a severe and often fatal dis- Marburg Popp (West Germany 1967) ease in humans. Onset sudden with Marburg Ratayczak (West Germany 1967) fever, head and limb pains, brady- Marburg Voege (West Germany 1967) cardia, diarrhea, vomiting, and confused aggressive mental state. Cardiac and renal failure with hemorrhages develop. First reported in 1967 when that the virus had not previously been 31 cases, 7 fatal, occurred in Germany detected in this part of Africa. Then and the former Yugoslavia, all traced in October 2004 the worst outbreak of to contact with tissues from a batch of Marburg virus infection occurred in African green monkeys, Cercopithecus the Uige province of northern Angola, aethiops, trapped in Uganda. Five sec- with 227 infections more than 90% of ondary cases occurred in hospital which were fatal. ( Table M2) workers due to contact with blood Marburg virus causes a uniformly from patients, and one case in which fatal infection in guinea pigs and mon- the virus was sexually transmitted 83 keys, and can be propagated in a variety days after the initial illness. The virus of cell cultures such as Vero, BHK2l, and appears to persist in the body for 2–9 HeLa. Diagnosis is by antigen capture months. A second outbreak occurred in ELISA or polymerase chain reaction of 1975 involving one primary case, a man acute sera, IgM or IgG ELISA of paired who hitch-hiked through Zimbabwe, (acute and convalescent) sera, or by and two women who nursed him. inoculation of Vero cells which develop Two further cases occurred in 1980 in eosinophilic inclusions and antigen western Kenya; the index case died demonstrable by immunofluorescence. and the physician, who was second- Virus is ether-sensitive and inactivated arily infected, survived. Then in 1987 in 30 min at 56°C. It has an outer enve- another fatal case occurred in western lope and is formed by budding through Kenya. Both the 1980 and 1987 cases the cell membrane. There are two had visited caves on Mount Elgon structural forms: filamentous and cir- 10–12 days before their fatal illness, cular. Both have an overall diameter of but intensive ecological studies in 70–100 nm and contain an internal helix the region have failed to identify the 40 nm in diameter. Some filaments may virus reservoir. In 1998 fatal cases of be several microns long, but the unit Marburg virus infection began to occur length associated with peak infectivity is in persons who had visited an illegal 790 nm. The virus has a negative-strand goldmine in Durba, in the Democratic RNA genome, 19.1 kb in length, encoding seven proteins, in the order 3 -NP, Republic of the Congo. Over 100 cases were described. Viruses isolated from VP35, VP40, GP, VP30, VP24, and L-5 . cases in Durba show wide sequence Synonyms: green monkey virus; variation, at least as great as all other Marburger Affenkrankheit virus; Marburg viruses isolated since 1967, vervet monkey disease virus. suggesting that genetic variants of Feldmann H and Klenk H-D (2005) In Marburg virus had been introduced at Topley & Wilson’s Microbiology and Microbial least seven times into the population Infections , vol. 2, Tenth edition, edited by in the Durba area. Although the mine BWJ Mahy and V ter Meulen. London: is clearly linked to the source of the Hodder Arnold, p. 1085 infection, no reservoir species has so Geisbert TW et al (2007) J Infect Dis 196 , S372 far been definitely identified. However, Towner JS et al (2007) PloS ONE 2 , e764 Marburg virus has been detected in a common African fruit bat (Rousettus Marchal bodies Inclusion bodies found aegypticus) in Gabon, despite the fact in cells infected with Ectromelia virus. 299 marmoset cell line (B95-8)

Marco virus (MCOV) An unassigned ver- Marin county virus A serotype of tebrate rhabdovirus. Isolated from the Human astrovirus in the genus Astro- lizard, Ameiva ameiva , in Para, Brazil. virus . Isolated from a gastroenteri- Pathogenic for newborn mice. Not tis outbreak in a Californian nursing reported to cause disease in humans. home, USA. Now classified as human Replicates well with CPE in Vero cells at astrovirus 5. 30°C. Not isolated from arthropods but Herrmann JE et al (1987) Lancet 2 , 743 considered to be arthropod-transmitted as it will replicate in experimentally Marituba virus (MTBV) infected mosquitoes. A species in the genus Orthobunyavirus , belong- Monath TP et al (1979) Arch Virol 60 , 1 ing to the group C serogroup. Isolated from sentinel Cebus monkey and Marcy virus Obtained from a case of mouse in Para, Brazil. Also found in gastroenteritis in an outbreak in New Culex sp. Has been associated with a York State, USA. Could be passed in febrile illness in humans. Antibodies humans causing anorexia, nausea, vomi- are frequent in arboreal opossums of ting, and diarrhea. This virus is now lost, Caluromys sp and Marmosa sp but not and hence can no longer be identified. so common in forest floor-dwelling Gordon I et al (1947) J Exp Med 86 , 409 rodents and marsupials.

Mardivirus A genus in the subfamily marker rescue See reactivation . Alphaherpesvirinae containing viruses found only in birds. Form a distinct genetic lineage and cross-react anti- marker rescue mapping A technique genically. They are the only mem- used for mapping the site of tempera- bers of the subfamily associated with ture-sensitive mutations on the genome, malignancy. The type species is Gallid used especially with poxviruses. Cells herpesvirus 2 (Marek’s disease virus infected with mutant virus are rescued type 1). Other species are Gallid her- by transfection with defined DNA mol- pes-virus 3 , representing non-oncogenic ecules from wild-type virus. When res- strains of Marek’s disease virus, and cue occurs by recombination with the Meleagrid herpesvirus 1 (turkey herpes- wild-type fragment, this maps the site virus), which has been used as a vac- of the temperature-sensitive lesion. cine against Gallid herpesvirus 2 . marmodid herpesvirus 1 (MarHV-1) mare abortion virus Synonym for Equid A tentative species in the genus Rhadino- herpesvirus 1 . virus, isolated from woodchuck hepatocytes cultured in vitro . The genome is Marek’s disease virus type 1 Synonym 160 kb in length. Replicates in a variety for Gallid herpesvirus 2 . of monkey, feline, and hamster cells, and also in WCH-17, a woodchuck Marek’s disease virus type 2 Synonym hepatoma cell line. Antibodies to the Gallid herpesvirus 3 for , a species in the virus have been found in woodchuck genus ‘ Marek’s disease-like viruses ’ sera. containing non-oncogenic strains. Synonyms : woodchuck herpesvirus; Marek’s disease virus type 3 (MDV3) herpesvirus marmota. See Meleagrid herpesvirus 1 . Gilles NG and Ogstron CW (1991) Virology 180 , 434 Maribavir (1263W94) A novel benzimi- Schechter EM et al (1988) J Gen Virol 69 , dazole compound that is active against 1591 human cytomegalovirus infection. Wang LH et al (2003) Antimicrob Agents marmoset cell line (B95-8) A cell line Chemother 47 , 1334 derived from marmoset peripheral blood lymphocytes which grows in Maridi virus Synonym for Ebola virus suspension and has frequently been Sudan. used for isolation of measles virus from marmoset cell line (B95-8) 300

clinical specimens. The cells have the spikes. The genome RNA is 8 kb in surface measles receptor SLAM (s ign- length with LTRs of 350 nt (U3-240, aling l ymphocyte a ctivation m olecule) R-15, U5-95). There are no genome- and strains isolated in them retain associated oncogenes. Transforms rhe- pathogenicity for monkeys. B95-8 cells sus foreskin cells in culture but injec- are transformed by EBV, and provide tion into monkeys has not resulted in a source of EBV to establish continu- any mammary tumors. Strains have ous lymphocyte cell lines from human been isolated from placental tissues donors. An adherent cell subline, B95a, of normal rhesus monkeys and from has been established. HeLa cell lines. Approximately 20% of the viral genetic sequences are present marmoset cytomegalovirus Synonym for in rhesus tissue as endogenous pro- callitrichine herpesvirus 2. virus. Similar sequences are present in other Old World monkeys but not in marmoset herpesvirus Synonym for the cell DNA of New World monkeys, Saimiriine herpesvirus 1 . apes, or humans. Differs from other retroviruses by assembling Gag polyproteins into procapsids in the marmoset lymphocryptovirus Synonym cytoplasm. for Callitrichine herpesvirus 3 . Sakalian M and Hunter E (1999) J Virol 73 , 8073 marmosetpox virus (MPV) An unassigned virus in the family Poxviridae . Masou salmon reovirus (MSRV) An Virion morphology like Yatapox virus. isolate of Aquareovirus A in the genus Aquareovirus . Isolated from land-locked Marrakai virus (MARV) A serotype of salmon, Oncorhynchus masou . See also Palyam virus in the genus Orbivirus . landlocked salmon reovirus . Isolated from the mosquitoes Culicoides Hsu YL et al (1989) Fish Pathol 24 , 37 schultzei and C perigrinus in Northern Territory, Australia. mast cell A connective tissue cell with numerous large basophilic meta- Marshall–Regnery myxoma virus A chromatic granules in the cytoplasm. strain of Myxoma virus in the genus Leporipoxvirus . Isolated from the Mastadenovirus A genus in the family California brush rabbit. Adenoviridae comprised of the species Regnery DC and Marshall ID (1971) Am J isolated from mammals. The type spe- Epidemiol 94 , 508 cies is Human adenovirus C . Twenty species have been recognized, which marsupial poxvirus See quokkapox all share a common antigen. The 12 virus . vertex capsomeres each have a single filament which varies in length in dif- MARU 10962 virus (GAMV) A mosquito ferent species. The double-stranded DNA genome is 36 kb in length, mol. isolate of Gamboa virus in the genus 6 Orthobunyavirus . wt. 20–25 1 0 . G C content 48–61% in different viruses. Many hemagglutinate. Lack of cross-neutralization com- masern virus Synonym for Measles virus . bined with a calculated phylogenetic distance of more than 10% separate Mason–Pfizer monkey virus (MPMV) two serotypes into different species. A species in the genus Betaretrovirus . Isolated from a spontaneous mam- mastocytoma A tumor of neoplastic mast mary carcinoma in an 8-year-old rhe- cells. sus monkey. Can be propagated in human and nonhuman primate cell Matariya virus An unclassified arbo- cultures. Morphologically type D, virus. The first member of the Matariya similar to mouse mammary tumor serogroup. Isolated from birds of Sylvia virus but has no prominent surface sp in Egypt. Probably also present in 301 Mayaro virus (MAYV)

Europe since the birds were viremic on Antibodies found in bats. Not reported arrival in Egypt. Not reported to cause to cause disease in humans. disease in the wild. maturation The final process in assembly maternal antibodies In many viral infec- of the mature progeny virion during tions the presence of antibodies in the replication. It may occur inside the cell female before conception protects both (e.g. picornaviruses, reoviruses, papova- her and her fetus against infection viruses, adenoviruses, herpesviruses, and disease. In the presence of certain and poxviruses), in which case cell vaccine-acquired antibodies such as lysis is needed for egress. Alternatively, measles, mumps, rubella, and varicella, maturation may be linked with egress silent reinfection of the mother may from the cell as seen with most envel- occur, but usually the fetus is protected oped viruses (e.g. negative-strand from infection or disease. viruses, togaviruses, and retroviruses). maternal rubella Infection of a pregnant Maus–Elberfeld virus A strain of woman with rubella virus may have Encephalomyocarditis virus in the genus serious consequences, including intra- Cardiovirus . uterine death or the birth of a malformed Synonym : Mouse Elberfield virus. infant. The greatest risk to the fetus is during the period of maternal viremia, Maxam and Gilbert method An early and the consequences are largely deter- method of sequencing DNA using mined by the gestational age at which chemical base-specific modification maternal infection occurs. In about 20% and cleavage. of cases maternal rubella may result in Maxam A and Gilbert W (1980) Methods spontaneous abortion. The advent of Enzymol 65 i, 497 universal vaccine given together with measles and mumps vaccines at 12–15 Mayaro virus (MAYV) A species in the months and again at 11–12 years of genus Alphavirus . First isolated from age, as recommended by the CDC, has febrile humans in Trinidad in 1954, resulted in a dramatic decline in cases and then from febrile humans and of congenital rubella syndrome . mosquitoes in Brazil. There is a sudden onset of fever, headache, epigastric mathematical modeling A technique pain, myalgia, arthralgia, rash, chills, used in experimental epidemiology to nausea, photophobia, and vertigo. The study, e.g., the effects of an intervention rash begins on the fifth day of the ill- such as vaccination on the progress of ness in children, less frequently in an epidemic within a community. See older persons. Recovery occurs within also Monte Carlo model . 10 days, except for some persistence of arthralgia. The disease is widely Hethcote HW et al (1991) Math Biosci 106, 203 distributed throughout Central and matrix protein A term used for several South America. Associated with an different types of protein. In ortho- and epidemic febrile illness in humans in paramyxoviruses and rhabdoviruses it Uruma colony, Bolivia. Antibodies are refers to the protein between the viral found in humans and monkeys, and membrane and the nucleocapsid. in rural communities in the Amazon up to 60% of the population have anti- Matruh virus (MTRV) A serotype of Tete bodies to MAYV. The principal vec- virus in the genus Orthobunyaviru s. tor is Haemagogus janthinomys , and the Isolated from birds in Egypt and Italy. main vertebrate hosts are nonhuman Not reported to cause disease in the primates. Birds can act as secondary wild. hosts. There is no vaccine, and prevention is by personal protection against Matucare virus (MATV) A tentative spe- mosquito vectors. cies in the genus Orbivirus , isolated Casals J and Whitman L (1957) Am J Trop from the tick, Ornithodoros boliviensis , Med Hyg 6 , 1004 in the San Joaquin area, Beni, Brazil. Tesh RB et al (1999) Clin Infect Dis 28 , 67 MB III (de Bruyn-Gey) cells 302

MB III (de Bruyn-Gey) cells (CCL 32) adult steer. Since 1982 the line has been Initiated as one of three strains from a free of contamination by bovine viral mouse lymphosarcoma. diarrhea virus (1 or 2).

Mboke virus (MBOV) A mosquito iso- MDCK (NBL-2) cells (CCL 34) A hetero- late of Bunyamwera virus in the genus ploid cell line derived from the kid- Orthobunyavirus . Isolated from Aedes ney of an apparently normal cocker (Finlaya ) ingrami . spaniel. Institut Pasteur, Dakar (1972) Annual Report ME virus Synonym for Encephalo- M cells Specialized cells found in the fol- myocardi tis virus . licle-associated epithelium of Peyer’s patches in the digestive tract or the Meaban virus (MEAV) A species in nasopharynx-associated lymphoid tis- the genus Flavivirus belonging to the sue. M cells are involved in the uptake Seabird tick-borne virus group. and transcytosis of macromolecules and microbes such as bacteria and Measles (Edmonston) virus (MeV) The viruses. Following transcytosis, anti- type species in the genus Morbillivirus . gens are released to cells of the immune A natural infection of humans caus- system in lymphoid aggregates beneath ing measles, an acute febrile illness of the epithelium where antigen process- children, which is the seventh leading ing and presentation and stimulation cause of childhood mortality world- of specific B and T lymphocytes occurs. wide. Onset with prodromal symptoms Hathaway LJ and Kraehenbuhl JP (2000) of cough, coryza, and conjunctivitis Cell Mol Life Sci 57 , 323 occurs about 10 days after infection. Prodromal stage 4–5 days, followed MC virus Synonym for Montgomery by mounting fever, the appearance of County virus. Koplik’s spots on the buccal mucosa and rash on head and neck spreading to MC2 virus Strain of Junín virus in the the trunk and limbs. Recovery usually genus Arenavirus . rapid but the disease can be fatal, especially in poorly nourished children. The rash is dependent on the presence of a MCF virus Synonym for mink cell focus specific immune response and is absent inducing virus. from certain immunodeficient patients. The patient is most infectious in the McDonough feline sarcoma virus prodromal period and transmission (SMFeSV) A strain of feline sarcoma is by airborne droplets. Respiratory virus in the genus Gammaretrovirus iso- complications and otitis media due lated by Susan McDonough. Carries to secondary bacterial infection are the v-fms oncogene. Transforms a vari- common. Encephalitis occurs rarely ety of cells, including 3T3, MDCK, but is a serious complication with and feline embryo fibroblasts in vitro . high mortality and incidence of Activating mutations of the human cel- sequelae. Subacute sclerosing pan- lular homolog, FMS , have been found encephalitis (SSPE), a rare progres- in 17% of patients with primary acute sive degenerative disease of the CNS, myeloblastic leukemia. is associated with chronic infection Besmer P et al (1986) J Virol 60 , 194 with onset of symptoms at age 5–11 Donner L et al (1982) J Virol 41 , 489 years. Virion ether-sensitive, roughly McDonough SK et al (1971) Cancer Res 31 , 953 spherical, 150 nm in diameter, buoyant density in CsCl about 1.27 g/ml, MCP M embrane c ofactor p rotein, a recep- and contains a helical nucleocapsid of tor present on many cell types. about 17 nm 1100 nm. The negative- stranded RNA genome (16 kb in MDBK (NBL-1) cells (CCL 22) Derived length, mol. wt. about 4.5 1 0 6 ) has from the kidney of an apparently normal a 3 leader sequence of 55 nt, and 303 Melaka virus

encodes eight known proteins in the Synonyms : masern virus; morbilli virus; order 3 -N-PCV-M-F-H, L-5 . Proteins rougeole virus; rubeola virus. P, C, and V are encoded in overlapping Forcic D et al (2004) Virus Res 99 , 51 reading frames. Measles virus can be Hsu EL et al (2001) Virology 279 , 9 grouped into a large number of genetic Liu X et al (2006) Virus Res 122 , 78 types based upon the sequences of the Manchester M et al (2000) Virology 274 , 5 NP or the H gene. This has enabled Tatsuo H et al (2000) Nature 406 , 893 molecular epidemiological studies to WHO (1998) WHO Wkly Epidemiol Rec 73 , 265 trace the movements of distinct genetic types globally, and is the basis for glo- mechanical transmission The trans- bal vaccine elimination programs. mission of a virus from one host to the Measles virus is related antigenically other by an arthropod in which the to canine distemper and rinderpest virus does not replicate. It is thus not viruses and antibodies to them can be a very efficient form of transmission demonstrated in patients with mea- when compared to true arthropod- sles. Virus hemagglutinates primate borne viruses which multiply in the erythrocytes only and lacks neurami- insect during transmission. nidase. Isolation from clinical specimens is easiest in B95-8 or the adherent median infective dose (ID50 ) The dose subline B-95a cells, or primary human of virus which, on average, will infect or monkey kidney cell cultures, but 50% of the individuals to whom it is the virus can be adapted to grow in a administered. These may be animals, number of cell lines (e.g. Vero) or eggs. humans, or eggs; with eggs the term The receptor for measles virus on B95a EID50 (egg infective dose) is often used. cells has been identified as a membrane glycoprotein known as SLAM median tissue culture infective dose (s ignaling l ymphocyte a ctivation m ol- (TCID50 ) The dose of virus which, on ecule), also called CDw150. Previous average, will infect 50% of susceptible work had established that another cell tissue culture cells. surface protein, CD46, is a receptor for the Edmonston strain of measles virus. Medical Lake macaque herpesvirus Following infection, multinuclear giant Synonym for Cercopithecine herpesvirus 9 . cells are formed, followed by gradual Blakely GA et al (1973) J Infect Dis 127 , 617 cell destruction. Changes are most easily seen in stained preparations. Monkeys are susceptible and develop a mel1-mel3 Serotype designation given disease similar to that seen in humans, to the avian adenovirus isolated from but many monkeys have antibodies the turkey, Meleagris gallopavo . Genus and are immune. Virus can be adapted Aviadenovirus , family Adenoviridae . to replicate in mice, ferrets, and hamsters. An effective attenuated vaccine Melaka virus A probable member of the produced in chick fibroblast cells is genus Orthoreovirus. Isolated from a recommended for use at about 39-year-old male patient in Melaka, 12 months of age, when maternal Malaysia who had a high fever and immunity has waned. Use of this vac- respiratory disease. Two family mem- cine in developed countries has almost bers developed similar symptoms and eliminated subacute sclerosing pan- later seroconverted to the same virus. encephalitis. In developing countries Genetic sequence analysis of the virus measles is usually contracted early, isolated in MDCK cells showed that it and the disease is more severe with was related to Pulau virus, an ortho- considerable mortality (estimated 1.5 reovirus isolated from a fruit bat on million deaths annually). Measles can Tioman island, Malaysia. It is possible be eliminated from populations by vac- that the patient was infected through cination of all susceptible individuals, contact with a bat which flew into his and is now a target for global eradi- house. cation, beginning with the Western Chua KB et al (2007) Proc Soc Nat Acad Sci hemisphere. 104 , 11424 Melao virus (MELV) 304

Melao virus (MELV) A mosquito isolate protein (e.g. rhabdovirus G protein or of California encephalitis virus in the influenza A hemagglutinin) or not (e.g. genus Orthobunyavirus . Isolated from paramyxovirus F protein). mosquitoes in Trinidad and Belem, Kielian M (1995) Adv Virus Res 45 , 113 Brazil. Not reported to cause disease in humans. membrane proteins A class of proteins that are synthesized on membrane- Meleagrid herpesvirus 1 (MeHV-1) A bound polyribosomes with an amino species in the genus Mardivirus in the terminal ‘ signal ’ peptide that directs subfamily Gammaherpesvirinae . Isolated the protein into the lumen of the endo- from a kidney cell culture of normal plasmic reticulum (ER). When the turkeys. Not pathogenic for turkeys. protein becomes incorporated into Causes viremia in chickens and protec- the cell or virion membrane the sig- tive immunity against Marek’s disease nal sequence is cleaved off by a sig- virus 1 (GaHV2). Used as a vaccine nal peptidase. Oligosaccharides are against Marek’s disease. The com- attached to glycoproteins in the lumen plete DNA sequence of MeHV-1 was of the ER and processed in the Golgi obtained recently and compared to that complex en route to the cell surface. of GaHV-2. Synonym: turkey herpesvirus 1; Marek’s disease virus type 3. membranoproliferative glomerulonephritis (MPGN) type 1 A condi- Alfonso CL et al (2001) J Virol 75 , 971 tion found in patients with hepatitis C Lee LF et al (1972) Avian Dis 16 , 799 infection, resulting from immune complex deposition in the glomerular capil- Melksham virus A strain of Norwalk virus laries. It is associated with the presence in the genus Norovirus . Isolated during of serum rheumatoid factor, comple- a gastroenteritis outbreak at a school in ment activation, and cryoglobulinemia. Melksham, Wiltshire, UK in 1989.The RNA sequence encoding the capsid memory cells Cells of the immune system protein is 98% similar to that of Snow which ‘ remember ’ their first encounter Mountain virus. with an antigen and facilitate a more Green SM et al (1995) Virus Res 37 , 271 rapid secondary response when the antigen is encountered on a second occasion. Lymphocytes circulate con- membrane cofactor protein Some viruses tinuously through lymphoid and non- require more than one cell membrane lymphoid tissues in search of antigens protein for attachment and entry into to provide specific protection against the host cell. For example, HIV requires pathogens. When naive T cells encoun- a CD4 molecule and also a membrane ter an antigen they are activated, dif- cofactor protein, a chemokine recep- ferentiate into effector or memory cells, tor, most frequently CCR5 or CXCR4 and proliferate. to enter cells. CCR5 is expressed on the surface of macrophages, while CXCR4 Moser B and Loetscher P (2001) Nat Immunol is expressed on T lymphocytes. 2 , 123 Doms RW and Peiper SC (1997) Virology 235 , 179 Menangle virus (MenV) A tentative Berger EA et al (1999) Annu Rev Immunol 17 , species in the genus Rubulavirus , iso- 657 lated in 1997 from stillborn piglets at a commercial piggery in Menangle, membrane fusion Fusion between viral New South Wales, Australia. Caused and cellular membranes is required for abortions and increased stillbirths, most enveloped viruses, after attach- with associated deformities and ment to the cell receptor, in order to severe degeneration of the brain and gain entry into the cell. Fusion is medi- spinal cord. In utero infection of pig- ated by a virus envelope protein, which lets is associated with severe skeletal may also be the receptor-binding and neurological malformations. Two 305 methisazone

humans in contact with the pigs devel- infectious and serves as mRNA upon oped influenza-like illness, and neu- entry into a susceptible cell. tralizing antibodies were detected in several species of fruit bats, Pteropus Metapneumovirus A genus in the family sp, in the vicinity. The virus multiplies Paramyxoviridae containing two spe- in Vero cells, and phylogenetic analy- cies, Avian metapneumovirus (Turkey sis of genome and expressed mRNA rhinotracheitis virus) and Human meta- confirmed a close relationship to other pneumovirus . Distinct from the genus members of the genus Rubulavirus , Pneumovirus because of differences with the exception of the deduced HN in genome organization, e.g. absence protein, which appeared to bear little of NS1 or NS2 genes, and reversal of relationship to attachment proteins of the placements of SH-G versus F-M2 other paramyxoviruses. So far, no other in the gene order. Although avian outbreak of pig disease caused by this metapneumovirus was first isolated in virus has been recorded. the 1970s, human metapneumovirus Bowden TR et al (2001) Virology 283 , 358 was unknown until 2001, since when Philbey AW et al (1998) Emerg Infect Dis 4 , it has been found to be an important 269 respiratory pathogen of infants in Philbey AW et al (2007) Aust Vet J 85 , 134 many countries from all continents. Seroprevalence studies show that the mengo virus A strain of virus has been circulating in the human Encephalomyocarditis virus in the genus population for more than 50 years and Cardiovirus . it infects virtually all children by the age of 5–10 years. In young children meractinomycin Synonym for actinomy- it causes bronchiolitis, pneumonitis, cin D. otitis media, and acute exacerbation of asthma. In one Australian study of Merkel cell polyomavirus ( MCPyV) 10,000 respiratory samples the aver- A novel polyomavirus with a DNA age annual incidence of human meta- genome of 5387 base-pairs, which was pneumovirus was 7.1%. In mice, human found integrated into the tumor cell metapneumovirus induced more sev- genome of 8 of 10 Merkel cell carcino- ere disease than respiratory syncytial mas studied. Merkel cell carcinomas are virus. an aggressive human skin cancer that Deffrasnes C et al (2007) Semin Respir Crit typically affects elderly and immuno- Care Med 28 , 213 compromised individuals. The possi- Hamelin ME and Boivin G (2005) Pediatr ble role of the virus in pathogenesis is Infect Dis J 24 , S203 unknown. Huck B et al (2007) Respir Res 8 , 6 Njenga MK et al (2003) Virus Res 91 , 163 Feng H et al (2008) Science 319 , 1096 Sloots TP et al (2006) Emerg Infect Dis 12 , 1263 Mermet virus (MERV) A strain of Manzanilla virus in the genus Orthobunyavirus , belonging to the Metaviridae A family of retrotransposons Simbu serogroup. Isolated from sev- that have been found in all studied lin- eral different species of birds and mos- eages of eukaryotes. Related to viruses quitoes in Illinois and Texas, USA. Not of the family Retroviridae by reverse reported to cause disease in humans. transcription and a viral core structure made up of Gag-like proteins. The mesogenic strains Virus strains of average family has three genera, Metavirus , virulence; particularly used to describe type species Saccharomyces cerevisiae strains of Newcastle disease virus. Ty3 virus ; Errantivirus, type species Drosophila malanogaster Gypsy virus; and messenger RNA (mRNA) A single- Semotivirus , type species Ascaris lumbri- stranded RNA molecule which carries coides Tas virus. the genetic information to the ribosomes for protein synthesis. The genome methisazone N -Methyl-isatin- β -thiosemi- RNA of most positive-strand viruses is carbazone. A synthetic antiviral agent. methisazone 306

Mode of action unclear but apparently for maintaining mRNA stability in the dependent upon presence of benzene cytoplasm and in the process of initia- ring and a side chain containing sulfur. tion of protein synthesis. Active in cell culture systems against poxviruses and adenoviruses. Inhibits N-methyl-isatin-β-thiosemicarbazone translation of late viral mRNA. Was See methisazone . used in the treatment of complicated vaccinia and in the short-term preven- 1-methyl spiro (adamantane-2,3-pyrro- tion of smallpox. lidine) maleate A derivative of aman- Fox MP et al (1977) Ann NY Acad Sci 284 , 533 tadine. A double-blind trial of this Pearson GD and Zimmerman EF (1969) drug against a placebo on volunteers Virology 38 , 641 infected with influenza virus A/Hong Kong/68 showed that the group receiv- 8-methoxypsoralen (Methoxsalen) A ing the drug, although not entirely pro- naturally occurring furocoumarin tected, had fewer clinical symptoms, which has photosensitizing proper- antibody rises, or virus secretions than ties in the skin of guinea pigs and those receiving the placebo. The drug humans. Albino guinea pigs with cuta- was apparently nontoxic, and routine neous infection by human herpesvirus liver function tests at the end of each showed significant favorable response trial were uniformly normal. to treatment with this drug and long- Beare AS et al (1972) Lancet 1 , 1039 wave UV light. Treatment was effective even after virus multiplication had methyl transferase Enzyme activity begun and lesions had appeared. found in some virus particles which is Oill PA et al (1978) J Infect Dis 137 , 715 involved in ‘ capping ’ of viral mRNAs.

α-methyl-1-adamantane-methylamine MEV See mink enteritis virus . hydrochloride See rimantadine hydro- Mexico virus (MXV) A strain of Norwalk chloride . virus in the genus Norovirus . 2-C-methyl -adenosine A selective inhib- MF59 adjuvant An oil–water emulsion itor of hepatitis C virus RNA polymer- now used in a split virion influenza ase. Not yet developed for clinical use. vaccine that increases immunogenicity Olsen DB et al (2004) Antimicrob Agents as compared with alum adjuvant. Chemother 48 , 3944 MG virus A strain of BK polyomavirus . methylation The addition of methyl Isolated in primary human fetal fibro- groups. Methylation of DNA at specific blasts from the urine of a renal allograft nucleotides within the target site of a patient in South Africa. restriction endonuclease (termed ‘ mod- Wright PJ et al (1976) J Virol 17 , 762 ification ’ ) can protect the DNA against cleavage by that enzyme and is the MH 2 virus Synonym for Avian carci- means by which bacteria protect their noma virus Mill Hill virus 2 . own DNA against the restriction endonucleases they encode. See S-adenosyl- MHC restriction See major histocompat- L -methionine (AdoMet, SAM) . ibility complex . methylene blue A photoreactive dye. MH1C1 cells (CCL 144) A clonal strain The oxidized form is blue, the reduced of epithelial cells derived from a transform colorless. See photodynamic plantable Morris hepatoma in a rat. inactivation . Mice minute virus (MMV) See Minute 7-methylguanylate(m7 G ) A modified virus of mice . nucleotide which is found at the 5 terminus of all cellular and most viral mice pneumotropic virus Synonym for RNAs, known as the 5 cap. Important Murine polyomavirus . 307 Mill Door virus (MDRV)

؊ .MiCl1 (S ؉ L ) (CCL 64.1) Developed exact homology with a target mRNA from the clonally derived substrain of Recently it has been shown that several the original parent mink lung line (CCL human herpesviruses express virally 64). It contains the murine sarcoma virus encoded miRNAs in infected cells, genome rescuable by super-infection and it is possible that these play an with compatible helper viruses. important role in replication, perhaps by silencing specific cellular RNAs. At microarrays Amplification/hybridization least 17 distinct miRNAs have been methods used in virus diagnosis that found in cells infected with human are miniaturized so that it is possible herpesvirus 3 (EBV), and they are also to assay, in a single run, specimens found in cells infected with herpes sim- against thousands of probes. The plex virus, where a miRNA encoded by probes consist of microspots of DNA or the latency-associated transcript (LAT) oligonucleotides bound to the surface gene protects infected neuronal cells of slides or microchips. from undergoing apoptosis. Cai X et al (2006) PloS Pathog 2 , e23 microbiological risk assessment (MRA) Gupta A et al (2006) Nature 442 , 82 See biosafety . microsomes Small particles, 16–150 nm in microcultures Growth of tissue culture diameter, obtained on cell fractionation. cells in 24- to 48- or 96-well plates They are fragments of endoplasmic with sealable lids. Used for growth of reticulum. On treatment with sodium viruses for neutralization tests, tissue deoxycholate they are disrupted into culture enzyme immunoassay, mono- two fractions. The first contains most clonal antibody testing, and other of the protein, phospholipid, pigment, screening assays. and enzymes. The second is a particulate fraction sedimentable at 100,000 g , microRNAs Single-stranded RNA mol- and containing nearly all the RNA of ecules 21–23 nt in length which regu- the cytoplasm; these are the ribosomes. late gene expression. They were first described in 1993, but the term micro- Microtus pennsylvanicus herpesvirus RNA was only introduced in 2001, and Synonym for murid herpesvirus 5. their importance has only been real- ized since 2003. Several hundred miR- Middelburg virus (MIDV) A species in NAs have been sequenced from a wide the genus Alphavirus. Isolated in South range of organisms, and show consid- Africa from Aedes sp mosquitoes, and erable homology, implying that they may be the cause of an epizootic in represent an old and important regu- sheep. Found in South Africa, Senegal, latory mechanism. In eukaryotic cells Central African Republic, Cameroon, miRNAs are transcribed from DNA as Kenya, and probably Mozambique and primary transcripts which are capped Angola. Not reported to cause disease and polyadenylated, then processed in humans. in the nucleus by the dsRNA-specific Jupp PG and Kemp A (1998) J Am Mosq ribonuclease Drosha to short 70-nt Control Assoc 14 , 40 stem loop structures, known as pre- Kokernot RH et al (1957) S Afr J Med Sci 22 , miRNAs. These are further processed 145 in the cytoplasm by the endonuclease milk factor Synonym for Bittner virus. Dicer, which cleaves the pre-miRNAs See Mouse mammary tumor virus . approximately 19 bp from the Drosha cut site. Dicer also initiates the forma- milker’s nodule virus Synonym for tion of the RNA-induced silencing Pseudocowpox virus . complex known as RISC. This complex is responsible for gene silencing and milk-pox virus Synonym for variola RNA interference. The function of miR- minor virus. See Variola virus . NAs is in gene regulation. Annealing of microRNA to mRNA inhibits trans- Mill Door virus (MDRV) A serotype of lation, and frequently the miRNA has Great Island virus in the genus Orbivirus . Mill Door virus (MDRV) 308

Isolated in 1979 from ticks (Ixodes uriae ) minimal essential medium (MEM) A from a seabird colony on the Isle of medium for the culture of vertebrate May, Scotland. Related serologically to cells. It differs from Eagle’s medium the Kemerovo virus group. mainly in its increased concentration of Spence RP et al (1985) Acta Virol 29 , 129 essential amino acids.

Mill Hill virus 2 See Avian carcinoma minireovirus Synonym for minirotavirus. virus Mill Hill virus 2 . minirotavirus A name sometimes used Mimivirus A free-standing genus con- for the 32 nm particles seen in feces. taining one species, a double-stranded Differ from other small, round virus- DNA virus of free living amoeba, like particles seen in feces in being Acanthamoeba polyphaga mimivirus , slightly larger. which has the largest viral genome Synonym : minireovirus. so far described, at 1.2 megabases. Middleton PJ and Szymanski MT (1977) Am Originally isolated in 1992 from a J Dis Child 131 , 733 water cooling tower in Bradford, Spratt HC et al (1978) J Pediatr 93 , 922 England during investigation of a suspected Legionnaire’s pneumonia out- mink calicivirus (MCV) A tentative spe- break, it was initially thought to be a cies in the genus Vesivirus . bacterium, and only in 2003 identified as a virus and shown to be cytolytic for mink cell focus-inducing virus (MCF the amoeba species. Virions are 400 nm MLV) A type C retrovirus which is in diameter. The name mimivirus associated with the development of was chosen for ‘ mimicking microbe. ’ lymphomas in mice. Replicates in cell Serological evidence suggests that the cultures of mouse or mink lung fibrob- virus has infected patients in Canada, lasts. Strains obtained from AKR mice England, and France. There is one produce cytopathic foci in mink cells report from France of a laboratory- but strains from BALB/c (Mo) mice acquired infection of a technician in cause foci in mink and mouse cells. The Marseilles linked to acute pneumonia, virus may have arisen by recombina- which showed that the mimivirus can tion between ecotropic and xenotropic cause clinical disease, but the public mouse viruses. Cause leukemia by health impact of the virus remains to stimulating proliferation of lymphoid be assessed. cells bearing the erythropoietin recep- La Scola B et al (2005) Emerg Infect Dis 11 , tor or related cytokine receptors such 449 as the interleukin-2 β chain receptor. Raoult D et al (2004) Science 306 , 1344 Synonym : MCF virus. Cell 32 mimotope A sequence of amino acids Cloyd MW (1983) , 217 Hartley JW et al (1977) Proc Natl Acad Sci 74 , which mimics the structure of an 789 epitope. Li JP and Baltimore D (1991) J Virol 65 , 2408 Nanua S and Yoshimura FK (2004) J Virol 78 , Minaçu virus A reovirus isolated from 12071 culicoides flies in Minaçu, Golás State, Brazil. Infected suckling mice, and rep- mink endogenous type C RNA virus A licated in Vero cells. The genome con- mammalian type C retrovirus. After sists of 10 segments of double-stranded about 100 generations in culture the RNA. mink cell line MV1LU produced sedi- Martins LC et al (2007) Int J Exp Pathol 88 , 63 mentable reverse transcriptase activity. This activity was the result of activa- Minatitlan virus (MNTV) A species tion of virus endogenous to the mink in the genus Orthobunyavirus . With cells. The DNA of normal mink cells Palestina virus forms the Minatitlan had extensive nucleotide sequence serogroup. Isolated from a sentinel homology with the viral nucleic acid, hamster in Veracruz, Mexico. Not demonstrating that the virus was an reported to cause disease in humans. endogenous virus of mink. 309 MM virus

Barbacid M et al (1978) J Virol 25 , 129 Astell CR et al (1996) Prog Nucleic Acid Res Sherr CJ et al (1978) J Virol 25 , 738 Mol Biol 55 , 245 Berns KI (1990) Microbiol Rev 54 , 316 mink enteric sapovirus A tentative spe- Choi EY et al (2005) J Virol 79 , 12375 cies in the genus Sapovirus . Kaufmann B et al (2007) J Virol 81 , 9851 Siegl G (1976) Virol Monogr 15 , 109 mink enteritis virus (MEV) A strain of Feline panleukopenia virus in the genus minute virus of mice (Cutter) (MVMc) A Parvovirus . strain of Minute virus of mice . Johnson RH (1967) J Small Anim Pract 8 , 319 minute virus of mice (immunosuppressive) (MVMi) A strain of Minute virus Minnal virus (MINV) A serotype of of mice . Umatilla virus in the genus Orbivirus . Isolated from Culex vishnui in Madras, Mirim virus (MIRV) A serotype India. Not known to cause disease in of Bertioga virus in the genus humans. Orthobunyavirus , belonging to the Guama serogroup. Isolated from sen- minus strand Synonym for negative tinel Cebus monkeys, and from mos- strand. quitoes in Para, Brazil. Not reported to cause disease in humans. minute virus of canines See Canine minute virus . missense mutants Mutants with an altered nucleotide sequence which Minute virus of mice (MVM) The results in the substitution of an incor- type species of the genus Parvovirus . rect amino acid in the polypeptide Isolated from a mouse adenovirus chain. preparation. A natural and probably silent infection in wild and laboratory Mitchell River virus (MRV) A sero- mice. Multiplies on injection into new- type of Warrego virus in the genus born mice, rats, and hamsters. In ham- Orbivirus. Isolated from Culicoides sp sters a disease similar to that caused in Queensland, Australia. Antibodies by latent rat virus is produced but in found in cattle, wallabies, and kanga- mice there is only retarded growth roos. Not reported to cause disease in and in rats a silent infection. Replicates the wild. in rat or mouse embryo cell cultures with CPE. Agglutinates guinea pig, mitogen An agent which stimulates qui- hamster, rat, and mouse erythrocytes escent cells to enter the cell cycle result- between 4°C and 37°C. The genome ing in DNA replication and growth of is negative-stranded DNA, 5084 bases the cells. in length, with hairpin structures at each end, 116 nt long at the 3 end and Miyagawanella Synonym for chlamy- 207 nt long at the 5 end. There are two dia. Named after Yoneji Miyagawa, mRNA promoters (map units 4 and 39) Japanese bacteriologist (1885–1959). and a single polyadenylation site at the 3 end. Multiplication occurs in the cell nucleus and requires the cell to go Mj virus A strain of human cytomegalo- through S phase in order to replicate virus isolated from a cell line derived the viral genome. For export from the from prostate tissue of a young boy. nucleus, the virus-encoded NS2 pro- Human embryo lung cells infected with tein interacts with the nuclear export Mj virus became persistently infected, protein Crm1. Recently the structure of and induced tumors when injected into MVM virions complexed with a mono- weanling athymic nude mice. clonal mouse antibody was solved by Geder L et al (1976) Science 192 , 1134 cryo-electron microscopy. Synonyms: MVM virus; minute virus of MM virus A strain of Encephalomyocarditis mice. virus . MM-2325 virus (BAKV) 310

MM-2325 virus (BAKV) A strain of Bakau sequenced. It is 10,600 nt in length and virus in the genus Orthobunyavirus . contains a single open reading frame from nucleotides 110 to 10,234 encod- MML virus A strain of Rio Bravo virus . ing 3374 amino acids. Isolated from bats of Myotis sp in Leyssen P et al (2002) Virology 293 , 125 Montana, USA. moesin M embrane o rganizing ex ternal MMR An abbreviation for combined s pike prote in , a putative receptor for measles–mumps–rubella vaccine. measles virus.

M-MSV-BALB/3T3 cells (CCL 163.2) A Moju virus (MOJUV) A strain of Guama Moloney murine sarcoma virus (M- virus in the genus Orthobunyavirus . MSV) transformed nonproducer cell Isolated from sentinel mice and forest line. rats, also mosquitoes in Para, Brazil. Not reported to cause disease in MMT 060562 cells (CCL 51) Derived humans. from a spontaneous mammary tumor which arose in a C57BL×Af F l hybrid Mojui dos Campos virus (MDCV) female mouse. A tentative species in the genus Orthobunyavirus , not assigned to any MMV virus A strain of BK virus. Isolated antigenic group. Isolated from an from a reticulum cell sarcoma of undetermined species of bat in Para, the brain of an 11-year-old boy with Brazil. Wiskott–Aldrich syndrome. Isolated by co-cultivation of brain tissue with Mokola virus (MOKV) A species in human fetal brain cells. the genus Lyssavirus . Antigenically Takemoto KK et al (1974) J Natl Cancer Inst related to, but distinguishable from, 53 , 1205 rabies virus. Isolated from a shrew of Crocidura sp and from children with Mobala virus (MOBV) A species in the CNS disease in Nigeria. Experimental genus Arenavirus belonging to the infection often fatal in shrews. Can be LCMV–LASV complex (Old World transmitted from shrews to mice by arena-viruses). Isolated in the Central biting. Pathogenic for dogs and mon- African Republic from rodents, Praomys keys given i.c. jacksonia . Kemp GE et al (1973) Am J Epidemiol 98 , 43 Bowen ME et al (2000) J Virol 74 , 6992 Tignor GH et al (1973) J Infect Dis 128 , 471 Gonzalez JP et al (1983) Intervirology 19 , 105 Peters CJ et al (1987) Curr Top Microbiol molecular mimicry The cross-reaction Immunol 134 , 5 of an antiviral immune response with host antigens. Believed to contribute to mock infection Inoculation of cells or an the induction of type 1 diabetes follow- organism with a solution not containing infection with group B coxsackie ing infectious virus. Used as a control viruses. in virus infection experiments to ascer- tain any possible effects of materials Hyoty H (2002) Ann Med 34 , 138 in the inoculum other than infectious particles. Molluscipoxvirus A genus in the family Poxviridae containing a sin- Modoc virus (MODV) A species in gle species, Molluscum contagiosum the genus Flavivirus belonging to virus. Virions are brick-shaped, the Modoc virus group. Found in 320 nm 250 nm 200 nm contain- California, Oregon, Montana, and ing double-stranded DNA, 200 kb in Colorado, USA. There is no known length; G C 60%. Produces localized arthropod vector. Isolated from the skin lesions containing enlarged cells deer mouse, Peromyscus maniculatus . with cytoplasmic inclusions. Probable Not known to cause disease in humans. species have been recognized in horses, The virus genome has been completely donkeys, and chimpanzees. 311 Monkeypox virus (MPXV)

Molluscum contagiosum virus A replication defective, acute trans- (MOCV) Type species of the genus forming virus which rapidly induces Molluscipoxvirus . An exclusively human sarcomas in mice and transforms cells infection, mostly of children, young in culture. Appears to be a recom- adults, or immunosuppressed patients. binant between the Moloney leukemia The DNA is 196–200 kb in length; G C virus and an endogenous mouse type is 60% with inverted terminal repeats. C virus. Carries the transduced v-mos There are 165 open reading frames, 104 gene that encodes a serine–threonine of which have homologs in other pox- protein kinase. A spontaneous dele- viruses such as vaccinia virus. Lesions tion mutant of MoMSV, known as R7, are confined to the skin. Incubation induces a distinct pathology with brain period 14–50 days. Pimples develop hemorrhage and angioendotheliomas. into nodules 2 mm in diameter which See Gammaretrovirus . become pearly white and may develop Lim KY et al (2000) J Neurovirol 6 , 106 an opening to reveal a white core. Moloney JB (1966) Natl Cancer Inst Monogr Lesions persist for months and may be 22 , 139 disfiguring when combined with bacterial infections. Transmission by direct Moloney sarcoma virus Synonym for and indirect contact can cause oppor- Moloney murine sarcoma virus . tunistic infection in immunocompromised (e.g. AIDS) patients. Virus does Moloney virus Synonym for Murine not replicate in eggs or cell cultures. leukemia virus . Not inhibited by vaccinia, cowpox or fowlpox antiserum and does not reac- monkey B virus Synonym for herpesvi- tivate heat-inactivated vaccinia virus. rus simiae; Cercopithecine herpesvirus 1 . Not inactivated by ether or chloroform but inactivated by exposure to pH monkey papillomavirus Synonym for 3. Does not hemagglutinate chick or simian papillomavirus. human erythrocytes. Monkeypox virus (MPXV) A species in Porter CD and Archard LC (1992) J Med Virol 38 , 1 the genus Orthopoxvirus. Has caused Senkevich TG et al (1996) Science 273 , 813 outbreaks of disease in captive mon- Senkevich TG et al (1997) Virology 233 , 19 keys, most commonly in cynomolgus and rhesus monkeys. The animals are not seriously ill except occasionally the molluscum-like pox virus (MOV) very young. Little is known about the An unassigned virus in the family infection in the wild, and the natural Poxviridae . Virions are brick-shaped, host is not established but it is prob- 320 nm 250 nm 200 nm. Genome ably a squirrel or other rodent. May be is double-stranded DNA, 188 kb in isolated from apparently normal mon- length; G C is 60%. Tentative spe- keys. Isolated from captive monkeys, cies in the group have been found in and from about 36 cases of human dis- horses, donkeys, and chimpanzees. ease in West and Central Africa, six of which were fatal. Mortality rate similar Moloney murine leukemia virus to that in variola virus infections and (MoMLV) A strain of Murine leukemia the disease is similar, but transmis- virus. Obtained from mouse sarcoma sion to human contacts is infrequent. S37. Produces lymphoid leukemia in Very similar to vaccinia but pocks on 3–4 months. Most active when injected the CAM may have hemorrhagic cent- into newborn mice. Also causes leuke- ers at 34.5°C and are not produced mia in rats. above 39°C. Inoculation of rabbit skin produces an indurated lesion with Moloney murine sarcoma virus (MoMSV) purple center. Adult mice susceptible. A species in the genus Gammaretrovirus , Replicates in various cell cultures but belonging to the mammalian virus CPE variable. In May and June 2003 group. Isolated from a BALB/c mouse the virus was introduced into the USA injected with Moloney leukemia virus. via a shipment of African rodents from Monkeypox virus (MPXV) 312

Ghana. They were initially housed in the same tick species in a swallow’s contact with prairie dogs (Cynomys nest in Texas, USA. Not reported to spp) who became infected with mon- cause disease in humans. keypox virus and were sold as pets. Initially 11 patients who had direct con- monolayers Single layers of tissue culture tact with the prairie dogs developed a cells growing on a solid surface (usu- febrile illness with skin eruptions, and ally glass or plastic) with a complete evidence of poxvirus infection was layer only one cell thick. obtained by immunohistochemistry or electron microscopy of skin lesion tis- Mononegavirales This was the first taxon sue. When animals from the original above the family level to be recog- importation were tested, three rodent nized by the ICTV. The order embraces species tested positive for monkeypox the families Bornaviridae , Filoviridae , virus: giant pouched rats, Cricetomys Paramyxoviridae , and Rhabdoviridae spp; rope squirrels, Funisciuris spp; and which are comprised of viruses with dormice, Graphiuris spp. As a result monopartite negative stranded RNA of this outbreak, a ban on importa- genomes. tion of wild rodents from Africa was Pringle CR (1997) Arch Virol 142 , 2321 introduced. Because the animals were housed together during transportation Monongahela virus A strain of Sin it was not possible to determine the Nombre virus in the genus Hantavirus . natural reservoir species as a result of Isolated from deer mice, Peromyscus this outbreak, so this is still unknown. maniculatus nubiterrae, in eastern USA Bernard SM and Anderson SA (2006) Emerg and Canada. Infect Dis 12 , 1827 Rhodes III LV et al (2000) Emerg Infect Dis 6 , et al Am J Trop Med Hyg Hutson CL (2007) 616 76 , 757 Jezek Z and Fenner F (1988) Human Monkeypox. Virology Monographs , vol. 17. monospot test A test for infectious mono- Basel: Karger nucleosis (caused by EBV infection) Reed KD et al (2004) N Engl J Med 350 , 342 which relies upon the presence from early in the course of infection of monocistronic Messenger RNAs which heterophile antibodies that agglutinate carry the information for a single pro- or lyse erythrocytes from sheep, goat, tein or polypeptide chain. See poly- horse, or bovines. See Paul–Bunnell– cistronic messenger RNA . Davidsohn test . monoclonal antibody An antibody prep- monovalent vaccine A vaccine contain- aration which contains only a single ing a single antigenic component or type of antibody molecule. Synthesized virus type. and secreted by clonal populations of hybrid cells (hybridoma) prepared by Montana myotis leukoencephalitis the fusion of individual B lymphocyte virus (MMLV) A species in the genus cells from an immunized animal (usu- Flavivirus , belonging to the Rio Bravo ally a mouse or a rat) with individual virus group. Isolated from a paralyzed cells from a lymphocytic tumor (e.g. little brown bat, Myotis lucifugus , in myeloma). western Montana, USA. There is no known arthropod vector. Not known to monocytes Large leukocytes (mono- cause disease in humans. nuclear phagocytes) circulating in the Charlier N et al (2002) J Gen Virol 83 , 1875; blood that will later migrate into tissue 1887 and differentiate into macrophages. Monte Carlo model A mathemati- Mono Lake virus A strain of Chenuda cal model used to study virus trans- virus in the genus Orbivirus . Isolated mission and epidemics and to identify from the tick Argas cooleyi, in California, the effects of possible interventions. USA. A similar virus was isolated from Abbey H (1952) Hum Biol 24 , 201 313 Mossman virus

Monte Dourado virus (MDOV) A sero- Morbillivirus (Latin: morbillus , diminu- type of Changuinola virus in the genus tive of morbus disease). A genus of Orbivirus . Isolated from the armadillo, the family Paramyxoviridae . Viral enve- Dasypus novemcinctus , in Para, Brazil. lope contains hemagglutinin but not neuraminidase. Nucleocapsid about Montgomery County virus A possi- 18 nm in diameter, with helical pitch of ble species in the genus Norovirus . 5–6 nm. All species contain a common Observed by electron microscopy in antigen, but are readily distinguishable. the feces in a family outbreak of gastro- Type species Measles virus . Other spe- enteritis in Montgomery County, cies are Canine distemper virus , Cetacean Maryland, USA. Antigenically related morbillivirus , Peste des petits ruminants to Norwalk virus. virus , and Rinderpest virus . Synonym : MC virus. Barrett T (2008) in BWJ Mahy and MHV van Dolin R (1978) J Am Vet Med Assoc 173 , 615 Regenmortel (eds) Encyclopedia of Virology , Thornhill TS et al (1977) J Infect Dis 135 , 20 Third edition, Oxford: Academic Press, vol. 4, p. 497 MOPC-31C cells (CCL 130) These cells Bellini WJ and Sever JL (2000) In Clinical Virology Manual , Third edition, edited by S were adapted to cell culture by alter- Specter et al . Washington: ASM Press, p. 501 nate passage in animals from a BALB/ c induced tumor. Moriche virus (MORV) A strain of Acara virus in the genus Orthobunyavirus . Mopeia virus (MPOV) A species in the Isolated from Culex amazonensis in genus Arenavirus, belonging to the Trinidad. Not reported to cause disease LCM–LASV complex (Old World arena- in humans. viruses). The natural rodent host is Mastomys natalensis, and virus isola- morphological alteration and disintegrations have been made in Mozambique tion test (MADT ) A test for the presence and Zimbabwe. A reassortant virus of hepatitis B virus involving electron between Lassa fever virus and Mopeia microscopy of purified virus, used to virus has been tested experimentally as test the inactivation of virus infectivity a vaccine against Lassa fever. by disinfectants before infectivity tests Synonym : Mozambique virus. using cell cultures became available. Emonet S et al (2006) Virology 350 , 251 Morro Bay virus (MBV) A possible spe- Johnson KM et al (1981) Am J Trop Med Hyg cies in the genus Orthobunyavirus , 30 , 1291 Moshkoff DA et al (2007) Virus Genes 34 , 169 isolated from salt marsh mosqui- Wulff H et al (1977) Bull World Health Organ toes, Aedes squamiger , in California, 55 , 441 USA. Antigenically a member of the California serogroup. Not shown to Moraten A strain of Edmonston B mea- cause disease in humans. sles virus that was further attenuated Fulhorst CF et al (1996) Am J Trop Med Hyg by multiple passages in chick embryo 54 , 563 fibroblast cells, and licensed in 1968. It is used in the USA as the measles Morumbi virus A possible species in the component of MMR vaccine. Moraten genus Phlebovirus , isolated from the means mor e at t enuated. The virus blood of a human case of febrile illness is closely similar if not identical to from the Amazon region of Brazil. Schwartz virus. Mosqueiro virus (MQOV) An unas- Rota JS et al (1994) Virus Res 31 , 317 signed species in the family Rhabdoviridae, belonging to the Hart Moravia/Ma5302V virus A strain of Tula Park serogroup. Isolated from mosqui- virus, in the genus Hantavirus . Isolated toes, Culex portesi, at Mosqueiro Beach, from the rodents Microtus arvalis and Colombia. Microtus rossiaemeridionalis . Mossman virus An unassigned species morbilli virus Synonym for Measles virus . or strain in the family Paramyxoviridae . Mossman virus 314

A rodent virus isolated on two occasions Mouse mammary tumor virus (MMTV) from wild rats trapped in Queensland, The type species of the genus Australia. The complete genome Betaretrovirus. The genome, in addition sequence resembles those of Salem to gag , pol , and env genes, has a gene virus and Tupaia paramyxovirus, and sag within the 3 LTR whose product is different from that of the Morbillivirus is a super-antigen. The LTR is 1330 nt and the Henipavirus genera. long (U3-1195, R-15, U5-120). The Miller PJ et al (2003) Virology 317 , 330 tRNA primer is tRNAlys-3. There are a number of exogenous strains (labora- Mosso das Pedras virus (MDPV) A spe- tory strains) transmitted via the milk cies in the genus Alphavirus . Isolated as well as endogenous viruses demon- in 1976 from culicine mosquitoes col- strable after the milk-transmitted virus lected at Sitio de Mosso das Pedras, Sao has been eliminated by foster nursing. Paulo State, Brazil. Phylogenetically No oncogenes are found in the genome related to Rio Negro virus and a mem- and the viruses cause mammary adeno- ber of the Venezualan equine encepha- carcinomas following infection of mam- litis virus complex, subtype I, variety F. mary epithelial cells by insertional An enzootic virus which has not been mutagenesis, especially by activating associated with disease in horses. the proto-oncogenes wnt-1 , int-2 , and wnt-3, which encode growth factors. Calisher CH et al (1982) Am J Trop Med Hyg However, there are other integration 31 , 1260 loci and endogenous mammary tumor Mossuril virus (MOSV) An unassigned virus DNA can be mapped to at least species of the family Rhabdoviridae , four separate chromosomes: 7, 11, 15, belonging to the Hart Park sero- and 17. In addition, MMTV can cause group. Isolated from Culex sp in South T lymphomas, although the puta- Africa, Central African Republic, and tive proto-oncogenes are not known. Mozambique. Not reported to cause As there is no agreed nomenclature, disease in humans. strains are probably best named after the mouse strain from which they were Mount Elgon bat virus (MEBV) A tenta- isolated. Often a letter is appended, tive species in the genus Vesiculovirus . e.g. MTV-S (S for standard). Other let- Isolated from a bat in Kenya. Virions ters are: L for low oncogenicity (this larger than most rhabdoviruses was formerly described as nodule- (mean 226 nm). Propagated in new- inducing virus and designated NIV); born mouse brain but not in cell cul- P for plaque-inducing; O for over- tures. Not reported to cause disease in looked; and X or Y for irradiation- humans. induced. The endogenous virus will often fail to infect the strain of mouse from which it came but will infect other mouse cytomegalovirus (MCMV) See strains. Can be propagated in cultures Murid herpesvirus 1 . of mammary tumor tissue, although the cells often lose their ability to pro- mouse Elberfeld virus Synonym for duce mammary tumor virus and start Encephalomyocarditis virus . to produce C-type viruses. Attempts to infect normal cells in culture have not mouse encephalomyelitis virus See been very successful. There are three Theilovirus . classes of laboratory mice: (1) Those that receive infectious virus mouse hepatitis virus See Murine hepa- in their mothers ’ milk, do not pro- titis virus . duce antibodies and have a high incidence of tumors. mouse herpesvirus strain 68 Synonym (2) Those that do not receive virus for Murid herpesvirus 4 . in their milk but have an intermediate tumor incidence, pre- mouse leukemia virus See Murine leuke- sumably due to endogenous virus mia virus . expression. 315 MTV-S virus

(3) Those that neither express virus MP 401 virus (NDV) A strain of Nyando nor develop tumors. virus in the genus Orthobunyavirus , iso- Synonym : MTV-S. lated from mosquitoes in Kenya and Coffin JM, Hughes SH, and Varmus H Central African Republic. Isolated also (Editors) (1997) Retroviruses . New York: from humans but not associated with Cold Spring Harbor Laboratory human disease. Cohen JC and Varmus HE (1979) Nature 278 , 418 MPC-11 cells (CCL 167) Established from Morris DW (1991) Rev Med Virol 1 , 223 the Merwin plasma cell tumor-11 car- Ross SR (2000) Microbes Infect 2 , 1215 ried in BALB/c mice. mouse papule agent (MPA) An uniden- MPK cells (CCL 166) Initiated from the tified infective agent in mice, causing trypsinized kidney tissue of a 118- a self-limiting papular lesion which day-old mini-pig obtained by hyster- heals with no obvious sequelae. The ectomy and processed in a germ-free condition can be serially transmit- environment. ted by injection of extracts of lesions, and there is an incubation period of M’Poko virus (MPOV) A species in the 4–5 days. Eosinophilic inclusion bodies genus Orthobunyavirus . Isolated from appear in the cytoplasm of epidermal mosquitoes. Not known to cause dis- cells. Mouse papule agent has neither ease in humans. been visualized nor cultured. M’Poko Yaba-1 virus See Yaba-1 virus . Kraft LM and Moore AE (1961) Z Versuchstierk 1 , 66 M protein See matrix protein .

Mouse parvovirus 1 (MPV) A species in MRC-5 cells (CCL 171) Derived from the the genus Parvovirus. Distinct from normal lung tissue of a 14-week-old mice minute virus on the basis of male fetus. Susceptible to a wide range genome sequence and hemagglutinat- of human viruses and used for the pro- ing properties. duction of viral vaccines. Tattersall P (2008) in BWJ Mahy and MHV van Regenmortel (eds) Encyclopedia of MRM31 virus (KOOV) A strain Virology , Third edition, Oxford: Academic of Koongol virus in the genus Press, vol. 4, p. 90 Orthobunyavirus . mouse poliovirus Synonym for Theiler’s MS-1 virus A strain of Hepatitis A virus in murine encephalomyelitis virus. the genus Hepatovirus . Originally identified as one of two hepatitis-induc- mousepox virus Synonym for Ectromelia ing viruses in the Willowbrook State virus . School for the Mentally Handicapped. The other virus, MS-2, was hepatitis B. mouse sarcoma virus See murine sar- MS-1 was demonstrated in the sera of coma viruses . hepatitis patients by inoculation into normal human volunteers, in whom mouse thymic herpesvirus Synonym for the disease could be passaged by the murid herpesvirus 3. fecal–oral route with a relatively short incubation period of about 4 weeks mouse-tropic strain Synonym for eco- compared to MS-2 virus. tropic murine type C virus. Feinstone SM et al (1973) Science 182 , 1026 mouse type C oncovirus See Murine Krugman S and Giles JP (1970) J Am Med leukemia virus . Assoc 212 , 1019 movar herpesvirus Synonym for Bovine MSD virus Marble spleen disease virus herpesvirus 4 . of pheasants.

Mozambique virus Synonym for Mopeia MTV-S virus Synonym for Mouse mam- virus . mary tumor virus . Mucambo virus (MUCV) 316

Mucambo virus (MUCV) A species in mule deerpox virus (DPV) An unas- the genus Alphavirus, closely related to signed virus in the family Poxviridae . Venezuelan equine encephalomyelitis Isolated in Wyoming, USA from the virus. Isolated from humans, rodents, mule-deer, Odocoileus hemionus . birds, and mosquitoes in São Paulo and Para, Brazil, and Trinidad, Surinam, Muleshoe virus (MULV) A species in the and French Guiana. Causes a febrile genus Hantavirus, genetically related illness with headache and myalgia in to Sin Nombre virus. Isolated from the humans. cotton rat Sigmodon hispidus (western Scherer WF and Pancake BA (1970) Am J form) in Texas. Not known to cause Epidemiol 91 , 225 hantavirus pulmonary syndrome. mucosal disease virus Synonym for multicomponent viruses Viruses in bovine diarrhea virus. which the complete viral genome is not present in a single virus particle but is Mucura virus A possible species in the divided between two or more particles. Phlebovirus genus . Isolated from phle- The partial genomes present in sepa- botomine sand flies in the Amazon rate particles are not completely over- region of Brazil. Not associated with lapping. The mixture of particles can disease in humans. replicate but the individual particles cannot. Defective viruses and helper Mudjinbarry virus (MUDV) A serotype viruses are thus excluded. There are or strain of Wallal virus in the genus many examples among the small RNA Orbivirus . Isolated from midges col- viruses of plants. lected in the Northern Territory of Australia. Antibodies have been found Reijnders L (1978) Adv Virus Res 23 , 79 in 5 of 30 wallabies, 2 of 12 dingoes, 1 of 30 domestic fowl, and in 1 of 53 Multifocal Castleman’s disease A rare human sera tested. Not known to cause nonmalignant progression of lym- disease in any of these species. phoid tissues which typically presents Doherty RL et al (1978) Aust J Biol Sci 31 , 97 as mediastinal masses. First described by Dr. Benjamin Castleman in 1956. Muhlbock virus A highly oncogenic Frequently associated with both HIV strain of Mouse mammary tumor virus infection and human herpesvirus 8, but isolated from GR mice. Also known as the precise etiology of the condition is GR virus or MTV-P. Differs from other still not clear. strains of high oncogenicity in being transmitted via the eggs and sperm. multimammate mouse papillomavirus (MmPV) A probable species in the Muir Springs virus (MSV) An genus Papillomavirus. Causes sponta- unassigned virus in the family neous papillomas in the skin of inbred Rhabdoviridae , belonging to the Bahia line ‘ GRA Giessen ’ of the multimam- Grande serogroup. Isolated from mos- mate mouse, Mastomys natalensis . quitoes, Aedes sp. Crystalline arrays of virus particles are Kerschner JH et al (1986) J Gen Virol 67 , 1081 present in the upper layers of the thickened stratum granulosum. Antiserum Muju virus A probable species in the can be prepared in rabbits. genus Hantavirus. Found in the arvi- Muller H and Gissmann L (1978) Med colid rodent Myodes regulus in Korea. Microbiol Immunol 165 , 93 The genome sequence is 77% similar Muller H and Gissmann L (1978) J Gen Virol to that of Puumala virus. Serological 41 , 315 evidence suggests that this virus may cause a proportion of cases of hemor- multinucleated syncytia formation A rhagic fever with renal syndrome in common cytopathic effect of infection Korea. by viruses which fuse with the cell Song KJ et al (2007) J Gen Virol 88 , 3121 membrane, such as paramyxoviruses. 317 Murid herpesvirus 1 (MuHV-1) multipartite genome A viral genome pancreatitis. Incubation period 18–21 divided between two or more nucleic days. Fatalities occur occasionally acid molecules. These may be encap- as a result of meningoencephalitis, sidated in the same particle, e.g. reo- with an overall case fatality ratio of virus, orthomyxovirus, or in separate 2–4 per 10,000. Virus present in saliva particles, in which case they are termed and urine. Patients are infectious 6 ‘ multicomponent. ’ See multicompo- days before onset and for 9 days after. nent viruses . Rhesus monkeys are susceptible and develop a similar disease to that seen multiple sclerosis-associated retrovirus in humans. Virus can be adapted to (MSRV) A virus isolated from the plasma hamsters, mice, and rats. Isolation is of MS patients which is phylogenetically by amnion inoculation, when it takes and experimentally related to human 4–5 days, or in primary monkey kid- endogenous retroviruses (HERVs). ney or human cell lines. Multiplication Blond JL et al (1999) J Virol 73 , 1175 is recognized by syncytia formation or hemadsorption. Human and multiple sclerosis virus Despite numer- fowl erythrocytes are agglutinated. ous claims, no virus has yet been A number of attenuated vaccine shown to be the cause of this disease. strains are now available, such as the Nevertheless, the disease has features Jeryl Lynn B strain produced in chick which suggest that a virus may be embryo fibroblast cells. This is admin- involved in the etiology. See carp virus . istered as part of the MMR (measles– mumps–rubella) vaccine given at about multiplicity of infection (moi) Ratio of age 12 months to provide long-term number of infectious virus particles protection against infection. added to a known number of cells in a Galazka AM et al (1999) Bull World Health culture. Organ 77 , 3 Knowles WA and Li Jin (2005) In Topley & multiplicity reactivation A form of reas- Wilson’s Microbiology and Microbial Infections , vol. 1, Tenth edition, edited by BWJ Mahy and sortment or complementation between V ter Meulen. London: Hodder Arnold, p. 744 two related viruses which have been Li Jin et al (2000) Virus Res 70 , 75 inactivated. The sites of inactivation must be in different parts of the Munguba virus (MUNV) An isolate of genomes of the two viruses. Bujaru virus in the genus Phlebovirus . Isolated from phlebotomine flies multiploid virus Virus containing a pop- Lutzomyia umbratilis in Para, Brazil. ulation of particles, most of which contain a variable number of genomes. The Mupapillomavirus A genus of the family number of genomes per particle may Papillomaviridae containing two species, depend on the host cell and other cul- Human papillomavirus 1 and Human tural conditions, but is independent of papillomavirus 63. Both cause cutane- the number of genomes contained by ous lesions in their host which are the infecting virus, i.e. it is not a geneti- distinguishable by histology and by cally determined characteristic. the presence of intracytoplasmic inclu- Simon EH (1972) Prog Med Virol 14 , 36 sion bodies. They also have a distinct genome organization with a larger con- Mumps virus (MuV) Type species of the trol region than other genera. genus Rubulavirus. Mumps virions have both hemagglutinating and neu- Murid herpesvirus 1 (MuHV-1) The type raminidase activities. Virion diameter species of the genus Muromegalovirus . 150 nm. The genome RNA is negative- A β -herpesvirus with a genome DNA stranded (15384 nt) with the gene order 235 kb in length; G C is 59%. Probably 3 -NP-P/V-M-F-SH-HN-L-5 ’ . A human a ubiquitous silent infection of wild infection commonly causing fever and mice but present in a minority only of parotitis, and occasionally meningo- laboratory stocks. Young uninfected encephalitis, orchitis, oophoritis, or mice can be infected by any route, the Murid herpesvirus 1 (MuHV-1) 318

virus localizing in the salivary glands, in Slovakia. Genome DNA 135 kb in in which tissue alone is serial passage length; G C content 45%. Establishes possible. Large doses of virus given i.p. infection in inbred laboratory mice, will kill mice in 4–7 days. Small doses a latent infection in B cells, and about produce focal hepatitis. Infection of 10% of the mice develop lympho- pregnant mice causes fetal infection. proliferative disease. Causes a persis- Replication occurs in primary mouse tent, latent infection in B-cell lines infec- fibroblasts with focal CPE in 9–12 days. ted in vitro . Virus is released into the medium Synonym : mouse herpesvirus strain 68. much more freely than with most Efstathiou S et al (1990) J Gen Virol 71 , 1355 cytomegaloviruses. Nash AA and Suril-Chandra NP (1994) Curr Synonyms : murid herpesvirus; mouse Opin Immunol 6 , 560 cytomegalovirus. Ebeling A et al (1983) J Virol 47 , 421 murid herpesvirus 5 (MuHV-5) An Moon NM et al (1979) J Gen Virol 42 , 159 unassigned species in the family Reddehase MJ (2000) Curr Opin Immunol 12 , Herpesviridae. Isolated from newborn 390 field mouse, Microtus pennsylvanicus , cell culture which showed spontane- Murid herpesvirus 2 (MuHV-2) A species ous CPE. Replicates in a wide variety in the genus Muromegalovirus . Present of cell lines, including Vero cells, with in the salivary glands of rats; can cause CPE. Some antigenic relationship to abortion in laboratory rats. Agglutinates human herpesvirus 1. rabbit erythrocytes at 4°C. Synonyms: field mouse herpesvirus; Synonyms: rat cytomegalovirus; rat Microtus pennsylvanicus herpesvirus. submaxillary gland virus. Melendez LV et al (1973) Lab Anim Sci 23 , Ashe WK (1969) J Gen Virol 4 , 1 385 Beisser PS et al (1998) Virology 246 , 341 Vink C et al (2000) J Virol 74 , 7656 murid herpesvirus 6 (MuHV-6) An unassigned species in the family murid herpesvirus 3 (MuHV-3) An Herpesviridae, isolated from a labora- unassigned species in the family tory colony of Egyptian sand rats, Herpesviridae. Injection into newborn Psammomys obesus, in which there had mice produces extensive necrosis of the been numerous deaths. Post-mortem thymus resulting in profound suppres- examination was inconclusive, and the sion of immunological functions medi- virus was isolated in primary rabbit ated by T cells. However, some T-cell kidney cell cultures from throat swabs functions, such as thymic cell reaction of live animals. CPE appeared after to mitogens, appear to be spared sug- 8 days and progressed to confluence by gesting that virus selectively destroys 11 days. Type A intranuclear inclusion sub-populations of T cells. The virus bodies are produced. Replicates with can be obtained from homogenates of CPE in HeLa, human amnion, sand mouse thymus during the acute phase rat, and squirrel monkey cells. Not of infection. In adult mice there is pathogenic for 5-day-old mice or rats. chronic infection of the salivary glands Probably not pathogenic for sand rats. without cell necrosis. Replication in cell Synonyms: sand rat nuclear inclusion cultures has not been reported, but the agent; sand rat herpesvirus. virus can be obtained from homoge- Melendez LV et al (1967) Lab Anim Care 17, 302 nates of mouse thymus during the acute phase of infection. murid herpesvirus 7 (MuHV-7) A prob- Synonym : mouse thymic herpesvirus. able species in the family Herpesviridae , Houba V et al (1976) J Immunol 117 , 635 isolated from mice in Slovakia. Parker J et al (1973) Infect Immun 7 , 305 Synonym : murine herpesvirus. Blaskovic D et al (1988) Acta Virol 32 , 329 Murid herpesvirus 4 (MuHV-4) A species in the genus Rhadinovirus, isolated from Murine adenovirus A (MAdV-A) A spe- a bank vole, Clethrionomis glareolus , cies in the genus Mastadenovirus . 319 Murine leukemia virus (MLV)

Usually a silent infection of mice but virus of rats. Often a silent infection may be fatal on injection into new- of laboratory mice which may be acti- born mice. Excretion of virus in the vated by passage of other viruses such urine can continue for a long time. as leukemia virus, or by cortisone, Multiplies with CPE in mouse embryo urethane or enterotoxin from gram- cell cultures but not in mouse cell lines negative bacteria, or by thymectomy. or rat, monkey or human cells. Some Injection of virus into mice infected strains do not agglutinate erythrocytes with the parasite Eperythrozoon coc- of human, monkey, rabbit, guinea pig, coides usually produces fatal hepatitis mouse, etc. but in its absence there is often no dis- Synonym : murine adenovirus 1. ease. Neurotropic strains infect cotton van der Veen J and Mes A (1974) Arch ges rats and hamsters when given i.c. All Virusforsch 45 , 386 strains are antigenically similar but vary in pathogenicity, and have been murine adenovirus B (MAdV-B) A numbered 1–4. Type 1 was the original tentative species in the genus Masta- isolate. Type 2 was activated on mouse denovirus . leukemia passage in Princeton mice Synonym : murine adenovirus 2. and produced hepatitis in the absence of E coccoides . Type 3 is also pathogenic murine adenovirus 1 (MadV-1) Synonym for weanling mice in the absence of E for Murine adenovirus A . coccoides , and causes ascites in older mice. Type 4 causes encephalomyeli- murine adenovirus 2 (MadV-2) Synonym tis with demyelination and some focal for Murine adenovirus B . liver necrosis in mice. Virus is present in excreta and is highly infectious. murine AIDS (MAIDS) Disease caused Transplacental transmission has not by a defective murine leukemia virus been demonstrated. Genome is positive, (e.g. BM5 def) characterized by lym- single-stranded RNA, 30 kb in length, phadenopathy, splenomegaly, hyper- with a 5 -cap and 3 poly A tract. gammaglobulinemia, profound T- and Synonyms: murine coronavirus; mouse B-cell anergy, late appearance of B- and hepatitis virus. T-cell lymphomas, and development of Cavanagh D et al (1994) Arch Virol 135 , 227 opportunistic infections. Wege H et al (1982) Curr Top Microbiol Immunol 99 , 165 Doyon L et al (1996) J Virol 70 , 1 Morse HL (1992) AIDS 6 , 607 murine herpesvirus Synonym for murid herpesvirus 7. murine astrovirus A tentative member of the family Astroviridae awaiting murine herpesvirus-68 A murine gamma- molecular characterization. herpesvirus which establishes a latent infection in B lymphocytes, dendritic murine coronavirus Synonym for Murine cells, macrophages, and lung epithe- hepatitis virus . lial cells of laboratory mice. The mice develop lymphoproliterative disease murine cytomegalovirus See mouse and an infectious mononucleosis-like cytomegalovirus . syndrome. Also called murid herpesvirus 4. murine encephalomyelitis (ME) virus Nash AA et al (2001) Philos Trans R Soc Lond See Encephalomyocarditis virus . B 356 , 569 murine gammaherpesvirus (MHV-68) murine K virus Synonym for Murine See Murid herpesvirus 4 . pneumotropic virus (Kilham polyomavirus). Murine hepatitis virus (MHV) A species in the genus Coronavirus. A species in Murine leukemia virus (MLV) A species group 2. Serologically related to Rat in the genus Gammaretrovirus . There coronavirus and sialodacryoadenitis are many strains and all mice probably Murine leukemia virus (MLV) 320

carry one or more viruses. The first murine minute virus See Minute virus of strain was isolated from AKR mice by mice . Gross. Strains can be grouped by their envelope antigens or the antigens they murine norovirus 1 A tentative species in induce on the surface of infected cells. the genus Norovirus . They vary in the type of leukemia that they induce, but this also depends on murine papovaviruses See K virus and the strain and age of the host animal. Murine polyomavirus . The Gross, Moloney, and Kaplan strains injected into newborn mice cause thy- murine parainfluenza virus type 1 See mus-dependent lymphocytic leukemia Sendai virus . after a latent period of 3–4 months. The mechanism of leukemogenesis involves Murine pneumonia virus (MPV) A spe- activation of cellular proto-oncogenes cies in the genus Pneumovirus . A com- after proviral insertion. Abelson and mon latent or mild respiratory virus Friend strains injected into adult mice infection of laboratory mice. Inbred cause splenomegaly and erythroblas- mouse strains differ in their suscepti- tic leukemia in a few weeks, as they bility to infection. Can be activated by carry oncogenes in the genome (v-abl , serial intranasal passage of lung tissue which encodes a tyrosine-specific pro- in uninfected mice at intervals of 7–9 tein kinase and gp52 , which encodes a days. Causes dense accumulations of viral glycoprotein). Sarcoma-inducing mononuclear cells around the bronchi strains which carry a transduced cel- and blood vessels. Lung lesions are lular oncogene can transform cells in produced in hamsters but the virus culture but are defective for virus repli- cannot be serially passaged. Virions are cation, requiring a helper leukemogenic 80–120 nm in diameter but filamentous virus to provide information for the forms occur. Envelope covered with viral coat proteins. Laboratory strains projections. There is a helical nucleo- and exogenous viruses are transmitted capsid 12–15 nm in diameter contain- mainly via the milk, but in nature the ing single-stranded RNA. Agglutinates endogenous virus probably passes to mouse and hamster erythrocytes. the young via the egg or sperm. There Replicates in hamster kidney cell cul- are three subspecies: tures and BHK21 cells, recognized by (1) Ecotropic murine type C viruses, hemadsorption. Antigenically related the laboratory strains of exogenous viruses have been isolated from ham- leukemia- and sarcoma-inducing sters, cotton rats, and rabbits. The virus. Members of this subspecies genome RNA has nine genes, in the show reciprocal viral interference. same order as respiratory syncytial (2) Xenotropic murine type C viruses. virus. (3) Amphotropic murine type C viruses. Synonyms : mouse type C oncovirus; Anh DB et al (2006) Am J Physiol Lung Cell Moloney virus. Mol Physiol 291 , L426 Chambers P et al (1990) Virus Res 18 , 263 Hung Fan (1994) In The Retroviridae , vol. 3, edited by JA Levy. New York: Plenum Press, Murine pneumotropic virus (MptV) p. 313 A species in the genus Polyomavirus Stockert E et al (1979) J Exp Med 149 , 200 which causes acute pneumonia in newborn mice. murine leukemia virus 1313 (MuLV- Synonym : Kilham polyomavirus. 1313) An amphotrophic murine leukemia virus, isolated from wild mice in murine polio-encephalomyelopathy California, which replicates efficiently virus An ecotropic strain of Murine in both mammalian and chicken cells. leukemia virus. Causes a noninflamma- Can be distinguished from most other tory spongiform degeneration of the murine leukemia virus isolates by phylo- CNS which results in paralysis of the genetic analysis. limbs. The condition occurs spontane- Howard TM et al (2006) Virol J 3 , 101 ously in wild mice between the ages 321 Murray Valley encephalitis virus (MVEV)

of 7 and 18 months. It can be produced sarcomas. Subsequently, other investi- in Swiss but not BALB/c (Mo) mice by gators made similar observations and i.c. injection of virus when they are less several viruses, each having differ- than 24 h old. The incubation period is ent transduced cellular oncogenes, are dose-dependent and can be as short as now recognized: Harvey-MSV (H-ras ), 3 weeks. Moloney MSV (mos ), Kirsten-MSV (Ki- Synonym : polio-encephalomyelopathy ras), FBJ MSV ( fos), FBR-MSV ( fos-fox ), of mice virus. 3611 MSV (raf ). The H-ras and Ki- ras genes are different but related cellu- Brooks BR et al (1979) Infect Immun 23 , 540 lar genes. Because the transduced cellular gene is inserted into the original murine poliovirus Synonym for Theiler’s murine leukemia virus genome, these murine encephalomyelitis virus. acute transforming murine sarcoma viruses are replication-defective. They Murine polyomavirus (MPyV) A species induce sarcomas in mice after a latent of the genus Polyomavirus . A natural period of only a few days and trans- infection of wild and laboratory mice. form fibroblasts in cell culture, but Causes no disease under natural con- are unable to produce infective prog- ditions, but if injected into newborn eny virus in the absence of a mouse mice or hamsters is highly oncogenic. leukemia virus which acts as a helper. Replicates with marked CPE in mouse The Harvey and Kirsten strains cause embryo cell cultures. Hamster cell marked erythroblastic splenomegaly cultures are not permissive for virus and progressively growing sarcomas. replication but are transformed by the The Moloney strain does not affect the virus. Hemagglutinates erythrocytes of erythroid cells and the tumors which it several species, e.g. guinea pig, at 4°C induces usually regress except in very by reacting with the neuraminidase- young or immunosuppressed mice. sensitive receptors, and nonenzymic de Vos AH et al (1988) Science 239 , 888 elution occurs at room temperature. The genome is circular DNA, 5.3 kb in murivirus Synonym for human rhino- length, and specifies three T antigens: virus. ST, MT and LT. Both MT and LT are involved in cell transformation induced Muromegalovirus A genus in the sub- by the virus. LT (785 amino acids) com- family Betaherpesvirinae. The type plexes with the tumor-suppressing species is Murid herpesvirus 1 (mouse protein, Rb, and inactivates it; MT (421 cytomegalovirus). Contains two spe- amino acids) complexes with three cell cies Murid herpesvirus 1 and Murid her- proteins, including c-src pp60, and pesvirus 2 (rat cytomegalovirus). The so contributes to cell transformation. genomes are large ( 200 kbp) and form Polyomaviruses have proved to be val- a distinct lineage within the subfamily uable models of virus replication and Betaherpesvirinae . cell transformation mechanisms. Consigli RA and Center MS (1978) CRC Crit Murray Valley encephalitis virus Rev Microbiol 6 , 263 (MVEV) A species in the genus Otte, J et al (2005) In Topley & Wilson’s Flavivirus, belonging to the Japanese Microbiology and Microbial Infections , vol.1, encephalitis serogroup. Natural host Tenth edition, edited by BWJ Mahy and V probably a bird. The major vector is ter Meulen. London: Hodder Arnold, p. 473 Culex annulirostris, but other mosquito species may be involved. Occurs in murine sarcoma viruses (MSV) Several Northern Territory and Queensland, species in the genus Gammaretrovirus Australia, and in Papua New Guinea. which carry transduced cellular onco- After the spring rains the virus is car- genes. The first to be described, Harvey ried south to Victoria, New South murine sarcoma virus (HaMSV), was Wales, and South Australia. In humans isolated from rats that had been injected it causes a mild fever and, in some with high-titered murine leukemia cases, encephalitis. There may be trou- virus preparations and had developed blesome sequelae. Epidemics occur Murray Valley encephalitis virus (MVEV) 322

and children are most often infected. virus. Obtained from the milk. Distinct Horses may be infected but do not from Mus cervicolor type C retroviruses. develop encephalitis. The major verte- Schlom J et al (1978) J Natl Cancer Inst 61 , brate hosts may be herons, especially 1509 the rufous night heron, Nycticorax caledonicus. Encephalitis follows i.c. Mus cervicolor type C retro- injection in mice, hamsters, monkeys, viruses Possible species in the genus sheep, and chicks. Rabbits, guinea pigs, Gammaretrovirus . Isolated from a lung and birds usually only have viremia. cell line of Mus cervicolor cells by treat- Antibody is present in the yolk of eggs ment with bromodeoxyuridine and laid by infected birds. Virus replicates co-cultivation with heterologous cell in eggs, producing pocks on the CAM. lines. Two viruses, CERV-CI and CERV- Synonym : Australian X-disease virus. CII, and endogenous multiple copies Johansen CA et al (2007) Virus Genes 35 , 147 are present in the cellular DNA. CERV- Mackenzie JS et al (1994) Arch Virol 136 , 447 CI replicates in SIRC rabbit cell line and is antigenically related to simian sar- Murray Valley virus Synonym for coma virus, simian sarcoma-associated Murray valley encephalitis virus . virus, and to the Gibbon ape leukemia virus. CERV-CII replicates in Mus mus- Murre virus (MURV) A strain of culus cell lines. It is related to, but dif- Uukuniemi virus in the genus ferent from, mouse leukemia viruses Phlebovirus . Isolated from Uria aalge . of Mus musculus. A third virus, M432, is an endogenous virus unrelated by morphology, antigenicity, or molecu- Murutucu virus (MURV) A strain lar hybridization to other retroviruses of Marituba virus in the genus except Mus caroli type C oncovirus. Orthobunyavirus , belonging to the group C viruses. Has been associ- Benveniste RE et al (1977) J Virol 21 , 849 ated with a febrile illness in humans. Callahan R et al (1977) Virology 80 , 401 Isolated from a sentinel Cebus monkey and mice in Para, Brazil. Has also been Muscovy duck parvovirus (MDPV) A isolated from the rodents, Nectomys species in the genus Parvovirus . squamipes and Proechimys guyannensis , the opossums, Didelphis marsupialis Mus dunni retrovirus (MDEV) An endo- and Marmosa sp, and the mosquitoes of genous murine retrovirus first identified Culex sp. during tests for possible helper viruses in human gene transfer experiments Mus caroli type C retrovirus A possible with retroviral vectors. The helper species in the genus Gammaretrovirus . assay cells from Mus dunni could be An endogenous xenotropic C-type activated by treatment with hydrocor- virus. Found in a cell line derived from tisone or IUDR to produce the virus, the Asian mouse, Mus caroli , on treat- which is unrelated to known murine ment with bromodeoxyuridine. The leukemia virus groups. It is able to reverse transcriptase and p30 antigen infect a wide range of cell types from are more closely related to the woolly different species. monkey sarcoma virus and Gibbon ape Miller AD et al (1996) J Virol 70 , 1804 (leukemia) virus, than to laboratory mouse viruses. mus1-mus2 Serotype designation given Lieber MM et al (1975) Proc Natl Acad Sci 72 , to the mastadenovirus isolated from 2315 the mouse, Mus musculus . Family Adenoviridae . Mus cervicolor mammary tumor virus A possible species in the genus Mustelid herpesvirus 1 (MusHV-1) A spe- Betaretrovirus. The major envelope cies in the genus Rhadinovirus . Isolated glycoprotein and internal protein are from pulmonary fibroblast cultures antigenically related to the equivalent established from a European badger proteins in mouse mammary tumor (Meles meles) from Cornwall, England. 323 myeloblastosis-associated viruses

The virus grew in NPL-7 cells (CCL64 MVC Synonym for Canine minute virus . cells), a line derived from fetal mink lung. A further study indicated that Mv 1 Lu (NBL-7) (CCL 64) A cell line infection with this virus is common in derived from the trypsinized lungs free ranging badgers in the UK. of several nearly full-term unsexed Banks M et al (2002) J Gen Virol 83 , 1325 fetuses of the Aleutian mink, Mustela vison . mutagen A chemical which induces change in the base composition of MVM virus Synonym for Mice minute a virus. Examples are base analogs virus . (5-fluorouracil and 5-azacytidine for RNA viruses and 5-bromodeoxyurid- Mx proteins The products of the Mx ine for DNA viruses); alkylating agents gene are proteins which are induced (ethylmethane sulfonate, diethyl sulfate); by interferon and have antiviral activ- intercalating agents (proflavine and ity against influenza virus, hepatitis C, N -nitro-N -nitrososguanidine); deami- and other viruses. nating agents (nitrous acid); hydroxy- Staeheli P (1990) Adv Virus Res 38 , 147 lamine, UV irradiation. Base analogs Staeheli P et al (1993) Trends Cell Biol 3 , 268 are incorporated into the genome and miscoding during replication produces MxA protein Name for the human pro- mutations. Other mutagens induce tein product of the Mx gene, which has mutation by direct chemical action on antiviral activity. the viral nucleic acid. myalgic encephalomyelitis (ME) Syno- Leppard KN and Pringle CR (1996) In nym for chronic fatigue syndrome. Virology Methods Manual , edited by BWJ Mahy and H Kangro. London: Academic mycoplasma Gram-negative bacteria Press, p. 231 which lack a cell wall and are resistant to many antibiotics. Have occasionally mutant A variant virus containing a been confused with viruses. Formerly gene that has undergone mutation known as pleuropneumonia-like organ- which may be expressed in the pheno- isms (PPLO). type. The standard virus is called the ‘ wild-type. ’ mycoviruses Viruses which replicate in the cells of fungi. mutation A change in the genotype of an organism not resulting from recombi- Hollings M (1978) Adv Virus Res 22 , 2 nation. In its simplest form it is the sub- myelin basic protein (MP) A protein stitution of one nucleotide for another found in brain and cerebrospinal leading to changes in the structure of fluid that appears to be the result of the protein coded for by a nucleotide myelin breakdown as a result of viral sequence or modifying gene regulation encephalitis. A lymphoproliferative sequences. response against MP has been detected in disorders such as measles encepha- mutation rate The error rate of a virus litis, varicella encephalitis and as a nucleic acid during replication. One consequence of post-exposure rabies of the highest rates is seen with HIV, immunization. which is estimated to have an error rate 5 of one base in every 3.4 1 0 . myeloblastosis-associated viruses A name suggested for the avian myelo- MVA M odified v irus Ankara. A strain of blastosis viruses (AMV); helper viruses, vaccinia virus that is under develop- present in ‘ standard ’ AMV but not ment as a gene vector, particularly in leukemogenic. Chicken leukosis sar- conjunction with DNA-based vaccines. coma viruses with surface properties of Antoine G et al (1998) Virology 244 , 365 subgroup A and B, respectively. Cause Hill AVS (2000) In Fighting Infections in the osteopetrosis on injection into day-old 21st Century . London: SGM, p. 87 chicks. Mykines virus (MYKV) 324

Mykines virus (MYKV) A serotype of show a difference. Produces pocks on Great Island virus in the genus Orbivirus , the CAM, but eggs are 2.5 times less belonging to the Kemerovo serogroup. sensitive than rabbit skin. Replicates Isolated from ticks, Ixodes ( Ceratixodes ) and produces CPE in tissue culture of uriae. Not associated with disease in rabbit, rat, hamster, and human cells; humans. also in cells of other species. Exists naturally in Uruguay and Brazil as a Mynahpox virus (MYPV) A species in silent or mild infection of wild rab- the genus Avipoxvirus. Isolated from a bits, Sylvilagus brasiliensis . Introduced captive Rothschild’s mynah, Leucospar into wild European rabbits, Oryctolagus rothschildii , shown to be pathogenic cuniculus, it causes at first a severe for starlings but not for chickens. The disease, 99% fatal, with inflamma- mynah is a member of the same family, tion and swelling of the eyelids, nose, Sturnidae, as starlings and may have genital, and anal openings. With pas- acquired starlingpox by direct contact. sage the disease becomes endemic and Landolt M and Kocan RM (1976) J Wildl Dis less severe due to selection of resist- 12 , 353 ant rabbits and attenuated strains of the virus. Hares, Lepus sp, are rarely Myoviridae A family of double-stranded naturally infected. The virus has been DNA viruses infecting bacteria, includ- propagated in suckling mouse brain. ing T4-like viruses. Transmission is by contact and by insects: mosquitoes in South America myristylation A post-translational modi- and Australia, fleas in the UK. Rabbits fication of virus proteins involving the can be protected by vaccination with covalent attachment of a myristic acid rabbit fibroma virus. Recently myxoma moiety. virus has been shown to selectively infect and kill human tumor cells, and mystery swine disease (Lelystad virus) is being considered as a candidate for Synonym for Porcine respiratory and use in oncolytic virotherapy. reproductive syndrome . Fenner F and Ross J (1994) In The European Rabbit , edited by HV Thompson and C King. myxoma subgroup viruses Synonym for Oxford: Oxford University Press, p. 205 leporipoxvirus. Stanford MM and McFadden G (2007) Expert Opin Biol Ther 7 , 1415 Myxoma virus (MYXV) The type species of the genus Leporipoxvirus . Similar to myxovirus multiforme A latinized name vaccinia virus but rather more sensitive for Newcastle disease virus, based on to inactivation by heat. Ether-sensitive the multiformity of both the disease but resistant to sodium desoxycholate. picture and the virus particles. Not in The DNA genome is 160 kb in length, common use. G C 40%. Antigenically similar to rabbit fibroma virus and California rab- myxovirus pestis-galli See influenza bit fibroma virus, yet diffusion tests virus A avian . N

NAAT N ucleic a cid a mplification in mice inoculated i.c. Transmitted by t echnique. the tick, Rhipicephalus appendiculatus . Virus replicates in cell cultures of N-acetylneuraminic acid A compound lamb and goat tissue with CPE. Cases derived from acetic acid, mannosine of human infection with fever and and pyruvic acid that is a major con- arthralgia have been reported, as well stituent of eukaryotic cell membranes. as cases of serological conversion with- See neuraminidase . out disease. Synonym: sialic acid. Davies FG et al (1978) J Comp Pathol 88 , 519

NAD N icotinamide a denine d inucleotide. Nairovirus A genus in the family Formerly known as diphosphopyrid- Bunyaviridae, type species Dugbe virus . ine nucleotide, DPN, coenzyme I, or There are 7 species in the genus, includ- cozymase. ing the important Crimean-Congo hemorrhagic fever virus. Nairoviruses are NADP N icotinamide a denine d inucle- morphologically similar and have no otide p hosphate, coenzyme II. antigenic cross-reaction with other genera in the family Bunyaviridae . Nagata strain of fecal virus A strain The genome is single-stranded nega- of virus found in Japan, related to tive-sense RNA in three segments, Norwalk virus. each of which have the sequence AGAGUUUCU at the 3 terminus and NA gene The neuraminidase gene (seg- UCUCAAAGA at the 5 terminus. ment 6) of influenza virus. In influenza B virus this gene gives rise to a bicis- naked viruses Viruses without a lipo- tronic mRNA containing two initiating protein envelope. AUG codons that are separated by four nucleotides. From the 5 AUG codon NANB hepatitis virus Synonym for non-A a 100 amino acid NB protein is trans- non-B hepatitis virus (e.g. Hepatitis C lated, and from the second AUG codon virus). the 466 amino acid neuraminidase (NA) protein is translated. The NB protein is Nandi virus A strain of Mouse mammary thought to function as an ion channel tumor virus, of low oncogenicity, in the during replication, but removal of the genus Betaretrovirus . Isolated from C3H NB gene by reverse genetics did not mice. Also known as MTV-L and NIV. affect virus replication in cell culture, though the NB-minus virus showed Naples virus infections The Sandfly fever restricted growth in mice. Naples virus has been recognized as a cause of militarily important disease Nairobi sheep disease virus (NSDV) since the Napoleonic wars. During both A strain of Dugbe virus in the genus world wars, foreign troops suffered sig- Nairovirus . Found in Kenya, Uganda, nificant incidence rates of sandfly fever Congo, Ethiopia, Somalia, and Tanzania. in the Mediterranean. The disease was Causes a non-contagious hemorrhagic first described by Sabin as an abrupt gastroenteritis in sheep and goats with fever onset lasting 2–4 days, headache, high mortality. There is splenic enlarge- eye pain, photophobia, back and joint ment and involvement of the female pain, anorexia and malaise. No deaths genital tract. Nephritis and myocardial have been documented. Sandfly fever degeneration occur. Causes encephalitis Sicilian virus causes similar symptoms. Naples virus infections 326

Sabin A (1951) Arch Gesamte Virusforsch T-cells and B-cells, but express CD16 4 , 367 and CD56 as characteristic markers. They appear to have no memory, MHC- Naranjal virus (NJLV) A strain of Aroa restriction or requirement for antibody. virus in the genus Flavivirus . Isolated They can be activated by cytokines from a sentinel hamster in coastal such as interleukin-2, and they secrete Ecuador. certain other cytokines such as interferon γ and tumor necrosis factor α . Nariva virus An unassigned virus in the Cytokine-activated NK cells are known family Paramyxoviridae isolated from for- as LAK cells. The mechanism of killing est rodents in Trinidad. Morphologically is similar to that of cytotoxic T-cells. and biologically similar to viruses of the They contain granzymes and release family. Kills newborn mice on i.c. injec- perforins which are inserted into the tion. Hamsters and guinea pigs pro- membrane of the target cell. NK cells duce antibodies but develop no disease. display a number of receptors involved No isolations from arthropods. Not in the activation and inhibition of NK reported to cause disease in humans. cell function. Activation receptors on Tikasingh ES et al (1966) Am J Trop Med Hyg human cells include NKp30, NKp44, 15 , 235 and NKp46, which recognizes the influenza virus hemagglutinin. To pro- nasal vaccination Intranasal vaccina- tect normal cells from damage, NK tion has the advantage of stimulating cells also carry inhibitory receptors, higher levels of mucosal immunity such as killer cell immunoglobulin-like (IgA antibodies) and is less invasive receptors (KIR), immunoglobulin-like than parenteral administration. A nasal inhibitory receptors (ILT) and lectin- spray influenza vaccine (FluMist) like heterodimers CD94-NKG2A. NK is now licensed annually for use in cells can also recognize IgG antibody- healthy people 2–49 years of age who coated cell surfaces via CD16 and kill are not pregnant. target cells. NK cells are rapidly mobi- lized following virus infection, and are nascent In the process of being detected at sites of infection within 2 synthesized. days of virus entry. Biron CA et al (1999) Annu Rev Immunol nascent cleavage Cleavage of a poly- 17 , 189 protein occurring at the same time as it Moretta A (2002) Nat Rev Immunol 2 , 957 is being synthesized by cell ribosomes. natural selection A principle put forward nascent RNA RNA in the process of by Charles Darwin in 1859 that natural being synthesized. processes favor members of a species that are best adapted to their environ- Nasoule virus (NASV) An unassigned ment and tend to eliminate those that animal rhabdovirus. Isolated from are unfitted to their environment. Synonym the bird, Andropadus virens, in Central : Darwinian selection. African Republic. Navarro virus (NAVV) An unassigned Sureau P (1974) Institut Pasteur Bangui, animal rhabdovirus. Isolated from Annual Report , 37 the turkey-vulture, Cathartes aura , in Colombia. Not reported to cause dis- natural cowpox virus Synonym for ease in humans. Pseudocowpox virus . NB41A3 cells (CCL 147) A cell line estab- natural killer (NK) cells Large granular lished by cloning a mouse neuroblas- lymphocytes which play a role in innate toma, C-1300. immunity, and can cause lysis of virus- infected cells and tumor cells without NB protein An integral membrane protein conventional immunological specificity. which forms an ion channel in influ- Lack the phenotypic markers of both enza virus B, structurally analogous 327 necrotic rhinitis virus

to the M2 protein of influenza virus A. subcutaneous areolar and adipose Synthesized by a bicistronic mRNA tissue of a normal 100-day-old male derived form segment 6 of the genome, C3H/An mouse, and was one of the which also encodes the neuraminidase. first cell lines to be established in continuous culture. Clone 929 was estab- NCTC 2071 cells (CCL 1.1) Derived lished from the 95th passage. from NCTC clone 929 and established on protein-free chemically defined Ndelle virus (NDEV) A subtype of medium. Mammalian orthoreovirus in the genus Orthoreovirus . Isolated from Mus minu- NCTC 3749 cells (CCL 46.1) The parent toides (musculoides) in Cameroon. Not line P388 D1 was derived from a methyl- known to cause disease in humans. cholanthrene-induced lymphoid neo- Institut Pasteur, Dakar (1975) Annual Report plasm (P388) originating in a DBA/2 mouse and converted to ascitic form in the first mouse transfer. Ndumu virus (NDUV) A species in the genus Alphavirus . Isolated from mos- NCTC 4093 cells (CCL 63) Developed quitoes in South Africa. No known from two individual cells of a mouse association with human disease. Kills embryo cell line isolated by the capil- newborn mice on injection. lary cloning technique. Kokernot RH et al (1961) Am J Trop Med Hyg 10 , 383 NCTC 4206 cells (CCL 14.2) Cell line adapted from a subline culture of nearest-neighbor base frequency analy- B14FAF28-G3 (ATCC 14), derived from sis A classical method of characteriz- an adult female Chinese hamster cul- ing DNA molecules, which compares tured in serum-supplemented medium. the frequencies with which any pair of adjacent bases occurs. Now largely NCTC clone 1469 cells (CCL 9.1) A cell supplanted by nucleotide sequence line derived from a subline of the analysis. original mouse liver strain NCTC 721, which was derived from the normal Nebraska bovine enteric calicivirus liver tissue of a 2-day-old male mouse. A strain of bovine enteric calicivirus which is related to the Lagovirus and NCTC clone 2472 cells (CCL 11) Derived Sapovirus genera, but may represent a from NCTC strain 1742 (subline VII) of new calicivirus genus. the parent NCTC clone 1328. The par- Smiley JR et al (2003) J Clin Microbiol 41 , ent line was derived from a culture 3089 of subcutaneous areolar and adipose connective tissue taken from a normal Nebraska calf diarrhea virus A strain of 82-day-old male C3H/HeN mouse. calf rotavirus. Associated with gastroenteritis in calves. Has been propa- NCTC clone 2555 cells (CCL 12) Derived gated in bovine embryonic kidney cell from NCTC strain 2049 (subline III) of cultures, where it is cell-associated. the parent NCTC clone 1328. The par- Agglutinates human O erythrocytes. ent line was derived from a culture of subcutaneous areolar and adipose Fauvel M et al (1978) Intervirology 9 , 95 connective tissue taken from a normal 82-day-old male C3H/HeN mouse. necrosis Death of cells in a circumscribed piece of tissue. This may be directly due NCTC clone 3526 cells (CCL 7.2) A heter- to virus infection or may occur in the oploid cell line derived from LLC-MK2 surrounding uninfected cells following cells and adapted to grow in a chemi- the release of toxic materials from the cally defined medium. infected cell. See also apoptosis .

NCTC clone 929 L cells (CCL 1) L strain necrotic rhinitis virus Synonym for is a heteroploid cell line derived from Bovine herpesvirus 1 . Neethling virus 328

Neethling virus Synonym for Lumpy skin Negri bodies Intracytoplasmic, acido- disease virus . philic, diagnostic, inclusion bodies seen in the neuronal cells of the cerebellum, Nef protein A protein (ne gative f actor) hippocampus, thalamus, hypothalamus, induced in cells infected by human and and brainstem of animals or humans simian immunodeficiency viruses. Its who have died of rabies. Rarely found exact function during replication has not in the cortex. been defined, but it plays an important Miyamoto K and Matsumoto S (1965) J Cell stimulatory role in the replication and Biol 27 , 677 pathogenesis of HIV, including modulation of cell activation and apoptosis, Nelfinavir A non-peptidic inhibitor of change in intracellular trafficking of cel- the HIV protease which interferes with lular proteins, such as CD4 and MHC the processing of viral polyprotein pre- class I, and increase of virus infectivity. cursors to yield non-infectious prog- Aiken C et al (1994) Cell 76 , 853 eny virions. Used in conjunction with Piguet V et al (2000) Nat Cell Biol 2 , 163 a reverse transcriptase inhibitor. Also known as Viracept. negative staining A method for visualizing virus particles by drying them in Nelson Bay orthoreovirus (NBV) A spe- a film of electron-dense material, such cies in the genus Orthoreovirus . Isolated as phospho-tungstic acid (PTA), which from the heart blood of a flying fox in outlines the particle and reveals surface New South Wales, Australia. Passage structures in the electron microscope. i.c. in suckling mice causes paralysis Although it has played an important and death. Causes cell fusion and CPE role in virus structural studies, it has in a line of pig kidney cells. now been replaced in many cases by Gard G and Compans RW (1970) J Virol cryo-electron microscopy, which avoids 6 , 100 dehydration, fixation, and staining artefacts. neo-antigens Antigens that only appear in cells after transformation. They are Adrian M et al (1984) Nature 308 , 32 Brenner S and Horne RW (1959) Biochim coded for by the transforming virus. Biophys Acta 34 , 103 Often called ‘ tumor antigens ’ or ‘ T antigens. ’ negative strand The RNA strand complementary to the positive (messenger- neonatal calf diarrhea coronavirus sense) strand. It forms the genome of (NCDV) A strain of Bovine coronavirus . several families of plant and animal Causes severe diarrhea, usually in RNA viruses which form an order calves less than 5 days old. Replicates termed the Mononegavirales or single- in bovine kidney cell cultures produc- stranded negative-strand viruses. ing syncytia. Some strains, such as There are also several families of seg- the highly pathogenic strain LY-138, mented negative strand viruses includ- have not been propagated in vitro and ing the Arenaviridae , Bunyaviridae and are replicated by passage in calves. The Orthomyxoviridae. strains can then be purified from the Synonym : minus strand. contents of the small intestines. Serial passage resulted in attenuation and Pringle CR (1997) Arch Virol 142 , 2321 this virus can induce resistance to viru- Roberts A and Rose JK (1998) Virology 247 , 1 lent virus. A similar disease is caused by calf rotavirus. Negishi virus (NEGV) A tick-borne virus Synonym : Bovine coronavirus . in the genus Flavivirus . Isolated from two fatal cases of human encephali- Hajer I and Storz J (1979) Arch Virol 59 , 47 tis in Japan, but one isolate was lost. Woode GN and Bridger JC (1975) Vet Rec 96 , 85 Sequence analysis of the existing isolate Zhang X et al (2007) Virology 363 , 1 indicates that it is a strain of Louping ill virus . neoplasm A localized population of pro- Okuno T et al (1961) Jpn J Med Sci Biol 14 , 51 liferating cells not governed by the 329 neuraminidase inhibitors

usual limitations of normal growth. viruses such as rabies, herpes sim- May be benign or become malignant if plex, and pseudorabies viruses, which it metastasizes. do not generally spread by viremia. Other viruses such as polio, reovirus neopterin A catabolite of GTP (2-amino-4- and mouse hepatitis may utilize both hydroxy-6-(D-erythro-1,2,3-trihydroxy- viremia and neural spread to dissemi- propyl)-pteridine). Increased amounts nate infection. of neopterin are produced by human monocytes/macrophages upon stimulation with the cytokine interferon-g. neuraminidase (NA) An enzyme present Only detected in humans and non- on the surface of influenza and para- human primates. Increased concentra- influenza viruses which splits N -acetyl- tions are found in serum and urine of neuraminic acid from glycoprotein patients infected with HIV, hepatitis B, viral receptors on the cell membrane. hepatitis C, influenza, measles, mumps, The crystal structure shows that NA rubella, and varicella viruses, and is a tetrameric molecule with each subunit containing six sheets of four appears to correlate with the severity β of the disease. It has also been reported -strands each, arranged like the blades to be elevated in various malignant of a propeller. The enzymatic site is a diseases. hydrophobic depression in the center of the β -sheets. Neuraminidase activ- Nephropathia epidemica virus Synonym ity is carried by one of the two types of for Hantaan virus . glycoprotein peplomer on the surface of members of the Orthomyxoviridae . Lee HW et al (1979) Lancet i , 186 The second type is a hemagglutinin or attachment site by which the virus is Nepuyo virus (NEPV) A serotype joined to the cell receptor site. This can Marituba virus of in the genus be destroyed by the neuraminidase, Orthobunyavirus , belonging to the releasing the virus. In most members group C viruses. Isolated from bats of the Paramyxoviridae , both neuramini- and mosquitoes in Trinidad, Honduras, dase and hemagglutinin activity are car- Mexico, Panama, and Belem, Brazil. ried on the same peplomer. However, Not associated with disease in humans. morbilliviruses such as measles lack neuraminidase activity. A new class of nerve growth factor A factor present in anti-influenza virus inhibitors which peripheral neurons required for the target the active enzymatic site based establishment of latent infection by upon the crystal atomic structure of the herpes simplex virus. neuraminidase has been developed and Wilcox CL et al (1990) J Neurosci 10 , 1268 licensed for use (see neuraminidase inhibitors) . Netivot virus (NETV) A serotype of Umatilla virus in the genus Orbivirus . Tisdale M (2000) Rev Med Virol 10 , 45 Recovered from a pool of Culex pipiens von Itzstein M et al (1993) Nature 363 , 418 captured in Israel. Tesh RB et al (1986) Am J Trop Med Hyg neuraminidase inhibitors A number of 35 , 418 neuraminidase inhibitors have now been developed for use against influ- Neudoerfl virus (NEUV) The proto- enza virus infection, and are being type strain of Tickborne encephalitis stockpiled for possible use in the event virus (European subtype) in the genus of a future influenza virus pandemic. Flavivirus . Isolated in Neudoerfl an der These include Oseltamivir (Tamiflu), Leitha, Austria. and Zanamivir (Relenza). Tamiflu is an ethyl ester that is orally bioavailable, Wallner G et al (1996) J Gen Virol 77 , 1035 whereas Relenza must be given intranasally or by inhalation. neural spread Dissemination of virus infection by spreading along periph- Oxford JS et al (2002) Antivir Chem Chemother eral nerves. Plays an essential role for 13 , 205 neuro-2a cells 330 neuro-2a cells (CCL 131) A cell line estab- combination of virus with specific anti- lished from a spontaneous tumor of a body. On the surface of the virus parti- strain A albino mouse. cle there may be several antigens, and antibodies to some of these can com- Neurocytes hydrophobiae Name given to bine without causing loss of infectivity. a supposed protozoan etiological agent However, such an infective antigen– of rabies. The structure is now known antibody complex may be neutralized as the ‘ Negri body. ’ if either complement or an antibody Calkins GN (1909) Protozoology . New York: against the first antibody binds to the Lea and Febiger complex. In such a case the neutralization may be due to virolysis of an envel- neurodegeneration Breakdown of neu- oped virus, or to steric hindrance due ronal structure and function, the end to the build-up of protein molecules. stage of prion disease. This type of neutralization is described as ‘ extrinsic ’ in contrast to ‘ intrinsic ’ neurolymphomatosis of fowls virus neutralization in which there is a direct Synonym for Gallid herpesvirus 2 . inactivation reaction between antibody and a vital site on the virus. Virus par- neuronal cell adhesion molecule ticles may be agglutinated without los- (NCAM) A probable receptor for rabies ing infectivity, but this will result in a virus on neuronal cells. fall in the titer of infectivity units. This process is described as ‘ pseudoneutrali- neuronal vacuoles Characteristic patho- zation.’ The initially formed virus–anti- logical lesions found in the brain in body union is dissociable, but with time patients with variant Creutzfeldt– the binding becomes firmer, although Jakob disease. The distribution of these some dissociation of a small amount of lesions within the brain is used as a infective virus can still occur. See also surrogate marker for possible strain non-neutralizable fraction and neu- differences in the prions. tralization test . neurotransmitters Small molecules lib- Cleveland SM et al (2000) Virology 266 , 66 erated at nerve endings that diffuse to Dimmock NJ (1995) Rev Med Virol 5 , 165 neighboring cells and trigger a specific response. Examples are acetylcholine, neutralization test Used to measure infec- epinephrine, norepinephrine, dopamine tivity neutralizing capacity, usually of and gamma-aminobutyrate (GABA). specific antibody. Virus and antiserum are mixed, and surviving infectivity, if neurotropism The ability of a virus any, is tested for by inoculation of ani- to infect specific cell populations of mals, eggs, or cell cultures. The amount the central nervous system. Families of virus or antibody is kept constant, of viruses that exhibit neuro- but the other component is varied to tropism include the Coronaviridae , obtain a titer of neutralizing activity Flaviviridae , Lentiviridae , Herpesviridae , which will be expressed as the serum Paramyxoviridae , Picornaviridae and dilution required to neutralize a certain Rhabdoviridae . number of infective doses of virus, or as the number of infective doses of virus neurovaccinia virus A strain of Vaccinia neutralized by a certain amount of anti- virus , in the genus Orthopoxvirus , more serum. A virus control with no antiserum virulent than other vaccinia viruses. is required, and the test is read when the Produces flat, ulcerated pocks on the control becomes positive because the CAM, with a tendency to hemorrhage. virus may later break through the neu- neurovirulence The capacity of a virus to tralizing action of the antibody, and the multiply and to extend infection after it apparent titer of the antiserum will fall. has invaded the nervous system. See also non-neutralizable fraction . neutralization Usually understood to neutralizing antibody An antibody mean neutralization of infectivity by which inhibits the infectivity of a virus. 331 Newcastle disease virus (NDV) neutral red A photoreactive dye. See high cleavability correlated with high photodynamic inactivation . virulence. Global panzootics occurred first in 1926 (originating in Java then neutroseron A name proposed for any spreading to Newcastle-upon-Tyne, UK) group of viruses which cross-react in then in 1973 and 1979 (both originating neutralization tests. No longer in use. in the Middle East). Highly pathogenic (velogenic) strains caused epidemics Nevirapine A non-competitive nucleo- among double-crested cormorants over side inhibitor of reverse transcriptase a wide area of central North America in that interacts with tyrosine residues 1990 and 1992, and in Saskatchewan, on the enzyme. In clinical trials against Canada in 1995. Large outbreaks HIV, resistant mutants rapidly devel- occurred in New South Wales, Australia oped, but it was effective when used in during 1998–2000. Disease produced is combination with AZT. Recommended primarily respiratory but signs of nerv- as a single dose to prevent perinatal ous system involvement may be seen. transmission of HIV. Has caused severe The eyes are closed, there is nasal dis- liver damage and other adverse events charge and watery diarrhea. Virus is in some healthcare workers taking high shed in all secretions and excretions for doses prophylactically. up to 4 weeks. Spasms and paralysis Synonyms : BI-RG-587; Viramune. may occur. Mild strains cause low mortality but reduce egg production. Most Anon (2001) MMWR 49 , 1153 avian species appear to be susceptible. Merluzzi VJ et al (1990) Science 250 , 1411 Infection of the conjunctiva has been reported in poultry and laboratory newborn pneumonitis virus Synonym workers. Experimental i.c. injection in Human parainfluenza virus type 1 for . hamsters and mice causes encephalitis which is not transmissible. Transmission Newbury agent-1 virus (NA-1V) A strain is through drinking water or inhala- of bovine enteric calicivirus, first iden- tion of dust. Control is by slaughter or tified in 1976. Recently, determination use of attenuated vaccines that can be of the complete sequence of the virus administered by aerosol or in drinking suggests that it is a member of a new water. Most strains replicate readily in genus, since it clearly does not belong eggs or chick cell cultures in which they to any of the present four calicivirus produce CPE. Replication also occurs genera. in many species of mammalian cells. Dastjerdi AM et al (1999) Virology 254 , 1 Cultures may remain latently infected Oliver SL et al (2006) Virology 350 , 240 for long periods. All strains agglutinate Woode GN and Bridger JC (1978) J Med fowl erythrocytes; some agglutinate a Microbiol 11 , 441 variety of avian and mammalian cells. Human erythrocytes treated with some Newbury agent-2 virus A strain of bovine strains are agglutinated by serum from enteric calicivirus which appears infectious mononucleosis patients. The to be genetically related to human virus has been shown to target tumors noroviruses. and to be oncolytic, killing cells by caspase-dependent apoptosis and late Newcastle disease virus (NDV) A spe- TRAIL-induced apoptosis. The external cies in the genus Rubulavirus . A highly spike glycoprotein on the virion con- contagious natural infection of fowls, tains both hemagglutinin and neurami- turkeys and other species of birds. nidase activities. Strains have antigenic differences and Synonyms : atypischen geflugelpest vary in virulence. The strains of low virus; avian pneumoencephalitis virus; pathogenicity are termed ‘ lentogenic, ’ ranikhet disease virus; fowlpest virus; those of medium pathogenicity ‘ mes- avian paramyxovirus 1. ogenic,’ and highly pathogenic strains ‘ velogenic. ’ These differences are Elankumaran S et al (2006) J Virol 80 , 7522 mostly determined by the sequence at Ferreira L et al (2004) J Gen Virol 85 , 1981 the cleavage site of the Fo protein, with Gould AR et al (2001) Virus Res 77 , 51 New Jersey virus 332

New Jersey virus See Vesicular stomati- Table N1 . New World arenaviruses tis New Jersey virus . Amapari Paraná New Minto virus (NMV) An unassigned Flexal Pichinde species in the family Rhabdoviridae , Guanarito * Pirital belonging to the Sawgrass group. Junín * Sabiá * Isolated from the tick, Haemaphysalis Latino Tacaribe leporispalustris , removed from snow- Machupo * Tamiami shoe hares, Lepus americanus, in central Oliveros Whitewater Arroyo * Alaska. Sensitive to sodium deoxycho- * Denotes highly pathogenic hemorrhagic late. Kills suckling mice on i.c. but not fever-inducing viruses. on i.p. injection. Weaned mice do not die after i.c. injection. Produces plaques in Vero cells but not in Pekin duck embryo cells. Antigenically related to Table N2. New World hantaviruses sawgrass virus and Connecticut virus. Anajatuba Juquitiba * Ritter DG et al (1978) Can J Microbiol 24 , 422 Andes * Laguna Negra Araraquara * Lechiguanas * newt viruses T6 to T20 Various viruses Bayou * Limestone Canyon have been isolated from newts which Bermejo Maciel may be strains of frog virus 3, but they Black Creek Canal * Monongahela * require further characterization. Bloodland Lake Muleshoe Blue River New York * new variant Creutzfeldt–Jakob disease Calabazo Oran * (nvCJD) A new disease announced by Caño Delgadito Pergamino an expert advisory committee to the Castelo dos Sonhos * Prospect Hill UK government on 20 March, 1996 Central Plata Rio Mamore based on the recognition of 10 per- Choclo * Rio Mearim sons with onset of a new form of CJD El Moro Canyon Rio Segundo with characteristic pathological brain Hu39694 * Seoul lesions. Molecular analysis showed Isla Vista Sin Nombre * that nvCJD is caused by a prion indis- * Denotes highly pathogenic hantavirus tinguishable from the causative agent pulmonary syndrome-inducing viruses. of bovine spongiform encephalopathy. By July 2007 the number of definite or probable cases of nvCJD was 165. As there is no treatment all are fatal, with pulmonary syndrome, a disease with the majority in young persons below about 40% mortality ( Table N2 ). the age of 30. There is evidence of a decline in the numbers of new cases New York virus (NYV) A species in the following a peak in 2000. The cases genus Hantavirus . Identified as the are widely distributed geographi- cause of fatal hantavirus pulmonary cally with no obvious common risk so syndrome in a Rhode Island patient far known, apart from one cluster in probably infected in Shelter Island, Queniborough, Leicestershire, UK. See New York. Genetically and antigeni- prion diseases . cally distinct from Sin Nombre virus . The rodent host is Peromyscus leucopus Will RG et al (1996) Lancet 347 , 921 (eastern haplotype). New World arenaviruses South American Hjelle B et al (1995) J Virol 69 , 8137 arenaviruses belonging to the Tacaribe complex ( Table N1 ). Nexo-Cendo configuration A trans-membrane protein with the N-terminus New World hantaviruses North and external to the membrane and the South American viruses belonging C-terminus endocytoplasmic. to the Hantavirus genus. Many are den Boon JA et al (1991) Virology 182 , 655 highly pathogenic, causing hantavirus Escors D et al (2001) J Virol 75 , 1312 333 Nile crocodilepox virus (CRV)

NF-1 protein Nuclear factor 1. A cellu- nicotinic acetylcholine receptor A host lar transcription factor that binds to molecule that may be a receptor for sites on the human polyomavirus (BK rabies virus. It is found on muscle cells and JC) genomes and enhances their but not on neurons, so if it is involved expression. it may enable the virus to multiply locally at the site of the bite in myo- NF-κ B A pleiotropic transcription factor tubes, which would facilitate subse- that is active in the nucleus of mature B quent penetration into neurons. lymphocytes, differentiated monocytes and some T-cell lines. Named because Nidovirales An order comprising the it was originally found to switch on Arteriviridae Coronaviridae κ families , , and genes for the class of immunoglobu- Roniviridae which share various struc- lins in B lymphocytes. Involved in the tural and replicative features. They response of lymphoid cells to a variety have linear, non-segmented, positive- of stimuli. The oncogene of avian retic- sense single-stranded RNA genomes. uloendotheliosis virus, v-rel , specifies a κ The gene order from 5 to 3 consists of v-REL protein that is related to NF- B, a replicase gene followed by the struc- and causes acute fatal neoplasia in tural and nonstructural protein genes. infected birds. Replication involves a 3 co-terminal nested set of four or more subgenomic Ngaingan virus (NGAV) An unassigned RNAs (which give the name Nido from vertebrate rhabdovirus. Isolated from Latin nidus meaning nest, to the Order). Culicoides sp in Queensland, Australia. The genomic RNA is the mRNA for Antibodies present in wallabies, kanga- translation of the replicase. There is a roos, and cattle. Not reported to cause virion envelope with an integral mem- disease in humans. brane protein that spans the membrane at least three times. There are two gen- Ngari virus (NRIV) A serotype of era, Coronavirus and Torovirus , in the Bunyamwera virus in the genus family Coronaviridae, one, Arterivirus , Orthobunyavirus. Isolated from male in the family Arteriviridae and one, mosquitoes, Aedes simpsoni, in Senegal. Okavirus in the family Roniviridae . Also found in Burkina Faso, Central African Republic, and Madagascar. Cavanagh D (1997) Arch Virol 142 , 629 Isolated from other Aedes sp and also Anopheles sp. Not reported to cause dis- Nigerian horse virus A virus isolated ease in humans. from the brain of a horse with sporadic Institut Pasteur, Dakar (1984) Annual Report meningo-encephalomyelitis (staggers) by i.c. inoculation of suckling mice. Ngoupe virus (NGOV) A serotype of Probably the cause of staggers in horses Eubenangee virus in the genus Orbivirus . in Nigeria; may be a strain of Borna Isolated from Aedes tarsalis in the disease virus . Not known to cause dis- Central African Republic. ease in humans. Institut Pasteur, Dakar (1975) Annual Report Porterfield JS et al (1958) Br Vet J 114 , 425 nick A break in a single strand of nucleic Nigg’s virus Not a virus. Synonym for acid, especially one strand of a double- mouse pneumonitis agent. A chlamy- stranded nucleic acid. dia of subgroup A. nickase An enzyme that introduces sin- Niigata virus A virus isolated from an gle-strand breaks into double-stranded outbreak of gastroenteritis in Japan. DNA. Could be passed in humans. Nicolau bodies Intranuclear or intracyto- Kojima S et al (1948) Jpn Med J 1 , 467 plasmic inclusion bodies found in cells infected with human herpesvirus 1, 2, Nile crocodilepox virus (CRV) An unas- or 3. signed virus in the family Poxviridae . Nipah virus 334

Nipah virus A species in the genus In 2001 an outbreak of infection Henipavirus. Isolated in March 1999 occurred in Siliguri, West Bengal involv- from the brain of a patient who died ing 9 human cases and in nearby in Sg Nipah village, Bukit Pelandok, Bangladesh outbreaks occurred in 2001 Malaysia during a large outbreak of and 2003. These outbreaks were caused human encephalitis with 105 fatali- by a virus slightly different from the ties amongst 265 cases. Because of an Malaysian isolate. The genome of Nipah associated outbreak of disease in pigs, virus is a negative-sense single-stranded the virus was initially thought to be RNA molecule 18,246 kb in length. The Japanese encephalitis virus, but was nucleotide sequence homology com- found to cause syncytia in Vero cells pared to Hendra virus ranges from 70% and to show positive fluorescence in an to 88% in the gene open reading frames’s IFA test using antiserum made against which are arranged 5-N-P-C-VM-F-G-L- Hendra virus. Subsequent molecular 3, and the intergenic regions are identical characterization confirmed that Hendra in the two viruses. Although Nipah and virus and Nipah virus are closely related, Hendra viruses are closely related, they though distinct, species that represent are clearly distinct from any of the estab- a new genus of the Paramyxoviridae . lished genera within the Paramyxoviridae . Nipah virus first emerged in domestic pigs in 1997 when a respiratory dis- Bellini WJ et al (2005) J Neurovirol 11 , 481 Chua KB et al (2000) Science 288 , 1432 ease was noted by farmers. The dis- Eaton BT et al (2006) Nat Rev Microbiol 4 , 23 ease spread amongst pigs during 1998, Harit AK et al (2006) Indian J Med Res becoming epizootic in September when 123 , 553 the first human cases of encephalitis Hsu VP et al (2004) Emerg Infect Dis 10 , 2082 were noted. 11 confirmed cases also occurred in abattoir workers handling Nique virus (NIQV) A strain of Candiru Malaysian pigs in Singapore, with virus in the genus Phlebovirus , member one fatality. The disease in pigs has of the sandfly fever serogroup, Candiru a relatively low mortality rate ( 5%) complex. Isolated from flies Lutzomyia compared to the disease in humans panamensis in Panama. Not reported to ( 40%). Pigs develop rapid, labored cause disease in humans. breathing and an explosive non-productive cough as the main symptoms, nitric oxide synthase but some neurological changes can be An inducible enzyme observed including lethargy or aggres- activity which has been reported to be sive behavior. In humans, who appear associated with brain infection by Borna to contract the disease only from direct disease virus and Rabies virus. contact with pigs or pig excretions (no Koprowski H et al (1993) Proc Natl Acad Sci transmission by human–human con- 90 , 3024 tact has been demonstrated), there is a febrile encephalitis which rapidly nitrocellulose A nitrated derivative of progresses to multisystem involvement cellulose which is used either as a with vasculitis and syncytial giant cell powder or is made into membrane fil- formation at various sites. Spread to ters of defined porosity. Used to bind the brain is by the vascular route and nucleic acids in Northern blotting and leads to diffuse small foci of necrosis Southern blotting procedures and pro- and neuronal degeneration. The disease teins in Western blotting. epidemic was stopped in May 1999 by the slaughter of more than a million Nkolbisson virus (NKOV) An unas- pigs. The reservoir of natural infection signed virus in the family Rhabdoviridae , seems to be the large fruit-eating bat, belonging to the Kern Canyon sero- Pteropus hypomelanus , but other species group. Isolated from mosquitoes near of Megachiroptera may also be involved, Yaounde, Cameroon. Not reported to as is the case with Hendra virus in cause disease in humans. Queensland, Australia. The reason for the sudden appearance in farmed pigs NMH10 A strain of Sin Nombre virus in is unknown. the genus Hantavirus . 335 non-ionic detergent

NMR-11 virus A strain of Sin Nombre nomenclature The names of viruses have virus in the genus Hantavirus . developed in a haphazard manner over the century since they were discovered, Nodamura virus The type species of the from descriptions of the disease (e.g. genus Alphanodavirus. Isolated from Yellow fever virus) to names of individual insects, but serological data suggests virologists (e.g. Rous sarcoma virus) or that it also naturally infects pigs and the place where they were first discov- perhaps herons. ered (e.g. Marburg virus) . In recent years, emphasis on the disease or the name Nodaviridae A family of RNA viruses of the investigator has generally been with two genera: Alphanodavirus , con- dropped, and newly discovered viruses sisting of viruses isolated only from have received a place name (e.g. Nipah insects, and Betanodavirus , consisting of virus), though a prominent exception viruses isolated from juvenile marine was the virus causing hantavirus pul- fish, in which they cause nervous monary syndrome, not welcomed by necrosis disease ( Table N3 ). the local affected population and finally called Sin Nombre (no name) virus . nodule-inducing mouse mammary For the last 30 years the ICTV (and its tumor virus A strain of virus in the predecessor the ICNV) has developed genus Betaretrovirus found in mice a framework of rules that govern the freed from mouse mammary tumor officially recognized names of viruses. virus by foster nursing. These mice A stable scheme using English for virus develop mammary tumors late in life. order, family, subfamily, genus, and spe- The virus is morphologically identical cies names has evolved: order names to mouse mammary tumor virus but have the suffix virales , family names viri- induces only hyperplastic nodules with dae , subfamily names virinae , and genus low neoplastic potential. names virus . This taxonomic scheme is both easy to use in practice, and valu- Nola virus (NOLAV) A serotype of Bakau able to the discipline of virology. virus in the genus Orthobunyavirus . Van Regenmortel MHV (2005) In Topley & Isolated from Culex perfuscus mosqui- Wilson’s Microbiology and Microbial Infections , toesin the Central African Republic. vol. 1, Tenth edition, edited by BWJ Mahy Not known to cause disease in and V ter Meulen, London: Hodder Arnold, humans. p. 24 non-allelic complementation See complementation . Table N3. Fish nervous necrosis viruses non-A non-B hepatitis virus Synonym Atlantic cod nervous necrosis for Hepatitis C virus . Atlantic halibut Barfin flounder non-bacterial regional lymphadenitis Dicentrarchus labrax (Sea bass) virus Synonym for cat-scratch disease Dragon grouper virus (not a virus, a rickettsial agent, Greasy grouper Bartonella henselae) . Grouper Halibut non-genetic reactivation See reactivation Japanese flounder and complementation . Lates calcarifer (Barramundi) Malabar grouper Nonidet P40 A non-ionic detergent com- Redspotted grouper prising octylphenol ethylene conden- Seabass sate. Used to disrupt cells and viral Striped jack membranes. Tiger puffer Umbrina cirrosa (Sea crow) non-ionic detergent A detergent with no All names italicized are species in the genus net surface charge, e.g. the triton series, Betanodavirus , others are tentative species. Nonidet P40. non-neutralizable fraction 336 non-neutralizable fraction Neutralization be introduced by a nonsense muta- by antibody is often not complete, a tion, converting a sense codon into a small fraction of the original infectiv- stop codon. The nonsense mutation ity resisting neutralization. This may can arise by base substitution or frame- be due to dissociation of the virus– shifting. It has been reported that in antibody union or to the formation of some contexts the UGA codon may infective complexes. The addition of encode the amino acid selenocysteine. anti-antibody neutralizes such com- Synonym : stop codon. plexes. See neutralization . Brenner S et al (1961) Nature 190 , 576 Synonym : persistent fraction. Taylor EW (1994) J Med Chem 37 , 2637 non-nucleoside reverse transcriptase nonstructural viral proteins Proteins inhibitors (NNRTIs) Compounds coded for by the viral genome but not which inhibit reverse transcriptase by incorporated into the viral particle. directly binding to the enzyme at an They have a functional role during allosteric site that influences the cata- viral replication. lytic site. All NNRTIs studied to date rapidly give rise to a high level of resistance both in patients and in vitro , non-templated nucleotide addition The but in combination with a nucleoside modification or editing of mRNAs inhibitor such as AZT, they appear to transcribed by paramyxoviruses to be much more effective. Detailed struc- alter mRNA coding. tural analysis has shown that these inhibitors bind to tyrosine residues on normal human immunoglobulin (NHIG) the enzyme surface, and the principal A prophylactic pre-exposure treatment mutation to resistance is Y181C. See used for travelers to regions where Delavirdine, Nevirapine . hepatitis A virus is endemic. About 80% effective, but the licensure of a hepatitis non-paralytic poliomyelitis About 1–2% A vaccine has made this the preferred of persons exposed to poliovirus experi- treatment for travelers. ence an illness lasting 2–10 days similar to ‘ aseptic ’ meningitis, often accompa- Norovirus A genus in the family nied by back pain and muscle spasm. Caliciviridae, with only one species, Recovery is usually rapid and complete. Norwalk virus. There are 7 recognized strains, which form a distinct clade non-permissive cells Cells in which a within the family Caliciviridae and virus will not replicate. They may be 5 tentative species infecting bovine, permissive for one virus but not for murine, or swine hosts. Formerly called another. The fact that they are non- ‘ small round structured virus (SRSV) permissive for virus replication may particles. ’ Associated with epidemic make them very suitable for demon- gastroenteritis, their importance has strating transformation. been increasingly recognized since the application of PCR-based diagnostic non-producer (NP) cells Cells usually assays in investigations of outbreaks transformed, carrying all or part of a worldwide, and it is estimated that they viral genome but not producing infec- cause 95% of cases of non-bacterial gas- tive virus particles. In the case of Rous troenteritis in the USA. Diagnosis using sarcoma virus-transformed NP cells, antigen detection ELISAs is also possi- non-infective virus may be produced. ble, but because these are less sensitive, Such cells are called ‘ L–R cells. ’ at least 6 samples should be tested per outbreak compared with 3 samples for nonsense codons Codons which do testing by PCR. On September 18, 1998, not code for an amino acid. They are the first documented case of transmis- UAA, UAG and UGA, sometimes sion of a gastroenteritis virus between referred to as the ‘ ochre, ’ ‘ amber ’ and players on a football field occurred dur- ‘ opal ’ codons, respectively. They are ing a match between Duke and Florida chain-terminating signals which can State University teams. Duke lost 62–13, 337 nuclear factor 1 (NF1)

but Duke players, vomiting after a ORF-2 encodes the major capsid pro- box lunch, infected several of their tein, and ORF-3 encodes a small virion opponents. protein of unknown function. Norwalk and related viruses are a major cause Becker KM et al (2000) N Engl J Med 343 , 1223 of acute non-bacterial gastroenteritis. It Duizer E et al (2007) J Clin Virol 40 , 38 is presently unclear whether protective immunity develops after Norwalk virus North Clett virus (NCLV) A serotype of infection, and no specific treatment or Great Island virus in the genus Orbivirus . antiviral therapy is available. A member of the Kemerovo antigenic Synonym: epidemic gastroenteritis virus. group. Berke T et al (1997) J Med Virol 52 , 419 Green J et al (2000) Virus Genes 20 , 227 North End virus (NEDV) A serotype of Jiang X et al (1993) Virology 195 , 51 Great Island virus in the genus Orbivirus . Lambden PR et al (1993) Science 259 , 516 A member of the Kemorovo antigenic group. nosocomial infections Hospital-acquired infections, usually applied to patients, Northern blotting A procedure analo- but includes hospital personnel as well. gous to Southern blotting but involving the transfer of RNA rather than Novirhabdovirus A genus in the family DNA on to nitrocellulose or activated Rhabdoviridae comprised of fish viruses. paper sheets. Optimum virus replication temperatures are 15–28°C. The genome nega- northern pike herpesvirus Synonym for tive single-stranded RNA is 11.1 kb in esocid herpesvirus 1. length with six genes in the order 3 -N- P-M-G-NV-L5 . The virus appears to be Northway virus (NORV) A serotype transmitted horizontally by waterborne of Bunyamwera virus in the genus virus. Egg-associated transmission has Orthobunyavirus . Isolated from Aedes sp also been demonstrated. There are four in Alaska. Not known to cause disease species in the genus: Hirame rhabdovirus , in humans. Infectious hematopoietic necrosis virus (the type species), Snakehead virus , and Viral Norvir An inhibitor of HIV protease, hemorrhagic septicemia virus (Egtved licensed for human use. See ritonavir . virus).

Norwalk virus (NV) A species in the genus NP cells See non-producer (NP) cells . Norovirus, the prototype of a group of related viruses, 27 nm in diameter, with NP protein Abbreviation for nucleo - a positive-strand RNA genome, found protein. in the feces of patients with gastroenteritis. There are seven recognized strains: Ntaya virus (NTAV) A species in the the Desert Shield, Hawaii, Lordsdale, genus Flavivirus . With Bagaza , Ilheus Mexico, Norwalk, Snow Mountain, and Israel turkey meningo-encephalomyelitis , Southampton agents. The viruses can- and Tembusu viruses forms the Ntaya not be propagated in cell cultures and serogroup. Isolated from mosquitoes no experimental animals have been in Uganda, Cameroon, and Central found susceptible to the virus, although African Republic. Not known to cause chimpanzees undergo subclinical infec- disease in humans or other animals. tion and shed antigen in the feces. Swine calicivirus is also closely related nuclear export protein (NEP) A protein genetically and is a tentative species involved in influenza virus replication, in the genus. The genome of Norwalk formerly known as NS2, encoded in virus is 7.75 kb in length with three open genome segment 8. reading frames (ORFs): ORF-1 encodes a polyprotein precursor of the nonstruc- nuclear factor 1 (NF1) A host cellular tural proteins including a replicase, transcription factor which binds as a nuclear factor 1 (NF1) 338

dimer to viral and cellular promoters material of viral origin is below the during replication of adenovirus. detection threshold, the advent of the polymerase chain reaction (PCR) nuclear factor 2 (NF2) A type II cellular has greatly improved studies of viral topoisomerase required for adenovirus nucleic acids. The template amplified DNA replication. Thought to relieve can be either viral DNA or RNA. Target overwinding of the template DNA dur- RNA must first be converted to cDNA ing replication. using reverse transcriptase (RT). nucleases Enzymes which break down nuclein The first name given to an nucleic acid molecules by hydrolysis of unusual phosphorus-containing com- phosphodiester bonds. They are present pound isolated in the last century from in almost all biological systems, are the nuclei of pus cells. Now known as usually specific for RNA or DNA, and ‘ nucleoprotein. ’ are called ribonucleases or deoxyribo- nucleases, respectively – although some nucleocapsid The viral nucleic acid are nonspecific. Nucleases can attack directly enclosed by the capsid. This the polynucleotide chain in two ways: simple arrangement is usual in the at points within the chain or stepwise rod-shaped plant viruses and a few from one end of the chain. Enzymes act- isometric virions, but with most ani- ing by the first method are called ‘ endo- mal viruses the capsid encloses a more nucleases ’ and work endolytically. They complex structure, the core. produce oligonucleotides and cause a rapid change in physical properties nucleoid A term used by electron micro- such as viscosity. Enzymes acting by scopists to describe the electron-dense, the second method are called ‘ exonucle- centrally placed structure observed in ases ’ and work exolytically. They pro- certain viruses. duce mononucleotides and change the physical properties rather more slowly. nucleoprotein A complex of nucleic acid Nucleases also differ in the point at and protein. The form in which DNA which they split the phosphodiester exists in the nucleus of eukaryotic cells bond: some cleave between the 3 -OH and virus particles. and the phosphate group while others cleave between the 5 -OH and the phos- Nucleorhabdovirus A genus in the fam- phate group. Certain endonucleases are ily Rhabdoviridae comprised of viruses highly specific for particular nucleotide which multiply in the nucleus of plant sequences and are known as ‘restriction cells. endonucleases.’ See also ribonuclease . nucleoside analogs Nucleosides that nucleic acid A compound consisting of a mimic the normal structural units of chain of alternate sugar (pentose) and DNA or RNA but differ in a way that phosphate molecules with one purine prevents their incorporation into these or pyrimidine base attached to each macromolecules, and thus stops DNA sugar molecule. The basic unit is thus or RNA synthesis by a process of chain- a nucleotide molecule. There are two termination. An example is AZT. types: DNA, in which the sugar molecules are d -2-deoxyribose; and RNA, in nucleoside reverse transcriptase inhibi- which they are d -ribose. In both types tors (NRTI ) The emergence of HIV as a of nucleic acid the sugar phosphate major human pathogen in the early 1980s chain is through the 3 and 5 carbons resulted in a major effort to develop anti- in the sugar. Thus the chain has a 3 viral agents, and the first to be produced and a 5 end. was 3 -azido-3 -deoxythymidine (AZT). This drug is phosphorylated in the nucleic acid sequence-based amplifica- cell and acts as a competitive inhibitor tion (NASBA) Since in the majority of with respect to thymidine triphosphate virus infections the amount of genetic and acts as a chain terminator of DNA 339 Nyabira virus (NYAV)

synthesis. Unfortunately, the virus rap- nucleotides Phosphoric esters of nucleo- idly develops resistant mutations, and sides. Those derived from ribonucleo- the drug is not selective for the HIV sides are called ‘ ribonucleotides, ’ and reverse transcriptase, but also inhibits those from deoxyribonucleosides are a variety of cellular enzymes, so caus- ‘ deoxyribonucleotides. ’ Sometimes the ing toxicity in the patient. A number abbreviations riboside, ribotide, deoxy- of other nucleoside analogs have been riboside and deoxyribotide are incor- developed and used with variable suc- rectly used. Since the ribonucleosides cess, but none has the selectivity of the have three free hydroxyl groups on anti-herpes drug acyclovir (which is the sugar ring, there are three possible only active in virus-infected cells) and so ribonucleoside monophosphates; e.g. are associated with discomfort or toxic- adenosine 2 , 3 or 5 monophosphate. ity in the patient. The adenosine 5 monophosphate is present in muscle and was formerly nucleosides Units composed of a purine known as muscle adenylic acid, while or pyrimidine base combined to a the 3 monophosphate obtained by pentose or deoxypentose sugar. Those hydrolysis of yeast RNA was called commonly found in RNA, derived yeast adenylic acid. In the same way from pentose, are adenosine, guano- guanosine, cytidine and uridine yield sine, cytidine, and uridine. They may guanylic acids, cytidylic acids and uri- be formed on partial hydrolysis of dylic acids, respectively. The adenosine RNA. The nucleosides found in DNA, 5 monophosphate (AMP) can be fur- derived from 2-deoxyribose, are known ther phosphorylated at position 5 to as deoxyadenosine, deoxyguanosine, give 5 di- and triphosphates. Thus deoxycytidine, and deoxythymidine. AMP becomes ADP and ATP. The triphosphates are used as precursor mol- nucleosomes Repeat units (beads) of ecules for the synthesis of nucleic acids chromatin consisting of DNA folded by virus or cellular polymerases. around a histone core joined by protein-free stretches of nucleic acid. There nucleus Most viruses with a DNA are two molecules each of histones genome multiply in the cell nucleus, H2A, H2B, H3, and H4 around which the exception being poxviruses, which is wrapped 140 bp of DNA. A fifth his- carry all their replication machinery tone (H1) interacts with an additional and replicate in the cytoplasm. Most 20 bp DNA to complete two turns of RNA viruses remain in the cytoplasm, DNA around the histone core. The the exceptions being influenza virus nucleosomes are connected by linker and the hepatitis delta virus. DNA (beads on a string). Nugget virus (NUGV) A serotype of nucleotide reverse transcriptase inhibi- Great Island virus in the genus Orbivirus , tors (NTRTI) These drugs require two belonging to the Kemerovo serogroup. intracellular phosphorylation events Isolated from nymphs of Ixodes uriae for activity, but one such drug now collected at Macquarie Island, 800 miles licensed in the USA, Tenofovir, is sup- south-east of Tasmania, in tussock grass plied in monophosphorylated form, and under planks on the shore near a and seems to have a favorable resist- rookery of Royal Penguins, Eudyptes ance profile compared to NRTI’s. chrysalophus schlegeli . Antibodies are found in these birds. Not known to nucleotide phosphohydrolase An infect humans. enzyme which converts nucleotide diphosphates to monophosphates. Present in virions of Reoviridae . May Nyabira virus (NYAV) A serotype of play a role in inhibition of host cell Palyam virus in the genus Orbivirus . DNA synthesis or in formation of the Isolated from an aborted bovine fetus 5 terminal cap structure on mRNA. An in the Nyabira region of Zimbabwe. enzyme which converts nucleotide to Swanepoel R and Blackburn NK (1976) Vet nucleoside by hydrolysis. Res 99 , 360 Nyamanini virus (NYMV) 340

Nyamanini virus (NYMV) An unas- Nyando virus (NDV) A species in the signed arbovirus. Isolated from cattle genus Orthobunyavirus . With Eret-147 egrets, Bubulcus ibis, and other birds, virus forms the Nyando group. Isolated and ticks of Argas sp in South Africa, from Anopheles funestus mosquitoes in Egypt, and Nigeria. On injection, kills Kisumu, Kenya and the Central African newborn mice. Not reported to cause Republic, and from culicine mos- disease in humans. Virions are envel- quitoes in Ethiopia. Antibodies have oped and contain RNA. been found in humans in Kenya and Uganda, but the virus is not known to Kemp GE et al (1975) J Med Entomol 12 , 535 cause disease in humans. O

‘ O ’ virus (O for o ffal) A strain of Rotavirus to detect, nevertheless the disease may A in the genus Rotavirus . Isolated from be transmitted if such patients donate the waste water that had been used to blood or organs, particularly liver. wash the intestines of slaughtered cattle and sheep. Indistinguishable from Oceanside virus (OCV) A serotype simian virus SA11 and human rota- of Uukuniemi virus in the genus virus by CFT, but electrophoresis of viral Phlebovirus. Isolated from ticks, Ixodes RNA reveals differences. Replicates in uriae. Not known to cause disease in monkey kidney cell cultures produc- humans. ing eosinophilic inclusions and cell Yunker CE (1975) Med Biol 53 , 302 destruction. Malherbe HH and Strickland-Cholmley M ochre codon UAA One of the three termi- (1967) Arch Gesamte Virusforsch 22 , 235 nation codons. See nonsense codons .

Oak-Vale virus (OVRV) An unassigned ochre mutant Virus with mutation result- vertebrate Rhabdovirus , isolated from ing in a chain termination codon UAA. culicoid mosquitoes in Australia in 1981. Antigenically related to lyssa- Ockelbo virus (OCKV) A serotype of viruses. Not known to cause disease in Sindbis virus in the genus Alphavirus . humans. Isolated from mosquitoes, Culiseta sp, in northern Europe. Causes epidemic obesity and viruses It has been reported polyarthritis (Karelian or Pogosta that a chicken adenovirus isolate from fever) in humans, sometimes with per- Mumbai, India, caused fat accumu- sistence of symptoms for months or lation in chickens, and that 10 of 52 years. obese persons tested had antibodies to Synonym : Karelian fever virus. the chicken adenovirus. In a study con- et al Virology 182 ducted in the USA, human adenovirus Shirako Y (1991) , 753 serotype 36 (a serotype of human adeno- Oct-1 protein A human transcription fac- virus D) caused obesity in chickens tor that can stimulate RNA polymerase and mice. These experiments have not II and III transcription from a variety of been independently confirmed. promoters. It stimulates the replication Dhurandhar NV et al (2000) Int J Obesity of adenovirus DNA in vivo . 24 , 989 Odocoileus adenovirus 1 A strain of Obodhiang virus (OBOV) An unas- Cervine adenovirus, a tentative species signed vertebrate Rhabdovirus isolated in the genus Atadenovirus . from Mansonia uniformis mosquitoes in Sudan and Ethiopia. Antigenically Odocoileus hemionus type C virus A related to Kotonkan virus and Mokola possible species in the genus Gamma- virus , but not to ephemeral fever virus. retrovirus. An endogenous virus Bauer SP and Murphy FA (1975) Infect isolated from the Columbian black- Immun 12 , 1157 tailed deer, Odocoileus hemionus ( Dama hemionus columbianus ), found on the occult infections Infections that do not western slopes of the Rocky Mountains, show any obvious symptoms. Patients USA. The distribution of DNA infected with hepatitis B virus, e.g., sequences related to the endogenous may show no elevated liver trans- virus in various species of deer is aminases, and so the virus is difficult compatible with the closeness of the Odocoileus hemionus type C virus 342

relationship between the different spe- contains only viruses which infect cies in the family Cervidae . crustaceans. The type species is Gill- Aaronson SA et al (1976) Cell 9 , 489 associated virus .

Odrenisrou virus (ODRV) A tentative spe- Okazaki fragments Short pieces of DNA, cies in the genus Phlebovirus, member of with RNA primers attached, produced the sandfly virus group not assigned to during DNA synthesis. Subsequently, any complex. Isolated from Culex (Eum) the primers are replaced by DNA and albiventris in the Ivory Coast. the fragments are ligated together. Named after the Japanese scientist Institut Pasteur, Dakar (1982) Annual who first described them. See semi- Report , 87 conservative replication .

Office International des Epizooties Okhotskiy virus (OKHV) A serotype of An international veterinary organi- Great Island virus in the genus Orbivirus , zation based in Paris which acts as a member of the Kemerovo serogroup. World Health organization for ani- Isolated from the tick, Ixodes putus , col- mal diseases. Now called the World lected from several islands in the sea Organization for Animal Health. of Okhotsk in the north of the former Soviet Far East. Antibodies are found O’Higgins disease virus Synonym for in guillemots, Uria aalge , fulmars, Junín virus . Fulmarus glacialis , and cormorants, Phalacrocorax pelagicus . Pathogenic for Oita strains Two measles virus strains suckling mice. Not reported to cause Oita-1 and Oita-2, isolated from disease in humans. the brain of a patient with immunosuppressive measles encephalitis. Lvov DK et al (1973) Arch Gesamte Virusforsch 41 , 160 Ohuchi M et al (1987) J Infect Dis 156 , 436 Okola virus (OKOV) A tentative species Oita virus (OITAV) An unassigned animal in the family Bunyaviridae . Related to rhabdovirus in the genus Rhabdoviridae . Tanga virus. Isolated from the mosquito, Eretmapodites chrysogaster , in Oka vaccine virus An attenuated Cameroon. Not reported to cause dis- Japanese strain of varicella-zoster virus, ease in humans. the basis of a licensed vaccine against chickenpox (Varivax®), which appears Old World arenaviruses Members of the to provide long-term cell-mediated and LCM–Lassa complex of arenaviruses humoral immunity, although break- ( Table O1 ). through infections after exposure to wild-type VZV are occasionally seen Old World hantaviruses Members of the in vaccinees. The vaccine has also been genus Hantavirus found in Old World shown to cause herpes zoster in up to rodents ( Table O2 ). 6% of immunocompromised vaccinees, such as children with leukemia. After Olifantsvlei virus (OLIV) A species in more than 20 years experience with the the genus Orthobunyavirus. With Bobia, vaccine in Japan, it was licensed and Dabakala, and Oubi virus forms the recommended for use in the USA in Olifantsvlei serogroup. Isolated in 1995. The genome DNAs of Oka vac- suckling mice from a pool of female cine and wild-type virus can be readily mosquitoes, Culex pipiens , trapped distinguished by a PCR-based test. at Olifantsvlei, Johannesburg, South Loparev VN et al (2000) J Clin Microbiol 38 , Africa. Also isolated from Mansonia 3156 uniformis in the Sudan and from Culex Vassilev V et al (2005) J Clin Microbiol 43 , poicilipes in Ethiopia. Not reported to 5415 cause disease in humans.

- Okavirus A genus of the family 2؅ ,5 ؅ -oligoadenylate synthetase (2؅ 5 ؅ Roniviridae (Order Nidovirales ) which OAS) An enzyme involved in the 343 Omo virus (OMOV)

Table O1. Old World arenaviruses Arenavirus Disease Host Location Ippy Arvicanthis sp Central African Republic Lassa Hemorrhagic fever Mastomys sp West Africa Lymphocytic Choriomeningitis Choriomeningitis Mus musculus Europe and the Americas Mobala Praomys sp Central African Republic Mopeia Mastomys natalensis Mozambique and Zimbabwe

Table O2 . Old World hantaviruses Hantavirus Disease Host Location Hantaan HFRS Apodemus agrarius Asia, Far East Russia Dobrava -Belgrade HFRS Apodemus flavicollis Balkans, Europe Saaremaa HFRS Apodemus agrarius Europe Seoul HFRS Rattus norvegicus Worldwide Thailand HFRS Bandicota indica Thailand Puumala Clethrionomys glareolus Europe Kamiiso Clethrionomys rufocanus Hokkaido, Far East Russia Topografov Lemmus sibiricus Arctic Russia Khabarovsk Microtus fortis Far East Russia Tula Microtus arvalis Europe

early stages of interferon induction. oligopeptide A short-chain linear peptide Upon activation by double-stranded containing from 2 to 10 amino acids RNA, 2 5 -OAS polymerizes ATP into joined by peptide bonds. pppA(2 p5 A)n(2’5’A). 2 5 A then activates a cellular endonuclease, 2 5 - Oliveros virus (OLI) A serologically dis- OAS-dependent RNase L, which then tinct species of the Tacaribe complex degrades both cellular and viral RNAs of the family Arenaviridae, isolated in at UU or AU nucleotides. 1990 in Oliveros, a village in Argentina from a trapped rodent, Bolomys obscu- Samuel CE (1991) Virology 183 , 1 rus. Phylogenetic analysis confirmed oligoastrocytoma A brain tumor caused that the virus is a distinct member of by JC virus infection even in immuno- the Tacaribe complex of New World competent patients. arenaviruses, most closely related to Latino virus. Role in human disease oligodendroglioma A brain tumor unknown. caused by JC virus in a patient with Bowen MD et al (1996) Virology 217 , 362 concomitant chronic lymphatic Mills JN et al (1996) Am J Trop Med Hyg 54 , leukemia and progressive multifocal 399 leukoencephalopathy. OLV virus Synonym for H-3 virus. oligonucleotide mapping A technique for RNA characterization based upon Omikron papillomavirus A genus in the the use especially of T1 ribonuclease family Papillomaviridae , containing to obtain RNA fragments separable papillomaviruses isolated from geni- by chromatography. Now replaced by tal warts in cetaceans. Type species is more specific sequence analysis. Phocoena spinipinnis (Burmeister’s porpoise) papillomavirus. oligonucleotides Short polynucleotides containing about 2–10 nt joined by Omo virus (OMOV) A serotype of Qalyub phosphodiester bonds. virus in the genus Nairovirus . Isolated Omo virus (OMOV) 344

from the rodent Mastomys erythroleucus seen at the genetic level. In describing in Ethiopia. oncogenes, the gene is italicized (e.g. jun ) and the protein product capital- Rodhain F et al (1985) Ann Inst Pasteur Virol 136E , 243 ized (e.g. JUN protein). The viral oncogene is written v- jun and the cellular gene counterpart c-jun . Omsk hemorrhagic fever virus (OHFV) A tick-borne species in the genus Flavivirus , Cooper GM (1995) Oncogenes , Second edi- member of the Mammalian tick-borne tion. Boston: Jones and Bartlett encephalitis serogroup. Vectors are ticks, Hesketh R (1994) The Oncogene Handbook . London: Academic Press Dermacentor pictus and D marginatus , Rasheed S (1995) In The Retroviridae, vol. 4, in which transovarian transmission is edited by JA Levy. New York: Plenum Press, reported. Man may also be infected by p. 293 direct contact with infected muskrats. Causes a biphasic illness in humans oncogenic Tumor-inducing. with fever, enlargement of lymph nodes, gastrointestinal symptoms, and hemor- oncogenic RNA virus Old name for rhages from nose, stomach, and uterus, group of RNA tumor viruses, now but little or no CNS involvement. included in the family Retroviridae . Mortality 1–2%. Disease occurs in the former central USSR. Causes fever in rhesus monkeys injected i.p. On first iso- oncolysis The lysis of transformed or lation does not infect adult mice. tumor cells. There is evidence that some viruses have this property. Transformed Netzky GI (1967) Jpn J Med Sci Biol (Suppl) cells are more permissive for reovirus 20 , 141 infection than untransformed cells, and kills transformed cells in animals through Omsk virus See Omsk hemorrhagic fever an activated Ras signaling pathway. The virus . avian paramyxovirus, Newcastle disease virus is also being investigated as a pos- Onchorhynchus masou herpesvirus sible oncolytic agent. Synonym for salmonid herpesvirus 2. Coffey MC et al (1998) Science 282 , 1332 Sinkovics JG and Horvath JC (2000) J Clin Oncorhynchus masou virus Synonym for Virol 16 , 1 salmonid herpesvirus 2. oncoproteins Proteins produced from oncogene A gene which encodes a protein oncogenes: tumor-inducing proteins. whose expression leads to cell transformation. Originally discovered during oncornaviruses Old name for the RNA studies of Rous sarcoma virus in the tumor viruses. No longer in use. 1970s when the src gene was isolated and shown to have a cell homolog. oncotropism An affinity shown by Since then, more than 100 viral onco- some viruses for replication in tumor genes and proto-oncogenes (cellular cells. Parvoviruses, e.g., need to grow genes that may become disregulated in dividing cells, and some, such as to cause cell transformation) have been Kilham rat virus have been shown to described. In general, viral oncogenes suppress leukemia induction in rats by are transduced genetic sequences Moloney murine leukemia virus. found in the genome of acutely transforming viruses which have cellular Oncovirinae An old name for a subfamily homologs (proto-oncogenes) from of oncogenic retroviruses. No longer which they were derived. Evolutionary in use. changes in the virus genome have led in some cases to considerable differ- one-hit kinetics In systems where ences between the products of the viral one particle can initiate infection, oncogenes and their cellular homologs, the number of plaques appearing is but their relationship can be clearly directly proportional to the first power 345 Orbivirus

of the concentration of the inoculum. opossum viruses A and B Not viruses. If this concentration is doubled, the Chlamydiae. number of plaques will be doubled. opportunistic infections Infections one-step growth curve An experiment in which become established due to which all the cells in a particular cul- immunosuppression of the patient, e.g. ture are infected simultaneously, so that as a result of chemotherapy or AIDS. events in individual cells can be inferred Glaser CA et al (1994) Clin Infect Dis 18 , 14 from events in the whole population. opportunistic pathogens A variety of Ontario encephalomyelitis virus infectious pathogens, including many Synonym for Porcine hemagglutinating viruses, cause opportunistic infec- encephalomyelitis virus . tions in immunosuppressed persons. Amongst the viruses, those that are O’Nyong-Nyong virus (ONNV) A species normally latent, such as cytomegalo- in the genus Alphavirus. The complete virus, herpes simplex virus, JC polyoma nucleotide sequence shows a close virus, and varicella-zoster virus, cause genetic relationship to Semliki Forest most common problems, especially in virus. In humans it causes a febrile ill- AIDS patients. ness with lymphadenitis, severe joint pains and rash. Epidemic spread occurs OPV Oral poliovirus vaccine, originally with anopheline mosquitoes as vector. developed by Albert Sabin. Occurs in Uganda, Kenya, Tanzania, Malawi, and Senegal. Pathogenic for suckling mice given i.c.; older mice are oral poliomyelitis vaccine The attenu- resistant. Infant mice which survive the ated vaccine against poliomyelitis infection are stunted and show patchy developed by Albert Sabin, which can alopecia. Virus is propagated in chick be administered to infants on a sugar embryo fibroblast cell cultures. lump, and has been the cornerstone of the polio eradication campaign. Chanas AC et al (1979) Arch Virol 59 , 231 Levinson R et al (1990) Virology 175 , 110 oral vaccination Vaccination by mouth, oophoritis Ovarian pain and pelvic rather than by injection. tenderness may occur in up to 7% of females infected with mumps virus, Oran virus A strain of Andes virus in but the symptoms are not as severe as the genus Hantavirus isolated from orchitis in males. Oligoryzomys longicaudatus in Argentina. Caused fatal hantavirus pulmonary opal codon UGA One of the three termi- syndrome in humans. nation codons. See nonsense codons . Calderon G et al (1999) Emerg Infect Dis 5 , 792 opal mutant Virus with mutation resulting in a chain termination codon UGA. orangutan herpesvirus Synonym for Pongine herpesvirus 2 . open circular DNA Circular DNA in which one or both strands are not cov- Orbivirus (Latin: orbis a ring) A genus alently closed. of the family Reoviridae containing 21 recognized species and 11 tentative open reading frame (ORF) A set of species. All multiply in insects and sev- codons for amino acids uninterrupted eral also in vertebrates. Virion 80 nm in by stop codons. Usually encodes one diameter, with a double protein shell: or more proteins. the outer one without readily definable capsomeres, the inner with 32 seemingly opossum adenovirus See Possum adeno- ring-shaped capsomeres arranged with virus . icosahedral symmetry and visible in Orbivirus 346

the presence of the outer shell. Only (20) Warrego virus (3 serotypes) slightly sensitive to lipid solvents, (Culicoides , anopheline and culi- inactivated at pH 3. The double- cine mosquitoes) stranded RNA genome consists of 10 (21) Wongorr virus (8 serotypes) segments between 822 and 3954 bp, (Culicoides , mosquitoes) total 19.2 kb. There are seven virion In addition there are 13 tentative spe- and three nonstructural proteins. The cies in the genus awaiting further single-shelled nucleocapsid having characterization. lost the outer shell has transcriptase Roy P (2008) in BWJ Mahy and MHV van activity with an optimum temperature Regenmortel (eds) Encyclopedia of Virology , of 28°C. Replicate with CPE in BHK21 Third edition, Oxford: Academic Press, cells. Kill newborn mice but not adults vol. 1, p. 328 on i.c. injection. Do not kill mice of any Roy P and Gorman B (Editors) (1990) Curr age on i.p. injection. The type species Top Microbiol Immunol 162 , 200pp is Bluetongue virus. They have no common antigen but the 153 known viruses orcinol reaction A colorimetric assay for are divided into 19 species on the basis the presence of carbohydrates, espe- of antigenic cross-reactions. The species cially pentose sugars. Used to deter- and their principal vectors are: mine RNA concentrations, e.g. the (1) African horse sickness virus (9 sero- amount of RNA in a virus preparation. types) ( Culicoides ) See diphenylamine reaction . (2) Bluetongue virus (24 serotypes) Lin RIS and Schjeide OA (1969) Anal Biochem (Culicoides ) 27 , 473 (3) Changuinola virus (12 serotypes) (phlebotomines or culicine Oregon sockeye disease virus Synonym mosquitoes) for Infectious hematopoietic necrosis virus (4) Chenuda virus (7 serotypes) (ticks) in the genus Rhabdovirus . (5) Chobar Gorge virus (2 serotypes) (ticks) (6) Corriparta virus (6 serotypes) (culi- ORF Abbreviation for o pen reading frame. cine mosquitoes) (7) Epizootic hemorrhagic disease virus Orf subgroup viruses Synonym for (10 serotypes) (Culicoides ) parapoxvirus. (8) Equine encephalosis virus (7 serotypes) ( Culicoides ) Orf virus (ORFV) The type species of the (9) Eubenangee virus (4 serotypes) genus Parapoxvirus. Virions are ovoid, (Culicoides , anopheline and culi- 220–300 140–170 nm, with a charac- cine mosquitoes) teristic surface structure that appears (10) Great Island virus (36 serotypes) in the electron microscope as a spiral (ticks) coil or ‘ ball of yarn.’ DNA genome ( 1 1 ) Ieri virus (3 serotypes) (mosquitoes) 130–150 kb, G C is 64%. The nucleo- (12) Lebombo virus (1 serotype) (culi- tide sequence shows that the virus has cine mosquitoes) 132 genes, 88 of which are present in (13) Orungo virus (4 serotypes) (culi- all chordopoxviruses. A real-time PCR cine mosquitoes) method has been developed for detec- (14) Palyam virus (13 serotypes) tion of the virus in clinical specimens. (Culicoides , culicine mosquitoes) Virus is inactivated by chloroform, (15) Peruvian horse sickness virus (mos- but ether may not inactivate. Causes quitoes, horses) disease predominantly in lambs and (16) St Croix River virus (ticks) kids. There are vesicles on the lips and (17) Umatilla virus (4 serotypes) (culi- nose, progressing to pustules, ulcers cine mosquitoes) and warty scabs. Malignant aphtha (18) Wad Medani virus (2 serotypes) is a severe form of the disease which (ticks) may be fatal. Papilloma of chamois (19) Wallal virus (3 serotypes) is caused by Orf virus, and there is (Culicoides ) evidence of natural infection in dogs, 347 Orthobunyavirus

goats, Himalayan tahr, musk-ox, rein- Domingo E and Holland JJ (2005) In Topley & deer, steenbok, and alpaca. Man may Wilson’s Microbiology and Microbial Infections , be infected from animals but human- vol. 1, Tenth edition, edited by BWJ Mahy and to-human infection is very rare. Does V ter Meulen, London: Hodder Arnold, p. 11 not naturally infect cattle but they can be infected experimentally as can rab- Orinoco sheldgoose hepadnavirus A bits, horses, dogs, and monkeys. Dried probable species in the genus Avihepad- scabs retain infectivity at room temper- navirus , cloned from a captive Orinoco ature for years. Virus does not replicate sheldgoose. Genome sequence analysis on the CAM but multiplies in ovine showed a close relationship to duck and bovine cell cultures. An active vac- hepatitis B virus. cine applied by scarification has been Guo H et al (2005) J Virol 79 , 2729 used. Synonyms : contagious pustular derma- Oriximiná virus (ORXV) A serotype of titis of sheep virus; contagious pustu- Candiru virus in the genus Phlebovirus , lar stomatitis of sheep virus; ecthyma belonging to the Candiru complex in contagiosum of sheep virus; infectious the sandfly fever serogroup. Isolated labial dermatitis virus; scabby mouth from Lutzomyia sp in Para, Brazil. virus; sore mouth virus. Oropouche virus (OROV) A species in the Haig DM (2006) Curr Opin Infect Dis 19 , 127 genus Orthobunyavirus. First isolated Mercer AA et al (2006) Virus Res 116 , 146 Nitsche A et al (2006) Clin Chem 52 , 316 from a febrile patient in Trinidad in 1955. In 1960 it caused an outbreak of organ culture A form of primary tis- fever affecting 7000 people near Belem sue culture in which the cells are not in Brazil, and has become a continu- trypsinized in order to maintain their ing problem in that region, associated interactions as close as possible to the with deforestation and increased con- situation in vivo . tact with the principal vector, Culicoides paraensis. The disease is an acute febrile illness with headache, myalgia, arthral- Oriboca virus (ORIV) A species in the gia, photophobia, retrobulbar pain, genus Orthobunyavirus, belonging to nausea, and dizziness. The virus has the group C serogroup. Isolated from been isolated from the three-toed sloth, the opossum, Didelphis marsupialis , Bradypus tridactylus , which appears and from rodents, Proechimys sp and to be involved in the sylvan cycle of Oryzomys capito . Mosquito-borne. Found transmission. in Brazil, Surinam, French Guiana, and Trinidad. Can cause a febrile illness in Anderson CR et al (1961) Am J Trop Med Hyg humans. 10 , 574 Pinheiro FP et al (1981) Am J Trop Med Hyg Causey OR et al (1961) Am J Trop Med Hyg 30 , 149 10 , 227 Wang H et al (2001) Virus Res 73 , 153 Karabatsos N and Shope RE (1979) J Med Virol 3 , 167 orphan virus Any virus which has not been identified as the cause of a dis- origin of viruses The question of where ease. See e.g. human echoviruses and viruses arose has been the subject of Reoviridae . much speculation. Most popular theories claim that viruses arose from cel- Orphanovirus A provisional name, that lular RNA or DNA, but some think was not adopted, for the genus in viruses evolved with primordial life the family Picornaviridae now called forms during the earliest origins of life Parechovirus . on earth. There are also claims by some well respected scientists that viruses Orthobunyavirus A genus in the fam- were seeded on earth from outer space. ily Bunyaviridae , most of which are It may never be possible to prove or arthropod-transmitted. The type species disprove any of these theories. is Bunyamwera virus. Orthohepadnavirus 348

Orthohepadnavirus A genus in the fam- the cell. The other envelope proteins ily Hepadnaviridae comprised of viruses are an abundant membrane (M) protein which infect mammals. The type species, and a small protein (M2 in influenza Hepatitis B virus, only infects humans A, NB in influenza B) that functions naturally, although it can be transmitted as an ion channel and is involved in experimentally to some nonhuman pri- cell fusion. In influenza C the single mates. There are also hepatitis B viruses surface glycoprotein is a hemagglutinin- of nonhuman primates. Other species esterase-fusion (HEF) protein that has in the genus are Ground squirrel hepatitis both cell attachment and receptor- virus, Woodchuck hepatitis B virus, Woolly destroying enzyme activity. The nucleo- monkey hepatitis B virus and, tentatively, capsid has helical symmetry 9 nm Arctic squirrel hepatitis virus. Virions in diameter, and is segmented into are spherical, 40–42 nm diameter, con- different-sized pieces, 150–130 nm in taining relaxed circular, partially double- length, each with a loop at one end. The stranded DNA, 3.2 kb in length, with 5 nucleocapsid contains the genome RNA cohesive ends. The genome contains a (eight segments in influenza A and B gene termed ‘ X ’ (involved in transcrip- and Isavirus; seven segments in influ- tional regulation) that is not found in enza C and Dhori virus; six segments in members of the genus Avihepadnavirus . Thogoto virus) as well as three polymer- All the viruses in the genus cause acute ase proteins, PA, PB1, and PB2, which are and chronic hepatitis which can lead to involved in genome trans cription and hepatocellular carcinoma. In the case of replication, together with at least two human hepatitis B, transmission occurs nonstructural proteins (NS1 and nuclear mainly by percutaneous contact with export protein or NS2). Transcription infected blood or body fluids, sexual occurs in the host cell nucleus and contact, and perinatal transmission from requires capped oligo nucleotide primer an infected mother, though this can be RNAs (10–13 nt long) that are derived prevented by appropriate use of vaccina- from newly synthesized host cell mRNAs tion if the infectious status of the mother by viral endonuclease activity of the PB2 is known before birth. protein. Some of the mRNAs have over- Gerlich WH and Kann M (2005) In Topley & lapping sequences and are spliced to pro- Wilson’s Microbiology and Microbial Infections , vide alternative products M2 and NS2 vol. 2, Tenth edition, edited by BWJ Mahy (nuclear export proteins), or are bicis- and V ter Meulen, London: Hodder Arnold, tronic (NA and NB of influenza B virus). p. 1226 Because of the intimate involvement of influenza replication with host cell tran- Orthomyxoviridae A family of nega- scription and splicing mechanisms, it is tive strand RNA viruses comprised of blocked by substances which inhibit these five genera: Influenza virus A; Influenza processes (actinomycin D, -amanitin). virus B; Influenza virus C; Thogotovirus Protein synthesis occurs in the cyto- and Isavirus . Virions are pleomorphic, plasm, and integral membrane proteins 80–120 nm in diameter, usually roughly migrate through the Golgi apparatus to spherical but filamentous forms up the plasma membrane where virions are to several microns in length occur. assembled and new virions are formed The virion envelope H is derived by budding. In mixed infections of from cell membrane lipids into which influenza A viruses gene reassortment virus-specific glycoproteins and non- may occur during replication or assem- glycosylated proteins are inserted bly but this does not occur between during virus assembly and budding. viruses of different genera. Influenza They form projections 10–14 nm in viruses agglutinate erythrocytes from length on the virion surface. In influenza many species, and antibody to the HA A and B viruses the surface glycopro- or HEF proteins neutralizes infectivity. teins are of two types: a hemagglutinin Hemagglutination is the basis for sero- (HA) that is involved in virus attach- typing because of the wide variation ment and cell fusion; and a neuramini- in HA structure that can be detected by dase (NA) (receptor-destroying enzyme) this technique. On this basis, 16 subtypes that participates in virus release from of HA are recognized (H1 to H16); the 349 Orthoreovirus

NA protein also undergoes variation Fenner F et al (Editors) (1989) The and 9 subtypes are recognized (N1 to Orthopoxviruses . New York: Academic Press N9). Within each subtype, multiple anti- Ropp SL et al (1995) J Clin Microbiol 33 , 2069 genic variations can be detected and Smith GL (2005) In Topley & Wilson’s Microbiology and Microbial Infections , vol. 1, used to differentiate strains of influenza Tenth edition, edited by BWJ Mahy and V virus. All 16H subtypes and all 9N sub- ter Meulen, London: Hodder Arnold, p. 578 types have been found in viruses from aquatic birds, which provide the genetic Orthopoxvirus bovis Name proposed for reservoir for generation of viruses infect- Cowpox virus , but not adopted. ing humans, horses, swine, domestic poultry, and sea-mammals. Much less variation is seen in influenza B and C Orthopoxvirus commune Name proviruses, which do not appear to have a posed for Vaccinia virus, but not reservoir in avian species. Thogoto virus adopted. and Dhori virus are transmitted between vertebrates by ticks and are not known to Orthopoxvirus officinale Name pro- cause disease in humans. Isaviruses only posed for Vaccinia virus, but not infect fish. adopted.

Cox NJ et al (2005) In Topley & Wilson’s Orthopoxvirus simiae Name proposed Microbiology and Microbial Infections , Tenth edition, BWJ Mahy and V ter Meulen, for Monkeypox virus , but not adopted. London: Hodder Arnold, 634 Orthoreovirus A genus of the family Orthopoxvirus A genus of the subfamily Reoviridae containing viruses which Chordopoxvirinae. Consists of viruses of infect only vertebrates and are spread by mammals, most of which cause gener- the respiratory or fecal–oral routes. The alized infections with a rash. Virions type species is Mammalian orthoreovirus . are brick-shaped, 220–450 nm long and Strains have been isolated from humans 140–260 nm wide, and ether-resistant. and many other mammals and birds. Different species undergo genetic Five species are currently recognized: recombination and exhibit serologi- Mammalian orthoreovirus, Avian ortho- cal cross-reactivity and nucleic acid reovirus, Baboon orthoreovirus, Nelson Bay homology. The DNA genome is a single orthoreovirus , and Reptilian ortho reovirus . molecule of covalently closed double- There are a number of group- and stranded DNA, 170–250 kb in length, type-specific antigens. Reoviruses have with G C about 36%. A hemaggluti- been isolated from humans and other nin (HA) is produced by infected cells; animals with a variety of febrile, enteric, it is serologically specific and is a lipid- and respiratory diseases, but the evi- rich pleomorphic particle 50–65 nm dence that they cause disease is not in diameter, separate from the virion. strong. They can often be isolated from They all produce pocks on the CAM the respiratory tract or feces of normal but some strains may be more eas- humans and other animals. Injection of ily isolated in animals or cell cultures. newborn mice with serotype 3 causes They replicate in chick embryo fibro- oily hair, jaundice and stunted growth, blasts, Vero cells and HeLa cell cultures. although often with recovery. Serotypes Because of the close serological relation- 1 and 2 more often cause cardiac and ship within the genus, species are iden- pulmonary lesions. Virus has been tified by biological characters. However, isolated from mosquitoes but replica- they can be accurately distinguished by tion probably does not occur in insects. polymerase chain reaction-based meth- Virus particle diameter: 60–75 nm; ods. Type species is Vaccinia virus. Other buoyant density: 1.38 g/ml, contains species are Variola , Camelpox, Cowpox , 14.6% RNA and 86% protein. RNA is Ectromelia, Monkeypox, Raccoonpox , double-stranded in 10 segments, catego- Taterapox , and Volepox viruses . Tentative rized in three size classes: large (L1–L3), species in the genus are skunkpox and about 3.8 kb; medium, (M1–M3), about Uasin Gishu disease viruses. 2.3 kb; and small (S1–S4), about 1.3 kb Synonym : vaccinia subgroup viruses. in length. The positive strands of each Orthoreovirus 350

duplex have a 5 -terminal cap structure Released from the cells by lysis or (type 1), and the negative strands have budding through membranes. Found 5-phosphorylated termini. There are in Uganda, Nigeria, Central African no 3 -poly A tracts. The virions also Republic, and Senegal. Has been iso- contain numerous oligonucleotides and lated from human blood and causes a single-stranded adenine-rich RNA mak- febrile illness in humans, with head- ing up 25% of the total encapsulated ache, conjunctivitis, myalgia, vomit- nucleic acid. Single-stranded RNA is not ing, and rash. Antibodies are found in required for infectivity and its function humans, other primates, sheep, and is unknown. Surrounding the nucleic cows in Nigeria. acid core are two protein shells with Brown SE et al (1991) J Gen Virol 72 , 1065 icosahedral symmetry, but the detailed Tomori O (1977) Microbios 19 , 157 structure remains to be elucidated. The Tomori O et al (1977) Arch Virol 55 , 181 outer shell can be removed with trypsin leaving a subviral particle of diam- Oryctes rhinoceros virus (OrV) An unas- eter 40–45 nm. Mammalian strains, but signed insect virus that infects scarab not avian, agglutinate human group beetles, not rhinoceroses. O erythrocytes. Infectivity is stable between pH 2.2 and pH 8.0, resists Oseltamivir A carbocyclic analog of sialic ether, 1% phenol, 3% formalin, and 20% acid with potent anti-influenza neurami- lysol but is inactivated by 70% ethanol. nidase activity. An orally administered Mammalian strains replicate with CPE neuraminidase inhibitor, effective in pre- in primary and continuous cell lines vention of influenza when given daily derived from a wide range of animal for 6 weeks during the influenza season. species. Avian strains replicate with Also active against human experimental CPE in chick embryo cell cultures. infection with Influenza B virus. Synonym : reovirus. Synonym : Tamiflu. Dermody TS and Desselberger U (2005) In Hayden FG et al (1999) N Engl J Med 341 , 1387 Topley & Wilson’s Microbiology and Microbial Hayden FG et al (2000) Antiviral Ther 5 , 205 Infections , vol. 2, Tenth edition, edited by Nicholson KG et al (2000) Lancet 355 , 1845 BWJ Mahy and V ter Meulen, London: Ong AK and Hayden FG (2007) J Infect Dis Hodder Arnold, p. 932 196 , 181

Orthoretrovirinae A subfamily of the osmiophilic inclusion bodies Inclusion family Retroviridae comprised of six bodies which contain myelin whorls genera, Alpharetrovirus , Betaretrovirus , and densely packed vesicles seen in Gammaretrovirus , Deltaretrovirus , Epsilon- human fetal, nasal, and tracheal organ retrovirus , and Lentivirus . cultures infected with rubella virus.

Orungo virus (ORUV) A species in the Ossa virus (OSSAV) A strain of Caraparu genus Orbivirus that has four serotypes virus in the genus Orthobunyavirus , (Orungo viruses 1–4). Isolated from belonging to the group C viruses. culicine mosquitoes, humans, camels, Isolated from humans, the spiny rat, cattle, goats, monkeys, and sheep. Proechimys semispinosus , and Culex sp in Panama. Associated with a febrile Orungo virus 1–4 (ORUV 1–4) Species in illness in humans. the genus Orbivirus , belonging to the de Rodaniche E et al (1964) Am J Trop Med Orungo serogroup. First isolated from Hyg 13 , 839 the mosquito, Anopheles funestus , in Uganda, and the mosquito, Aedes den- osteolytic syndrome A developmen- tatus , and humans in Nigeria. Can be tal abnormality syndrome in rodents passed in newborn mice by i.c. injection. caused by parvovirus infection of the Replicates with CPE in BHK21 cells. fetus late in pregnancy or soon after Develops in the cytoplasm associated birth. Cytolytic replication in osteo- with a specific viral granular matrix genic tissues results in animals with and accompanying filaments. Virion mongoloid features, small flat faces, diameter: 63 nm; core diameter: 34 nm. microcephalic heads, protruding eyes 351 Ovine adeno-associated virus (OAAV)

and tongue, missing or abnormal teeth Inhibits the growth of mouse cell lines and fragile bones. May also result in K3b and JLS-V9, and the production persistent dwarfism. in them of mouse type C retroviruses. Low concentrations of ouabain inhibit osteopetrosis virus A strain of avian leu- the growth of some enveloped viruses kosis virus in the genus Alpharetrovirus such as influenza and parainfluenza which can cause osteopetrosis, a disease viruses in chick embryo cells. which occurs naturally in fowls, and Tomita Y and Kuwata T (1978) J Gen Virol is characterized by enlargement of the 38 , 223 bones, especially the leg bones. There is an increase of hard bone due to hyper- Ouango virus (OUAV) An unassigned trophic activity of the periosteum. It can vertebrate rhabdovirus. Isolated in 1970 be transmitted by inoculation of day-old from a bird, Sitagra melanocephala , in chicks or the amnion of chick embryos Central African Republic. Not reported with blood from birds with osteopetro- to cause disease in humans. sis or from some cases of lymphomatosis. Osteopetrosis is often accompanied Oubangui virus An unclassified arbo- by soft tissue tumors. Infected tissue virus. Isolated in suckling mice from contains large amounts of virus-specific a pool of female mosquitoes, Culex DNA, much of which is unintegrated. guiarti, collected in Bangui, Central Turkeys do not naturally develop the African Republic from human bait. disease but it can be transmitted to Infectivity resistant to chloroform. them. See Avian myeloblastosis virus . Hemagglutinates goose erythrocytes. Synonyms: big bone disease virus; mar- Not reported to cause disease in ble bone disease virus; thick leg disease humans. virus. Foster RG et al (1994) Virology 205 , 179 Oubi virus (OUBIV) A strain of Olifantsvlei Robinson HL et al (1986) J Virol 59 , 45 virus in the genus Orthobunyavirus . Smith RE (1982) Curr Top Microbiol Immunol Isolated from Culex (Eumelanomyia ) rima 101 , 75 sp in the Ivory Coast. Not reported to cause disease in humans. osteosarcomas Tumors of the bone. Institut Pasteur, Dakar (1982) Annual Report , 84 ostreid herpesvirus 1 (OsHV-1) An Ourém virus (OURV) A serotype unassigned member of the family of Changuinola virus in the genus Herpesviridae which infects oysters. Orbivirus . Isolated from Lutzomyia sp in Experimental transmission to the Para, Brazil. Not reported to cause dis- Pacific oyster, Crassostrea gigas , has been ease in humans. reported. Le Deuff RM et al (1994) Bull Eur Assoc Fish outlier (O) subtype A subtype of HIV, Pathol 14 , 69 named on the basis of its sequence to distinguish it from the main (M) ostrich adenovirus A virus isolated from subtype. ostriches with respiratory disease. Gough RE et al (1997) Vet Rec 140 , 402 ovine AAV See Ovine adeno-associated virus . otofuke agent A small round-structured virus detected by electron microscopy Ovine adeno-associated virus (OAAV) in the feces of Japanese patients with A species in the genus Dependovirus . gastroenteritis. Isolated from ovine feces, in association with an adenovirus in lamb kidney ouabain A cardiac glycoside obtained cell cultures. Agglutinates guinea pig from the seeds of Strophanthus gratus and human erythrocytes. Replicates in or the wood of Acokanthera schimperi or association with the cell nucleus and A ouabio . An inhibitor of Na - and K - requires the presence of an adenovirus. dependent adenosine triphosphatase. Clarke JK et al (1979) Arch Virol 60 , 171 Ovine adenoviruses A to C (OadV A–C) 352

Ovine adenoviruses A to C (OadV A–C) in epithelial cells of the small intestine Species in the genus Mastadenovirus by immunofluorescence. Crystalline which cause mild upper respiratory arrays of 29 nm diameter particles can tract or enteric infections in sheep. Six be seen by electron microscopy in the serotypes are identified: ovine adeno- cytoplasm of cells in microvilli. virus 1 (Ovine adenovirus B ), ovine adeno - Herring AJ et al (1981) J Gen Virol 53 , 47 viruses 2–5 (Ovine adenovirus A ), and ovine adenovirus 6 (ovine adenovirus ovine catarrhal fever virus Synonym for C), a tentative species. Because few Bluetongue virus . sheep give a positive reaction in the gel diffusion test for precipitating antibod- ovine encephalomyelitis virus ies it was suggested that the infection Synonym was not common, but the majority of for louping-ill virus. sheep in Scotland have neutralizing ovine herpesvirus 1 (OvHV-1) antibodies to four serotypes of this An unas- Herpesviridae virus. The infection must therefore be signed species in the family . common and, as the virus can be recov- Found during studies on sheep pulmo- ered from the feces of normal sheep, its nary adenomatosis (jaagsiekte) but not isolation from disease outbreaks must the cause of that disease. Synonym be viewed with caution. Multiplies in : sheep pulmonary adenoma- sheep kidney cell cultures. Agglutinates tosis-associated herpesvirus. rat erythrocytes. Martin WB et al (1976) Nature 264 , 183 Belak S (1990) In Virus infections of Rumi- nants , edited by Z Dinter and B Morein. Ovine herpesvirus 2 (OvHV-2) A spe- Amsterdam: Elsevier Science, p. 171 cies in the genus Rhadinovirus . Causes Sharp JM (1977) Vet Rec 101 , 524 sheep-associated malignant catarrhal fever of cattle, bison and deer. The virus ovine adenoviruses 1–6 (OadV-1 to -6) has not been grown in cell culture. Originally six serotypes of sheep adeno- Synonyms : sheep-associated malignant virus were recognized. These are now catarrhal fever of cattle virus; herpes- grouped into three species (A–C); Ovine virus ovis. The complete sequence adenovirus A incorporates serotypes 2–5 of the genome (130,930 bp) has been as well as bovine adenovirus 2. Ovine determined. An excellent model has adenovirus B is the original serotype 1, been established in rabbits, which and the third putative species, Ovine develop a disease that resembles that adenovirus C , is the original serotype 6. in ungulates. Anderson IE et al (2007) J Comp Pathol 136 , ovine adenovirus isolate 287 (OadV-287) 156 A strain of Ovine adenovirus D in the Bridgen A and Reid HW (1991) Res Vet Sci genus Atadenovirus . 50 , 38 Hart J et al (2007) J Gen Virol 88 , 28 Harrach B et al (1997) Virology 229 , 302 Plowright W (1986) Rev Sci Tech 5 , 897

Ovine astrovirus (OAsTV) A species in ovine lentivirus There are two dis- the genus Mamastrovirus , detected in eases associated with a lentivirus in stool samples of sheep. The disease sheep, Visna / maedi virus in the genus is generally a mild self-limiting acute Lentivirus . In 1933, 20 Karakul sheep gastroenteritis of young animals. In were imported into Iceland from experimental infection of lambs, there Germany, and within 2 years the two was a destruction of mature enterocytes diseases visna (meaning wasting) and on the apical two-thirds of villi, leading maedi (meaning dyspnea) emerged to villus atrophy and crypt hypertrophy. and became a major problem to the sheep industry in Iceland. Following ovine astrovirus 1 (OAstV-1) A sero- a massive slaughter of some 600,000 type of Ovine astrovirus in the genus sheep, the disease was declared eradi- Mamastrovirus , associated with acute cated from Iceland in 1965, though enteritis in lambs. Virus can be detected it continues in some other European 353 owl’s eye intranuclear inclusions

countries and in the USA where it is ovine progressive pneumonia See called progressive pneumonia of sheep. Visna / maedi virus . See Visna/maedi virus . Campbell BJ and Avery RJ (1996) J Gen Virol Ovine pulmonary adenocarcinoma virus 77 , 2999 (OPAV) a synonym for Jaagsiekte sheep Clements JE and Zink MC (1996) Clin Micro- retrovirus . biol Rev 9 , 100 Karr BM et al (1996) Virology 225 , 1 ovine pulmonary adenomatosis A disease of sheep caused by jaagsiekte sheep ret- Ovine papillomavirus 1 (OvPV-1) A spe- rovirus in the genus Betaretrovirus . cies in the genus Deltapapillomavirus . There are two genotypes. Papillomas occur in some flocks but are not com- ovine respiratory syncytial virus Isolates mon. They may become malignant. of respiratory syncytial virus have been Virus inoculated into the skin of sheep made from sheep which appear to be produces papillomas. Cattle and distinct from the human or bovine goats are resistant, but slowly grow- viruses, but little is known of the clini- ing fibromas are produced in neonatal cal characteristics or significance of hamsters. Causes a cutaneous fibro- natural infection with this virus. papilloma of sheep, found on the eye- lid, muzzle, vulva, and legs. owl hepatosplenitis herpesvirus Syno- Synonym : sheep papillomavirus. nym for strigid herpesvirus 1. Gibbs EPJ et al (1975) J Comp Pathol 85 , 327 Vanselow BA et al (1982) Vet Rec 110 , 561 owl monkey herpesvirus Synonym for cebine herpesvirus 1 and 2. ovine papillomaviruses 1 and 2 (OPV-1 and OPV-2) Two serotypes of Ovine owl’s eye intranuclear inclusions Char- papillomavirus 1 have been reported in acteristic large inclusion bodies seen the literature, but their taxonomic stain the nucleus of cells infected with tus is uncertain. cytomegaloviruses. P

P1 kinase An enzyme induced by inter- fever with renal syndrome (HFRS) in feron which is a protein-serine/threo- western Russia. Phylogenetic analy- nine kinase, designated the P1/eIF-2α sis showed it to be related to Puumala kinase. Plays a central role in transla- virus; the probable rodent host is the tional control and in the antiviral action vole, Clethrionomys glareolus , from of interferon. which a closely similar isolate was made. P1-1-Ut (NBL-9) cells (CCL 74) A hetero- Xiao S-Y et al (1993) Virus Res 30 , 97 ploid cell line derived from trypsinized uterine tissue of an adult raccoon, 13p2 virus Synonym for American oyster Procyon lotor. Developed for studies of reolike virus. canine and feline viruses, since the raccoon is one of the few animals known PAA See phosphonoacetic acid . to be susceptible to both groups. Pacajá virus A probable species in the p2b-2 virus A strain of Tacaribe virus in genus Orbivirus isolated from phle- the genus Arenavirus . botomine sand flies in the Amazon region of Brazil. Antigenically related P4 laboratory An old name for a high to Changuinola virus . containment laboratory designed for safe work with viruses that cause seri- Pacheco’s disease virus Synonym for ous human disease for which no vac- psittacid herpesvirus 1. cines or other treatment are available (e.g. Ebola virus, Lassa fever virus) and thus present a serious hazard to the Pacific pond turtle herpesvirus Synonym laboratory worker. Now referred to as for chelonid herpesvirus 2. a BSL4 (Biosafety level 4) laboratory. Paclitaxel A mitotic inhibitor used in can- p53 A multifunctional protein which cer chemotherapy, originally obtained plays a role in modulating gene tran- from the bark of the Pacific yew (Taxus scription, control of the cell cycle, acti- brevifolia ). It is used in the treatment of vating apoptosis, controlling DNA advanced forms of Kaposi’s sarcoma. replication and repair, and maintaining genomic stability and tumor Pacora virus (PCAV) An unassigned virus suppression. Several groups of onco- in the family Bunyaviridae. Isolated from genic viruses (adenoviruses, SV40, Culex dunni in Panama. Not reported to and papillomaviruses) produce pro- cause disease in humans. teins that interact with and inactivate p53 to cause cell transformation and Pacora-like virus A possible member neoplasia. of the family Bunyaviridae . Isolated from birds Automolus chrolaemus and Elledge RM and Lee W-H (1995) Bioessays 17 , 923 Phlegopsis nigromaculata and from Culex Grinstein E and Wernet P (2007) Cell Signal mosquitoes in the Amazon region of 19 , 2428 Brazil. Not associated with disease in Levine AJ (1993) Annu Rev Biochem 62 , 623 humans. Liebermann DA et al (1995) Oncogene 11 , 199 pactamycin A drug that selectively p360 virus (P360V) A strain of Puumala inhibits initiation of protein synthesis. virus in the genus Hantavirus . Isolated Interferes with attachment of the ini- from a human case of hemorrhagic tiator tRNA to the initiation complex. 355 Papillomaviridae

Nascent polypeptides are completed. pandemic A widespread epidemic of Has proved useful experimentally in human disease occurring throughout mapping the gene order of positive- more than one country, or a continent strand viruses since, after drug addi- or globally. tion, the first polypeptides to be affected will be coded close to the 5 end panhandle structure A structure present and the last affected near the 3 end of in the RNA genome of several seg- the genome. mented negative-stranded viruses (e.g. arenaviruses, bunyaviruses, and Pacui virus (PACV) A tentative species orthomyxoviruses) that is formed in the genus Phlebovirus, member of by base-pairing of complementary the sandfly fever virus group. Isolated sequences, e.g. in influenza virus, each from rodents of Oryzomys sp in Para, of the eight RNA genome segments Brazil, and Trinidad. Not reported to has the first 12 nt at the 3 end comple- cause disease in humans. mentary to the last 13 nt at the 5 end, which form a panhandle structure that Pahayokee virus (PAHV) A serotype is involved in the initiation of tran- of Patois virus in the genus Orthobunya - scription. DNA viruses such as adenovirus. Isolated from mosquitoes in virus also utilize panhandle structures Florida, USA. Not known to cause dis- during their replicative process. ease in humans. Fodor E et al (1994) J Virol 68 , 4092 painted turtle herpesvirus Synonym for pan herpesvirus See Pongine herpesvirus 1 . chelonid herpesvirus 3. panleukopenia virus See Feline panleuko- pale bird syndrome A disease affecting penia virus . poultry throughout the world. Caused by avian reovirus infections occurring pantomorphic pattern of disease One of during the first week of age. See Avian a number of disease patterns such as orthoreovirus . neuromorphic (e.g. encephalitis), viscero- Montgomery RD et al (1986) Avian Dis 30 , morphic (e.g. hemorrhagic fever), 460 and pantomorphic (e.g. nonspecific febrile illness) coined by Monath. Palestina virus (PLSV) A serotype of There is some doubt as to the value of Minatitlan virus in the genus Orthobunya - this classification. virus . With Minatitlan virus forms Monath TP (1986) In Togaviridae and the Minatitlan virus group. Isolated Flaviviridae, edited by S Schlesinger and MJ from Culex ( Mel ) paracrybda in Schlesinger. New York: Plenum Press, p. 375 Ecuador. Pan troglodytes rhadinovirus 1 A proba- Palyam virus (PALV) A species in the ble member of the genus Rhadinovirus genus Orbivirus , belonging to the isolated from wild-caught chimpanzees Palyam serogroup. Isolated from in Gabon and Cameroon. Culex vishnui in Tamil Nadu, India, from Culicoides sp in Australia and panzootic A widespread epidemic of from Culicoides sp and mosquitoes in disease affecting many animals over a Abadina, Nigeria. wide area.

Pampa virus (PAMV) A tentative species papilloma of chamois A parapox virus in the genus Arenavirus . Isolated from infection. See chamois contagious rodents, Bolomys sp, in Argentina. ecthyma virus .

PAn 18400 virus A strain of Pampa virus. Papillomaviridae A family of viruses containing 16 genera. Virions are non- pancreatic necrosis virus of fish See enveloped, 55 nm in diameter. The Infectious pancreatic necrosis virus . icosahedral capsid has 72 capsomeres Papillomaviridae 356

in a skewed (T 7) arrangement. parainfluenza virus type 2 human See The genome is a circular dsDNA, Human parainfluenza virus type 2 . 6800–8400 bp in length, encoding 8–10 proteins. parainfluenza virus type 3 human See Human parainfluenza virus type 3 . papillomavirus sylvilagi See Cottontail rabbit papillomavirus . parainfluenza virus type 4 human See Human parainfluenza virus type 4 . papio Epstein–Barr herpesvirus See Cercopithecine herpesvirus 12 . parainfluenza virus type 1 murine See Papovaviridae A discontinued family. Sendai virus . Now split into the families Papillo ma- viridae and Polyomaviridae . parainfluenza virus type 5 simian See Simian virus 5 . Papovaviruses An old name for the species first included in the former family Parakaná virus A possible species in the Papovaviridae. They were pa pilloma- genus Phlebovirus , isolated from mos- virus, po lyomavirus, and va cuolating quitoes in the Amazon region of Brazil. virus. No longer in use. Not associated with human disease. pappataci fever viruses Synonym for paralytic poliomyelitis The major illness phlebotomus fever viruses. following poliovirus infection, which occurs in 0.1–2% of infected persons. PA protein A component of the RNA tran- It is a flaccid paralysis resulting from scriptase complex of influenza virus, lower motor neuron damage. It can encoded by genome RNA segment 3. be spinal, or, more seriously bulbar PA stands for polymerase acidic, to dis- if the upper spine and brainstem are tinguish it from PB1 and PB2, which affected, since this affects the respira- are polymerase basic proteins. tory system. papular stomatitis of cattle virus paralytic rabies A clinical course seen Synonym for Bovine papular stomatitis in about 20% of patients with acute virus, although other viruses may cause rabies infection characterized by par- the same clinical picture. esthesia, weakness, and flaccid paralysis. Contrasts with furious rabies, the Papura virus An unclassified virus iso- commonest form. Also called ‘ dumb lated once from culicine mosquitoes in rabies. ’ the Amazon region of Brazil. Paramaribo virus An Alphavirus first PARA P article a iding r eplication of a den- isolated in Paramaribo, Surinam, ovirus. See adenovirus-SV40 hybrids . from the blood of an adult male. Pathogenic for suckling mice, hamsters Pará virus (PARAV) A strain of Simbu rats, and guinea pigs by intracerebral virus in the genus Orthobunyavirus . inoculation. Isolated from a sentinel mouse in Brazil Metselaar D et al (1964) Arch Virol 14 , 336 and from a pool of Culex ocossa mosqui- Young NA and Johnson KM (1969) Am toes in Argentina. J Epidemiol 89 , 286 Saeed et al (2001) J Gen Virol 82 , 2173 Paramastadenovirus A proposed name paracrine signals Chemical signals that for a third genus of the family Adeno- originate from neighborhood cells in viridae , containing viruses with an the vicinity of the target cell. AT-rich genome. Now called the Atadenovirus genus. parainfluenza virus type 1 human See Benkö M and Harrach B (1998) Arch Virol Human parainfluenza virus type 1 . 143/144 , 830 357 Parapoxvirus

Paramushir virus (PMRV) A serotype syncytium formation. Most species or strain of Sakhalin virus in the genus also replicate in embryonated eggs. Nairovirus . Also called Avalon virus. Paramyxoviruses cause a wide spec- Isolated from the ticks, Ixodes signa- trum of disease in humans and other tus and I putus , collected on Tyuleniy, animals. They can cause persistent Bering, and Paramushir Islands in infection in cultured cells and in ani- the north of the former Soviet Far mals and humans, as well as cell fusion East, where there are colonies of guil- and lysis of erythrocytes. The ability to lemots, Uria aalge , and cormorants, fuse cells and produce hybrids is used Phalacrocorax pelagicus . Pathogenic in cell genetics and in the study of cel- for suckling mice on i.c. injection. lular regulation of growth. Replicates without CPE in chick Rota PA et al (2005) In Topley & Wilson’s embryo fibroblasts, human and BHK21 Microbiology and Microbial Infections , vol. 1, cells. Not reported to cause disease in Tenth edition, edited by BWJ Mahy and humans. V ter Meulen, London: Hodder Arnold, p. 699 Lvov DK et al (1976) Arch Virol 51 , 157 Russell CJ et al (2001) EMBO J 20 , 4024 Paramyxoviridae A family of large Paramyxovirinae A subfamily of the negative-strand RNA viruses within family Paramyxoviridae containing the order Mononegavirales comprising five genera: Respirovirus , Morbillivirus , two subfamilies: (1) Paramyxovirinae , Rubulavirus , Avulavirus , and Henipavirus . containing five genera, Respirovirus , Virion nucleocapsid diameter is 18 nm, Morbillivirus , Rubulavirus , Avulavirus , and the length of the surface spikes is and Henipavirus and (2) Pneumovirinae , 8 nm. The genome includes up to seven containing the two genera, Pneumovirus transcriptional elements, in contrast and Metapneumovirus. Differ from to the subfamily Pneumovirinae , which Orthomyxoviridae in that: (1) they con - have 10. tain a larger helical nucleocapsid, 18 nm in diameter and up to 1000 nm Paraná virus (PARV) A species in the in length; (2) they have a genome con- genus Arenavirus , a New World arena - sisting of a single continuous RNA virus belonging to the Tacaribe molecule 15,156–18,246 nt (Nipah complex. Isolated from the rodent, virus) in length; and (3) their replica- Oryzomys buccinatus, in Paraguay. Not tion is not inhibited by actinomycin D. reported to cause disease in humans. Virions consist of a helical ribonucleoprotein nucleocapsid contained in Paranati virus An unclassified virus iso- an approximately spherical envelope lated from sand flies in the Amazon 150–200 nm in diameter derived from region of Brazil. Not associated with the host cell plasma membrane during disease in humans. budding. Larger virions of 500–600 nm in diameter and filamentous forms Parapox of red deer in New Zealand are occasionally observed. There are virus (PVNZ) A species in the genus two surface glycoproteins, 8–12 nm in Parapoxvirus , isolated from farmed red length: the fusion (F) and the attach- deer in New Zealand. The genome ment protein (G or H, or HN). The HN DNA differs by restriction analysis has both hemagglutinating and neur- from other parapoxviruses, and on this aminidase activity. There is also a non- basis PVNZ is a separate species in the glycosylated enveloped protein, M. genus. There are three internal nucleocapsid- Robinson AJ and Mercer AA (1995) Virology associated proteins, an RNA-binding 208 , 812 protein (N or NP), a phosphoprotein (P), and a large polymerase protein (L). Parapoxvirus A genus of the subfamily Virus replication occurs in the cyto- Chordopoxvirinae . Consists of viruses plasm. The RNA genome is transcribed of ungulates which may infect humans. into 6–10 separate mRNAs. Virus can Infectivity ether-sensitive. Virion ovoid, be propagated in tissue cultures, usu- 220–300 140–170 nm. External coat and ally with CPE, hemadsorption, and filaments are thicker than in vaccinia Parapoxvirus 358

and are arranged in a regular spiral coil Stanway G and Hyypia T (1999) J Virol 73 , consisting of a single thread. Species 5249 show serological cross-reactivity. Hema- Stanway G et al (2000) Rev Med Virol 10 , 57 gglutinin not produced. Type species Orf virus. Other species are Bovine papu- paresis Slight or incomplete paralysis, an lar stomatitis virus , Pseudocowpox virus , early symptom of clinical rabies. Parapoxvirus of red deer in New Zealand , and Squirrel parapoxvirus . Parixá virus An unclassified virus iso- Synonym : orf subgroup viruses. lated from a bat, Lonchophylla thomasi , in the Amazon region of Brazil. Not associated with disease in humans. pararetroviruses A name suggested but not adopted for two families of DNA Parma wallaby herpesvirus See macro- viruses that use reverse transcrip- podid herpesvirus 1 . tion during their replication cycle; Hepadnaviridae and the plant virus fam- Paroo River virus (PRV) A serotype of ily, Caulimoviridae . Wongorr virus in the genus Orbivirus . Isolated from Culex annulirostris in Parauapebas virus A possible species New South Wales, Australia. in the genus Orbivirus isolated from phlebotomine sand flies in the Amazon Parramatta virus A virus seen by elec- region of Brazil. Antigenically related tron microscopy in fecal extracts from to Changuinola virus. Not known to 14 patients with gastroenteritis dur- cause disease in humans. ing an outbreak in a primary school near Parramatta, New South Wales, Australia. Diameter: 23–26 nm; mor- paravaccinia virus See Pseudocowpox phologically like a parvovirus. virus . Christopher PJ et al (1978) Med J Aust i , 121

Parechovirus A genus in the family parrot herpesvirus See psittacid herpes- Picornaviridae, containing two spe- virus 1 . cies, Human parechovirus , with three serotypes, human parechovirus 1 and parrot papillomavirus A possible mem- human parechovirus 2 (formerly these ber of the Papillomavirus genus, family were called human echovirus 22 and Papovaviridae . 23), and human parechovirus 3, and Ljungan virus. A fourth serotype of Parry Creek virus (PCRV) An unas- human parechovirus has been reported signed vertebrate rhabdovirus. Isolated but not yet confirmed. The genus was from mosquitoes, Culex annulirostris , formed to recognize distinct biologi- in Australia. Some antigenic rela- cal and molecular properties of these tionship to members of the genus viruses compared with other entero- Lyssavirus . Not known to cause disease viruses in the family. Parechoviruses in humans. have characteristic effects on their Calisher CH et al (1989) Intervirology 30 , 241 host cell and cause a somewhat different spectrum of clinical disease with parsimony (stinginess, niggardliness, gastrointestinal or respiratory illness. miserliness) An approach to the con- In contrast to other picornaviruses, struction of a phylogenetic tree that the protein 1AB is not cleaved during selects the phylogeny that minimizes maturation, and is not myristylated. the number of evolutionary changes The capsid has only three proteins, required to explain the data. Used 1AB, 1C, and 1D. Ljungan virus mainly extensively to build trees based upon infects rodents. nucleotide sequences. Benschop KS et al (2006) Emerg Infect Dis 12 , Hillis DM et al (1993) Meth Enzymol 224 , 456 1572 Joki-Korpela P and Hyypia T (2001) Ann Paru virus A possible member of Med 33 , 466 the genus Orbivirus , isolated from 359 passage

phlebotomine flies in the Amazon viruses that infect vertebrates or verte- region of Brazil. Antigenically related brate cell cultures. There are five genera: to Changuinola virus. Not known to Parvovirus , Erythrovirus , Depen do virus , cause disease in humans. Amdovirus , and Bocavirus .

PARV 4 and 5 viruses A novel human Parvovirus A genus of the family parvovirus found mainly in patients Parvoviridae . Type species Minute virus with hemophilia or infected with HIV. of mice. Parvoviruses replicate in sus- Appears to be transmitted parenterally, ceptible cell cultures without a helper e.g. by injection drug use. An analysis virus. Mature virus particles contain of the tissue distribution in autopsy only positive strands of DNA which specimens from the UK indicated a have a hairpin structure at both the 5 change in genotype of the virus, with and 3 ends of the otherwise single- PARV 4 virus found in subjects born stranded molecule. There are many since 1958, but a variant virus (known species infecting a variety of verte- as PARV 5) found in older subjects. Both brates and most are host-specific. A PARV 4 and PARV 5 have been found in number of parvoviruses have been iso- factor VIII concentrates manufactured lated from cell lines, and their origins in the UK. The disease significance of are therefore doubtful. Only the feline this virus is unknown. and related viruses are known to cause Fryer JF et al (2007) Vox Sang 93 , 341 disease in the wild. The pathogenicity Manning A et al (2007) J Infect Dis 195 , 1345 of the rodent viruses in the laboratory suggests that parvoviruses may cause Parvoviridae (Latin: parvus small) A non-acute disease in humans and other family of small DNA viruses. Virion animals. Replication in cell culture is diameter, 18–26 nm, non-enveloped, best in rapidly dividing cells derived composed of 60 copies of the capsid from tissues of the natural host, but protein with icosahedral symmetry. usually produces only slight or tran- Buoyant density (CsCl): 1.39–1.42 g/ml. sient CPE, making detection difficult. Genome consists of one molecule of However, many species hemaggluti- linear single-stranded DNA, 4–6 kb in nate, and infected cells hemadsorb. No length, mol. wt. 1.5–2.2 1 0 6 . G C group antigen and no antigenic rela- content: 41–53%. In some genera the tionship to other DNA viruses. single strands from virions are com- Berns KI (1990) Microbiol Rev 54 , 316 plementary and, after extraction, these Lukhov VV and Goudsmit J (2001) J Virol 75 , come together to form a double strand. 2729 Infectivity ether-resistant and relatively heat-stable. Replication occurs in parvovirus B19 See Human parvovirus the nucleus and is dependent on either B19 . certain functions of the host cell or on a helper virus. There are two subfamilies: parvovirus initiation factor (PIF ) A cellu- (1) Parvovirinae, genera Parvovirus , lar factor required for the initiation of Erythrovirus , Dependovirus , Amdovirus , replication of parvovirus DNA. Forms and Bocavirus and (2) Densovirinae , a complex with the viral NS1 protein genera Densovirus , Iteravirus , Brevidenso- which binds to the DNA that is capable virus , and Pefudensovirus . The mem- of nicking and subsequently initiating bers of the Densovirinae subfamily only replication at the origin. infect arthropods. Paschen bodies Elementary bodies found Siegl G et al (1985) Intervirology 23 , 61 Tattersall P and Cotmore SF (2005) In in cells infected with variola or vaccinia Topley & Wilson’s Microbiology and Microbial viruses. See also Buist bodies . Infections , vol. 1, Tenth edition, edited by Synonym : chlamydozoa variolae. BWJ Mahy and V ter Meulen, London: Hodder Arnold, p. 407 passage Infection of a host with a virus or a mixture of viruses with subse- Parvovirinae A subfamily of the fam- quent recovery of the virus from that ily Parvoviridae which includes only host (usually after one infection cycle). passage 360

Can be used to separate a specific virus Patois virus (PATV) A species in the from a mixture of viruses, or (through genus Orthobunyavirus , belonging to a series of passages) a virus can be the Patois serogroup. Isolated from adapted to grow well in a host in which cotton rats, Sigmodon hispidus , and it originally grew poorly. mosquitoes of Culex sp in Panama and Mexico. Not reported to cause disease Passatempo virus A possible member of in humans. the genus Orthopoxvirus , isolated during a zoonotic outbreak in Brazil. pattern recognition receptors (PRRs) Leite JA et al (2005) Emerg Infect Dis 11 1935 Recognition of pathogens by phagocytic cells is achieved by pattern rec- passenger virus Any non-pathogenic ognition receptors encoded in the virus. When isolated from diseased tis- germline. The PRRs have broad specifi- sue it has no causal relationship to the city and recognize pathogen-associated disease process. molecular patterns that differ between pathogens but are not present in the passive hemagglutination A serological host. There are two major groups of test which can be used to detect a virus- PRRs – those secreted in the blood and specific antibody by coating red blood lymph, and those on the surface of cells with viral antigen. If viral anti- cells. body is present in test samples, the red blood cells will agglutinate. See also Paul–Bunnell antibodies Heterophile reverse passive hemagglutination . antibodies characteristic for infectious mononucleosis (glandular fever), passive immunization The use of anti- described by Paul and Bunnell in 1932. bodies purified from human or animal They are predominantly IgM, but their serum to prevent infection or to mod- mode of formation is not fully under- ify the course of infection when given stood. In the Paul–Bunnell test, they either before or after exposure to a are detected by agglutination of sheep virus. Since 1998 a new immunoglobu- erythrocytes. lin product, humanized monoclonal antibodies against respiratory disease Paul–Bunnell–Davidsohn test The Paul– virus, has been licensed for use in the Bunnell test for glandular fever as USA. originally described consisted of a simple titration of the patient’s serum for Pasteur virus A strain of Rabies virus . sheep erythrocyte-agglutinating antibodies. In order to distinguish between Pata virus (PATAV) A serotype of infectious mononucleosis, Forssman Eubenangee virus in the genus Orbivirus . and serum-sickness antibodies (all With Eubenangee, Ngoupe, and of which agglutinate sheep cells), Tilligerry viruses forms the Eubenangee Davidsohn introduced an absorption serogroup. Isolated from Aedes pal- step into the test, the patient’s serum palis in Central African Republic. Not being absorbed with: (a) guinea pig reported to cause disease in humans. kidney tissue and (b) ox cells before titration of agglutinins. Guinea pig kid- patas monkey herpesvirus Synonym for ney absorbs both Forssman and serum- cercopithecine herpesvirus 9. sickness antibodies; ox cells absorb McCarthy K et al (1968) Lancet ii , 856 serum-sickness and infectious mononucleosis antibodies, so that it is pos- pathogen An organism or virus which sible to distinguish between the three causes a disease. agglutinins.

Pathum Thani virus (PTHV) A strain Paul Ehrlich Institute units (PEIs) of Dera Ghazi Khan virus in the genus Diagnostic units used to measure hep- Nairovirus. Isolated from a tick, Argas atitis B surface antigen (HbsAg) stand- robertsi, in Thailand. Not reported to ardized by the Paul Ehrlich Institute, cause disease in humans. Frankfurt, Germany. 361 pentamers

Paxillin A scaffolding protein which modu- Peaton virus (PEAV) A serotype lates network interactions amongst a of Shamonda virus in the genus variety of proteins related to cell adhe- Orthobunyavirus , belonging to the sion. Transduces signals from the Simbu serogroup. Isolated from the plasma membrane to focal adhesions mosquito, Culicoides brevitarsis , in and the actin cytoskeleton. Queensland, Australia. Paver virus A small virus 22 nm in diam- St George TD et al (1980) Aust J Biol Sci 33 , eter, seen by immunoelectron micro- 235 scopy in the feces of patients with gastroenteritis and in normal subjects. PEG Abbreviation for p oly e thylene g lycol. Very similar in morphology and den- A chemical used for phase partitioning sity on CsCl gradients to porcine parvo- that is often employed to concentrate virus and mink enteritis virus. viruses present in a suspension. Paver WK et al (1974) J Gen Virol 22 , 447 pellet The material concentrated at the PB1-F2 protein A small protein (79–101 bottom of a centrifuge tube after cen- amino acids) induced in cells infected trifugation. See supernatant . with influenza A viruses, which specifically targets and destroys mono- penciclovir A drug related to acyclovir cytes and alveolar macrophages. It is which is selectively phosphorylated by encoded in a second (alternate) 1 the HSV thymidine kinase and inhib- reading frame of influenza A genome its replication of herpesviruses such as RNA segment 2, the primary product VZV and CMV as well as HSV. Because of which is the PB1 protein. The func- it was not effective orally, the 6-deoxy- tion of PB1-F2 appears to kill host diacetyl ester derivative famciclovir immune cells following influenza virus was developed as an orally delivered infection. It has been called the influ- prodrug of penciclovir and is now enza death protein. licensed for treatment, especially of Chen W et al (2001) Nat Med 7 , 1306 herpes zoster infections in immunosuppressed patients. It is also effective PB1 protein A basic component of the against hepatitis B. influenza virus RNA transcriptase complex, involved in RNA-dependent RNA Vere Hodge RA (1993) Antivir Chem polymerase activity and capped mRNA Chemother 4 , 67 endonuclease activity. Encoded by RNA segment 2 of the virus genome. penetration The second stage of infection of a cell by a virus. After attachment to PB2 protein A basic component of the a receptor area on the cell surface, the influenza virus RNA transcriptase virus is taken into the cell either by complex involved in host cell capped fusion with the plasma membrane or mRNA recognition and binding. by viropexis. The mechanism differs Encoded by RNA segment 1 of the from one virus/cell system to another. genome. Lonberg-Holm K and Philipson L (1974) PCR See polymerase chain reaction . Monogr Virol 9 , 148pp peacockpox virus (PKPV) A tenta- penguinpox virus (PEPV) A tentative tive species in the genus Avipoxvirus . species in the genus Avipoxvirus iso- Isolated from captive peacocks, Pavo lated from jackass penguins, Speniscus cristatus sp, in Baghdad Zoo in 1978 demercus . during an outbreak in which there was 75% morbidity and 13% mortal- Stannard LM et al (1998) J Gen Virol 79 , 1637 ity. Pathogenic for chickens infected experimentally. Antigenically related to pentamers Protein units on the virion fowlpox and pigeonpox virus. surface which in groups of five form Al Fallugi MM et al (1979) J Wildl Dis 15 , 597 pentons. penton 362 penton Group of five protein units at the peptide vaccines Synthetic peptides were 12 vertices of a virus capsid with icosa- produced initially in 1963 and thought hedral symmetry. In members of the to have great promise as vaccines, as Adenoviridae , the pentons are complex the development of gene sequence structures with base, fiber and knob, analysis led to precise descriptions of and are involved in cell attachment. the amino acid sequences of presumed See icosahedral symmetry . antigenic binding sites (epitopes). Although clear evidence was obtained Pependana virus A probable species that protection against a major animal in the genus Orbivirus , isolated from pathogen, foot and mouth disease phlebotomine sand flies in the Amazon virus, could be obtained by peptide region of Brazil. Antigenically related immunization, albeit only at very high to Changuinola virus . Not associated peptide doses in cattle, other efforts with disease in humans. with human disease-inducing viruses have met with less success. Peptides peplomers Knob-like structures (spikes) are poorly immunogenic, do not nec- projecting from the surface envelope essarily adopt the same configuration of a virus particle. They may have as the natural epitope, and seem to hemagglutinating activity and act as give rise to binding but not neutral- cell receptors, or have enzyme activity izing antibodies. A number of ways to such as neuraminidase. They are sur- improve the response have included face antigens. coupling to peptide carrier molecules incorporating T-cell epitopes, but so far these remain at the stage of develop- peplos (Greek: peplos an outer robe or mental research. cloak as worn by women in ancient Greece.) Synonym for envelope. Meloen RH et al (1995) Vaccine 13 , 885 peptabody A new type of high-avidity peptidyl transferase An enzyme, part of binding protein. A short peptide lig- the 50S subunit of the ribosome, which and was fused via a semi-rigid hinge forms the peptide bonds between region with the coiled-coil assembly amino acids. domain of cartilage oligomeric matrix protein to produce a multivalent bind- Peramivir A potential neuraminidase ing molecule. inhibitor that appeared to be more effective in inhibiting influenza virus Terskikh AV et al (1997) Proc Natl Acad Sci growth in cell culture than Relenza or 94 , 1663 Tamiflu. However clinical trials of the drug were disappointing, and it is not peptide A compound of two or more yet licensed for use. amino acids joined by peptide bonds. perch hyperplasia virus (PHV) A tentative peptide bond A chemical link formed species in the genus Epsilonretrovirus . by the reaction between the carboxylic Sequence analysis of the pol gene acid group of one amino acid and the showed a phylogenetic similarity to amino group of another, thus uniting the type species, Walleye dermal sarcoma the carbon atom of one with the nitro- virus . gen atom of the other. perch iridovirus An iridovirus was iso- peptide fusion inhibitors Some small lated from pike-perch, Stizostedion molecule inhibitors have been designed lucioperca, fingerlings with no sign of to inhibit fusion by the F protein of disease. paramyxoviruses for development Tapiovaara H et al (1998) Dis Aquat Org 32 , as antiviral agents. However though 185 promising, this approach has not so far yielded any licensed drugs. perch rhabdovirus An unassigned mem- de Clercq E (2002) Mini Rev Med Chem 2 , 163 ber of the genus Lyssavirus , family 363 periodic disease

Rhabdoviridae . Isolated from yearling Table P1. Viruses causing perinatal infection perch, Perca fluviatilis, in France. Causes a CNS disorder with low mortality in Cytomegalovirus young perch inoculated intracranially. Herpes simplex virus 1 and 2 Human immunodeficiency virus 1 and 2 Betts AM et al (2003) Dis Aquat Organ 57 , 201 Hepatitis B virus Dorson M et al (1984) J Fish Dis 7 , 241 Hepatitis C virus Hepatitis E virus percid herpesvirus 1 (PeHV-1) An unas- Human papillomavirus signed virus in the Herpesviridae fam- Varicella-zoster virus ily. First reported in 1980 in association with skin lesions (epidermal hyperpla- Enders G (2005) In Topley & Wilson’s Microbiology and Microbial Infections , vol. 2, sia) in spawning walleyes, Stizostedion Tenth edition, edited by BWJ Mahy and vitreum vitreum, in Canada. So far only V ter Meulen, London: Hodder Arnold, found in fish from Bad Carrot River, p. 1443 Saskatchewan. Significance in fish populations is unclear. Synonyms: Walleye epidermal hyper- of harmful viruses during passage plasia virus; herpesvirus vitreum. through the birth canal or from breast Yamamoto T et al (1985) Fish Pathol 20 , 361 milk. perdicid herpesvirus 1 (PdHV-1) An perinatal transmission Transmission of unassigned virus in the family Herpes- a virus to the products of conception viridae. Isolated in 1979 from the liver during delivery or shortly there- of bobwhite quail, Colinus virginianus , after. Occurs commonly if the mother is in Germany. The quail had probably infected with cytomegalovirus, herpes died from Clostridium colinum infection simplex virus, human immuno- (quail disease). A herpesvirus was iso- deficiency virus, hepatitis B, C, or E virus, lated from the birds in chick embryo human papillomavirus, or varicella- fibroblast cultures. Using neutraliza- zoster virus. tion tests, there was no cross-reaction between this virus and antisera against Perinet virus (PERV) A tentative species Marek’s disease virus, or herpesviruses in the genus Vesiculovirus. Isolated from from duck, turkey, chicken, parrot, owl, Culex antennatus in Madagascar. Could falcon, pigeon, cormorant, or stork. be the same as Andasibe virus. The only cross-reaction was with crane Clerc Y et al (1983) Ann Inst Pasteur Virol herpesvirus. The role of this virus in 134E , 61 disease in quail or other species is not known. perinuclear Situated in the region Synonyms : herpesvirus colinum; bob- between the two membranes of the white quail herpesvirus. nucleus or close to the outer mem- Kaleta EF et al (1980) Arch Virol 66 , 359 brane. Herpesviruses develop in the nucleus from immature capsids con- Pergamino virus A strain of Andes virus taining newly synthesized DNA which in the genus Hantavirus. Isolated in can then be observed budding through Pergamino, Argentina, from the rodent the inner nuclear membrane into the Akadon azarae . perinuclear space. Certain rhabdo - viruses which are thought to bud through perinatal infections Virus infections one nuclear membrane accumulate in acquired by the neonate during deliv- the perinuclear space; members of the ery or shortly thereafter. The placenta Reoviridae replicate in the cytoplasm provides a powerful barrier that allows near the outer nuclear membrane. transit of maternal IgG to the fetus whilst preventing a large number of periodic disease A syndrome involving infective agents from reaching the episodes of relatively benign symptoms embryo or the fetus. However, the (fever, lymphadenopathy) at intervals neonate may be exposed to a number of 2–3 weeks for more than 10 years. periodic disease 364

Associated with chronic Epstein–Barr Ahmed R and Chen ISY (1999) Persistent virus infection. Viral Infections. New York: John Wiley and Sons, 725pp Lekstrom-Himes JA et al (1995) Clin Infect Mahy BWJ et al (Editors) (1982) Virus Persist- Dis 22 , 22 ence, SGM Symposia, vol. 33. Cambridge: Cambridge University Press peripheral blood mononuclear cells ter Meulen V (1994) Semin Virol 5 , 259 (PBMCs) Human herpesviruses-6, -7, and -8 have all been isolated from personal protective equipment (PPE ) PBMCs, and unlike herpes simplex The equipment required to protect viruses, their interaction with CNS scientists from infection when work- cells is unclear. ing with hazardous viruses. This may include a mask that supplies high effi- permissive cells Cells in which replica- ciency particulate air (HEPA) filtration tion of a particular virus can take place. to the individual or (as in a biosafety level 4 ‘ suited ’ laboratory, a fully Peromyscus virus A possible species encapsulating suit supplied with air in the genus Paramyxovirus . Isolated from outside of the laboratory). from the pooled tissues of four wild white-footed mice, Peromyscus leucopus , Peruvian horse sickness virus (PHSV ) A trapped in Virginia, USA. Replicates species in the genus Orbivirus. Isolated in embryonated eggs and a variety of in 1997 from a horse with severe cell cultures with CPE. Infected cells meningo-encephalitis, from an out- hemadsorb guinea pig erythrocytes. break in which more than 100 horses Kills suckling mice on i.c. injection died. Then in 1999 the same virus was and hamsters on i.c. or i.p. injection. isolated in Australia from two horses Antibodies are found in some humans with neurological signs. A serologi- and in wild Peromyscus sp but not in cal survey of 411 stored equine sera laboratory mice or other wild animals. revealed 40 positive sera in horses from Morris AJ et al (1963) Proc Soc Exp Biol Med the Northern territories of Australia. 113 , 276 Sera that were antibody-positive to PHSV were also found in about half peroxidase An enzyme that catalyzes of the sera tested from fruit bats (black reactions in which hydrogen perox- and also red ﬂ ying foxes). A database ide is an electron acceptor. Used as a established at the Institute for Animal reporter molecule in immunodiagnos- Health, Pirbright, UK was used to show tic techniques such as ELISA. that several of the ten gene segments of the Australian virus were almost 100% persistent fraction Synonym for non- identical to the Peruvian virus. neutralizable fraction. Peste des petits ruminants virus (PPRV) persistent infection The term encom- A species in the genus Morbillivirus . passes a wide range of pathological Causes an important disease of sheep processes in cell cultures and whole and goats in West and Central Africa, animals. In general the term is best the Middle East and southern India. described as including infections in Similar to rinderpest but cattle are not which a degree of equilibrium is estab- affected. There is pyrexia with nasal lished between the virus and the host. and ocular discharge, necrotic stomati- In an animal host the chronic patholog- tis, and later severe enteritis and pneu- ical process may or may not progress, monia. Mortality may be over 90%. with or without fluctuation in sever- Antigenically related to distemper, ity. Infective virus is intermittently or measles, and rinderpest viruses but always recoverable. In cell cultures distinct from them. Rinderpest vaccine there are three types of persistent can be used to prevent this disease. infection: carrier cultures, steady-state Non-pathogenic for cattle, although infection, and a third in which the infection gives immunity to rinderpest viral genome is integrated with the cell virus. Replicates with CPE in a vari- genome. ety of cell cultures but sheep and goat 365 Peyer’s patches

cells are more sensitive. CPE develops Bovine diarrhea virus 1 . Other species 6–15 days after infection. Syncytia are in the genus are Bovine diarrhea virus 2 , formed. There are cytoplasmic and Border disease virus , and Classical swine intranuclear inclusions. fever virus . As of 2008, PPR is present in the Middle Becher P et al (1997) J Gen Virol 78 , 1357 East and the Indian sub-continent Meyers G and Thiel H-J (1996) Adv Virus Res as far east as Bangladesh. It recently 47 , 53 spread through Afghanistan into van Rijn PA et al (1997) Virology 237 , 337 Central Asia (Uzbekistan, Tajikistan and Turkmenistan) and has now also pestivirus bovis See bovine diarrhea spread to Tibetan China. Historically virus . associated with West Africa where it was first described, the virus is actu- pestivirus diarrhea virus See Bovine ally distributed across the continent diarrhea virus , diarrhea virus of from west to east. The virus is cur- bovines , and mucosal disease virus . rently spreading south from Sudan and Ethiopia into Kenya and Uganda. On pestivirus of giraffe A tentative species the North African coast the infection is in the genus Pestivirus . now reported from Morocco. Based on the understanding that rinderpest did pestivirus ovis See Border disease virus . not commonly infect small ruminants, and that following the recent demise pestivirus suis See hog cholera virus . of rinderpest a clearer picture of PPR’s distribution has emerged, many epi- Petevo virus (PETV) A serotype of Palyam demics historically ascribed to rinder- virus in the genus Orbivirus . Isolated pest were very likely due to PPR. On from the tick, Amblyomma variegatum , this basis it is highly likely that PPR in Central African Republic. Only one evolved on the Asiatic landmass and virus isolation was made from more only spread to Africa later. than 2500 pools of several tick species. Synonyms : Kata virus; pseudorinderpest Saluzzo JF et al (1982) Ann Inst Pasteur Virol virus; stomatitis/pneumoenteritis 133E , 215 complex virus. Petuluma virus (FIV-P) A strain of Feline Bailey D et al (2005) Virus Res 110 , 119 immunodeficiency virus in the genus Lentivirus . Pestivirus A genus of the family Flaviviridae . Antigenically unrelated Peyer’s patches Secondary lym- to viruses in other genera of the fam- phoid organs named after Johann ily. No invertebrate host. Spherical Conrad Peyer, a Swiss anatomist. virions 40–60 nm diameter. The virion Lymphoepithelial follicles in the sub- envelope has 10–12 nm ring-like subu- mucosa of the small intestine. More nits on its surface. Contain a positive- prominent in the ileum than the jeju- strand RNA genome about 12.3 kb num. Contain lymphocytes, germinal long with no 3 poly A or 5 -terminal centers, and thymus-dependent areas. cap. There is a single large open read- B lymphocytes predominate in the gering frame which encodes a polypro- minal centers, and T lymphocytes are tein of about 4000 amino acids. Four found in the zones between follicles. structural proteins are encoded toward They are separated from the lumen of the 5 terminus: a basic nucleocap- the intestine by a single layer of colum- sid protein, p14; and three envelope nar epithelium containing specialized glycoproteins, gp48, gp25, and gp53. microfold (M)-cells which take up In addition, at least seven nonstruc- antigen from the lumen and transport tural proteins are formed by cleav- it into the lymphoid areas. The prin- age of the polyprotein. Only species cipal site for the induction of intesti- affecting domestic and wild mam- nal immune responses. Hypertrophy mals (pigs and ruminants) have so of Peyer’s patches is associated with far been recognized. Type species prion diseases and it appears that Peyer’s patches 366

Peyer’s patches represent the main phenotype The outward, observable portal of entry for orally administered characteristics of a virus determined by prions such as scrapie, and perhaps its genotype. also bovine spongiform encephalopathy. See M cells . phenotypic mixing The production, in a mixed infection, of progeny virus Pferdestaupe virus See Equine arteritis with phenotypic characters from two virus . genetically different viruses, but with a genome derived from only one of phage See bacteriophage . them. The phenomenon is particularly common in viruses which mature by phagocyte A cell which is capable of budding through the cell membrane. phagocytosis. In mammals these are The defective avian and mammalian neutrophils and macrophages. sarcoma viruses are usually phenotypi- cally mixed since their envelope proteins are coded for by a helper virus. phalacrocoracid herpesvirus 1 (PhHV1) Another form of phenotypic mixing An unassigned virus in the family is the formation of pseudotype virus, Herpesviridae . Isolated on the CAM where nucleocapsids of one virus are from a young little pied cormorant, enclosed in the envelope of another Phalacrocorax melanoleucos. Other birds virus. In an extreme case, where the and rodents are resistant. No evi- genome of one virus is enclosed in dence of pathogenicity for cormorants. a capsid coded for by another, it is Replicates on the CAM, producing described as transcapsidation (see pocks. adenovirus-SV40 hybrids) or genomic Synonyms: cormorant herpesvirus 1; masking. Phenotypic mixing has also Lake Victoria cormorant herpesvirus. been accomplished artificially by French EL et al (1973) Avian Pathol 2 , 3 physical encapsidation of the genome of mouse sarcoma virus with the enve- phasianid herpesvirus 1 Synonym for lope of feline leukemia virus by ultra- Gallid herpesvirus 1 . centrifugation of a mixture of the two viruses. phasianid herpesvirus 2 Synonym for Boettiger D (1979) Prog Med Virol 25 , 37 Gallid herpesvirus 2 . Fischinger PJ and O’Connor TE (1969) Science 165 , 714 pheasant adenovirus 1 (PhAdV-1) A virus Zavada J (1976) Arch Virol 50 , 1 in the genus Siadenovirus . Causes n-phenylacetoaminomethylene-DL-p- marble spleen disease in the ring- nitrophenylalanine An antiviral agent. necked pheasant, Phasianus colchicus . An amino acid-related compound. Can be transmitted experimentally to Prolongs survival time and reduces turkeys. splenomegaly in mice injected with Iltis JP and Daniels SB (1977) Infect Immun Friend leukemia virus. Reduces the 16 , 701 yield of Friend and Moloney leukemia virus in mouse cell cultures. pheasant leukosis virus A possible Mechanism of action not clear. species in the genus Alpharetrovirus . Fujita H et al (1979) J Natl Cancer Inst 62 , 565 Endogenous virus present in pheasant cells and capable of acting as helper phlebotomus fever serogroup An old virus for defective Rous sarcoma virus. grouping of viruses within the genus Envelope-characterized specificities Phlebovirus which included Rift Valley of the virus from ring-necked pheas- fever virus , Sandfly fever Naples virus , ants, Phasianus colchicus , were group F, and others which are serologically while those of the virus from golden related. pheasants, Chrysolophus pictus , were group G. phlebotomus fever viruses Six viruses Fujita DJ et al (1974) Virology 60 , 558 (Alenquer, Candiru, Chagres, Punta 367 phosphatase

Toro and Sandfly fever viruses of phocid distemper virus See Phocine dis- Naples, and Sicilian types) can cause temper virus . phlebotomus fever in humans, although only SF-Naples virus and Phocid herpesvirus 1 (PhoHV-1) A spe- SF-Sicilian virus have caused large out- cies in the genus Varicellovirus iso- breaks. They are antigenically distinct lated from harbor seals, Phoca vitulina , from each other. Isolated from humans during an outbreak of pneumonia in Italy, Egypt, Iran, and Pakistan. and focal hepatitis in a seal orphan- Cause short, sharp fever after an incu- age in The Netherlands in which half bation period of 2–4 days. Fever may the seals died. Subsequently, related be recurrent, there is infection of the viruses were isolated from harbor seals conjunctiva and pain in the eyes, head, in Germany and the USA, and from the back, and limbs. Gastro-intestinal European gray seal, Halichoerus grypus . symptoms occur. There is leukope- Synonym : harbor seal herpesvirus. nia. No fatalities have been reported. Osterhaus ADME et al (1985) Arch Virol 86 , Not known to cause disease except 239 in humans. The sandfly, Phlebotomus papatasi, is the vector. Mouse-adapted phocid herpesvirus 2 (PhHV-2) A group virus given i.d. to humans produces of herpesviruses were isolated from immunity but no disease. However, seals, Phoca vitulina , and a Californian prevention is usually by control of the sea lion, Zalophus californicus . These vector. Replicate in human, mouse, and all bear closest sequence relation- hamster kidney cell cultures with CPE. ship to members of the subfamily Diagnosis confirmed by rising antibody Gammaherpesvirinae . titer or by virus isolation from blood in early stages of disease. Harder TC et al (1996) J Gen Virol 77 , 27 Synonyms : hundskrankheitvirus; pappataci fever viruses; sandfly fever Phocine distemper virus (PDV) A spe- viruses. cies in the genus Morbillivirus , w hich appears to be endemic in the harp seal Phlebovirus A genus in the family (Phoca groenlandica ). Caused an epi- Bunyaviridae containing nine species, zootic in harbor seals, Phoca vitulina , Bujaru , Candiru , Chilibre , Frijoles , Punta in the Baltic and North seas in 1988. Toro , Rift Valley fever , Salehabad , Sandfly Related to canine distemper virus, fever Naples, and Uukuniemi viruses . about 70% by nucleotide sequence The surface morphology is distinct analysis. Another outbreak occurred in in having small round subunits with 2002. Sampling of Scottish gray seals a central hole. Transmission may be (Halichoerus grypus) showed that many by flies, mosquitoes or ticks, depend- gray seals were infected with PDV ing on the species. The nine species but showed no obvious symptoms of are based upon antigenic groups, but infection, so may be an important vec- all Phleboviruses have common cod- tor species for transmission of PDV ing and transcriptional strategies. The to harbor seals, with which they are negative-stranded RNA genome seg- sympatric. ments have a 3 -terminal UGUGUUUC Synonym : seal distemper virus. and 5 -terminal ACACAAG sequence. Barrett T et al (1993) Virology 193 , 1010 The S RNA is ambisense and encodes Hammond JA et al (2005) J Gen Virol 86 , 2563 the N protein in negative sense and a Kennedy S (1998) J Comp Pathol 119 , 201 nonstructural protein (NSs) in positive Mahy BWJ et al (1988) Nature 336 , 115 sense. Visser IKG et al (1993) J Gen Virol 74 , 631

Phnom Penh bat virus (PPBV) A spe- phocine morbillivirus See Phocine dis- cies in the genus Flavivirus in the Rio temper virus . Bravo virus group. Isolated from bats in Cambodia. No known arthropod phosphatase An enzyme that catalyzes vector. Not reported to cause disease in the hydrolysis and synthesis of phos- humans. phoric acid esters and the transfer of phosphatase 368

phosphate groups from phosphoric widely used mechanisms for transduc- acid to other compounds. ing messages from receptor interactions at cell surfaces to effector molecules in phosphatidylserine A lipid molecule that the cell nucleus. has been proposed to be a receptor for vesicular stomatitis virus. phosphotungstic acid A negative stain Schlegel R et al (1983) Cell 32 , 639 used for electron microscopy of viruses. It consists of dodecatungstophosphoric phosphodiester bond Link formed acid dissolved in water to give a 1–2% between the nucleotides of polynucle- solution and adjusted to about pH 7 otide chains by covalent bonding of the with NaOH. phosphoric acid with the 3 -hydroxyl photodynamic inactivation Inactivation group of one ribose or deoxyribose of viruses by visible light in the pres- molecule and the 5 -hydroxyl group of ence of certain photoreactive dyes: acri- the next ribose or deoxyribose ring. See dine orange, acriflavine, brilliant cresyl nucleic acid . blue, methylene blue, neutral red, pro- phospholipase An enzyme that catalyzes flavine, and toluidine blue. The dye is the hydrolysis of a phospholipid, e.g. able to pass through the cell membrane lecithinase that acts on lecithin. Used and become associated with the DNA. in the study of viral membranes. The antiviral effect is seen when light energy absorbed by the dye causes phospholipids Components of many photochemical oxidation of any viral viruses, but especially the members of DNA with which it has become associ- the order Mononegavirales , which are ated. Clinical use limited to treatment composed of up to 25% lipids, of which of superficial herpetic lesions. 60% are phospholipids. Myers MG et al (1975) N Engl J Med 293 , 945 Yen G and Simon EH (1978) J Gen Virol 41 , phosphonoacetic acid (PAA) An antiviral 273 agent, used as the sodium salt. Inhibits replication of some DNA viruses by photoreactivation The repair of UV- blocking DNA synthesis. Superseded inactivated viral DNA by cellular by phosphonoformate. enzymes activated by exposure to long- wave light. Such enzymes are found in Felsenfeld AD et al (1978) Antimicrob Agents Chemother 14 , 331 the cells of bacteria, birds, and frogs but not placental mammals. phosphonoformate See trisodium Pfefferkorn ER and Boyle MK (1972) J Virol phosphonoformate (foscarnet). 9 , 474 Synonym : fosfonet. Photo-Shootur virus A synonym for phosphonoformic acid Foscarnet, a Camelpox virus . direct inhibitor of the DNA polymerase phylogenetic tree A diagrammatic rep- of herpes simplex virus. resentation of the interrelationships phosphoprotein A protein that has had between biological attributes of organ- one or more amino acids phosphory- isms or genes. For viruses, it is usu- lated by a protein kinase. The amino ally based on genome nucleic acid acids most commonly phosphorylated sequences. The relationship between are serine, threonine, and tyrosine. the common ancestor, intermediate ancestors, and contemporary species phosphorylation An important post- are depicted as the root, branches, and translational modification of viral leaves of a tree. See parsinomy . proteins. A mechanism for inducing Synonym : cladogram. reversible conformational changes in Hillis DM et al (1993) Meth Enzymol 224 , 456 proteins, so enabling them to act as molecular switches. Cascades of phos- phylogeny The relationship between bio- phorylation and dephosphorylation are logical lineages by common descent. 369 Picornaviridae

Viruses change through generations Picodna virus A name proposed but not over time by insertion, deletion, or adopted for the family Parvoviridae . substitution of genome nucleotides, and the resultant mutations may be advantageous, neutral, or deleteri- Picola virus (PIAV) A serotype of ous to survival and replication of the Wongorr virus in the genus Orbivirus . virus. Therefore a mutant virus line- Isolated from Culex annulirostris mos- age may proliferate in a particular quitoes in Victoria, Australia. Not environment or else become extinct. known to cause disease in humans. Phylogenetic analysis compares nucleotide sequences of viruses to determine Picornaviridae (Latin pico small) A their relationship, and the results are family of naked, ether-resistant viruses usually expressed in the form of a phy- with icosahedral capsids 22–30 nm in logenetic tree. diameter. The capsid is composed of physical particle:plaque-forming unit several different polypeptides whose ratios Not all virus particles are capa- apparent size can vary between closely ble of forming a plaque and for most related strains, but whose aggregate animal viruses the particle:pfu ratios molecular weight lies between 80,000 are seldom lower than 10 and some- and 120,000. Typically there are equal times much higher. amounts of four major capsid polypeptides, three (1B, 1C, and 1D) of mol. phytohemagglutinin A plant-derived wt. 20,000–40,000 and one (1 A) of lectin that agglutinates mammalian mol. wt. 5000–10,000. Proteins 1A, 1B, erythrocytes, and is a mitogen that 1C, and 1D are commonly known as stimulates predominantly T lympho- VP4, VP2, VP3, and VP1, respectively. cytes. The most commonly used phyto- Proteins 1B, 1C, and 1D each pos- hemagglutinin is derived from the red sesses a core structure comprising an kidney bean, Phaseolus vulgaris . eight-stranded beta sandwich (BETA- barrel). One molecule of each makes up Phytoreovirus A genus of the family the capsid structural unit. The capsid is Reoviridae containing species which composed of 60 structural units, with infect plants and insect vectors. T 1, pseudo T 3 icosahedral symmetry. The genome RNA of 7–8.5 kb Pichinde virus (PICV) A species in the in length is infectious. It is the mes- genus Arenavirus . One of the New sage for protein translation and car- World arenaviruses, belonging to the ries a poly A tract at the 3 end added Tacaribe serogroup. Isolated from transcriptionally. A small protein, VPg, the rodents, Oryzomys albigularis and is linked covalently to the 5 termi- Thomasomys fuscatus, mosquitoes of nus. Virus multiplication occurs in the Ixodes sp and mites of Gigantolaelaps sp cytoplasm and functional proteins are in Colombia. Not reported to cause dis- mainly produced by processing and ease in humans. post-translational cleavage of a nas- Leung WC et al (1979) J Virol 30 , 98 cent precursor polyprotein. There are nine genera: Aphthovirus , Enterovirus , picobirnavirus A virus first identified Erbovirus , Cardiovirus , Hepatovirus , in rat feces and later in humans and Kobuvirus , Parechovirus , Rhinovirus , in other animals. Contains a genome and Teschovirus. Distinguished by consisting of two small segments of sensitivity to acid, buoyant density of double-stranded RNA, 2.6 and 1.5 kb in the virion, genome structure, and clini- length. A probable cause of diarrhea in cal features of infection in susceptible humans. A study of 150 animal species hosts. in a zoo in Argentina showed widespread infection of the animals, but not Hyypia T et al (1997) J Gen Virol 78 , 1 associated disease. Minor P (2005) In Topley & Wilson’s Microbiology and Microbial Infections , vol. Gallimore CI et al (1995) J Med Virol 45 , 135 2, Tenth edition, edited by BWJ Mahy and Masachessi G et al (2007) Arch Virol 152 , 989 V ter Meulen, London: Hodder Arnold, p. 857 picornavirus epidemic conjunctivitis 370 picornavirus epidemic conjunctivitis pigeon adenovirus (PiAdV) A tentative virus A name proposed for acute hem- species in the genus Aviadenovirus . orrhagic conjunctivitis virus since the Isolated from pigeons with inclusion severity and frequency of the sub- body hepatitis. The virus kills 1-day- conjunctival hemorrhages is very old specific pathogen-free chicks within variable. This term would include con- 3 days. Adult racing pigeons were junctivitis due to coxsackie virus A24. resistant. The genome DNA restriction Not adopted. enzyme profile resembles that of fowl adenovirus D. Lim KH and Yin-Murphy M (1977) Singapore Med J 18 , 41 Wang CH and Chang CM (2000) J Vet Med Sci 62 , 989 pig cytomegalovirus See suid herpesvirus 2 . pigeon circovirus (PiCV) A virus similar to but distinct from beak and feather disease virus that has been diagnosed pig herpesvirus 1 See pseudorabies virus . retrospectively by electron microscopy and histological examination pig herpesvirus 2 See suid herpesvirus 2 . in 12 pigeons from the USA, 4 from Australia, and 1 from Canada that died pig infertility virus May be the same as from 1986 to 1993. Inclusions consist- Porcine parvovirus . ing of paracrystalline arrays of tightly packed, non-enveloped icosahedral vir- pig papillomavirus A possible species ions 14–17 nm in diameter were found in the genus Papillomavirus . Isolated in splenic, bursal, gut-associated, and from swine pancreatic trypsin in a line bronchus-associated lymphoid tissue of pig kidney cells. Causes a slowly macrophages and in bursal epithelial developing CPE over 3 weeks. Infected cells. cells can be identified by immunofluo- Woods LW et al (1994) J Vet Diagn Invest 6 , rescence using guinea pig antiserum. 156 Antiserum to SV40 or polyomavirus did not react with infected cells. Virion pigeon herpesvirus See columbid her- diameter is 36–44 nm. Replicates in pri- pesvirus 1 . mary embryo pig kidney and a pig kidney cell line. Pigeonpox virus (PGPV) A species in the Newman JT and Smith KO (1972) Infect genus Avipoxvirus . Highly pathogenic Immun 5 , 961 for pigeons, but produces a milder disease in chickens or turkeys. Attenuated pig pox virus See Swinepox virus . mutant viruses have been isolated from naturally infected pigeons, and these appear to provide some protection pig rotaviruses See porcine rotavirus . when tested in challenge experiments using fully virulent pigeonpox virus. pig’s blue eye disease Viral-induced disease of swine with characteristic Tanatawi HH et al (1979) Acta Virol 23 , 249 corneal opacity accompanied by neurological, reproductive, and respiratory pike fry rhabdovirus (PFRV) A tenta- alterations. See Porcine rubulavirus . tive species in the genus Vesiculovirus . Synonym: La-Piedad-Michoacan-Mexico Causes hemorrhagic lesions in the mus- virus (LPMV). cles and kidneys of young pike, Esox lucius , and red swollen areas are visible on the trunk, usually above the pelvic pig-tailed macaque parvovirus A species fins. Caused a severe fatal disease in in the genus Erythrovirus. Isolated from Dutch fish hatcheries. Isolated from pig-tailed macaques (Macaca nemes- diseased pike fry in FHM cells. CPE trina ) with anemia following experi- evident in 40 h. Pathogenic on injection mental immunosuppression. into young pike. In the presence of Green SW et al (2000) Virology 269 , 105 complement it can be neutralized 371 plaque-forming units (pfu)

by antiserum to spring viremia of carp Pirodavir An antiviral compound that virus. inhibits picornavirus replication by Synonyms : grass carp rhabdovirus; red binding to a hydrophobic pocket of disease virus; hydrocephalus of pike capsid protein VP1 beneath the canyon virus. floor and inhibiting binding or uncoat- Ahne W (1975) Arch Virol 48 , 181 ing of the virion. de Kinkelin P et al (1973) Nature 241 , 465 Andries K et al (1992) Antimicrob Agents Chemother 36 , 10 pike type C oncovirus A possible species in the genus Epsilonretrovirus . The Piry virus (PIRYV) A species in the genus Northern pike, Esox lucius, suffers epi- Vesiculovirus . Antigenically related to zootics of a benign lymphosarcoma. vesicular stomatitis virus. Isolated from There are cutaneous lesions and there an opposum in Para, Brazil. Laboratory is evidence of horizontal transmission human infections have resulted in a during spawning. A C-type virus and febrile illness with myalgia, arthralgia, reverse transcriptase have been demon- and abdominal tenderness. strated in the tumor cells. Tumors often regress during the summer season. Piscivirus A proposed name, not adopted, Sonstegard RA (1976) Nature 261 , 506 for the genus of fish rhabdoviruses Yamamoto T et al (1984) Arch Virol 79 , 255 now known as Novirhabdovirus . pilot whale morbillivirus The long- Pixuna virus (PIXV) A species in the genus finned pilot whale (Globicephala melas ) Alphavirus . Closely related to and perhaps is host to a novel cetacean morbilli- a serological type of Venezuelan virus, distinct from the dolphin or por- equine encephalitis virus. Isolated from poise morbilliviruses. The virus has rodents, Proechimys guyannensis oris , caused considerable mortality to pilot and mosquitoes in Brazil. No known whales in the Mediterranean Sea. association with human disease. Taubenberger JK et al (1996) Emerg Infect Dis Shope RE et al (1964) Am J Trop Med Hyg 13 , 2 , 213 723 PK (15) cells (CCL 33) A heteroploid pink-eye virus See Equine arteritis virus . cell line from the kidneys of a pig, Sus scrofa . Pindobaí virus A possible species in the genus Orbivirus, isolated from phle- Pl 1 Ut (NBL-9) cells (CCL 74) Initiated botomine flies in the Amazon region from the trypsinized uterine tissue of of Brazil. Antigenically related to an adult female raccoon, Procyon lotor . Changuinola virus. Not associated with disease in humans. plaque The area of lysis or ‘hole ’ formed in a lawn of cells due to infection with pinocytosis See viropexis . a single infectious unit of a cytopathic virus. Piratuba virus A possible species in the genus Orbivirus , isolated from mosqui- plaque assay An assay in which the toes in the Amazon region of Brazil. concentration of infective particles in Antigenically related to Changuinola a virus solution is determined as the virus . Not associated with disease in number of plaques induced on a lawn humans. of bacteria or eukaryotic cells. It can also be used to distinguish between Pirital virus (PIRV) A species in the different strains or distinct viruses by genus Arenavirus isolated in Venezuela. the features of the plaque. Natural rodent host is the cane rat, Sigmodon alstoni. Not known to cause plaque-forming units (pfu) The number human disease. Genetically related to of plaques formed per unit of volume Tacaribe complex viruses. or weight of a virus suspension. plaque mutants 372 plaque mutants Mutants producing vesicles. These migrate within the plaques different in size or appearance cytoplasm toward the nucleus becom- from those produced by the wild-type. ing acidified during the process. The Plaque size may be affected by speed reduced pH within the endosomal vesi- of replication, sensitivity or resistance cles triggers conformational changes in to inhibitors in the agar, and pH. the haemagglutinin (HA). The globular head domains of the HAS move away plaque neutralization test (plaque from the central stalk region, which is reduction test) A method either for reorganized into an elongated alpha identifying a virus (or serotype) or for helical structure terminating in a short titrating an antiserum by analyzing the hydrophobic region, the fusion pep- inhibitory effect of antibodies on the tide. Insertion of this peptide into the infectivity of the virus using the plaque membrane of the endosomal vesicle assay. and subsequent conformational change brings the viral and cellular mem- plaque picking The selection of indi- branes into close proximity and initiate vidual plaques which are formed by the fusion of the two membranes. a single infection event. Clones of a virus can thus be selected and further Weissenhorn W et al (1999) Mol Membr Biol studied. 16 , 3 plaque-reduced neutralization (PRN) Plasmaviridae A family of viruses which See plaque neutralization test. infect mycoplasma. plasma-derived vaccine The produc- plasmids Genetic elements composed of tion of inactivated hepatitis B vac- double-stranded circular DNA which cine derived from the blood plasma of replicate separately from the bacte- hepatitis B virus carriers is an impor- rial chromosome within the bacte- tant source of hepatitis B vaccine used rial cell wall. Some can induce their in millions of recipients worldwide, direct transmission to other bacteria, without any known transmission of although they differ from viruses in virus. Although most developed coun- having no extracellular infective parti- tries have now adopted the genetically cle. Some plasmids are under stringent engineered recombinant hepatitis B control and as little as one copy is repli- vaccine made in yeast or mammalian cated per genome. Others, under more cells as standard because of its proven relaxed control, replicate many copies reliability and absolute safety, the high per cell. They may carry genetic deter- cost means that many countries can minants which can be translocated only afford to immunize with plasma- from the plasmid to the bacterial chro- derived vaccine. mosome. Some of these determinants mediate antibiotic resistance. See also plasma membrane The external lipid episomes . bilayer membrane of cells. Enveloped Lederberg J (1952) Physiol Rev 32 , 403 viruses transfer their nucleocapsids across membranes by fusing with plasmid vector A plasmid which is used them, resulting in simultaneous enve- for cloning ‘ foreign ’ DNA. The plas- lope removal and cell entry. mid is often manipulated to contain desirable features such as resistance to plasma membrane fusion The process two or more antibiotics, ability to pro- by which enveloped viruses gain entry duce multiple copies, single-cutting into the host cell. The best studied is restriction enzyme sites, and strong influenza virus. Binding of the virus to promoters. host-cell receptors initiates a process of endocytosis which draws the attached platelet derived growth factor (PDGF ) virus particles into clathrin-coated The major mitogen in serum for growth pits. The pits invaginate into the cell in culture of cells of connective tissue and are pinched off to form endosomal origin. Consists of two different but 373 pneumonia virus of rats

homologous peptides A and B. The B occurs. The only signs of infection are chain is almost identical in sequence to pathological changes in epithelial cells the transforming protein of SV40 virus. of the skin and gills where giant cells Appears to be involved in wound heal- are seen containing herpesvirus-like ing. Serves as the primary receptor for particles. Virus isolation has not been the parvovirus AAV5. reported. Di Pasquale G et al (2003) Nat Med 9 , 1306 Synonyms : herpesvirus scophthalmus; turbot herpesvirus. plating efficiency See efficiency of Buchanan JS and Madeley CR (1978) J Fish plating . Dis 1 , 283 Buchanan JS et al (1978) Vet Rec 102 , 527 Hellberg H et al (2002) Dis Aquat Organ 49 , 27 Playa de Oro virus (PDOV) A probable species in the genus Hantavirus . Identified by cloning the S and M seg- plus strand The RNA strand which ments of the hantavirus from Oryzomys acts as messenger RNA. See positive couesi rodents in Mexico. The sequences strand . of these segments showed the closest relationship to Bayou and Catacamas Plymouth virus A strain of ‘ classic ’ viruses. human calicivirus from a 16-month-old girl. Identified by genomic sequence Chu YK et al (2008) Virus Res 131 , 180 analysis. Playas virus (PLAV) A serotype of Buny- Liu BL et al (1995) Arch Virol 140 , 1345 amwera virus in the genus Ortho- bunyavirus. Isolated from mosquitoes, PML-2 virus See JC polyomavirus . Aedes taeniorhynchus , in Ecuador. Not known to cause disease in humans. PMSF (phenyl methanesulfonyl fluoride , α -toluenesulfonyl fluoride) An pleconaril An antiviral compound that inhibitor of proteases. Used in studies inhibits picornavirus replication in vitro on viral proteins where proteolysis is and is active in mice by oral admin- to be avoided. istration. Binds to the hydrophobic pocket of the virion capsid. Clinical tri- pneumonia virus of mice (PVM) See als against enterovirus and rhinovirus Murine pneumonia virus . infections are complex, and not all rhinovirus species react in the same way to pneumonia virus of rats This very com- the drug. mon chronic pulmonary disease of old Barrett B et al (2007) Ann Fam Med 5 , 216 laboratory rats probably has no specific Ledford RM et al (2005) Antiviral Res 68 , 135 cause, but a multiple etiology in which mycoplasma play a part. The many pleocytosis Increased leukocyte count in descriptive names given to chronic a bodily fluid, such as cerebrospinal lung disease in rats do not describe fluid. Commonly occurs in measles specific diseases but rather clinical find- virus infection and also in more than ings. Disease starts as a silent pneumo- 50% of cases of mumps virus infection. nia and progresses slowly, often with the formation of pus-filled cavities and pleurodynia See Bamble disease . fibrosis. The nose and middle ears may be mainly involved. Pleuropneumonia- pleuronectid herpesvirus (PiHV-1) like organisms and bacteria, as well as An unassigned virus in the family mycoplasma, are regularly isolated. Herpesviridae . First recognized in 1978 Synonyms: atypical pneumonia of rats; among young turbot, Scophthalmus endemic pneumonia of rats; enzootic maximus , in a fish farm in Scotland, bronchiectasis of rats. but since then also recognized in Wales Brennan PC et al (1969) Lab Anim Care 19 , and in Norway. The fish develop ano- 360 rexia and lethargy and heavy mortality Nelson JB (1946) J Exp Med 84 , 7, 15 Pneumovirinae 374

Pneumovirinae A subfamily in the fam- virus infection, the fraction of cells (P ) ily Paramyxoviridae with two genera, that actually receive any given number Pneumovirus and Metapneumovirus . (k ) of infectious virus particles at an Differ from members of the subfamily average multiplicity of infection (m ) is Paramyxovirinae in having 10 sepa- described by the Poisson equation as: rate transcriptional units, extensive P ( k ) ( e m m k )/ k ! O-linked glycosylation of the G protein, a smaller nucleocapsid diameter polarized epithelial cells The plasma and longer surface peplomers. membrane of epithelial cells is divided Pneumovirus A genus of the subfamily into two discrete domains with dis- Pneumovirinae. Virions contain no tinct protein and lipid compositions. neuraminidase. Nucleocapsid about Tight junctions serve to preserve these 14 nm in diameter with a helical pitch domains and prevent mixing between of 7 nm. The negative-strand RNA them. Several cell lines maintain the genome encodes 10 separate genes. The properties of epithelial cells in culture glycoprotein (G protein) has extensive (polarized cells), and some viruses are O-linked glycosylation. Type species found to bud asymmetrically from Human respiratory syncytial virus . Other these cells, e.g. vesicular stomatitis, species are Bovine respiratory syncytial and retroviruses are released from the virus and Murine pneumonia virus . basolateral domain, whereas influenza virus, paramyxovirus, and respiratory Kingsbury DW et al (1978) Intervirology 10 , syncytial virus bud from the apical 137 domain. pock assay Viruses which replicate on the Tucker SP and Compans RW (1993) Adv CAM often produce local lesions called Virus Res 42 , 187 pocks, visible to the naked eye. Each one originates from a single infected pol gene One of the genes in the genome cell, so a count of the pocks produced of the retroviruses. Codes for the is a measure of the infective dose reverse transcriptase of the virus: the applied to the membrane. Analogous polymerase, hence the name. to a plaque assay. Katz RA and Skalka AM (1994) Annu Rev Pogosta disease Name given to an out- Biochem 63 , 133 break of disease in Finland caused by a variant of Sindbis virus. A Finnish pol II-dependent transcriptase The major name for the disease usually called cellular RNA transcriptase enzyme Karelian fever, caused by Ockelbo that makes mRNAs and defines gene virus. See Ockelbo virus . expression patterns within cells. point-of-care tests Rapid tests for virus Pol protein The reverse transcriptase pro- infection which can be carried out in a tein of HIV. physicians office. point-source outbreaks An outbreak of polio-encephalomyelopathy of mice disease, such as norovirus enteritis, in virus Synonym for Theiler’s murine which all case patients became infected encephalomyelitis virus. at the same place and time. poliomyelitis The disease caused by Poisson distribution An important sta- poliovirus. See Poliovirus . tistical concept used in virology to model experimental infections and to poliomyelitis virus type 4 A name used estimate the multiplicity of infection. It at one time for coxsackie virus A7. provides a distribution of the number of randomly occurring events over a Chumakov MP et al (1956) Probl Virol 1 , 16 given period of time; times between successive events are exponential ran- Poliovirus (PV) The type species of dom variables. In an experimental the genus Enterovirus. There are three 375 polycistronic messenger RNA

serotypes: human poliovirus 1, human occurs at the 3 end of eukaryotic cell poliovirus 2, and human poliovirus 3 mRNA (histone mRNA is an excep- (PV-1 to 3). Causes poliomyelitis, a dis- tion) and of most virus mRNAs which ease of great antiquity, and was one of have been studied, including the virion the first viruses to be isolated in 1909. RNA of most positive-strand viruses. Following infection the virus may It is believed to increase the stability of multiply in mucosal surfaces, tonsils, mRNA by making it more resistant to and Peyer’s patches of the small intes- nuclease digestion. tine before spreading to more distant lymph nodes and through viremia to polyacrylamide gel electrophoresis other sites including peripheral nerves (PAGE) A technique for separation of and spinal cord. The outcome of the nucleic acids or proteins present in a infection may be inapparent (90–95%); mixture. An important diagnostic tech- an abortive or minor illness (respira- nique for rotaviruses, where a sam- tory, gastrointestinal, or influenza-like) ple is separated by PAGE followed by from which the patient soon recov- silver-staining. The pattern of the sepa- ers (4–8%); non-paralytic poliomyeli- rated dsRNAs of the virus can be used tis, with symptoms of back pain and in outbreak investigations to distin- muscle spasm from which the patient guish the causative virus. soon recovers (1–2%); or paralytic poliomyelitis where following a minor polyadenylation The addition of ade- prodromal illness as above, the pre- nylate residues, usually to the 3 end of dominant feature is flaccid paralysis RNA molecules. resulting from lower motor neuron damage. Poliomyelitis is termed spi- poly A polymerase An enzyme which nal if the lower spine is involved or adds adenylate residues to the 3 end bulbar if the upper spine and brain of RNA. stem are involved. About 10% of cases, especially of bulbar poliomyelitis, are fatal and paralysis remains significant poly AU A synthetic double-stranded or severe in 80% of cases, with about polynucleotide. Under the influence of 10% of cases recovering with only polynucleotide phosphorylase, adenos- minor paralysis. Poliomyelitis was ine diphosphate molecules polymerize uncommon until standards of hygiene to form a polyribonucleotide contain- improved in the twentieth century. ing only adenine bases. This is usu- Formerly it was a common infection ally referred to as ‘ poly A.’ Similarly, and the presence of maternal antibody using uridine diphosphate, poly U can probably protected infants from neuro- be obtained. If equimolar amounts of logical disease. Once maternal infec- poly A and poly U are mixed in dilute tion became uncommon, infants were aqueous solution, they form a complex exposed to poliovirus and infantile in which the adenine and uridine bases paralysis became a major problem in link together to make a double strand, the Western world. Safe and effective known as ‘ poly AU. ’ polio vaccines were developed in the 1950s, which have led to a major ini- poly C Polycytidylic acid. A stretch of tiative by WHO to eradicate poliovirus polycytidylic acid about 100 nt long, globally. See human polioviruses (PV- of unknown function, is found in the 1 to 3). genome RNA of some picornaviruses. In foot-and-mouth disease virus there poliovirus hominis See Poliovirus . is a poly C tract about 360 nt from the 5 -terminus of the genome RNA. It polioviruses 1–3 See human polioviruses varies in length from 100 to 400 nt, but (PV-1 to 3) its exact function is unclear. polka fever virus See dengue viruses 1–4 polycistronic messenger RNA An mRNA poly A Polyadenylic acid. A stretch of which contains the coding sequences polyadenylic acid up to 300 bases long for two or more proteins. polyclonal antibody 376 polyclonal antibody A preparation con- polynucleotide kinase An enzyme iso- taining antibodies against more than lated from bacteriophage T4-infected one epitope of an antigen. Escherichia coli, which transfers a phosphate group from ATP and phospho- Polydnaviridae A family of viruses rylates the 5 -OH termini of RNA or affecting invertebrates whose genome DNA chains. Used experimentally to has multiple double-stranded DNAs label RNA prior to sequencing. of variable size and which infect para- Lockard RE et al (1978) Nucl Acids Res 5 , 37 sitic wasps in which the viral genome is integrated within the wasp genome. polynucleotide ligase Generic term for enzymes which catalyze the linking or polyhedral cytoplasmic deoxyribovirus repair of either DNA or RNA strands. See Iridoviridae . See DNA ligase and RNA ligase . poly I poly C A duplex of polyriboinos- Polyomaviridae A family of double- inic acid and polycytidylic acid, formed stranded DNA viruses with only in the same way as poly AU. A potent one genus, Polyomavirus. The family interferon inducer. description corresponds to the genus description. polymerase See DNA polymerase and Polyomavirus The only genus in the RNA polymerase . family Polyomaviridae. The type species is Simian virus 40 . Virions are non- polymerase-based viral classification enveloped 40 nm in diameter with 72 The sequence encoding the viral RNA- capsomeres arranged in a right-handed dependent RNA polymerase can be skew icosahedral lattice. The genome a valuable indicator of relationships is a single molecule of circular double- between viruses. stranded DNA which is 5243 bp in Zanotto PM et al (1996) J Virol 70 , 6083 length for the type species. Virus-specific proteins are encoded on both DNA strands. Several species hemaggluti- polymerase chain reaction (PCR) The nate by reacting with neuraminidase- selective amplification of DNA by sensitive receptors. Replication occurs repeated cycles of: (1) heat denatura- in the cell nucleus. During replication, tion of the DNA; (2) annealing of two transcription of the genome is divided oligonucleotide primers that flank the into early and late stages which are DNA segment to be amplified; and under the control of separate promoters (3) the extension of the annealed prim- and occur on opposite strands. During ers with the heat-insensitive Taq DNA replication 2–3 nonstructural protein polymerase. The product is termed antigens are expressed, which include the ‘ amplicon. ’ Can be used to deter- large T, middle T and small t for mouse mine the sequence of the amplicon, for and hamster polyomaviruses and large virus diagnosis, and in the amplifica- T and small t for other viruses such as tion of low copy number sequences. SV40, JC, and BK viruses. Replication Amplification of RNA can be achieved of the viral genome is initiated by bind- by first reverse transcribing the RNA to ing of the T antigen at a specific site on DNA, followed by PCR. This is referred the DNA and its interaction with host to as ‘ RT-PCR. ’ cell DNA polymerases that are used Barnes WM (1994) Proc Natl Acad Sci 91 , to replicate the DNA. In their natural 2216 hosts most species cause silent infec- Saiki RK et al (1988) Science 239 , 487 tions, but on injection into newborn animals (hamsters, mice, etc.) most polynucleotide A linear polymer of are oncogenic. Viral DNA is integrated nucleotide units linked by phosphodi- into the cellular DNA of transformed ester bonds betwe]en the 3 and 5 and tumor cells. There are five rec - positions on the sugar ring. Long poly- ognized human polyomaviruses: BK, nucleotides are nucleic acids. JC, KI, MC and WU. The most serious 377 Ponteves virus (PTVV)

of these is JC virus, which can infect and lineage and occupies a particular eco- destroy oligodendrocytes of the cen - logical niche. ’ tral nervous system, and in immunosuppressed patients (e.g. those with poly U Polyuridylic acid. See poly AU . AIDS) can cause fatal demyelinating disease known as progressive multi- Pongine herpesvirus 1 (PoHV-1) A spe- focal leukoencephalopathy (PML). So cies in the genus Lymphocryptovirus , far, no specific diseases have been isolated from lymphoid cell lines of the attributed to infection with BK, KI, chimpanzee, Pan troglodytes . A primate or WU polyomaviruses. MC virus is B-lymphotropic herpesvirus sharing found in the Merkel cell carcinoma, a 40% well-conserved DNA sequence rare neuroendocrine-derived skin can- relatedness with Epstein–Barr virus cer, more frequently found in immuno- (Human herpesvirus 4 ) and herpesvirus suppressed patients. MC polyoma papio (Cercopithecine herpesvirus 12). virus is clonally integrated into the All three viruses cross-react antigeni- tumor cells, and is believed to be the cally. Cultivation restricted to B cell cause of this cancer. lymphocytes. No evidence of clinical disease in chimpanzees. polyomavirus papionis 1 A polyoma- Synonyms: chimpanzee agent; chimpan- virus isolated from the baboon. See zee herpesvirus 1; herpesvirus pan. Baboon polyomavirus 2 . Gerber P et al (1976) J Virol 19 , 1090 Heller M et al (1982) J Virol 41 , 931 polypeptide A chain of amino acids linked together by peptide bonds Pongine herpesvirus 2 (PoHV-2) A species obtained by synthesis or by partial in the genus Lymphocryptovirus , iso- hydrolysis of a protein. Can also refer lated from a cell line established from a to the primary structure of a protein, leukemic orangutan, Pongo pygmaeus . e.g. polypeptide chain. Synonym : orangutan herpesvirus. polyprotein A large polypeptide that Rasheed S et al (1977) Science 198 , 407 gives rise to two or more proteins by Pongine herpesvirus 3 (PoHV-3) A spe- enzymatic cleavage. For example, the cies in the genus Lymphocryptovirus . A human poliovirus genome codes for virus associated with a B lymphoid cell a large protein which is subsequently line established from a gorilla, Gorilla cleaved to produce all the virus struc- gorilla. Cross-reacted in DNA hybridi- tural and nonstructural proteins. zation studies to 30–40% with EBV DNA. Transformed lymphocytes from polyribosomes See polysomes . gibbon apes, Hylobates lar , in vitro . No evidence that the virus causes disease polysomes Ribosomes attached to mRNA in the host species. at intervals of 50–10 nm. In the process Synonyms : gorilla herpesvirus. of protein synthesis the ribosomes pass Neubauer RH et al (1979) J Virol 31 , 845 along the mRNA strand, each forming a polypeptide chain as it goes. Pongola virus (PGAV) A sero- Synonym : polyribosomes. type of Bwamba virus in the genus Orthobunyavirus . With Bwamba polythetic class A class defined in terms virus forms the Bwamba serogroup. of a broad set of criteria that are neither Isolated from mosquitoes in South necessary nor sufficient. Each mem- Africa, Uganda, Ethiopia, Kenya, ber of the class must possess a certain Mozambique, and Central African minimal number of defining character- Republic. On injection kills newborn istics, but none of the features has to mice. Natural hosts sheep, cattle and be present in each member of the class. donkeys. Antibodies found in humans In the definition of a virus species but disease not reported. accepted by the ICTV in 1991, a virus species is defined as ‘ a polythetic class Ponteves virus (PTVV) A serotype of of viruses that constitutes a replicating Uukuniemi virus in the genus Phlebovirus . Ponteves virus (PTVV) 378

Isolated from a tick, Argas reflexus , in be widespread in pigs in Europe and southern France. Not reported to cause North America. An antigenically dis- disease in humans. tinct serotype (PCV-2) was isolated in France in 1997 from young pigs with Poovoot virus (POOV) A serotype of wasting disease and is associated with Great Island virus in the genus Orbivirus . a newly recognized disease known as Member of the Kemerovo serogroup. post-weaning multisystemic wasting Isolated from ticks, Ixodes uriae . Not syndrome (PMWS). Pigs with PMWS reported to cause disease in humans. show debility, dyspnea, palpable lym- Yunker CE (1975) Med Biol 53 , 302 phadenopathy, diarrhea, and pallor or icterus. The lesions can be reproduced Porcine adenoviruses A–C (PadV-A to C) experimentally in piglets, though their Three species in the genus Mastade- full expression may involve co-infection novirus. There are at least five with porcine parvovirus or PRRS virus. serological types. Some strains agglu- There is only about 80% homology tinate erythrocytes of several species. between the genomes of PCV-1 and Replicate in a wide range of cell cul- PCV-2. The precise genetic and pheno- tures (pig, cattle, dog, hamster and typic characteristics of these two cir- humans). Commonly found in the coviruses, and whether more than one digestive tract of pigs. On inoculation virus is involved in disease causation intranasally into colostrum-deprived requires further investigation. newborn pigs, the tonsils and lower Allan GM and Ellis JA (2000) J Vet Diagn intestine become infected but no symp- Invest 12 , 3 toms or diseases are produced. Meehan BM et al (1998) J Gen Virol 79 , 2171 Derbyshire JB (1989) In Virus Infections Niagro FD et al (1998) Arch Virol 143 , 1723 of Porcines , edited by MB Pensaert. Amsterdam: Elsevier, p. 73 porcine endogenous retrovirus (PoEV) A possible species in the genus Lentivirus . porcine adenovirus 1–5 (PadV-1 to 5) An Mitogenic activation of peripheral original serotype classification of por- blood mononuclear cells (PBMCs) from cine adenoviruses. Now porcine adeno- a minipig and a Yucatan pig resulted in viruses 1–3 are Porcine adenovirus A , 4 activation and release of an infectious is Porcine adenovirus B, and 5 is Porcine C-type virus that was shown to infect adenovirus C . both human and pig cell lines. The virus is of concern regarding the pos- Porcine astrovirus (PastV) A species in sible use of pigs as organ transplanta- the genus Astrovirus. There is a single tion donors for humans. Analysis of serotype, porcine astrovirus 1 (PastV-1). genome sequences of a range of por- Isolated from pigs with gastroenteritis. cine species suggests that many other The virus grows in a porcine embry- endogenous viruses remain to be onic kidney cell line. characterized. Shimizu M et al (1990) J Clin Microbiol 28 , 201 Paradis K et al (1999) Science 285 , 1236 Porcine circovirus 1 and 2 (PCV-1 and Patience C et al (2001) J Virol 75 , 2771 Wilson CA et al (1998) J Virol 72 , 3082 PCV-2) Species in the genus Circovirus , PCV-1 was first isolated in Germany in 1974 from a persistently infected PK15 porcine enteric sapovirus-Cowden A cell line. The DNA genome is 1.76 kb in tentative species in the genus Sapovirus , length. Does not cross-react antigeni- associated with diarrhea in young pigs, cally or by DNA sequence homology from which it was first isolated. Can with avian circovirus species (e.g. beak be propagated in primary pig kidney and feather disease virus or chicken cells or a pig cell line (LLC-PK-2 cells). anemia agent). Of various species Genome sequence analyses reveal tested, only domestic pigs, minipigs, a close relationship to human sapo- and wild boar were found to have anti- virus isolates, The virus has one major bodies to the virus, which appears to structural protein, mol. wt. 58,000, that 379 Porcine parvovirus (PPV)

does not cross-react with feline calici- Porcine hemagglutinating encephalomy- virus proteins. elitis virus (HEV) A group 2 species in the genus Coronavirus. There is only Wang QH et al (2005) J Clin Microbiol 43 , 5963 one serotype. Found in North America Parwani AV et al (1990) Arch Virol 112 , 41 and Europe, associated with outbreaks of encephalomyelitis in suckling pigs. porcine enteroviruses Eleven species in Antibodies have been reported in herds the genus Teschovirus , originally classi- in Australia and Japan. Most infec- fied into 13 serotypes of enterovirus on tions are subclinical and the economic the basis of cross-neutralization tests importance of the disease is low. There in cell culture. Using genome nucleo- is high mortality in young animals but tide sequence analysis, serotype 1 was older animals may survive although renamed porcine teschovirus 1, and all they grow less well. Adults may vomit porcine enteroviruses were reclassi- and eat poorly but recover or merely fied into a new genus called Teschovirus have a silent infection. Virus is found within the family Picornaviridae . in the respiratory tract but not in other Usually harmless inhabitants of the tissues. It may be excreted for up to 10 intestinal tract of pigs. Do not replicate days. No evidence of infection in utero . in cells from other species, and do not Virus replicates in primary pig kidney hemagglutinate. May cause Teschen cell cultures causing the formation of (or Talfan) disease (polioencephalo- multinucleate giant cells. Agglutinates myelitis) which varies in severity. rat, chicken, turkey, mouse, and ham- Synonyms : Ansteckende schweineläh- ster erythrocytes at 22°C. mung virus; enteric cytopathic porcine Synonyms: Canadian vomiting and orphan virus (ecpovirus); enteric cyto- wasting disease of pigs virus; hemag- pathic swine orphan virus (ecsovirus). glutinating encephalomyelitis virus of pigs; Ontario encephalomyelitis virus; Kaku Y et al (2001) J Gen Virol 82 , 417 vomiting and wasting disease virus of piglets. Porcine enteroviruses A and B (PEV-A Andries K and Pensaert MB (1980) Am J Vet and B) Species in the genus Enterovirus . Res 41 , 1372 PEVA is serotype 8 in the original clas- Mengeling WL and Cutlip RC (1976) J Am sification of porcine enteroviruses. Vet Med Assoc 168 , 236 PEV-B is comprised of serotypes 9 and 10 in the original classification of porcine enteroviruses. Porcine parvovirus (PPV) A species in the genus Parvovirus. A natural infection of pigs in most countries of the world. porcine enzootic pneumonia Not a virus Has been recovered from infertile pigs disease. A disease of swine caused by and aborted piglets. Causes repro- Mycoplasma hyopneumoniae . ductive failure with prenatal death but without maternal clinical disease. Porcine epidemic diarrhea virus (PEDV) Surviving piglets have virus in various A group 1 species in the genus organs up to 9 weeks of age. Has been Coronavirus which infects swine world- isolated from cell cultures of kidney wide including Europe, Australia, tissue from normal pigs. The mecha- and the USA, causing acute watery nism of pathogenicity is unknown. diarrhea. Symptoms are similar to Agglutinates chick, rat, guinea pig, cat, those of transmissible gastroenteritis rhesus, patas monkey and human O virus (TGEV), but develop more slowly, erythrocytes. Replicates in pig kidney with a longer incubation period, and cell cultures. Very similar or identical lowered mortality (up to 50%) in baby to KBSH virus. Antigenically different piglets. Older pigs usually recover from all other parvoviruses. An attenu- after a week. A sensitive ELISA test is ated strain (NADL-2) is used as a live available. Control is by sanitary meas- vaccine; it causes limited viremia and ures, since the virus is mainly spread does not cross the placenta. Complete by human and animal traffic. genome sequences of the vaccine strain Porcine parvovirus (PPV) 380

and a virulent field strain (Kresse) have eye) in pigs in Michoacan state, Mexico. been determined. The disease mainly affects piglets and is now endemic in Mexico. Infects cells Bergeron J et al (1996) J Virol 70 , 2508 Mengeling WL (1989) In Virus Infections through a sialo-glycoprotein receptor. of Porcines , edited by MB Pensaert. The structural proteins that have been Amsterdam: Elsevier, p. 83 sequenced are related to those of SV5 Zeeuw EJL et al (2007) J Gen Virol 88 , 420 and mumps viruses, and the pattern of transcriptional editing during P gene porcine poliomyelitis virus See Porcine expression, which involves insertion of teschovirus 1-11 . two non-templated G residues at a specific site to produce the P mRNA, also Porcine reproductive and respiratory occurs during mumps virus replication. syndrome virus (PRRSV) A species The complete genome sequence shows in the genus Arterivirus. Pigs are the the closest relationship to Mapuera only species affected. The disease was virus, isolated from a bat in Brazil. first reported in the USA in 1987, then Synonyms: La-Piedad-Michoacan-Mexico rapidly recognized in Europe and else- virus (LPMV); pig blue eye disease. where. Characterized by reproductive Mendoza-Magana ML et al (2007) Vet J failure, pneumonia in young piglets, 173 , 428 and increased preweaning mortal- Wang LF et al (2007) Arch Virol 152 , 1259 ity. The genome RNA is about 15 kb in length with eight open reading frames Porcine teschovirus (PTV) A species in (ORFs). Viruses isolated in Europe and the genus Teschovirus. Originally called the USA have quite distinct genotypes porcine enterovirus serotype 1, the and differences in pathogenicity, and only serotype to cause serious disease. within the USA at least three genotypes First recognized in 1930 in Teschen, in have been identified. the Czech Republic, and described as a Synonyms: porcine epidemic abortion virulent, highly fatal non-suppurative and respiratory syndrome (PEARS); encephalomyelitis in which lesions were porcine respiratory and reproductive present throughout the central nervous syndrome virus; Lelystad virus; blue system. Later, a less severe form of the ear disease; swine infertility and respi- disease was recognized in the UK and ratory syndrome virus. called Talfan disease after the name of a hill in Wales. The disease includes Halbur PG et al (1996) Vet Pathol 33 , 159 fever, convulsions, and paralysis which Nelsen CJ et al (1999) J Virol 73 , 270 may be permanent in survivors. There Rowland RR et al (1999) Virology 259 , 262 is a diffuse encephalomyelitis. Many outbreaks of severe disease caused by porcine respiratory coronavirus Teschen strains occurred in the 1930 to (PRCoV) A strain or serotype of 1950s, but since that time these highly Transmissible gastroenteritis virus in the virulent strains have been replaced by genus Coronavirus . less virulent Talfan-like strains. Disease is produced in no other species. Strains porcine rotavirus C/Cowden (PoRVC/ causing significant disease are all in one Cowden) A strain of Rotavirus C in the serological group which can be divided genus Rotavirus isolated in the USA into three subgroups: (1) includes from a suckling pig with diarrhea. Teschen (Konratice) and Bozen strains; Jiang B et al (2000) Virus Genes 20 , 193 (2) includes Talfan and Tyrol strains; and (3) includes Reporyje strain. The porcine rotavirus E/DC-9 (PoRV-E/DC9) serological subgroups are not corre- A strain of Rotavirus E in the genus lated with virulence. Virus replicates Rotavirus . with CPE in pig kidney cell culture and strains can be types T or V accord- Porcine rubulavirus (PoRV) A species in ing to the type of CPE. In general, does the genus Rubulavirus, isolated during not replicate in cells from other species. an outbreak of encephalitis, reproduc- Does not hemagglutinate. A vaccine has tive failure and corneal opacity (blue been used successfully. 381 positive strand

Synonym: infectious porcine encephalo- to demonstrate parvovirus as the etio- myelitis virus; infectious porcine polio- logical agent. myelitis virus; Talfan disease virus; Frelier PF et al (1984) J Am Vet Med Assoc Teschen disease virus. 185 , 1291

Porcine teschovirus 1–11 Eleven sero- porcupine papillomavirus A novel types of porcine teschovirus. Of these, papillomavirus was found in a papil- the most severe disease is caused by lomatous lesion of the skin of a north viruses of serotype 1, and also serotype American porcupine (Erethizon dorsa- 11, which includes the new Dresden tum) and completely sequenced. The strain. sequence differed from other papilloma- Zell et al (2001) J Virol 75 , 1620 viruses, and it was suggested that a new genus, Sigma papillomavirus, Porcine torovirus (PoTV) A species in should be established to accommo- the genus Torovirus . Immunoelectron date the porcupine virus. These results microscopy of piglet feces revealed await confirmation. elongated particles 120 55 nm. The Rector A et al (2005) Virology 331 , 449 genome RNA was amplified and sequences related to those of bovine and equine toroviruses were detected. porpoise distemper virus A strain of Pigs also had serum neutralizing anti- Cetacean morbillivirus , in the genus bodies to equine torovirus. In a survey Morbillivirus, first identified in 1988 in of 200 weaned pigs in Hungary, toro- harbor porpoises, Phocoena phocoena , in virus was found in 10 healthy weaned the North Sea. Closely related geneti- pigs by electron microscopy and cally to dolphin morbillivirus. RT-PCR. Bolt G et al (1994) Virus Res 34 , 291 Kennedy S (1998) J Comp Pathol 119 , 201 Kroneman A et al (1998) J Virol 72 , 3507 Matiz K et al (2002) Acta Vet Hung 50 , 293 porpoise morbillivirus The first Cetacean porcine transmissible gastroenteritis morbillivirus to be described was identi- virus Synonym for Transmissible gastro- fied in a porpoise caught off the coast enteritis virus . of Northern Ireland. Partial genome sequences show the closest relationship Porcine type C oncovirus (PCOV) A spe- to dolphin morbillivirus. cies in the genus Gammaretrovirus . Banyard AC et al (2008) Virus Res 132 , 213 Identified in a cell line, originating from a lymph node of a leukemic pig, after Portillo virus A possible species in the treatment with 5-bromo-deoxyuridine genus Arenavirus. A member of the and dimethyl sulfoxide. The porcine Tacaribe serogroup, and closely related lymphoma cell particle (PLCP) is sero- antigenically to Junín virus. Has been logically distinct from mouse and cat isolated from infants in Buenos Aires oncoviruses but contains mammalian with hemolytic-uremic disease. interspecies antigen B only. It is thus more like woolly monkey and Gibbon Porton virus (PORV) A tentative spe- ape leukemia viruses than the cat and cies in the genus Vesiculovirus . Isolated mouse oncoviruses. It is endogenous from mosquitoes, Mansonia uniformis , and the DNA sequences are present in in Malaysia. Not reported to cause dis- the cellular DNA of domestic pigs and ease in humans. other species of the family Suidae . positive strand One of the two possible porcupine parvovirus The investigation RNA strands. The one which func- into the deaths of six porcupines over tions as mRNA is known as the ‘posi- a 142-day period, showed histological tive strand: ’ the complementary strand findings indicative of parvovirus infec- is called the ‘negative strand.’ See tion. However, electron microscopy, genome . serology, and virological studies failed Synonym : plus strand. Possum adenovirus 382

Possum adenovirus A species in the fatal, and 20–40% of those who recover genus Atadenovirus . Originally isolated have lasting sequelae. from an opossum, Didelphis marsupialis , primary kidney cell culture show- post-polio syndrome A rare condition in ing spontaneous CPE. Did not replicate which persons who have survived an in rabbit, hamster, human or rhesus initial attack of poliomyelitis suffer a monkey kidney cell cultures, or in degeneration of function many years HeLa or HEP-2 cells. Adenovirus-like later. Infectious virus cannot be iso- particles were subsequently observed lated. Sequences of poliovirus-related by electron microscopy in the intes- RNA have been detected in the CSF in tinal contents of brushtail possums such cases, which may indicate that the Trichosurus vulpecula ( ) in New Zealand. virus can persist in neural cells, but this Using specific primer PCR, sequences is not proven. were amplified of the adenovirus penton base gene that strongly suggested Stone R (1994) Science 264 , 909 that the brushtail possum adenovirus belongs to the genus Atadenovirus . post-transcriptional modification Alter- Thomson D et al (2002) Virus Res 83 , 189 ations in the structure of RNA transcripts prior to utilization as mRNA. These may include splicing, capping possum papillomavirus An unassigned with a blocked methylated structure virus in the family Papillomaviridae at the 5 end, addition of poly A at identified by electron microscopy and the 3 end, or methylation of certain PCR analysis of epithelial tissues from bases internally in the RNA, particu- a brushtail possum, Trichosurus vulpec- larly adenylic and cytidylic acids. ula . Phylogenetic analysis suggests that These modifications all occur in most this is a new papillomavirus. species of eukaryotic cell mRNA, Perrott MR et al (2000) Arch Virol 145 , 1247 and are apparently accomplished by enzymes in the cell nucleus. Viral RNA transcripts synthesized within post-exposure prophylaxis People bit- the cell nucleus (e.g. Orthomyxoviridae , ten by a mammal acting abnormally in Retroviridae , Papovaviridae , Adenoviridae , an area where rabies virus is known to and Herpesviridae ) may be modi- occur must be considered as exposed, fied by these cell enzymes. Viruses and should receive post-exposure treat- replicating wholly in the cytoplasm ment. It consists of passive immuniza- (e.g. Paramyxoviridae , Rhabdoviridae , tion with rabies immunoglobulin and Picornaviridae , Togaviridae , and vaccination. Properly administered, Poxviridae) carry similar enzymes in it is highly effective. If the animal is the virion or induce their synthesis in available, it should be euthanized and infected cells. examined for the presence of rabies virus in the brain. post-translational cleavage Division of a large protein molecule (polyprotein) post-herpetic neuralgia A common com- after translation from the viral genome. plication of herpes zoster, involving The polyprotein is cleaved at specific pain lasting longer than 1 month after sites to produce smaller functional the lesions heal. It occurs in 50% of proteins; this may involve a series of patients older than 60 and can last for cleavages as with human poliovirus. a year or more. Post-translational cleavage is a necessary feature of the replication of post-infectious encephalitis ( PIE) A sev- positive-strand RNA viruses, since it ere complication of measles reported appears that eukaryotic ribosomes only in about 0.1% of measles cases. Usually recognize one initiation site on these develops within 8 days of measles polycistronic genomes. Most other onset when the exanthem is still viruses induce the formation of mono- present. Some 15% of cases of PIE are cistronic mRNAs during replication, 383 Poxviridae

so that functional proteins may be California, Colorado, and New York formed directly. Cleavage may also State, USA, and Dermacentor ander- occur within the assembled virion. soni in South Dakota, USA. Antibodies Cleavage of the surface glycoproteins occur in squirrels and chipmunks in appears to increase virulence in the Ontario. Newborn mice can be infected case of Paramyxoviridae . experimentally, but not adults. Ehrenfeld E (1993) Semin Virol 4 , 199 Wilson MS et al (1979) Can Med Assoc J 121 , 320 post-translational modification A variety of different modifications have Poxviridae A family of large double- stranded DNA viruses. Brick-shaped or been found to occur after translation of a viral protein, for reasons that are not ovoid virions, 220–450 140–260 nm, fully understood. These include cleav- with external coat containing lipid and age into smaller fragments, glycosyla- tubular or globular protein structures, tion, phosphorylation, prenylation, and enclosing one or two lateral bodies, myristylation. and a core which contains the genome. Buoyant density (CsCl): 1.25 g/ml. post-viral fatigue syndrome See chronic There are about 100 virion proteins and fatigue syndrome . many virus-induced proteins, including a DNA-dependent RNA polymer- Potiskum virus (POTV) A serotype of ase. The genome is a single molecule Saboya virus Flavivirus of covalently closed double-stranded , in the genus , belonging to the Yellow fever virus DNA 130–375 kb in length. G C con- antigenic group. tent of vertebrate species is 35–40%. Genetic recombination occurs within Potosi virus (POTV) A serotype of genera; non-genetic reactivation occurs Bunyamwera virus in the genus Ortho- both within and between genera of bunyavirus belonging to the Bun yam- vertebrate species. There are at least wera virus group which appears to 10 major antigens in the virion, one of circulate in several regions of the US. which cross-reacts with most vertebrate Isolated from mosquitoes, Aedes albopic- species, and there is extensive sero- tus . Whilst antibodies have been found logical cross-reactivity within genera. in white-tailed deer the virus is not Replication occurs in the cytoplasm associated with disease in humans. with type B (viral factories) and type A (cytoplasmic accumulation) inclusion Blackmore CGM and Grimstad PR (1998) bodies. Virus is released from micro- Am J Trop Med Hyg 59 , 704 villi or by cellular disruption. Ether- sensitivity varies between genera. A Poult enteritis mortality syndrome hemagglutinin separate from the virion (PEMS) A disease of turkey poults is produced by species in the genus characterized by enteritis, growth Orthopoxvirus. There are two sub- depression, lymphoid atrophy, and families: Chordopoxvirinae (poxviruses immunosuppression. Caused by a tur- of vertebrates) and Entomopoxvirinae key astrovirus. (poxviruses of insects) and 11 genera: Tang Y et al (2006) Avian Dis 50 , 526 Orthopoxvirus , Avipoxvirus , Capripox- virus , Leporipoxvirus , Mollus cipoxvirus , poultry rotaviruses See Rotavirus D , and Parapoxvirus , Suipoxvirus , Yatapoxvirus tentative species rotavirus F and rota- and three genera of Entomopoxvirus , A , virus G . B , and C . Powassan virus (POWV) A tick-borne Buller RM and Palumbo GJ (1991) Microbiol species in the genus Flavivirus , mem- Rev 55 , 80 Fenner F et al (Editors) (1989) The Orthopox ber of the mammalian tick-borne Viruses . New York: Academic Press virus group. Isolated from a human Smith GL (2005) In Topley & Wilson’s case of fatal encephalitis in Ontario, Microbiology and Microbial Infections , vol. Canada. The virus has been recovered 1, Tenth edition, edited by BWJ Mahy and from ticks, Ixodes marxi and I cookei , in V ter Meulen, London: Hodder Arnold, p. 578 poxvirus avium 384 poxvirus avium Synonym for Fowlpox prevalence The number of existing cases virus . of disease in a population at risk. Point prevalence is the number of cases at poxvirus officinalis Name proposed for an arbitrary point in time. Differs from Vaccinia virus , but not adopted. incidence, which is the number of new cases of disease arising during a fixed pRB A cellular tumor suppressor gene period in a population of given size. (retinoblastoma tumor suppressor) that functions in a similar way to the p53 primary culture The establishment of gene in growth control and apoptosis. cells in culture from fresh tissue. The organized tissue needs dissociation Pietenpol JA et al (1996) Proc Natl Acad Sci into single cells, usually by various 93 , 8390 proteolytic enzymes (e.g. pronase or trypsin). It does not include cultures PR8 virus. An early strain of human started from explants of tumors devel- influenza A virus, isolated in Puerto oped by injecting cultured cells into Rico in 1934. animals. Precarious Point virus (PPV) A tick- borne serotype in the genus Phlebovirus , primary rhesus monkey kidney cells member of the Uukuniemi serogroup. (PMKC) Cells used for isolation of a Isolated from Ixodes ( Ceratixodes ) uriae wide range of human viruses includ- in Southern Ocean, Australia. Not asso- ing influenza and other respiratory ciated with human disease. viruses. precipitin An antibody which reacts with primate adeno-associated viruses an antigen to form a precipitate (visible Four possible species in the genus complex) in a precipitin reaction. Dependovirus referred to as types 1, 2, 3, and 4. Antigenically distinguishable pre-exposure prophylaxis Pre-exposure by neutralization, CF, and precipitin rabies immunization, recommended tests. There is some antigenic relation- for persons working with rabies in the ship between types 2 and 3. Type 3 laboratory, veterinarians, and animal species has strains K, H, and T, distin- control and wildlife officers. guishable by neutralization. Unrelated antigenically to bovine or avian adeno- associated viruses. The presence of pre-integration complex (PIC) A com- antibodies suggests that types 2 and 3 plex formed early in the replication are human viruses and that types 1 and cycle of HIV. Following virus entry, 4 are monkey viruses. No evidence of the genome RNA is reverse transcribed pathogenicity. into dsDNA, and the PIC is formed, consisting of three virus proteins, the Berns KI (1990) Microbiol Rev 54 , 316 integrase, matrix, and Vpr, together Hoggan MD (1971) In Comparative Virology , with the dsDNA. This complex then edited by K Maramorosch and E Kurstak. moves into the nucleus. New York: Academic Press, p. 43 prenylation Addition of a carbon moi- primate calicivirus (Pan-1) (VESV/ ety derived from mevalonic acid to a Pan1) A tentative species in the genus cysteine located at the C-terminus of a Vesivirus causing mucosal vesiculation protein. So called because prenyltrans- and persistent infection in chimpan- ferases are responsible for the transfer zees. Related genetically to vesicular of the moiety on the cysteine. exanthema of swine virus.

Pretoria virus (PREV) A serotype of Primate T-lymphotropic viruses 1–3 Dera Ghazi Khan virus in the genus (PTLV-1 to 3) Species in the genus Nairovirus . Isolated from the tick, Deltaretrovirus . The viruses HTLV-1 and Argas africolumbae, in South Africa. Not STLV-1 are not clustered genetically reported to cause disease in humans. according to host species but according 385 probe

to geographical origin. Primate Tenth edition, edited by BWJ Mahy and T-lymphotropic viruses 1–3 each include V ter Meulen, London: Hodder Arnold, a simian and a human isolate as strains. p. 1346 Human T-lymphotropic virus type 3 Prusiner SB (Editor) (1999) Prion Biology and Diseases. Cold Spring Harbor Laboratory was identified in 2006 in two primate Monograph 38. New York: Cold Spring hunters from Africa. The nucleotide Harbor Laboratory Press sequence of this virus genome is closer Safar J et al (1998) Nat Med 4 , 1157 to that of STLV-3 than to that of HTLV-1 or HTLV-2, and it is assumed to have a prion diseases The following diseases are nonhuman primate origin. probably caused by prions: Switzer WM et al (2006) J Virol 80 , 7427 bovine spongiform encephalopathy (cattle) prime and realign model A model pro- chronic wasting disease (mule deer and posed to explain the transcription of elk) the hantavirus genome. Creutzfeld–Jakob disease (CJD) Garcin D et al (1995) J Virol 69 , 5754 (humans) exotic ungulate encephalopathy (nyala and kudu) primer A short RNA sequence that pairs fatal familial insomnia (humans) with one strand of DNA and provides feline spongiform encephalopathy (cats) a free 3 -OH terminus at which a DNA Gerstmann–Sträussler–Scheinker syn- polymerase starts DNA synthesis, or drome (humans) pairs with a strand of RNA and allows kuru (humans) reverse transcriptase to start copy- variant CJD (humans) ing the RNA into DNA. Retroviruses scrapie (sheep and goats) use specific transfer RNAs to initiate transmissible mink encephalopathy reverse transcription of the genome (mink) RNA into DNA. Endogenous human retroviruses are classified according to There is evidence that the newly recog- which tRNA primer is specified in the nized fatal neurodegeneration known LTR sequence of the provirus. as variant CJD is caused by the same prion strain as causes bovine spongi- prion Sigla from pr oteinaceous and form encephalopathy (BSE). However in fectious particle. Term used for the the mode of transmission from cattle pathogens which induce fatal neuro- to humans has not been elucidated. A logical diseases (transmissible enceph- method for prion strain discrimination alopathies) of vertebrates, e.g. scrapie using luminescent conjugated poly- disease of sheep and goats, bovine mers has been described. spongiform encephalopathy in cattle, chronic wasting disease of deer and Collinge J et al (1996) Nature 383 , 685 elk, and Creutzfeld–Jakob disease and Hill AF et al (1997) Nature 389 , 448 kuru in humans. Prions are oligom- Scott MR et al (1999) Proc Natl Acad Sci 96 , ers composed of a 27–30 kDa protein, 15137 PrPsc, that has the same amino acid Sigurdson CJ et al (2007) Nat Methods 4 , 1023 sequence as a normal cell protein, PrPc, prion protein (PrP) A normal cellular which occurs in neuronal membranes, protein of 33–35 kD which is protease and is encoded in the PRNP gene on the sensitive and soluble in non-denaturing short arm of chromosome 20. All cases detergents, the function of which is of familial prion disease cosegregate unknown. Encoded by a chromosomal with PRNP missense mutations. Prions gene. When infected by a prion, PrP are therefore isoforms of a normal is converted into a protease-resistant protein, capable of self-replication, but disease-specific protein that is insolu- they are not viruses since they do not ble in detergents. contain nucleic acid. Aguzzi A (2005) In Topley & Wilson’s probe A sequence of DNA or RNA which Microbiology and Microbial Infections , vol. 2, is labeled (e.g. with radioactivity or probe 386

biotin) and used to detect complemen- USA, in 1923. Caused by Visna/maedi tary sequences by hybridization. virus.

Proboscivirus Proposed name for a new Synonyms : progressive interstitial pneu- genus in the subfamily Betaherpesvirinae monia virus; visna-maedi. that would include endotheliotropic elephant herpesviruses. Not yet projectile vomiting A symptom associ- adopted. ated with acute outbreaks of norovirus infection. The vomitus contains Ehlers B et al (2006) J Gen Virol 87 , 2781 infectious virus, and in addition to Wellehan JF et al (2008) Vet Microbiol 127 , 249 fecal-oral transmission, projectile vomiting has been shown to be an impor- procapsid In some virus infections, a tant means of virus spread in explosive stage before virion formation. A viral outbreaks of norovirus associated with capsid without nucleic acid. catering establishments. procyonid herpesvirus 1 See lorisine herpesvirus 1 . prokaryote An organism of the kingdom Prokaryotae, including bacteria and productive infection Infection of a cell in cyanobacteria. which complete infectious virus particles are formed. prokaryotic expression systems Exp- ression of virus genes in E coli or other productive transformation The RNA prokaryotic systems provides a conven- tumor viruses integrate their entire ient and inexpensive method of obtain- genome into the host cell, and by ing virus polypeptides for study, but expressing oncogenes transform their the processing pathways in prokaryo- host cells to a state of uncontrolled cell tes are different from eukaryotes. division and continuous release of viral Consequently antigens produced in progeny. these systems may lack correct amino- terminal modifications, disulfide bond proflavine A photoreactive dye derived formation and glycosylation needed from acriflavine. See photodynamic for full activity. inactivation . proliferative ileitis of hamsters See ham- progressive interstitial pneumonia virus ster enteritis . See Visna/maedi virus (strain 1514 ). ProMED The Pro gram for M onitoring progressive multifocal leukoencepha- E merging Diseases. It has inaugurated lopathy (PML) PML is a rare, slowly an e-mail conference system on the progressive, fatal and non-inflam- Internet to encourage timely informa- matory demyelinating disease of the tion sharing and discussion on emerg- human CNS caused by reactivation ing disease problems worldwide. of latent JC polyomavirus infection in patients with immunodeficiency, http://www.fas.org/promed/ usually due to disease (e.g. AIDS) or immunosuppressive therapy. See JC promoter A region of DNA, usually polyomavirus . upstream of a coding sequence which directs RNA polymerase to bind and progressive pneumonia of sheep A initiate transcription. As well as the chronic pulmonary disease in sheep, defined sequences (e.g. TATA box) which affects older ewes and consists there are upstream and sometimes of slowly progressive loss of weight downstream sequences which attenu- and dysphagia over a period of 3–12 ate or modulate transcription. months. There is nodular proliferation of lymphocytes in the lungs and promyelocyte leukemia protein (PML) thickening of interalveolar septa. First A host cell nuclear oncoprotein. The reported to affect sheep in Montana, Z protein of arenaviruses interacts 387 protein folding

with PML to modulate the interferon Prostate virus A novel gammaretrovirus response to infection. identified in prostate tissue RNA from Borden KL et al (1998) J Virol 72 , 758 human cancer patients homozygous for a reduced activity variant of the antiviral enzyme RNase L. See XMRV . pronase A nonspecific proteolytic enzyme isolated from a fungus, Streptomyces Dong B et al (2007) Proc Natl Acad Sci 104 , griseus . Useful as a preliminary treat- 1655 ment during extraction of intact RNA molecules from cells or virus because protamines Simple basic proteins rich of its strong inhibitory action on in arginine. Present in cell nuclei. See ribonuclease. histones . Huppert J and Semmel M (1965) Biochim Biophys Acta 108 , 501 protease A generic term for an enzyme such as pepsin or trypsin which cleaves a polypeptide chain by hydrolysis. An propagated epidemics Epidemics of viral important mechanism of pathogenic- disease in which spread from host to ity associated with enveloped virus host continues to expand as long as infections. In retroviruses, a protease each infection gives rise to more than that is essential for virus replication is one new infection. Distinct from com- encoded within the gag-pol gene, and mon source epidemics. is a target for anti-retroviral inhibitory drugs. See protease inhibitors . propagation Growth of a virus with suc- Katz RA and Skalka AM (1994) Annu Rev cessive passages in cell cultures, fertile Biochem 63 , 133 eggs, or animals. Nagai Y (1993) Trends Microbiol 1 , 81 prophage The bacteriophage genome protease inhibitors As antiviral agents, integrated into the genome of a lys- compounds which block the post- ogenic bacterial cell. translational cleavage of the gag and gag- pol polypeptides of human immuno- Lwoff A and Gutmann A (1950) Ann Inst deficiency virus type 1, and so inhibit Pasteur 78 , 711 virus assembly and maturation. Several such compounds have been licensed β -propiolactone A chemical widely used for treatment of AIDS patients, usually to inactivate viruses because it gener- in combination with other drug treat- ally does not interfere with antigen test ments such as zidovudine (AZT). They systems. include ritonavir and saquinavir. prop-pox Synonym for scrum-pox. proteid An alternative name for protein.

Prospect Hill virus (PHV) A species in the protein A high-molecular-weight poly- genus Hantavirus, isolated by Carlton peptide of L-amino acids that is syn- Geiduschek from the meadow vole, thesized by living cells. The principal Microtus pennsylvanicus , in Maryland, constituent of the cytoplasm. USA. The first hantavirus found in a protein A A cell wall protein of some US rodent. Not known to cause disease strains of Staphylococcus aureus , which in humans. Related viruses have been has the ability to bind strongly to the Fc isolated in North Dakota and Nevada, portion of an antibody molecule when USA, from the voles, Microtus pennsyl- that antibody is bound to an anti- vanicus , M ochrogaster, and M montanus . gen; used to collect antigen–antibody Their disease potential is unclear. complexes. Lee PW et al (1985) J Infect Dis 152 , 826 protein folding The capsid proteins of Prospect Hill-like virus Strains isolated icosahedral viruses frequently undergo from Microtus pennsylvanicus , M monta- folding during the final assembly pro- nus , and M ochrogaster . cess, often into an eight-strand β -barrel. protein folding 388

Rossmann MG and Johnson JE (1989) Ann 1, Tenth edition, edited by BWJ Mahy and Rev Biochem 58 , 533 V ter Meulen, London: Hodder Arnold, p. 202 Kozak M (1992) Annu Rev Cell Biol 8 , 197 protein kinase (PKR) An enzyme which catalyzes the phosphorylation of a pro- proteinaceous infectious particles See tein, usually in the presence of a cyclic prion . nucleotide, cyclic AMP, or cyclic GMP, although kinases utilizing ATP or proteolytic cleavage Enzymatic cleav- GTP are known. There are three main age of a protein at specific site(s), classes: serine-, threonine-, or tyrosine- e.g. the cleavage of a polyprotein to kinase, depending upon the amino yield structural proteins. acid that is phosphorylated. Several oncogene products have protein kinase proteolytic enzyme An enzyme that cata- activity. lyzes the hydrolysis of peptide bonds. Rubin CS and Rosen OM (1975) Annu Rev proteosome A complex of proteolytic Biochem 44 , 831 enzymes where peptide cleavage of intracellular pathogens such as viruses protein synthesis Initiation of protein occurs before transport via peptide synthesis in eukaryotic cells usually transporters (TAP1 and -2) to the endo- involves the recognition by the cel- plasmic reticulum where they associate lular initiation factor eIF-4F of the with the MHC class I heavy chain. The 7-methylguanylate cap at the 5 end light chain ( β 2 microglobulin) associ- of cellular and many viral mRNAs. ates with the heavy chain and the tri- A complex then forms between the molecular complex is transported to mRNA and the 40S ribosomal subunit, the cell surface for antigen presenta- met-tRNA, and another initiation fac- tion. The complex of MHC class I with tor, eIF-2. With a small number of cel- a foreign (virus) peptide is recognized lular RNAs and with uncapped viral by CD8 T lymphocytes which then RNAs (such as picornaviral RNAs), destroy the infected cell. initiation occurs at an internal ribos- Lehner PJ and Cresswell P (1996) Curr Opin omal entry site (IRES), and eIF-4F is Immunol 8, 59 not involved. To start protein synthesis the 40S ribosomal subunit scans protomers Protein units which polymer- in a 3 direction until it encounters an ize to form a capsomere. AUG (methionine) codon, and if the local sequence context is favorable proto-oncogenes The cellular genes that for initiation a 60S ribosomal subunit are transduced by retroviruses and joins the complex, eIF-2 dissociates, are recognized by sequence similar- and synthesis of the polypeptide chain ity to the viral oncogenes. For exam- begins. The most favorable sequence ple v-src, the first viral oncogene to be at the ribosomal binding site is A/ recognized, was found to be present in GCCACCAUGG, also known as the normal chicken cell DNA, and it is now Kozak sequence, but various other believed that all vertebrate cells con- factors such as secondary structure of tain highly conserved proto-oncogenes. the mRNA may play a role. In some The transduced oncogene may become viral mRNAs the first AUG is ignored modified by mutation so that, for for these reasons and a later AUG is example, the tyrosine kinase activity of used. Depending on the virus, various v-SRC greatly exceeds that of the cellu- modifications of the cellular machinery lar gene product SRC. See oncogene . for protein synthesis may occur during replication, which may favor virus ‘ protovirus ’ hypothesis A hypothesis protein synthesis over cellular, and in promulgated by Howard Temin that some cases cellular protein synthesis is retroid elements, present in all cells, shut off completely and the cell dies. may acquire additional genes encoding Ball LA (2005) In Topley & Wilson’s envelope proteins and so develop into Microbiology and Microbial Infections, vol. retroviruses. 389 psittacid herpesvirus 1 (PsHV-1) provirus The viral genome integrated pseudorabies virus (PRV) See Suid her- as DNA into the cell genome with pesvirus 1 . which it replicates. The term is usually applied to retroviruses. Can be pseudorinderpest virus Synonym activated spontaneously or in response for Peste des petits ruminants virus to certain stimuli to produce complete (Katavirus) in the genus Morbillivirus . virus. Analogous to a prophage.

C pseudotype virus A virus which may PrP The normal cellular prion protein. result from a mixed infection. The genome of one virus is enclosed in PrPCJD The protease-resistant prion an envelope coded for by another protein found in cells infected with virus. Can be used to introduce virus Creutzfeldt–Jakob disease. genomes into cell types in which they would not normally replicate. In con- PrPSc The protease-resistant prion protein junction with recombinant genomes found in cells infected with the scrapie which carry dominant selectable mark- prion. Also used as a general term for ers for drug resistance or metabolism, protease-resistant prions of other trans- can be used to clone receptor genes, missible encephalopathy agents. e.g. of retroviruses. See phenotypic mixing . prune belly syndrome A syndrome Cosset FL et al (1992) J Virol 66 , 5671 which results in a thin-walled protruding abdomen with wrinkled skin that occurred in a human neonate in asso- Pseudovirida e A family of retrotrans- ciation with hydrops fetalis following posons found in yeast, insects, and maternal infection with B19 virus. plants. They form an intrinsic part of the genome, but are not infectious, Walther J-U et al (1994) Monatsschr in the virological sense, in normal Kinderheilkd 142 , 592 conditions. pseudo-aphthous stomatitis of cattle virus Synonym for Bovine papular sto- pseudovirions Virus particles that con- matitis virus, although other viruses tain host nucleic acid in place of viral may cause the same clinical picture. genome nucleic acid. A type of defective particle. Pseudocowpox virus (PCPV) A species in the genus Parapoxvirus. Similar to Orf psittacid herpesvirus 1 (PsHV-1) An virus. Causes hemispherical cherry-red unassigned virus in the subfamily papules on the udders of cows and the Alphaherpesvirinae. Isolated in Brazil hands of milkers. Can be propagated in in 1931 from parrots especially of the human cell cultures. genus Amazona, in which it causes Synonyms: natural cowpox virus; para- weakness, diarrhea, coma, and death vaccinia virus; milker’s nodule virus. in 3–7 days. Budgerigars, Melopsittacus sp, are also highly susceptible. Disease pseudoknot A secondary structure in is not produced on experimental inoc- a viral mRNA which slows move- ulation of guinea pigs, mice, pigeons, ment of the ribosome and may cause a chickens, or turkeys. The virus has also frameshift that allows entry to an alter- been isolated from aviary birds in the native reading frame. Important espe- USA. It can be propagated on the CAM cially in the replication of members of where it produces white plaques and the order Nidovirales . kills the embryo. Replicates with CPE in chick kidney cell cultures. pseudolumpy skin disease Synonym for Synonyms : Pacheco’s disease virus; par- Bovine herpesvirus 2 . rot herpesvirus; Pacheco’s parrot virus. Simpson CF et al (1975) J Infect Dis 13 , 390 pseudolymphocytic choriomeningitis Tomazewski E et al (2001) J Clin Microbiol virus Synonym for Ectromelia virus . 39 , 533 Psittacinepox virus (PSPV) 390

Psittacinepox virus (PSPV) A species the causative agent has not been defi- in the genus Avipoxvirus , isolated nitely established. from pox-like lesions on blue-fronted Nuttall PA et al (1985) J Wildl Dis 21 , 120 Amazon parrots. Stoker MGP and Miles JAR (1953) J Hyg McDonald SE et al (1981) J Am Vet Med Assoc (Camb ) 51 , 195 179 , 218 pullet disease virus Synonym for infec- Psorophora columbiae The dark ricefield tious enteritis virus. See also blue mosquito, found in Mexico, Central comb virus . America, the Caribbean, and South America to Argentina. An important vector for Venezuelan equine encepha- pulmonary adenomatosis of sheep virus litis and West Nile virus transmission. See Ovine pulmonary adenocarcinoma virus (jaagsiekte virus). Pt K1 (NBL-3) cells (CCL 35) A cell line derived from the kidney of an appar- pulse-chase analysis An experiment in ently normal adult female potoroo, which a radioactively labeled precur- Potorous tridactylis. Perhaps the first sor compound is added to cells or a permanent marsupial cell line to be cell extract in vitro for a short period established. (pulse), after which a large excess of unlabeled compound is added to Pt K2 (NBL-5) cells (CCL 56) A cell line dilute and prevent further significant derived from the kidney of an appar- incorporation of radioactivity. Samples ently normal adult male potoroo, are taken at various times to follow the Potorous tridactylis . course of the radioactive precursor as it is metabolized (chase period). Puchong virus (PUCV) A tentative species in the genus Ephemerovirus , Puma lentivirus ( PLV-14 ) A species in antigenically related to bovine ephem- the genus Lentivirus in the feline lenti- eral fever, Berrimah, Kimberley, and virus group. Infectious puma lentivirus Malakal viruses. Isolated from the mos- has been isolated from several Florida quito, Mansonia uniformis , in Malaysia panthers. The virus can be grown in a in 1965. Not reported to cause disease. domestic cat lymphoma cell line 3201. Genomic DNA from a Florida puma, Pueblo Viejo virus (PVV) A sero- Felis concolor coryi, infected with the type of Gamboa virus in the genus PLV 14 strain of PLV was sequenced Orthobunyavirus. Isolated from Aedeo- and found to be closely related to myia squamipennis in Ecuador. Not feline immunodeficiency virus. The reported to cause disease. virus has also been found in cougars (Felis concolor ) sampled in several other US States, including Arizona, Puffin Island virus (PIV) A serotype of California, Colorado, Idaho, Oregon, Hughes virus in the genus Nairovirus . and Washington. Isolated from ticks, Ornithodoros maritimus, from a colony area of herring gulls, Evermann JF (1997) J Wildl Dis 33 , 316 Larus argentatus, in North Wales, UK. Langley RJ et al (1994) Virology 202 , 853 van deWoude S et al (1997) Virology 233 , 185 Converse JD et al (1976) Acta Virol 20 , 243 Puna teal hepadnavirus An avihepadna- puffinosis An avian disease, probably virus cloned from a captive Puna teal. caused by a virus, which is character- The genome sequence is closely related ized by blisters on the foot-web of to duck hepatitis B virus. some seabirds, particularly shear- Guo H et al (2005) J Virol 79 , 2729 waters. Occurs in UK as an epizootic in late August to mid-September. Species Punchana virus A strain of Rio Mamore of Poxviridae and Coronaviridae have virus. The rodent vector is Oligoryzomis been isolated from infected birds but microtis . 391 pyrophosphate analogues

Punta Salinas virus (PSV) A strain of pustular dermatitis of camels virus A Hughes Virus in the genus Nairovirus , possible species in the genus Parapox- belonging to the Hughes serogroup. virus. Causes pustular dermatitis in Isolated from the tick, Ornithodoros camels. amblus , in Peru. Not reported to cause Roslyakov AA (1972) Vopr Virusol 17 , 26 disease in humans. Puumala virus (PUUV) A species in the Punta Toro virus (PTV) A species in the genus Hantavirus. A natural infection genus Phlebovirus, belonging to the of the bank vole, Clethrionomys glareolus Punta Toro complex, sandfly fever and C rufocanus, from which the virus serogroup. Isolated from Lutzomyia was isolated in Vero cells. In humans sp and from humans in Panama and causes nephropathia epidemica, an Colombia. In humans it can cause a acute fever with renal involvement febrile illness with myalgia, enlarged recognized in Scandinavia and several liver and spleen, and increased protein other countries in Europe, including in the CSF. Russia. Mortality is low (less than 1%). Robeson G et al (1979) J Virol 30 , 339 Cross-reacts antigenically more strongly with North American hantaviruses Punta Toro virus D-4021A (PTV) A strain (Prospect Hill and Sin Nombre viruses) of Punta Toro virus . than with Hantaan or Seoul viruses. Brummer-Korvenkontio M et al (1980) J purine A basic heterocyclic nitrogen- Infect Dis 141 , 131 containing compound. Adenine and Plyusnin A et al (1996) J Gen Virol 77 , 2677 guanine are the purine components of nucleic acids. pyrimidine A basic heterocyclic nitrogen- containing compound. Cytosine, puromycin A nucleoside antibiotic thymine, and uracil are the pyrimidine produced by Streptomyces alboniger , components of nucleic acids. and an inhibitor of protein synthesis. Structurally similar to amino-acyl python orthoreovirus (PRV) A tentative transfer RNA, so can act as acceptor species in the genus Orthoreovirus iso- of the nascent polypeptide chain of lated from a moribund snake, Python ribosome-bound peptidyl-tRNA, which regius . The genome contained 10 seg- is then released prematurely as pepti- ments of double-stranded RNA. dyl puromycin. Ahne W et al (1987) Arch Virol 94 , 135 Bablanian R (1968) J Gen Virol 3 , 51 pyrophosphate analogues A series of Purus virus (PURV) A serotype of compounds that interact directly with Changuinola virus in the genus Orbi- the DNA polymerase of herpesviruses. virus, member of the Changuinola virus They include phosphonoformate (PFA) – group. Isolated from Psorophora ( Jan ) foscornat – which has been developed albipes in Acre, Brazil. Not reported to for treatment of cytomegalovirus disease cause disease in humans. in immunocompromised hosts. Q

Qalyub virus (QYBV) A species in the diseases such as rinderpest and rabies in genus Nairovirus . With Bakel, Bandia, non-endemic regions. The UK had a 6- and Omo virus forms the Qalyub month quarantine period for imported serogroup. Isolated from the tick, cats and dogs, but this has been relaxed Ornithodoros erraticus, in Egypt. Not given the wide availability of effective reported to cause disease in humans. vaccines for pet animal immunization Clerx JPM and Bishop DHL (1981) Virology and tests for verifying an adequate anti- 108 , 361 body response. DEFRA are currently reviewing policies on quarantine of pet quail adenovirus See fowl adenovirus 1 . animals. http://www.defra.gov.uk/animalh/ quail bronchitis A disease of quail quarantine. caused by Fowl adenovirus A, a species in the genus Aviadenovirus . An acute, quasi-equivalence highly contagious respiratory disease A theory invoked to account for the surface morphology of in captive and wild bobwhite quail, spherical viruses. It requires that sub- Colinus virginianus . In young birds less units forming the icosahedral capsid than 4 weeks old there may be 100% should be capable of assembling into fatality. Identical to CELO virus, fowl both hexamers and pentamers. The adenovirus 1. See fowl adenovirus 1 . insertion of 12 pentamers produces cur- Synonym : quail adenovirus. vature in the sheet of hexamers where they are inserted, resulting in a closed quail parvovirus Synonym for a strain of Avian adeno-associated virus , genus icosahedral shell that is not strictly Dependovirus . equivalent, but forms a more stable structure. Thus icosahedral viruses have a capsid composed of 12 pentamers and Quailpox virus (QUPV) A species in the a variable number of hexamers, e.g. genus Avipoxvirus . Member of a group herpesviruses have 150 hexamers and of antigenically related viruses which 12 pentamers, making up the capsid; infect a number of different hosts, e.g. adenoviruses have 240 hexamers and 12 canary, pigeon, and sparrow. pentamers. See icosahedral symmetry . Winterfield RW and Reed W (1985) Poult Sci 64 , 65 Caspar DLD and Klug A (1962) Cold Spring Harbor Symp Quant Biol 27 , 1 Quaranfil virus (QRFV) An unassigned vertebrate virus, pathogenic for a variety of laboratory hosts. Related antigen- quasi-species A term that describes the ically to Johnston Atoll virus. Isolated nature of most RNA viruses, but is from argasid ticks collected near Cairo, particularly well studied with hepa- Egypt in 1953. Subsequent isolates were titis C and HIV, which are popula- made from humans, the cattle egret, tions of genetic variants within which Bubulcus ibis, and from pigeons in Egypt one (the quasi-species) predominates. and South Africa. The virions appear to There is also evidence that a minority be enveloped but the virus has not been population exists containing variants fully characterized. Has been associated that were dominant at an earlier stage with a febrile illness in humans. of the evolutionary lineage, and these may influence the subsequent evolu- quarantine A quarantine and slaughter tion of the quasi-species population. policy is used to control serious animal Briones C et al (2006) Gene 384 , 129 393 quokkapox virus (QPV)

Domingo E and Gomez J (2007) Virus Res from the cytoplasm of cells in the dor- 127 , 131 sum of the tails of the quokka, Setonix Domingo E et al (2002) Virus Res 82 , 39 brachyurus . Electron microscopy studies revealed the presence of poxvirus-like quat A quaternary ammonium com- particles in the cytoplasm of cells in pound, such as CTAB. the stratum granulosum of papillomas on the dorsum. The infected popula- quinacrine A substance that has been tion of this marsupial has been isolated suggested as a treatment for prion for 7000 years on the small island of diseases, but it was not effective in a Rottnest off the coast of West Australia. mouse model of CJD disease. Synonyms : marsupial papillomavirus; Collins SJ et al (2002) Ann Neurol 52 , 503 marsupialpox virus. McKenzie RA et al (1979) Aust Vet J 55 , 188 quokkapox virus (QPV) An unassigned Papadimitriou JM and Ashman RB (1972) J virus in the family Poxviridae . Isolated Gen Virol 16 , 87 R

R61837 compound A drug that inhibits closely related to myxoma virus. Can replication of rhinovirus by preventing be propagated on the CAM but no virus uncoating, probably by binding to lesions are produced and the embryo the virus capsid. Effective intranasally in is not infected. In cultures of rabbit, clinical trials against human rhinovirus. guinea pig, rat and human cells, replication occurs with CPE. The natu- R plasmid A bacterial plasmid containing ral host is the wild cottontail rabbit, a gene for drug resistance. Sylvilagus floridanus , in which the virus causes benign fibromas, commonly on Rab5 A membrane-bound intracellular the feet. Virus can be extracted from protein which is a positive regulator the tumors and produces fibromas in of endocytosis. Rab5 transfected cells wild and domestic rabbits on injection. acquire abnormally large endosomes See Berry–Dedrick phenomenon . immunoreactive for Rab5. Synonyms : fibroma virus of rabbits; Shope fibroma virus. rabbit calicivirus (RCV) A strain of Rabbit hemorrhagic disease virus in the genus Fenner F (1994) In Virus Infections of Rodents Lagovirus . In contrast to RHDV, RCV and Lagomorphs , edited by ADME Osterhaus. Amsterdam: Elsevier Science, p. 71 appears to be apathogenic, causing an intestinal infection but not the severe Rabbit hemorrhagic disease virus (RHDV) liver and spleen necrosis seen with The type species of the genus Lagovirus . RHDV infection. The sequence of the A relatively new and economically capsid protein gene and antigenicity important viral disease of wild and studies show that the two viruses are domestic rabbits and hares, outbreaks of closely related, however. which have appeared worldwide since Capucci L et al (1996) J Virol 70 , 8614 1984. Characteristic hemorrhages occur in many organs, e.g. trachea, lung, liver, rabbit coronavirus (RbCoV) A tenta- kidney, spleen, peritoneum, and pleura. tive species in the genus Coronavirus , Antigenically related to European first detected in 1970 in Sweden. brown hare syndrome virus, with which Serologically related to porcine hemag- it shares a similar genome organization. glutinating encephalomyelitis virus, Genome is 7437 bases long and includes mouse hepatitis virus, bovine corona- two open reading frames (ORFs), one virus, turkey blue comb coronavirus, encoding a large polyprotein which sialodacryoadenitis virus of rats, and includes the capsid protein as well as the human coronaviruses 229E and OC- nonstructural proteins, and a smaller one 43. Causes severe symptoms in the eye encoding part of the polymerase gene (dullness of the scleras and congestion and the complete capsid protein gene. of conjunctivae), the heart (myocarditis This organization is different from other with extensive calcification and dila- calicivirus genomes, which have three tion of the ventricles), and respiratory ORFs. There is a genome-linked protein symptoms (pulmonary edema, pleural (VPg) at the 5 terminus. Animal trans- effusion) in rabbits. The rabbit corona- mission studies have been performed virus has been proposed as an experi- to establish the host range of RHDV mental model for studies of myocarditis since it was considered in 1995 for use and congestive heart failure. in a rabbit control program in Australia Edwards S et al (1992) J Infect Dis 165 , 134 and New Zealand. More than 30 species were inoculated with large doses of virus Rabbit fibroma virus (SFV) A species in but none showed evidence of infection the genus Leporipoxvirus . Antigenically except kiwis, which made antibodies 395 rabbit type C endogenous virus

against the virus but showed no clini- Rabbit oral papillomavirus (ROPV) A cal signs. During experimental testing of species in the genus Kappapapillomavirus . the effectiveness of RHDV in controlling Antigenically and genetically different rabbits on Wardang Island off Southern from cottontail rabbit papillomavirus. Australia, the virus was accidentally A natural infection of domestic rabbits transmitted to the mainland, where large causing papillomas, usually beneath the numbers of rabbits have died of the calici- tongue, which regress in a month or two. virus infection. Overall rabbit numbers On inoculation into the oral mucosa, have declined 60% in Australia. In papillomas appear in 6–38 days. Will not New Zealand, the virus was introduced produce skin warts. Sylvilagus and Lepus illegally, probably by irate farmers who can be infected. The genome proteins objected to a decision of the govern- E6, E7, and E8 are oncogenic. ment not to allow legal importation of Christenson ND et al (2000) Virology 269 , 451 the virus until more was known about Hu J et al (2004) Virology 325 , 48 its potential host range. The emergence Parsons RJ and Kidd J (1943) J Exp Med of rabbit calicivirus, which appears to be 77 , 233 a non-pathogenic mutant of RHDV, may diminish the value of RHDV as a biolog- rabbit papillomavirus See Cottontail ical control agent. rabbit papillomavirus . Landstrom C (2001) Soc Stud Sci 31 , 912 Meyers G et al (2000) Virology 276 , 349 rabbit parvovirus Synonym for Lapine Studdert MJ (1994) Aust Vet J 71 , 264 parvovirus . Wirblich C et al (1996) J Virol 68 , 5164 rabbit plague virus Synonym for rabbit- rabbit herpesvirus Synonym for leporid pox virus. herpesvirus 1. rabbitpox virus (RPXV) A strain of Rabbit kidney vacuolating virus (RKV) Vaccinia virus in the genus Orthopoxvirus . A species in the genus Polyomavirus . A laboratory artifact due to infection of A natural and latent infection of cot- colonized rabbits with vaccinia virus. tontail rabbits, Sylvilagus floridanus. Not No natural reservoir. After infection known to be pathogenic in any spe- by the respiratory route causes usually cies. Agglutinates guinea pig erythro- lethal generalized disease. cytes at 4°C and 20°C, by reacting with Synonym : rabbit plague virus. neuraminidase-sensitive receptors. Replicates, producing cell vacuolation, in domestic and cottontail rabbit kidney rabbit reticulocyte lysate A cell-free sys- cell cultures but not in other species. tem prepared from lyzed rabbit reticu- Synonyms : polyomavirus sylvilagus; locytes which is used for the translation rabbit vacuolating virus. of eukaryotic mRNAs. The rabbit is made anemic by injection with acetyl Hartley JW and Rowe WP (1964) Science 143 , phenylhydrazine and the reticulocytes 258 extracted. After lysis the endogenous mRNAs are destroyed using micrococ- Table R1 . Strains of rabbit hemorrhagic cal nuclease which is then inactivated disease virus by the addition of EGTA (ethylene- rabbit calicivirus glycolbis (aminoethylether) tetra-acetic rabbit hemorrhagic disease virus-AST89 acid). Translation in the extract is then (RHDVAST89) totally dependent upon added mRNA. rabbit hemorrhagic disease virus-BS89 See wheat germ extract . (RHDV-BS89) McCrae M (1985) In Virology : A Practical rabbit hemorrhagic disease virus-FRG Approach, edited by BWJ Mahy. Oxford: IRL (RHDV-FRG) Press, p. 167 rabbit hemorrhagic disease virus-SD (RHDV-SD) rabbit type C endogenous virus A possi- rabbit hemorrhagic disease virus-V351 ble species in the genus Gammaretrovirus . (RHDV-V351) When primary lymphosarcoma cell rabbit type C endogenous virus 396

cultures from WH/J rabbits were treated In the former USSR the wolf, in South with idoxuridine, C-type virus particles Africa the mongoose, in India the jackal, were produced which contained RNA- and in South and Central America the dependent DNA polymerase and a p30 vampire bat are important reservoirs structural protein. These proteins shared of infection. In the USA skunks, foxes, antigenic homologies with other mam- bats, coyotes, and raccoons spread malian group viruses but also possessed rabies. Insular areas such as Hawaii and unique antigenic determinants. There Australia are kept free of rabies by a was evidence that viral genetic infor- 6-month quarantine of imported cats mation was present in the WH/J rabbit and dogs. This was also the situation in cells. These rabbits developed spontane- the UK until recently, when the Channel ous lymphosarcomas. Tunnel was opened. All laboratory animals can be infected and die. There is Bedigian HG et al (1978) J Virol 27 , 313 encephalitis, especially of the midbrain, cerebellum, and medulla. Negri bod- rabbit vacuolating virus Synonym for ies are present in nerve cells. Diagnosis Rabbit kidney vacuolating virus . is with fluorescein-labeled antirabies antibody staining of brain tissue taken rabies immunoglobulin (HRIG) An at necropsy. The virus can be replicated immunoglobulin prepared from human in most tissues of the embryonated egg serum that is used for post-exposure and in a wide range of primary and rabies prophylaxis together with rabies continuous cell cultures. CPE is rare. vaccine. The animal reservoir of an isolate of rabies virus can be determined using Rabies virus (RABV) The type species of monoclonal antibody panels and nucleo- the genus Lyssavirus. The natural hosts tide sequence analysis. Early diagnosis are many bat species and terrestrial in humans depends primarily on a his- carnivores, but most mammals can be tory of a bite from an animal proved to infected. In dogs the incubation period have rabies, since the first clinical signs is from 10 days to more than 6 months; are nonspecific. Late in the course of the in humans usually from 15 days to 5 infection, virus replicates in other tis- months, but incubation periods as long sues besides the CNS and brain, and can as several years have been documented. be demonstrated by fluorescent anti- Dogs at first show excitement or aggres- body or by isolation in saliva, corneal sion with hypersalivation and often epithelium and cutaneous nerves in bite (furious rabies), later depression skin biopsy, but this is too late for post- and paralysis (dumb rabies) and soon exposure treatment. On passage by i.c. die. The same two stages may be seen injection in laboratory rabbits the wild in humans, with about 20% of cases virus ( ‘ street ’ virus) becomes lethal in being mainly paralytic, especially 4–6 days. This now called ‘ fixed ’ virus after exposure to bat strains. In cattle was attenuated by desiccation of the spi- and horses the signs are variable and nal cord to become the vaccine used by diagnosis may be difficult. In all spe- Louis Pasteur. Modern rabies vaccines cies recovery from disease is extremely for humans and animals are chemically rare. The saliva of infected animals is inactivated purified virus from cell cul- highly infectious, and bites are the usual ture. For prevention of rabies after ani- means of transmission although infec- mal bites (post-exposure prophylaxis, tion through superficial skin lesions is PEP) rabies immunuglobulin (RIG) is possible. The natural reservoir of infec- given with five doses of vaccine. tion varies: in Europe the fox is the most Synonyms : hydrophobia virus; lyssa important. After the Second World War virus; rage virus; tollwut virus; wut foxes became much more numerous virus. and the virus was able to spread slowly Mebatsion T et al (1999) J Virol 73 , 242 from Poland across Europe. Control Rupprecht CE et al (1994) Curr Top Microbiol measures involving the use of oral bait Immunol 187 , 352pp vaccines have been effective in reducing Tordo N et al (2005) In Topley & Wilson’s fox rabies in most European countries. Microbiology and Microbial Infections , vol. 2, 397 raft culture system

Tenth edition, edited by BWJ Mahy and consist of periodic high amplitude V ter Meulen, London: Hodder Arnold, slow-wave complexes in the EEG that p. 1102 are synchronous with myoclonic jerks recurring at 3.5–20 s intervals. In asso- Rabok virus See Tanjong Rabok virus . ciation with very high titers of measles antibodies in the serum these EEG raccoon papillomavirus A possible species changes are considered pathognomic in the family Papillomaviridae . During a for SSPE. field trial of vaccinia-rabies recombinant oral vaccine for raccoons, on Parramore Radi virus (RADIV) A tentative spe- Island, VA, USA, two of 53 live-trapped cies in the genus Vesiculovirus . Isolated raccoons had papillomatous lesions of from Phlebotomus perfiliéwi in central the skin. Papillomavirus antigens were Italy. Not reported to cause disease in detected in the skin lesions by immu- humans. nohistochemistry using a broadly cross- reactive rabbit polyclonal antibody. radiation An effective means of inactivat- Hamir AN et al (1995) J Vet Diagn Invest ing virus infectivity. Clarified virus/ 7 , 549 antigen is exposed in an unbreakable container in an ice bath to a 60 Co- γ raccoon parvovirus (RPV) A strain of source. The dosage required for com- Feline panleukopenia virus in the genus plete inactivation of the virus is esti- Parvovirus . mated from dose-response curves. Gamble WC et al (1980) J Clin Microbiol raccoon rabies virus A strain of Rabies 12 , 676 virus that was introduced by hunters from Florida to Virginia, has become radial immunodiffusion A serological test enzootic in raccoons, Procyon lotor , and in which the antigen, placed in wells in has spread throughout the eastern USA agar gel containing antibody, diffuses during the past 20 years. radially into the agar and the resulting Biek R et al (2007) Proc Natl Acad Sci 104 , antigen–antibody complex forms a halo 7993 or ring of precipitate around the well. Rupprecht CE and Smith JS (1994) Semin Virol 5 , 155 radioimmunoassay (RIA) A serological test in which one of the reactants, Raccoonpox virus (RCNV) A species in usually the antibody, is labeled with the genus Orthopoxvirus . Isolated from a radioisotope; 125I is the most com- raccoons, Procyon lotor , in Maryland, monly used. As the amount of isotope USA. Experimental inoculation of rac- precipitated with the antigen–antibody coons resulted in a silent infection and complex can be measured, it is a very antibody production. Injection into accurate and sensitive method for quan- the footpads of suckling mice caused tifying immunoprecipitation. swelling and paralysis of the hind legs. Replicates on the CAM produc- radioimmunoprecipitation (RIP) A ing pinpoint white pocks, but growth method used for characterizing virus- decreases on passage. Replicates in Vero induced protein synthesis in infected cells with CPE. Analysis of the genome cells. Immune complexes are separated suggests that both raccoonpox and vole- by polyacrylamide gel electrophoresis pox are phylogenetically distant from and characterized on the basis of gel other orthopoxvirus species that are not mobility. Radioactive amino acids used indigenous to the Americas. for RIP are usually 35 S-methionine or Knight JC et al (1992) Virology 190 , 423 35 S-cysteine for general labeling, or 3 H-mannose for labeling glycoproteins. Radermecker complexes A brain disturbance that occurs in cases of sub- raft culture system A system for cell cul- acute sclerosing panencephalitis, a rare ture which allows differentiation of complication of measles virus. They epithelial cells and has allowed growth raft culture system 398

of human papillomaviruses in vitro for which may affect up to 90% of the pop- the first time. ulation. The disease is seasonal, and Ozbun MA (2002) J Virol 76 , 11291 frogs excrete virus in urine when they are kept at 4°C but not if kept at 25°C. The virus also induces tumors experi- rage virus Synonym for Rabies virus . mentally in the kidneys of Rana pipiens , R clamitans, and R palustris . The DNA rainbow trout virus (RTV) A strain of genome is 220 kbp in length, with a Epizootic hematopoietic necrosis virus in G C content of 45%, and is extensively Ranavirus the genus . methylated. The virus has not been grown in cell cultures. Raji cells (CCL 86) A lymphoblastoid cell Synonym : Lucké frog herpesvirus. line established from a Burkitt’s tumor of the maxilla in an 11-year-old black Davison AJ et al (2006) J Gen Virol 87 , 3509 boy. Grows in suspension. The majority McKinnell FG and Carlson DL (1997) J Cell Physiol 173 , 115 of the cells in culture contain the EBNA antigen. ranid herpesvirus 2 (RaHV-2) An unassigned virus in the family Herpesviridae . Raltegravir An anti-HIV drug which Isolated from the urine of Lucke tumor- inhibits the integration of HIV DNA by bearing frogs, but genetically and blocking the viral integrase. Approved antigenically distinct from ranid herpes- for use by the FDA in September 2007. virus 1. Not oncogenic. The DNA genome is 231 kbp in length, with a Ranavirus A genus in the family G C content 56% and is extensively Iridoviridae , type species Frog virus 3 . methylated. Grows in frog embryo cell Particle diameter is 150 nm in thin sec- cultures at 25°C. tion. The capsid has skew symmetry Synonyms : frog herpesvirus 4; frog with T 133 or 147. Viruses grow in virus 4. avian, fish, or mammalian cells and rapidly shut off host cell macromolecular Davison AJ et al (2006) J Gen Virol 87 , 3509 Granoff A (1999) In Encyclopedia of Virology , synthesis. The linear dsDNA genome is Second edition, edited by A Granoff and 170 kbp in length, G C is 53%; highly RG Webster. London: Academic Press, p. 51 methylated (25% of dC residues). It is terminally redundant and circularly ranikhet disease virus Synonym for permuted. There are six recognized Newcastle disease virus . species in the genus, and there are three tentative species members which share RANTES Regulation upon A ctivation sequences with the gene for the major N ormal T cell E xpressed and S ecreted. capsid protein of FV-3, and await fur- A beta chemokine. ther investigation. Primers for PCR amplification of Ranavirus sequences Rat coronavirus (RtCoV) A species in the have been described. genus Coronavirus, belonging to group Goorha RM and Granoff A (1999) In 2 species. Isolated from rats with sialo- Encyclopedia of Virology , Second edition, dacryoadenitis. A natural infectious edited by A Granoff and RG Webster. disease of rats, easily missed as it has London: Academic Press, p. 582 a low mortality. There is fullness of the Mao J et al (1997) Virology 229 , 212 neck due to enlargement of the salivary glands. The submaxillary, salivary, Rangifer tarandus herpesvirus Synonym and Harderian glands are the glands for Cervid herpesvirus 2 . mainly involved and there is necrosis of the ductal epithelium with acute ranid herpesvirus 1 (RaHV-1) An unas- lymphocytic infiltration and gelatinous signed virus in the family Herpesviridae . edema. Seromucinous glands are not A natural infection of Rana pipiens in affected. Red tears and staining of the north, central, and north-eastern parts of fur around the eyes due to porphyrins, the USA and adjacent southern Canada. excreted in tears, may occur. There is Causes renal carcinoma in these frogs, repair of the tissues which is complete 399 rate zonal centrifugation

in 2 weeks. Injection i.c. into newborn rat type C retrovirus A possible spe- mice causes ataxia, paralysis and death cies in the genus Gammaretrovirus . 10 days after injection. Four-week-old An endogenous non-oncogenic virus mice are resistant. Antigenically related designated WF-1 was spontaneously to mouse hepatitis virus. Does not released by a cell line WF derived from agglutinate erythrocytes of mice, chick- a normal Wistar–Furth rat embryo. ens, humans, sheep, rabbit, guinea pig Closely related to two viruses (R-35 and or goose. Contains an acetyl esterase RMTDV) produced by cell lines derived specific for 4-O acetylated sialic acid. from rat mammary tumor tissue. All The genome RNA sequence is distinct three are morphologically and anti- from those of the mouse coronaviruses genically similar but only rat mammary MHV JHM and MHV A59. Can be tumor-derived virus (RMTDV) causes propagated in suckling rats and mice leukemia on injection into rats. Can pro- or adapted to replicate in primary rat vide coat proteins which renders mouse kidney cell cultures with the formation sarcoma virus oncogenic for the rat. of multinucleate giant cells. Bronson DL et al (1976) Proc Soc Exp Biol Med Synonym : sialodacryoadenitis virus of 152 , 116 rats. Bhatt PN et al (1972) J Infect Dis 126 , 123 rat virus R Synonym for Kilham rat virus Stasser P et al (2004) Glycoconj J 20 , 551 in the genus Parvovirus . Yoo D et al (2000) Clin Diagn Lab Immunol 7 , 568 rational drug design The development of antiviral drugs originally depended rat cytomegalovirus (RCMV) Synonym upon random screening of large num- for Murid herpesvirus 2 . bers of random compounds against virus growth in cell culture. With a rat encephalomyelitis virus (REV) better understanding of the molecular A strain of Theilovirus in the genus basis of replication of many important Cardiovirus . viruses, including the availability of the entire nucleotide sequences of their rat minute virus 1 A tentative species in genomes and the three dimensional the genus Parvovirus . protein structure derived from X-ray diffraction analysis, compounds can be rat parvovirus See Kilham rat virus . designed to interact with and inhibit specific targets in the replication cycle. rat parvovirus 1 A tentative species in the This has greatly improved the treat- genus Parvovirus . ment and control of virus infections. rat polyomavirus A possible species in the genus Polyomavirus . Isolated from rattlesnake orthoreovirus (RRV) A pos- an athymic nude rat, in which it caused sible species in the genus Orthoreovirus . sialoadenitis. Isolated from the brain of a rattle- Ward JM et al (1984) Lab Anim 18 , 84 snake with central nervous symptoms. Causes syncytial giant cells in Vero cell rat rotavirus See Rotaviruses group A culture. The genome contains 10 seg- and B . ments of double-stranded RNA. Vieler E et al (1994) Arch Virol 138 , 341 rat sarcoma virus A possible species in the genus Gammaretrovirus . A stable rate zonal centrifugation A form of den- transforming virus of rat origin formed sity gradient centrifugation. As the by the combination of a rat ecotropic particles begin to sediment under the type C virus with src genes. influence of centrifugal force, they Rasheed S et al (1978) Proc Natl Acad Sci become separated into different zones, 75 , 2972 each containing particles of similar sedimentation rate. Particles studied rat submaxillary gland virus Synonym by this technique should have a den- for Murid herpesvirus 2 . sity greater than that at any point in rate zonal centrifugation 400

the supporting gradient column, and Razdan virus (RAZV) A tentative virus the run should be brought to an end in the family Bunyaviridae . Isolated in before any separate zone reaches the suckling mice from a pool of female bottom. See also isopycnic gradient ticks, Dermacentor marginatus , collected centrifugation . from sheep in Razdansk Region, Armenian SSR. Diameter 100 nm by Rauscher leukemia virus A strain of electron microscopy. Infectivity not Murine leukemia virus in the genus sensitive to 5-bromo-2deoxyuridine Gammaretrovirus, related to Friend but sensitive to ether. Hemagglutinates murine leukemia virus. Obtained by goose erythrocytes. Not reported to passage of filtrates of leukemia mouse cause disease in humans. tissue in newborn BALB/c mice. A Lvov DK et al (1978) Acta Virol 22 , 506 defective virus requiring a helper leukemia virus for replication. Genome Rb tumor suppressor proteins Retino contains specific leukemia-producing blastoma susceptibility gene products, a sequences very like or identical to those family of 100–130 kDa proteins involved in Friend virus spleen focus-forming in cell growth regulation and tumor virus and other sequences related to a suppression. Oncogenic viruses such as different helper virus. adenovirus, SV40 and papillomaviruses produce proteins which bind to Rb pro- RAV R ous- a ssociated v irus. teins and inactivate them, resulting in cell transformation and neoplasia. See RAV-0 virus An endogenous chicken pRB . virus which replicates in some chicken cells, but very poorly, and does not R2C cells (CCL 97) A steroid-secreting cause any disease in chickens; this is cell line derived from a transplantable probably related to its LTR sequences. Leydig cell tumor of a rat. Belongs to subgroup E of chicken leukosis sarcoma viruses. Recombination RD cells (CCL 136) A cell line estab- with horizontally transmitted wild lished from a malignant embryonal strains of virus leads to a very virulent rhabdomyosarcoma of the pelvis of a virus. Indistinguishable from RAV-60 7-year-old Caucasian female. in its envelope properties. Robinson HL et al (1980) Cold Spring Harbor RD114 virus A strain in the genus Symp Quant Biol 44 , 1133 Gammaretrovirus isolated when a human rhabdomyosarcoma cell line (RD) was RAV-60 virus A Rous-associated virus passed in cat fetuses developing in utero . in the genus Alpharetrovirus , isolated One kitten at birth contained RD tumor when fowl cells with the chick helper cells which were producing a type C factor were infected with an avian oncovirus. Differs in major species spe- leukemia virus RAV-1 or RAV-2. It cific protein and reverse transcriptase belongs to subgroup E of the chicken from exogenous cat type C virus, but leukosis sarcoma virus. Propagates very similar or identical to feline endo- very much more rapidly in quail cells genous type C retrovirus. It is a xeno- than RAV-0 does but is indistinguish- tropic virus unable to infect cat cells able from RAV-0 in its envelope prop- and was at first thought to be a human erties. Unlike RAV-0, it causes leukosis virus. No association with disease. in chickens. Multiple copies of RD114 viral genomes are found in all domestic cat cells, but Hanafusa T et al (1970) Proc Natl Acad Sci these are not normally expressed. Most 67 , 1797 wild cats do not have these endogenous sequences. The virus is related to Raza virus (RAZAV) A serotype of an endogenous retrovirus of baboons, Hughes virus in the genus Nairovirus . and may have been acquired from Isolated from ticks. Not reported to African primates several million years cause disease in humans. ago. Receptors for RD114 virus have 401 recombinant DNA

been identified on human cells, and viruses which have multisegmented the virus is being used experimentally genomes, such as Orthomyxoviridae and as a pseudotype vector for therapeutic Reoviridae . gene transfer into human hematopoietic stem cells. reassortment Closely related viruses with Germain E et al (2005) J Gene Med 7 , 389 segmented genomes under genetic Kelly PF et al (2000) Blood 96 , 1206 reassortment during dual infections. Niman HL et al (1977) Nature 226 , 357 Different subtypes of influenza virus can reassort to produce new viruses in RDE R eceptor- d estroying e nzyme. See this way, but no reassortment occurs neuraminidase . between influenza types A, B, and C. The basis for this restriction is not known. reactivation A special type of recombi- Each of the major human pandemics of nation. When one or more of the virus influenza in the twentieth century (1918, particles involved in a multiple infec- 1957, and 1968) was caused by reassort- tion is inactivated, and unable alone to ment between an existing human virus initiate a productive cycle, it neverthe- and an avian influenza virus. less can contribute to the production of progeny virus with the assistance of receptor A site or structure on the cell the other viruses, i.e. it is ‘ reactivated. ’ surface to which a virus binds. A host When the virus particles involved dif- surface component that participates in fer genetically, the process is known as virus binding and facilitates viral infec- ‘ cross-reactivation ’ or ‘ marker rescue. ’ tion. See CD . If they are genetically identical and all Haywood AM (1994) J Virol 68 , 1 are inactivated, the process is known as ‘ multiplicity reactivation. ’ ‘ Non- receptor-binding protein The protein on genetic reactivation ’ is in fact not a case the surface of the virion which binds to of reactivation, but a special example of the host cell receptor. complementation of a virus inactivated by damage to its proteins rather than its genome nucleic acid. See Berry– receptor-destroying enzyme (RDE) See Dedrick phenomenon . neuraminidase . reading frame A sequence of codons in receptor-mediated endocytosis A proc- RNA or DNA beginning with the ini- ess by which many viruses gain entry tiation codon AUG. See open reading to the cell. In order to fuse with the cell frame . membrane a mildly acid environment is required (pH 5–6). The virus enters the cell by receptor-mediated endocytosis readthrough The reading of an mRNA into endosomes whose contents become through a stop codon. A suppressor acidified by an ATP-driven proton tRNA causes the insertion of an amino pump. The virus fuses with the mem- acid into a growing polypeptide chain brane of the endosome and releases the in response to the stop codon. The viral genome into the cytoplasm. The readthrough protein is thus longer than membrane-bound protein Rab5 is a the usual polypeptide. See also protein positive regulator of endocytosis. synthesis , ribosomal frameshifting . recognizate A term proposed, but not reannealing The coming together of widely adopted, to describe the demon- complementary strands of nucleic stration of the presence of a virus, and acid after separation by melting. See its recognition without it having been hybridization . cultivated in pure culture. Cf . isolate . reassortants Viruses which have derived Madeley CR and Kay CJ (1978) Lancet ii , 733 parts of their genomes from two viruses involved in a mixed infection. This proc- recombinant DNA DNA molecules ess is particularly likely to occur with containing novel sequences, formed recombinant DNA 402

by in vitro recombination with non- in the genus Megalocytivirus, family homologous molecules. Iridoviridae. Causes an acute highly contagious disease with high mortality in recombinant RNA RNA virus genomes cultured red sea bream and at least 30 constructed by genetic engineering of other marine fish species in south-west infectious DNA clones. Intertypic chi- Japan. Affected fish have severe anemia, meras of polioviruses have been con- petechiae of the gills, and enlargement of structed in this way. the spleen. A virus which appears to be an iridovirus can be cultured from dis- recombinants Viruses containing nucleic eased fish on various fish cell lines. An acid sequences from two or more dif- effective formalin-killed vaccine given ferent virus genomes. They may be by i.p. inoculation has been described. formed naturally in either of two ways: Lua DT et al (2005) J Virol 79 , 15151 intramolecular recombination, which Matsuoka S et al (1995) Fish Pathol 31 , 115 involves transfer of sequences within Nakajima K et al (1999) Dis Aquat Org 36 , 73 single molecules of nucleic acid, or genetic reassortment, in which viruses redfin perch virus (RFPV) A strain of whose genomes are fragmented into Epizootic hematopoietic necrosis virus , a a number of pieces (e.g. Arenaviridae , species in the genus Ranavirus . Bunyaviridae , Orthomyxoviridae, or Reo- viridae ) exchange whole pieces (seg- Redspotted grouper nervous necrosis ments) of RNA. virus (RGNNV) A species in the genus Betanodavirus . recombination The exchange of genetic material from two or more virus par- Redwood Park virus (RPV) A strain of ticles into recombinant progeny virus Frog virus 3 in the genus Ranavirus . during a mixed infection. Reed Ranch virus (RRV) An unassigned recurrent respiratory papillomatosis species in the family Rhabdoviridae , (RRP) A common benign laryngeal belonging to the Bahia Grande sero- tumor, caused by human papilloma- group. Isolated from mosquitoes, Culex virus. There may be significant morbid- salinarius , in Texas, USA in 1974. Not ity due to an effect on patency of the associated with disease in humans. airways. Treatment is often difficult, usually requiring multiple surgical Reed–Sternberg cells Giant histiocytes, interventions. The incidence in chil- typically multinucleate or binucleate dren is estimated at 4.3 per 100,000 and that contain prominent nucleoli and in adults 1.8 per 100,000. are commonly seen in Hodgkin’s dis- Dickens P et al (1991) J Pathol 165 , 243 ease, where they may be infected by Epstein–Barr virus. red deer herpesvirus Synonym for Cervid herpesvirus 1 . reference types Virus strains held in culture collections such as the American red disease of pike virus Synonym for Type Culture Collection (ATCC) or pike fry rhabdovirus. under the auspices of the World Health Organization (WHO) and which may red kangaroo poxvirus An unassigned be used in the identification and typing virus in the family Poxviridae . of new virus isolates. Bagnall BG and Wilson GR (1974) Aust J Dermatol 15 , 115 regina ranavirus (RRV) A strain of Ambystoma tigrinum virus in the genus red nose virus Synonym for Bovine her- Ranavirus . pesvirus 1 . regressive evolution One theory of the red sea bream iridovirus A strain of origin of viruses is that they were the Infectious spleen and kidney necrosis virus result of regressive evolution of more 403 replicative form

complex microbial forms that had cel- Dermody TS and Desselberger U (2005) In lular organization and metabolism. Topley & Wilson’s Microbiology and Microbial Infections , vol. 2, Tenth edition, edited by reindeer herpesvirus BWJ Mahy and V ter Meulen, London: Synonym for cervid Hodder Arnold, p. 932 herpesvirus 2. Nibert ML et al (1991) J Clin Invest 88 , 727 reindeer papillomavirus (RePV) A repeated sequence A nucleotide sequence strain of European elk papillomavirus in which occurs more than once in a DNA the genus Deltapapillomavirus , isolated or RNA molecule either in the same from the epithelial layer of a cutaneous (direct repeats) or opposite (inverted fibropapilloma on a Swedish reindeer, repeats) orientation. Rangifer tarandus . Morphologically Synonym : reiterated sequence. indistinguishable from other papillomaviruses, but the restriction enzyme cleavage pattern of its genome is dif- repetitive DNA DNA sequences which ferent. RePV is oncogenic for ham- occur repeatedly and comprise 20–50% sters and transforms C127 mouse cells of the chromosomal DNA of animal in vitro . Transforming properties corre- genomes. Can be divided into slightly late with a highly conserved E5 region. repetitive DNA (1–10 copies per haploid genome), moderately repetitive DNA Moreno-Lopez J et al (1987) J Virol 61 , 3394 (ten to thousands of copies per haploid genome) and highly repetitive DNA reiterated sequence See repeated (thousands to millions of copies per hap- sequence . loid genome). Highly repetitive DNA is often found as spacer DNA between Relenza An inhaled anti-neuraminidase structural genes (unique DNA). drug effective against influenza. Britten RJ and Kohne DE (1968) Science 161 , 529 Reoviridae (acronym: respiratory, e nteric, Davidson EH and Britten RJ (1971) Q Rev o rphan) A family of unenveloped Biol 46 , 111 double-stranded RNA viruses infecting vertebrates, invertebrates, higher replicase An RNA-dependent RNA plants, bacteria, and fungi. The term polymerase catalyzing the formation ‘ reovirus ’ was coined by Albert Sabin of new virion RNA from a comple- in 1959 because the human r espiratory mentary strand template. See RNA- and enteric isolates were ‘ o rphans, ’ i.e. dependent RNA polymerase . not associated with disease. Virion has an isometric two-layered capsid with replication The multiplication of a virus, icosahedral symmetry, 60–80 nm in involving virus attachment to a host diameter, usually naked but pseudo- cell, entry, gene expression, mRNA membranes, probably of host origin, synthesis, protein synthesis, genome are described. Buoyant density (CsCl): replication, and virus assembly and 1.31–1.38 g/ml; 630S. Virion contains an release. RNA-dependent RNA polymerase. The RNA genome is in 10, 11, or 12 segments Rowlands DJ (2005) In Topley & Wilson’s of total mol. wt. 12–20 1 0 6, all within Microbiology and Microbial Infections , vol. 1, a single virion. G C content 42–44%. Tenth edition, edited by BWJ Mahy and V ter Meulen, London: Hodder Arnold, p. 105 Resist lipid solvents. Genetic reassortment occurs readily within genera. Viral synthesis and maturation occur replication and transcription activator in the cytoplasm. Inclusion bodies with A key immediate-early regulator pro- crystalline arrays of virus particles are tein of human herpes virus 8 encoded often seen. The vertebrate species are by ORF50 and required for the switch in five genera: Orthoreovirus, Orbivirus , from latent infection to lytic replication. Rotavirus, Coltivirus, Seadornavirus, and Aquareovirus . There are also six genera replicative form See RNA-dependent of insect and plant viruses. RNA polymerase . replicative intermediate 404 replicative intermediate See RNA- Table R2. Examples of reptilian viruses dependent RNA polymerase . boid herpesvirus 1 replicons One theory of the origin of Chaco virus viruses is that they are descendents chelonid herpesvirus 1–4 of primitive precellular replicons that corn snake retrovirus existed before the first cellular forms Elaphe virus existed on earth. elapid herpesvirus 1 fer-de-lance virus representational difference analysis gecko virus (RDA) A virus detection technique in green lizard papillomavirus which DNA molecules are compared. iguanid herpesvirus 1 Used e.g. to find human herpes- lacertid herpesvirus 1 virus 8 (Kaposi’s sarcoma-associated Marco virus herpesvirus). python orthoreovirus Lisitsyn N et al (1993) Science 259 , 946 rattlesnake orthoreovirus reptile calicivirus repressor A protein which prevents DNA- Vesicular stomatitis virus dependent RNA polymerase from Viper retrovirus starting RNA synthesis by binding to a specific DNA sequence upstream of the transcription initiation site. Rescriptor An anti-HIV drug which is reptile calicivirus (Cro-1) (VESV/Cro-1) a reverse transcriptase inhibitor. The A strain of Vesicular exanthema of swine generic name is delaviridine . virus in the genus Vesivirus . rescue See reactivation . reptilian virus group A group in the genus Gammaretrovirus. Natural host reservoirs Many vertebrate viruses exist in range restricted to reptiles. Type spe- reservoir species from which they may cies Viper retrovirus . Another probable emerge to cause a disease outbreak. species is corn snake retrovirus. There This is particularly true of alphaviruses, is also morphological evidence for ret- which are maintained in zoonotic trans- roviruses associated with neoplasia in a mission cycles in birds or mammals California king snake, Lampropeltis get- from which they are transmitted to the ulus californiae, and a four-lined chicken human population by arthropods. The snake, Elaphe obsoleta quadrivittata . reservoir for influenza viruses is mainly Poulet FM et al (1994) In The Retroviridae , aquatic birds. There remain serious vol. 3, edited by JA Levy. New York: Plenum human diseases for which the reservoir Press, p. 1 species remains unknown, e.g. Ebola virus and Marburg virus. reptilian viruses Serological evidence of infection with several virus species has Resiquimod An immune response mod- been found in snakes, alligators, liz- ulator which was considered as a pos- ards, and turtles. Japanese B virus and sible anti-herpes virus drug, but was Western equine encephalitis virus may not successful in clinical trials. over-winter in snakes and turtles, and Tomai MA et al (1995) Antiviral Res 28 , 285 Eastern equine encephalitis virus may do so in lizards. resistance-inducing factor (RIF) A fowl Lunger PD and Clark HF (1978) Adv Virus leukemia virus which interferes with Res 23 , 159 the growth of Rous sarcoma virus.

Reptilian orthoreovirus A species in the Resistencia virus (RTAV) An unas- genus Orthoreovirus isolated from a signed virus in the family Bunyaviridae . python. One of a subgroup of orthoreo- Grouped with Antequera virus and viruses which are fusogenic, capable of Barranqueras virus . Isolated from Culex inducing syncytium formation. (Melanoconion) delpontei in Argentina. Duncan R et al (2004) Virology 319 , 131 Not associated with disease in humans. 405 restriction fragment length polymorphism

Resolvase An enzyme encoded by vac- disease symptoms. Two further out- cinia virus which is required during breaks occurred in association with replication for resolution of concate- monkeys imported from the same meric DNA into monomers. Phillippines facility, in Siena, Italy in Garcia AD and Moss B (2001) J Virol 75 , 6460 1992, and in Texas, USA in 1996. The company providing the monkeys has respiratory infection virus Synonym for now ceased trading. The Reston Ebola Equid herpesvirus 4 . virus is highly pathogenic for monkeys, and whilst it has been suggested that it may be less pathogenic for humans this respiratory enteric orphan viruses The has not been put to the test, for obvious full name for reoviruses. reasons. The following strains of Reston Ebola virus are recognized: Reston Ebola respiratory and reproductive syndrome virus Philippines 1989; Reston Ebola virus virus See Porcine respiratory and Reston 1989; Reston Ebola virus Siena reproductive syndrome virus . 1992; Reston Ebola virus Texas 1996 . respiratory syncytial virus of bovines restriction endonuclease mapping A Synonym for Bovine respiratory syncytial technique for relatively rapid compari- virus . son of DNA molecules based upon the size of fragments yielded in a standard respiratory syncytial virus of humans reaction with a restriction endonucle- Synonym for Human respiratory syncy- ase. The procedure may identify restric- tial virus . tion sites and the sites of insertions or deletions in the DNA under test Respirovirus A genus in the family relative to a standard DNA of known Paramyxoviridae. The type species is sequence. A practical application is Sendai virus . Members of the genus Pulse-Net, a system for analyzing the have a hemagglutinin and a neurami- relationship between food-borne bacte- nidase and have six transcriptional rial pathogens, by restriction fragment elements. All members encode a C pro- length polymorphism (RFLP). tein. The unedited P mRNA encodes P and C, and editing by insertion of a single G accesses the open reading restriction endonucleases Bacterial frame for V. In addition to Sendai virus , enzymes with a role in the host spe- Simian virus 10 , Bovine parainfluenza cificity of bacteriophages. They are virus 3, and Human parainfluenza viruses highly specific, recognizing a particu- 1 and 3 are members of the genus. lar sequence of nucleotides, where they attach and cut the nucleic acid chain. Thus they can inactivate foreign, Restan virus (RESV) A serotype of incoming DNA. Host cell DNA is not Marituba virus in the genus Orthobunya- cut because the specific sites have been virus belonging to the C-group viruses. methylated, and thus protected, by a Isolated from Culex sp mosquitoes in methylase enzyme also present in the Trinidad and Surinam. Can cause a bacterial cell. In response to host cell febrile illness in humans. restriction systems, some bacteriophages encode inhibitory proteins that can bind Reston Ebola virus (REBOV-Res) A spe- to and inactivate restriction endonu- Ebolavirus cies in the genus . In 1989 cleases. Others, such as T-even phages, an outbreak of hemorrhagic fever modify their own DNA by methylation occurred in a nonhuman primate facil- corresponding to that occurring in cer- ity in Reston, Virginia, USA involving tain bacterial hosts, so are not restricted. Cynomolgus monkeys imported from the Phillippines. Ebola virus was diag- Roberts RJ (1978) Nature 271 , 502 nosed as the cause of the outbreak and all monkeys were destroyed. At least restriction fragment length polymor- one case of demonstrated human infec- phism A useful method for comparing tion occurred, but with no associated the relatedness of genomes. The virion restriction fragment length polymorphism 406

nucleic acid is digested with specific Synonym : avian reticuloendotheliosis restriction endonucleases and the frag- virus. ments separated using a polyacryla- Payne LN (1992) In The Retroviridae, vol. 1, mide gel. Especially useful in outbreak edited by JA Levy. New York: Plenum Press, studies of rotaviruses. p. 365 restriction fragments Fragments of viral retinoic acid A compound which or cellular DNA produced by non-ran- inhibits the replication of human dom cleavage of the DNA with specific papillomavirus. endonucleases. The fragments of a Bartsch D (1992) EMBO J 11 , 2283 particular DNA are characteristic. See restriction endonuclease mapping . retroid elements Reverse transcription Reticuloendotheliosis virus (REV) A is not uniquely associated with retro- species in the genus Gammaretrovirus . viruses and hepadnaviruses. A variety Avian viruses for which related endo- of transposable elements from yeast, genous sequences have been found Drosophila, Dictyostelium and maize only in mammals, not birds. There are have structural similarities to integrated two recognized strains, A and T, and forms of the retrovirus genome and two related species: Chick syncytial virus these are known as retroid elements or and Trager duck spleen necrosis virus . All retrotransposons. three are closely related, but cross-react hardly at all with chicken leukosis sar- retroperitoneal fibrosis (RF) An infre- coma viruses. They do not have nucle- quent disease syndrome occurring in otide sequences in common with them. immunosuppressed macaques. Lesions The viruses are all non-defective except consist of aggressively proliferating REV strain T, which has a smaller fibrous tissue with a high degree of genome length (5.7 kb) than the other vascularization. Two novel rhadino- viruses (9.0 kb), and requires a helper viruses were identified in separate virus for its replication. However, strain macaques by PCR, but attempts to iso- T carries the oncogene v-rel which spec- late these viruses in cell culture were ifies a DNA-binding transcription factor unsuccessful. in infected cells and is highly oncogenic, Rose TM et al (1997) J Virol 71 , 4138 inducing immature B-cell lymphomas. All three viruses are probably widely retrotransposons Mobile elements in distributed among turkeys and wild the cell genome which contain reverse water fowl, especially ducks and geese. transcriptase or other ‘ retroid elements’ The only spontaneous disease produced such as LTR, capsid protein, ribo- appears to be turkey leukosis. Chickens, nuclease, integrase, and protease. There quail, ducklings, goslings, turkeys, is a wide variety of ‘ viral ’ and ‘ non- pheasants, and guinea keets are sus- viral’ retrotransposons and related ele- ceptible to virus injected by any route. ments including Gypsy, CopiaLINES A large dose causes death in 3 days. and LINE-like elements, mitochondrial With smaller doses or in older animals introns and retroplasmids. The ‘ viral ’ death is not so rapid and some animals retrotransposons more closely resem- survive. Such birds are often thin, ane- ble retrovirus genomes, and all have mic, retarded, and have poor feather the reverse transcriptase enzyme. Most development. Histological changes proof these elements represent jumping duced by the virus are visceral or neu- genes which sometimes cause useful ral, proliferative lesions and necrotizing rearrangements of host cell DNA lesions. The proportion of each varies sequences and sometimes damage or with the strain of virus. In the prolifera- kill their host by DNA alterations. All tive lesions the cells are histiocytoid and complete eukaryotic genomes sequenced probably malignant. The virus replicates so far have been found to contain trans- in chick embryo fibroblast cultures, pro- posons, and 44% of the human genome ducing a transitory CPE followed by consists of transposons or transposable transformation. elements. 407 reverse transcription polymerase chain reaction (RT-PCR)

Bushman F (2002) Lateral DNA Transfer. retroviruses Species in the family Mechanisms and Consequences . New York: Retroviridae . Cold Spring Harbor Laboratory Press. rev protein An RNA-binding nuclear protein induced in cells infected by Retrovir The first effective inhibitor of human immunodeficiency viruses retroviruses, better known as AZT (azi- (HIV-1 and HIV-2) that regulates virus dothymidine). See AZT . gene expression and is essential for virus replication. Binds to a region of Retroviridae A family of large single- HIV RNA known as the rev response stranded RNA viruses which have a vir- element (RRE), and controls splicing so ion RNA-dependent DNA polymerase. that single-spliced and unspliced RNA Virions are spherical, about 80–100 nm species are produced in the later stages in diameter. Lipoprotein envelope of virus replication. encloses an icosahedral core shell within which there is a helical nucleocapsid. rev response element (RRE) A specific The envelope has glycoprotein surface sequence in the retroviral RNA genome projections 8 nm long. The genome is a that interacts with a sequence in the dimer of two hydrogen-bonded posi- host cell genome. tive single-stranded RNAs, each monomer 7–11 kb in length. Viral RNA is reverse genetics The recovery of negative- transcribed by the virion transcriptase strand virus RNA as DNA to facilitate into a covalently linked circle of genome studies. double-stranded DNA (provirus) which Nagai Y and Kato A (1999) Microbiol becomes integrated into the cellular Immunol 43 , 613 DNA. Viral RNA serving as mRNA Roberts A and Rose JK (1998) Virology 247 , 1 and virion RNA for progeny particles is transcribed from the integrated DNA reverse passive hemagglutination A sen- provirus. Replication is sensitive to sitive serological test in which red blood inhibitors of DNA synthesis during the cells are coated with virus-specific first 6 h after infection, and to actino- antibody and used to test for the pres- mycin D at any time. Maturation occurs ence of antigen. If virus antigen is by budding from the cytoplasmic mem- present, the red blood cells are aggluti- branes. Provirus DNA extracted from nated. See passive hemagglutination . infected cells is infective. There are two subfamilies, Orthoretrovirinae with six reverse-transcribing viruses Members of genera: Alpharetrovirus, Betaretrovirus , the families Retroviridae and Hepadna- Gammaretrovirus, Deltaretrovirus, Epsilon- viridae, both of which contain an RNA- retrovirus, Lentivirus , and Spumavirinae dependent DNA polymerase which is with a single genus: Spumavirus. Retro- central to their process of replication. viruses are associated with a variety There are also families of plant, fungal, of animal diseases as well as human and invertebrate viruses which contain diseases including AIDS, autoimmune the enzyme ( Pseudoviridae , Metaviridae , disease and lower motor neurone dis- and Caulimoviridae ), but no bacterial ease, but may also be non-pathogenic. viruses have been found which utilise Retroviruses may be present as endo- reverse transcriptase. genous DNA in all vertebrate genomes. Synonym : ribodeoxy virus. reverse transcriptase Synonym for RNA- dependent DNA polymerase. Coffin JM (1992) In The Retroviridae, vol. 1, edited by JA Levy. New York: Plenum Press, reverse transcription Transcription of p. 19 RNA into DNA. Dezutti CS et al (2005) In Topley & Wilson’s Microbiology and Microbial Infections , vol. 2, Tenth editionn, edited by BWJ Mahy and reverse transcription polymerase chain V ter Meulen, London: Hodder Arnold, reaction (RT-PCR) A frequently emp- p. 1284 loyed method to prepare DNA copies Herniou E et al (1998) J Virol 72 , 5955 of an RNA virus, which can then be reverse transcription polymerase chain reaction (RT-PCR) 408

amplified by PCR and used for seq- RFL-6 cells (CCL 192) A fibroblast-like uence analysis or other purposes. cell line derived from the lung of a normal, germ-free, Sprague–Dawley reversion A change in nucleotide rat fetus. sequence that reverses the mutation at the original site and restores the origi- RFLP Restriction fragment length nal phenotype. polymorphism. rex protein An RNA-binding protein, Rhabdoviridae A large family of RNA induced in cells infected with human negative-strand viruses comprising T-cell leukemia viruses types 1 and 2, six genera and 75 species which infect which influences splicing and trans- vertebrates, invertebrates, and plants. port of viral mRNAs. Many species are pathogenic and transmitted by arthropods. Rhabdoviruses are rod-shaped, varying in length Reyataz An HIV protease inhibitor. See (100–430 nm) but more uniform in diam- Atazanavir . eter (45–100 nm). The animal species are bullet-shaped, being flattened at one Reye’s syndrome A neurological and end and pointed at the other, whereas metabolic disease of children and ado- the plant species are rounded at both lescents first described in Australia in ends. They all have a membranous 1963. Characterized by encephalo- envelope with spikes 5–10 nm long. The pathy and fatty degeneration of the liver. envelope is disrupted by lipid solvents. The syndrome has been observed to Wound inside the envelope is a helical follow infections with influenza virus nucleocapsid with a diameter of 50 nm. A, influenza virus B, parainfluenza There is one molecule of negative-sense virus, human adenovirus, human her- single-stranded RNA 11–15 kb in length pesvirus 3, and human respiratory which is not infective and is transcribed syncytial virus. An acute encephalopa- by an RNA-dependent RNA polymer- thy with fatty degeneration of the vis- ase in the nucleocapsid into at least five cera with a fatality rate of about 20%. messenger RNA species. Most of the Cause is uncertain but some 40% of viruses which have been studied con- cases have an association with various tain five proteins designated: L (large), drugs or chemicals, especially aspirin G (glycoprotein), N (nucleoprotein), NS and related salicylates. There are often or M1 (nonstructural), and M and M2 symptoms and signs of upper respira- (matrix). The NS protein was originally tory tract infection, and an association thought to be nonstructural but is a with various viruses has been sug- component of the virion RNA polymer- gested, but the evidence is not very ase required for transcription. Defective strong and environmental or constitu- truncated virions (T virions) with a tional factors may be important. The portion of the RNA genome deleted addition of warning notices on aspirin arise frequently during replication. bottles in the USA has helped to reduce Virus attaches to cells by the G protein the number of cases. and enters by endocytosis via coated Synonym : Reye–Johnson syndrome. pits. After uncoating, the virion RNA- dependent RNA polymerase transcribes Belay ED et al (1999) N Engl J Med 340 , 1377 Reye RDK et al (1963) Lancet 2 , 749 capped and polyadenylated RNA species which are translated into virus proteins. RNA replication occurs entirely in RF virus A strain of BK virus. Isolated the cytoplasm and requires no nuclear in human embryo kidney cell culture functions. Nucleocapsids are assem- from the urine of a renal transplant bled and enveloped, then bud from patient. Antigenically indistinguishable the plasma membranes. Four genera, from BK virus and their DNAs have an Lyssavirus , Vesiculovirus , Ephemerovirus , 88% homology. and Novirhabdoviruses contain the Miao R and Dougherty RM (1977) J Gen viruses which replicate in vertebrate Virol 35 , 67 species. 409 ribonuclease B

Tordo N et al (2005) In Topley & Wilson’s RI-1 virus Rhode Island-1 virus, a Microbiology and Microbial Infections , vol. 2, strain of New York virus in the genus Tenth edition, edited by BWJ Mahy and Hantavirus . V ter Meulen, London: Hodder Arnold, p. 1102 ribavirin 1-␤-D-ribofuranosyl-1, 2, 4-triazole-3-carboxamide rhabdovirus 903/87 A novel fish patho- An antiviral agent. genic rhabdovirus. A synthetic nucleoside analog of guanosine. Interferes with biosynthesis of guanylic acid nucleotides. An inhibitor rhabdovirus carpio See spring viremia of IMP dehydrogenase. Active in vitro of carp virus . against a wide range of RNA viruses including picornaviruses, influenza, rhabdovirus entameba See entamoeba parainfluenza, arenaviruses and bunyavirus . viruses. May act by increasing the rate of RNA mutations, in addition to its Rhadinovirus A genus in the subfamily inhibitory effect on RNA synthesis. Also Gammaherpesvirinae . The type species is inhibits cytomegalovirus and vaccinia Saimiriine herpesvirus 2, which has a dis- virus growth. Placebo-controlled studies tinctive genome structure and 13 addi- of hepatitis A patients in Brazil showed tional species and 4 tentative species that the drug accelerated the return to are currently included in the genus. normal of elevated serum bilirubin and of liver enzymes in the serum. A com- rheumatoid arthritis In an attempted parable study in the USA of hepatitis gene therapy for this condition, an B patients revealed no such beneficial adeno-associated virus containing the effect. Has proved useful in treatment gene encoding the receptor for tumor of arenavirus infections, especially necrosis factor (TNF-alpha) was devel- Lassa fever and some South American oped for injection directly into the arenavirus infections. Used in combina- arthritic joint, but in a clinical trial of tion with alpha-interferon to treat 100 patients, a 36-year-old woman died hepatitis C infection, and for severe in August 2007, and the trial has been respiratory syncytial virus infection. discontinued. Animal studies suggest that ribavirin can produce anemia, immunosuppression rhinocerospox virus A possible species in and, in rodents, teratogenesis. the genus Orthopoxvirus isolated from a Synonym : Virazole. Ceratotherium simum white rhinoceros ( ) Crotty S et al (2000) Nat Med 6 , 1375 from a zoo. Oxford JS (1975) J Gen Virol 28 , 409 Pilaski J et al (1986) Arch Virol 88 , 135 Stephen EL and Jahrling PB (1979) Lancet i , 268 Rhinovirus A genus of the family ␤ Picornaviridae , type species is Human 1- -D -ribofuranosyl-1, 2, 4-triazole-3- rhinovirus A . Distinguished from entero- carboxamide See ribavirin . virus by: (1) being unstable below pH 5–6; (2) density (CsCl) of 1.38–1.42 g/ml; ribonuclease A The principal active com- and (3) the disease produced. Species ponent of bovine pancreatic ribo- are Human rhinovirus A (75 serotypes), nuclease; cleaves phosphodiester bonds Human rhinovirus B (25 serotypes), and between pyrimidines and adjacent 3 tentative species, bovine rhinoviruses nucleotides. Pure purine polymers are 1, 2, and 3. They infect the respiratory relatively resistant to attack by RNase tract and transmission is by airborne A, but sufficiently high enzyme con- droplets or contamination with respira- centrations will degrade poly A, for tory tract secretions. example. Minor P (2005) In Topley & Wilson’s Microbiology and Microbial Infections , vol. 1, ribonuclease B A component of bovine Tenth edition, BWJ Mahy and V ter Meulen, pancreatic ribonuclease which can London: Hodder Arnold, p. 857 be separated from ribonuclease A by ribonuclease B 410

ion-exchange chromatography, and is the cytoplasm stimulates 2 ,5 -oligo-A- present in ten-fold lower concentration. synthetase to produce strands of It is a glycosylated form of ribo- 2 ,5 adenylic acid which induce activa- nuclease A, having 6 mannose residues tion of RNase L which then degrades and 2 N-acetylglucosamine residues cellular RNA to produce small RNA per molecule. cleavage products. These small RNAs Gotte G et al (2003) J Biol Chem 278 , 46241 activate the genes that encode alpha- and beta-interferon. Signalling occurs through the interaction of the RNAs ribonuclease C Isolated from human pla- with either of two pathogen recogni- centa. Preferentially cleaves RNA on tion receptors: retinoic acid inducible the 5 side of cytidine residues. gene-1(RIG-1) or melanoma differentiation associated gene 5 (MDA5). ribonuclease D A ribonuclease that removes nucleotides from precursor Malathi K et al (2007) Nature 448 , 816 Silverman RH (2007) Cytokine Growth Factor tRNA molecules, generating the 3 ter- Rev 18 , 381 minus of mature tRNA. ribonuclease P A processing enzyme, ribonuclease from Bacillus cereus A involved in tRNA biosynthesis in ribonuclease which cleaves the 3 - Escherichia coli , which cleaves tRNA phosphodiester bonds after pyrimidine precursor molecules, removing 5 prox- residues. imal nucleotides to generate the 5 termini of mature tRNA molecules. ribonuclease H A ribonuclease which specifically cleaves the RNA strand ribonuclease Phy 1 A ribonuclease iso- present in an RNA–DNA hybrid, but lated from a slime mold, Physarum does not digest free single-stranded or polycephalum. Preferentially cleaves the double-stranded RNA. Can be isolated phosphodiester bonds between gua- from Escherichia coli; also present as nine, adenine, and uracil and adjacent part of the reverse transcriptase activity residues. Used in RNA sequence analy- of retroviruses. Used experimentally to sis to discriminate between cytosine remove poly A tails from mRNA after and uracil. hybridization with poly dT. Simoncsits A et al (1977) Nature 269 , 833 Oberhaus SM and Newbold JE (1995) J Virol 69 , 5697 ribonuclease Phy M A ribonuclease that cleaves on the 3 side of adenine and ribonuclease III A ribonuclease that uridine residues. hydrolyzes double-stranded RNA. An endoribonuclease from Escherichia coli ribonuclease S A preparation of ribo- which cleaves double-stranded RNA nuclease A in which the main peptide to single-stranded RNA of approxi- chain has been cleaved with subtilisin mately 15 nt chain length. The enzyme (proteinase from Bacillus subtilis ). The also cleaves specific sequences in single- enzyme activity of RNase S is very sim- stranded RNA, and is responsible for ilar to that of RNase A. processing large RNA transcripts of ribonuclease T1 A ribonuclease isolated bacteriophage T7 DNA into individual from taka-diastase which cleaves early mRNAs. phosphodiester bonds between 3 Dunn JJ (1975) J Biol Chem 251 , 3807 guanylic acid groups and the 5 hydroxyl groups of adjacent nucleotides. It has ribonuclease L A 2 ,5 -oligoadenylate- also been termed ‘ guanyloribonuclease. ’ dependent ribonuclease which is central to the innate cellular defense ribonuclease T2 A ribonuclease isolated mechanism induced by type 1 interfer- from taka-diastase which cleaves ons during virus infection. Present in phosphodiester bonds between any pair minute quantities during the normal of nucleotides, but displays a preference cell cycle, but the presence of dsRNA in for adenylic acid bonds. 411 ribosomal skipping ribonuclease U2 A ribonuclease found ribonucleoprotein A complex comprising in culture broth of the smut fungus, ribonucleic acid and protein, usually Ustilago sphaerogena , which cleaves an linked by electrostatic bonds. RNA molecule at the phosphodiester bonds of purine nucleotides to yield 3 ribonucleoside A purine or pyrimidine nucleotides with intermediary forma- d base covalently bound to a -ribose tion of purine nucleoside 2 -3 cyclic sugar molecule. See nucleic acid . phosphates. ribonucleic acid (RNA) A polymer of ribonucleotide A ribonucleoside with one or more phosphate groups esteri- ribonucleotides which differs in several respects from DNA. Three of the bases, fied to the 5 position of the sugar moi- adenine, guanine and cytosine, are the ety. See nucleic acid . same as in DNA but uracil replaces thymine and a few minor or modified ribonucleotide reductase (RR) An bases are present, especially in tRNA. enzyme that catalyzes the conversion The d-ribose sugar of RNA differs from of ribonucleoside diphosphates to the 2-deoxy-d -ribose of DNA in having deoxyribonucleoside diphosphates. four rather than three hydroxyl groups. Complementarity of the bases seen in ribose The sugar of ribonucleotides. See DNA is not evident. Thus long dou- nucleic acid . ble strands are not formed but short stretches of base-pairing occur, producing loops. See poly AU. In all types of ribosomal frameshifting A purposeful cells there are three kinds of RNA: shift in reading frame whereby ribo- (1) Ribosomal RNA ( rRNA ) forms about somes respond to signals in the mRNA 80% of the total. It is of high molec- and move into a new reading frame ular weight and is metabolically (usually –1) at a specific point and con- stable. It is of two main types. See tinue translation in the new reading ribosomes . frame. Used in several virus mRNAs, (2) Transfer RNA ( tRNA) forms about especially by retroviruses, astroviruses, 15% of the total mol. wt. 23,000– and coronaviruses. May involve a 28,000. There are tRNA molecules pseudoknot in the mRNA or a shifty or specific for each amino acid and on slippery sequence, or both. each molecule is an anticodon that Brierley I (1995) J Gen Virol 76 , 1885 locates to a codon on the mRNA and so brings the amino acids into ribosomal RNA See ribonucleic acid and correct sequence in the polypeptide ribosomes . being formed. (3) Messenger RNA (mRNA) forms about 5% of the total and has a base com- ribosomal scanning model The hypoth- position corresponding very closely esis that initiation of protein synthesis to DNA. The DNA base sequences involves scanning of the 40S ribosomal transcribed and processed into subunit along the mRNA molecule until it encounters the first AUG from mRNA determine the sequence of amino acids in the polypeptide the 5 terminus in order to begin pro- chain. tein synthesis with a methionine. Almost all plant virus species, and a majority of animal virus species, use ribosomal skipping A variation on the RNA rather than DNA as their gen- normal translation process demon- etic material. RNA may also play a strated to occur during the replica- regulatory role in expression at the tion of foot-and-mouth disease virus. post-transcriptional level. Translation continues along the viral Altuvia S and Wagner GH (2000) Proc Natl RNA but fails to create a peptide bond Acad Sci 97 , 9824 at a specific site. This has the same effect Eaton BE and Pieken WA (1995) Annu Rev as proteolytic processing of the poly- Biochem 64 , 837 protein translation product of the virus. ribosomal skipping 412

Skipping occurs after the structural pro- polymerases, although 3-oxime deriv- tein coding region, at a 18 amino acid atives with some inhibitory action sequence known as protein 2A. against mammalian RNA polymerases Donnelly ML et al (2001) J Gen Virol 82 , 1013 have been described (rifamycins AF/05 and AF/013). Rifamycin will prevent focus formation by Rous sarcoma virus ribosome binding site A sequence of and inhibits maturation of Poxviridae . 4–7 nt in mRNA to which ribosomes bind. In prokaryotes this is termed a Pennington TH et al (1970) J Gen Virol 9 , 225 ‘ Shine–Dalgarno sequence ’ and is a Vaheri A and Hanafusa H (1971) Cancer Res sequence complementary to the 3 end of 31 , 2032 the 16S rRNA. In eukaryotes there is considerable sequence variation at the 5 end Rift Valley fever virus (RVFV) A species of the mRNA, and a consensus ribosomal in the genus Phlebovirus. Member of the binding sequence is not found. Rift Valley fever complex in the sandfly Shine J and Dalgarno L (1974) Proc Natl Acad fever serogroup. Found in central and Sci 71 , 4734 southern Africa. Causes abortion and many deaths in pregnant and newborn sheep, goats, and cattle. Lambs develop ribosomes Small, round, electron-dense fever, vomiting, mucopurulent nasal particles, 10–20 nm in diameter, found discharge, and bloody diarrhea. Cattle on the outer surface of the limiting are less seriously affected. Herdsmen membrane of the rough-surfaced endo- and slaughtermen often become infected plasmic reticulum. Also found free in and develop a biphasic illness which is the cytoplasm, sometimes in the nucleus usually mild, although retinal damage and mitochondria, and present in all may occur. Buffalo, camels, and ante- types of living cells. Name introduced in lopes may be naturally infected and die. 1957 to distinguish the particulate ribo- Infection is mosquito-borne, but contact somes from membrane-associated ribo- infection probably also occurs. Large somes known as ‘ microsomes, ’ obtained mosquito-borne epizootics occurred in on cell fractionation. They contain 40% 1977–1978 in Egypt, in 1987–1988 in East protein and 60% RNA, and play a vital Africa. Absent for a decade, it returned part in protein synthesis, attaching to to Egypt in 1993. Then in 1999 a large mRNA to form polysomes. In mamma- outbreak began in the Kingdom of lian cells the ribosome consists of two Saudi Arabia and Yemen, the first out- subunits that sediment at 40S and 60S, break ever recorded outside the African respectively. continent, with more than a thousand human cases and more than 150 deaths ribovirus RNA-containing virus. by the end of 2000. Control is by protection from mosquitoes and by vac- ribozyme An RNA molecule with cata- cination of livestock with formalinized lytic activity. or attenuated vaccines. Mice die of hepatitis when infected experimentally. Rida virus An Icelandic strain of the Guinea pigs, ferrets, and young dogs prion disease scrapie, probably can also be infected, but birds are resist- imported from Scotland, UK. ant. Virus replicates in cultures of chick, Peturson G (1994) Ann NY Acad Sci 724 , 43 rat, mouse, and human cells, and on the CAM, causing thickening. The virus RIF See resistance-inducing factor . hemagglutinates day-old chick cells at pH 6.5 and 25°C. rifampicin See rifamycin . LeBeaud AD et al (2007) Am J Trop Med Hyg 76 , 795 rifamycin An antibiotic which inhibits RNA synthesis in sensitive strains of RIG-1 Retinoic acid-inducible gene 1. Escherichia coli by binding to a sub- A cytosolic viral RNA receptor that unit of the bacterial RNA polymerase. interacts with a mitochondrial antiviral Does not inhibit eukaryotic cell RNA signaling adaptor (MASVS) to induce 413 Rio Bravo virus (RBV)

type 1 interferon mediated host pro- Nations (FAO) is underway with the tective innate immunity against virus aim of global eradication by 2010. As infection. Some viruses such as hepa- of 2008, it seems as though the Global titis C virus, evade this mitochondrial Rinderpest Eradication Programme immune response by cleaving MASVS has succeeded, and the aim is now for- from the mitochondrial membrane. mally to declare that the world is free Paz S et al (2006) Cell Mol Biol 52 , 17–28 of rinderpest in the year 2010. Synonyms: cattle plague virus; peste bovina; peste bovine. RIID 3229 virus A strain of Oliveros virus in the genus Arenavirus . Roeder PL and Taylor WP (2002) Vet Clin North Am Food Anim Pract 18 , 515 Riley virus Synonym for Lactate dehydrogenase-elevating virus . RING finger motif See RING finger proteins . rimantadine hydrochloride (␣-methyl- 1adamantane-methylamine hydro- RING finger proteins Proteins having chloride) A derivative of amantadine a sequence related to the zinc finger hydrochloride licensed as an antiviral but containing additional cysteine and agent. Good results have been reported histidine residues that help the mol- from its use in the treatment of patients ecule to form a ring. Play a critical role with influenza virus A infections. Toxic in mediating the transfer of ubiquitin effects include anxiety, nightmares, both to heterologous substrates and to and vomiting, but are less than with the RING finger proteins themselves. amantadine. Ubiquitination is a first step in protein degradation, and some viruses, notably Galegov GA et al (1979) Lancet i , 269 human papillomavirus, express ring finger proteins that cause ubiquitina- Rinderpest virus (RPV) A species in the tion leading to degradation of cellular genus Morbillivirus. A serious natural proteins, such as p53. infection of wild and domestic ani- Freemont PS (2000) Curr Biol 10 , R84 mals in many parts of Asia and Africa. Joazeiro CAP and Weissman AM (2000) Cell Ox, zebu, buffalo, yak, sheep, goats, 102 , 549 pigs, camels, hippopotamus, warthog, giraffe, and several other wild animals ring-necked pheasant adenovirus See are naturally infected. In the acute dis- marble spleen disease virus . ease there is high fever and constipation followed by diarrhea. Mortality may ring-necked pheasant leukosis virus A exceed 90%. Inflammation and ulcera- strain of avian leukosis sarcoma virus. tion of the whole alimentary tract is the An endogenous virus present in nor- main pathological lesion, but patchy mal ring-necked pheasant, Phasianus pneumonia may occur. Transmission is colchicus , cells, with genetic information by direct contact and outbreaks usually to give group F host-range specificity start by the introduction of an infected to virus particles produced with it as animal with up to 100% infection of helper virus. the affected herd. Control is by slaugh- Hanafusa T and Hanafusa H (1973) Virology ter and use of tissue-culture attenu- 51 , 247 ated vaccines, which generate life-long immunity. The virus is closely similar in ring vaccination Vaccination of contacts structure to measles and canine distem- and people who may come into contact per virus and contains cross-reacting with an infected person, so as to form a antigens. Serum from rinderpest virus- ring of non-susceptibles. A method used infected cattle prevents hemaggluti- to control diseases such as smallpox. nation by measles virus. Several live virus-attenuated vaccines are avail- Rio Bravo virus (RBV) A species in the able and a global rinderpest eradication genus Flavivirus , member of the Rio campaign coordinated by the Food and Bravo virus serogroup. Isolated from Agriculture Organization of the United the salivary gland of a bat caught in Rio Bravo virus (RBV) 414

California, USA. Similar viruses have Synonyms: harvest mouse virus 2; been isolated in Texas, USA and Mexico. HMV-2. No known arthropod vector and did Hjelle B et al (1995) Virology 207 , 452 not replicate in any of several mosquito species. In mice shows tropism for kid- risk The probability of a disease-free per- ney, mammary, and salivary gland tis- son developing the disease of interest sue. Has caused laboratory infections over a defined time, and not dying from associated with orchitis. No evidence any other disease during that period. that it causes disease in bats. Synonym : bat salivary virus. ritonavir A protease inhibitor active in therapy of HIV infection. Used in com- Rio Grande cichlid virus (RGRCV) bination with lopinavir, in the form of An unassigned virus in the family single capsules called Kaletra. Ritinovir Rhabdoviridae . increases the amount of lopinavir in the blood, making it more effective against Rio Grande virus (RGV) A tentative spe- HIV. cies in the genus Phlebovirus , belonging Synonym: Norvir. to the sandfly fever virus serogroup. Isolated from pack rats, Neotoma micro- RK13 cells (CCL 37) A cell line initiated pus , in Texas, USA. On injection into from trypsinized kidney cells of a suckling mice causes death in 5–6 5-week-old rabbit. days. Replicates in Vero cells with CPE. Serological surveys suggest the pack RKV (rabbit) polyomavirus See Rabbit rat is the principal natural host. Has kidney vacuolating virus . not been isolated from hematophagous insects, so the vector is uncertain. Levels R-loop mapping A technique for map- of viremia are low in experimentally ping single-stranded RNA by hybridi- infected pack rats, so mode of transmis- zation to the complementary strand of sion is uncertain. Not reported to cause partially denatured double-stranded disease in humans. DNA. The formation of the RNA–DNA hybrid displaces the opposite DNA Calisher CH et al (1977) Am J Trop Med Hyg strand as a loop which can be visual- 26 , 997 ized under the electron microscope. Rio Mamore virus (RIOMV) A species RM-97 virus A strain of El Moro Canyon in the genus Hantavirus identified in virus in the genus Hantavirus . Bolivia in rodents, Oligoryzomys microtis, by nucleotide sequence analysis of RML 105355 virus (RMLV) A strain kidney-derived RNA. Closest North of Uukuniemi virus in the genus American relative appears to be Bayou Phlebovirus . Transmitted by ticks. virus. Hjelle B et al (1996) Lancet 347 , 57 RNA See ribonucleic acid .

Rio Mearim virus (RIME) A probable RNA-dependent DNA polymerase An species in the genus Hantavirus iden- enzyme present in the virion of all tified in Maranhao State, Brazil, in retroviruses and in cells during repli- Wagner’s marsh rats (Holochilus sciu- cation of hepadnaviruses. It copies sin- reus). The nucleotide sequence of the N gle-stranded RNA into DNA, and has gene shows a close relationship to that ribonuclease H activity which digests of Rio Mamore virus. RNA present in RNA–DNA hybrid Rosa ES et al (2005) Vector Borne Zoonotic Dis molecules. Like DNA polymerase, the 5 , 11 enzyme requires a primer for DNA synthesis. The natural primer in retrovi- Rio Segundo virus (RIOSV) A spe- ruses is a species of tRNA bound some cies in the genus Hantavirus , isolated 100 or so nucleotides from the 5 end from a Costa Rican harvest mouse, of the genome RNA; the primer var- Reithrodontomys mexicanus . Not known ies according to the virus species. It is to cause disease in humans. tRNATrp for avian sarcoma virus and 415 RNA polymerase

probably for all avian leukosis viruses; from the cells, particularly if phenol is tRNAPro for Moloney leukemia virus, used. In those viruses where the RNA all other murine leukemia viruses which is a negative strand, the viral ribonu- have been examined, simian sarcoma cleoprotein has RNA-dependent RNA virus and avian reticuloendotheliosis polymerase activity which means that virus; tRNALys3 for mouse mammary mRNAs are made as well as comple- tumor virus; and tRNALys 1,2 for HIV. mentary RNA to act as template for Hepadnavirus replication involves a new virion negative-strand RNA syn- reverse transcription step primed by a thesis. RNA-dependent RNA polymer- polypeptide. During establishment of ases which catalyze the transcription of retrovirus infection, the end-product mRNA from the virion negative strand of RNA-dependent DNA polymer- are also termed ‘ RNA transcriptases ’ ; ase activity is a linear double-stranded those which catalyze the formation of DNA molecule containing terminal new virion RNA of whatever polarity repeats which subsequently becomes are called ‘ RNA replicases. ’ RNA syn- integrated into the cell genome. The thesis does not require a primer, in con- enzyme has been purified free of the trast to DNA synthesis. natural template and in this form is Synonyms : RNA replicase; RNA syn- widely used for genetic manipulation thetase; RNA transcriptase. and nucleic acid sequencing. The crystal structure shows that the enzyme is a RNA–DNA viruses A name sometimes heterodimer of full-length reverse tran- used for the family Retroviridae . scriptase (p66) and a cleavage product RNA ligase An enzyme isolated from (p51) that lacks the C-terminal ribonu- bacteriophage T4-infected Escherichia clease H domain. coli which adds single residues (pNp) Synonym : reverse transcriptase. to the 3 termini of RNA chains. Used Katz RA and Skalka AM (1994) Annu Rev experimentally to label RNA prior to Biochem 63 , 133 sequencing by the gel method. Koehlstaedt LA et al (1992) Science 256 , 1783 RNA polymerase An enzyme which RNA-dependent RNA polymerase An catalyzes either the formation of RNA enzyme unique to viruses, encoded by from a DNA template, in which case the viral genome, which is specific for it is DNA-dependent RNA polymer- its own viral RNA and does not func- ase, or the formation of RNA from tion for other virus species. It brings an RNA template, in which case it is about transcription of viral RNA to RNA-dependent RNA polymerase. In complementary RNA and this is in turn eukaryotic cells three main types of transcribed back to form more viral DNA-dependent RNA polymerase are RNA. In viruses where the RNA is a identified, dependent on the type of positive strand and can act as mRNA, RNA they synthesize: translation commences immediately on infection and RNA-dependent RNA RNA polymerase I synthesises 45s pre- polymerase molecules are produced. RNA, which matures into the ribos- These attach to the viral RNA and pass omal RNAs, 28s, 18s, and 5.8s. along from the 3 end producing a com- RNA polymerase II synthesises precur- plementary strand which is thus syn- sors of mRNAs, and most snRNAs and thesized from 5 to 3 . This complex of microRNAs. It also binds the 5 -cap one positive strand with about six or synthesising and cap-binding com- seven complementary strands growing plex at its C-terminal domain, which on it is called the ‘ replicative interme- is also the site for spliceosome factors diate.’ The presence of positive strands involved in the removal of introns dur- and complementary strands results ing DNA transcription. in the formation of some double- RNA polymerase III synthesises tRNAs, helical strands, which are known as the 5s ribosomal RNA and other small ‘ replicative form. ’ This is an irrelevant RNAs found in the cell. end-product and much of it may be Hurwitz J (2005) J Biol Chem 280 , 42477 formed during the process of extraction Willis IM (1993) Eur J Biochem 212 , 1 RNA processing 416

RNA processing See post-transcriptional Paulo, Brazil. Caused an epidemic of modification . encephalitis in several coastal counties of São Paulo in 1975: there were 462 RNA replicase Synonym for RNA- cases, 61 of which were fatal. No dependent RNA polymerase. virus was isolated from 420 sera from patients with encephalitis but 9 isola- RNA segment A distinct piece of genomic tions were made from brain tissue. RNA; the genome segments of seg- Isolated from 2 of 395 sentinel mice mented genome viruses, e.g. the double- exposed in the epidemic area, and from stranded RNAs of reovirus or the a rufous collared sparrow, Zonotrichia single-stranded RNAs of orthomyxo- capensis , collected in the same area. virus, arenavirus, or bunyavirus. Epidemiology suggests mosquito vectors, Psorophora ferox and Aedes scapu- RNA splicing See splicing . laris , and possibly a bird natural host. Hemagglutinin present in mouse brain RNA synthetase Synonym for RNA- tissue. Serologically related to other dependent RNA polymerase. flaviviruses but distinguishable from them. Pathogenic on injection into RNA transcriptase Synonym for RNA- or mice and hamsters. Replicates in Vero DNA-dependent RNA polymerase. cells and BHK21 cells with CPE. RNA tumor viruses See retroviruses . Lopes OS et al (1978) Am J Epidemiol 107 , 444; 108 , 394 RNA viruses Viruses having RNA Monath TP et al (1978) Am J Trop Med Hyg genomes. 27 , 1251 rodent paramyxoviruses Rodent species RNase Abbreviation for ribonuclease. in the genus Paramyxovirus include: RNase H1 An endonuclease which J virus cleaves the RNA primer during repli- Mossman virus cation of SV40 virus. Murine pneumonia virus Nariva virus roan antelope herpesvirus Synonym for Parainfluenza virus type 1 murine Hippotragine herpesvirus 1 . Peromyscus virus Pneumonia virus of mice Rochambeau virus (RBUV) A tentative Sendai virus species in the genus Lyssavirus . Isolated Tupaia virus from mosquitoes, Coquillettidia albicosta , in French Guiana in 1973. Not reported Jun MH et al (1977) Aust J Exp Biol Med Sci 55 , 645 to cause disease in humans. rodent parvoviruses Can be divided by Rock bream iridovirus A probable species their serological properties into five in the genus Megalocytivirus . Causes species with their respective strains as epizootics of disease affecting the spleen follows: and kidney among cultured rock bream (1) Kilham rat virus (Oplegnathus fasciatus ) in Korea. The H-3 virus complete genome sequence showed Kilham rat virus that it consists of 112,080 bp of dsDNA, (2) H-1 virus similar in sequence and organization to H-1 parvovirus red sea bream iridovirus and infectious (3) HB parvovirus spleen and kidney necrosis virus. HB parvovirus Do JW et al (2004) Virology 325 , 351 (4) Minute virus of mice Minute virus of mice (Cutter) Rocio virus (ROCV) A strain of Ilhéus Minute virus of mice (immuno- virus in the genus Flavivirus , Ntaya suppressive) virus group. Isolated by i.c. injection Minute virus of mice (prototype) into newborn mice from the brain of a (5) Mouse parvovirus 1 patient who died of encephalitis in São Mouse parvovirus 1 417 Ross River virus (RRV)

H-1, and HB may be human viruses as Roniviridae A family in the order Nido- they were apparently recovered from virales containing a single genus, human tissues, but antibodies to them Okavirus, consisting of viruses which are rare in humans and they are patho- infect crustaceans. The type species is genic to newborn hamsters, a feature Gill-associated virus . not yet observed with parvoviruses outside the rodent group. roseola infantum Synonym for exanthem subitum . A childhood disease appear- rodent (wild in Turkmenia) poxvirus ing rarely before 6 months of age and A possible species in the genus usually before age 4 years sporadically Orthopoxvirus, isolated in 1974 from the or in limited size outbreaks. There is kidneys of a wild big gerbil, Rhombomys an exanthema preceded by a fever that opimus , caught in Turkmenia. It resem- subsides with the appearance of a mac- bles cowpox virus and carnivora pox- ular-papular rash on the trunk and to viruses but was markedly different a lesser extent on the face and extremi- from Ectromelia virus . Apparently iden- ties. Rarely persists more than 24 h. tical to viruses isolated from Felidae in Caused by human herpesvirus 6 in the Moscow Zoo. Experimental infection of genus Roseolovirus . the natural hosts, big gerbil, and yellow suslik, Citellus fulvus, caused severe dis- Roseolovirus A genus in the subfamily ease and high mortality. Transmission Betaherpesvirinae. Viruses are isolated between cage mates occurred. Virus from lymphocytes and have a unique is present in urine for at least 3 weeks DNA structure. There are two spe- and in kidneys for at least 5 weeks. cies in the genus, Human herpesvirus 6 which causes roseola infantum (also Marennikova SS et al (1978) Arch Virol 56 , 7 called exanthem subitum or sixth disease), and Human herpesvirus 7 which rolling circle A model for DNA replica- has not so far been clearly associated tion which involves a circular inter- with a human disease. There are two mediate molecule. One strand remains variants of HHV6, HHV-6A, and HHV- as a circular template and may be cop- 6B, and only HHV6-B has been clearly ied repeatedly but the other parental associated with roseola infantum. strand is only copied once, so replication is asymmetric. Ross’ goose hepatitis B virus (RGHBV) Gilbert W and Dressler D (1968) Cold Spring An unassigned virus in the family Harbor Symp Quant Biol 33 , 473 Hepadnaviridae . Isolated from Ross ’ goose (Anser rossi ). Genome sequence rolling hairpin replication (RHR) A resembles that of several other modification of the rolling circle model avihepadnaviruses. of DNA replication applied to parvo- Guo H et al (2005) J Virol 79 , 2729 virus DNA replication. The genome is replicated through a series of mono- Ross River virus (RRV) A species in the meric and concatemeric duplex rep- genus Alphavirus. Isolated from birds licative form intermediates. The 3 and mosquitoes in Fiji, and Queensland terminus of the virion DNA strand is and New South Wales, Australia. thought to fold back on itself to pro- Causes a febrile illness and rash with vide a primer for initiating synthesis of arthralgia in humans. The vectors are the complementary strand resulting in the mosquitoes, Aedes vigilax and Culex a linear duplex intermediate. annulirostris . Antibodies are present in Berns KI (1990) Microbiol Rev 54 , 316 horses, cattle, sheep, dogs, rats, bats, and kangaroos in northern and eastern Rondônia virus A probable strain in the Australia, New Guinea, and northern genus Phlebovirus, isolated in Samuel, Solomon Islands. Rondônia State, Brazil, from phlebot- Aaskov JG and Davies CEA (1979) J Immunol omine sand flies in 1988. Not associ- Methods 25 , 37 ated with human disease. Mackenzie JS et al (1994) Arch Virol 136 , 447 Rost Island virus (RSTV) 418

Rost Island virus (RSTV) A serotype of intestinal enterocytes. Virus particles are Great Island virus in the genus Orbivirus , found in the columnar epithelial cells, belonging to the Great Island complex. goblet cells, phagocytic cells, and M cells in the small intestine. In children, Rotavirus A genus of the family Reoviridae . rotavirus gastroenteritis commonly Virion diameter 100 nm. The icosa- occurs between the ages of 3 months hedral capsid is triple-layered with a and 2 years, and in other age groups clearly defined outer layer, appear- infections may be asymptomatic. In ing like the rim of a wheel, while the developing countries, it is estimated inner layers gives the appearance of that rotavirus infections cause about spokes, hence the name. 132 channels 870,000 deaths annually, especially in extend inward from the surface to the malnourished children. Oral rehydra- core, and 60 short spikes extend out- tion therapy is very effective where it ward from the surface. The outer layer has been used, but the development of is often removed spontaneously so that an attenuated rotavirus vaccine is prob- both triple-layered smooth and double- ably the best long-term strategy for con- layered rough virions are found in the trolling rotavirus disease. One vaccine, gut contents. Virus replicates in intes- known as Rotashield™, was licensed tinal epithelial cells. Genome RNA for a short period in the USA but with- is double-stranded with 11 segments drawn in 2000 because of a higher than from 667 bp to 3.3 kb long. Buoyant normal incidence of intussusception in density of 1.36 g/ml in CsCl. On poly- recipients. Because of the high burden of acrylamide gel fractionation, eight struc- rotavirus disease, vaccine development tural polypeptides and six nonstructural has continued, and two new vaccines, polypeptides can be identified. There Rotarix, an attenuated human vaccine are strains causing acute gastroenteritis and Rotateq, a human-bovine recom- in humans, especially infants, and in binant vaccine, have been licensed for numerous other hosts such as calves, live oral administration. mice, piglets, lambs, foals, rabbits, ante- Synonyms: duovirus; reovirus-like agent; lope, hares, chickens, turkeys, and chim- stellavirus. panzees. Very large numbers of particles are present in the feces and are detected Desselberger U (1996) Adv Virus Res 46 , 69 Desselberger U et al (2005) In Topley & by electron microscopy. Many species Wilson’s Microbiology and Microbial Infections , are difficult to replicate in cell cultures, vol. 1, Tenth edition, edited by BWJ Mahy but most strains can be grown in sim- and V ter Meulen, London: Hodder Arnold, ian kidney cell lines if trypsin is added p. 946 to the culture medium. Rotaviruses can Ramig RF (1994) Curr Top Microbiol Immunol be divided into seven groups (A–G) on 185 , 380pp the basis of the VP6 inner capsid protein, the group-specific antigen. Within Rotavirus A (ROTAV A) A species in the group A, a subdivision into subgroups genus Rotavirus. The most common I and II is possible using anti-VP6 mon- cause of diarrheal disease in infants and oclonal antibodies. There are two type- young children, constituting more than specific antigens: VP4 (HA) and VP7. At 95% of currently identified strains in least 14 serotypes of group A (G1–G14) humans worldwide. All group A rota- based on VP7, and 11 serotypes of group viruses share a common antigen, VP6, A (P1–P11) based on VP4 are recog- the major inner capsid protein. There nized. These can be discriminated either are two subgroups, I and II, compris- by reactions with monoclonal antibod- ing a number of serotypes. In addition, ies or polyclonal antibodies to baculo- the viruses are classified as G types virus expressed proteins or to rotavirus (G from g lycoprotein) based on VP7, reassortants, or using sequence-specific and P types (P from p rotease-sensitive nucleic acid probes or genotype-specific protein) based on VP4. At least 15 dif- reverse transcription-polymerase chain ferent G types and more than 20 differ- reaction primers. Rotaviruses cause ent P types have so far been described. diarrhea in the species that they infect These designations are partly based which may be due in part to lysis of on sequence analysis (genotypes) 419 Rous sarcoma virus (Schmidt–Ruppin B) (RSV-SR-B)

where complete serology is missing, identified in birds. The prototype is the so the G or P types may fall into dif- chicken A4 strain. ferent subgroups. For example, the G (VP7) serotypes 1, 3, or 4 are in sub- rotavirus G (ROTAV G) A tentative spe- group II, whilst subgroup I rotaviruses cies in the genus Rotavirus, only so far belong to G serotype 2. Group A rota- identified in birds. The prototype is the viruses are ubiquitous worldwide, and chicken 555 strain. include the prototype rotavirus, simian rotavirus SA11. rougeole virus Synonym for Measles virus . Bellamy AR and Both GW (1990) Adv Virus rough membrane Endoplasmic reticu- Res 38 , 1 lum encrusted with ribosomes. See also Burke B and Desselberger U (1996) Virology smooth membrane . 218 , 299 Ramig RF (Editor) (1994) Curr Top Microbiol Rous-associated virus (RAV) An avian Immunol 185 , 380pp leukemia virus which acts as a helper for replication defective avian sarcoma Rotavirus B (ROTAV B) A species in virus, providing information for the the genus Rotavirus. A major cause of coat and thus controlling surface anti- severe adult diarrheal disease in China, gens and host range. See Avian leuko- where it was identified in 1983. In the sis virus , genus Alpharetrovirus . USA, only 5% of the population is seropositive for group B rotaviruses. Causes Rous sarcoma virus (RSV) A species infectious diarrhea of infant rats, and in the genus Alpharetrovirus . The first has also been found in pigs, calves, and virus demonstrated, in 1911, to cause sheep. a solid malignant tumor; Peyton Rous Synonym: adult diarrheal rotavirus waited 55 years before receiving the (ADRV). Nobel Prize for his work in 1966, shortly before his death at the age of Rotavirus C (ROTAVC) A species in the 91 in 1970, the year that reverse tran- genus Rotavirus. Originally termed scriptase was discovered. There are ‘ pararotaviruses ’ when found in 1982– a number of strains of RSV varying 1983 in infants in Australia, Brazil, in their oncogenicity and host range. and France. Other than humans, pigs, Some will produce tumors in mammals and cattle have been found to be such as rats, cotton rats, guinea pigs, infected. The prototype is the porcine mice, hamsters, and monkeys. Some Cowden strain. Epidemic outbreaks are replication-competent, but most are have occurred in England and Japan. defective and require a leukemia virus Agglutinate human and sheep eryth- to code for the viral envelope which rocytes. Closely related to group A determines host range. Transforms cells rotaviruses. in culture which do not produce infective virus unless also infected with a Rotavirus D (ROTAV D) A species in the leukemia virus. Transformed cells can- genus Rotavirus, only identified so far not be maintained indefinitely in cul- in birds. The prototype is the chicken ture. The genome of the virus contains 132 strain. Most natural infections the v-src oncogene sequence which is involve young birds, less than 6 weeks not present in leukemia viruses and of age. Turkeys, chickens, pheasants, is responsible for cell transformation and ducks are all susceptible. leading to solid tumor formation. Rous P (1911) J Exp Med 13 , 397 Rotavirus E (ROTAV E) A species in the genus Rotavirus, found in pigs. Rous sarcoma virus (Prague C) (RSV- Unrelated antigenically to groups A, B, Pr-C) A replication competent strain of C, D, and F. The prototype is the por- Rous sarcoma virus . cine DC-9 strain. Rous sarcoma virus (Schmidt–Ruppin B) rotavirus F (ROTAV F) A tentative spe- (RSV-SR-B) A replication competent cies in the genus Rotavirus, only so far strain of Rous sarcoma virus . Rous sarcoma virus (Schmidt–Ruppin D) (RSV-SR-D) 420

Rous sarcoma virus (Schmidt–Ruppin D) with extensive malignant tumor of the (RSV-SR-D) A replication competent nasal septum. strain of Rous sarcoma virus . RPMI 7666 cells (CCL 114) One of a large Rousettus lyssavirus (RLV) A strain of series of apparently permanent cell European bat lyssavirus 1 in the genus lines derived from leukocytes in the Lyssavirus, isolated in Denmark from a peripheral blood of donors with and colony of apparently healthy Egyptian without malignancies. flying foxes, Rousettus aegypticus , originating from Rotterdam zoo where RPMI 8226 cells (CCL 155) One of a they had inhabited an artificial cave series of human hematopoietic cell for more than 6 years. Caused clini- lines obtained from the peripheral cal rabies when inoculated intracrani- blood of a 61-year-old male with mul- ally into bats of the same species and tiple myeloma. caused neurological signs in mice. The virus is closely similar by genetic anal- R protein A low abundant protein ysis to EBLV-1, isolated from free-living expressed from the P mRNA of measles Eptesicus serotinus bats. virus by frameshifting. Van der Poel WHM et al (2000) Arch Virol Liston P and Briedis DJ (1995) J Virol 69 , 145 , 1919 6742

Rowson–Parr virus A strain of RPV See raccoon parvovirus . Murine leukemia virus in the genus Gammaretrovirus, isolated from a Friend RR 1022 cells (CCL 47) A heteroploid cell murine leukemia virus preparation by line derived from a tumor induced in endpoint dilution. Causes a very minor inbred Amsterdam rats, Rattus norvegi- degree of splenomegaly but is a potent cus , by i.m. injection of the Schmidt– depressor of the immune response. Ruppin strain of Rous sarcoma virus. After a long latent period of 6–8 months, neoplastic lymphoid cells appear in the RTG-2 cells (CCL 55) A heteroploid cell germinal centers of the spleen and later line derived from pooled trypsinized in other lymphoid tissues. male and female gonadal tissue of Rowson KEK and Parr IB (1970) Int J Cancer yearling rainbow trout, Salmo gairdneri . 5 , 96 The first cell line to be established from poikilothermic animals. Cells do not Royal Farm virus (RFV) A species in survive above 26°C. the genus Flavivirus , belonging to the mammalian tick-borne virus group. RT parvovirus (RTPV) A species in the Isolated from a tick, Argas hermanni , in genus Parvovirus . Isolated from a line Afghanistan. Not reported to cause dis- of rat fibroblasts, RT. Natural host ease in humans. unknown. Antigenically different from KBSH, tumor virus X, and LU III RPMI 1788 cells (CCL 156) One of a viruses. series of hematopoietic cell lines origi- Hallauer C et al (1971) Arch Virol 35 , 80 nating from the peripheral blood leukocytes of an apparently normal RT parvovirus An autonomous parvo- 33-year-old Caucasian male. virus isolated from suckling mice that had been inoculated with human syno- RPMI 1846 cells (CCL 49) A melanotic vial tissue from cases of rheumatoid cell line derived from a malignant arthritis. A rodent strain of parvovirus melanoma which arose spontaneously unrelated to the human disease. The in an aged golden hamster and was reported association with arthritis was carried for 41 animal passages by sub- erroneous. cutaneous implantation. Simpson RW et al (1984) Science 223 , 1425 RPMI 2650 cells (CCL 30) Established from the pleural effusion of a patient RT-PCR See polymerase chain reaction . 421 Rubulavirus

R-type virus particles ( ‘ R ’ because of often results in infection of the fetus the structures radiating from the core). and congenital abnormalities such as Enveloped virus particles 100 nm in cataract, hearing loss, cardiac and den- diameter with a clear space between tal malformations, and microcephaly. the central core and outer envelope. Congenitally infected children have Across this space are radial threads. rubella virus antibodies but continue to First described in BHK21 cells, and excrete virus for many months. A very later in calf kidney cells and hamster few cases of progressive rubella panen- tumors induced by polyoma, SV40, cephalitis have occurred, commencing Rous sarcoma and mouse sarcoma in the second decade of life and slowly viruses. They are usually seen singly progressing. Monkeys, rabbits, ham- or in small groups within cell vacuoles. sters, guinea pigs, rats, and mice can They have reverse transcriptase and be infected experimentally, but show can rescue mouse sarcoma virus from no disease except leukopenia and rash non-producer cells. No known patho- in some rhesus monkeys, and growth logical role. retardation in congenitally infected ani- Albu E and Holmes KV (1973) J Virol 12, 1164 mals. Major epidemics occur every 9–10 Bergman DC et al (1977) J Natl Cancer Inst years and 80–90% of young adults have 58 , 295 antibodies. Several excellent live attenuated vaccines have been developed; Rubarth’s disease virus Synonym for the RA27/3 strain is used most widely. Canine adenovirus . Rubella vaccine is normally given as a component of measles–mumps–rubella Rubella virus (RUBV) The type species (MMR) vaccine to all preschool chil- of the genus Rubivirus in the family dren at about 12–15 months of age. A Togaviridae . The virus is 50–70 nm in booster should be given to all girls at diameter, has a triple-layered enve- the age of 12–l4 years, if they were not lope 8 nm thick surrounding an elec- immunized at 12–15 months. tron-lucent layer 11 nm thick and an Synonym : German measles virus. electron-dense core 30 nm in diameter. Best JM et al (2005) In Topley & Wilson’s Matures by budding from cytoplas- Microbiology and Microbial Infections , vol. mic membranes. The nucleic acid is 1, Tenth edition, edited by BWJ Mahy and positive single-stranded RNA 9757 nt V ter Meulen, London: Hodder Arnold, p. 959 in length, with an additional 3 poly Pugachev KV et al (1997) Arch Virol 142 , 1165 A tract. There are three structural pro- Reef SE (1999) MMWR 48 , Suppl, 199 teins: two glycoproteins (E1 and E2) and a capsid protein, C. The gene order rubeola virus Synonym for Measles virus . in the precursor polyprotein is NH2- CP-E2-E1-COOH. Infectivity sensitive Rubini strain A strain of mumps virus to lipid solvents and pH 3. Day-old attenuated by passage in human dip- chick erythrocytes are agglutinated at loid cells and licensed in Switzerland 4°C. No antigenic differences have been for use as a vaccine. It has since observed between strains. Replication appeared to induce only a low level of occurs in duck eggs and a wide vari- protective immunity. ety of cell cultures, in which it may be demonstrated by interference with Rubivirus A genus of the family growth of another virus, by CPE or by Togaviridae. Antigenically unrelated to immunofluorescence. For isolation, pri- viruses in other genera of the family. mary African green monkey kidney cell Rubella virus is the only species in this cultures or BHK21 cells are best. Causes genus, and the structural proteins have a mild illness in humans with general- no amino acid similarity to those of ized rash and enlarged lymph nodes. alphaviruses. Some homology can be Usually little fever or constitutional dis- observed within the nonstructural pro- turbance. Incubation period 16–18 days. teins, however. Meningo-encephalitis and other complications are rare but infection dur- Rubulavirus A genus in the subfamily ing the first 3–4 months of pregnancy Paramyxovirinae. Mumps virus is the Rubulavirus 422

type species. Other species include neutralize virus infectivity completely. human parainfluenza viruses 2 and 4, Replicates with CPE in BHK21 cells. Mapuera virus, porcine rubulavirus, Hemagglutinates chicken erythrocytes. simian virus 5, and simian virus 41. Antibodies are present in sea birds. Not All species have hemagglutination and reported to cause disease in humans. neuraminidase activities. Tioman virus and Menangle virus are tentative spe- Traavik T (1979) Acta Pathol Microbiol Scand B 87 , 1 cies in the genus. Traavik T and Brunvold E (1978) Acta Pathol Microbiol Scand B 86 , 349 rule of six A phenomenon discovered by Calain and Roux whilst investigat- Russian autumn encephalitis virus ing the replication of deletion mutants Synonym for Japanese encephalitis virus . of Sendai virus. Efficient genome replication only occurred when the total Russian spring–summer encephalitis length of the genome was a number virus (RSSEV) Tick-borne of bases divisible by six. It is believed A strain of encephalitis virus Flavivirus to result from the notion that the NP in the genus . protein of paramyxoviruses covers The principal tick vector for RSSEV is Ixodes persulcatus precisely six nucleotides when bind- , and several small ing to the RNA. The phenomenon has rodents and insectivores appear to stood the test of time as new para- be amplifying hosts. The disease is myxoviruses have been discovered acquired in humans by tick bites, and and sequenced. For example, the two onset is gradual with fever, headache, most recently sequenced paramyxo- nausea, vomiting, and photophobia. viruses follow this rule: the Hendra The case fatality rate is about 20% in virus genome has 18,234 nt and the rural areas where the disease is Nipah virus has 18,246 nt. prevalent, with children most affected. Survivors may suffer from neurologi- Calain P and Roux L (1993) J Virol 67 , 4822 cal sequelae such as mild paralysis. Peeters BPH (2000) Arch Virol 145 , 1829 A formalin-inactivated vaccine made from virus grown in chick embryo cells Runde virus A possible species in the is available in Russia. genus Coronavirus . An arbovirus iso- Synonym: tick-borne encephalitis virus lated from the tick, Ixodes uriae , col- (Far Eastern subtype). lected in sea bird colonies at Runde, Norway. No antigenic relationship to Kuno G et al (1998) J Virol 72 , 73. avian infectious bronchitis virus or to Roehrig JT and Gubler DJ (2005) in Topley & major arbovirus groups. Lethal to new- Wilson’s Microbiology and Microbial Infections , born mice but in 2-week-old mice pro- vol. 2, Tenth edition, edited by BWJ Mahy and V ter Meulen, London: Hodder Arnold, duces a persistent infection and chronic p. 993 disease may result. Antibodies fail to S

S 1954-847-32 virus (TURV) A strain Sabethes A genus of forest-dwelling mos- of Turlock virus in the genus quitoes found in Central and South Orthobunyavirus . America. Important in the transmission of yellow fever virus, especially SA virus A designation given to a virus Sabethes chloropterus . isolated in hamster brain from nasal washings of a patient with a ‘ common Sabiá virus (SABV) A species in the cold. ’ A strain of parainfluenza virus genus Arenavirus . Isolated in São Paulo, type 5. Pathogenic for hamsters on i.c. Brazil in 1990 from a fatal case of viral injection. Antigenically identical to hemorrhagic fever. A laboratory tech- Simian virus SV5 and DA virus. nician handling the agent became infected with the virus and a third case SA virus series See simian viruses (SA occurred in a research worker at Yale series). University, USA during characterization of the virus. All recovered, the Schultz EW and Habel K (1959) J Immunol 82 , 274 third case after treatment with ribavirin. The presumed rodent vector of the SA6 virus Synonym for cercopithecine virus is not known. herpesvirus 3. Synonym : SP H 114202 virus. Barry M et al (1995) N Engl J Med 333 , 294 SA8 virus Synonym for Cercopithecine Lisieux T et al (1994) Lancet 343 , 391 herpesvirus 2 . Vasconcelos PFC et al (1993) Rev Inst Med Trop , São Paulo 35 , 521 SA10 virus Synonym for simian para- Sabin vaccine A live attenuated polio- influenza virus type 3. myelitis vaccine containing all three serotypes of poliovirus. Given orally. SA11 virus A strain of Rotavirus A in the genus Rotavirus . Sabin virus (SFNV) A strain of Sandflyfever Naples virus in the genus SA12 virus Synonym for polyoma- Phlebovirus . virus papionis 1. Virus isolated from baboons. Sabo virus (SABOV) A strain of Akabane virus in the genus Orthobunyavirus . SA15 virus Synonym for cercopithecine Belongs to the Simbu serogroup. herpesvirus 4. Isolated from cattle, goats, and flies of Culicoides sp in Nigeria. Not reported SAAAr5133 virus (OLIV) A strain to cause disease in humans. of Olifantsvlei virus in the genus Orthobunyavirus . Saboya virus (SABV) A species in the genus Flavivirus. Member of the Yellow SAAn3518 virus (TETEV) A strain of Tete fever virus serogroup. Isolated from virus in the genus Orthobunyavirus . Kemp’s gerbil, Tatera kempi , in Senegal. Antibodies present in many mammals, SAAr53 virus (SIMV) A strain of Simbu birds, and reptiles. Not reported to virus in the genus Orthobunyavirus . cause disease in humans.

Saaremaa virus (SAAV) A strain of Sacramento River chinook salmon dis- Dobrava-Belgrade virus in the genus ease virus See Infectious hematopoi- Hantavirus . etic necrosis virus . saddle back fever 424 saddle back fever A pattern of fever fre- AIDS-like disease caused by Simian quently seen during infections with immunodeficiency virus in the genus arthropod viruses. The patient’s tem- Lentivirus . perature rises and falls, then a second Brody BA et al (1992) J Virol 66 , 3950 wave of fever occurs, often with no Stromberg K et al (1984) Science 224 , 289 other disease symptoms. Saimiriine herpesvirus 1 (SaHV-1) A S-adenosyl-L -homocysteine (AdoHcy , SAH) An inhibitor of methylation, e.g. species in the genus Simplexvirus , sub- of nucleic acids, as it is an analog of family Alphaherpesvirinae . Originally S -adenosyl- l -methionine. isolated from throat swabs and autopsy material from marmosets, Tamarinus S-adenosyl-L -methionine (AdoMet , nigricollis, in which it causes a fatal dis- SAM) A high-energy compound ease. Also isolated from owl monkeys, derived from ATP that is an intracel- Aotus sp, and squirrel monkeys, Saimiri lular source of activated methyl groups sp. The latter is the natural host for the including those used for RNA or RNA virus, and may excrete virus intermit- methylation. Stimulates transcription tently for long periods. Only minor in some viruses, e.g. cytoplasmic poly- disease is caused in spider and squirrel hedrosis viruses. Also required by class monkeys, but experimental or labora- I restriction endonucleases for their ini- tory contact infection of owl monkeys tial binding. and marmosets causes a generalized fatal disease. The virus replicates in Saffold virus (SAFV) A probable picorna- mouse and chick embryo and rabbit virus of the genus Cardiovirus isolated and marmoset kidney cell cultures. from a stool sample of an 8-month-old Pocks are produced on the CAM. It is female with fever of unknown origin. pathogenic for rabbits and adult mice The virus replicated in HFDK cells and given i.c., and for suckling mice and suckling mice. The diameter by elec- hamsters given i.p. tron microscopy was 28–30 nm. RNA Synonyms : callitrichid herpesvirus; her- sequence analysis showed that it was pesvirus M; herpesvirus platyrrhinae related to Theilovirus , a species in the type; herpesvirus T (tamarinus); mar- genus Cardiovirus. A second isolate of moset herpesvirus; M virus. the virus has been made from a child in Leib DA et al (1987) Arch Virol 93 , 287 Canada, but the global distribution and significance of this new human virus awaits further investigation. Saimiriine herpesvirus 2 (SaHV-2) Type species of the g enus Rhadinovirus . Jones MS et al (2007) J Clin Microbiol 45 , 2144 A ubiquitous infection of squirrel Drexler JF et al (2008) Emerg Inf Dis 14 , 1398 monkeys, Saimiri sciureus , in South America, first isolated in 1968 from an Sagiyama virus (SAGV) A strain of Ross owl monkey kidney tumor, and also River virus in the genus Alphavirus . from primary kidney cells derived Isolated from mosquitoes in Japan. from a squirrel monkey. Not patho- On injection kills newborn mice. Not genic in squirrel monkeys, in which reported to cause disease in humans. there is a lifelong persistent infection, SAH Abbreviation for S-a denosyl-l- but highly oncogenic causing leukemia h omocysteine. and lymphosarcomas in many New World primates including marmosets, SAIDS-D S imian a cquired i mmuno d efi- owl monkeys, capuchin monkeys, ciency syndrome-D. A rare fatal immu- spider monkeys, and howler mon- nosuppressive disease syndrome which keys. Peripheral blood lymphocytes occurs naturally and can be induced from these species are immortalized experimentally in macaque monkeys in vitro to interleukin-2 independent (Macaca mulatta) by infection with growth. The virus genome is unusual Mason–Pfizer monkey virus, a D-type and occurs in two forms, M and H, in retrovirus in the genus Betaretrovirus . the virion. M genome is linear double- The disease is distinct from the simian stranded DNA, 150 kb in length, with a 425 saliva

unique region of L-DNA (112 kb; 36% from a Kaiser’s rat, Aethomys kai- G C) flanked by multiple 1.4 kb repeat seri medicatus, in the Central African units (72% G C) of H-DNA. The H Republic. genome is also 150 kb long, but contains only repeat units of H-DNA arranged Salanga virus (SGAV) An unas- head-to-tail. Immortalized cells may signed ungrouped virus in the family contain only the left-hand end of Bunyaviridae. Isolated in suckling mice L-DNA, implicating these sequences in from the blood of a young specimen oncogenicity. Natural infection is per- of the Kaiser’s rat, Aethomys kaiseri petuated by horizontal transmission medicatus , trapped in Salanga, Central from adult to young monkeys through African Republic. No hemagglutinin saliva. No human infections have been detected. Not reported to cause disease reported. in humans. Synonyms : herpesvirus saimiri 2; squirrel monkey herpesvirus. Salehabad virus (SALV) A species in the Trimble JJ and Desrosiers RC (1991) Adv genus Phlebovirus, the first member of Cancer Res 56 , 335 the Salehabad complex, sandfly fever serogroup. Isolated from female sand Saint-Floris virus (SAFV) A tenta- flies of Phlebotomus sp in Iran. Probably tive species in the genus Phlebovirus , also present in Pakistan, as antibodies belonging to the sandfly fever virus are present in humans and sheep. Not group. Isolated from a gerbil of Tatera reported to cause disease in the wild. sp in the Central African Republic. Not reported to cause disease in humans. Salehabad I-81 virus (SALV) A strain of Salehabad virus . Saint Louis encephalitis virus (SLEV) See St Louis encephalitis virus . Salem virus (SaLV) An apparently novel paramyxovirus isolated from the mono- Sakhalin group viruses Antigenically nuclear blood cells of a horse involved related species in the genus Nairovirus , in a disease outbreak that occurred isolated from ixodid ticks taken from simultaneously at three race tracks in sea birds ’ nests. New Hampshire and Massachusetts, USA in 1992. Not clearly associated Sakhalin virus (SAKV) A species in the with the disease, so the isolation of the genus Nairovirus . The first member virus may have been fortuitous. of the Sakhalin serogroup. Isolated Glaser AL et al (2002) Vet Microbiol 87 , 205 from the tick, Ixodes putus , collected Renshaw RW et al (2000) Virology 270 , 417 on Tyuleniy Island off the south-east coast of Sakhalin Island where there Salisbury virus Synonym for human is a colony of guillemots, ( Uria aalge ). rhinovirus. Named for the MRC Com- Geographical distribution of the virus mon Cold Unit, near Salisbury, Wiltshire, appears to coincide with that of I UK where a great deal of valuable putus and probably involves the Kuril, work on respiratory viruses was car- Commodore and Aleutian Islands and ried out in human volunteers, under the northern coasts of Europe, Canada, the direction of David Tyrrell. Sadly and USA. Antibodies are present the Unit was closed by the government in guillemots but not in a variety of in 1990. other birds. Replicates in experimen- Tyrrell D and Fielder M (2002) Cold Wars . tally infected Culex modestus mosqui- Oxford: Oxford University Press. 253pp. toes. Not reported to cause disease in humans. saliva Some viruses such as cytomegalo- Lvov DK et al (1972) Arch Gesamte Virusforsch virus, rabies virus, and mumps virus, 38 , 133 replicate in the salivary glands and are discharged into the saliva to enter Salanga poxvirus (SGV) An unassigned the oral cavity and can be passed to virus in the family Poxviridae . Isolated another individual by a bite or other saliva 426

oral contact. HIV has been found in saltatory DNA replication Lateral ampli- very low concentrations in the saliva of fication of DNA resulting in repetitive some AIDS patients. However, contact copies of a DNA segment. See repeti- with saliva has never been shown to tive DNA . result in transmission of HIV. Sal Vieja virus (SVV) A species in the salivary gland virus See cytomegalo- genus Flavivirus , belonging to the virus group . Modoc virus group. Isolated from a rodent in Texas. Salk vaccine A formalin-inactivated poliomyelitis vaccine containing all SAM See S-adenosyl-L -methionine . three serotypes of poliovirus. Given by injection. Samford virus A strain of Aino virus. See Aino virus . Salmon pancreas disease virus (SPDV) A species in the genus Alphavirus . First recorded in 1995 in Atlantic salmon San Angelo virus (SAV) A strain of (Salmo salar) in Ireland. Subsequently California encephalitis virus in the genus a closely related strain of the virus Orthobunyavirus , belonging to the was isolated in France from rainbow California serogroup. Isolated from trout (Oncorhynchus mykiss ) suffering mosquitoes in Texas, USA. Antibodies from sleeping disease. Finally the virus found in raccoons and opossums. Not was also isolated from farmed Atlantic reported to cause disease in humans. salmon and rainbow trout in Norway. Comparison of all three virus isolates Sanban virus A TTV-like virus detected by genome sequence analysis, mono- in serum of a patient in Japan. A single- clonal antibody reactivity, and cross stranded circular DNA virus with only infection indicated that all are strains 57% homology to TTV. of a single species, SPDV. Hino S and Miyata H (2007) Rev Med Virol McLoughlin MF and Graham DA (2007) J 17 , 45 Fish Dis 30 , 511 Mushahwar IK (2000) J Med Virol 62 , 399 Okamoto H et al (1998) J Med Virol 56 , 128 salmon reovirus (SSRV) A strain in the genus Aquareovirus , related to Atlantic Sandfly fever Naples virus (SFNV) A spe- salmon reovirus. cies in the genus Phlebovirus , belonging to the Salehabad complex, sandfly salmonid herpesvirus 1 (SaHV-1) fever serogroup. Antigenically distinct An unassigned virus in the family from sandfly fever Sicilian virus. Strains Herpesviridae . Isolated from a post- or serotypes include Karimabad virus, spawning steelhead trout, Salmo gaird- Sabin virus, Tehran virus, and Toscana neri . Chum salmon fry, Oncorhynchus virus. keta , are also susceptible. Synonym : herpesvirus salmonis. sandfly fever Sicilian virus (SFSV) A tentative species in the genus Wolf K et al (1978) J Virol 27 , 659 Phlebovirus, member of the sandfly salmonid herpesvirus 2 (SaHV-2) fever serogroup. Antigenically distinct An unassigned virus in the family from Sandfly fever Naples virus. Herpesviridae . Isolated from salmon, Robeson G et al (1979) J Virol 30 , 339 Oncorhynchus masou, in Japan. Causes renal failure and liver atrophy in sandfly fever viruses Synonym for phle- yamame (landlocked O masou ). botomus fever viruses. Synonym : Oncorhynchus masou herpesvirus. Sandjimba virus (SJAV) An unassigned Kimura T et al (1983) Fish Pathol 17 , 251 vertebrate rhabdovirus. Isolated from a bird Acrocephalus schoenobaenus in the salmonis virus Synonym for Viral hemor- Central African Republic. Not reported rhagic septicemia virus . to cause disease in humans. 427 Sapovirus sand rat herpesvirus Synonym for murid distributed in Pacific marine mammals herpesvirus 6, an unassigned virus in including northern fur seals. May be a the family Herpesviridae . cause of abortions in sea lions. Causes a vesicular disease in swine, and it has sand rat nuclear inclusion agent been suggested that marine mammals Synonym for murid herpesvirus 6, may be a reservoir of virus for terres- an unassigned virus in the family trial mammals. There are five serologi- Herpesviridae . cal types. Antigenically very close to VESV. Antibodies have been found sandwich technique See indirect fluo- in laboratory workers but there is no rescent antibody test . evidence that disease is produced in Sangassou virus A probable species in humans. Isolated in Vero cells. Causes the genus Hantavirus , identified from CPE in primary porcine and human the African wood mouse ( Hylomyscus ) cells but not in cells of marine mam- in Equatorial Guinea. mals and rodents. et al Intervirology 10 Klempa B et al (2006) Emerg Inf Dis 12 , 878 Burroughs JN (1978) , 51 Smith AW and Boyt PM (1990) J Zoo Wildl Med 21 , 3 Sanger method A widely used technique McClenahan SD et al (2008) Virus Res in for sequencing DNA by terminating press . in vitro synthesis using dideoxy ribonucleotides. See DNA sequencing . San Miguel sea lion virus, serotype 1 Sanger F et al (1977) Proc Natl Acad Sci 74 , (VESV/SMSV-1) A strain of VESV. 5463 San Miguel sea lion virus, serotype 4 Sango virus (SANV) A strain of Shamonda (VESV/SMSV-4) A strain of VESV. virus in the genus Orthobunyavirus , San Miguel sea lion virus, serotype 17 belonging to the Simbu serogroup. (VESV/SMSV-17) A strain of VESV. Isolated from cattle and from flies of Culicoides sp in Nigeria and Kenya. Not San Perlita virus (SPV) A species in reported to cause disease in humans. the genus Orbivirus, belonging to the Modoc virus serogroup. Isolated from sanitation The quality of public health the cotton rat, Sigmodon hispidus , in sanitation is particularly important in Texas, USA. Not reported to cause dis- the control of virus infections spread ease in humans. by the fecal-oral route, such as enteroviruses and hepatitis viruses A and E. Santarém virus (STMV) An unassigned In poorly developed countries, infec- virus in the family Bunyaviridae . tion with these viruses is acquired at Isolated from Oryzomys sp in Para, an early age, with little disease in older Brazil. people. However when travelers from highly developed countries travel to Santa Rosa virus (SARV) A serotype other regions they may be at high risk of Bunyamwera virus in the genus of infection. Orthobunyavirus. Isolated from the mosquito, Aedes angustivittatus , in Durango, San Juan virus (SJV) A strain of Alujuela Mexico. virus in the genus Orthobunyavirus , belonging to the Gamboa virus sero- Santee-Cooper ranavirus (SCRV) A spe- group. Isolated from Aedeomyia squami- cies in the genus Ranavirus . Related pennis in Ecuador. viruses include largemouth bass virus, doctor fish virus, and guppy virus 6. San Miguel sea lion virus (SMSV) A strain of Vesicular exanthema of swine Sapovirus A genus in the family virus (VESV) in the genus Calicivirus . Caliciviridae , type species Sapporo virus , Isolated from sea lions on San Miguel the members of which form a distinct Island off the coast of California, USA, phylogenetic clade within the family. and St Paul Island, Alaska. Serological The viruses are associated with out- evidence suggests the virus is widely breaks of gastroenteritis in humans and Sapovirus 428

in pigs. There is also a tentative species in Oriximina, Brazil. Not reported to causing mink enteritis in the genus. cause disease in humans.

Sapphire II virus (SAPV) A serotype of sarcoma viruses Sarcoma viruses have Hughes virus in the genus Nairovirus . been isolated from a number of spe- Isolated from swallow ticks, Argas cies of birds and mammals but the cooleyi. Not reported to cause disease in most extensively studied are the fowl, humans. mouse, and cat viruses. Since they con- Yunker CE et al (1972) Acta Virol 16 , 415 tain a transduced cellular oncogene, one of the virus genes is usually absent Sapporo virus (SV) The type species or nonfunctional, so they are gener- of the genus Sapovirus in the fam- ally defective and require a helper ily Caliciviridae . Strains isolated from leukemia virus for replication. They many parts of the world form a distinct transform cells in culture and pro- phylogenetic clade within the family. duce infective virus if a helper virus Has been isolated from cases of severe is present. Sarcomas are produced on gastroenteritis in Japan, (Sapporo injection into animals, usually after a strain) Australia, (Parkville strain), the short latent period of a few days. Some UK (London 29845 and Manchester tumors spontaneously regress. strains), and the USA (Houston 86 and Houston 90 strains). sarkosyl A detergent used to break up virus particles or infected cells. Sapporo virus-Houston/86 (Hu/SV/Hou/ 1986/US) A strain of Sapporo virus in the SARS See severe acute respiratory syn- genus Sapovirus . drome . satellite Sapporo virus-Houston/90 (Hu/SV/Hou A subviral agent which lacks the genes required for replication. 27/1990/US) A strain of Sapporo virus Multiplication is dependent upon in the genus Sapovirus. Related phylo- co-infection of the host cell with a genetically to Manchester and Parkville helper virus. Two major types of satel- strains. lite are satellite nucleic acids and satellite viruses. Satellite nucleic acids may Sapporo virus-London/29845 (Hu/SV/ or may not encode nonstructural pro- Lon29845/1992/UK) A strain of Sapporo teins, and are encapsidated by the cap- virus in the genus Sapovirus . sid protein of the helper virus. Satellite viruses have nucleic acid genomes that Sapporo virus-Manchester (Hu/SV/Man/ encode a structural protein that encap- 1993/UK) Sapporo virus A strain of in sidates the genome. the genus Sapovirus . satellite RNAs Small RNA molecules Sapporo virus-Parkville (Hu/SV/Park/ found in some plant viruses which may 1994/US) A strain of Sapporo virus in the resemble viroids but can only replicate genus Sapovirus . in the presence of the ssRNA helper virus. Also called ‘ virusoids. ’ Saquinavir A licensed antiviral drug, an inhibitor of the HIV protease, also Francki R (1986) Annu Rev Microbiol 39 , 151 known as Ro31-8959. Extremely active satellite virus inhibitor of HIV replication in cell cul- An absolutely defective ture. A peptide-based transition state virus which, in nature, depends on the mimetic of the protein cleavage site. presence of a helper virus to provide Synonym : Invirase. some factor necessary for its replication. See Dependovirus . Roberts NA et al (1990) Science 248 , 358 Sathuperi virus (SATV) A species in the Saracá virus (SRAV) A serotype of genus Orthobunyavirus , belonging to Changuinola virus in the genus the Simbu serogroup. Isolated from Orbivirus . Isolated from Lutzomyia sp mosquitoes in Madras, India, and 429 scrapie agent

Ibadan, Nigeria. Also isolated from use as a live measles vaccine in 1965. cattle. Not reported to cause disease in Used in many parts of the world as the humans. measles vaccine component. Closely similar genetically to Moraten virus. Saumarez Reef virus (SREV) A spe- Rota JS et al (1994) Virus Res 31 , 317 cies in the genus Flavivirus belong- Schwarz AJF (1962) Am J Dis Child 103 , 256 ing to the Seabird tick-borne virus group. Isolated from sea bird ticks, sciurid herpesvirus 1 (ScHV-1) An un- Ornithodoros capensis , collected from assigned virus in the subfamily the nests of sooty terns on islands off Betaherpesvirinae . Seventeen of 81 the coast of Queensland, Australia and European ground squirrels, Citellus Ixodes eudyptidis from dead silver gulls citellus , trapped in the Timisoara area in northern Tasmania. Antigenically of Romania were found to have typi- most closely related to Tyuleniy virus . cal cytomegalovirus inclusion bodies Causes paralysis on i.c. injection in in their salivary gland cells. These ani- newborn mice. Not known to cause mals appeared to be in good health. disease in humans, or other animals. Cultures of the salivary gland cells St George TD et al (1977) Aust J Exp Biol Med developed foci of round refractile Sci 55 , 493 cells after 4 days, and in 10 days these spread to involve half the culture. The sawgrass virus (SAWV) An unassigned virus could not be passaged in mouse, species in the family Rhabdoviridae , rat, or human cultures. belonging to the Sawgrass serogroup. Synonyms : ground squirrel cytome- Isolated from the eastern dog tick, galovirus; European ground squirrel Dermacentor variabilis , and the rabbit cytomegalovirus. tick, Haemaphysalis leporis-palustris , in Florida, USA. Antigenically related Barahona HH et al (1975) Lab Anim Sci 25 , 725 Diosi P and Babusceac L (1970) Am J Vet Res to New Minto virus and Connecticut 31 , 157 virus. Not reported to cause disease in Diosi P et al (1967) Arch Gesamte Virusforsch humans. 20 , 383 Ritter DG et al (1978) Can J Microbiol 24 , 422 sciurid herpesvirus 2 (ScHV-2) An unas- scabby mouth virus Synonym for Orf signed virus in the family Herpesviridae . virus . Isolated from a spontaneously degenerating primary kidney cell culture from scaffold The assembly of large virus cap- a North American species of ground sids, such as the icosahedral capsid of squirrel, Citellus citellus , in which it is herpes simplex virus, occurs with the a latent infection. Kills suckling mice aid of a scaffold that is later removed on injection i.c. Produces pocks on the by virus-encoded protease. The scaffold CAM. Replicates well in rabbit, ham- itself is not icosahedral, but enforces ster, marmoset, and owl monkey cells, the correct curvature on the assembling but poorly in dog fetal lung and Vero subunits. In the herpes virus genome, cells. Virus is readily released from the scaffold and protease genes over- cells. Neutralized by herpesvirus san- lap, ensuring colocalization of the guinus antiserum but there is no recip- protease with the substrate scaffold rocal neutralization. Not neutralized by protein, which is removed before final antiserum to sciurid herpesvirus 1. assembly of the virus. Synonym : American ground squirrel herpesvirus. Scandinavian epidemic nephropathy Barahona HH et al (1975) Lab Anim Sci 25 , 735 virus Synonym for Puumala virus . Diosi P et al (1975) Pathol Microbiol 42 , 42

Schwarz virus A strain of Edmonston B scrapie See prion diseases . measles virus that was further attenuated by multiple passages in chick scrapie agent A prion which causes a embryo fibroblast cells and licensed for natural disease of sheep and goats, scrapie agent 430

mainly in the northern hemisphere. important cell surface molecules. The Affected animals are usually about scraped cells are vigorously pipetted 3 years old and have intense pruri- to obtain a single cell suspension, then tis and ataxia, which becomes severe replated in fresh medium. before death. They scratch themselves by scraping against fences, hence the scrum-pox A contagious disease of the name. Recovery in mild cases may be facial skin of rugby players. Etiology explained by misdiagnosis, which, to variable but human herpesvirus 1 and be certain, is by histological examina- vaccinia virus have been implicated. tion of the brain post-mortem. Mode Almost exclusively confined to for- of transmission is not clear, but prob- wards. A survey of 30 rugby clubs ably often vertical. In primates causes a identified 48 infected players, of whom condition similar to Creutzfeldt–Jakob 47 were forwards and one a scrum-half. disease. Chimpanzees are not sus- Of the 47 forwards, 32 played in the ceptible, and there is no evidence of front row, 8 in the second row, and 7 in infection in humans. Goats are more the back row of the scrum. 23 of them uniformly susceptible to experimental reported direct contact with opponents disease than sheep. Injections of brain who had obvious facial lesions; 34 tissue extract by any route will trans- had similar contacts within their own mit the disease, but the shortest incu- teams. Some of the cases were treated bation period (6–9 months) follows with idoxuridine. See also Norwalk i.c. injection. Mice and hamsters can virus . be infected, the virus replicating first Synonyms : herpes gladiatorum; her- in lymphoid tissue, then in the brain. pes rugbeiorum; herpes venatorum; Injection into mink causes a disease prop-pox. similar to transmissible mink encepha- Shute P et al (1979) BMJ 4 , 1629 lopathy, but the agent so passed will not reinfect mice. Different strains of SDAV Sialodacryoadenitis virus. See Rat agent cause slightly different signs coronavirus . and symptoms. Infectivity appears to reside in a small proteinaceous infec- SDS S odium d odecyl sulfate, an anionic tious particle called a ‘ prion. ’ Prions detergent. can be recovered from scrapie-infected brain as a 27–30 kDa protein called SDS-polyacrylamide gel electrophoresis PrPsc. They appear to be isoforms of a Electrophoresis in polyacrylamide gels cellular protein of the same amino acid of proteins denatured with the ani- sequence called PrPc. The sequence of onic detergent sodium dodecyl sulfate the PrPsc gene has been determined. (SDS). The treatment with SDS usually It is believed that the PrPsc, which in gives proteins equal charge per unit contrast to PrPc is highly resistant to molecular weight and thus the proteins proteolysis, can undergo autocatalytic are separated according to their molec- replication without the involvement of ular weight. nucleic acid. See prion . Chesebro BW (Editor) (1991) Curr Top sea-bass virus-1 (SBV) An unassigned Microbiol Immun 172 , 228 virus in the family Picornaviridae . Prusiner SB (Editor) (1999) Prion Biology and Diseases. Cold Spring Harb Lab Monogr 38 , seal distemper virus A strain of New York: Cold Spring Harbor Laboratory Phocine distemper virus in the family Press Paramyxoviridae . Sakudo A et al (2007) Vet Med Sci 69 , 329 Seadornavirus A genus in the family scraping Propagation of cells by scraping Reoviridae , type species Banna virus . The is a method for physically removing virus genome consists of 12 segments adherent cells from the culture vessel, of dsRNA. Virions are non-enveloped and is used when enzymatic removal with a diameter of 60–70 nm with two (e.g. by trypsin) may be toxic to the concentric capsid shells and a core of cells or destroy receptors or other 40–50 nm diameter. The viruses are 431 sedimentation coefficient

mainly found in China and Indonesia, Sebokele virus An unclassified virus. and are transmitted to their hosts by Isolated in suckling mice from the mosquito vectors. The genus contains pooled brain, liver, spleen and heart three species, Banna virus , Kadipiro virus tissue of an adult female rodent of and Liao ning virus , and there are at Hylomyscus sp, trapped in a banana least 15 other tentative species await- plantation at Botambi, Central African ing characterization. Republic. Infectivity not sensitive to Attoui H et al (2000) J Gen Virol 81 , 1507 chloroform. Not reported to cause disease in humans. seal influenza virus Occasional isolations Digoutte JP (1978) Am J Trop Med Hyg 27 , of influenza viruses have been made 424 from seals and whales and the viruses are found to be close relatives of avian secondary attack rate (SAR) A measure influenza viruses. In 1980, an epizootic of the infectivity of an agent. of H7N7 influenza occurred in seals Longini IM et al (1982) Am J Epidemiol 115 , on the coast near Boston, USA with 736 more than 100 deaths. Transmission of this virus to a laboratory worker was secretory immunoglobulin A (sIgA) The recorded. From other disease outbreaks form of IgA found in external body in seals H4N5 influenza viruses have secretions such as intestinal mucus, been recovered, and H1N1 viruses colostrum, milk, saliva, sweat, and have been isolated from Pacific ocean tears. It is mainly in the form of a dimer whales. In so far as these viruses have with secretory piece bound to it. been characterized, they appear to be closely related to avian influenza virus secretory piece A polypeptide of mol. wt. species. 60 kDa which is attached to secretory Webster RG et al (1992) Microbiol Rev 56 , 152 IgA. Synthesized by epithelial cells in the gut, lung, mammary, or other secretory tissues. Has strong affinity seal morbillivirus See Phocine distemper for mucus. It is part of a larger mole- virus . cule, a receptor for polymeric immunoglobulin on the surface of certain sealpox virus A tentative species in the epithelial cells. The receptor binds to genus Parapoxvirus . Morphologically IgA and transports it through the cell similar to Orf virus . Isolated from a after endocytosis. The receptor is then captive Californian sea lion, Zalophus cleaved, releasing IgA, with the remain- californianus . Causes a severe disease in ing part of the molecule (the secretory wild and captive sea lions. piece) still attached, into the lumen Wilson TM and Poglayen-Neuwall I (1971) of the gut, mammary gland or other Can J Comp Med 35 , 174 organ. seasonal outbreaks Many virus diseases sedimentation coefficient The sedi- seem to be seasonal, e.g. respiratory mentation rate of a protein or other viruses are more common in the win- macromolecule per unit of applied ter months, and in temperate zones gravitational force is termed the ‘ sedi- rotavirus transmission takes place mentation coefficient’ or constant, s , mainly during cold weather. Rubella and is defined by the equation: virus infections tend to increase in the spring, and hepatitis A virus infections dr s 1 peak in the fall. These seasonal varia- ω2 tions are not entirely understood, but rtd comparison of data concerning influenza virus outbreaks in the southern where r radius (the distance in cm and northern hemispheres shows that between the particle and the center the winter is the peak influenza season of rotation); ω the angular velocity in both regions. in radians/s of the centrifuge head; sedimentation coefficient 432

dr/dt the rate of movement of the selectins A family of cell adhesion mol- particle in cm/s. ecules. They are type I transmembrane The units of s are reciprocal seconds; proteins containing an N-terminal C- for convenience, the basic unit is taken type lectin domain, which can bind as 10 13 s, and termed one Svedberg carbohydrates, an epidermal growth unit (S). With this unit, the sedimenta- factor-like domain and two to nine tion coefficients of most proteins fall short repeat units homologous to between 1 and 50 S. domains found in complement bind- Svedberg T and Pedersen KO (1940) The ing proteins. There are three subsets of Ultracentrifuge . London: Oxford University selectins: L-selectin (CD62L), expressed Press in leucocytes, E-selectin (CD62E) expressed on vascular endothelia, and P-selectin (CD 62P) expressed on sedimentation rate The velocity at which platelets. The function of selectins is a particle (assumed to be approxi- to mediate neutrophil, monocyte, and mately a sphere) settles under a given lymphocyte rolling along the venular set of conditions. It is proportional to wall, thus influencing the extent and the square of the particle diameter and severity of the inflammatory response. to the difference between the particle density and the density of the sus- Tedder TF et al (1995) FASEB J 9 , 866 pending medium. It decreases as the viscosity of the suspending medium Seletar virus (SELV) A serotype of Wad increases and increases with the gravi- Medani virus in the genus Orbivirus , tational force. belonging to the Wad Medani com- d ()ρρ g plex of the Kemerovo virus serogroup. Sedimentation rate 2 PL μ Isolated from the tick, Boophilus micro- 18 plus , in Singapore and peninsular where d diameter of the particle; Malaysia. Antibodies found in cattle, ρ P particle density; carabao (water buffalo), and pigs. Not ρ L suspending medium density; reported to cause disease in humans. μ viscosity of suspending medium; Sembalam virus An unclassified arbo- g gravitational force. virus. Isolated from a night heron (Nycticorax nycticorax) in Vellore, India It is measured in cm/s. in 1963, and 4 further strains from the grey heron (Ardea cinerea ). Not reported Sedlec virus A possible arbovirus of the to cause disease in humans. family Bunyaviridae . Isolated from the reed warbler, Acrocephalus scirpaceus semi-conservative replication A model (Herm ), in Nesyt fishpond in Southern for DNA replication as it normally Moravia. occurs in nature. The double strand Hubalek Z et al (1990) Acta Virol 34 , 339 becomes separated and each base in the single strand becomes attached to Seewis virus A probable species in the complementary nucleotides to form genus Hantavirus, detected by PCR two new double strands, in which one analysis in tissues of the common shrew strand of each daughter molecule will (Sorex araneus) captured in Switzerland. be derived from the original DNA. Song JW et al (2007) Virol J 4 , 114 Semliki Forest virus (SFV) (Semliki means ‘I do not know ’ – the reply segmentation Segmentation of the virus given by natives when asked the genome is seen in the arenaviruses, name of the forest.) A species in the birnaviruses, bunyaviruses, orthomyxo- genus Alphavirus. The natural host and viruses, and reoviruses. One advantage vector are not known but antibodies this offers to the virus is the potential are found in humans and wild pri- for increasing genetic heterogeneity by mates in Uganda, Mozambique, reassortment of the segments during Cameroon, Central African Republic, replication and morphogenesis. Kenya, Nigeria, northern Borneo, and 433 Seoul virus (SEOV)

Malaysia. Multiplies in the mosquito, is available to protect laboratory Aedes aegypti. First isolated in Uganda mice. Not generally accepted as caus- in 1942. Not associated with any dis- ing human infection. Injected i.c. in ease, and generally considered to be mice it causes a fatal infection but non-pathogenic for humans although serial passage by this route is not pos- a single fatal case of encephalitis in a sible. Causes inapparent infection in laboratory worker has been associated ferrets, monkeys, and pigs and occurs with the virus. Causes encephalitis in as a latent infection of laboratory mice, adult mice on experimental injection rats, and guinea pigs. Used extensively by various routes and on i.c. injection as a model paramyxovirus for mole- in guinea pigs, rabbits, and rhesus cular studies, and to induce cell fusion. monkeys. Most inoculated animals The virus is also being developed as a have shown kidney damage. Viremia gene vector. Removal of the envelope occurs in inoculated birds of several genes results in a virus with reduced species and in hamsters. Virus can be cytopathogenicity and immunogenicity propagated in eggs killing the embryo that can be used to deliver other genes and in cell cultures of many species for clinical therapeutic applications. with CPE. The virus has proved to be Synonyms : hemagglutinating virus a valuable model for the investigation of Japan; murine parainfluenza virus of pathogenesis, and the SFV replicon type 1. can be used to express heterologous Faisca P and Desmecht D (2007) Res Vet Sci genes and so to generate recombinant 82 , 115 proteins for research, vaccination, gene Ishida N and Homma M (1978) Adv Virus therapy, and cancer therapy. Res 23 , 349 et al J Gene Med 9 Lundstrom K (2005) Gene Ther 12 , Suppl 1, Yoshizaki M (2006) , 1151 S92 McKimmie CS and Fazakerley JK (2005) sentinel animal An animal exposed cap- J Neuroimmunol 169 , 116 tive in the wild to contact infection Willens WR et al (1979) Science 203 , 1127 from the environment, usually from insects. Used to test for the presence Sena Madureira virus (SMV) An unas- of viruses in a particular geographical signed virus in the family Rhabdoviridae , area. belonging to the Timbo serogroup. Isolated from lizards ( Ameiva ameiva sentinel case An isolated case of some ameiva) in Brazil in 1976. Not reported infectious disease or the first case in an to cause disease in humans. outbreak.

Sendai virus (SeV) The type species in sentinel virus (SNNV) A TTV-like virus the genus Respirovirus. Causes a highly isolated from sera of non-A-G hepatitis transmissible respiratory tract infec- patients by representational difference tion of mice, hamsters, and guinea analysis. pigs. Found in mouse colonies world- Mushahwar IK (2000) J Med Virol 62 , 399 wide, usually in suckling or young adult mice. Can be isolated in embryo- Sen virus (SENV) A strain of Torque teno nated eggs, the allantoic cavity being virus . Isolated from an HIV patient (ini- the most sensitive, or in primary cul- tials SEN). tures of mouse, human, chick, and Mushahwar IK (2000) J Med Virol 62 , 399 other species. Less exacting in require- ments for replication and causing a Seoul virus (SEOV) A species in the more definite CPE than parainfluenza genus Hantavirus . First isolated from virus type 1 human. Produces plaques rats in 1980 in Korea. Infects Rattus rat- in human, bovine, and simian cells. tus and Rattus norvegicus and has been Inactivated virus is used experimen- isolated in Japan, Korea, China, Egypt, tally to cause cell fusion. On mouse and the USA. Human infection can to mouse passage there is an increase occur where there is close contact with in virulence resulting in pneumonia. rats, and there is evidence of nephro- An inactivated chick embryo vaccine pathy following these infections. The Seoul virus (SEOV) 434

virus is probably distributed world- serogroups Groups of viruses that cross- wide in Rattus species, but does not react with each other antigenically. cause disease in the natural host. serological surveillance The measure- Sepik virus (SEPV) A species in the genus ment of serum antibody levels in a Flavivirus, in the Yellow fever virus population to look for the presence of a group. Isolated from mosquitoes in particular infectious agent. New Guinea. Causes a febrile illness in humans. serology The study of antibody levels in septicemia anserum exudative virus sera of an individual or a population. Synonym for Derzsy’s disease of geese, caused by goose parvovirus. seron A name proposed but not adopted for any group comprised of viruses sequence The order of nucleotides in with antigenic similarity which can be RNA or DNA or of amino acids in a demonstrated serologically. polypeptide. seroprevalence The number of existing sequencing Determination of the cases of a virus disease as measured sequence of nucleotides in a nucleic serologically. acid or of amino acids in a protein. See DNA sequencing , Sanger method , serpins Serine protease inhibi- Maxam–Gilbert method, and Edman tors encoded by poxviruses. degradation . Orthopoxviruses and leporipoxviruses encode three, whereas avipoxviruses serine-threonine kinase A cellular can encode five or more. enzyme related to protein kinase C which is a target for several oncogenes, serpin SP-1 An orthopoxvirus serine including Akt , Raf, and Ras. Akt is the protease which inhibits cathepsin G, a transforming gene of AKT8 virus, Raf major constituent of neutrophils, and is is the transforming gene of murine a host range gene required for the virus retrovirus 3611-MSV, and Ras is the to grow on human and swine cells. transforming gene of the Harvey and Kirsten species of murine sarcoma virus. serotype A measure of the antigenic properties of a virus, important in virus seroarcheology The use of sera from characterization. Determined by rais- older adults to detect what viruses ing antisera against the virus in rabbits may have been circulating in the past. or other species, and using the antisera Especially valuable for studies of influ- in neutralization, immunofluorescence enza pandemics in the nineteenth and or tests to compare it to other viruses. early twentieth century. Serra do Navio virus (SDNV) A strain Mulder J and Masurel N (1958) Lancet 1 , 810 of California encephalitis virus in the seroconversion Acquisition of detectable genus Orthobunyavirus , belonging to antibodies to a virus in the serum of an the California encephalitis virus group. individual. Isolated from mosquitoes, Aedes fulvus, in Amapa Territory of Brazil. Not serodiagnosis Diagnosis of a virus reported to cause disease. infection based upon seroconversion. Confirmation of infection ide- Serra Norte virus An unassigned virus ally requires two samples of serum, isolated from the blood of febrile one early (within a few days) after patients in the Amazon region of exposure and one convalescent (a few Brazil. weeks) after exposure. A four-fold rise in serum antibody titer against the Serra Sul virus An unassigned virus iso- infecting virus is taken as confirmation lated from phlebotomine sand flies in of infection with that virus. Brazil. 435 sexually transmitted diseases serum-free medium Cell culture medium acute respiratory syndrome corona- without serum. After a period in virus, that had not previously been serum-free medium, cells can be syn- seen in the human population, and was chronized by addition of serum, stimu- only distantly related to other animal lating cell division. and human coronaviruses. Guo Y et al (2008) Virus Res 133 , 4 serum hepatitis virus An old name for Hepatitis B virus. The name derives Severe acute respiratory syndrome corona- from the fact that the virus was often virus (SARS-CoV) A species in the genus transmitted by serum in blood trans- Coronavirus , belonging to group 2. fusions or repeated needle use in the The cause of severe acute respiratory process of giving injections, skin scari- syndrome. The S protein of the virion is fication (vaccination), ear-piercing, or uncleaved, in contrast to other group 2 tattooing with inadequately sterilized viruses. There is evidence that the instruments. natural host of the virus is the Chinese horseshoe bat (Rhinolophus sinicus ), serum neutralization Inhibition of virus and an intermediate host in transmis- infectivity by antiserum. sion of the virus to humans was the masked palm civet (Paguma larvata ). In severe acute respiratory syndrome a remarkable international public health (SARS) A life-threatening and rapidly response to the outbreak of SARS, the progressing form of pneumonia that virus was characterized and completely first emerged in Guangdong province, sequenced by several different groups China in November 2002. The clinical by May 2003, and the spread of the dis- course includes early nonspecific influ- ease was controlled by July 2003. enza-like symptoms, including persistent fever, chills, rigors, myalgia, and Cheng VC et al (2007) Clin Microbiol Rev 20 , general malaise. After several days, 660 Lau SK et al (2005) Proc Natl Acad Sci 102 , respiratory symptoms (dry cough, 14040 rhinorrhea, shortness of breath) and Wang LF and Eaton BT (2007) Curr Top gastrointestinal symptoms (nausea, Microbiol Immunol 315 , 325 vomiting, diarrhea) develop in most cases. About 8 days after the onset of severe combined immunodeficiency A fever, there is a rapid, often bilateral genetic disorder which results in a progression of ground glass opacities life-threatening syndrome of recurrent on chest radiographs, and multifocal infections. Both arms of the immune and diffuse consolidation develops. The response (B cells and T cells) are lung pathology may show pneumo- blocked and antibody production is mediastinum and pneumothorax, and impaired severely. The condition can about 20% of the patients deteriorate arise from mutations in any one of 10 with evidence of an acute respiratory known genes. The condition has been distress syndrome requiring intensive cured successfully by gene therapy care and mechanical ventilation. The using a retroviral vector derived from disease was taken to Hong Kong by Moloney murine leukemia virus. But a sick Chinese doctor who stayed in two of the patients developed a leuke- room 9/11 of a hotel where he trans- mia-like disease because of the site of mitted the infection to other resi- integration of the vector. dents from various parts of the world. Hacein-Bey-Abina et al (2003) N Engl J Med Between February and July 2003 the 348 , 255 disease was spread to some 8,400 persons worldwide, of whom more than sexually transmitted diseases Several 800 (10%) died. Dr. Carlo Urbani, of viruses cause sexually transmitted dis- the WHO, who first described the new ease, usually acquired by exposure to disease while working in Vietnam, virus-containing secretions. Infection became infected and died from it. The occurs through breaks in the skin or cause was identified in March 2003 as a direct invasion of the superficial epi- new human coronavirus, named severe thelium of mucous membranes. The sexually transmitted diseases 436

viruses include hepatitis B and HIV sheep pulmonary adenomatosis-associ- (associated with persistent viremia). ated herpesvirus Synonym for ovine herpesvirus 1. SH genes The genes encoding the small hydrophobic proteins of sheep rotaviruses Members of group A paramyxoviruses. and B rotaviruses, family Reoviridae . Shamonda virus (SHAV) A species in the sheldgoose hepadnavirus Avian hepad- genus Orthobunyavirus , belonging to the naviruses have been isolated and char- Simbu serogroup. Isolated from cat- acterized from the Orinoco sheldgoose tle and Culicoides sp in Nigeria. Not (Neochen jubatus ) and the ashy-headed reported to cause disease in humans. sheldgoose (Chloephaga poliocephala ) Shark River virus Strain of Patois virus in Guo H et al (2005) J Virol 79 , 2729 the genus Orthobunyavirus , belonging to the Patois serogroup. Isolated from mos- Shiant Islands virus (SHIV) A strain of quitoes in Florida, USA, Mexico, and Great Island virus in the genus Orbivirus , Guatemala. Not reported to cause dis- belonging to the Great Island complex. ease in humans. shifty sequence A short nucleotide sheatfish iridovirus An iridovirus-like sequence motif which facilitates ribo- agent affecting sheatfish, Silurus glanis . somal frameshifting during expression Siwicki AK et al (1999) Virus Res 63 , 115 of viral mRNA. Present in the replicase gene of coronaviruses, where the sheep-associated malignant catarrhal sequence is UUUAAAC. See also slip- fever of cattle virus Synonym for pery sequence . Ovine herpesvirus 2 . Shine–Dalgarno sequence A short stretch sheep papillomavirus (SPV) See Ovine of nucleotides on a prokaryotic mRNA papillomavirus 1 . molecule, upstream of the translational start site, that binds to ribosomal sheeppox subgroup virus Synonym for RNA and brings the ribosome to the capripoxvirus. initiation codon on the mRNA. The sequence is AGGAGG. See ribosome Sheeppox virus (SPPV) A species in the binding site , Kozak sequence . genus Capripoxvirus . Causes a generalized pox disease in sheep, often shingles A painful local condition with with tracheitis and involvement of the rash in the region (dermatome) served lungs. Mortality 5–50%. Economically by one nerve root. May follow expo- important. Occurs in parts of Africa, sure to human herpesvirus 3 (varicella- Asia, Middle East, southern Europe, zoster virus), but usually occurs as a and Iberian peninsula. Only sheep reactivation of latent infection with are infected naturally. Virus is closely varicella virus, especially in patients related antigenically to goatpox virus who are immunocompromised. and lumpy skin disease virus. Difficult Synonym : zona; zoster. to adapt to growth in eggs. Replicates in sheep, goat, and calf cell cultures with CPE. An attenuated virus vac- shingles vaccine A live attenuated vari- cine is used successfully, and has cella-zoster vaccine (Zostavax-Merck) been genetically engineered to protect which was approved by the FDA in against rinderpest, in addition to sheep 2006 for prevention of herpes zoster and goatpox and lumpy skin disease. (shingles) in persons 60 years of age Synonyms : Clavelee virus; Isiolo virus; or older. Contains a much higher dose Kedong virus; variola ovina virus. of varicella-zoster virus than does Varivax, which was licensed in 1995 for Gershon PD et al (1989) J Virol 63 , 4703 use in infants to vaccinate against vari- Kitching RP (1999) In Encyclopedia of Virology , cella (chicken-pox). Second edition, edited by A Granoff and RG Webster. London: Academic Press, p. 1376 Holcomb K and Weinberg JM (2006) J Drugs Romero CH et al (1993) Vaccine 11 , 737 Dermatol 5 , 863 437 signal recognition particle (SRP) shipping fever virus A bovine strain of sialic acid Synonym for N -acetyl- the species parainfluenza virus type neuraminic acid. See neuraminidase . 3. Causes respiratory disease in cattle, especially under stress. sialodacryoadenitis virus (SDAV) A Synonym : SF-4 virus. strain of Rat coronavirus . Frank GH and Marshall RG (1973) J Am Vet Med Assoc 163 , 858 sialoglycoconjugates Receptors for paramyxoviruses on the surface of cells. shipyard eye Epidemic keratoconjunctivitis, associated with human adeno- sialoglycoproteins Receptors for parvovirus types 8, 19 and 37. viruses on the surface of cells.

Shokwe virus (SHOV) A strain of sialyloligosaccharides Sugar-contain- Bunyamwera virus in the genus ing receptor molecules on the surface Orthobunyavirus , belonging to the of normal cells to which influenza Bunyamwera virus group. Isolated virus hemagglutinins bind. They α from Aedes cumminsii mosquitoes in include: ( 2, 6)sialyl lactose; N -acetyl- α Natal, South Africa. Not reported to neuraminic acid- 2, 6-galactose; and α cause disease in humans. ( 2, 3)sialyl lactose. Influenza viruses vary in their receptor specificity. Most avian influenza viruses bind prefer- Shope fibroma growth factor A pro- α tein encoded by Shope fibroma virus entially to the NeuAc 2, 3Gal linkage but most human influenza viruses and malignant rabbit fibroma virus. α Responsible for tumor induction of prefer binding to the NeuAc 2, 6 Gal tumors in host species such as rabbits. linkage. Couceiro JN et al (1993) Virus Res 29 , 155 Shope fibroma virus Synonym for Rabbit fibroma virus . siamese cobra herpesvirus Synonym for elapid herpesvirus 1. Shope papillomavirus Synonym for rabbit oral papillomavirus. sigla A contracted form of sigilla (Latin: sigillum seal). In virology, a device Show fever virus Synonym for Feline formed of letters, especially initials, or panleukopenia virus . other characters taken from the principal words in a compound term, e.g. SH protein A s mall h ydrophobic protein papovavirus. expressed in cells by paramyxoviruses. In human respiratory syncytial virus it sigma virus (SIGMAV) An unassigned is a short (64 amino acids) glycoprotein animal rhabdovirus in the family anchored to the membrane. It’s func- Rhabdoviridae which infects insects. The tion is unknown. virions are slightly smaller than vesicular stomatitis virus (VSV). Congenitally Shuni virus (SHUV) A species in the transmitted through the germinal cells, genus Orthobunyavirus, belonging to and confers carbon dioxide sensitivity the Simbu serogroup. Isolated from upon Drosophila melanogaster . mosquitoes, culicoid flies, humans, sheep, and cattle in Nigeria and South signal peptide A short amino acid Africa. Has been associated with dis- sequence at the N-terminus of a pro- ease in humans. Strains of Shuni virus tein, which is recognized by cellular are Aino virus and Kaikalur virus. membranes as the signal for glycosylation of the protein. The signal peptide Siadenovirus A genus in the family is cleaved off by a signal peptidase Adenoviridae. The type species is Frog during maturation of the protein. adenovirus , and the other species in the Lingappa VR et al (1979) Nature 281 , 117 genus is Turkey adenovirus A . They are serologically distinct from members of signal recognition particle (SRP) A other adenovirus genera. multi-subunit protein present in the signal recognition particle (SRP) 438

cytosol that binds to ribosomes shortly Simbu virus (SIMV) A species in the after they have synthesized the signal genus Orthobunyavirus , belonging to peptide and serves to bind the ribo- the Simbu serogroup. Isolated from somes to the endoplasmic reticulum. mosquitoes in South Africa, Central Recognizes both the N-terminal amino- African Republic, and Cameroon. acid of the nascent polypeptide chain Antibodies found in humans but the and a receptor on the rough endoplas- virus is not reported to cause disease. mic reticulum (RER) and halts further translation until the ribosome has simian adenoviruses 1 to 18 and become bound to the RER. 20 (SAdV-1 to -18 and SAdV-20) Strains or serotypes in the genus signal sequence A specific sequence, Mastadenovirus . Many strains were immediately following the initiation found because they produced CPE in codon, in the mRNA coding for a secre- kidney cell cultures of Asian, African, tory protein. It encodes a hydrophobic and New World monkeys. Several peptide of 15–30 amino acids which types have been associated with respi- binds to the signal recognition particle ratory and enteric disease in baboons, and is directed to the RER. As transla- rhesus, Erythrocebus, and Cercopithecus tion continues, the signal sequence is monkeys, but most infections are silent. extruded across the membrane of the Some strains are oncogenic in newborn RER, and the signal peptide is cleaved hamsters. Division into groups is pos- off the nascent protein by a specific sig- sible on the basis of agglutination of rat nal peptidase enzyme. and rhesus erythrocytes. Kalter SS et al (1979) Personal commu- signal transducing activators of tran- nication scription (STATs) Cellular transcrip- Merkow LP and Slifkin M (1973) Prog Exp tion factors that become activated in Tumor Res 18 , 67 response to interferons and migrate to the cell nucleus where they bind to simian adenovirus 19 (SadV-19) A mon- specific interferon response elements key adenovirus type reclassified on the in promoter sites upstream of inter- basis of sequence data, as a strain of feron-inducible genes, activating their human adenovirus-F. transcription. Simian enterovirus A A species in the genus Enterovirus . Contains four sero- Sikhote-Alyn virus (SAV) An unas- types: A1, A2 plaque virus, SV4, and signed virus in the family Picornaviridae SV28. related to Syr-Darya Valley fever virus. Isolated from Ixodes persulcatus simian enterovirus 1 to 17 . Tentative spe- ticks collected from a wild boar in the cies in the genus Enterovirus , with 17 Primorie region of Russia. The virus serotypes,. They are A13, N125, N203, had properties of a cardiovirus. SA5, SV16, SV18, SV19, SV2, SV 26, Lvov DK et al (1978) Acta Virol 22 , 458 SV35, SV42, SV43, SV44, SV45, SV47, SV49, and SV6. Isolated from monkey silent infection An infection with no tissues or excreta in cell cultures. Often apparent signs or symptoms. latent infections appearing in kidney cell cultures from apparently normal silurid herpesvirus 1 See Ictalurid animals. They have been given num- herpesvirus . bers in the SA and SV series of simian viruses. See simian viruses . Silverwater virus (SILV) An unas- Synonym : ecmoviruses. signed virus in the family Bunyaviridae . Serologically related to Kaisodi and Simian foamy virus Six species are rec- Lanjan viruses. Isolated from ticks ognized in the genus Spumavirus on removed from hares in Manitoulin the basis of nucleotide sequence stud- Island and the Powassan district ies. Three are simian, and the other of northern Ontario, Canada. Not three are bovine, equine and feline reported to cause disease in humans. viruses. No natural human infections 439 simian papillomavirus

are known, but a few have been docu- Hepatovirus. Isolated from cynomolgus mented as a result of rare zoonotic and African green monkeys. Related transmission from non-human pri- to human hepatitis A virus, and cross- mates. No diseases have been associ- reacts serologically, but genetically dis- ated with spumavirus infections, and tinct from all human isolates. no oncogene-containing member of the Nainan OV et al (1991) J Gen Virol 72 , 1685 genus has been found. Foamy viruses Tsarev SA et al (1991) J Gen Virol 72 , 1677 can be identiﬁ ed by recognition of foamy degeneration and syncytia for- Simian immunodeficiency virus (SIV) mation in cell cultures. A species in the genus Lentivirus . The first isolate was in 1984 from rhesus Table S1. Species of Spumavirus macaques, Macaca mulatta, and is called SIVmac. Subsequently, strains have 1 . Macaque simian foamy virus 1 , from rhesus been isolated from a range of simians monkey, Macaca mulatta , (simian foamy including sooty mangabey (SIVsm), virus 1) Sykes monkey (SIVsyk), stump-tailed 2 . African green monkey simian foamy virus . (Simian foamy virus 3 ) macaque (SIVstm), pig-tailed macaque 3 . Simian foamy virus . Human isolate. (SIVmne), mandrill (SIVmnd), chim- Chimpanzee foamy virus. Simian foamy panzee (SIVcpz), grivet (SIVgr-1), and virus, chimpanzee isolate. African green monkey (SIVagm). Causes 4 . Bovine foamy virus . a disease known as simian AIDS in rhe- 5 . Equine foamy virus . sus (Asian) or cynomolgus monkeys 6 . Feline foamy virus . which is remarkably similar to AIDS in humans. In their own species of monkey, SIV viruses appear to be non-path- Simian foamy virus 1 (SFV-1) A species ogenic. Human infection with SIV has in the genus Spumavirus. Isolated from occurred in persons working with the a macaque monkey. virus or handling monkeys. There is no Simian foamy virus 3 (SFV-3) A species evidence of disease in these cases, the in the genus Spumavirus. Isolated from virus of which is called SIVhu ( Table S2 ). an African green monkey. Khabbaz RF et al (1994) N Engl J Med 330 , 172 Simian hemorrhagic fever virus (SHFV) Novembre FJ et al (1998) J Virol 72 , 8841 A species in the genus Arterivirus . Causes asymptomatic acute or per- simian papillomavirus A natural infec- sistent infection in several species of tion of cebus monkeys causing papil- African monkey including patas mon- lomas. Papillomas can be transmitted keys, Erythrocebus patas, a natural host, experimentally to both New and Old but severe, often fatal, disease in rhesus World monkeys. After injection of tis- macaque monkeys. There is high fever, sue extract, hyperemic patches appear facial edema, splenomegaly, and severe within 2 weeks and then the epidermis hemorrhagic diathesis. Replicates in becomes thickened to form a papil- rhesus monkey cell cultures with CPE. loma. No evidence of invasion of nor- Non-pathogenic on injection into mice. mal tissue, and regression occurs in Genome is positive single-stranded 4–6 months. Papilloma viruses have RNA, about 15.7 kb in length. There been isolated from a penile lesion of a are four structural proteins. Diameter Colobus monkey and shown to cause 50–60 nm, with a lipid envelope and papillomas in other Colobus monkeys, 12–15 nm surface peplomers. Labile at and several different papillomavirus pH 3 and inactivated by chloroform. types were isolated from the genital regions of rhesus monkeys. It is likely Brinton MA and Snijder EJ (2008) in BWJ that a systematic study of these pri- Mahy and MHV van Regenmortel (eds) Encyclopedia of Virology, Third edition, mate species would yield a wealth of Oxford: Academic Press, Vol 1, p. 176 simian papillomaviruses for study. Synonym : monkey papillomavirus. simian hepatitis A virus (SHAV) A Chan SY et al (1997) J Virol 71 , 4938 strain of Hepatitis A virus in the genus Chan SY et al (1997) Virology 228 , 213 simian parainfluenza virus type 5 (SV5) 440

Table S2. Strains of Simian immunodeficiency virus

simian immunodeficiency virus African green monkey 155 (SIV-agm.155) simian immunodeficiency virus African green monkey 3 (SIV-agm.3) simian immunodeficiency virus African green monkey Sab-1 (SIV-agm.sab) simian immunodeficiency virus African green monkey Tan-1 (SIV-agm.tan) simian immunodeficiency virus African green monkey TYO (SIV-agm.TYO) simian immunodeficiency virus African green monkey (SIV-agm) simian immunodeficiency virus chimpanzee SIV (SIV-cpz) simian immunodeficiency virus pig-tailed macaque (SIV-mne) simian immunodeficiency virus red capped mangabey SIV (SIV-rcm) simian immunodeficiency virus rhesus (Maccaca mulatta ) (SIV-mac) simian immunodeficiency virus sooty mangabey SIV-H4 (SIV-sm) simian immunodeficiency virus stump-tailed macaque (stm) (SIV-stm) simian immunodeficiency virus Sykes monkey SIV (SIV-syk)

Lucke B et al (1950) Fed Proc Am Soc Exp Biol strain of Gibbon ape leukemia virus . The 9 , 336 pet woolly monkey lived in the same Reszka AA et al (1991) Virology 181 , 787 household as a gibbon ape, and it is now clear that the woolly monkey simian parainfluenza virus type 5 (SV5) was infected by a virus it caught from See Simian virus 5 . the gibbon. A non-transforming virus simian parainfluenza virus type 10 which acts as helper for the replica- (SPIV10) See Simian virus 10 . tion-defective woolly monkey sarcoma virus. Several strains can be isolated simian parainfluenza virus type 41 from gibbons. Probably originated (SV41) See Simian virus 41 . from an endogenous C-type virus of the Asian mouse, Mus caroli, to which simian polyomaviruses There are three it is very closely related. species: African green monkey polyomavi- Synonym : gibbon ape lymphosarcoma rus , Simian virus 12 (SA12), and Simian virus. virus 40 (SV40). Ting YT et al (1998) J Virol 72 , 9453 simian retrovirus 1 (SRV-1) A serotype of Mason–Pfizer monkey virus in the genus simian spumavirus See Simian foamy Betaretrovirus . virus . simian retrovirus 2 (SRV-2) A serotype of simian T-cell leukemia virus (STLV-1) Mason–Pfizer monkey virus in the genus See simian T-lymphotropic virus . Betaretrovirus . simian T-lymphotropic virus 1–3 (STLV- simian rotavirus See Rotaviruses group A . 1 to -3) Strains of Primate T-lympho- simian rotavirus SA11 (SiRV-A/SA11) tropic viruses 1–3 , species in the genus The type species of the genus Rotavirus , Deltaretrovirus. STLV isolates have been prototype strain of Rotavirus A . Isolated made from various nonhuman primate from the rectum of a healthy vervet species including Japanese macaque, monkey. Can be propagated in primary pigtailed macaque, bonnet macaque, vervet monkey cell cultures or in a cell stump-tailed macaque, Taiwanese line from this species. Produces CPE. macaque baboon, African green mon- Morphologically similar to and anti- key, tantalus monkey, and chimpanzee. genically related to human and bovine The viruses from African monkeys and rotaviruses. Can be used as antigen to chimpanzees are closely related geneti- titrate human and bovine rotavirus cally (more than 95% sequence homol- antibodies. ogy) to human T-lymphotropic virus 1 (HTLV-1) whereas the Asian STLV simian sarcoma virus (WMSV) A syno- strains are only 90% related genetically nym for Woolly monkey sarcoma virus to Asian HTLV-1 strains. Two more dis- in the genus Gammaretrovirus origi- tantly related STLV isolates have been nally isolated from a fibrosarcoma in made, from a bonobo, Pan paniscus , a pet woolly monkey, but actually a and a baboon, Papio hamadryas , from 441 simian viruses

Eritrea. The latter isolate was called required for virus replication in cell STLV-PH 969, and both isolates appear culture. more closely related genetically to HTLV-II rather than HTLV-I. Because it Simian virus 10 (SV-10) A species in is difficult to distinguish the STLVs and the genus Respirovirus . Isolated from HTLVs genetically, the species name is the mouth of a samango monkey, now Primate T-lymphotropic virus 1 , 2, Cercopithecus mitis . Agglutinates human, and 3. Leukemia has not been observed bovine, and guinea pig erythrocytes. in the monkeys from which STLV iso- Synonym : SA10 virus. lates were made. Simian virus 12 (SV-12) A species in the van Brussel M (1996) J Gen Virol 77 , 347 Yanigahara R (1994) Adv Virus Res 43 , 147 genus Polyomavirus . A natural infection of the chacma baboon, Papio ursinus , in simian type D virus (SRV-1) A strain Africa. of Mason–Pfizer monkey virus in the Valis JD et al (1977) Infect Immun 18 , 247 genus Betaretrovirus. Causes immunosuppression (SAIDS-D) in macaque Simian virus 40 (SV40) A species in the monkeys and originally called simian genus Polyomavirus . A natural and AIDS D-type (SAIDS-D) virus. Virions silent infection of rhesus, cynomol- have a type D morphology and lack gus and Cercopithecus monkeys. Often prominent spikes. The genome is typi- isolated from kidney cell cultures. cal of a retrovirus, with the structure Replicates in a variety of cell cultures 5-gag-pro-pol-env- 3 and no oncogenes. but when first isolated was cytopathic Reverse transcriptase has a preference for grivet monkey kidney cell cultures for magnesium rather than manganese only, producing vacuolation of the cyto- and is primed by tRNA-Lys. Virions plasm. Does not hemagglutinate. Foci induce cell fusion in cell cultures, and of transformed cells appear in human, the virus can be titrated on the basis bovine, porcine, hamster, rabbit, and of induction of syncytia. Infection of mouse cell cultures inoculated with macaque monkeys is associated with the virus. A silent infection in humans, severe and often fatal immunosuppres- although antibodies are formed. sion. Some strains also induce retro- Produces tumors, mainly sarcomas on peritoneal fibromatosis, a proliferation injection into newborn hamsters, griv- of vascular fibrous tissue with some ets, baboons, and rhesus monkeys. Was similarities to Kaposi’s sarcoma seen in a contaminant of certain early batches human AIDS patients. of polio vaccine but caused no disease. Fine D and Schochetman G (1978) Cancer Has been used extensively for stud- Res 38 , 3123 ies of viral oncogenesis, but there is no van der Kuyl AC et al (1997) J Virol 71 , 3666 convincing evidence of a role for SV40 virus in any human cancers. simian vacuolating virus Synonym for Synonym : simian vacuolating virus. Simian virus 40 . Shah KV (2000) Rev Med Virol 10 , 31 simian varicella virus Synonym for Simian virus 41 (SV-41) Cercopithecine herpesvirus 9 . A species in the genus Rubulavirus. A virus isolated Simian virus 5 (SV-5) A species in the from primary cultures of monkey kid- genus Rubulavirus . A virus originally ney cells. Antibodies to SV41 are found isolated from primary cultures of mon- in the human population, and the virus key kidney cells which has been used appears to be closely related to human extensively as a model paramyxovirus. parainfluenza virus 2. Monkeys are probably not the natu- Tsurodome M et al (1990) Virology 179 , 738 ral host, and the close relationship to canine parainfluenza virus suggests simian viruses Viruses isolated from non- that SV5 may be a canine virus or a human primates. Some have been iso- subtype of human parainfluenza virus lated from excreta or diseased tissues 2. The genome encodes a small hydro- but most have appeared as cytopathic phobic (SH) protein, but this is not viruses in cultures of normal tissues. simian viruses 442

The large number of monkeys used relation between virus inoculum and to provide cell cultures has resulted infectivity is linear. Thus a single virion in many isolates which form a mixed suffices to infect a cell. group of DNA and RNA viruses. Two numbered series have been described: single radial hemolysis (SRH) A type simian agents (SA viruses) mainly from of hemagglutination inhibition test, African monkeys, and simian viruses in which the virus–red cell combina- (SV viruses) mainly from Asian mon- tion is immobilized in an agarose gel. keys. Differentiation into groups was Test sera are added to the wells cut in at first made according to the type of the gel and antibody diffuses radially. CPE produced, but most have now been With addition of complement, anti- assigned to various families. See Table bodies lyse the red cells, and the zone S3 . See also simian adenoviruses . Some of hemolysis has an area proportional SV and SA numbers are missing because to the amount of antibody. Widely they referred to isolates which were later used for diagnosis of anti-rubella virus found not to be viruses, or identical to antibodies. previous isolates, or have been lost. Kurtz JB et al (1980) J Hyg ( Camb ) 84 , 213 Hull RN (1968) Virol Monogr 2 , 124 Kalter SS et al (1995) Personal com- single-stranded DNA viruses There munication are two families of vertebrate viruses: Malherbe H and Herwin R (1963) S Afr Med Parvoviridae, in which the genome in J 37 , 407 most species is strand but in some there are and strands in differ- Simplexvirus A genus in the family ent viral particles (on extraction the Herpesviridae , subfamily Alpha- strands can form double strands); and herpesvirinae. Human herpesvirus 1 is the Circoviridae, where the single-stranded type species and the Human herpes- DNA genome is circular. There is also viruses 1 and 2 , Bovine herpesvirus 2 a free-standing genus, Anellovirus , and simian herpesvirus B are members. which contains the TT virus one of the Contains viruses of mammals which most ubiquitous species known. In show a considerable degree of serolog- plants there are three important fami- ical cross-reactivity (including neutrali- lies of ssDNA viruses, the Inoviridae , zation) and some genetic homology. Microviridae, and the Geminiviridae . Synonym : herpes simplex virus group. Sindbis virus (SINV) A species in the single-stranded RNA viruses The fami- genus Alphavirus. Probably a natural lies containing viruses which affect infection of birds but antibodies are vertebrates are: found in humans and domestic ungulates. May be associated with fever in Arenaviridae ambisense, 2 segments humans. Closely related to Whataroa virus and Western equine encephalo- Arteriviridae strand, unsegmented Astroviridae strand, unsegmented myelitis virus. Replicates in eggs kill- Bornaviridae strand, unsegmented ing the embryo, and in cell cultures of Bunyaviridae or ambisense, chick, human, and monkey tissues with 3 segments CPE. Experimentally lethal for suckling Caliciviridae strand, unsegmented mice. Causes myositis and encephalitis Coronaviridae strand, unsegmented in infant mice. Found in Egypt, South Filoviridae strand, unsegmented Flaviviridae strand, unsegmented Africa, India, Malaysia, the Philippines, and Australia. Nodaviridae strand, 2 segments Orthomyxoviridae strand, 6, 7, or 8 Griffin DE (2005) Curr Top Microbiol segments Immunol , 289, 57 Paramyxoviridae strand, unsegmented Picornaviridae strand, unsegmented Singapore grouper iridovirus A tentative Retroviridae strand, species in the genus Ranavirus . unsegmented, diploid Rhabdoviridae strand, unsegmented s i ngle hit relationships Virus infection Togaviridae strand, unsegmented is a single hit phenomenon, since the 443 single-stranded RNA viruses

Table S3. Simian viruses

SA series of viruses (simian agents): SA1 Foamy virus SA2 Produces CPE similar to SA1 but there are nuclear inclusions SA3 Reovirus type 1 SA4 Enterovirus. Isolated from the intestinal tract of Cercopithecus aethiops . Serologically related to SV4 and SV28. (Enterovirus S-16) SA5 Enterovirus. Isolated from the intestinal tract of Cercopithecus aethiops . (Enterovirus S-17) SA6 Cercopithecine herpesvirus 3 SA7 Adenovirus (Adenovirus S-16) SA8 Cercopithecine herpesvirus 2 SA9 Isolated from the mouth of a monkey. CPE resembles that produced by a reovirus but it does not agglutinate human O erythrocytes SA10 Parainfluenza virus type 3. Isolated from the mouth of a samango monkey, Cercopithecus mitis . Agglutinates human O, guinea pig and bovine erythrocytes (Simian virus 10) SA11 Rotavirus SA12 Vervet monkey virus SA13 Is now lost. CPE resembles that of measles virus SA14 Is now lost SA15 Is now lost. Herpesvirus isolated from baboons, Papio ursinus SA16 Is now lost. Isolated from vervet monkeys. Produced eosinophilic cytoplasmic inclusions and was difficult to passage SA17 Adenovirus. (Adenovirus S-17) SA18 Adenovirus. (Adenovirus S-18) SV series of viruses (simian viruses): SV1 Adenovirus. Type 1 of hemagglutination group 3. (Adenovirus S-1) SV2 Enterovirus. Isolated from intestinal tract of Macaca mulatta . (Enterovirus S-1) SV4 Enterovirus. Serologically related to SA4 and SV28 viruses SV5 Parainfluenza virus type 5. Antigenically identical to SA virus and DA virus. (Simian virus 5) SV6 Enterovirus. Isolated from intestinal tract of Macaca mulatta . (Enterovirus S-2) SV11 Adenovirus. Type 2 of hemagglutination group 3. (Adenovirus S-2) SV12 Reovirus type 1 SV13 Foamy virus SV15 Adenovirus. Type 3 of hemagglutination group 2. (Adenovirus S-3) SV16 Enterovirus. Isolated from intestinal tract of Macaca mulatta . (Enterovirus S-3) SV17 Adenovirus. Type 4 of hemagglutination group 2. Isolated from a monkey, Erythrocebus patas . (Adenovirus S-4) SV18 Enterovirus. Isolated from intestinal tract of Macaca mulatta . (Enterovirus S-4) SV19 Enterovirus. Isolated from intestinal tract of Macaca fascicularis . (Enterovirus S-5) SV20 Adenovirus. Type 5 of hemagglutination group 3. (Adenovirus S-5) SV21 Enterovirus. Identical to SV4 SV22 Proved not to be a new isolate SV23 Adenovirus. Type 6 of hemagglutination group 2. (Adenovirus S-6) SV24 An amoeba of the genus Acanthamoeba SV25 Adenovirus. Type 7 of hemagglutination group 3. (Adenovirus S-7) SV26 Enterovirus. Isolated from intestinal tract of Macaca mulatta . (Enterovirus S-6) SV27 Adenovirus. Similar or identical to SV31 SV28 Enterovirus. Isolated from normal kidney cell culture of Macaca mulatta . Serologically related to SA4 and SV4 viruses. (Enterovirus S-7) SV29 Proved not to be a new isolate SV30 Adenovirus. Type 8 of hemagglutination group 3. (Adenovirus S-8) SV31 Adenovirus. Type 9 of hemagglutination group 2. (Adenovirus S-9) SV32 Adenovirus. Type 10 of hemagglutination group 2. (Adenovirus S-10) SV33 Adenovirus. (Adenovirus S-11) SV34 Adenovirus. Type 12 of hemagglutination group 3. (Adenovirus S-12) SV35 Enterovirus. Isolated from intestinal tract of Macaca mulatta . (Enterovirus S-8) SV36 Adenovirus. Type 13 of hemagglutination group 1. (Adenovirus S-13) SV37 Adenovirus. Type 14 of hemagglutination group 2. (Adenovirus S-14) SV38 Adenovirus. Type 15 of hemagglutination group 3. (Adenovirus S-15) SV39 Identical to SV23 (continued) Sin Nombre virus (SNV) 444

Table S3 ( continued ) SV40 Polyomavirus species. See Simian virus 40 SV41 Similar to SV5 (Simian virus 41) SV42 Enterovirus. Isolated from intestinal tract of Macaca fascicularis . (Enterovirus S-9) SV43 Enterovirus. Isolated from intestinal tract of Macaca fascicularis . (Enterovirus S-10) SV44 Enterovirus. Isolated from intestinal tract of Macaca mulatta . (Enterovirus S-11) SV45 Enterovirus. Isolated from intestinal tract of Macaca fascicularis . (Enterovirus S-12) SV46 Enterovirus. Isolated from intestinal tract of Macaca sp . (Enterovirus S-13) SV47 Enterovirus. Isolated from intestinal tract of Macaca fascicularis .(Enterovirus S-14) SV48 Enterovirus SV49 Enterovirus. Isolated from intestinal tract of Macaca mulatta . (Enterovirus S-15) SV50–SV58 Probably isolated by Heberling, but not included in the final SV series. (Heberling RL and Cheerer FS (1965) Am J Epidemiol 81, 106) SV59 Reovirus type 3. Sent to Hull as agent 59 and numbered SV59 even before SV41 was reached.

Sin Nombre virus (SNV) A species in Elliott LH et al (1994) Am J Trop Med Hyg 51 , the genus Hantavirus. The principal 102 etiologic agent of hantavirus pulmo- Ksiazek TG et al (1995) Am J Trop Med Hyg nary syndrome, first recognized in 52 , 117 the Four Corners region of the USA Nichol ST et al (1993) Science 262 , 914 in June–August, 1993. Antigenically Zaki SR et al (1995) Am J Pathol 146 , 552 related to, but distinct from, other hantaviruses. Isolated from the deer Siniperca chuatsi rhabdovirus (SCRV) A mouse, Peromyscus maniculatus (grass- rhabdovirus isolated from mandarin land form), which is the natural res- fish (Siniperca chuatsi) in China. The ervoir species in North America. The complete genome sequence of the virus infection is spread to humans (who are has been determined. dead-end hosts and do not transmit Tao J-J et al (2008) Virus Res 132 , 86 the virus), through contact with urine or other excreta of chronically infected mice. Infection results in an acute pul- sinus histocytosis syndrome A rare dis- monary distress syndrome with a fatal- ease with massive lymphadenopathy ity rate of 50%. No specific treatment is (also known as Rosai-Dorfman dis- available. Control is by avoiding close ease), which might be linked to human contact with infected deer mice or herpesvirus 6 infection. their excreta. A Mid-western variant, Levine PH et al (1992) J Infect Dis 166 , 291 Blue River virus, has also been recognized in Indiana and Oklahoma, where SIRC cells (CCL 60) A heteroploid cell the host is the white-footed mouse, line derived from the cornea of a nor- Peromyscus leucopus . Another variant mal rabbit. strain, Monongahela virus, was identified in P maniculatus from the north- eastern USA. Numerous related viruses siRNA Small interfering RNA (some- are now also recognized in Canada, times known as silencing RNA). A class and Central and South America. of double-stranded RNA molecules Classification is by phylogenetic analy- 20–25 nt long which interfere with the sis, requiring evidence of at least 7% expression of a specific gene. Originally difference in amino acid sequence of discovered as a gene silencing mecha- the precursor glycoprotein and nucleo- nism in plants, and later found in capsid protein sequences to describe a mammalian cells. It has become clear new species. that siRNAs may have therapeutic Synonyms: Four Corners virus; Muerto applications. Canyon virus. Tuschl T et al (1999) Genes Dev 13 , 3191 445 skunk calicivirus (VESV/SCV)

SISPA (s equence-i ndependent, s ingle- belonging to the Chenuda complex of p rimer a mplification) A method for the Kemerovo serogroup. Isolated from non-selective cloning of sequences the tick, Argas cooleyi, in Texas and present in minute quantity that was Colorado, USA. Not reported to cause used in the discovery of the hepatitis E disease in humans. and hepatitis G virus genomes. cDNA molecules are blunt-ended and ligated Sjögren’s syndrome A complex disease to a 5 -staggered, 3 -blunt-ended of unknown etiology named after double-stranded oligonucleotide. A Swedish ophthalmologist HSC Sjögren single primer (one of the linker/primer which might be linked to human her- strands) is used for non-selective pes virus 4 (EBV) infection. Occurs amplification of all the cDNA molecules, mainly in middle-aged or older women which can then be expressed in vitro involving keratoconjunctivitis, dry and their protein products screened eyes, dry mouth, and connective tis- with a specific antiserum (immuno- sue disease (usually rheumatoid arthri- screening). Recently a method was tis). Both salivary and lacrimal glands developed that included pretreatment involved contain large numbers of of the sample with DNAse and this infiltrated B cells. was used to detect several new human viruses such as the human bocavirus Jones DT et al (1994) Invest Ophthalmol Vis and human polyomaviruses KI and Sci 35 , 3493 WU in human respiratory samples. Allander T et al (2001) Proc Natl Acad Sci 98 , SKIF (PKR inhibitory factor) A s pe- 11609 cific k inase i nhibitory f actor which Reyes GR and Kim JP (1991) Mol Cell Probes is induced in cells infected with vac- 5 , 473 cinia virus and inhibits the interferon- induced RNA-activated protein kinase Si SV See simian sarcoma virus . (PKR). site-specific mutagenesis With the avail- Akkaraju et al (1989) J Biol Chem 264 , 10321 ability of a DNA clone of the virus genome, it is possible to alter the Skinner Tank virus A novel arena- nucleotide sequence at specific sites by virus isolated from Mexican woodrats deletion or mutagenesis. The technique (Neotoma mexicana) isolated near Skinner is superior to random mutagenesis, Tank, Arizona, USA. Genetically related and has largely replaced it for genetic to Whitewater Arroya virus. Not known studies. to cause human disease. Sitiawan virus A probable species in the Cajimat MN et al (2008) Virus Res 133 , 211 genus Flavivirus, isolated from chicks at a broiler farm in Sitiawan, Perak, skin-heterogenizing virus of mice An Malaysia suffering from a disease unclassified virus present in many characterized by stretching the legs mouse tumors and capable of inducing and impaired mobility. A virus was strong transplantation antigens in the isolated in SPF chicken eggs which skin. Some leukemia viruses may have was enveloped, 41 nm in diameter, this property. with hemagglutinating activity against goose erythrocytes. It cross-reacted Salaman MH et al (1973) Transplantation 16 , with Japanese encephalitis virus by HA 583 tests, but not by neutralization. The Svet Moldavsky GJ et al (1970) J Natl Cancer genome sequence was 92% homolo- Inst 45 , 475 gous to Tembusu virus, a mosquito- borne flavivirus. skunk calicivirus (VESV/SCV) A strain of Vesicular exanthema of swine virus in Sixgun City virus (SCV) A serotype of the genus Vesivirus . A virus was iso- Chenuda virus in the genus Orbivirus , lated in human 293 cells. skunkpox virus (SKPV) 446 skunkpox virus (SKPV) A tentative spe- incubation period, particularly measles cies in the genus Orthopoxvirus . Isolated virus, JC polyomavirus, murine polio- from the North American striped encephalomyelopathy virus, rabies skunk, Mephitis mephitis . DNA avail- (depending on the infection route), and able for six of eight Old World viruses the transmissible spongiform encepha- and three New World viruses was used lopathy agents (prions). to detect amplicons from 510 to 1673 bp Narayan O (1992) Semin Virol 3 , 135 depending upon the species. SLAM protein S ignaling l ymphocytic smallpox virus Synonym for Variola virus . a ctivating m olecule. A cellular mem- Originally named to distinguish it from brane protein whose truncated form largepox (syphilis). lacks the transmembrane domain and can act as a self-ligand that has been small round adeno-associated viruses shown to induce the activation of See Adeno-associated virus . T lymphocytes. Acts as a cellular receptor for measles virus. small round structured viruses (SRSV) Yanagi Y (2001) Rev Med Viro l 11 , 149 A variety of agents about 30 nm in diameter associated with gastroenteri- slapped cheek disease A popular name tis; originally identified by electron for the infectious childhood rash called microscopy. Many are now classified as Erythema infectiosum , the major mani- astroviruses, caliciviruses, etc. festation of infection with B19 virus. An exanthematous rash illness that gives the child the appearance of hav- small round viruses See ‘Norwalk virus . ’ ing slapped cheeks. In 1905 Cheinisse gave it the name ‘ fifth disease ’ of six small secreted glycoprotein A nonstruc- erythematous rash diseases of child- tural glycoprotein of Ebola virus which hood which he described. Other names is generated by transcriptional editing. given to it include ‘ academy rash’ and It is a carboxyterminal truncated vari- ‘ Sticker’s disease. ’ ant of the secreted glycoprotein. Brown KE (1997) In Human Parvovirus B19 , Sanchez A et al (1996) Proc Natl Acad Sci USA edited by LJ Anderson and NS Young. Basel: 93 , 3602 Karger, p. 42 Smedi A syndrome (s tillbirths, m ummifi- SLE virus See St Louis encephalitis cation, e mbryonic death and i nfertility) virus . in pigs caused by porcine enterovirus. slippery sequence A sequence which Smedi enteroviruses are only clinically causes ribosomes to move with varying apparent when they infect the fetus, and frequency to an alternative open read- the pathological effects depend on the ing frame. In the case of astroviruses it stage of gestation. At early stages, there is a heptanucleotide with an adjacent may be embryonic death and return stem–loop structure that allows access to estrus. Fetuses infected between 40 to a –1 reading frame with an efficiency and 70 days die, and those infected of about 25%. It does not require a later may survive or are stillborn. The pseudoknot as in the case of retroviral viruses are recognized worldwide, and or coronaviral frameshifting. control involves encouraging infection of gilts before they reach sexual matu- Belew AT et al (2008) BMC Genomics 9 , 339 rity, with as many enterovirus serotypes Cao S and Chen SJ (2008) Phys Biol 5 , 16002 as possible. The Smedi syndrome can slow virus infections A poor term for also be induced by porcine parvovirus viruses causing slowly progressive infection, however. disease, originally coined to describe slow progressive retrovirus diseases smelt papilloma herpesvirus Causes of sheep in Iceland. They include the seasonal neoplasms of smelt, Osmerus lentiviruses but viruses of other fami- eperlanus , in Europe, taken from the lies may cause diseases after a long Elbe estuary. External growths appear 447 SnRNPs

on the head and fins. A herpesvirus showed several unique features, was identified by electron microscopy. including the use of an arginine tRNA primer binding site. Anders K and Möller H (1985) J Fish Dis 8 , 233 Hart D et al (1996) J Virol 70 , 3606 smelt reovirus (SRV) A strain of Snakehead virus (SHV) Aquareovirus A. Isolated in Canada A species in Novirhabdovirus from the viscera of diseased rainbow the genus , isolated smelt, Osmerus mordax . from internal organs of striped snake- heads, Ophicephalus striatus , with Marshall SH et al (1990) J Fish Dis 13 , 87 severe ulcerative disease and mortality. Other South-East Asian fish spe- smelt virus-1 (SmV-1) An unassigned cies, the swamp eel, Fluta alba , and the virus in the family Picornaviridae . Australian barramundi, Lates calcarifer , also seem to suffer from this infection. smelt virus-2 (SmV-2) An unassigned The virus is antigenically distinct from virus in the family Picornaviridae . other fish rhabdoviruses. Kasornchandra J et al (1992) Dis Aquat Org smoldering adult T-cell leukemia A clin- 13 , 89 ical subtype of adult T-cell leukemia caused by human T-cell leukemia virus Snotsiekte virus Synonym for alcelaphine 1 (HTLV-1) which has the best progno- herpesvirus 1 . sis with a mean survival time of more than 2 years. Characterized by skin and snow goose hepatitis B virus (SGHBV) lung infiltration of leukemic cells. A probable species in the genus Avihepadnavirus isolated from snow SMON virus See subacute myelo-optico- geese, Anser caerulescens . neuropathy virus . Chang SP et al (1999) Virology 262 , 39 smooth dogfish herpesvirus A virus Snow Mountain virus (SMV) identified by electron microscopy A distinct Norwalk virus of skin lesions of smooth dogfish, serotype of isolated from Mustelus canis . a water-borne outbreak of gastroenteritis in a resort camp in Colorado, USA. et al J Fish Dis 8 Leibovitz L (1985) , 273 Dolin R et al (1986) J Med Virol 19 , 11 King AD and Green KY (1997) Virus Genes smooth membrane The region of the 15 , 5 endoplasmic reticulum to which few or no ribosomes are attached. See rough Snowshoe hare virus (SSHV) A strain membrane . of California encephalitis virus in the genus Orthobunyavirus , belonging to snake adenovirus (SnAdV-1) A tenta- the California serogroup. Originally tive species in the genus Atadenovirus . isolated from the blood of an emaci- An adenovirus-like agent was isolated ated hare, Lepus americanus , caught from a moribund royal python, Python in Bitterroot Valley, Montana, USA. regius . Adenovirus-shaped particles Isolated from lemmings, hares and measuring 67–79 nm diameter were mosquitoes in various parts of northern seen in the nucleus. A similar isolate USA, Canada, and Alaska. Antibodies of an adenovirus-like agent was made found in humans and other animals. from a moribund corn snake, Elaphe Distinguished from other members of guttata . the group by CFT, HAI, or neutraliza- Juhasz A and Ahne W (1993) Arch Virol 130 , tion tests. Causes subclinical infection 429 in humans. Farkas SL et al (2008) Virus Res 132 , 132 Gentsch J and Bishop DHL (1976) J Virol 20 , 351 snakehead retrovirus (SnRV) A tentative species in the genus Epsilonretrovirus . SnRNPs S mall n uclear r ibo n ucleo p rotein s The complete sequence of this virus which participate in cellular and viral SnRNPs 448

RNA processing. Complexes of small Soldado virus (SOLV) A strain of Hughes nuclear RNA molecules, 100–300 nt in virus in the genus Nairovirus belong- length and specific nuclear proteins. ing to the Hughes virus serogroup. Isolated from a mixed pool of ticks of Snyder–Theilen feline sarcoma virus Ornithodoros sp infesting a common (STFeSV) A species in the genus noddy, Anous stolidus, on Soldado Gammaretrovirus . Isolated from a feline rocks in the Caribbean Sea. Also from fibrosarcoma and induces the tumor O capensis associated with sea birds in on inoculation into cats. Contains the Ethiopia, Seychelles, USA, and Senegal; transduced oncogene v-fes (expressing and from O maritimus in France, tyrosine kinase) which is also present Ireland, and North Wales, UK. Kills in the Gardner–Arnstein feline sarcoma suckling mice but not known to cause virus, and in the Hardy–Zuckerman disease in humans or other animals. feline sarcoma virus. Chastel C et al (1979) Arch Virol 60 , 153 sockeye salmon virus Synonym for Oregon sockeye disease virus, a strain soluble antigen An antigen which is of Infectious hematopoietic necrosis virus . virus-specific but not the virion itself, e.g. the nucleoprotein of influenza sodium dodecyl sulfate (SDS) A deter- virus. It often comprises isolated struc- gent used to break up virus particles or tural subunits but can also be nonstruc- infected cells. tural virus-coded proteins. Soehner–Dmochowski murine sarcoma soluble RNA RNA that is soluble in virus A strain of Moloney murine sarcoma strong salt solutions, e.g. 3 M sodium virus in the genus Gammaretrovirus . acetate. Consists mainly of small spe- Obtained from bone tumors induced in cies such as transfer RNA and ribos- New Zealand black rats by injection of omal 5S RNA. mouse sarcoma virus (Moloney strain). Ohtsuki Y et al (1978) Cancer Res 38 , 901 somatostatin A growth hormone- inhibiting hormone produced by soft-shelled turtle iridovirus (STIV) A neuroendocrine neurons. Greatly possible species in the genus Iridovirus . reduced in the acute phase of infection Isolated on a farm in Shenzhen, China with Borna disease virus. from soft-shelled turtles, Trionyx sinen- Lipkin WI et al (1988) Brain Res 475 , 366 sis , with ‘ redneck ’ disease. Chen ZX et al (1999) Virus Res 63 , 147 Somerville virus 4 A strain in the genus Orthoreovirus, belonging to the avian Sofyin virus (SOFV) The prototype orthoreovirus group. strain of Tick-borne encephalitis virus , Far Eastern subtype, in the genus Somone virus An unclassified arbovirus. Flavivirus. No known vector. An iso- Isolated from the tick, Ambylomma vari- late from a human patient with TBE in egatum , in Senegal. Also isolated from Far Eastern Russia in 1937. Differences Boophilus decoloratus in Nigeria. in sequence and pathogenicity have Institut Pasteur, Dakar (1986) Annual Report , been reported between Sofyin virus p. 9 and related Far Eastern TBE strains, Siberian TBE strains, and European sonication Use of high frequency sound TBE strains. wave energy to release virus from Bakhvalova VN et al (2000) Virus Res 70 , 1 associated cells for antigen preparation Pletnev AG et al (1990) Virology 174 , 250 or, at higher energy input, to inactivate virus infectivity. Sokoluk virus (SOKV) A strain of Hierholzer JC et al (1996) In Virology Methods Entebbe bat virus in the genus Flavivirus . Manual , edited by BWJ Mahy and HO Isolated from pipistrelle bats and vari- Kangro. London: Academic Press, p. 47 ous birds in Kirghizia. No known vector. Not reported to cause disease in sore mouth virus An infection of sheep. humans. A poor term because it has been used 449 spaceflight and viruses

as a synonym for both Bluetongue virus Sp-1 transcription factor A DNA-bind- and Orf virus . ing transcription factor that binds to JC virus DNA. Sorivudine (bromovinylarabinosyl- uracil: brovavir) A nucleoside analog Henson JW (1994) J Biol Chem 269 , 1046 that is an active inhibitor of human herpesvirus 3 (VZV). Unfortunately one Sp 104 virus A strain of Murine leukemia of its metabolites in vivo is bromovinyl- virus . A B-tropic virus, weakly onco- uracil which is a potent inhibitor of the genic but efficient at stimulating the liver enzyme dihydrothymine dehy- production of antinuclear antibodies in drogenase, responsible for degrading mice. Isolated from an established cell drugs such as 5-fluorouracil (5-FU) line derived from a plasmacytoma in a used in cancer chemotherapy. When mouse. This animal had been injected both 5-FU and sorivudine were used when newborn with a cell-free filtrate together, a number of deaths resulted, from the spleen of a dog with systemic so the compound will not be licensed lupus erythematosus. The virus shared in the USA. a cross-reacting antigen with the surface of blood lymphocytes of human Sororoca virus (SORV) A strain of or canine patients with systemic lupus Wyeomyia virus in the genus Orthobun- erythematosus. yavirus , belonging to the Bunyamwera serogroup. Isolated from mosquitoes Quimby FW et al (1978) Clin Immunol of Sabethini sp in Para, Brazil. Not Immunopathol 9 , 194 reported to cause disease in humans. SpAr 2317 virus (VRV) A strain of Sotkamo virus A strain of Puumula virus Tacaiuma virus in the genus Bunyavirus , isolated from the rodents, Clethrionomys belonging to the Anopheles A sero- glareolus and Clethrionomys rufocanus , in group. Isolated from mosquitoes, Finland. Anopheles sp. Not reported to cause Vapahlati O et al (1992) J Gen Virol 73 , 829 disease in humans. Now called Virgin River virus . Southampton virus (SHV) A serotype of Norwalk virus , in the genus Norovirus . Calisher CH et al (1980) Bull Pan Am Health Isolated from a family outbreak of Organ 14 , 386 gastroenteritis in Southampton, UK in 1991. Sp 1 K (NBL-10) cells (CCL 78) A heteroploid cell line derived from trypsinized Southern blotting Technique invented kidney tissue of the Atlantic spotted by Prof. EM Southern which involves dolphin, Stenella plagiodon . The line was transfer of denatured single-stranded developed to allow viral studies to be DNA fragments separated on an agar- made in the cells of a marine mammal. ose gel to a nitrocellulose filter which is then analyzed by hybridization to radioactive or biotinylated DNA or spaceflight and viruses The stress of RNA probes. Later forms of blotting being launched into space followed for RNA and protein were named by microgravity can cause reactiva- Northern and Western, respectively, tion and shedding of latent herpesvi- to distinguish them from the original ruses such as EBV, CMV and VZV. 71 technique. astronauts studied by PCR showed increased shedding of CMV in their South River virus (SORV) A strain urine before spaceflight compared to of California encephalitis virus in the ground-based controls. After return to genus Orthobunyavirus, belonging to earth the astronauts had up to eight the California serogroup. A variety of fold higher antibody titers to CMV than Jamestown Canyon virus. Isolated from controls, suggesting that CMV replica- mosquitoes, Anopheles crucians . Not tion occurred during spaceflight. reported to cause disease in humans. Mehta SK et al (2000) J Infect Dis 182 , 1761 Sudia WD et al (1971) Mosq News 31 , 576 Cohrs RJ et al (2008) J Med Virol 80 , 1116 Sparrowpox virus (SRPV) 450

Sparrowpox virus (SRPV) A species in sphenicid herpesvirus 1 (SpHV-1) An the genus Avipoxvirus . unassigned virus in the family Herpes- Giddens WE et al (1971) Vet Pathol 8 , 260 viridae. Isolated from a black-footed penguin, Spheniscus dermersus . Synonym: black-footed penguin species A virus species is defined by the herpesvirus. International Committee on Taxonomy of Viruses (ICTV) as a polythetic class Kincaid AL (1988) J Wildl Dis 24 , 173 of viruses that constitutes a replicating lineage and occupies a particular spider monkey herpesvirus Synonym ecological niche. Members of a poly- for Ateline herpesvirus 1 a species in the thetic class are defined by more than genus Simplexvirus . one property and no single property is essential or necessary. Does not depend spike A projection from the surface of a on a unique characteristic. virus particle, usually associated with binding of the particle to the cell sur- van Regenmortel MHV (1990) Intervirology face. Also known as S protein. See 31 , 241 peplomers . species barrier Host properties that limit spiking disease Turkey enteritis caused the ability of most viruses to replicate by turkey coronavirus. in more than one species. Many viruses display increased pathogenicity in a spleen focus-forming virus A strain of new species, but few of the factors that Murine leukemia virus which causes limit the ability of viruses to cross spe- countable foci of tumor cells in the cies barriers are well understood. spleen. Used as a name for the compo- Mahy BWJ and Brown CC (2000) Rev Sci nent of Friend murine leukemia virus Tech 19 , 33 which is defective and which requires Mahy BWJ and Murphy FA (2005) In lymphoid leukemia virus as a helper. Topley & Wilson’s Microbiology and Microbial The proliferative effect of the virus Infections , vol. 2, Tenth edition, edited by appears to be mediated by binding of BWJ Mahy and V ter Meulen. London: the virus glycoprotein (gp55) to EpoR, Hodder Arnold, p. 1646 the erythropoietin receptor on the cell membrane. species jumping The ability of viruses to Kabat D (1989) Curr Top Microbiol Immunol cross the species barrier as a result of 148 , 1 mutations. Li J-P et al (1990) Nature 343 , 762 Mahy BWJ and Brown CC (2000) Rev Sci Tech 19 , 33 spleen necrosis virus See Trager duck spleen necrosis virus . spectacled caiman poxvirus (SPV) spliceosome A multicomponent ribo- An unassigned virus in the family nucleoprotein complex that carries out Poxviridae . the splicing of RNA. Jacobson ER et al (1979) J Am Vet Med Assoc Harvey Lect 90 175 , 937 Guthrie C (1994) , 59 splice sites The sites at which the intron SPH114202 virus A strain of Sabiá virus is excised. The 5 -splice site is adjacent in the genus Arenavirus. Isolated in São to the GU end of the donor junction. Paulo State, Brazil, in 1990 from a fatal The 3 - splice site is adjacent to the AG case of Brazilian hemorrhagic fever. end of the acceptor junction. Caused a laboratory infection, apparently acquired by aerosol transmission. splicing Ligation of non-contiguous The presumed rodent host of the virus cleaved portions of an RNA molecule is unknown. to remove introns from precursor RNA Vasconcelos PFC et al (1993) Rev Inst Med transcripts and produce functional Trop , São Paulo 35 , 521 mRNA. Originally discovered from 451 Spumavirinae

work on adenovirus mRNA synthesis. spongothymidine (Ara T) 1- β -d -Arabino Now known to be a common feature of furanosylthymidine. A nucleoside anti- mRNA production in eukaryotic cells. viral agent isolated from the sponge, Functional mRNA in eukaryotic cells Cryptotethia crypta. A selective inhibitor can be made up of spliced transcripts of human herpesviruses 1 and 2. originating from widely separated regions of the DNA. The sequences spongouridine (Ara U) 1- β - d - of DNA, transcripts of which are Arabinofuranosyluracil. A nucleo- present in mRNA, are termed ‘exons ’ side antiviral agent isolated from the (expressed regions); the intervening sponge, Cryptotethia crypta . DNA sequences, transcripts of which are removed by splicing, are termed spontaneous mutant A virus with a ‘ introns. ’ Plays an important role in mutation that arises naturally during the expression of virus genomes whose virus replication. replication involves the cell nucleus, such as influenza virus, retroviruses, sporadic Creutzfeldt-Jakob disease and several DNA viruses. (sCJD) The most common form of Darnell JE (1978) Science 202 , 1257 prion disease in humans, though the Sharp PA (1994) Cell 77 , 805 exact cause is unknown. It appears to result from spontaneous conversion of c split vaccines Vaccines prepared from cellular prion protein PrP to the pro- sc disrupted virus particles and purified tease resistant isoform PrP . to remove the toxic fraction of viral proteins which may cause side-effects. sporadic infections Virus infections that This permits the injection of a larger occur at low frequency and in a seem- dose of the useful antigen. ingly unconnected way.

Spondweni virus (SPOV) A strain of Zika spring viremia of carp virus (SVCV) virus in the genus Flavivirus . Isolated A tentative species in the genus from culicine mosquitoes in South Vesiculovirus. Causes disease and death Africa, Nigeria, Mozambique, and in fish farms in the USA and Europe. Cameroon. Can cause a febrile illness Primarily a pathogen of carp but can with hepatitis in humans. Antibodies infect pike. Replicates in FHM cells, present in cattle, sheep, and goats, in most rapidly at 20–22°C, but can rep- which it causes disease. licate at 31°C. Antigenically related to swim-bladder inflammation virus. The spongiform encephalopathy agents disease erupts in the spring, and both Prions causing scrapie, kuru, bovine adults and young fish are killed by the spongiform encephalopathy, trans- infection which results in hemorrhage missible mink encephalopathy, of the internal organs, including the Creutzfeldt–Jakob disease, and related swim-bladder. Lethargy, loss of bal- neurological disorders with a common ance, and aimless swimming are clini- pathological picture and progressive cal signs of the disease. course to death. There is spongiform Synonyms: infectious dropsy of carp degeneration of the brain but absence virus; rhabdovirus carpio. of inflammatory reaction. They are Bucke D and Finlay J (1979) Vet Rec 104 , 69 caused by prions, self-replicating cel- Roy P and Clewley JP (1978) J Virol 25 , 912 lular protein isoforms that are not true viruses, as they contain no detectable Spumaretrovirinae A subfamily of the nucleic acid. See prion . family Retroviridae containing a single Synonym : subacute spongiform enceph- genus, Spumavirus . alopathy viruses. Prusiner SB (Editor) (1999) Prion Biology and Spumavirinae An old name for a sub- Diseases. Cold Spring Harb Lab Monogr 38 , family in the family Retroviridae that New York: Cold Spring Harbor Laboratory no longer exists. It is now named Press Spumaretrovirinae . Spumavirus 452

Spumavirus (Latin: spuma foam) A Squirrel monkey retrovirus (SMRV) A genus of the family Retroviridae . Usually species in the genus Betaretrovirus . An cause persistent but silent infections endogenous xenotropic virus of squir- in their natural host. Only exogenous rel monkeys, Saimiri sciureus. Similar to species have been detected, and no Mason–Pfizer virus, although virions diseases have been associated with have a central electron-dense nucle- spumavirus infections. Viruses have a oid while the Mason–Pfizer virus has widespread distribution and are found a bipolar tubular nucleoid. Isolated in many mammals. No oncogenes have from lung cells by co-cultivation with been detected in the genome. Often canine cells. Infectious for cells of found in primary tissue cultures, espe- human, mink, mouse, dog, bat, chim- cially on prolonged passage. In cell cul- panzee, rabbit, and rhesus monkey, but tures syncytium formation is induced not marmoset, owl monkey, baboon, and the cells develop a foamy appear- or howler monkey cells. Buds with an ance. There are six recognized species: intact nucleoid through the cell mem- African green monkey simian foamy virus , brane and has a reverse transcriptase Bovine foamy virus , Equine foamy virus , with a magnesium cation preference. Feline foamy virus , Macaque simian foamy Viral RNA hybridized with the DNA virus, and Simian foamy virus . Several from all the squirrel monkey tissues serotypes have been isolated from tested, but not with DNA from other monkeys of various species. No natu- New and Old World monkeys or apes. ral human infections are known, and Not immunologically related to onco- the former human isolate is a simian viruses of baboon, woolly monkey, virus. Viruses of simian, bovine, and rhesus monkey, cat, cattle, horse, rat, feline origin do not cross-react in neu- hamster, or mouse. tralization tests. Virions, 100–140 nm in Fine D and Schochetman G (1978) Cancer diameter, have a distinctive morphol- Res 38 , 3123 ogy with an electron-lucent nucleoid Sommerfelt MA and Hunter E (1999) In core and envelope with prominent sur- Encyclopedia of Virology , Second edition, face projections. The genome is 11 kb edited by A Granoff and RG Webster. in length (one monomer) and the LTR London: Academic Press, p. 1518 is 1150 nt long (V3,800-R,200-U5,150). Maturation by budding. Inactivated Squirrel parapoxvirus (SPPV) A species by lipid solvents and pH 3. Replicate in the genus Parapoxvirus . Implicated slowly in a wide range of dividing in the decline of the red squirrel in cells. See Simian foamy virus . the UK. Causes severe skin lesions in the squirrels. Typical parapoxvirus Loh PC (1993) In The Retroviridae, vol. 2, morphology. edited by JA Levy. New York: Plenum Press, p. 361 Thomas K et al (2003) J Gen Virol 84 , 3337 Rethwilm A (2005) In Topley & Wilson’s Microbiology and Microbial Infections , vol. 2, SR-11 virus (SR11V) A strain of Seoul Tenth edition, edited by BWJ Mahy and virus in the genus Hantavirus , isolated V ter Meulen. London: Hodder Arnold, from rats, Rattus norvegicus, in Japan. p. 1304 Member of the Seoul virus antigenic group. Squirrel fibroma virus (SQFV) A species in the genus Leporipoxvirus , serologi- Src gene The oncogene of Rous sar- cally related to rabbit fibroma virus. coma virus, inserted at the 3 - end of Causes multiple fibromas in grey the genome, which encodes a tyrosine squirrels, Sciurus carolinensis , in North kinase (pp60), anchored to the inner America. Produces fibromas in domes- surface of the plasma membrane by tic rabbits but cannot be passaged in a myristilated N-terminus. The v-src them. protein differs from the cellular protein Synonym : fibroma virus of squirrels. (c-src) by several mutations, which result in an activated form of the enzyme squirrel monkey herpesvirus Synonym in transformed cells, with increased for herpesvirus saimiri 2. amounts of phosphotyrosine-containing 453 stellavirus

proteins that presumably contribute to is sometimes used. See also Kozak the transformed state. The crystal struc- sequence . ture of Src has been obtained. Synonym : initiation codon. Brugge JS et al (1979) J Virol 29 , 1196 Superti-Furga G and Gonfloni S (1997) STAT S ignal transducer and a ctivator BioEssays 19 , 447 of transcription. Induced in cells by interferon. Bind to DNA and cause Sripur virus (SRIV) An unassigned ver- transcription of interferon-stimulated tebrate rhabdovirus, isolated from genes. Sergentomyia sp in India in 1973. Not reported to cause disease in humans. statolon A fermentation product of the fungus, Penicillium stoloniferum , and a SSV See simian sarcoma virus . potent interferon inducer. This activity is due to the presence of a double-stranded St Abbs Head virus (SAHV) A RNA viral genome. Electron microscopy strain of Uukuniemi virus in the studies have demonstrated the presence genus Phlebovirus, belonging to the of numerous particles of typical virus Uukuniemi serogroup. Isolated from morphology, about 30 nm in diameter. engorged female ticks, Ixodes uriae , col- They are reported to be serologically lected from a sea bird colony at St Abbs unrelated to helenine particles. Head, Scotland, UK. Kleinschmidt WI et al (1968) Nature 220 , 167 Watret GE and Elliott RM (1985) J Gen Virol 66 , 1001 stavudine 2 ,3 -Didehydro-2 -deoxythymidine (D4T). A potent inhibitor of St Abbs Head virus (SAHV) A sero- HIV-1 reverse transcriptase in vitro . type of Great Island virus in the genus Similar in its action to AZT. Orbivirus . Isolated from ticks (Ixodes uriae ) collected from a seabird colony Hitchcock MJM (1991) Antivir Chem at St Abb’s Head, Scotland, UK. Chemother 2 , 125 Riddler SA et al (1995) Antiviral Res 27 , 189 Eley SM et al (1985) Arch Virol 85 , 47 steady-state infection staging system A standardized system Infection in a cell for describing Kaposi’s sarcoma, which culture where both virus and cell multi- occurs in several different clinical plication proceed. Most or all of the presentations. Developed by the AIDS cells are infected, virus is released con- Clinical Trial Group. tinuously from the cells, but there is no CPE. The infection cannot be cured by Krown SE (2001) Curr Opin Oncol 13 , 374 adding antiviral antibody to the culture medium. Superinfection with another standard virus Term introduced by von virus is possible but there may be com- Magnus to describe complete virus as plementation or interference. Examples opposed to defective virus. are numerous among viruses which mature by budding: Paramyxoviridae , Starlingpox virus (SLPV) A species in the Rhabdoviridae , Togaviridae, and genus Avipoxvirus . Retroviridae . See carrier cultures . Landolt M and Kocan RM (1976) J Wildl Dis Hotchin J (1974) Prog Med Virol 18 , 81 12 , 353 start codon The trinucleotide in a mRNA steelhead picornavirus A virus iso- at which ribosomes start the process of lated in Washington State, USA translation and which sets the reading from the ovarian fluid of steelhead, frame for the translation. In eukaryotes Oncorhynchus mykiss. Grows in a vari- it is AUG which is decoded as methio- ety of fish cell lines. No known associa- nine. In prokaryotes AUG (giving tion with disease. N -formylmethionine) is the most common start codon but GUG (valine) stellavirus Synonym for rotavirus. stickleback virus (SBV) 454 stickleback virus (SBV) A strain of Frog virus, although other viruses may cause virus 3 in the genus Ranavirus . Isolated the same clinical picture. from the three-spine stickleback, Gasterostelus aculeatus . An apparently stomatitis–pneumoenteritis complex identical virus was isolated from an virus Synonym for Peste des petits rumi- amphibian, the redlegged frog, Rana nants virus . aurora , suggesting that fish iridoviruses can also infect amphibia. stop codon The trinucleotide sequence Mao J et al (1999) Virus Res 63 , 45 at which protein synthesis is terminated. There are three stop codons: sticky ends The single-stranded ends UAA (Ochre), UAG (Amber), and UGA on DNA produced by many type II (Opal). There is evidence that UGA restriction endonucleases. They may be may signal the insertion of seleno- either 5 or 3 extensions of the DNA cysteine in some contexts, especially in molecule. HIV replication. See suppressor . Synonym : nonsense codon. stimulon A putative factor produced in Brenner S et al (1961) Nature 190 , 576 adenovirus-infected cell cultures which Taylor EW (1994) J Med Chem 37 , 2637 increases the replication of latent rat virus and H-1 virus. Can be demon- stork hepatitis B virus A strain of strated in virus-free extracts of infected Heron hepatitis B virus in the genus cells but not in extracts of uninfected Avihepadnavirus . Found in white storks cells. Stimulon is inactivated by in Germany. trypsin, but not by DNase or RNase. Pult I et al (2001) Virology 289 , 114 Brailovsky C and Chany C (1965) CR Acad Sci Paris 260 , 2634; 261 , 4282 Stoxil Trade name for idoxuridine eye St Louis encephalitis virus (SLEV) A drops. species in the genus Flavivirus , belonging to the Japanese encephalitis virus strain An isolate of a virus which resem- serogroup. The wild host is birds. bles the type virus in the major proper- Transmission is by mosquitoes, Culex ties that define the type, but differs in sp. Occurs in Canada, eastern and west- minor properties such as vector species ern USA, Central and South America, specificity, symptoms induced, serolog- sometimes causing serious outbreaks ical and genetic properties. In practice, of encephalitis. In humans many infec- it is often very difficult to differenti- tions result in brief febrile illness but ate the boundaries between variants, encephalitis may occur, especially strains, and species of viruses. In the in urban outbreaks in eastern USA, past antigenic cross-reactivity (sero- which reappear about every 10 years. typing) was the main means of distin- Sequelae are uncommon. Disease in guishing closely related viruses. With horses not reported. Injection i.c. into the advent of PCR and sequence analy- certain strains of mice causes encepha- sis, relationships between viruses as litis. Virus can be propagated in eggs. determined by genome sequence com- Causes diffuse edematous lesions with parison is considered the most appro- proliferative and necrotic elements on priate way to decide upon species and the CAM. Replication occurs in cell strains. See serotype . cultures of chick, mouse, and other species with CPE. No effective vaccine strand polarity The organization of available for humans. the nucleotide sequence in a single- Synonym : Saint Louis encephalitis virus . stranded nucleic acid. Positive ( ) Monath TP and Tsai TF (1987) Am J Trop Med polarity refers to single-stranded mol- Hyg 37 , 40s ecules that contain the same sequence as mRNA. Negative ( ) polarity mol- stomatitis papulosa of cattle virus ecules have sequence complementary Synonym for Bovine papular stomatitis to the ( )-sense strand. 455 subacute myelo-optico-neuropathy virus (SMON)

Stratford virus (STRV) A strain of turbot (Scophthalmus maximus ), smelt Kokobera virus in the genus Flavivirus , (Osmerus mordax ), and Atlantic salmon belonging to the Japanese encephalitis (Salmo salar ). The virus genome con- virus group. Isolated from Aedes vigilax sists of 11 segments of double-stranded in Queensland, Australia. Not reported RNA. to cause disease in humans or other animals. Striped jack nervous necrosis virus A species in the genus Betanodavirus isolated street virus A virulent wild strain. A term from striped jack, Pseudocaranx dentex . applied to rabies virus isolated from animals. After attenuation through pas- Mori KI et al (1992) Virology 187 , 368 sage in rabbits, the virus is called ‘ fixed virus. ’ striposomes See uncoating . streptavidin A protein which binds to Strongyloplasma hominis An old name biotin. When coupled to an enzyme for the molluscum body produced in such as horseradish peroxidase, it will epidermal cells of patients infected detect biotin in biotinylated DNA so with Molluscum contagiosum virus . An providing a nonradioactive detection inclusion body. technique for specific sequences. Hiller Y (1987) Biochem J 248 , 167 S-tropic viruses See xenotropic virus . streptovitacin A A glutarimide antibiotic. structural protein A protein which forms A potent reversible inhibitor of protein part of the structure of a virus particle. synthesis. stump-tailed macaque virus See Bovine stress proteins A small group of cel- polyomavirus . lular proteins that are synthesized in response to metabolic stress including stutter site The site at which paramyxo- heat-shock. Viral infection often sup- viruses insert guanylate residues during presses the synthesis of host cell pro- RNA transcriptional editing. Believed teins, but the stress protein genes, to occur through the polymerase ‘stut- along with the genes encoding inter- tering,’ resulting in pseudo-templated feron proteins, are generally activated transcription. The addition of gua- by virus infection. nylate residues to the mRNA gives Jindal S and Malkovsky M (1994) Trends access to additional reading frames Microbiol 2 , 89 within the P gene. strigid herpesvirus 1 (StHV-1) An unas- Hausmann S et al (1999) J Virol 73 , 5568 signed virus in the family Herpesviridae . Isolated from owls of several species subacute myelo-optico-neuropathy virus in which it may cause hepatosplenitis (SMON) A herpesvirus isolated from and paralysis. Disease can be produced the feces and CSF of patients with experimentally in owls but not other subacute myelo-optico-neuropathy. birds. Replicates on the CAM and in Seen mainly in Japan, the disease is the allantois. characterized by sensory disturbance, Synonym: owl hepatosplenitis especially of the lower part of the legs, herpesvirus. abdominal symptoms, decreased mus- Burtscher H and Sibalin M (1975) J Wildl Dis cle strength, and bilateral impairment 11 , 164 of visual acuity. There are no changes in the blood or CSF. There is degen- striped bass reovirus (SBRV) A strain eration of posterior and lateral tracts of Aquareovirus A in the genus of the spinal cord. The virus was iso- Aquareovirus. Isolated from striped lated in BAT-6 cells and causes a thin- bass, Morone saxatilis, with hemorrhagic ning of the cell sheet. On injection into lesions in Chesapeake Bay, Maryland, newborn C57Bl/6 mice it is reported USA. Viruses were also isolated from to cause paralysis of the hind legs. It is subacute myelo-optico-neuropathy virus (SMON) 456

claimed that the virus can be derived sub-families The family is the bedrock on passage of avian infectious laryngo- of universal virus taxonomy, but in tracheitis virus on the CAM or in new- some instances subfamilies have been born C57BL/6 mice. It is antigenically introduced to accommodate new related to this virus but is said to dif- knowledge of the relationships of gen- fer from it in being non-pathogenic era within families. Subfamilies are for fowls, less unstable at low pH and indicated by the suffix -virinae , e.g. pathogenic for C57BL/6 mice. The role Alphaherpesvirinae . of the virus in subacute myelo-optico- neuropathy has been questioned and subgenomic RNA A species of RNA it is suggested that the disease is due of less than genome length found in to the administration of clioquinol, RNA virus infected cells. The genome an antidiarrheal drug. When the use RNA of several groups of positive- of clioquinol was stopped in Japan strand RNA viruses (e.g. Togaviridae , in September 1970, incidence of the Retroviridae, and Caliciviridae ) con- disease fell dramatically. However, tains more than one initiation site, but clioquinol is used outside Japan and early in infection only the portion of appears to cause little disease. the RNA coding for RNA-replicating Synonym : Inoue–Melnick virus. enzymes is translated. Subsequently, Inoue YK (1975) Prog Med Virol 21 , 35 subgenomic RNA containing the pre- Kono R (1975) Lancet ii , 370 viously masked initiation site is synthesized and can then be translated subacute sclerosing panencephalitis into virus coat proteins. In togaviruses, (SSPE) Measles virus is the cause this RNA has been termed ‘interjacent of SSPE, a slow virus disease of the RNA.’ The several monocistronic spe- brain. Vaccinated children are much cies of mRNA synthesized by tran- less likely to develop SSPE than the scription from full-length genome RNA unvaccinated. The disease is more in cells infected with some negative- common in patients who have measles strand viruses, such as Paramyxoviridae before the age of 2 years. The majority and Rhabdoviridae , may also be termed of cases appear 6–8 years after acute subgenomic RNA. See interjacent measles, with an incidence in unvac- RNA . cinated children of 1 in 1 million cases. More likely to occur in boys than girls. submaxillary virus See Cytomegalovirus . Measles virus can be isolated from the brain in cases of SSPE but only by subunit reassortment The process by co-cultivation of brain cells with sus- which all segmented RNA viruses ceptible target cells. It seems that host of vertebrates exchange genetic cell-dependent attenuation of measles information. virus gene expression and function occurs which determines persistence subunit vaccine A vaccine composed of in the brain. Some SSPE strains of a purified antigenic determinant that is measles virus are neurovirulent in separated from the virulent virus. ferrets and these strains are strongly cell-associated. suckling mouse cataract virus Not Baczko K et al (1993) Virology 197 , 188 a virus. An unusual type of myco- Schneider-Schaulies J et al (2005) In Topley & plasma isolated from rabbit ticks, Wilson’s Microbiology and Microbial Infections , vol.2, Tenth edition, edited by BWJ Mahy Haemaphysalis leporis-palustris , in and V ter Meulen. London: Hodder Arnold, Georgia, USA. Replicates to high titer p. 1403 in chick embryos. Causes cataracts in Schneider-Schaulies S et al (1994) Semin Virol suckling mice after 20 days, sometimes 5 , 273 with signs of neurological involvement and stunting of growth. Passes through subacute spongiform encephalopathy filters of APD 220 nm but not at APD viruses Synonym for spongiform 100 nm. Electron microscopy reveals encephalopathy viruses. mycoplasma-like bodies. 457 supercoiled DNA

Bastardo JW et al (1974) Infect Immun 9 , 444 suid herpesvirus 2 (SuHV-2) An unas- Clark HF (1974) Prog Med Virol 18 , 307 signed species in the family Herpes- viridae . Causes rhinitis and destruction Sudan Ebola virus (SEBOV) A species in of the turbinates, with distortion of the the genus Ebolavirus, which caused a snout, epistaxis, and sneezing, notably major outbreak of Ebola hemorrhagic in 2- week-old piglets, when death is fever in Sudan in 1976. See Ebolavirus . common. Transmission is possible in piglets but not in adult pigs. Disease occurs in outbreaks and inclusions are Sudan Ebola virus Boniface An isolate of present in the cells of many organs. Sudan Ebola virus from a patient named Can be cultivated in primary pig cell Boniface. cultures, replicating better in epithelial than in fibroblastic cells. Sudan Ebola virus Maleo An isolate of Synonyms: inclusion-body rhinitis Sudan Ebola virus from a patient named virus; swine cytomegalovirus. Maleo. Edington N et al (1976) J Hyg ( Camb ) 77 , 283

Suid herpesvirus 1 (SuHV-1) A species in Suipoxvirus A genus in the subfamily the genus Varicellovirus. A natural infec- Chordopoxvirinae containing the Swinepox virus. Virions are brick- tion, mainly of pigs, but cattle, horses, sheep, dogs, cats, foxes, and mink are shaped, about 300 250 200 nm. also susceptible. Endemic in pig popu- DNA 175 kb in length with inverted ter- lations throughout the world. In pigs minal repeats of 5 kb.Virus forms foci the infection is usually silent, but in in pig kidney cell cultures and plaques 5–10% the virus infects the tonsils from in swine testes cell cultures. The only which it spreads to the CNS. There are species in the genus is Swinepox virus . nervous symptoms and fever but the pigs recover. Causes abortion in up to suit laboratory A biosafety level 4 labora- 50% of pregnant sows. In cattle, sheep, tory which isolates the workers from and carnivores the disease is usually the laboratory environment by means fatal, with intense pruritis (known of fully encapsulating suits which are as ‘ mad itch’ ). There are reports of supplied by air from outside the labo- infection in laboratory workers who ratory. Used for work with viruses for developed aphthae of the mouth and which there are no known protective local pruritis. Rabbits, guinea pigs, vaccines or therapeutic agents, such and many other species are suscepti- as Ebola, Marburg, and Lassa fever ble experimentally. Monkeys infected viruses. intranasally develop ataxia, salivation and have convulsions, but there is no sulfation A post-translational modifica- pruritis. Virus replicates on the CAM tion of some virus proteins, the func- with plaque production and CPE in tion of which is not well understood. cultures of chick, rabbit, guinea pig, and dog cells. All strains appear anti- Sunday Canyon virus (SCAV) An unas- genically similar. The virus has pro- signed virus in the family Bunyaviridae . vided a valuable model for studies of Isolated from the tick1, Argas cooleyi , the spread of viruses through the nerv- collected in south-western USA in ous system. areas frequented by cliff swallows, Synonyms : herpesvirus suis; Aujeszky’s Petrochelidon pyrrhonota. Sensitive to disease virus; infectious bulbar paral- ether and to low pH. Pathogenic for ysis virus; mad itch virus; pig her- suckling mice. Not reported to cause pesvirus 1; porcine herpesvirus 1, disease in humans. pseudorabies virus. Yunker CE et al (1977) Acta Virol , Prague 21 , 36 Enquist LW et al (1999) Adv Virus Res 51 , 237 Pensaert MB and Kluge JP (1989) In Virus Infections of Porcines , edited by MB Pensaert. supercoiled DNA A conformation that New York: Elsevier a double-stranded DNA molecule can supercoiled DNA 458

adopt. When both strands of a double- of Brazil. Antigenically related to stranded molecule are covalently Changuinola virus. Not associated with closed, one of the strands becomes disease in humans. over- or under-wound in relation to the other. The torsional strain causes Sustiva An alternative name for the molecule to coil into a characteris- Efavirenz , an inhibitor of HIV reverse tic shape. transcriptase. supergroup The picornavirus supergroup SV virus series See Simian viruses SV includes the family Picornaviridae plus series . the three families of plant viruses which share common features of organ- SV5 See Simian virus 5 . ization, the Comoviridae , Potyviridae , and Sequiviridae . SV40-PML virus See JC polyomavirus . superhelical DNA See supercoiled SV40 virus See Simian virus 40 . DNA . Svedberg units See sedimentation superinfection exclusion The inability of coefficient . two related viruses to replicate in the same cell. The presence of one repli- SV41 virus See Simian virus 41 . cating virus prevents the second virus SW-13 cells (CCL 105) Initiated from the from replicating. May be the result of biopsy tissue of a small-cell carcinoma entry exclusion because the cell recep- originating in the adrenal cortex of a tors are blocked, although other fac- 55-year-old Caucasian female. tors such as immunity may also play a role. swamp fever virus Synonym for Equine infectious anemia virus . supernatant The liquid above sedimented material or above a precipitate. swan circovirus A novel circovirus Cygnus olor supplemental essential genes A term detected in mute swans ( ) used to describe 37 genes in the her- found dead in Germany in 2006. The pes simplex virus genome which are genome sequence was most closely not needed for virus replication in related to goose circovirus. The clini- cell culture, but may be required for cal significance of the infection is other virus properties such as neuro- unknown. invasiveness. Halami MY et al (2008) Virus Res 132 , 208 suppressor A gene that can partially or Sweetwater Branch virus (SWBV) An completely reverse the effect of a muta- unassigned vertebrate rhabdovirus, tion in another gene. antigenically related to lyssaviruses. Isolated from Culicoides insignis in suppressor tRNA A tRNA molecule, Florida, USA in 1982. Not known to produced by a suppressor gene, that cause disease in humans. can pair with a nonsense (termination) codon so that the correct amino acid Gibbs EPJ et al (1989) Vet Microbiol 19 , 141 is incorporated into the polypeptide chain. swim-bladder inflammation virus A possible species in the genus Vesiculovirus . surfactant proteins-A and D Collectins, Causes a severe and fatal disease in soluble proteins in lung that act as a carp. Experimental infection of carp, first line of defense against virus infec- Cyprinus carpio and Carassius auratus , tion. See collectins. caused reduced reflex activity and in some cases loss of balance, swelling Surubim virus A possible species in the of the anus and abdomen, petechiae genus Orbivirus, isolated from phle- on the skin and muscles. Death occurs botomine flies in the Amazon region 4–8 days after signs of infection appear. 459 syncytial viruses

Antigenically very similar to infec- infection by rotaviruses of many spe- tious hematopoietic necrosis virus. cies, but only the calf and pig rotavi- Replicates in FHM cells at an optimal ruses cause disease. temperature of 20–22°C. Bachmann PA and Ahne W (1974) Arch swine vesicular disease virus (SVDV) Gesamte Virusforsch 44 , 261 A porcine variant of human coxsackie virus B5, a serotype of human entero- swine calicivirus (SwV-43) A tentative virus B in the genus Enterovirus . Swine species in the genus Norovirus . vesicular disease was first observed in Italy in 1966. Subsequent outbreaks swine cytomegalovirus Synonym for occurred in Hong Kong in 1971, Europe suid herpesvirus 2. and Japan in 1972–1975. There are several antigenically different strains of the swine fever virus Synonym for hog virus. Infectivity can be neutralized by cholera virus. Not to be confused with human coxsackie B5 antiserum but the African swine fever virus. viruses can be distinguished by immunodiffusion, neutralization, and RNA swine hepatitis E virus (swine HEV) A hybridization. Causes a disease similar virus closely similar to human hepati- to foot-and-mouth disease in pigs. Fever tis E virus, isolated from swine in the and vesicular lesions on the feet and USA and elsewhere. A study of natu- snout. Replicates with CPE in pig kid- rally infected swine in Spain showed a ney cell cultures. Injected i.c. in newborn prevalence rate of 23%, highest in pigs mice causes paralysis and death in 5–10 less than 12 weeks old. days. Donkeys, cattle, rabbits, guinea Fernandez-Barredo S et al (2007) Can J Vet pigs, and chickens develop no disease Res 71 , 236 on exposure to virus. Laboratory infec- Meng X-J et al (1998) Arch Virol 143 , 1405 tions in humans with aseptic meningitis are reported. Coxsackie B5 injected into swine infertility and respiratory syn- pigs does not cause disease. A novel drome virus (SIRSV) See Porcine diagnostic reagent was recently devel- reproductive and respiratory syndrome oped by engineering diagnostic primer virus . sets for swine vesicular disease and foot-and-mouth disease viruses into swine influenza virus See Influenza recombinant cowpea mosaic virus par- virus . ticles which can be grown in cowpea swine norovirus A tentative species in plants (Vigna unguiculata ) to produce a the genus Norovirus . thermostable noninfectious reparation. King DP et al (2007) J Virol Methods 146 , 218 Swinepox virus (SWPV) The type species Seechurn P et al (1990) Virus Res 16 , 255 of the genus Suipoxvirus. Cultivation in eggs not reported. Replicates with CPE swine vesicular exanthema virus in pig kidney, testis, brain and embryo Synonym for Vesicular exanthema of lung cell cultures. Affects chiefly very swine virus . young pigs causing a generalized disease. Injected i.d. in rabbits it causes swollen baby syndrome A syndrome papular lesions, but cannot be passed. with widespread edema, abdominal Guinea pigs, suckling mice, calves, distension and bleeding seen in chil- sheep and goats are insusceptible. dren in Liberia in response to Lassa The pig louse transmits the disease, fever virus infection. although it may also occur in absence Monson MH et al (1987) Am J Trop Med Hyg of lice. Pigs may also suffer from infec- 36 , 408 tion with vaccinia virus. Synonyms : pigpox virus; variola suilla. Symmetrel™ Trade name for amantadine hydrochloride as 100 mg capsules. swine rotavirus See porcine rotavirus . Causes acute enteritis with diarrhea syncytial viruses Viruses which in in young pigs. Pigs are susceptible to cell cultures induce the formation of syncytial viruses 460

syncytia. These include bovine, ham- active in the presence of dsRNA. They ster, human, feline, and simian species activate an endoribonuclease, RNase L, belonging to the genus Spumavirus which is constitutive but latent in most as well as other viruses such as res- cell types. piratory syncytial virus, herpesvirus and measles virus, which also induce Syr-Daria Valley fever virus (SDFV) syncytia. Often used as an alternative An unassigned virus in the fam- name for foamy viruses. ily Picornaviridae . Caused outbreaks of infection in the Syr-Daria Valley, Loh PC (1993) In The Retroviridae, vol. 2, edited by JA Levy. New York: Plenum Press, Kazakhstan. Isolated from the blood of p. 361 a febrile patient. Antigenically related to mengo virus and Sikhote-Alin virus. syndrome A group of symptoms or signs Tick vectors Hyalomma asiaticum and which together characterize a disease. Dermacentor deghestanicus have been implicated in spreading the infection. 2–5 A synthetases Enzymes induced in cells by interferon treatment, and are T

Taarbaek disease Synonym for Bamble of these transforming proteins involves disease. binding to cell proteins that normally act as tumor suppressors, e.g. p53 and pRB Taastrup virus An unclassified virus (retinoblastoma) proteins. The adeno- detected by electron microscopy in the virus E1A and E1B proteins, the SV40T leafhopper, Psammotettix alienus, reared antigen and the human papillomavirus on healthy Festuca gigantea plants. The E6 and E7 proteins all bind to p53 and particles closely resemble those of filo- pRB, leading to their functional inactiva- viruses, Marburg and Ebola viruses. The tion. The polyoma middle T and large T particles are straight, slightly curved proteins also target cell proteins that or flexuous, sometimes with one end normally regulate cell growth; middle T curved into a ring, with an outer diam- binds to tyrosine kinase, phosphatidyli- eter of 200 nm. The median length was nositol-3 kinase and phosphatase 2A; 600–1100 nm. The morphological resem- and large T binds to pRB, but not to p53. blance to filoviruses is striking. Neil JC and Wyke JA (2005) In Topley & Lundsgaard T (1997) Virus Res 48 , 35 Wilson’s Microbiology and Microbial Infections , vol. 2, Tenth edition, edited by BWJ Mahy and V ter Meulen, London: Hodder Arnold, Tanganya virus A probable species in p. 330 the genus Hantavirus identified from Therese’s shrew (Crocidura theresae ) in T-cell epitopes Immunoreactive regions Guinea, West Africa. on virus proteins which interact with Klempa B et al (2007) Emerg Infect Dis 13, 520 T cells.

T antigens Tumor antigens, demonstrated T-cell lymphomas Many types of T-cell by immunofluorescence or CF test using lymphoma are found to contain EBV sera from tumor-bearing animals, that DNA, including angioimmunoblastic appear in the nucleus and in some cases lymphadenopathy, nasal lymphoma also in the cytoplasm of virus-induced and peripheral T-cell lymphomas. The tumor cells or cells transformed in vitro. T role of EBV in the pathogenesis of T-cell antigens are specific for the inducing lymphomas remains to be elucidated. virus, but not for cell species, and are encoded in the viral genome. In cells T cells Lymphocytes of thymic origin transformed by papovaviruses or certain that have been through thymic process- adenoviruses (e.g. types 12, 18, or 31) ing. They bear T-cell antigen receptors virus is not produced, and only certain (CD3) and lack Fc or C3b receptors. early genes are transcribed. The prod- Major T-cell subsets are CD4 (helper ucts of these genes act as oncogenes to cells) and CD8 (cytotoxic cells). transform the host cells to altered growth properties. Polyomavirus induces three T T-cell receptors (TcR) Antigen-specific proteins: small T, middle T, and large T; receptors on the surface of T lym- SV40 virus induces two: large T and phocytes which recognize peptides small T. The analogous gene products bound either to class I or class II major of adenoviruses and papillomaviruses histocompatibility complex (MHC) are no longer referred to as T proteins. molecules. Upon specific interaction of For adenovirus they are E1A and E1B, the T-cell receptor with antigen, signals and for papillomaviruses they are E6 are transduced through the plasma and E7, named after the E (early) genes membrane, activating the T cell to ini- that specify them. The action of several tiate cell division, secrete lymphokines T-cell receptors (TcR) 462

(T-helper cells) or lyze its target cells Taggert virus (TAGV) A strain of Sakhalin (cytotoxic T-lymphocytes). virus in the genus Nairovirus . Isolated in suckling mice from nymphs of the Parham P (1992) Nature 357 , 538 tick, Ixodes uriae , collected in tussock grass and under planks on the shore T6–T20 See newt viruses T6 to T20 . near a rookery of the royal penguin, Eudyptes chrysolophus schlegeli, on Mac- T21 See Xenopus virus T21 . quarie Island, 800 miles South East of Tasmania. Antibodies were found by Tb 1 Lu (NBL-12) cells (CCL 88) A cell plaque reduction test in four of 31 pen- line initiated from the trypsinized lung guin sera. Not reported to cause dis- of an adult female bat. ease in humans.

3T3-L1 (ATCC CL 173) cells (CCL Doherty RL et al (1975) Am J Trop Med Hyg 24 , 521 92.1) A continuous substrain of 3T3 (Swiss albino) derived through clonal isolation. Tahyna virus (TAHV) A strain of California encephalitis virus in the genus Orthobunyavirus , belonging to Tacaiuma virus (TCMV) A species in the California serogroup. Isolated in the genus Orthobunyavirus , belong- the former Czechoslovakia, Germany, ing to the Anopheles A serogroup. former Yugoslavia, France, and Italy. Isolated from sentinel Cebus monkeys Can cause a febrile illness in humans. and mosquitoes in Para and São Paulo, Serologically indistinguishable from Brazil. Not reported to cause disease in Lumbo virus isolated in Mozambique. humans. Bardos V and Danielova V (1959) J Hyg Tacaribe complex viruses A group of Epidemiol Microbiol 3 , 264 morphologically identical and sero- Drilganescu N and Glrjabu E (1979) Virol , Bucuresti 30 , 91 logically related viruses. Members of the New World group of the genus Arenavirus , named after the first one Tai virus (TAIV) An unassigned virus to be described: Tacaribe virus. Mostly in the family Bunyaviridae . Isolated transmitted by rodents. from Culiciomyia nebulosus in the Ivory Coast. Tacaribe virus (TACV) A species in the Institut Pasteur, Dakar (1980) Annual genus Arenavirus . One of the New Report , 60 World arenaviruses. Isolated from two species of Artibeus bats, A litura- Taiassui virus (TAIAV) A strain of tus and A jamaicensis in Trinidad, and Wyeomyia virus in the genus Orthobunya- named after a pre-Colombian tribe of virus , belonging to the Bunyamwera Trinidadian Indians. Has been recov- virus serogroup. Isolated from a pool ered on one occasion from mosqui- of Sabethini mosquitoes. toes. A silent infection can be induced Gerrard SR et al (2004) J Virol 78 , 8922 experimentally in the guinea pig. Not reported to cause disease in humans. taka-diastase A crude preparation from Martinez-Peralta LA et al (1993) In The the fungus, Aspergillus oryzae , which Arenaviridae , edited by MS Salvato. New has α-amylase activity and from which York: Plenum Press, p. 281 several nucleases can be isolated, including ribonuclease T1, ribonucle- tadpole edema virus (TEV) A strain of ase T2 and S1 nuclease. Frog virus 3 in the genus Ranavirus . Isolated from Rana catesbiana . Talfan disease virus Synonym for a strain Wolf K et al (1968) J Infect Dis 118 , 253 of porcine teschovirus in the genus Teschovirus. Talfan disease is a mild tadpole virus 2 A strain of Frog virus 3 in form of Teschen disease, first observed the genus Ranavirus . in Denmark and England in 1955–1957. 463 TATA box-binding protein

Tamana bat virus (TABV) A tentative Tanga virus (TANV) An unassigned virus species in the genus Flavivirus , with no in the family Bunyaviridae . Grouped known vector. with Okola virus. Isolated from mosquitoes, Anopheles funestus , in Tanzania Tamdy virus (TDYV) An unassigned and Uganda. Not reported to cause dis- virus in the family Bunyaviridae . Iso- ease in humans. lated from ticks, Hyalomma asiaticum asiaticum and H plumbeum plumbeum , Taniguchi bodies Inclusion bodies found collected from humans working with in epithelial cells of monkeys infected sheep and camels in the desert regions with Japanese encephalitis virus. of Turkmenistan and Uzbekistan. As these ticks parasitize rodents, hedge- Tanjong Rabok virus (TRV) A strain of hogs, hares, and birds, they may also be Bakau virus in the genus Orthobunya- hosts for the virus. No antigenic rela- virus. Isolated from sentinel rhesus tionship found to a range of viruses. monkeys, Macaca nemestrina , in penin- Does not agglutinate goose erythro- sular Malaysia. Antibodies found in cytes. Pathogenic for suckling and wild rodents, humans, birds, and bats. 3-week-old mice on i.c. injection. Not May cause a febrile illness with hemor- reported to cause disease in humans. rhagic signs in humans. Lvov DK et al (1976) Arch Virol 51 , 15 Tapiropé virus A possible species in the Tamiami virus (TAMV) A species in the genus Orbivirus isolated from phlebo- genus Arenavirus . One of the New tomine sand flies in the Amazon World arenaviruses, belonging to the region of Brazil. Antigenically related Tacaribe complex. Isolated from cot- to Changuinola virus. Not reported to ton rats, Sigmodon hispidus, in Florida, cause human disease. USA. Not reported to cause disease in humans. Tas proteins Proteins induced in cells by foamy viruses, which are trans- Tamiflu Generic name oseltamivir phos- activators of gene expression. They phate. Drug effective against both A bind directly to DNA motifs in the LTR. and B types of influenza. Acts by inhibiting influenza virus neuraminidase Tat protein A transcriptional activating activity, and was the first orally active protein induced in cells infected with drug for the treatment of influenza. human immunodeficiency virus (HIV). Increases the rate of transcription of Tanapox virus (TANV) A species in the HIV-specific mRNAs by binding to genus Yatapoxvirus causing local skin a specific RNA sequence in the LTR lesions in children in Kenya. The virus known as the Tat response element has also been isolated in Zaire. The (TAR). illness is non-fatal and of short dura- Cullen BR (1990) Cell 63 , 655 tion. Appears serologically the same as a virus causing disease in captive TATA box A nucleotide sequence in monkeys. Antigenically related to Yaba eukaryotic DNA, 25–30 bp upstream monkey tumor virus. Does not grow in from the transcriptional start site. It has eggs but replicates in monkey kidney the consensus sequence: cell cultures. Causes single and multiple lesions in monkeys. Monkey han- 5 -T A T A A A A-3 dlers have also been infected, but the - - natural host remains unclear. T T Synonyms: benign epidermal monkeypox virus; Yaba-like disease virus. TATA box-binding protein The TATA box is the site which binds a common tran- tandem repeats Multiple copies of a gene scription factor, promoting transcription. present in some virus DNAs such as Removal of the TATA box prevents or Orthopoxvirus genome DNA. greatly reduces transcription frequency. TATA box-binding protein 464

An analogous sequence (Pribnow Box) Tax protein A 40-kDa nuclear protein is found in prokaryotes. induced in cells infected with human T-cell leukemia viruses types I and II, Tataguine virus (TATV) An unassigned which is a transcriptional regulator of virus in the family Bunyaviridae . viral gene expression and necessary for Isolated from humans and mosqui- replication. Activates many cell genes toes (both Culex and Anopheles sp) in including IL-Z and proto-oncogenes Senegal, Cameroon, Central African c-fos and c-jun. Also acts to increase Republic, Nigeria, and Ethiopia. Does expression of several gene promoters; it not appear to be a cause of significant activates the cellular transcription factor disease in humans. NFκ B, which in turn can enhance HIV transcription in dually infected cells. Taterapox virus (GBLV) A species in the Hirai H et al (1994) Proc Natl Acad Sci 91 , genus Orthopoxvirus. Isolated from the 3584 African gerbil, Tatera kempi . Synonym : gerbilpox virus. taxonomy The theories and techniques Fenner F (2000) FEMS Microbiol Rev 24 , 123 of describing, naming, and classifying viruses or organisms. Taunton virus A strain of Norwalk virus in the genus Norovirus . Isolated from a Tchoupitoulas virus A strain of Seoul hospital outbreak of gastroenteritis in virus , isolated in New Orleans from the Taunton, UK in 1979. pancreas of a rat, Rattus norvegicus . This was the first Hantaan-related virus iso- Pether JVS and Caul EO (1983) J Hyg ( Camb ) lated from a feral rat in the USA. 91 , 343 Shi X et al (2003) J Med Virol 71 , 105 Tsai TF et al (1985) J Infect Dis 152 , 126 Taura syndrome virus (TSV) A virus isolated from marine penaeid shrimp, an TCMK-1 cells (CCL 139) One of a series unassigned virus in the family Dicistro- of SV40 virus-transformed cell lines, viridae. First described in 1992 from aqua- derived from C3H/Mai mice. The cells cultured shrimp in Ecuador it has now carry the SV40 T antigen. spread to most shrimp-producing countries in the western hemisphere and Asia, tegument In the Herpesviridae it is the and has been described as ‘The Ebola structure located between the capsid virus of the shrimp industry. ’ The virion and the envelope. It has been called an is a 32 nm icosahedral particle, buoyant ‘inner membrane’ although it does not density 1.338. The genome is a single- have the trilaminar unit structure char- stranded polyadenylated RNA 10,205 acteristic of true membranes. bases in length with two open reading frames. The virion consists of three major Tehran virus (TEHV) A strain of and one minor polypeptides. There is Sandfly fever Naples virus in the genus some variability in disease suscepti- Phlebovirus, belonging to the Salehabad bility among penaeid shrimp, but the complex of the sandfly fever virus major species cultivated in the Americas, serogroup. Isolated from Phlebotomus Penaeus vannamei , is highly susceptible papatasi in Iran. in post-larval, juvenile and adult shrimp with extensive cuticular epithelial necro- Tekupeu virus A possible species in the sis of the gut, gills, appendages, and gen- genus Orbivirus, isolated from phle- eral body. Transportation of shrimp from botomine sand flies in the Amazon South America to Asia resulted in the region of Brazil. Antigenically related first disease outbreak in Taiwan in 1998. to Changuinola virus. Not reported to Some disease-free shrimp lines have cause human disease. been bred, and restocking and careful management is currently the only way Tellina virus 2 A species in the genus to control the disease. Aquabirnavirus . Isolated from the Mari J et al (2002) J Gen Virol 83 , 915 bivalve mollusc, Tellina tenuis . 465 terminal transferase, deoxynucleotide transferase

Telok Forest virus (TFV) A strain of Bakau in Florida, Alabama, and Georgia, virus in the genus Orthobunyavirus , USA. Can cause a febrile illness with belonging to the Bakau virus serogroup. encephalitis in humans. Virus has been Isolated from the wild monkey, Macaca isolated from dogs and marsh rabbits. nemestrina , in Selangor, Malaysia. Antibodies present in cows, chickens, and humans. telomeres Terminal chromomeres of a chromosome; a DNA sequence required teratogenesis Two human viruses, for stability at the end of eukaryotic rubella virus, and human herpes- chromosomes. virus 5, may cause severe damage to the fetus. Three species of pestivirus Tembe virus (TMEV) A tentative species cause economically important terato- in the genus Orbivirus. Isolated from genesis in domestic animals: bovine virus mosquitoes, Anopheles nimbus , in Para, diarrhea virus, Border disease virus, and Brazil. Not reported to cause disease in hog cholera virus. The arterivirus, humans. porcine respiratory and reproductive syndrome virus causes fetal mortal- Tembusu virus (TMUV) A species in ity and reproductive failure in pigs. A the genus Flavivirus , belonging to the severe disease of the fetus called Smedi Ntaya serogroup. Isolated in Malaysia, syndrome is caused by porcine entero- Thailand, and Sarawak. Mosquito- virus. The bunyavirus, Akabane virus, borne. Not reported to cause disease in may cause epizootic bovine congenital humans. arthrogryposis and hydranencephaly. Atkins GJ et al (1995) Rev Med Virol 5 , 75 temperate bacteriophage Phage which can establish a lysogenic relationship Termeil virus (TERV) A tentative species with the host bacterial cell without kill- in the genus Orthobunyavirus . Isolated ing it. A form of latent infection. from Aedes camptorhynchus mosquitoe s in New South Wales, Australia. temperature-sensitive mutant A conditional lethal virus mutant that repli- terminal inverted sequence region cates at the ‘ permissive ’ temperature, A structure in the DNA genome of but fails to replicate at the ‘ restrictive ’ poxviruses. temperature at which wild-type virus replicates normally. N-terminal myristylation See myristylation . template A nucleic acid or other parent molecule which determines the compo- terminal redundancy Presence of identi- sition of a new molecule as it is being cal nucleotide sequences at both ends synthesized. of a genome nucleic acid. Occurs frequently in linear DNA virus genomes. tench reovirus (TNRV) A tentative species in the genus Aquareovirus , isolated terminal repetition A form of terminal from tench, Tinca tinca, in Germany. redundancy. Presence of nucleotide Not associated with disease in the fish. sequences at both ends of a genome Ahne W and Kolbl O (1987) J Appl Ichthyol nucleic acid which are either identical 3 , 129 or may be inverted. Inverted terminal repeats occur, e.g., in the adenovirus tenofovir (R)-9-(2-phosphonylmethoxy- genome and if such a DNA molecule propyl)adenine. An inhibitor of reverse is denatured and reannealed, the for- transcriptase. mation of structures known as ‘pan- Synonym : PMPA. handles ’ may be seen in electron micrographs of the molecules. Tensaw virus (TENV) A strain of Bunyamwera virus in the genus Ortho- terminal transferase, deoxynucleotide bunyavirus . Isolated from mosquitoes transferase An enzyme which will add terminal transferase, deoxynucleotide transferase 466

deoxyribonucleotide triphosphates the genus Orthobunyavirus . All isolated (dNTP) to the 3 hydroxyl group of a from birds. They are: DNA fragment. Usually isolated from calf thymus. Used in 3 end labeling of Bahig DNA and in tailing DNA fragments Matruh with complementary dNTPs to facili- Tete tate cloning. Tsuruse Weldona termination codon Any of the stop Tete virus (TETEV) A species in the genus codons, UAA, UAG, or UGA, which sig- Orthobunyavirus , belonging to the Tete nal termination of a growing polypep- serogroup. Isolated from a number of tide chain. See nonsense codons . species of birds in South Africa and Nigeria. Not reported to cause disease Teschen disease virus A strain of porcine in humans. teschovirus in the genus Teschovirus . Teschen is a region of the former tetradecanoylphorbol acetate (TPA) A Czechoslovakia where the first out- phorbol ester that is required for the breaks of the disease were observed transcription of the structural proteins in 1929–1930. However, the disease of human herpesvirus 8. Three classes may already have occurred in Moravia of transcript are identified in HHV8- in 1913. infected cells. Class 1 and class II genes can be transcribed without TPA in BC- 1 cells, but class III genes (primarily Teschovirus A genus in the family structural protein genes) are only tran- Picornaviridae containing a single spe- scribed in the presence of TPA. cies, Porcine teschovirus . At least 11 strains are recognized within the spe- tetraonine endogenous retrovirus cies. They are distinguished from other (TERV) A probable species in the picornaviruses in the internal ribos- genus Alpharetrovirus, identified by RT- ome entry site (IRES) which is shorter PCR sequence analysis of the genome (290 nt) and functional in the absence of a ruffled grouse, Bonasa umbellus . of eIF-4G. Analysis of the complete proviral DNA sequence showed that it occurred in Testudo iridovirus A tentative species in Tetraoninae (grouse and ptarmigan) but the genus Ranavirus. Isolated from tis- not in other subfamilies of galliform sues of two diseased Hermann’s tor- birds such as chickens or ducks. toises (Testudo hermanni) . Replicates Dimcheff DE et al (2001) J Virol 75 , 2002 in avian, mammalian, or reptilian cell lines at 28°C but not at 37°C tempera- Tettnang virus A strain of Murine hepatitis ture. Genome nucleotide sequence virus in the genus Coronavirus , probably related to that of Frog virus 3. originating from the mice used for isolation of what was originally thought Marschang RE et al (1999) Arch Virol 144 , 1909 to be an arbovirus. Smith AL et al (1983) Am J Trop Med Hyg 32 , Tet An inducible gene expression system 1172 developed in E coli which requires the addition of doxycycline for activation. TH-1, subline B1 cells (CCL 50) A ter- Allows inducible expression in a vari- rapene heart line established from the ety of viral vector systems. Has been trypsinized heart of a box turtle. introduced into HIV-1 in the development of candidate live vaccines. Thailand virus (THAIV) A species in the genus Hantavirus. Strain Thai 605 Verhoef K et al (2001) J Virol 75 , 979 was isolated from Rattus norvegicus and strain 749 from Bandicota indica , in Tete serogroup viruses Five serologi- Thailand. Human infections have been cally related subtypes of Tete virus in detected serologically. 467 Thiafora virus (TFAV)

Elwell MR et al (1985) Southeast Asian J Trop Theiler’s murine encephalomyelitis Med Public Health 16, 349 virus; Theiler’s virus.

THCAr virus A strain of Tembusu virus Lipton HL et al (2005) Virus Res 111 , 214 Monteyne P et al (1997) Immunol Rev 159 , in the genus Flavivirus, isolated in 163 Northern Thailand. Lipton HL (2008) Rev Med Virol 18 , 347 Kuno G et al (1998) J Virol 72 , 73 T-helper cell A thymus-derived lym- Theiler’s murine encephalomyelitis phocyte, which is usually CD4 and virus See Theilovirus . class II MHC antigen restricted, whose help is required for the production of normal levels of antibody by B lym- Theiler’s virus See Theilovirus . phocytes (Th-2) and for the normal development of cell-mediated immu- Theilovirus (ThV) A species in the genus nity (Th-1). Cardiovirus . Strains include Theiler’s murine encephalomyelitis virus, T-helper cells type 1 (Th1) T-cells with Theiler-like virus of rats, and Vilyuisk a cytokine expression profile in which human encephalomyelitis virus. The IFN-γ, interleukin -2, and interleukin-12 RNA genome lacks a poly C tract, in are produced, leading to cell-mediated contrast to other species in the genus. immunity. A common and usually inapparent infection of laboratory and wild T-helper cells type 2 (Th2) T-cells with rodents. Occasional animals develop a cytokine expression profile in which flaccid paralysis of the hind limbs. In interleukin (IL)-4, IL-5, IL-6, and IL-10 the 1930s Theiler showed that injection are produced, leading to production of of mice i.c. may produce flaccid paraly- virus-specific antibodies. sis after 12–29 days. Mice injected with the virus and recovering may later therapeutic index The ratio of the median develop demyelinating disease. Mice lethal dose of a drug to the median from a colony free of infection respond effective dose. more regularly to experimental infection. Replicates in mouse kidney cell Thetalymphocryptovirus A name pro- cultures and, after adaptation, proposed but not adopted for a genus in duces CPE. Strains vary in the cells in the family Herpesviridae . It would con- which they will replicate. There are tain Gallid herpesvirus 2 (Marek’s disease several strains: BeAn, DA, TO, FA, and virus) as its type species and Meleagrid GD I–VII; FA and GD VII on i.m. injec- herpesvirus 1 as a second member. The tion produce local myositis in mice. genus has now been established and FA causes encephalitis. The BeAn and named Mardivirus . DA strains are less virulent and produce persistent infections with chronic thiacytidine A nucleoside analog inhibi- demyelination. FA and GD VII are anti- tor of reverse transcriptase which is genically related. The virus has been synergistic with AZT, and active against purified and its crystallographic struc- resistant strains of HIV and against ture determined. Recently a closely hepatitis B virus infection. Also called related virus, Saffold virus, was found lamivudine or Epivir. It is marketed in in a child with fever of unknown origin combination with AZT as Combivir. in California. A second case has been described in a child in Canada, The Thiafora virus (TFAV) A species in the virus nucleotide sequence is closely genus Nairovirus . Erve virus is a strain related to the murine virus, and Saffold of Thiafora virus. Isolated from the virus may represent a new group of shrew, Crocidura sp, in Senegal. Has human neurotropic viruses. also been isolated in Cameroon. Synonyms: mouse poliovirus; murine poliovirus; mouse encephalomyeli- Institut Pasteur, Dakar (1977) Annual Report. tis virus; polio-encephalomyelopathy; Reference Center on Arboviruses 6 thick leg disease virus 468 thick leg disease virus Synonym for and Rhipicephalus sp in Egypt, Kenya, osteopetrosis virus, a strain of Avian Nigeria, and Sicily. Antibodies present leukosis virus . in sheep, goats, cattle, and camels. Causes disease in humans, sometimes Thimiri virus (THIV) A species in the severe. Optic neuritis and meningoen- genus Orthobunyavirus , belonging to cephalitis are reported. the Simbu serogroup. Isolated from Neumann G et al (2004) Curr Top Microbiol the pond heron or paddybird, Ardeola Immunol 283 , 121 grayii , in Tamil Nadu, India and from the lesser whitethroat, Sylvia curruca , in Thormódseyjarklettur virus (THRV) Egypt. Not reported to cause disease in A serotype of Great Island virus in the humans. genus Orbivirus , belonging to the Great Island complex of the Kemerovo sero- thiobenzimidazolones (TIBO) The first group. Isolated from a pool of ticks, non-nucleoside reverse transcriptase Ixodes uriae, from a sea bird colony in inhibitors to be discovered, in the early west Iceland in 1982. 1990s. They have relatively low toxicity but their oral bioavailability is poor. Thottapalayam virus (TPMV) The site on reverse transcriptase at A species Hantavirus which they bind is called the TIBO site. in the genus . Isolated from Suncus murinus Drug resistant mutants of HIV develop the Asian house shrew, , rapidly, but they have a role when used in Vellore, North Arcot District, Tamil Nadu, India. Not reported to cause in combination with other drugs. disease in humans. Genetically distant Pauwels R et al (1990) Nature 343 , 470 from the rodent-borne hantaviruses. Yadav et al (2007) Virol J 4 , 80 Thogotovirus A genus in the family Orthomyxoviridae of which Thogoto virus is the type species. The only other spe- three-day stiff-sickness virus Synonym cies in the genus is Dhori virus . Virions for Bovine ephemeral fever virus . contain six (THOV) or seven (DHOV) segments of linear negative-strand Thylaxoviridae (Saclike viruses. Greek: RNA, with a total genome size about thylax sac) A name suggested for the 10 kb. The conserved sequences of the RNA tumor viruses by Prof. Gilbert RNA segments are reminiscent of those Highet, Head of Department of Greek found in the genome segments of influ- and Latin, Columbia University, New enza virus. The viruses are transmit- York. Not adopted. ted between vertebrate hosts by ticks. Dalton AJ et al (1966) J Natl Cancer Inst A virus closely related genetically to 37 , 395 Dhori virus is Batken virus, isolated from mosquitoes and ticks from Russia. thymidine analogues Inhibitors of DNA It has been proposed that Araguari polymerase enzymes, including reverse virus should be assigned to the genus. transcriptase. The first analogs to be Da Silva EV et al (2005) Am J Trop Med Hyg used changed the thymidine mol- 73 , 1050 ecule by replacing the 3 -hydroxyl of the deoxyribose sugar with an azido Thogoto virus (THOV) A species in the group to produce 3 -azido-3-deoxythy- genus Thogotovirus . Virions contain six midine, known as Zidovudine or AZT. segments of linear negative-strand single- The presence of the azido group causes stranded RNA. Total genome 10 kb in chain termination. length. Sequences at the ends of each segment resemble those of influenza thymidine kinase (TK) An enzyme virus. The virus proteins are not related which is induced in cells infected with antigenically to those of influenza some DNA viruses and which cata- viruses, and there is no receptor-destroy- lyzes the phosphorylation of thymi- ing activity. Isolated from humans, cat- dine to thymidylic acid. In polyoma tle, camels, and from ticks of Boophilus virus-infected cells the induced enzyme 469 tick-borne encephalitis virus (Far Eastern subtype)

is cellular in origin, but the thymidine amino acid sequence across the strains kinase induced by herpesviruses is which make up the subtype. These specified by the virus genome and dif- include Absettarov virus, Hanzalova fers in several properties from the cell virus, Hypr virus, Kumlinge virus, and enzyme. This is the basis for the specifi- Neudoerfl virus, which are strains vary- city of the inhibition of herpesviruses ing in virulence and epidemiology. The by acycloguanosine (acyclovir) which is main vector is a tick, Ixodes ricinus , but phosphorylated by the virus-specified, mosquitoes and mites may be involved but not the host, thymidine kinase. in transmission. The tick is the most important reservoir of infection. Disease thymidylate kinase A transferase enzyme in humans is biphasic; a febrile illness of which converts thymidine 5 -phosphate 4–10 days is followed by meningitis or (dTMP) to thymidine 5 -diphosphate meningoencephalitis. Mild or inappar- (TDP) in the process of preparing nucleo- ent infections occur but in severe cases tide pools as substrates for DNA synthe- there is transient or permanent paralysis. Most poxviruses encode this enzyme sis. Infection occurs in central Europe and induce its synthesis in cells. from Scandinavia to the Balkans and from Germany to the former western thymidylate synthase (TS) An enzyme USSR. Experimentally the virus often which synthesises thymidine mono- kills mice; guinea pigs develop fever. phosphate (dTMP) from deoxyuridine Virus is often excreted in the milk of monophosphate and N5,N10-methylene goats, sheep and cows and may be a tetrahydrofolate by reductive methyla- source of infection for humans. It is also tion yielding dihydrofolate as a second- excreted in the urine. Control is by elim- ary product. ination of ticks. A vaccine is available. Synonyms: biphasic milk fever virus; TIBO See thiobenzimidazolones . biundulant meningoencephalitis virus; diphasic milk fever; Central European Tibrogargan virus (TIBV) An unassigned encephalitis virus. vertebrate rhabdovirus, antigenically Ecker M et al (1999) J Gen Virol 80 , 179 related to lyssaviruses. Isolated from the mosquito, Culicoides brevitarsis , in Australia in 1976. Not reported to tick-borne encephalitis virus (Far cause disease in humans. Eastern subtype) A subtype in the Mammalian tick-borne virus group in the genus Flavivirus. Strains within Tick-borne encephalitis virus (TBEV) A the subtype include Crimea virus, species in the genus Flavivirus , in the Karshi virus, Negishi virus, Oshima Mammalian tick-borne virus group. virus, and Russian spring–summer There are three recognized subtypes: encephalitis (RSSE) virus (Sofyn is the European, Far Eastern, and Siberian. The prototype strain). The species vary in viruses are distinguished as subtypes virulence and epidemiology but dif- or strains on the basis of nucleotide fer from each other by predicted amino sequence data, antigenic characteristics, acid sequences by only 2.2%. Humans geographical association, vector associa- become infected by tick-bite or con- tion, host association, disease association, sumption of milk from infected animals. and ecological characteristics. Overall, The clinical onset is an acute influenza- the maximum predicted amino acid like illness with mild fever, headache, sequence difference between strains and malaise that lasts for a week and is included in this species is less than 6%. followed by an asymptomatic period. A See Sofyin virus . second phase of illness involving men- Ecker M et al (1999) J Gen Virol 80 , 179 ingitis occurs in about 25% of infections, and usually resolves, but there is an tick-borne encephalitis virus (TBEV) overall case fatality rate of about 1%. A (European subtype) A subtype of formalin-inactivated vaccine is available Tickborne encephalitis virus in the genus for persons at high risk of infection. The Flavivirus, which vary by only 2.2% in vectors are ticks, Ixodes persulcatus and tick-borne encephalitis virus (Far Eastern subtype) 470

I ricinus. A severe human infection caus- Tillamook virus (TILLV) A strain of ing flaccid paralysis and 30% mortality. Sakhalin virus in the genus Nairovirus Disease may also occur in naturally belonging to the Sakhalin serogroup. infected rodents and birds. The disease Has been isolated from cattle in which is found in the former eastern USSR but it causes respiratory symptoms. a few isolations have been made in St Petersburg and elsewhere in the former Tilligerry virus (TILV) A serotype of western USSR, as well as China and Eubenangee virus in the genus Orbivirus , Japan. Experimentally the virus causes belonging to the Eubenangee virus sero- encephalitis in mice and fever in guinea group. Isolated from Anopheles annulipes pigs. Injected i.c. it causes encephalitis caught in the Port Stephens Peninsula in rhesus monkeys, sheep, goats, and of New South Wales, Australia. Not some wild rodents but not in others. reported to cause disease in humans. Control is by elimination of ticks. A vac- Gorman BH and Taylor J (1978) Aust J Exp cine is available. Biol Med Sci 56 , 369 Synonyms : Far East Russian encephalitis virus; Russian spring–summer tilorone hydrochloride 2,7-bis-[2- encephalitis virus. (Diethylamino)ethoxy]-fluoren-9-one dihydrochloride. An interferon inducer, tick-borne encephalitis virus (Siberian active by mouth, with maximum inter- subtype) A subtype of Tick-borne encepha- feron titres 24 h after administration. litis virus in the genus Flavivirus . Includes Inactive in cell cultures. Protects mice two strains from Central Siberia, Aina against vesicular stomatitis virus. Toxic virus, and Vasilchenko virus, for which for the hematopoietic and reticulo- nucleotide sequence information is avail- endothelial systems. able and shows a predicted amino acid Fitzwilliam JF and Griffith JF (1976) J Infect sequence difference from the European Dis 133 , Suppl, 221 subtype of 3.6–5.6% and from the Far Eastern subtype of 3.8–5.6%. Timbo virus (TIMV) An unassigned spe- Bakhvalova VN et al (2000) Virus Res 70 , 1 cies in the family Rhabdoviridae . With Chaco and Sena Madureira viruses forms the Timbo serogroup. Isolated tick-borne hemorrhagic fever A seri- from the lizard, Ameiva ameiva ameiva , ous febrile hemorrhagic disease with in Para, Brazil. Not isolated from significant mortality caused by sev- arthropods but considered to be arthro- eral strains of Tick-borne encephalitis pod-transmitted as it will replicate in virus , as well as Omsk hemorrhagic fever experimentally infected mosquitoes. virus , Kyasanur Forest disease virus , and Will kill suckling mice but they are Alkhurma virus. In different outbreaks, not as sensitive as Vero cells in which mortality has generally ranged from it replicates with CPE at 30°C. Not 10% to 25%. reported to cause disease in humans. tiger frog virus A strain of Frog virus 3 in Monath TP et al (1979) Arch Virol 60 , 1 the genus Ranavirus . Timboteua virus (TBTV) A species in the genus Orthobunyavirus. Isolated from a Tiger puffer nervous necrosis virus sentinel mouse in Brazil. (TPNNV) A species in the genus Betanodavirus, isolated from the marine Timbozal virus A possible species in the tiger puffer, Takifugu rubripes . genus Orbivirus . Isolated from phlebotomine flies in the Amazon region tiger salamander iridovirus See regina of Brazil. Antigenically related to ranavirus . Changuinola virus. Not associated with disease in humans. Tillamook virus (TLMV) A serotype of Great Island virus in the genus time-resolved fluoroimmunoassay Orbivirus . (TRFIA) A technique for solid phase 471 Togaviridae

antigen detection that has especially enzymes, before cultivation. Cell cul- been applied to respiratory viruses. tures are usually adherent to a solid Scalia G et al (1995) Clin Diagn Virol 3 , 351 surface (glass or plastic), but some cells group in suspension in liquid media. Tinaroo virus (TINV) A serotype of See culture medium . Akabane virus in the genus Orthobunya- George VG et al (1996) In Virology Methods virus , belonging to the Simbu sero- Manual , edited by BWJ Mahy and HO group. Isolated from Culicoides in Kangro. London: Academic Press, p. 3 Queensland, Australia. titer (titre) The concentration of infectious Tindholmur virus (TDMV) A serotype of virus present in a preparation measured Great Island virus in the genus Orbivirus , by bioassay or a relative measure of the belonging to the Great Island complex, concentration of a specific antibody in Kemerovo virus serogroup. Isolated antiserum. from Ixodes (Ceratixodes) uriae in the Faeroe Islands, Denmark. TK See thymidine kinase . Tlacotalpan virus (TLAV) A strain Tioman virus A probable species in the of Bunyamwera virus in the genus genus Rubulavirus , isolated from fruit- Orthobunyavirus , belonging to the eating bats (Peromyscus sp) on Tioman Bunyamwera serogroup. Isolated in island off the eastern coast of peninsu- Mexico from mosquitoes of Anopheles lar Malaysia. The virus is pleomorphic, and Aedes sp and Mansona titillans . Not 100–350 nm diameter, compatible with reported to cause disease in humans. viruses in the family Paramyxoviridae . The virus failed to cross-react with Tm melting temperature The tempera- antibodies against many known para- ture at which a transition occurs (e.g. myxoviruses, but did react by immuno- double-stranded to single-stranded fluorescence with Menangle virus, DNA) when temperature is the inde- although antiserum to Menangle pendent thermodynamic variable. virus did not neutralize Tioman virus. Sequencing of the nucleocapsid and TO virus A strain of mouse encephalo- phosphoprotein genes shows that myelitis virus. both Tioman and Menangle viruses are closely related to members of the Tobetsu-60Cr-93 virus A possible strain of Rubulavirus genus. Disease potential for Puumala virus in the genus Hantavirus animals or humans is unknown. isolated from the rodent Clethrionomys Chua KB et al (2001) Virology 283 , 215 rufocanus in Japan. Clement J et al (1997) Emerg Infect Dis 3 , 205 Tipranavir A non-peptidic protease inhib- Kariwa H et al (1999) Virus Res 59 , 219 itor approved by the FDA to treat HIV- 1 infection in 2005. Available as 250 mg Tocantins virus A possible species in the capsules under the name Aptivus®. genus Orbivirus , isolated from phlebotomine sand flies in the Amazon region tissue culture The growth or mainte- of Brazil. Antigenically related to nance of living tissue in a liquid or soft Changuinola virus. Not associated with gel medium in vitro. A large number disease in humans. of techniques have been described, and these can be classified under three Tocaxá virus A possible species in the general headings: (1) organ culture, in genus Orbivirus , isolated from phlebot- which the organization of the tissue omine sand flies in the Amazon region is maintained, e.g. culture of kidney of Brazil. Antigenically related to slices; (2) tissue culture in the strict Changuinola virus. Not associated with sense of the term, in which a fragment disease in humans. of tissue is cultured; and (3) cell culture, in which the tissue is broken down into Togaviridae (Latin: toga mantle or individual cells, usually by proteolytic cloak) A family of enveloped viruses Togaviridae 472

containing positive single-stranded lin- Tollwut virus Synonym for Rabies virus . ear RNA, genome 9.7–11.8 kb in size. Virions yield infective RNA which is toluidine blue A photoreactive dye. See capped at the 5 terminus and polyade- photodynamic inactivation . nylated at the 3 end. Isometric, probably icosahedral (T 4), nucleocapsids Tonate virus A possible strain of surrounded by a lipoprotein envelope Venezuelan equine encephalitis virus , a containing host cell lipid and virus- species in the genus Alphavirus . Isolated specified polypeptides, including one from mosquitoes in French Guiana. or more glycoproteins. Inactivated by Powers AM et al (2001) J Virol 75 , 10118 lipid solvents. Multiplication occurs in the cytoplasm where the genome RNA Topografov virus (TOPV) A species in the serves as mRNA for a polyprotein genus Hantavirus initially identified by comprising the nonstructural proteins, immunoblotting liver samples from which participate in RNA replication, Siberian lemmings, Lemmus sibiricus , col- including the formation of a 26S RNA lected near the Topografov river in the species termed ‘ subgenomic RNA ’ that Taymyr peninsula, Siberia. Following encodes the viral structural proteins passage of the material in laboratory- involved in encapsidation and envel- bred Norwegian lemmings, Lemmus lem- opment of the virus. Both viral and cel- mus , the virus was amplified by RTPCR lular proteinase enzymes participate in and sequenced. Topografov virus is cleaving the precursor polyproteins. 76–96% related by RNA sequence anal- The virus matures by budding from the ysis to other European hantaviruses plasma membrane. Agglutinate goose present in voles such as Puumula and and newly hatched chick erythrocytes. Khabarovsk viruses. Both lemmings There are two genera: Alphavirus and and voles are members of the rodent Rubivirus . subfamily Arvicolinae , but lemmings are Frey TK (1994) Adv Virus Res 44 , 69 a more ancestral species. The disease Strauss JH and Strauss EG (1994) Microbiol potential of Topografov virus in humans Rev 58 , 491 is not known. ten Dam E et al (1999) J Gen Virol 80 , 1879 Vapalahti O et al (1999) J Virol 73 , 5586 toll-like receptors (TLRs) A family of topoisomerase An enzyme that catalyses receptors found on macrophages, and guides the unknotting of DNA. dendritic cells, and polymorpho- Important in the replication cycle of nuclear cells. They have an extracellular most DNA viruses, and also for the domain containing leucine-rich repeats insertion of DNA into the host cell and an intracellular domain homolo- chromosome. gous to the cytoplasmic tail of the interleukin (IL)-1 receptor. TLRs form an Champoux JJ (2001) Ann Rev Biochem 70 , 369 early surveillance mechanism against TORCH T o r virus infection. Signaling through TLRs oxoplasmosis; ther; ubella; c h leads to activation of NF-κ B resulting ytomegalovirus; erpes simplex. A in the production of pro-inflammatory popular acronym devised in 1971 by cytokines. 11 TLRs are recognized, Prof. Andre Nahmias, now viewed but the two most important for virolo- with some despair by microbiologists. gists are TLR-3 (which recognizes Nahmias AJ et al (1971) Am J Obstet Gynecol dsRNA) and TLR-7 (which recognizes 110 , 825 ssRNA). Toll was originally discovered in Drosophila as a factor crucial for Toronto virus A strain of human calici- dorsoventral axis formation within the virus originally isolated from fecal Drosophila embryo. When Nobel lau- specimens of small children with reate Christine Nüsslein-Volhard dis- diarrhea in Toronto and called a covered this factor, she shouted ‘ Toll! ’ ‘ minireovirus. ’ Now known to be a which in German means ‘ great! ’ , giving strain in the genus Norovirus . it a name. Leite JPG et al (1996) Arch Virol 141 , 865 473 transcriptases

Torovirus A genus in the family Corona- describe the circular genome DNA. On viridae . The type species is Equine torovirus . the basis of sequence analysis, there Virions are enveloped, peplomer- are some 70 human isolates divided bearing particles containing an elon- into five groups, and at least ten ani- gated tubular nucleocapsid with helical mal isolates. TT virus particles in the symmetry. The capsid may bend into an blood are bound to immunoglobulin open torus, conferring a biconcave disk G forming immune complexes. Little or kidney-shaped morphology to the is known concerning pathogenesis, but virion (120–140 nm) or the capsid may there is evidence from some popula- be a rod-shaped particle (35–170 nm). tions that infection is acquired early in The virus genome is positive-sense life, perhaps by fecal-oral or salivary single-stranded RNA, about 28 kb in spread. See TT virus. length, with a polyadenylic acid tract at its 3 end. The RNA is surrounded by Torque teno mini virus ( TTMV) A group a major nucleocapsid phosphoprotein, of more than 20 tentative species in the mol. wt. ca. 20 1 0 3, which in turn is genus Anellovirus . The virus is 30 nm in enveloped by a membrane containing diameter and contains circular ssDNAs one major protein, mol. wt. 26 1 0 3 which range in size from 2.8 to 2.9 kb. and a phosphoprotein, mol. wt. In addition to humans, virus has been 37 1 0 3 . The peplomers, ca. 20 nm in found in cats, dogs, dourocouli, pigs, length, carry determinants for neutrali- tamarins, and Tupaia. The biologi- zation and hemagglutination, and con- cal significance of these viruses is not sist of a polydispersed N -glycosylated known. protein, mol. wt. 75–100 1 0 3 . Infected cells contain four subgenomic poly- Toscana virus (TOSV) A strain of adenylated RNAs with mol. wt. 3.0, 0.71, Sandfly fever Naples virus in the genus 0.46, and 0.26 1 06 . All the toroviruses Phlebovirus, belonging to the Salehabad identified so far cause enteric infection. complex of the sandfly fever Naples Species in the genus include Bovine complex. torovirus (Breda virus, cattle), Equine torovirus (Berne virus, horses) as well Toure virus An unclassified virus. as Porcine torovirus and Human toro- Isolated from Kemp’s gerbil, Tatera virus. The viruses that occur in humans, kempi, in Senegal. Not reported to cause horse, and cattle are serologically disease in humans. related. Tracambé virus An unclassified virus iso- Cavanagh D et al (1994) Arch Virol 135 , 227 Koopmans M and Horzinek MC (1994) Adv lated from culicine mosquitoes in the Virus Res 43 , 233 Amazon region of Brazil. tortoise virus 5 (TV5) A strain of Frog Trager duck spleen necrosis virus virus 3 in the genus Ranavirus . (TDSNV) A species in the genus Gamma- retrovirus, avian (reticuloendotheliosis) Torque teno virus ( TTV) The type species virus group. Related antigenically to of the genus Anellovirus. Originally reticuloendotheliosis virus (strain T), detected in the blood of a patient (ini- chick syncytial virus and duck infec- tials TT) and thought to be a cause of tious anemia virus. hepatitis, but this was not proven. Trager W (1959) Proc Soc Exp Biol Med Recent studies indicate that this is 101 , 578 one of the most ubiquitous of known viruses. The small non-enveloped viri- transcapsidation See phenotypic mixing . ons contain circular negative-stranded ssDNAs, 3.5–3.8 kb in length. The name transcriptases Enzymes which bring was adopted to replace the original about transcription. They may be patient initials and is derived from the DNA-dependent or RNA-dependent Latin Torques meaning necklace and according to the template nucleic acid Tenuis meaning thin, and is meant to being transcribed. transcription 474 transcription The process of transferring chemical carcinogens. But it is most the information encoded in the base usually observed after infection with sequence of a template nucleic acid oncogenic viruses, retroviruses, and molecule to another. It can be either the DNA tumor viruses. Retroviruses which formation of mRNA from DNA, or transform cells in vitro usually carry the production of a complementary a transduced oncogene, such as v-src , strand from single-stranded RNA. The which transforms the cells with very enzymes involved are DNA-dependent high efficiency. Alternatively, they may RNA polymerase, RNA-dependent RNA activate cellular proto-oncogenes follow- polymerase, and RNA-dependent DNA ing integration into the DNA genome. polymerase (reverse transcription). Both RNA and DNA tumor viruses may also induce proteins in infected cells transduction Transfer of host DNA genetic (e.g. T antigens) which combine with material from one cell to another by a tumor suppressor genes and inactivate transducing bacteriophage or an ani- them, leading to cell transformation. mal virus such as a retrovirus. The Transformed cells can be maintained introduction of oncogenes into cells indefinitely in culture, unlike non-trans- by retroviruses is a specialized form of formed cells. See immortalization . transduction. De Duve C (2007) Nat Rev Genet 8 , 395 transfection Direct transmission of transformation assay When viruses have genetic material (viral infectivity) a sufficiently high transforming activ- into cells using isolated nucleic acid ity (e.g. Rous sarcoma virus), it is pos- extracted from cells or virus particles, sible to assay the frequency with which in contrast to transmission by infective cells are transformed by observing the virus particles. effect of the virus on a monolayer cul- Miller G et al (1979) Proc Natl Acad Sci 76 , ture. Transformed cells grow in a man- 949 ner different from that of normal cells, forming small, heaped-up colonies transferases A large class of enzymes (foci) of morphologically altered cells. A catalyzing the transfer of groups from focus assay for transformation is analo- one molecule to another. Included gous to a plaque assay for infectivity. are enzymes transferring one-carbon β β groups (e.g. transmethylases) alde- transforming growth factor (TGF- ) hyde residues (e.g. transketolase), acyl A cytokine which is the most potent groups (e.g. transacetylase), sugars (e.g. known inhibitor of cell cycle progres- transglucosylase), nitrogenous groups sion of normal mammary epithelial β (e.g. transaminases), phosphorus-con- cells. Increased levels of TGF- are taining groups (e.g. protein kinase), found in the sera of AIDS patients. and sulfur-containing groups (e.g. CoA transferases). transgenic mice Strains of mice that have a deliberately altered genome. Used for transfer RNA See ribonucleic acid . experimental purposes. Genes may be added (e.g. adding human genes speci- transferrin receptor The receptor for the fying poliovirus receptors) or deleted binding of transferrin, a blood plasma (e.g. deleting the prion gene), in which protein for delivering iron to cells, is case they are called ‘ knock-out ’ mice. also the receptor for canine, feline, and human parvoviruses. transition temperature Temperature at which double-stranded nucleic acid transformation An alteration in cell mor- dissociates into single strands. phology and/or behavior, involving loss of contact inhibition and usually translation The process of making a pro- the acquisition of neoplastic potential. tein chain from the information in the Transformation may occur sponta- mRNA. The four-letter language of neously or after exposure to certain the nucleic acid (sequence of bases) 475 Tree shrew adenovirus (TSAdV)

is translated into a 20-letter protein transmissible spongiform encephalo- (sequence of amino acids). pathies See prion diseases . translocation A chromosomal aberration transmissible virus-dementia virus . which results in a change in the posi- Synonym for Creutzfeldt–Jakob disease. tion of a gene within the genome. transmethylase Enzyme catalyzing the transmission Transfer of a virus infection addition of methyl groups, e.g. to RNA from an infected organism to an unin- or DNA. Present in virions of Reoviridae fected one. Horizontal transmission is or Rhabdoviridae and involved in forma- most effective when it occurs by the tion of the 5 cap structure on mRNA. respiratory route, as with influenza or measles viruses, but the viral determi- transmissible enteritis of turkeys nants which promote respiratory trans- Synonym for Turkey coronavirus . mission are poorly understood. See epidemiology . transmissible gastroenteritis of pigs virus Synonym for Transmissible gastro- Mims CA (1995) Epidemiol Infect 115 , 377 enteritis virus . transovarian transmission A rare form Transmissible gastroenteritis virus of transmission to the embryo in which (TGEV) A species in the genus infected sperm carrying cytomegalo- Coronavirus, with worldwide distribu- virus or HIV-1 can infect very early in tion. Causes a commonly fatal disease the gestational period. of young pigs and occasionally has infected dogs. There is diarrhea, vomiting, dehydration, and death after 5–7 transport medium A sterile liquid used to days. In pigs over 3 weeks there is more prevent or reduce inactivation in speci- chronic diarrhea but recovery is usual. mens taken for virus isolation. Most Virus replicates in the small intestine transport media, of which there are and can be demonstrated in the feces many, contain a protein such as albu- by electron microscopy. Infectivity min, in which viruses are less readily survives drying at room temperature inactivated than in solutions with low for 3 days. Spread by direct and indi- protein concentrations. They also pre- rect contact: starlings may play a role vent drying and change of pH. in the mechanical spread. Replicates in pig kidney cell cultures. An inactivated transposons A transposable genetic ele- vaccine prepared in dog cell cultures ment in bacteria. Certain transposable has been used. Colostrum from recov- phages such as Mu move frequently ered sows protects the young. There from one insertion site to another posi- appears to be only one antigenic type. tion in bacterial chromosomal DNA. Synonyms: transmissible gastroenteri- The mechanism of this transposition tis of pigs virus; porcine respiratory is similar to that used by HIV when coronavirus. integrating into the eukaryotic chromo- Horzinek MC et al (1982) Infect Immun 37 , 1148 somal DNA. Pensaert MB et al (1986) Vet Quart 8 , 257 Hedges RW and Jacob AE (1974) Mol Gen Genet 132 , 31 transmissible mink encephalopathy virus The same as or a variant of scrapie virus. Present in mink brain tissue and trans-synaptic transport Spread of vir- infectivity will survive storage in 10% uses within the nervous system is usu- neutral formalin. On injection into mink ally within the neuronal axoplasm, but causes progressive neurological disease spread from one nerve cell to another after an incubation period of 6 months. seems to occur mainly at synapses. See also prion diseases . Chesebro BW (Editor) (1991) Curr Top Tree shrew adenovirus (TSAdV) A spe- Microbiol Immun 172 , 208pp cies in the genus Mastadenovirus . tree shrew adenovirus 1 (TSAdV-1) 476 tree shrew adenovirus 1 (TSAdV-1) A Trinidad rabies A form of rabies follow- strain of Tree shrew adenovirus in the ing the bite of an infected vampire genus Mastadenovirus . bat, most commonly Desmodus rotun- Brinckmann U et al (1983) EMBO J 2 , 2185 dus murinus . Clinically the infection Faissner A et al (1980) Intervirology 14 , 272 takes the form of an acute ascending myelitis. The disease occurs in cattle tree shrew herpesvirus Synonym for in Trinidad and South America, whose tupaiid herpesvirus 1. blood is the normal food of the bat. Humans are attacked only if the live- tree shrew paramyxovirus A cytopathic stock are shut away. virus from kidneys of a tree shrew, Tupaia belangeri, proved to be a para- Triniti virus An unclassified virus. Iso- myxovirus with some homology to lated from adult mosquitoes of Tricho- Hendra virus. prosopon sp in Trinidad. Not reported to cause disease in humans. Tidona CA et al (1999) Virology 258 , 425 tree shrew retrovirus A possible spe- trisodium phosphonoformate An anti- cies in the genus Gammaretrovirus . An viral agent. A pyrophosphate analog endogenous virus demonstrable in the that is a non-competitive inhibitor of placenta of a prosimian tree shrew, the viral DNA polymerase. Inhibits Tupaia belangeri , at full term. Could be both herpesviruses and hepatitis B activated in embryo skin cultures by virus. Used to treat cytomegalovirus treatment with idoxuridine. infections of the eye; effective against acyclovir-resistant viruses. Flugel RM et al (1978) Nature 271 , 543 Synonym : foscarnet. tree shrew rhabdovirus See tupaia virus . Helgstrand E et al (1978) Science 201 , 819 Jacobson MA et al (1991) J Infect Dis 163 , triangulation number See icosahedral 1348 symmetry . Trivittatus virus (TVTV) A strain of Tribec virus (TRBV) A serotype of Great California encephalitis virus in the genus Island virus in the genus Orbivirus , Orthobunyavirus . Isolated from Aedes belonging to the Kemerovo complex of trivittatus mosquitoes in North Dakota, the Kemerovo serogroup. Isolated from Iowa, Wisconsin, Illinois, Ohio, Florida, the tick, Ixodes ricinus , mice, Clethriono- Alabama, and Minnesota, USA. Probably mys glareolus and Pitymys subterraneus , not a significant cause of human disease. and goats in Slovakia. Antibodies are found in humans but the virus is not tRNA Abbreviation for transfer RNA. known to cause human disease. Trocará virus An unclassified virus iso- TRIC agent Not a virus. Synonym for lated from culicine mosquitoes in the chlamydia. Amazon region of Brazil. Not associated with human disease. trifluorothymidine (Trifluridine) An antiviral agent and an analog of thymidine. Acts like idoxuridine in inhibiting Trombetas virus (TRMV) A serotype of uptake of thymidine into DNA, and Anopheles A virus in the genus Ortho- has a similar range of antiviral activ- bun yavirus , belonging to the Anopheles ity although it is more potent. It is also A serogroup. more soluble and hence more dangerous in topical application. tropism The movement response of a virus toward particular cells or tissues in McNeill JI and Kaufman HE (1979) Arch which it replicates, e.g. Rabies virus has Ophthal 97 , 727 tropism for nerve cells (is neurotropic). Trinidad donkey virus A strain of Tropism may be modified by proteases. Venezuelan equine encephalomyelitis virus . Nagai Y (1993) Trends Microbiol 1 , 81 477 tumor necrosis factor (TNF)

Trubanaman virus (TRUV) An unas- TTM virus See Torque-teno minivirus . signed virus in the family Bunyaviridae , related to Gan Gan, Mapputta, and TT virus (TTV) A non-enveloped single- Maprik viruses. Isolated from mosqui- stranded circular DNA virus, genome toes, Anopheles annulipes . Antibodies length about 3.8 kb. A species in the are found in humans, cattle, sheep, genus Asnellovirus from the initials pigs, goats, horses, wallabies, etc. Not (TT) of a patient who developed non- reported to cause disease in humans. A, B, C, D, E, or G transfusion-related hepatitis. Representational difference T.RVL.II 573 virus A strain of Tacaribe analysis of DNA was originally used to virus in the genus Arenavirus . isolate a clone from serum comparing before and after transfusion. Using spe- trypsin An enzyme catalyzing the hydrol- cific primers to screen sera by PCR it ysis of peptide bonds on the carboxyl was found that more than 90% of some side of arginine, lysine and aminoethyl human populations are infected, and cysteine residues. the virus is distributed world-wide. However, multiple variants are found tryptic peptide A peptide formed from a in the human population, and also in protein by the action of trypsin. It has nonhuman primates. It seems likely arginine, lysine or aminoethyl cysteine that transmission may occur by the at the C-terminus. fecal–oral route as well as parenterally. Many of the TT viruses appear to be ts mutants Abbreviation for t emperature recombinants. So far TTV has not been s ensitive mutants. associated with any human or nonhuman primate disease. See Torque teno Tst-1 A member of the tissue-specific and virus and Torque teno mini virus . developmentally regulated POU family of transcription factors. Regulates early Davidson F et al (1999) J Infect Dis 179 , 1070 Khudyakov YE et al (2000) J Virol 74, 2990 and late gene promoters of human Mushahwar IK et al (1999) Proc Natl Acad Sci polyomavirus JC. 96 , 3177 Verschoor EJ et al (1999) J Gen Virol 80 , 2491 Tsuruse virus (TSUV) A strain of Tete Worobey M (2000) J Virol 74 , 7666 virus in the genus Orthobunyavirus , belonging to the Tete virus group. TTM virus See Torque teno mini virus . Isolated from a bird, Cyanopica cyanus , in Japan. Not reported to cause disease Tucunduba virus (TUCV) A serotype in humans. of Bunyamwera virus in the genus Orthobunyavirus . 3T3-Swiss albino cells and 3T6 cells (CCL 92) and (CCL 96) Two of a number Tula virus (TULV) A species in the genus of cell lines obtained by repeated pas- Hantavirus isolated from European sage of random-bred Swiss mouse common voles, Microtus arvalis and M embryo cells. The first number indicates rossiaemeridionalis . There is evidence of the passage interval, the second x l000 human infection in Moravia. the number of cells plated per 20 cm 2 . Passage at low multiplicity resulted in Vapalahti O et al (1996) J Gen Virol 77 , 3063 3T3 cells, which have not lost contact inhibition and cease dividing at low tumor necrosis factor (TNF) A cytokine cell density. They are therefore valuable produced by macrophages in response to for detecting the transforming abilities infection. Preferentially kills tumor cells of oncogenic viruses. Passage at a high and causes necrosis of some transplanted cell density resulted in 3Tl2 cells, which tumors in mice. Human TNF-α is a pro- have lost contact inhibition and grow to tein of 157 amino acids. Tumor necrosis a high cell density. factor β is a cytokine of mol. wt. 25 kD Todaro CT and Green H (1963) J Cell Biol which shares some sequence homology 17 , 299 and many functions with TNF-α . tumor suppressor genes 478 tumor suppressor genes Genes that It has proved difficult to induce malig- encode a product which negatively nant lymphomas experimentally in tree regulates the cell cycle. shrews with this virus. Probably non- pathogenic for tree shrews, but produces tumor viruses Synonym for oncogenic lymphoid granulomas experimentally in viruses. rabbit lung and spleen. Synonym : tree shrew herpesvirus. Tumor virus X (TVX) A species in the Bahr U et al (1999) Virus Res 60 , 123 genus Parvovirus. Isolated from a line Darai G et al (1979) J Gen Virol 43, 541 of human amnion cells originating from a Hamburg laboratory. turbot herpesvirus Synonym for pleuro- Hallauer C et al (1971) Arch Virol 35 , 80 nectid herpesvirus 1.

Tumucumaque virus A possible species turbot reovirus (TRV) A strain of Aqua- in the genus Orbivirus isolated from reovirus E in the genus Aquareovirus . phlebotomine flies in the Amazon Isolated from cultured turbot, Scoph- region of Brazil. Not associated with thalmus maximus , in a population with human disease. chronic mortality. Caused cytopathic effects in CHSE cells. Not associated TUNEL assay T erminal transferase- with disease in the fish. mediated dU TP n ick e nd l abeling. An Lupiani B et al (1989) J Aquat Anim Health assay for DNA degradation, e.g. in 1, 197 apoptosis. turbot virus-1 (TuV-1) An unassigned tunicamycin A compound which blocks virus in the family Picornaviridae . the formation of N -glycosidic protein– carbohydrate linkages. Turkey adenovirus A A species in the genus Siadenovirus, which causes tur- Tunis virus (TUNV) A serotype key hemorrhagic enteritis. A wide- of Uukuniemi virus in the genus spread infection of turkeys recognized Phlebovirus . in Europe, North America, Asia, and Australia. Disease usually apparent tupaia virus (TUPV) A tentative species in 4- to 12-week-old turkeys. There in the genus Vesiculovirus, isolated from is short-term depression with bloody a captive tree shrew, Tupaia belangeri , in droppings, followed by death or recov- Heidelberg, Germany. Antigenically ery. Mortality varies from 1 to 60%. related to vesiculovirus. The intestine is filled with blood, and death probably results from blood loss. tupaiid herpesvirus 1 (TuHV-1) An unas- Incubation period 5–6 days. Intestinal signed virus in the family Herpesviridae . contents are infectious only during Originally isolated from a spontane- the acute phase of disease. In chick- ously degenerating lung tissue culture ens the infection is usually subclinical. from an apparently normal tree shrew, Pheasants are susceptible with con- Tupaia glis . Appears to be a common siderable mortality. See also marble and silent infection of tree shrews as it spleen disease of pheasants . can often be isolated from mouth swabs Synonym: turkey hemorrhagic enteritis and from cell cultures of various organs. virus. Replicates in a tree shrew fibroblast cell Ianconescu M et al (1984) Avian Dis 28 , 677 line. Another strain was isolated from a McFerran JJ and Adair BM (1977) Avian malignant lymphosarcoma in an 8-year- Pathol 6 , 189 old tree shrew, a third from a Hodgkin’s Picovsski J et al (1998) Virology 249 , 307 sarcoma in a 9-year-old tree shrew, and four others from moribund animals aged turkey adenoviruses B (TadV-1, 2) 4–11 years. They are probably all strains Tentative species in the genus Avi- of the original virus isolate, as they adenovirus , originally isolated from tur- share close DNA sequence homology. keys with respiratory disease in 1972. 479 Tursiops truncatus parainfluenza virus 1

Antigenically distinct from other fowl Virus replicates in chick embryo cell adenoviruses. cultures with CPE, and in eggs kill- Scott M and McFerran JB (1972) Avian Dis ing the embryo. Virus attenuated by 16 , 413 egg passage can be used as a vaccine. Injected i.c. in mice causes encephalitis. Chicks and other birds are resistant. turkey adenovirus 3 (TadV-3) A strain of turkey adenovirus A. turkey parainfluenza virus See Avian paramyxovirus 3 . Turkey astrovirus (TastV) A species in the Astrovirus genus , with two serotypes. Turkeypox virus (TKPV) A species in the genus Avipoxvirus . An economically turkey astrovirus 1 (TastV-1) A serotype important pathogen of turkeys, closely of Turkey astrovirus . related to fowlpox virus. A fowlpox live attenuated vaccine is available and turkey astrovirus 2 (TastVB-2) A sero- can be used to vaccinate turkeys. type of Turkey astrovirus . turkey pseudo enterovirus 1 and 2 Turkey coronavirus (TCoV) A species in (TPEV1, 2) Unassigned viruses in the the genus Coronavirus : Group 3 . Causes family Picornaviridae . diarrhea in young turkeys. Replicates in embryonated turkey eggs producing Turkey rhinotracheitis virus (TRTV) A damage to the lining of the embryo gut species in the genus Metapneumovirus . similar to that seen in poults. The term Causes swelling of the orbital sinuses, ‘blue comb’ is also used to describe a torticollis (swollen head syndrome) in similar clinical disease caused by infec- broiler birds, and in young birds rales, tious enteritis virus. sneezing and nasal discharge are accom- Synonyms: transmissible enteritis of tur- panied by submandibular edema and keys virus; turkey blue comb disease. swelling of the orbital sinuses. The virus Dea S et al (1990) J Virol 64 , 3112 has been found in several European countries as well as South Africa, where the disease is called ‘ dikkop. ’ turkey entero-like virus (TELV) An unassigned virus in the family Lister SA and Alexander DJ (1986) Vet Bull Picornaviridae . 56 , 637 turkey strain T reticuloendotheliosis turkey hemorrhagic enteritis virus virus A strain of Reticuloendotheliosis (HEV) (TAdV-3) Synonym for turkey virus . adenovirus 3. Turlock serogroup viruses A group of turkey hepatitis virus (THV) An antigenically related viruses in the unassigned virus in the family genus Orthobunyavirus. They include Picornaviridae. Infectious only for tur- Turlock , Lednice, Umbre, and S 1954- keys. Multiplies in fowl embryos. 847-32 viruses. Klein PN et al (1991) Avian Dis 35 , 115 McFerran JB (1993) In Virus Infections Turlock virus (TURV) A species in the of Birds , edited by JB McFerran and MS genus Orthobunyavirus , belonging to McNulty. Amsterdam: Elsevier, p. 515 the Turlock serogroup. Isolated from birds, rabbits, mosquitoes of Culex turkey herpesvirus 1 Synonym for tarsatis and other Culex sp in Alberta, Meleagrid herpesvirus 1 . Canada and California, Texas and New Mexico, USA, Trinidad and Brazil. turkey meningoencephalitis virus A possible species in the genus Flavivirus . Tursiops truncatus parainfluenza virus 1 Caused a progressive and fatal paraly- Proposed name for a novel para- sis with enteritis of turkeys in Israel. influenza virus isolated from a male Tursiops truncatus parainfluenza virus 1 480

Atlantic bottlenose dolphin (Tursiops trun- type B oncovirus group See catus ) with respiratory disease. Limited Betaretrovirus . sequence analysis of the genome showed some relation to bovine parainfluenza 3. type B virus particles See B-type virus particles . Nollens HH et al (2008) Vet Microbiol 128, 231 type C oncovirus group See turtle papillomavirus Green turtle fibro- Gammaretrovirus . papilloma is a life-threatening disease of turtles, an endangered species. Serial type C virus particles See C-type virus cultivation of cell lines derived from particles . tissues of a green turtle, Chelonia mydas , yielded a papilloma virus-like trans- type D virus particles See D-type virus missible agent. particles . Lu Y et al (2000) J Virol Methods 86 , 25 type-specific antigen An antigen which Turuna virus (TUAV) A strain of Candiru defines a particular virus serotype, virus in the genus Phlebovirus , belong- normally on the basis of a neutraliza- ing to the Candiru complex of the tion test. sandfly fever virus group. Isolated from Lutzomyia sp in the tropical rain- Tyuleniy virus (TYUV) A species in the forest in Brazil. genus Flavivirus, a member of the Seabird tick-borne virus group. Isolated from a tick, Ixodes putus , collected from rifts in TVX virus See Tumor virus X . the rocks on Tyuleniy Island, Patience Bay, south-east of Sakhalin Island, in the Twinrix A combination vaccine for north of the former Soviet Far East, where protection against hepatitis A and B there is a colony of guillemots, Uria aalge . viruses. Has also been isolated from Ixodes uriae collected in Three Arch Rocks National two-dimensional electrophoresis A tech- Wildlife Refuge, on the Oregon coast, nique in which the constituents of a USA and from Ixodes putus collected sample are separated by electrophore- from pelagic cormorants, Phalacrocorax sis in one dimension on one property pelagicus , on the Commodore Islands. and in a second dimension, usually Pathogenic for suckling mice i.c. and at right angles to the first, on another i.p. but kills adult mice only on i.c. injec- property. It is used to resolve complex tion. Replicates with CPE in pig kidney mixtures of molecules. cell cultures. Antibodies are found in humans, fur seals, and several species of Ty elements Transposable elements of birds. The mosquito, Aedes aegypti , can be the yeast Saccharomyces cerevisiae . infected experimentally. Not reported to cause disease in humans. type A virus particles See A-type virus Lvov DK et al (1972) Arch Gesamte Virusforsch particles . 331 , 139 U

U1 One of a group of small nuclear ribo- Bundibugyo, Uganda, in August 2007. nucleoproteins (snRNP) that function The genome sequence of the virus was in RNA splicing. Others include U2, very different from all previous isolates U4, U5 and U6. of the virus (from Sudan, Zaire, Reston, Madhani HD and Guthrie C (1994) Annu and the Ivory Coast). In the outbreak, Rev Genet 28 , 1 more than 40 patients died, including 5 hospital staff, of more than 140 persons U virus Synonym for echovirus 11. Found infected. in children with acute upper respiratory illness. May cause rashes, uveitis, Uganda S virus (UGSV) A species in and gastrointestinal disturbances. the genus Flavivirus, a member of the Yellow fever virus serogroup. Isolated 63U11 virus A species in the genus in Uganda, Nigeria, and Central Orthobunyavirus serologically related to African Republic from mosquitoes of C group viruses. No known association Aedes sp and birds. Antibodies found in with disease. humans and Cercopithecus sp but there is no evidence that the virus causes human disease. Uasin Gishu disease virus (UGDV) A ten- Synonym : Makonde virus. tative species in the genus Orthopoxvirus. An infection of African wildlife, occa- UL The ‘ unique long ’ region of the DNA sionally transferred to horses. First genome of herpes viruses. described as a skin disease of horses on the Uasin Gishu plateau in Kenya ulcerative dermatosis virus Probably a in 1934. Similar clinical cases have strain of Orf virus . been described in horses in Burundi, Rwanda, Zambia, and the Democratic Trueblood MS (1966) Cornell Vet 56 , 521 Republic of the Congo. Lesions resembling papillomas are found on many ulcerative disease rhabdovirus (UDRV) parts of the body. The presumed wild- A tentative species in the genus life source of the virus is unknown. Vesiculovirus.

Uatumã virus A possible species in the ulcerative stomatitis of cattle virus genus Orbivirus, isolated from phle- Synonym for Bovine papular stomatitis botomine sand flies in the Amazon virus, although other viruses may cause region of Brazil. Antigenically related the same clinical picture. to Changuinola virus. Not reported to cause human disease. ultracentrifuge A high-speed centrifuge that will generate forces up to 500,000g . Used to separate and purify virus ubiquitin A protein containing 76 amino particles. acids that is highly conserved in all eukaryotic organisms. Addition of multi- ultrastructure The fine structure of cells ple ubiquitin molecules to a cellular pro- or viruses as revealed by the electron tein (ubiquitination) targets it for rapid microscope. degradation by the 26S proteasome. Muller S and Schwartz LM (1995) Bioassays Umatilla virus (UMAV) A species in 17 , 677 the genus Orbivirus; with Llano Seco, Minnal, and Netivot viruses forms the Uganda ebolavirus A new strain of Ebola Umatilla virus serogroup. Isolated from virus that struck the western district of the sparrow, Passer domesticus, and Umatilla virus (UMAV) 482

from mosquitoes of Culex sp in Oregon, acid; (3) the facility for reverse transcrip- Utah, Colorado, and Texas, USA. Not tion; and (4) the polarity of the genome reported to cause disease in humans. (negative or positive). There are seven categories of virus: single-stranded DNA Umbre virus (UMBV) A strain of Turlock viruses, double-stranded DNA viruses, virus in the genus Orthobunyavirus reverse transcribing viruses, double- belonging to the Turlock serogroup. stranded RNA viruses, negative polarity Isolated from mosquitoes and birds in single-stranded RNA viruses, positive Maharashtra, formerly Bombay State, polarity single-stranded RNA viruses, India. Not reported to cause disease in and unconventional agents such as pri- humans. ons, naked RNA viruses, viroids, and satellite viruses. UNAIDS United Nations Programme on Van Regenmortel MHV (2005) In Topley & HIV/AIDS. Wilson’s Microbiology and Microbial Infections , vol. 1, Tenth edition, edited by BWJ Mahy Una virus (UNAV) A species in the genus and V ter Meulen. London: Hodder Arnold, Alphavirus. Isolated from mosquitoes p. 24 of Psorophora , Aedes , Culex , Anopheles , and Coquillettidia sp in Brazil, Panama, Upolu virus (UPOV) An unassigned Trinidad, Colombia, French Guiana, virus in the family Bunyaviridae. Surinam, and Argentina. Not reported Isolated from a tick, Ornithodoros capen- to cause disease in humans. sis, on the Great Barrier Reef, Australia. Antigenically related to Aransas Bay Causey OR et al (1963) Am J Trop Med Hyg virus. Antibodies have been found in 12 , 777 cattle and kangaroo. Not reported to cause disease in humans. uncoating The release of nucleic acid from a virus in the process of viral UR2 (University of Rochester sarcoma infection of a cell. With some bacte- virus 2) sarcoma virus (UR2SV) A riophages the viral coat remains out- species in the genus Alpharetrovirus. A side the cell, and only the nucleic acid defective virus in which the RNA genome enters; thus penetration and uncoat- is only 3.2 kb in length, and carries the ing take place together. In animal cells ros oncogene, encoding a phospho- uncoating may occur as a result of tyrosine kinase. Transforms chicken fusion of the virus and cell membranes and rat embryo fibroblasts and induces or in the cytoplasm. In some viruses, sarcomas and fibromas in chickens. e.g. reoviruses, only partial uncoating occurs, and virus gene expression Balduzzi PC et al (1981) J Virol 40 , 268 and release of mRNAs occurs without Urabe Am 9 strain release of the genome from the core. A live attenuated strain of mumps virus, developed Helenius A (1995) Cell 81 , 651 in Japan and licensed as a vaccine in 1979. However it was associated with unconventional viruses See prion . a number of adverse events, including meningitis, so was subsequently with- ungulate transmissible encephalopathy drawn from many markets in the 1990s. Prion diseases in ungulate animals. Galazka et al (1999) Bull World Health Organ 77 , 3 unique DNA DNA sequences that occur only once in the genome. uracil A pyrimidine base, one of two principal bases in RNA. Can be formed in Universal System of Virus Taxonomy The DNA by the deamination of cytosine, ICTV has adopted a universal system of which in turn gives a point mutation taxonomy based upon the four principal as it is replicated to give adenine. See characteristics that define the strategy nucleic acid . of genome replication. These are:(1) the nature of the virus genome (DNA or uracil-DNA glycosylase A highly con- RNA); (2) the strandedness of the nucleic served gene in all herpesviruses except 483 Uxituba virus

channel catfish virus, which removes Utinga virus (UTIV) A strain of Oropouche deaminated cytosine and misincorpo- virus in the genus Orthobunyavirus , rated uracil residues from DNA. belonging to the Simbu serogroup. Isolated from the three-toed sloth, uracil N glycosylase An enzyme involved Bradypus tridactylus , in Para, Brazil. Not in herpes virus DNA replication, which associated with disease in humans. removes uracil from DNA and may be involved in DNA repair. Utive virus (UVV) A strain of Oropouche virus in the genus Orthobunyavirus , uridine The nucleoside of uracil and belonging to the Simbu serogroup. ribose. See nucleic acid . Isolated from a three-toed sloth, Bradypus tridactylus. ؅ Not associated with uridine 5 -triphosphate (UTP) A pyrimi- disease in humans. dine nucleotide, one of the four major constituents of RNA. See nucleic acid . Uukuniemi serogroup viruses Thirteen serologically related, tick-borne viruses Uriurana virus A possible species in the in the genus Phlebovirus , which form the genus Phlebovirus , isolated from phlebot- species Uukuniemi virus. Type species omine sand flies in the Amazon region Uukuniemi virus strain S23. They are: of Brazil near Tucurui. Antigenically related to Changuinola virus. Not EgAn 1825-61 virus reported to cause human disease. Fin V-707 virus Grand Arbaud virus Urmurtia/338Cg/92 virus A strain of Manawa virus Puumala virus in the genus Hantavirus. Murre virus Oceanside virus Uruará virus An unclassified virus iso- Ponteves virus lated from culicine mosquitoes in the Precarious Point virus Amazon region of Brazil. RML 105355 virus St Abbs Head virus Urucuri virus (URUV) A tentative spe- Tunis virus cies in the genus Phlebovirus , belonging Uukuniemi virus strain S23 to the sandfly fever virus serogroup. Zaliv Terpeniya virus Isolated in suckling mice from the blood of an apparently normal male rodent, Uukuniemi virus (UUKV) A species in the Proechimys guyannensis, found in Utinga genus Phlebovirus. First member of the Forest, Brazil. Antibodies were found Uukuniemi virus serogroup. Isolated in the sera of members of this species in from a rodent, Apodemus flavicollis , pas- Para and Amapa, Brazil. Not reported serine birds, a thrush, Turdus merula , to cause disease in humans. and Ixodes sp in Finland, the former Czechoslovakia, Poland, Hungary, the Uruma virus A strain of Mayaro virus. former USSR, Lithuania, and Oregon, Caused an outbreak of fever and head- USA. Antibodies are found in humans ache in Bolivia. but there is no evidence that the virus causes disease. US The ‘ unique short ’ region of the genome DNA of herpes viruses. Uukuniemi virus S23 (UUKV) The proto- Usutu virus (USUV) A species in the genus type strain of Uukuniemi virus. Flavivirus, belonging to the Japanese encephalitis virus group. Isolated from Uxituba virus A possible species in the mosquitoes of Culex , Mansonia , and genus Orbivirus , isolated from phlebot- Aedes sp, also birds, in South Africa, omine sand flies in the Amazon region Uganda, Cameroon, Congo, Central of Brazil. Antigenically related to African Republic, Nigeria, and Austria Changuinola virus. Not associated with and Hungary in Central Europe. Not human disease. reported to cause disease in humans. V vaccination The induction of protective of the group. A heat-resistant lipoprotein immunity against infection by admin- hemagglutinin, separable from the virus istration of a vaccine. particle, agglutinates turkey erythrocytes and those of some fowls. Virus suspen- vaccine A prophylactic or therapeutic sions are inactivated in 10 min at 60°C material containing antigens derived but dried virus withstands 100°C in the from a pathogen such as a virus which same time. Also inactivated by potas- on administration will stimulate active sium permanganate, ethylene oxide, and immunity and protect against infection chloroform, but resistant to ether. The with that pathogen. first virus to be grown in cell culture. Replicates in many cell types, including vaccine virus markers Characters which chick embryo, rabbit kidney, bovine, and can be used to distinguish vaccine human cells. Pocks are produced on the strains of a virus from wild strains of CAM up to 40.5°C, are large and white, the same virus. Growth characters, viru- and have a slight tendency to become lence, antigenic, and biochemical mark- hemorrhagic. Scarification of the virus ers can be used, but nucleotide sequence into the skin produces a local lesion differences provide the most definitive and immunity, but can cause spreading marker. and generalized infection in patients with skin disease or impaired immune vaccinia immunoglobulin (VIG) A prep- responsiveness. Vaccinia virus is under aration given by intramuscular injec- development as a recombinant vector tion to treat complications of vaccinia for other virus genes, and is being used such as disseminated vaccinia. to vector the rabies glycoprotein gene for use in animals. The use of vaccinia, vaccinia subgroup virus Synonym for especially strain MVA (modified virus orthopoxvirus. Ankara), as a recombinant vaccine vector in humans for HIV, malaria, and vaccinia subspecies Synonym for buffalo- other viral antigens, especially following pox virus. priming by DNA vaccination, is presently being evaluated in phase II clinical vaccinia variolae Synonym for Vaccinia trials. Similar local lesions are caused in virus . calf, sheep, and rabbit skin, and these tissues, as well as cell cultures, can be Vaccinia virus (VACV) Type species of used to produce vaccine. the genus Orthopoxvirus. Used as a live Synonym : vaccinia variolae. attenuated vaccine for protection against Alcami A and Smith G (1995) Immun Today smallpox. The origin of vaccinia virus 16 , 474 is not clear. The nucleotide sequences Blanchard TJ (1998) J Gen Virol 79 , 1159 of vaccinia and variola viruses show Moss B (1991) Science 252 , 1662 that they have about 150 genes in com- Smith GL (2005) In Topley & Wilson’s mon, with some 37 additional genes Microbiology and Microbial Infections, vol. that are unique to variola. Both viruses 1 Tenth edition, edited by BWJ Mahy and V ter Meulen. London: Hodder Arnold, p. 578 encode a number of growth factors and cytokines involved in the establishment vacuolating viruses Species of the genus of infection and suppression of host Polyomavirus. See Simian virus 40 and immune responses. Several antigens are Rabbit kidney vacuolating virus . demonstrable by precipitation, and neutralizing antibodies can be used to dif- valacyclovir (VACV ) A prodrug. The ferentiate vaccinia from other members l -valylester of acyclovir. It is rapidly 485 variola ovina virus

converted to acyclovir after oral admin- vesicles. The vesicles crust within a istration and provides higher plasma few days and heal, usually without levels of acyclovir than with oral acy- scarring, in a few weeks. Infection of clovir itself. adults is generally more severe than in Beutner KR (1995) Antiviral Res 28 , 281 children, and in immuno-compromised patients it may be so severe as to be life- valganciclovir A derivative of acyclo- threatening. A vaccine (Varivax-Merck) guanosine that is a potent inhibitor of is available for protection of infants. human cytomegalovirus (HHV5) repli- varicella-zoster immunoglobulin A cation, and is active by the oral route. preparation given by intramuscular It is metabolized to ganciclovir in vivo injection as postexposure prophylaxis and has comparable activity against for susceptible infants, adults, and cytomegalovirus. Used to treat CMV immunocompromised persons. retinitis, a common problem in AIDS patients. See ganciclovir varicella-zoster virus 1 (VZV) Synonym for Human herpesvirus 3 . Vand endogenous type C virus A possible species in the genus Gammaretrovirus . varicellavirus See Human herpesvirus 3 . Released spontaneously by a kidney cell culture from the long-tailed tree Varicellovirus A genus in the subfamily mouse, Vandeleuria oleracea , after 24 Alphaherpesvirinae, the type species of weeks in culture and 12 passages. The which is Human herpesvirus 3 . Form a virus-associated reverse transcriptase distinct lineage within the subfamily. and major internal protein p30 are Establish latent infection in cells of the immunologically related to the analo- sensory nervous system. There are 17 gous proteins of the simian sarcoma species assigned to the genus and one virus complex. The viral genome is tentative species ( Table V1 ). present in V oleracea cellular DNA, in multiple copies. variola major virus See Variola virus . Callahan R et al (1979) J Virol 30 , 124 variola minor virus See Variola virus . variant Creutzfeld–Jakob disease (vCJD) variola ovina virus Synonym for Sheeppox A distinct clinicopathological disease virus . that is apparently caused by bovine spongiform encephalopathy prion infection of humans. The onset of the disease is characterized by psychiatric Table V1. Species in the genus Varicellovirus abnormalities, sensory symptoms, and ataxia, eventually leading to dementia Bovine herpesvirus 1 and death. The age of patients is low Bovine herpesvirus 5 (19–39 years) as compared to spon- Bubaline herpesvirus 1 Canid herpesvirus 1 taneous CJD (55–70 years). There is a Caprine herpesvirus 1 distinct brain pathology characterized Cercopithecine herpesvirus 9 by abundant ‘ florid plaques’ decorated Cervid herpesvirus 1 with a daisy-like pattern of vacuola- Cervid herpesvirus 2 tion. By early 2008 a total of 166 cases Equid herpesvirus 1 had been reported, but the incidence of Equid herpesvirus 3 the disease appears to be declining. Equid herpesvirus 4 Equid herpesvirus 8 varicella A common childhood exan- Equid herpesvirus 9 them known as chickenpox caused by Felid herpesvirus 1 Human herpesvirus 3 human herpesvirus 3 (varicella-zoster Phocid herpesvirus 1 virus). The incubation period is about Suid herpesvirus 1 2 weeks, after which the characteris- Tentative species tic rash appears, composed of macules equid herpesvirus 6 which rapidly develop into fluid-filled variola suilla virus 486 variola suilla virus Synonym for VAV-488 virus A strain of Pirital virus , in Swinepox virus . the genus Arenavirus .

Variola virus (VARV) A species in the VAV-499 virus A strain of Pirital virus , in genus Orthopoxvirus . Causes human the genus Arenavirus . smallpox, a severe and frequently fatal disease with often confluent rash, Vearoy virus (VAEV) A serotype of Great fever, and prostration. Eliminated from Island virus in the genus Orbivirus , the world population in 1977. This was belonging to the Great Island complex possible because silent human carriage of the Kemerovo serogroup. Isolated of the virus does not occur and there from a pool of ticks, Ixodes uriae , col- is no natural animal reservoir. There lected from a shag, Phalacrocorax aris- were two main types of smallpox. One, tolesis , in the Lofoton Islands, Norway. occurring in Asia and the Middle and Far East, had a high mortality (20–30%) Vectavir Synonym for Penciclovir, an and was caused by variola major virus. anti-herpesvirus drug. The other, occurring in South America and West Africa, had a low mortal- vector-borne infections Virus infections ity (1–5%) and was caused by vari- transmitted to humans by vectors, ola minor virus. Both viruses produce which include arthropods (especially small white-domed pocks on the CAM mosquitoes and ticks), bats, and many after 72 h. Variola major produces pocks species of rodent. The vectors may at 38.5°C whereas variola minor does acquire the virus from a reservoir host, not produce pocks above 38°C. Variola such as a species of bird in the case of major is more lethal for chick embryos many alphavirus diseases. than variola minor. Replication is demonstrated with ease only in suckling vector control Since there are no licensed mice. African strains may be difficult to human alphavirus vaccines, inter- differentiate. Control is by immuniza- ruption of transmission by mosquito tion with vaccinia virus and isolation control is the only approach to control of cases and contacts. All remaining most alphavirus diseases. For viruses stocks of variola virus are now held in such as arenaviruses and hantaviruses two repositories: one in Atlanta, USA which are transmitted to humans by and the other in Novosibirsk, Russia. rodents, the best approach is to educate Their destruction would finally eradi- the public about the dangers of close cate variola virus, and this has been contact with rodents. recommended by WHO. Synonyms: Alastrim virus; Amaas virus; vectors In biological terms, virus vec- kaffir-pox virus; milk-pox virus; small- tors can be any of a variety of animals, pox virus; variola minor virus. arthropods or birds in which virus Fenner F et al (1988) Smallpox and Its Eradica- multiplication can occur, and which tion . Geneva: World Health Organization may then pass the virus infection on to Massung RF et al (1993) Nature 366 , 748 another human or other species. Many viruses persist in insects, rodents or variolation The deliberate inoculation of ticks, which may spread the infection infectious variola virus taken directly to humans in contact with them. In from a patient with smallpox. Practised genetic terms, viruses may act as vectors in the period before smallpox vaccina- to deliver genes to an organism which tion became available. Introduction of they infect, and because of the tropism the virus through the skin, rather than of viruses for certain specific cells or tis- natural infection via the respiratory sues they are considered to be promising tract, gave a less serious disease, but vehicles for gene delivery as an impor- even so about 1% of variolated patients tant component of medical treatment in developed smallpox and died. So vari- the future. See viruses as gene vectors . olation was only tolerated because of the fear of contracting smallpox vector transmission Normally, viruses naturally. multiply in the vector host, such as an 487 Vesicular exanthema of swine virus (VESV)

arthropod, which then transmits them and develop disease. Natural reservoir to another host, but mechanical trans- probably mammalian, with mosquitoes, mission is also possible. In this case the Aedes taeniorhynchus , A scapularis , and vector is only involved in providing Culex (Melanconion) sp, as the main transport for the virus, and no multi- vectors. Replicates with CPE in a wide plication of the virus occurs, so it is not range of primary cell cultures and con- an efficient form of transmission. tinuous cell lines of mammalian origin, and in embryonated eggs killing the VEE virus Abbreviation for Venezuelan embryo in less than 48 h. Diagnosis is equine encephalitis virus . usually by antibody detection (IgG or IgM ELISA). A formalinized vaccine Vellore virus (VELV) A serotype of has been used. Palyam virus in the genus Orbivirus , Johnson BJB et al (1986) J Gen Virol 67 , 1951 belonging to the Palyam virus sero- Yuill TM (1999) In Encyclopedia of Virology , group. Isolated from Culex pseudovish- Second edition, edited by A Granoff and RG nui and other species in Vellore, North Webster. London: Academic Press, p. 1967 Arcot District, Tamil Nadu, India. Not reported to cause disease in humans. Venezuelan hemorrhagic fever A severe disease caused by rodent-borne Gua- Vellore virus-like particles An unclas- narito virus, one of the South American sified fecal virus type IV. No proven hemorrhagic fever viruses. association with human gastroenteritis. Described from India. Morphologically Venkatapuram virus An unclassified arbo- enveloped particles similar to influenza virus. Isolated from mosquitoes, Culex virus. vishnui , in North Arcot District, Tamil Nadu, India. Not reported to cause dis- velogenic strains A term used to describe ease in humans. virulent virus strains, particularly in relation to Newcastle disease virus. Vero cells (CCL 81) A heteroploid cell line derived from the kidney of a normal Venezuelan equine encephalitis virus African green monkey, Cercopithecus (VEEV) A species in the genus aethiops . Used widely in virus replica- Alphavirus. Found in Venezuela, Brazil, tion studies and plaque assays. Colombia, Ecuador, Panama, Trinidad, and in recent years also in Mexico, Texas vertical transmission Spread of infection and Florida, USA. Antigenically very from parent to the young via the egg, closely related to Mucambo and Pixuna sperm, or in utero . Transmission via the viruses found in the Amazon area. A maternal milk is sometimes included. number of antigenic and genetic subtypes can be differentiated which have vervet monkey disease virus Synonym differing epidemiologies, geographical for Marburg virus . distribution, and disease importance. Causes disease in horses, donkeys, and vervet monkey herpesvirus Synonym for humans. Is more viscerotropic than Cercopithecine herpesvirus 5 . neurotropic. There is damage to blood Clarkson MJ et al (1967) Arch Gesamte vessels, and lesions are produced in Virusforsch 22 , 219 many organs. In horses and donkeys there is fever, loss of condition, diarrhea, vervet monkey virus See African green and in some cases signs of CNS involve- monkey polyomavirus . ment and often death (up to 80% case fatality). In humans the incubation period vesicle A closed membrane shell derived is 2–5 days. There is fever, severe head- from membranes by budding. A coated ache, tremors, diplopia, and a death rate vesicle is one which is surrounded by a of up to 1%. Laboratory infections occur basket of clathrin. readily, probably by inhalation. Horses, dogs, cats, sheep, and goats, but not cat- Vesicular exanthema of swine virus tle, are readily infected experimentally (VESV) The type species in the genus Vesicular exanthema of swine virus (VESV) 488

Vesivirus . There are at least 13 serologi- may be infected and have an influenza- cal types identified by letters A, B, C, like disease, and serum antibodies are etc. First recognized in California, USA relatively common among people liv- in 1932; spread in 1952 to most of the ing in rural areas where the viruses are USA but never observed in any other endemic. Almost all animals, including country. Controlled by quarantine, birds, can be infected experimentally. slaughter, and cooking of pig food, Virus has been recovered from arthro- which may have contained sea lion pods but their role as vectors is doubt- carcasses, as the virus is closely similar ful. Virus replicates on the CAM and to San Miguel sea lion virus. Spreads in the allantoic cavity. Also in chick by contact. Replicates in cell cultures embryo cell cultures and primary cul- of swine, horse, dog, and cat with CPE. tures of bovine, pig, and monkey cells. Only infects pigs causing a disease There are a number of antigenically similar to foot-and-mouth disease, but different strains that can be clearly dis- milder. Vesicles appear on the snout, tinguished genetically by nucleotide tongue, feet, and teats. Some strains sequence analysis of the phospho- injected into the tongues of horses protein gene. The New Jersey and and dogs cause local lesions. See San Indiana viruses share a common nucleo- Miguel sea lion virus . capsid (N) protein antigen. The glyco- Neill JD et al (1996) J Virol 69 , 4484 protein (G) antigen allows differentiation Studdert MJ (1999) In Encyclopedia of between these two viruses. Virology , Second edition, edited by A Calisher CH et al (1989) Intervirology 30 , 241 Granoff and RG Webster. London: Academic Rodriguez LL and Nichol ST (1999) In Press, p. 217 Encyclopedia of Virology , Second edition, edited by A Granoff and RG Webster. vesicular exanthema of swine virus-A48 London: Academic Press, p. 1910 (VESV-A48) A serotype of Vesicular exanthema of swine virus . Vesicular stomatitis New Jersey virus (VSNJV) A species in the genus Vesicular stomatitis Alagoas virus Vesiculovirus. The commonest species (VSAV) A species in the genus of VSV, first isolated in 1926. Causes Vesiculovirus . Isolated from a mule in periodic outbreaks of disease in cattle, Alagoas, Brazil. Antigenically distinct horses, and pigs in the USA with peaks from VSV New Jersey and VSV Indiana of activity every 10–15 years. The virus viruses. persists in wild pigs on Ossabaw Island, Georgia, USA. Frequent disease epizoot- Vesicular stomatitis Indiana virus (VSIV) ics occur in Mexico, Central America, Type species of the genus Vesiculovirus , and northern South America. Virus prop- first isolated from cattle in Richmond, erties similar to VSIV, the type species. Indiana, USA in 1925. A closely related species is New Jersey virus (VSNJV) iso- Vesiculovirus A genus of the family lated in 1926. Natural hosts are horses, Rhabdoviridae. Type species Vesicular cattle, sheep, and pigs, in which a dis- stomatitis Indiana virus . Vesiculoviruses ease resembling a mild form of foot- have five major polypeptides (L, G, N, P, and-mouth disease is produced. The and M) and a negative single-stranded disease is confined to North and Central RNA genome, 11.2 kb in length, includ- America, and the northern region of ing a 47 nt leader sequence. There are two small nonstructural proteins called South America. There are small papules or vesicles in the mouth and excess C and C (55 and 65 amino acids, respec- salivation. Lesions last only a few days. tively) which are highly basic, arginine Lesions on feet are uncommon except rich, and of unknown function, speci- in pigs. Teat lesions may occur in cat- fied in a second reading frame within the P gene. The genes are located in the tle. Raccoons and deer may constitute a reservoir of infection. Antibodies have order 3 -N, P, M, G, L-5 . Found in a been found in the turtle, Trionyx spini- variety of animals including mammals, fer and the snake, Natrix erythrogaster . fish, and insects. Laboratory workers and cattle handlers Brown F et al (1979) Intervirology 12 , 1 489 viral deformity virus (VDV)

Vesivirus A genus in the family vinblastine An alkaloid derived from the Caliciviridae . Type species is Vesicular periwinkle plant, Vinca rosea . Inhibits exanthema of swine virus . Strains of this synthesis of cellular RNA and protein. virus include bovine calicivirus, ceta- Mitosis is arrested in metaphase and cean calicivirus, primate calicivirus, San there is attachment and uncoating of Miguel sea lion virus, and skunk calici- infecting viruses but no viral macro- virus. Other species are Feline calicivirus , molecules are made. and tentatively mink calicivirus. Vinca alkaloids Cytotoxic agents used in VESV See Vesicular exanthema of swine the systemic chemotherapy of Kaposi’s virus . sarcoma lesions. vFLIPs See v iral F LICE i nhibitory Vinces virus (VINV) A strain of Caraparu proteins . virus in the genus Orthobunyavirus , belonging to the group C virus V genes Genes coding for segments of group. Isolated from mosquitoes in the variable region of the heavy chain Ecuador. or light chain of immunoglobulin molecules. Eukaryotic germ line DNA con- Calisher CH et al (1983) Am J Trop Med Hyg tains many different V exons. 32 , 877

VH2 cells (CCL 140) A cell line estab- Vindeln/L20Cg/83 virus A strain of lished from the heart of a normal Puumala virus in the genus Hantavirus . female Russell’s viper, Vipera russelli . Viper retrovirus (VRV) The type spe- viable virus Virus capable of replication cies of the reptilian virus group, in the when introduced into a suitable host cell. genus Gammaretrovirus. First observed in a cell line VSW, established from vidarabine See adenine arabinoside . the spleen of an Asian pit viper, Vipera russelli. Two further strains have been Videx Trade name for didanosine, or obtained from two different viper 2 3 -dideoxyinosine (DDI), a nucleo- heart cell lines, neither of which were side analog inhibitor of HIV reverse producing virus particles at first but transcriptase. spontaneously commenced to do so. These strains have been designated McLaren C et al (1991) Antivir Chem VV-VH-2 and VV-VH-3 to distinguish Chemother 2 , 321 them from the original strain VV-VSW. Characterization of VRV showed a vif protein A protein (v irion i nfectivity reverse transcriptase with a preference factor) encoded by human immuno- for Mg2 and a genome similar in size deficiency virus type 1 which plays a to that of murine retroviruses. role in HIV replication in peripheral blood lymphocytes. Andersen PR et al (1979) Science 204 , 318 Lunger PD and Clark HF (1978) Adv Virus Vilyuisk human encephalomyelitis Res 23 , 159 virus (VHEV) A strain of Theilovirus in the genus Cardiovirus . Encephalitis Viracept Trade name for an orally active occurs in northern Siberia and subse- antiviral drug, nelfinavir, which inhib- quently one-third of patients develop its HIV protease. chronic progressive disease similar to amyotrophic lateral sclerosis. Virus viral deformity virus (VDV) A disease isolated from CSF, blood, and brain first seen in 1993 in young yellowtail resembles encephalomyocarditis virus. fish in Japan. A birnavirus that is not Relation to disease not established. See infectious pancreatic necrosis virus also Saffold virus . (IPNV), isolated from diseased fish. Casals J (1963) Nature 200 , 339 Nakajima K and Sorimachi N (1996) Fish Lipton HL (2008) Rev Med Virol 18 , 347 Pathol 31 , 87 viral epidermal hyperplasia 490 viral epidermal hyperplasia A disease Table V2. Examples of viral hemorrhagic of young flounder Paralichthys oli- fever viruses of humans vaceu . First described in 1989 in Japan. Herpesvirus-like particles have been Virus Means of seen in diseased tissues. transmission Alkhurma virus Tick-borne viral erythrocytic infection A seasonal Chikungunya virus Mosquito-borne infectious disease of Mediterranean sea Dengue virus Mosquito-borne bass, Dicentrarchus labras, which is prob- Rift Valley fever virus Mosquito-borne ably caused by a species in the family Yellow fever virus Mosquito-borne Retroviridae . Crimean–Congo hemorrhagic Tick-borne fever viruses Pinto RM et al (1995) Arch Virol 140 , 721 Kyasanur Forest disease Tick-borne virus viral factories Discrete granular foci, Omsk hemorrhagic fever Tick-borne which are the sites of virus replication, virus seen in the cytoplasm of cells infected Guanarito virus Rodent-borne with poxviruses. They contain replicat- Junín virus Rodent-borne ing DNA surrounded by cellular mem- Hantaan virus Rodent-borne Lassa fever virus Rodent-borne branes. As infection progresses they Machupo virus Rodent-borne increase in size. Sin Nombre virus Rodent-borne Tolonen N et al (2001) Mol Biol Cell 12 , 2031 Sabià virus Rodent-borne Lake Victoria Marburg virus Unknown viral FLICE inhibitory proteins (vFLIPs) Zaire Ebola virus Unknown Viral inhibitors of cell death by apoptosis induced by several herpesviruses and poxviruses. F add-l ike I nterferonL1β - C onverting Enzyme [FLICE] caspase- Viral hemorrhagic septicemia virus 8-inhibitory proteins [vFLIPS] prevent (VHSV) A species in the genus the activation of apoptosis by FLICE. Novirhabdovirus , first recognized in the Cellular homologs of vFLIPS [cFLIPS] Danish village of Egtved. Formerly were subsequently identified which believed to be confined to portions of render cells resistant to apoptopic signals the European continent, but in 1988 but their exact physiological function is VHSV was isolated from adult chinook, unclear. Oncorhynchus tshawytsha , and coho, French LE and Tschopp, J (1999) J Exp Med O kisutch , salmon returning to two 190 , 891 hatcheries in the north-western part of Wu Z et al (2004) J Immunol 172 , 6313 Washington State, USA. Subsequently isolated from many species from the viral hemorrhagic fever viruses of North Sea and the Baltic Sea. European humans A term with no precise mean- and USA isolates are thought to be of ing applied to a group of viruses which independent origin. Causes a severe cause diseases characterized by fever and often fatal hemorrhagic septicemia and hemorrhagic phenomena resulting in salmonids, both young and sexu- from various forms of capillary dam- ally mature fish. Rainbow trout are age. They usually have natural animal severely affected but other trout are hosts (arthropod- or rodent-borne), and susceptible to inoculation. Pike, Esox humans may become infected through lucius, turbot, Scophthalmus maximus , venturing into the ecological domain of Atlantic cod, Gadus morhua , Pacific the virus and its natural host. The term cod, Gadus macrocephalus, and her- began to appear in the literature in the ring, Clupea harengus pallasi, and sev- early 1950s in discussions of Korean eral other fish species are susceptible. hemorrhagic fever virus. Examples are They lose appetite, become apathetic given in Table V2 . or swim abnormally. Abdomen is swol- Shelokov A (1970) J Infect Dis 122 , 560 len, gills pale, with hemorrhages in the Simpson DIH (1978) Bull World Health Org gill filaments, around the eyes, and at 56 , 819 fin bases. VHSV can be propagated in 491 virolysis

cultures of trout ovarian cells: best at virogenic cells Cells carrying a latent 12–14°C and not above 22°C. Complete viral genome and not producing infec- sequence analysis of several strains tive virus, but able to do so on being shows that the marine and freshwater grafted into an animal of a suitable spe- viruses differ by as few as 10 amino cies, or on co-cultivation or fusion with acid substitutions. a cell of a different species, or induction Synonyms: hemorrhagic septicemia by irradiation or certain chemicals. virus of fish; salmonis virus; viral hemorrhagic septicemia of trout virus; viroid A term introduced by Altenburg Egtved virus; Atlantic cod ulcus syn- (1946) to designate hypothetical symbi- drome virus. onts, akin to viruses, supposed to occur Bernard J et al (1992) J Gen Virol 73 , 1011 universally within the cells of animals, Betts AM and Stone DM (2000) Virus Genes and to give rise by mutation to viruses. 20 , 259 Experimental verification of this theory has not materialized. Diener (1971) Viramun e Trade name for nevirapine. proposed that the term be redefined and used as a name for agents such as Virazole Trade name for ribavirin. potato spindle tuber ‘ virus, ’ a small infective nucleic acid with no capsid Viread Trade name for tenofovir. protein and too little nucleic acid to code for its own replication. No helper viremia The presence of virus infectiv- virus has been demonstrated. Viroids ity in the blood. May occur as free are the smallest known agents of infec- infectious particles in the plasma or as tious disease, consisting of a highly infected peripheral blood cells. Occurs structured RNA molecule 246–375 nt transiently in many infections but virus in length in different viroids. More is rarely isolated, probably because the than 20 viroids have been completely viremia only occurs very early in the sequenced. The RNA is not translated infection. Chronic viremia occurs in a but is replicated by pre-existing host few infections: e.g., in Aleutian disease enzymes. Viroids have as yet only been of mink, hepatitis B or C or human found in plants where they induce immunodeficiency virus infections in economically important diseases. One humans, and lactate dehydrogenase- example of a small RNA genome in elevating virus infection in mice. vertebrate virology is the hepatitis delta Infective virus may also circulate as virus, but this does not replicate auton- virus–antibody complexes. omously, and requires a helper virus (hepatitis B virus). Virgin River virus (VRV) A strain of Altenburg E (1946) Am Nat 80 , 559 Tacaiuma virus in the genus Orthobun ya- Diener TO (1979) Science 205 , 859 virus , isolated from Anopheles Taylor JM (2005) In Topley & Wilson’s mosquitoes. Microbiology and Microbial Infections , vol. 2, Tenth edition, edited by BWJ Mahy and virion Synonym for virus particle. V ter Meulen. London: Hodder Arnold, p. 1269 viroceptors Virus gene products, especially of large DNA viruses, that have virokines Virus gene products that have homology with cellular receptors for functional homology with cytokines cytokines, and inhibit cytokine-induced and affect cellular function in a similar defense mechanisms. way. McFadden G and Graham K (1994) Semin Palumbo GJ et al (1994) J Virol 68 , 1737 Virol 5 , 421 virolysis Irreversible structural damage virogenes Cell DNA sequences carrying which may go as far as complete dis- information for production of com- integration of virus particles. When ponents of virus particles. See endog- certain enveloped viruses are exposed enous retrovirus to specific antiserum in the presence of virolysis 492

complement at 2°C there is neutrali- viruses Infectious units (obligate intra- zation of infectivity but no gross struc- cellular parasites) consisting of either tural damage. However, at 37°C there RNA or DNA enclosed in a protective is virolysis, presumably mediated by protein coat. Viruses are not organisms, late-acting components of complement. and contain no functional ribosomes or Radwan AI et al (1973) Virology 83 , 372 other cellular organelles and no energy- producing enzyme systems, although viropexis The engulfment of virus parti- many viruses contain enzymes involved cles by cells. A form of pinocytosis. An in nucleic acid transcription. They can- active process by the cell and an impor- not grow in size but their nucleic acid tant method of virus penetration. contains the necessary information for Synonym : engulfment. their replication in a susceptible host cell. This cell may provide some of the viroplasms A term used to describe enzymes necessary for viral replication inclusion bodies in rotavirus-infected but its main function is to provide the cells which are the sites for assembly of energy-producing systems. The host subviral particles enclosing the 11 seg- cell may or may not be destroyed in the ments of mRNA which are then repli- process of viral replication and release. cated to form double-stranded RNA The Latin noun virus is defective, i.e. molecules. does not have a full set of case-forms, singular and plural. Ancient gram- viroporin A term introduced by Carrasco marians used only the singular form. to describe virus proteins that enhance Modern usage has made the word a cell membrane permeability. countable entity and modern languages each pluralize it in their own fashion. Carrasco L (1993) Adv Virus Res 45 , 61 Smutny RJ (1999) ASM News 65 , 388 virosomes Liposomes with viral pro- van Regenmortel MHV (2005) In Topley & teins on their surfaces. For example, Wilson’s Microbiology and Microbial Infections , vol. 1, Tenth edition, edited by BWJ Mahy the hemagglutinin and neuraminidase and V ter Meulen. London: Hodder Arnold, surface units of influenza virus A can p. 24 be removed from the virus and relo- cated on the surface of liposomes. Such virus III Synonym for leporid herpes- virosomes can be used as antigens. virus 2. Liposomes are particles consisting of aqueous dispersions of phospholipid in Nesburn AB (1969) J Virol 3 , 59 the form of either multi- or unilamellar lipid bilayers. They are formed when virus assembly Formation of a virus a dried film of a phospholipid such as particle from its constituent parts. The lecithin is shaken with buffer and then process can vary from the autoassembly sonicated. of protein subunits around viral nucleic acid, to assembly of complex viruses at Morein B and Simons K (1985) Vaccine 3 , 83 cell membranes. Morein B et al (1978) Nature 276 , 715 Guo P (Editor) (1994) Semin Virol 5 , 1 virostatic A substance able to prevent viral replication. viruses as gene vectors With greater understanding of the human genome it virucidal A substance causing inactiva- is now clear that many human diseases tion of a virus. have a genetic basis. Gene therapy aims to correct these defects, and provided Virudox Trade name for idoxuridine. the normal gene sequence is known, viruses appear to provide the means to virulence The capacity of a virus to cause deliver new genetic information to the disease in the host. cell, since they have tropism for particular cells or tissues, they naturally deliver viruria Presence of infectious virus in the genes into the cell, and are expressed urine. in a regulated manor, and may persist 493 Visna/maedi virus (strain 1514) (VISNA)

for extended periods of time. The four a culture of BHK21 hamster cells and most commonly used viruses have thought to have been derived from the been retroviruses, adenoviruses, herpes bovine serum in the medium. simplex virus, and adeno-associated Verwoerd DW (1970) Prog Med Virol 12 , 192 virus. However, there has also been some recent work with alphaviruses virusoid A term used to describe sin- such as Semliki Forest virus. To date, a gle-stranded RNA satellite viruses. number of advances have been made Consist of a single-stranded RNA with all these systems, but it will take genome encapsidated in protein struc- more fundamental research in animal tures which are provided by the helper models despite an early rush to pursue virus. Most species are found in plants clinical trials none of which were com- but one, hepatitis delta virus, infects pletely successful. Recently vesicular humans. stomatitis virus was chosen as a selective killer of glioblastoma cells is a possible therapy for malignant gliomas in virus transport medium A solution used the brain. Following adaptation to these to preserve virus infectivity as much cells in vitro, the virus was tested in as possible during transport from a mouse brain, where it rapidly destroyed field location to a virus laboratory. It implanted glioblastomas after intranasal contains sterile buffered salt (e.g. 0.9% inoculation leading to olfactory nerve saline in phosphate buffer at pH 7.4), transport of the virus into the brain. protein (usually bovine serum albumin), antibiotics, and a pH indicator. Linden RM and Berns KI (2005) In Topley & See transport medium . Wilson’s Microbiology and Microbial Infections , vol. 2, Tenth edition, edited by BWJ Mahy and V ter Meulen. London: Hodder Arnold, visceral disease virus Synonym p. 1590 for Human herpesvirus 5 (human Luindstrom K (2005) Gene Ther 12, Suppl 1, cytomegalovirus). 592 Ozduman K et al (2008) J Neurosci 28 , 1882 visceral lymphomatosis of fowls An old Yoshizaki M et al (2006) J Gene Med 8 1151 term for the leukoses involving the viscera: Marek’s disease and the leukosis– virus de rue renforcé A term applied to sarcoma group of diseases. certain strains of rabies virus (street virus) of unusual virulence. Visna /maedi virus (strain 1514) (VISNA) (Icelandic: visna shrinking or wasting) virus induction Activation of a provirus A species in the genus Lentivirus, in the to replicate complete virus. May occur ovine/caprine lentivirus group. Causes spontaneously or be promoted by a slowly progressive demyelinating dis- various factors, e.g. exposure to com- ease of the CNS in sheep. Early signs are pounds such as idoxuridine. lip tremor and abnormal carriage of the head. Later there is progressive paraly- virus-like particle Structure resembling sis and death. The disease was imported a virus particle but which has not been into Iceland in 1933 with a shipment demonstrated to be infectious. of 20 Karakul rams from Germany, intended to provide a new gene pool for virus N A strain of avian influenza A the relatively isolated Icelandic sheep. virus. Within 2 years the two diseases maedi (dyspnea) and visna (wasting) emerged. virus replication The process of form- Sporadic cases were reported between ing progeny virus from input virus. It 1935 and 1951 in Iceland, when an involves the expression and replication extensive slaughter policy was started of the viral genomic nucleic acid and to eliminate pulmonary adenomatosis the assembly of progeny virus particles. and maedi as well as visna . Localized outbreaks of maedi occurred again virus X of bovine serum A possible spe- between 1958 and 1965 but there was cies in the genus Orbivirus . Found in no recurrence of the other two diseases. Visna/ maedi virus (strain 1514) (VISNA) 494

Virion is 85 nm in diameter with a dense vomiting and wasting disease of pigs core. Genome RNA is related to that virus Synonym for Porcine hemaggluti- of caprine arthritis encephalitis virus nating encephalitis virus . with homologies of 75–78% in the gag and pol genes, and 60% in the env gene. v-onc General term for a viral oncogene. Transmission requires close contact and seldom occurs between sheep out- von Magnus phenomenon A phenome- doors. Antibodies are formed but virus non observed during repeated passage is not eliminated from the animal. The of influenza virus A at high multiplic- virus undergoes antigenic change and ity. Results in a progressive increase in antibodies are formed to the new anti- the proportion of defective virus parti- genic type. This can occur several times. cles produced. These changes appear to be limited and von Magnus P (1954) Adv Virus Res 2 , 59 are probably not due to mutations so much as selection of strains expressing VPg Abbreviation for virion protein, various alternative antigens. Replicates genome-linked. A small virus-coded in cultures of sheep and human cells. protein attached through a phospho- Giant cells and CPE occur in 2–3 weeks. diester linkage from an amino acid (e.g. Transforms mouse cells in vitro . On the phenolic hydroxyl group of a tyro- injection into mice these cells will form sine residue in poliovirus) to the 5 sarcomas from which the virus can be end of the virion nucleic acid of certain rescued. See also progressive pneumo- viruses, e.g. picornaviruses. Plays an nia of sheep . essential role in RNA replication. Synonyms: chronic progressive pneumonia of sheep virus; Zwoegerziekte Vpr protein A protein induced in virus; La Bouhite; Graaf Reinet; ovine cells infected with human immuno- progressive pneumonia. deficiency virus type 1 which upregulates Narayan O et al (1993) In The Retroviridae , HIV gene expression. vol. 2, edited by JA Levy. New York: Plenum Press, p. 229 Vpu protein A nonstructural membrane- Sonigo P et al (1985) Cell 42 , 369 associated protein induced in cells Zink MC (1992) Semin Virol 3 , 147 infected by human immunodeficiency virus (HIV) type 1 which functions vistide Trade name for cidofovir. during virus release and also acts to downregulate the CD4 glycoprotein by vole poxvirus (VPV) An uncharacter- causing its retention and degradation ized virus in the family Poxviridae . in the endoplasmic reticulum. Isolated in Turkmenia from Microtus oeconomus and in Canada from Microtus Vpx protein A protein component of the pennsylvanicus . virion of human immunodeficiency virus (HIV) type 2. Nonessential for Volepox virus (VPXV) A species in the replication in vitro. Plays a role in genus Orthopoxvirus isolated from nuclear translocation of the HIV pre- a Pinon mouse, Peromyscus truei , integration complex. and a vole, Microtus californicus , in California, USA. More closely related Vranica virus A strain of Puumala virus in to raccoon poxvirus than to Old World the genus Hantavirus. Isolated from a poxviruses. bank vole in Bosnia. et al Virology 190 Knight JJ (1992) , 423 Reip A et al (1995) Arch Virol 140 , 2011 Regnery DC (1987) Arch Virol 94 , 159 W

W10777 virus A strain of Tamiami virus in fingerlings produced tumors in 87% of the genus Arenavirus . the fish within a 14-week period. Zhang Z and Martineau D (1999) J Virol 73 , W virus Abbreviation for Wollan virus. 8884 Also used in 1932 by Gay and Holden Zhang Z et al (1996) Virology 225 , 406 to designate a virus which in all probability was identical with herpesvirus walleye epidermal hyperplasia See B (Cercopithecine herpesvirus 1 ). percid herpesvirus 1 . Gay FP and Holden M (1932) Proc Soc Exp Biol Med 39 , 1051 Walleye epidermal hyperplasia virus 1 (WEHV-1) A species in the genus Epsilonretrovirus. Causes discrete epi- Wad Medani virus (WMV) A species in dermal hyperplasia on the skin of wall- the genus Orbivirus belonging to the eyes distinct from the diffuse epidermal Wad Medani complex of the Kemerovo hyperplasia caused by percid herpesvirus serogroup. Isolated from vari- virus 1. The virus is distinguishable ous ticks, Rhipicephalus sanguineus , from Walleye dermal sarcoma virus by Hyalomma sp, Amblyomma sp, and phylogenetic analysis of the genome. Boophilus sp, in Sudan, India, Jamaica, and Pakistan. Not reported to cause Walleye epidermal hyperplasia virus disease in humans. 2 (WEHV-2) A species in the genus Epsilonretrovirus. Causes discrete epi- Wallal virus (WALV) A species in the dermal hyperplasia on the skin of wall- genus Orbivirus , belonging to the Wallal eyes. Nucleotide sequence analysis virus serogroup. There are three mem- shows that this virus is distinct from bers of the serogroup: Mudjinbarry Walleye epidermal hyperplasia virus virus, Wallal virus, and Wallal K virus. 1 and from Walleye dermal sarcoma Isolated from Culicoides sp flies and virus, but the exact role played by marsupials in Queensland, Australia. each of the three retroviruses in patho- Antibodies found in wallabies, kanga- genesis and tumor formation remains roos, and other vertebrates. Has been to be determined. isolated from cases of kangaroo blind- LaPierre L et al (1999) J Virol 73 , 9393 ness and been shown to reproduce the disease experimentally. Probably the walleye herpesvirus percid herpes- causative agent of this disease. Not See virus 1 reported to cause disease in humans. . Doherty RL et al (1978) Aust J Biol Sci 31 , 97 walrus calicivirus (WCV) An unassigned virus in the family Caliciviridae , isolated Walleye dermal sarcoma virus (WDSV) from the feces of the walrus, Odobenus The type species of the genus rosmarus , collected off sea ice in the Epsilonretrovirus . A retrovirus that is Chukchi Sea. etiologically associated with a multi- Smith AW and Boyt PM (1990) J Zoo Wildl focal skin tumor of the fish, walleye Med 21 , 3 (Stizostedion vitreum vitreum). The dis- Smith AW et al (1983) J Wildl Dis 19 , 86 ease has been observed in up to 10% of adult walleyes in Oneida Lake, Wanowrie virus (WANV) An un- New York and in several lakes in assigned virus in the family Bunyaviridae . Canada. Cell-free filtrates of tumor tis- Isolated from the tick, Hyalomma mar- sue injected into 9-week-old walleye ginatum isaaci , and the mosquito, Culex Wanowrie virus (WANV) 496

fatigans , in India, Sri Lanka and Egypt. related to water exposure. Many of Has been isolated from the brain of a these viruses have also been spread by patient with hepatitis and hemorrhagic ingestion of uncooked shellfish, which disease of the gut. Pathogenic for new- filter and concentrate the virus. born but not for adult mice. Replicates with CPE in BHK21 cells. water buffalo herpesvirus Synonym for Khorshed M et al (1976) Indian J Med Res 61 , Bubaline herpesvirus 1 . 557 Wavre virus Described originally as Warrego virus (WARV) A species in the a picornavirus which agglutinated genus Orbivirus. With Mitchell River erythrocytes of several species includ- virus forms the Warrego virus sero- ing monkeys, guinea pig, swine, and group. Isolated from flies of Culicoides chicken. However, it is probably a sp in Queensland, Australia. Antibodies strain of Porcine parvovirus . Replicates found in wallabies, kangaroos, and cat- with CPE in pig kidney cell cultures tle. Has been isolated from cases of and was originally isolated from a pig kangaroo blindness. Not reported to kidney cell monolayer from an appar- cause disease in humans. ently normal pig. Cartwright SF et al (1969) J Comp Pathol Warrego K virus (WARKV) A strain of 78 , 37 Warrego virus isolated from a kangaroo. Huygelen C and Peetermans J (1968) Arch Gesamte Virusforsch 20 , 26 Warthin–Finkeldey cells Syncytial lymphoid cells found in measles virus WB virus A strain of parainfluenza virus infection in the human tonsil, Peyer’s type 5. Isolated in WI-38 cells from two patches, lymphoid tissue of the appen- patients with infectious hepatitis. dix, lymph nodes and spleen. Liebhaber H et al (1965) J Exp Med 122 , 1135, 1151 wart-hog disease virus Synonym for African swine fever virus . WC3 virus A strain of bovine rotavirus that has been evaluated in clinical tri- wart virus Synonym for papillomavirus. als in infants in Philadelphia, USA as a candidate human vaccine. wasting disease A chronic transmissible wasting disease of captive mule Clark HF et al (1986) Am J Dis Child 140 , 350 deer and elk which resembles scrapie of sheep. Presumably caused by a WE virus A strain of Lymphocytic chorio- transmissible prion, but the exact rela- meningitis virus (LCM) in the genus tionship to other transmissible spongi- Arenavirus . form encephalopathies is not known. Recently, the disease has also been WEE virus Abbreviation for Western identified in free-ranging moose (Alces equine encephalomyelitis virus . alces shirasi ) in Colorado, USA. Weldona virus (WELV) A strain of Tete Baeten LA et al (2007) J Wildl Dis 43 , 309 virus in the genus Orthobunyavirus , Williams ES and Young S (1993) Vet Pathol 30 , 36 belonging to the Tete antigenic virus group. Isolated from ceratopogonid water-borne transmission Some virus midges collected in northern Colorado, infections are spread through water, USA. although this mode of transmission is Calisher CH et al (1990) Am J Trop Med Hyg more common with bacteria, that can 43 , 314 survive in the environment for prolonged periods. Most enteric viruses Wesselsbron virus (WESSV) A spe- (astroviruses, noroviruses, picorna- cies in the genus Flavivirus , belong- viruses, and sapoviruses) can be spread ing to the Yellow fever virus group. in water, and large outbreaks of hepati- Epizootic in sheep causing abortion tis A and hepatitis E viruses have been and death of lambs and pregnant ewes. 497 West Nile virus (WNV)

Hemorrhages and jaundice occur in the analysis shows that most of the WEE ewes and meningoencephalitis in the virus genome is closely related to that fetuses. May cause abortion in cat- of EEE virus, but the genes encoding tle. Infects humans causing fever and virion coat proteins and the 3 untrans- muscular pains. Transmission is by lated region are closely similar to those mosquito bites. Found in South Africa, of Sindbis virus. It is highly likely that Namibia, Zimbabwe, Mozambique, Cam- WEE virus arose as a recombinant eroon, Nigeria, Uganda, Mada gascar, between EEE and Sindbis viruses. Botswana, and Thailand. Injection of Hahn CS et al (1988) Proc Natl Acad Sci 85 , suckling mice i.c. causes encephalitis. In 5997 rabbits and guinea pigs it causes abor- Reisen WK and Monath TP (1989) In The tions. Replicates in lamb kidney cell cul- Arboviruses : Epidemiology and Ecology , vol. tures and in eggs. 5, edited by TP Monath. Boca Raton: CRC Press, p. 90 West-Caucasian bat virus (WCBV) A ten- Strauss JH and Strauss EG (1994) Microbiol tative species in the genus Lyssavirus . Rev 58 , 491 Isolated in 2002 from a bat in the Krasnodar region of the former Soviet West Nile virus (WNV) A species in Union. the genus Flavivirus belonging to the Japanese encephalitis virus serogroup. Kuzmin IV et al (2005 ) Virus Res 111 , 28 First isolated in 1937 in Uganda. A silent or short febrile infection in humans Western blotting The transfer of pro- especially children, but a more severe teins which have been separated on a disease which can be fatal occurs in polyacrylamide gel to an immobilizing elderly people. There is a short incuba- matrix, commonly nitrocellulose. The tion period of a few days followed by proteins on the matrix can be probed fever, headache, and myalgia. A rash with, e.g., specific antibodies to iden- occurs in about half the cases. After 3–6 tify a particular protein species. See days there is usually complete recovery. Southern blotting . Occurs in Egypt, Uganda, South Africa, Towbin H et al (1979) Proc Natl Acad Sci 76 , Israel, India, the south of France and, 4350 since 1999, in the USA. There are strain differences between viruses from India Western equine encephalitis virus and the Far East and those from Africa, (WEEV) A species in the genus Europe, and the Middle East. The virus Alphavirus . Maintained in the wild that appeared in New York in 1999 as an endemic harmless infection of was phylogenetically similar to an iso- birds and mosquitoes, especially Culex late from geese in Israel. After causing tarsalis, but Culex stigmatosoma , Aedes more than 60 clinical cases with seven melanimon and Aedes dorsalis are also deaths in older people in New York, vectors. Man and horses are infected and the deaths of many crows and by mosquito bites. Disease produced other birds in the New York region, is similar to that caused by Eastern the virus overwintered and returned equine encephalomyelitis virus (EEE) in 2000 to cause 18 clinical cases with but milder. Mortality in horses 20–30% two deaths in older people. The strain and in humans 10%. Sequelae are that moved to New York appears to uncommon. Virus found in most of be particularly virulent for birds, but USA (except the eastern seaboard), has also caused deaths in horses and southern Canada and S America as has infected a variety of other mam- far as Argentina. Injection i.c. causes mals. Experimentally, West Nile virus meningoencephalomyelitis in a range causes encephalitis on i.c. injection into of rodents, monkeys, rabbits, pigs, and rodents, chicks, and rhesus monkeys. birds. Hamsters, mice and guinea pigs Virus is propagated in eggs causing can be infected by i.p. and i.m. injec- plaques on the CAM and in cell culture tion. Virus can be propagated in eggs of chicks and many mammals as well and cell cultures of many types in as mosquitoes. Birds are probably the which it causes a CPE. RNA sequence natural host, the virus being spread to West Nile virus (WNV) 498

humans by mosquitoes of many dif- White spot syndrome virus 1 A spe- ferent species. No vaccine is available cies in the genus Whispovirus . Highly for human use, and control is by use infectious for most known species of of insecticides and personal protection cultivated penaeid shrimp. Natural against mosquitoes. infections have been recorded in Lanciotti RS et al (1999) Science 286 , 2333 black tiger shrimp ( Penaeus monodon ), Kuruma shrimp (P japonica ), Chinese white shrimp ( P chinensis ), banana wet-tail of hamsters See hamster enteritis . prawns (P meguiensis and P inducus ), Wexford virus (WEXV) A serotype of and white shrimp (P vannamei ). Causes Great Island virus in the genus Orbivirus , cessation of feeding and increased mor- belonging to the Great Island complex, tality. There may be white inclusions Kemerovo serogroup. Isolated from visible on the cuticle. The genome of a pool of engorged ticks, Ixodes uriae , the virus is double-stranded DNA removed from a murre, Uria aalge , on 293,000 bp in length. Virions are ovoid Great Saltee Island, Eire in 1980. or ellipsoid in shape, 150 nm in diameter and 280 nm in length. Replication Whataroa virus (WHAV) A species in occurs in the nucleus, where virions the genus Alphavirus. Isolated from are assembled. mosquitoes and birds in New Zealand. Yang F et al (2001) J Virol 75 , 11811 No known association with disease. Closely related to Sindbis virus. white sturgeon adenovirus 1 An unas- Miles JAR et al (1971) Aust J Exp Biol Med Sci signed virus in the family Adenoviridae . 49 , 365 white sturgeon herpesvirus 1 and 2 wheat germ extract A preparation used Synonym for acipenserid herpesvirus for cell-free translation of messenger 1 and 2. RNAs from viruses or cells. Inglis SC et al (1977) Virology 78 , 522 white sturgeon iridovirus An unassigned virus in the family Iridoviridae . Whispovirus A genus of large double- stranded DNA viruses that cause white Whitewater Arroyo virus (WWAV) A spe- spot syndrome, a serious disease of cies in the genus Arenavirus discovered farmed shrimp worldwide. The only in 1995 in pack rats, Neotoma albigula , genus in the family Nimaviridae . The in New Mexico, USA. One of the New type species is White spot syndrome World arenaviruses. Associated with virus 1 . cases of human disease in California in van Hulten MCW et al (2000) J Gen Virol 81 , 2000. 307 whooping cough viruses Although there whitepox virus Variola virus pro- is no doubt that Bordetella pertussis is duces white pocks on the chick CAM, the major cause of whooping cough, whereas monkeypox normally proviruses may sometimes cause a similar duces red (hemorrhagic) pocks. A syndrome. Virus-associated cases are virus isolated in a Russian laboratory most common in England in the win- from healthy cynomolgus monkeys in ter and the viruses incriminated most Holland, a chimpanzee shot in West often are adenovirus, respiratory syn- Africa, a sun squirrel, and a multi- cytial virus, parainfluenza, and influ- mammate mouse, Mastomys natalensis , enza virus types A and B. produced CPE in cell cultures and white pocks on the CAM like variola virus. WI-1003 cells (CCL 154) One of several The observation could not be repeated, fibroblast-like strains derived from and later was found to have arisen by human adult lungs. contamination with variola virus. Marennikova SM and Shelukhina EM (1978) WI-26 VA4 cells (CCL 95.1) This cell Nature 276 , 291 line is an SV40 virus-transformed 499 Wongal virus (WONV)

derivative of WI-26 cells, a human Philapelphia, USA, in 1906 for use in diploid cell line established from medical research. It has a wide head, the embryonic lung tissue of a male long ears, and a tail length that is Caucasian. always less than its body length. It was from the Wistar rat strain that Sprague– WI-38 cells (CCL 75) A diploid cell line Dawley rats were developed. derived from normal female embryonic lung tissue. Has one of the broadest Wistar-King-Aptekman-Hokudai rats A spectra for human viruses of any cell strain of rats which provide a model line tested, and is particularly useful in for HTLV-1-associated myelopathy/ the isolation of rhinoviruses. Has been tropical spastic paraparesis. Following used for the preparation of a number HTLV-1 infection, these rats develop of human virus vaccines. the disease with paraparesis of the Synonym : Wistar Institute 38 cells. hind limbs after an incubation period of 15 months. wild strains Isolations made from natu- Kasai T et al (1999) Acta Neuropathol 97 , 107 rally infected hosts. Such strains may be different from laboratory strains. Witwatersrand virus (WITV) An unas- See vaccine virus markers . signed virus in the family Bunyaviridae . Isolated from a mosquito, Culex rubi- rotus , and rodents in Uganda, Mozam- wild-type The reference or original geno- bique, and South Africa. Kills mice on type, used for comparison with mutant injection. Probably non-pathogenic for or laboratory-adapted strains which humans. have arisen from it. WM 1504 E virus A possible species of wildebeest herpesvirus Synonym for the genus Gammaretrovirus . A non- Alcelaphine herpesvirus 1 . transforming mouse virus. Wild mice trapped in Los Angeles County, USA WIN-54954 An antiviral drug developed were found to have progressive lower to bind to the hydrophobic pocket in motor neuron paralysis of the hind VP1 of picornaviruses and inhibits legs. They also showed a high inci- their replication. See Pirodavir . dence of lymphomas. A C-type virus was present and the condition could windpocken Synonym for chickenpox. be passed by inoculation of newborn See Human herpesvirus 3 . wild or laboratory mice with extracts of tissue containing the virus. The neu- winter vomiting disease virus A better rological disease and the lymphoma name is ‘ acute epidemic gastroenteritis appeared to be caused by the same virus ’ as diarrhea may be as important virus. a symptom as vomiting which may be Oldstone MBA et al (1977) Am J Pathol 88 , absent, and the winter season is not a 193 well-documented feature. wobble hypothesis See anticodon . wish cells (CCL 25) A heteroploid human amnion cell line derived from colo- Wollan virus An unclassified fecal virus nies of ‘ altered ’ cells which appeared associated with acute epidemic gastro- after the primary cell culture had been enteritis in humans. Passed in volun- passaged 35 times. Has been used to teers given fecal extracts collected from differentiate virulent from avirulent a boarding-school outbreak in the UK. strains of measles virus. Clarke SKR et al (1972) BMJ 3 , 86

Wistar Institute 38 cells See WI-38 cells . Wongal virus (WONV) A serotype of Koongol virus in the genus Wistar (WI) rats An outbred strain of Orthobunyavirus . With Koongol virus albino rats of the species Rattus norvegi- forms the Koongol virus serogroup. cus developed at the Wistar Institute, Isolated from Culex annulirostris and Wongal virus (WONV) 500

Table W1. Serotypes in the Wongorr virus Woolly monkey sarcoma virus (WMSV) group A species in the genus Gammaretrovirus . Isolated from a fibrosarcoma of a Paroo river virus woolly monkey. Causes sarcomas on Picola virus Wongorr virus CS131 (WGRV-CS131) injection into marmosets. It is a trans- Wongorr virus MRM13443 (WGRV-MRM13443) forming defective virus requiring Wongorr virus V1447 (WGRV-V1447) simian sarcoma-associated virus as a Wongorr virus V195 (WGRV-V195) helper to produce infective virus. The Wongorr virus V199 (WGRV-V199) genome contains a single open reading Wongorr virus V595 (WGRV-V595) frame encoding a single protein v-sis, which is a fusion protein product of the virus env gene and the transduced c-sis oncogene of the woolly monkey. Coquillettidia crassipes in Queensland, Other genes (pol and gag ) were deleted Australia. Not reported to cause dis- when the genome was formed, so rep- ease in humans. lication is not possible in the absence of the SSAV helper virus. The sis gene β Wongorr virus (WGRV) A distinct species is derived from the cellular gene for - in the genus Orbivirus . First isolated platelet-derived growth factor which from mosquitoes, Aedes lineato pen- is involved in wound repair and will nis , trapped in Queensland, Australia stimulate growth of fibroblasts in vitro . in 1970. Related to Parou River virus Synonym : simian sarcoma virus. and Picola virus which together form the Wongorr serogroup. Experimental woolly monkey type C virus A strain of transmission to cattle, detected sero- Gibbon ape leukemia virus, in the genus logically, has been reported. Gammaretrovirus , a simian sarcoma- Parkes H and Gould AR (1996) Virus Res associated virus (SSAV), seen by elec- 49 , 11 tron microscopy and isolated from a spontaneous fibrosarcoma of a pet Woodchuck hepatitis B virus (WHV) A woolly monkey of Lagothrix sp. The species in the genus Orthohepadnavirus , woolly monkey lived in the same affecting woodchucks, Marmota monax , household as a pet gibbon, Hylobates over a broad area of the eastern USA. lar , and the woolly monkey must have Does not infect other species such been infected by the gibbon ape leuke- as ground squirrel or other rodents. mia virus, since the isolates are of the Woodchucks can be infected experi- same virus species. mentally with the ground squirrel hepatitis virus; however, the virus has not Wut virus Synonym for Rabies virus . been grown in cell culture. Summers J et al (1978) Proc Natl Acad Sci 75 , Wyeomyia virus (WYOV) A species in 4533 the genus Orthobunyavirus , belonging to the Bunyamwera serogroup. woodchuck herpesvirus Synonym for Isolated from birds and mosquitoes in marmodid herpesvirus 1. Colombia, Panama, Brazil, Trinidad, and French Guiana. Causes a febrile ill- woodchuck herpesvirus marmota 1 ness in humans. Synonym for marmodid herpesvirus 1. X

X14 virus Synonym for Tumor virus X . xenotropic murine type C viruses A subspecies of the genus Gammaretrovirus Xaraira virus A possible species in the which is endogenous in mice, and genus Orbivirus , isolated from phlebot- infects and replicates efficiently only omine sandflies in the Amazon region in cells from a species other than the of Brazil. Antigenically related to Chan- mouse. The restriction is based on the guinola virus. Not associated with dis- surface glycoprotein (gp70) of the virus, ease in humans. for which mouse cells do not carry available receptors on their surface. XC cells (CCL 165) A cell line derived This may be because the receptors are from a transplantable tumor which was blocked by glycoproteins of the endo- induced in newborn outbred Wistar genous xenotropic virus. The first xeno- rats by the intramuscular injection tropic virus was identified in NZB mice, of the Prague strain of Rous sarcoma but all strains may carry such viruses. virus. They all show similar interference tests, p30 and reverse transcriptase. They dif- Xenopus virus T21 (XV-T21) A probable fer, however, in p12, gp70, and nucleic species in the genus Ranavirus . acid sequences. α or class II viruses Essani K and Granoff A (1989) Intervirology are inducible by idoxuridine. May be 30 , 187 present in mouse strains which also have an ecotropic virus. β or class III xenotransplantation A shortage of human viruses are not inducible and are pro- organs for transplantation therapy duced spontaneously, but only by a few has led to studies of other species that mouse strains and these strains do not might become suitable organ donors produce ecotropic viruses. for humans. Initially, nonhuman pri- Levy JA (1978) Curr Top Microbiol Immunol mates were considered suitable, and 79 , 111 more recently pigs. However, in each case there has been concern about xenotropic virus An endogenous virus possible transmission to the recipient which will not replicate complete virus of endogenous or exogenous viruses particles in cells of the species in which found in the donor species. it occurs naturally. Sometimes called X tropic or S tropic viruses. Boveva RS et al (2001) Clin Microbiol Rev 14 , 1 Xiburema virus (XIBV) An unassigned xenotropic murine-leukemia virus- vertebrate rhabdovirus, antigenically related virus (XMRV) A virus genome unrelated to other rhabdoviruses. identified in prostate cancer tissue from Isolated from Sabethes (Sbn) intermedias patients homozygous for a reduced in Brazil in 1977. activity variant of RNase L. The virus was reconstructed from prostate tissue Xingu virus (XINV) A strain of Bunya- RNA and replicated in a prostate can- mwera virus in the genus Ortho- cer cell line DU145, and shown to be bunyavirus . Isolated from a human susceptible to inhibition by interferon (blood). Not known to cause human and its downstream effector, RNase L. disease. The significance of this virus in the Calisher CH et al (1986) Am J Trop Med Hyg pathogenesis of prostate cancer is 35 , 429 unknown. Dong B et al (2007) Proc Natl Acad Sci USA Xiphophorus retrovirus A C-type virus 104 , 1655 found in melanoma tissue of platyfish, Xiphophorus retrovirus 502

Xiphophorus maculatus . Probably an XJ virus A strain of Junín virus in the endogenous virus. The reverse tran- genus Arenavirus . scriptase of the virus was found to cross-react with the feline leukemia X-linked lymphoproliferative disease virus reverse transcriptase. The possible (XLP) See Duncan’s disease . role of Xiphophorus retrovirus in tumor formation has not been established. X-ray crystallography A method which Petry H et al (1992) Virology 188 , 785 has been applied to study the structure of viruses and subviral virion compo- Xiwanga virus A possible species in the nents which have been crystallized. genus Orbivirus, isolated from phle- Johnson JE and Chiu W (2000) Curr Opin botomine sand flies in the Amazon Struct Biol 10 , 229 region of Brazil. Antigenically related to Changuinola virus. Not associated X-tropic virus Xenotropic viruses. with human disease. XJ-cl 3 strain of Junín virus An avirulent strain, non-pathogenic for guinea pigs. Y

Y-1 cells (CCL 79) Clone Y-1, a steroid- goose and duck embryo cell cultures secreting cell strain, was initiated from without CPE. Pathogenic for newborn an adrenal cortex tumor of a male mice. Not reported to cause disease in LAF1 mouse. humans. Y 62-33 virus A species in the genus Marhoul Z et al (1976) Acta Virol ( Prague ) 20 , 499 Alphavirus , related to Western equine encephalitis virus, probably of Old Yaba-7 virus (Y7V) A serotype of Akabane World origin. virus in the genus Orthobunyavirus , Y73 sarcoma virus (Y73SV) A species in belonging to the Simbu virus sero- the genus Alpharetrovirus. group. Isolated from mosquitoes, Mansonia africana in Nigeria. Yaba-like disease virus Synonym for Tanapox virus. Yacaaba virus (YACV) An unassigned virus in the family Bunyaviridae. Yaba monkey tumor virus (YMTV) A Isolated from Aedes vigilax in New species in the genus Yatapoxvirus caus- South Wales, Australia. ing benign fibrous tumors of the head and limbs of rhesus and cynomolgus Yaounde virus (YAOV) A species in the monkeys, which may ulcerate before genus Flavivirus , a member of the regressing. First seen in captive rhe- Japanese encephalitis virus group. sus monkeys in 1958 in Yaba, Nigeria. Isolated from Culex nebulosus in Natural host probably an African pri- Cameroon. Has also been isolated in mate, with transmission by insect vec- the Central African Republic. tors in the wild. Workers in contact with Robin Y (1970) Annual Report Centre infected animals often become infected Collaborateur OMS de Reference pour les and have local disease, with fever in Arbovirus en Afrique de l’Ouest , Dakar, some cases. Experimental infection in Senegal humans causes a small nodule which regresses. Virus genome DNA is 145 kb Yaquina Head virus (YHV) A serotype of long; G C content 32.5%; density of Great Island virus in the genus Orbivirus , DNA 1.69 g/ml in CsCl. Inactivated belonging to the Great Island complex after 1 h at 56°C or by pH 3 at room tem- of the Kemerovo serogroup. Isolated perature. Replicates on the CAM and in from Ixodes uriae in Oregon, USA. The primary human kidney, Cercopithecus same or a closely related virus has kidney and MK2 cells. been isolated in Alaska. Not reported Rouhandeh H (1999) In Encyclopedia of to cause disease in humans. Virology , Second edition, edited by A Granoff and RG Webster. London: Academic Yata virus (YATAV) An unassigned ver- Press, p. 1971 tebrate rhabdovirus. Isolated from the mosquito, Mansonia uniformis, in the Yaba-1 virus (Y1V) A serotype of M’Poko Central African Republic. Not reported virus in the genus Orthobunyavirus. to cause disease in humans. Antigenically related to the Turlock virus serogroup. First isolated in Yatapoxvirus A genus in the fam- 1962 in Nigeria. A very similar virus, ily Poxviridae , containing two spe- Lednice virus, was isolated near the cies: Tanapox virus and Yaba monkey small town of Lednice in southern tumor virus. Virions are brick-shaped, Moravia from the tick, Culex modes- 300 250 200 nm. Genome DNA tus. Resistant to high pH. Replicates in about 146 kb long, G C content 32.5%. Yatapoxvirus 504

The two species differ according to Monath TP (2006) Bull Soc Pathol Exot 99 , 341 restriction enzyme analysis. Probably Monath TP (2007) Antiviral Res 78 , 116 maintained in the wild by insect transmission between various monkey yellow fever virus serogroup A group species. of nine serologically related viruses in the genus Flavivirus. They are: Banzi , Yellow fever virus (YFV) A species in the Boubai , Edge Hill , Jugra , Saboya , Sepik , genus Flavivirus. Jungle yellow fever is Uganda S , Wesselbron , and Yellow fever. an infection of wild primates in forests of Africa and South America. Yellow yellow head virus A strain of Gill- fever is endemic in Africa, south of the associated virus in the genus Okavirus. Sahara, and as far south as northern Isolated from the black tiger prawn Zimbabwe. Epidemics have occurred (Penaeus monodon ), the natural host, in Sudan and Ethiopia. Spreads occa- but many other prawn species can be sionally from South America to Central infected experimentally. Causes high America and Trinidad. In the African mortality in cultured prawn in Asia tree tops the virus is spread by the mos- and Australia. quito, Aedes africanus and A simpsoni , Sittidilokratna N et al (2002) Dis Aquat Org carrying infection from the monkeys 50 , 87 to humans in the villages. In South America Haemagogus sp are the main yellowtail ascites virus A birna- vectors in the sylvan cycle. In the urban virus isolated from cultured Seriola area Aedes aegypti is the vector carrying quinqueradiates. the human disease. The virus may be Nakajima K and Hara T (1985) Fish Pathol maintained in mosquitoes by trans- 19 , 231 ovarial transmission. Incubation period is 3–6 days. Infection in humans may be YLD Yaba-like disease. inapparent (in natives) or a fulminating, often fatal infection, with high fever, Yogue virus (YOGV) An unassigned albuminuria, jaundice, black vomit, and virus in the family Bunyaviridae. other hemorrhages. In children it may Isolated from a bat, Rousettus aegyptia- be difficult to diagnose. Macaque mon- cus, in Senegal. Related antigenically to keys, marmosets, and howler monkeys Kasokero virus. Not reported to cause develop an illness similar to humans, disease in humans. and may die after experimental inoculation. In most African primates there Yoka poxvirus (YKV) An unassigned is only viremia. The virus is fatal to virus in the family Poxviridae. Isolated hedgehogs. Replicates in cultures of from Aedes (Stegomyia) simpsoni in the chick and mouse embryo cells and Central African Republic. after adaptation will infect eggs. The Sureau P (1972) Institut Pasteur Bangui, attenuated 17D strain was obtained by Annual Report , 16 passage in chick embryo cells, and is produced in embryonated eggs; it gives Yokose virus (YOKV) A species in the few reactions when used as a vaccine. It genus Flavivirus, belonging to the gives protection for several years, and Ntaya virus serogroup. has been given to more than 200 million people. Urban yellow fever is best con- Yonban virus A virus related to TTV, trolled by elimination of Aedes aegypti. but phylogenetically distinct. Disease The endemic prevalence of dengue potential is unknown. and other related viruses may prevent Erker JC et al (1999) J Gen Virol 80 , 1743 spread to Asia, because immunity to dengue affords cross-protection against Yucaipa virus See Avian paramyxovirus 2 . yellow fever. Synonyms : fiebre amarilla virus; flavi- Yug Bogdanovac virus (YBV) A tenta- virus febricus. tive species in the genus Vesiculovirus. Chambers TJ et al (1990) Annu Rev Microbiol Isolated from Phlebotomus perfiliewi in 44 , 649 Serbia. Not reported to cause disease. Z

Z-DNA A form of DNA with a left- Zaysan virus A possible species in the handed (zig-zag) double helix and a genus Alphavirus, related to Semliki single groove. Specific Z-DNA-binding Forest virus. Isolated from mosquitoes proteins may control gene activity, since in the Far East of the former USSR. Z-DNA provides recognition signals in Slavik I et al (1976) Acta Virol ( Prague ) the regulation of gene transcription. 20, 177 Wang A et al (1979) Nature 282 , 680 Zegla virus (ZEGV) A species in the Zaire Ebola virus (ZEBOV) See Table Z1 . genus Orthobunyavirus . Isolated from the cotton rat, Sigmodon hispidus , in Zalcitabine (DDC) An antiviral drug, Almirante, Panama, also in Honduras, dideoxycytidine, which is active Guatemala, and Mexico. Not reported against HIV including some strains to cause disease in humans. which are resistant to AZT. Similar in potency to AZT. Main toxic side effect Zerit A trade name for stavudine, dide- is peripheral neuropathy. hydrodideoxyuridine (D4T), a thymi- Bozzette SA et al (1995) JAMA 273 , 295 dine kinase analog similar to AZT. Hitchcock MJM (1991) Antivir Chem Chemother Zaliv Terpeniya virus (ZTV) A strain of 2, 125 Uukuniemi virus in the genus Phlebovirus , belonging to the Uukuniemi virus sero- ZH548 virus A strain of Rift Valley fever group. Isolated from ticks, Ixodes putus , virus, isolated in Egypt. Developed as collected on Tyuleniy Island in Patience a candidate vaccine against Rift Valley Bay (Zaliv Terpeniya), Sakhalin region, fever. and Commodore Island, Kamchatka region in the former USSR where there Brown JL et al (1981) Infect Immun 33 , 848 are sea bird colonies. Pathogenic for suckling mice. Not reported to cause zidovudine See AZT (azidothymidine) . disease in humans. Zika virus (ZIKV) A species in the genus Lvov DK et al (1973) Arch Gesamte Virusforsch Flavivirus . Related antigenically to 41 , 165 Spondweni virus. Isolated from humans, Zanamivir An anti-influenza virus drug wild monkeys, and mosquitoes, Aedes that acts by inhibiting the virus neura- africanus , in Uganda, Nigeria, Central minidase. Can be used prophylactically African Republic, Senegal, and Malaysia. and is effective by oral inhalation. See Experimental infection of rhesus mon- Oseltamivir, Tamiflu . keys causes a fever. In humans there is a febrile illness with rash. Hayden FG et al (2000) N Engl J Med 343 , 1282 Fagbami AH (1979) J Hyg ( Camb ) 83 , 213 zinc finger A structural region in a protein formed by folding a polypeptide Table Z1. Strains of Zaire Ebola virus chain with a 28–30 amino acid repeating motif into a loop centered on a zinc Zaire Ebola virus Eckron ion. Found mainly in DNA-binding Zaire Ebola virus Gabon proteins and believed to be involved in Zaire Ebola virus Kikwit DNA-binding. Zaire Ebola virus Mayinga Zaire Ebola virus Tandala Zaire Ebola virus Zaire zinc salts Inhibit the cleavage of enterovirus proteins in vitro. This led to a clinical zinc salts 506

trial of zinc gluconate lozenges, but no Fiennes R (1967) Zoonoses of Primates . London: beneficial effects were found when pla- Weidenfeld and Nicolson, p. 2 cebo-controlled trials were done. zoonotic viruses Viruses transmitted Zinga virus An unclassified arbovirus. between animals and humans. Isolated from Mansonia africana and Aedes sp mosquitoes, and also from zoster See varicella-zoster virus 1 . humans. Antibodies found in birds, rodents, elephants, hogs, buffalo, zoster immunoglobulin (ZIG) Passive and hartebeest in the Central African immunization with immunoglobulin Republic. Causes a febrile illness in derived from donors with high anti- humans. VSV titers has been used to modify Zingilamo virus An unclassified arbo- infection in high-risk patients. The treat- virus. Antigenically related to Boteke ment is effective if administered within virus. Isolated from a bird, Bycanistes 2 days after contact with a disease case. sharpei, in the Central African Republic. Not reported to cause disease in zoster sineherpete A term proposed for humans. atypical reactivation of varicella-zoster virus in which there is neuropathic pain Zirqa virus (ZIRV) A serotype of Hughes but no cutaneous rash, and a typical virus in the genus Nairovirus . Isolated amnestic antibody response is found. from Ornithodoros muesebacki ticks col- Lewis GW (1958) BMJ 2 , 418 lected on Zirqa Island, Persian Gulf. Not reported to cause disease in humans. Zostrex Trade name for brivudin. zona Synonym for shingles. Zovirax Trade name for acycloguanosine. zoonosis A disease or an infection naturally transmitted between vertebrate Z-RNA A left-handed RNA double helix. animals and humans. However, the term has been frequently misunder- Hall K et al (1984) Nature 311 , 584 stood. See Fiennes (1967) for a discussion of the etymology of this term and Zwoegerziekte virus Synonym for Visna / the various interpretations which have maedi virus in the Netherlands. been placed upon it. Appendix

Source: Reproduced with permission from Virus Taxonomy: Eighth Report of the International Committee on Taxonomy of Viruses. CM Fauquet et al. (eds). London: Elsevier, 2005, p. 14. Appendix 508

Table of families and genera of viruses affecting vertebrates

Family Subfamily Genus Type species

DNA viruses – single-stranded DNA viruses

Circoviridae Circovirus Porcine circovirus 1 Gyrovirus Chicken anemia virus Unassigned genus Anellovirus Torque teno virus Parvoviridae Parvovirinae Parvovirus Minute virus of mice Erythrovirus Human parvovirus B19 Dependovirus Adeno-associated virus 2 Amdovirus Aleutian mink disease virus Bocavirus Bovine parvovirus DNA viruses – double-stranded DNA viruses

Poxviridae Chordopoxvirinae Orthopoxvirus Vaccinia virus Parapoxvirus Orf virus Avipoxvirus Fowlpox virus Capripoxvirus Sheeppox virus Leporipoxvirus Myxoma virus Suipoxvirus Swinepox virus Molluscipoxvirus Molluscum contagiosum virus Yatapoxvirus Yaba monkey tumor virus Asfarviridae Asfivirus African swine fever virus Iridoviridae Ranavirus Frog virus 3 Lymphocystivirus Lymphocystis disease virus 1 Megalocytivirus Infectious spleen and kidney necrosis virus Herpesviridae Alphaherpesvirinae Simplexvirus Human herpesvirus 1 Varicellovirus Human herpesvirus 3 Mardivirus Gallid herpesvirus 2 Iltovirus Gallid herpesvirus 1 Betaherpesvirinae Cytomegalovirus Human herpesvirus 5 Muromegalovirus Murid herpesvirus 1 Roseolovirus Human herpesvirus 6 Gammaherpesvirinae Lymphocryptovirus Human herpesvirus 4 Rhadinovirus Saimiriine herpesvirus 2 Unassigned genus Ictalurivirus Ictalurid herpesvirus 1 Adenoviridae Mastadenovirus Human adenovirus C Aviadenovirus Fowl adenovirus A Atadenovirus Ovine adenovirus D Siadenovirus Frog adenovirus Polyomaviridae Polyomavirus Simian virus 40 Papillomaviridae Alphapapillomavirus Human papillomavirus 32 Betapapillomavirus Human papillomavirus 5 Gammapapillomavirus Human papillomavirus 4 Deltapapillomavirus European elk papillomavirus Epsilonpapillomavirus Bovine papillomavirus 5 Zetapapillomavirus Equine papillomavirus 1 Etapapillomavirus Fringilla coelebs papillomavirus Thetapapillomavirus Psittacus erithacus tinneh papillomavirus Iotapapillomavirus Mastomys natalensis papillomavirus Kappapapillomavirus Cottontail rabbit papillomavirus Lambdapapillomavirus Canine oral papillomavirus 509 Appendix

Table of families and genera of viruses affecting vertebrates (continued)

Family Subfamily Genus Type species Mupapillomavirus Human papillomavirus 1 Nupapillomavirus Human papillomavirus 41 Xipapillomavirus Bovine papillomavirus 3 Omikronpapillomavirus Phocoena spinipinnis papillomavirus Pipapapillomavirus Hamster oral papillomavirus Unassigned Mimivirus Acanthamoeba polyphaga genus mimivirus

DNA and RNA transcribing viruses

Hepadnaviridae Orthohepadnavirus Hepatitis B virus Avihepadnavirus Duck hepatitis B virus Retroviridae Orthoretrovirinae Alpharetrovirus Avian leukosis virus Betaretrovirus Mouse mammary tumor virus Gammaretrovirus Murine leukemia virus Deltaretrovirus Bovine leukemia virus Epsilonretrovirus Walleye dermal sarcoma virus Lentivirus Human immunodeficiency virus 1 Spumavirinae Spumavirus Simian foamy virus

RNA viruses – dsRNA viruses

Reoviridae Orthoreovirus Mammalian orthoreovirus Orbivirus Bluetongue virus Rotavirus Rotavirus A Coltivirus Colorado tick fever virus Seadornavirus Banna virus Aquareovirus Aquareovirus A Birnaviridae Aquabirnavirus Infectious pancreatic necrosis virus Avibirnavirus Infectious bursal disease virus

RNA viruses – negative stranded ssRNA viruses

Bornaviridae Bornavirus Borna disease virus Rhabdoviridae Vesiculovirus Vesicular stomatitis Indiana virus Lyssavirus Rabies virus Ephemerovirus Bovine ephemeral fever virus Novirhabdovirus Infectious hematopoietic necrosis virus Filoviridae Marburgvirus Lake Victoria marburgvirus Ebolavirus Zaire ebolavirus Paramyxoviridae Paramyxovirinae Rubulavirus Mumps virus Avulavirus Newcastle disease virus Respirovirus Sendai virus Henipavirus Hendra virus Morbillivirus Measles virus Pneumovirinae Pneumovirus Human respiratory syncytial virus Metapneumovirus Avian metapneumovirus Orthomyxoviridae Influenzavirus A Influenza A virus Influenzavirus B Influenza B virus Appendix 510

Table of families and genera of viruses affecting vertebrates (continued)

Family Subfamily Genus Type species Influenzavirus C Influenza C virus Thogotovirus Thogoto virus Isavirus Infectious salmon anemia virus Bunyaviridae Orthobunyavirus Bunyamwera virus Hantavirus Hantaan virus Nairovirus Dugbe virus Phlebovirus Rift Valley fever virus Arenaviridae Arenavirus Lymphocytic choriomeningitis virus Unassigned genus Deltavirus Hepatitis delta virus

Positive-stranded ssRNA viruses

Picornaviridae Enterovirus Poliovirus Rhinovirus Human rhinovirus A Cardiovirus Encephalomyocarditis virus Aphthovirus Foot-and-mouth disease virus Hepatovirus Hepatitis A virus Parechovirus Human parechovirus Erbovirus Equine rhinitis B virus Kobuvirus Aichi virus Teschovirus Porcine teschovirus Caliciviridae Lagovirus Rabbit hemorrhagic disease virus Norovirus Norwalk virus Sapovirus Sapporo virus Vesivirus Vesicular exanthema of swine virus Unassigned genus Hepevirus Hepatitis E virus Astroviridae Aviastrovirus Turkey astrovirus Mamastrovirus Human astrovirus Nodaviridae Betanodavirus Striped jack nervous necrosis virus Coronaviridae Coronavirus Infectious bronchitis virus Torovirus Equine torovirus Arteriviridae Arterivirus Equine arteritis virus Flaviviridae Flavivirus Yellow fever virus Pestivirus Bovine viral diarrhea virus 1 Hepacivirus Hepatitis C virus Togaviridae Alphavirus Sindbis virus Rubivirus Rubella virus

Source: Adapted from Virus Taxonomy: Eighth Report of the International Committee on Taxonomy of Viruses. CM Fauquet et al. (eds). London: Elsevier, 2005. Abbreviations

A adenine IFA immunofl uorescent antibody ADP adenosine diphosphate IgA immunoglobulin A AMP adenosine monophosphate IgG immunoglobulin G araA adenosine arabinoside IgM immunoglobulin M araC cytosine arabinoside i.m. intramuscular ATP adenosine triphosphate i.p. intraperitoneal BHK baby hamster kidney IRES internal ribosome entry site bp base pair IUDR iododeoxyuridine C cytosine i.v. intravenous CAM chorioallantoic membrane kb kilobase CCC certifi ed cell culture kbp kilobase pair cccDNA covalently closed circular kDa kilodalton deoxyribonucleic acid LTR long terminal repeat CCL certifi ed cell line MCF mink cell focus-forming CD4 cell bearing a CD4 receptor MgCl2 magnesium chloride cDNA a deoxyribonucleic acid MHC major histocompatibility that is complementary to complex mRNA molecule ml milliliter CF complement fi xation mRNA messenger RNA CFT complement fi xation test mol. wt. molecular weight CMA cell membrane antigen NA neuraminidase CMP cytidine monophosphate NK natural killer CNS central nervous system nm nanometer CPE cytopathic effect nt nucleotide/nucleotides CsCl caesium chloride NTP nucleoside triphosphate CSF colony stimulating factor NTR non-translated region CTL cytotoxic T lymphocyte ORF open reading frame CTP cytidine triphosphate PCR polymerase chain reaction dA deoxyadenosine PM postmortem DNA deoxyribonucleic acid RNA ribonucleic acid dsRNA double-stranded rRNA ribosomal ribonucleic acid ribonucleic acid sp a species of dT deoxythmidine ssDNA single-stranded ECG electrocardiogram deoxyribonucleic acid ELISA enzyme linked immuno ssRNA single-stranded ribonucleic sorbent assay acid Fc receptor receptor for Fab C (fraction T thymine antibody-binding C) TMP thymidine monophosphate g gram tRNA transfer ribonucleic acid G guanine ts temperature sensitive GMP guanosine monophosphate U uracil GTP guanosine triphosphate UMP uridine monophosphate HA hemagglutinin UTP uridine triphosphate HAI hemagglutination inhibition UTPase uridine triphosphatase HAT hypoxanthine, aminopterin, UV ultraviolet and thymidine VPg genome-linked virus protein i.c. intracerebral wt wild type