Cambridge University Press 978-0-521-82714-0 - Human Herpesviruses: Biology, Therapy, and Immunoprophylaxis Edited by Ann Arvin, Gabriella Campadelli-Fiume, Edward Mocarski, Patrick S. Moore, Bernard Roizman, Richard Whitley and Koichi Yamanishi Excerpt More information Part I Introduction: definition and classification of the human herpesviruses Edited by Bernard Roizman © Cambridge University Press www.cambridge.org Cambridge University Press 978-0-521-82714-0 - Human Herpesviruses: Biology, Therapy, and Immunoprophylaxis Edited by Ann Arvin, Gabriella Campadelli-Fiume, Edward Mocarski, Patrick S. Moore, Bernard Roizman, Richard Whitley and Koichi Yamanishi Excerpt More information 1 Overview of classification Andrew J. Davison MRC Virology Unit, Institute of Virology, Glasgow, UK Introduction and Iltovirus genera), Betaherpesvirinae (containing the Cytomegalovirus, Muromegalovirus and Roseolovirus gen- Taxonomy aimstostructure relationships among diverse era) and Gammaherpesvirinae (containing the Lym- organisms in order to provide abroader understanding of phocryptovirus and Rhadinovirus genera). In addition, Naturethan is affordedbyconsideration of organisms in there is agenus(Ictalurivirus) unattached to any subfamily isolation. Since biologicalsystems areshapedbyevolution, and a large number of species not assigned to genera. The which is not influencedbythe human desiretoimpose currentlist is giveninTable 1.1. All butone of the viruses order, any taxonomical scheme is bound to be incomplete assigned to taxa infect mammals or birds, although a sub- and to someextent arbitrary. The criteria appliedare ne- stantial number of unassigned herpesviruses havelower cessarily confined to what is technicallypossible, and thus vertebrate (reptilian, amphibian and fish) or invertebrate taxonomy has an importanthistoricalcomponent. In addi- (bivalve) hosts. tion,taxonomy developsconservatively,since striving for the ideal must betemperedbythe need to maintain utility. It is also an unfortunate fact thattaxonomy provides fer- Morphological criteria tile soilfordebate among a fewbut isoflittle interest to most. However, it is beyond dispute that the setting of her- The primary criterion forinclusion of an agent in the family pesviruses in a taxonomicalframework is vitalfor under- Herpesviridae isthatofvirionmorphology. The virionis standing the origins and behavior of thisfascinating family spherical, and comprises four major components: the core, of organisms. the capsid, the tegument and the envelope (see Chapter3). Historically,herpesvirustaxonomy has been addressed The diameter of the viriondependson the viralspecies, but since 1971 by the International Committee on Taxonomy is approximately200 nm. The coreconsists of a single copy of Viruses (ICTV)(Wildy, 1971).Aprovisional approach of a linear, double-strandedDNAmolecule packagedat to endowing herpesviruses with formal names (Roizman high densityinto the capsid. The capsid is an icosahedron, et al., 1973) was followedbygrouping into subfami- and has an external diameter of 125–130 nm. It consists of lies largely on the basisofbiologicalcriteria (Roizman 162 capsomeres, 12 of which are pentons and 150hexons, et al., 1981). This effort was rather successful, but not each containing five and six copies, respectively, of the free fromwhatturned out in hindsight to be a fewmis- major capsid protein. The capsomeres are joined via the classifications(Roizman et al., 1992).Further division of triplexes, each of which containstwocopies of one protein the subfamilies into genera utilizedmolecular data to a and onecopy of another. The tegument, which surrounds greater extentthan before, primarilyinrelation to genome the capsid,contains perhaps 30ormore viral protein characteristics such assize and structure(Roizman et species and is poorlydefined structurally. In the tegument, al., 1992). In the latest report of the ICTV Herpesvirus structures positionedwith symmetry corresponding to Study Group (Davison et al., 2005), the family Herpesviri- that of the capsid are detectable onlyinthe region close to dae consists of three subfamilies: Alphaherpesvirinae the capsid. The lipid envelopesurrounds the exterior of the (containing the Simplexvirus, Varicellovirus, Mardivirus tegument, and isstuddedwith atleast ten viral membrane 3 © Cambridge University Press www.cambridge.org Cambridge University Press 978-0-521-82714-0 - Human Herpesviruses: Biology, Therapy, and Immunoprophylaxis Edited by Ann Arvin, Gabriella Campadelli-Fiume, Edward Mocarski, Patrick S. Moore, Bernard Roizman, Richard Whitley and Koichi Yamanishi Excerpt More information 4 A. J. Davison Table 1.1. Herpesvirustaxonomy and nomenclature Formal namea Abbrev. Common nameb Abbrev.c Subfamily Alphaherpesvirinae Genus Simplexvirus Atelineherpesvirus 1AtHV-1 Spidermonkey herpesvirus Bovineherpesvirus 2BoHV-2Bovine mamillitis virus Cercopithecineherpesvirus 1CeHV-1B-virusHVB Cercopithecineherpesvirus 2CeHV-2 SA8 virus Cercopithecineherpesvirus 16 CeHV-16 Herpesvirus papio 2 Human herpesvirus 1 HHV-1 Herpes simplex virus[type] 1 HSV-1 Human herpesvirus 2 HHV-2 Herpes simplex virus[type] 2 HSV-2 Macropodid herpesvirus 1 MaHV-1 Parma wallaby herpesvirus Macropodid herpesvirus 2 MaHV-2Dorcopsis wallaby herpesvirus Saimiriineherpesvirus 1SaHV-1 Herpesvirustamarinus Genus Varicellovirus Bovineherpesvirus 1BoHV-1 Infectious bovine rhinotracheitis virus BHV-1 Bovineherpesvirus 5BoHV-5Bovineencephalitis virus BHV-5 Bubalineherpesvirus 1BuHV-1Waterbuffalo herpesvirus Canid herpesvirus 1CaHV-1 Canineherpesvirus Caprineherpesvirus 1CpHV-1Goatherpesvirus Cercopithecineherpesvirus 9CeHV-9 Simian varicella virus SVV Cervid herpesvirus 1CvHV-1Reddeer herpesvirus Cervid herpesvirus 2CvHV-2Reindeer herpesvirus Equid herpesvirus 1EHV-1Equine abortion virus Equid herpesvirus 3EHV-3Equinecoitalexanthema virus Equid herpesvirus4 EHV-4 Equine rhinopneumonitis virus Equid herpesvirus 8EHV-8Asinineherpesvirus 3 Equid herpesvirus 9EHV-9Gazelle herpesvirus Felid herpesvirus 1FeHV-1Feline rhinotracheitis virus Human herpesvirus 3 HHV-3 Varicella-zoster virus VZV Phocid herpesvirus 1 PhoHV-1 Harbour sealherpesvirus Suid herpesvirus 1SuHV-1 Pseudorabies virusPRV Tentative species in genus Varicellovirus Equid herpesvirus 6EHV-6Asinineherpesvirus 1 Genus Mardivirus Gallid herpesvirus 2 GaHV-2 Marek’s disease virustype 1 MDV-1 Gallid herpesvirus 3GaHV-3 Marek’s disease virustype 2 MDV-2 Meleagrid herpesvirus 1 MeHV-1 Turkey herpesvirusHVT Genus Iltovirus Gallid herpesvirus 1 GaHV-1 Infectiouslaryngotracheitis virusILTV Unassigned species in subfamily Alphaherpesvirinae Psittacid herpesvirus 1 PsHV-1 Parrot herpesvirus Subfamily Betaherpesvirinae Genus Cytomegalovirus Cercopithecineherpesvirus 5CeHV-5African greenmonkey cytomegalovirus SCMV Cercopithecineherpesvirus 8CeHV-8Rhesus monkey cytomegalovirus RhCMV Human herpesvirus 5 HHV-5 Human cytomegalovirusHCMV Pongineherpesvirus4 PoHV-4 Chimpanzee cytomegalovirus CCMV Tentative species in genus Cytomegalovirus Aotineherpesvirus 1AoHV-1 Herpesvirus aotus 1 Aotineherpesvirus 3AoHV-3 Herpesvirus aotus 3 © Cambridge University Press www.cambridge.org Cambridge University Press 978-0-521-82714-0 - Human Herpesviruses: Biology, Therapy, and Immunoprophylaxis Edited by Ann Arvin, Gabriella Campadelli-Fiume, Edward Mocarski, Patrick S. Moore, Bernard Roizman, Richard Whitley and Koichi Yamanishi Excerpt More information Overview of classification 5 Table 1.1. (cont.) Formal namea Abbrev. Common nameb Abbrev.c Genus Muromegalovirus Murid herpesvirus 1 MuHV-1 Mouse cytomegalovirusMCMV Murid herpesvirus 2 MuHV-2Ratcytomegalovirus RCMV Genus Roseolovirus Human herpesvirus 6 HHV-6 Human herpesvirus 7 HHV-7 Unassigned species in subfamily Betaherpesvirinae Caviid herpesvirus 2CavHV-2 Guinea pig cytomegalovirus GPCMV Tupaiid herpesvirus 1 TuHV-1 Tree shrew herpesvirusTHV Subfamily Gammaherpesvirinae Genus Lymphocryptovirus Callitrichineherpesvirus 3CalHV-3 Marmoset lymphocryptovirusMarmoset LCV Cercopithecineherpesvirus 12 CeHV-12 Herpesvirus papio Cercopithecineherpesvirus 14 CeHV-14 African greenmonkeyEBV-like virus Cercopithecineherpesvirus 15 CeHV-15 Rhesuslymphocryptovirus Rhesus LCV Human herpesvirus 4 HHV-4 Epstein-Barr virus EBV Pongineherpesvirus 1 PoHV-1 Herpesvirus pan Pongineherpesvirus 2 PoHV-2 Orangutan herpesvirus Pongineherpesvirus 3 PoHV-3Gorilla herpesvirus Genus Rhadinovirus Alcelaphineherpesvirus 1AlHV-1 Malignantcatarrhalfever virus AHV-1 Alcelaphineherpesvirus 2AlHV-2 Hartebeest malignantcatarrhalfever virus Atelineherpesvirus 2AtHV-2 Herpesvirus ateles HVA Bovineherpesvirus4 BoHV-4 Movar virus BHV-4 Cercopithecineherpesvirus 17 CeHV-17 Rhesus rhadinovirus RRV Equid herpesvirus 2EHV-2 Equid herpesvirus 5EHV-5 Equid herpesvirus 7EHV-7Asinineherpesvirus 2 Hippotragineherpesvirus 1 HiHV-1Roan antelopeherpesvirus Human herpesvirus 8 HHV-8 Kaposi’ssarcoma-associated herpesvirusKSHV Murid herpesvirus4 MuHV-4 Murine gammaherpesvirus 68 MHV-68 Mustelid herpesvirus 1 MusHV-1 Badger herpesvirus Ovineherpesvirus 2 OvHV-2Sheep-associatedmalignantcatarrhal fever virus Saimiriine herpesvirus 2 SaHV-2 Herpesvirussaimiri HVS Tentative species in genus Rhadinovirus Leporid herpesvirus 1LeHV-1Cottontail rabbitherpesvirus Leporid herpesvirus 2LeHV-2 Herpesviruscuniculi Leporid herpesvirus 3LeHV-3 Herpesvirussylvilagus Marmodid herpesvirus 1 MarHV-1Woodchuck herpesvirus Unassigned species in subfamily Gammaherpesvirinae Callitrichineherpesvirus 1CalHV-1 Herpesvirussaguinus Unassigned genus Ictalurivirus in family Herpesviridae Ictalurid