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Chapter 21. Orthomyxoviridae

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Chapter 21. Orthomyxoviridae Chapter 21 Orthomyxoviridae Chapter Outline Properties of ORTHOMYXOVIRUSES 392 AVIAN INFLUENZA VIRUSES 403 Classification 392 Human Disease 407 Virion Properties 394 CANINE INFLUENZA VIRUSES 407 Virus Replication 395 BAT INFLUENZA VIRUSES 408 Molecular Determinants of Pathogenesis 397 BOVINE INFLUENZA D VIRUSES 408 MEMBERS OF THE GENUS INFLUENZAVIRUS A 398 HUMAN INFLUENZA VIRUSES 408 EQUINE INFLUENZA VIRUSES 398 MEMBERS OF THE GENUS ISAVIRUS 408 SWINE INFLUENZA VIRUSES 400 INFECTIOUS SALMON ANEMIA VIRUS 408 Human Disease 403 Other ORTHOMYXOVIRUSES 409 The family Orthomyxoviridae includes viruses with pandemic influenza viruses in wild and domestic animal genomes composed of several (six to eight) segments of species. Lastly, widespread use of sensitive and specific single-stranded RNA. The most important members of the RT-PCR assays that detect all influenza viruses has facili- family are the influenza viruses, which are included in four tated detailed surveillance without the need for virus iso- genera (Influenzavirus A, B, C, and D). Influenza viruses lation. Once a sample is identified as influenza A-positive that are pathogenic to wild and domestic animals and birds with this assay, the infecting virus rapidly can be further are included in the genus Influenzavirus A, whereas viruses characterized by gene-specific RT-PCR assays to deter- in the two other genera (B and C) circulate continuously in mine its hemagglutinin/neuraminidase subtype and cul- humans. Influenza A viruses infrequently are transmitted tured for subsequent antigenic analyses. from their animal hosts to humans, but human epidemics Although the original isolation of an influenza virus did and pandemics caused by influenza A viruses typically not occur until 1930 (from swine), associated diseases pre- have no animal involvement beyond the initial incursion. viously had been recognized in both animals and humans. Human influenza viruses sporadically are transmitted to Indeed, human influenza was described by Hippocrates swine, leading to establishment of virus lineages adapted to some 2400 years ago. Human pandemics have occurred this host species. Continuing surveillance, therefore, is throughout history, and the “Spanish flu” pandemic of 1918 essential to identify and detect influenza A virus variants was especially dramatic. The causative agent of highly that are capable of infecting humans while they are still pathogenic avian influenza (HPAI), “fowl plague,” was rec- confined either to their animal host or to a limited number ognized in the late 19th century as a filterable agent (ie, of human contacts. virus), but was not identified as an influenza virus until Recent developments have advanced significantly the 1955. High-pathogenicity avian influenza (HPAI) virus was understanding of the biology of influenza viruses. First, first isolated from wild birds in 1961—specifically, com- increased virologic surveillance and the rapid characteri- mon terns (Sterna hirundo) in South Africa—but until zation of viruses by sequencing have confirmed extensive recently this highly lethal virus has since been rarely genomic rearrangements between different influenza detected in wild birds. Low-pathogenicity avian influenza viruses. Secondly, the emergence of the highly pathogenic (LPAI) virus was first isolated from wild birds in 1972, and Eurasian H5N1 virus in Southeast Asia in 1997 and the such low-virulence viruses are common in aquatic species H1N1 virus pandemic of 2009 led to establishment of of wild birds. Aquatic birds (orders Anseriformes and worldwide surveillance programs to identify potentially Charadriiformes), especially ducks, shorebirds, and gulls, Fenner’s Veterinary Virology. DOI: http://dx.doi.org/10.1016/B978-0-12-800946-8.00021-0 © 2017 Elsevier Inc. All rights reserved. 389 390 PART | II Veterinary and Zoonotic Viruses are the essential reservoir hosts of low-pathogenicity influ- influenza A virus from wild aquatic birds into domestic enza A viruses (Table 21.1). Influenza viruses replicate in poultry occur much more frequently, but until recently the intestinal and upper respiratory epithelium of these birds most of these events went undetected as the immediate without producing overt disease, and are excreted in high consequences were limited. There have been only a very concentrations in feces and oral secretions. The viruses effi- limited number of outbreaks of highly pathogenic avian ciently are transmitted by the fecalÀoral route, and migrat- influenza in domestic poultry worldwide that resulted in ing aquatic birds carry viruses between their summer and high death losses or regulatory action to eradicate or man- winter habitats, which may span continents. Feeding stops age the infection, such as the recent epizootic of Eurasian along the flyways during the migrations provide further H5N1 HPAI virus infection. In addition, many human opportunity for spread of the viruses to resident contact infections with low-pathogenicity H7N9 avian influenza wild and domestic bird populations, and facilitate the con- virus reported by China in 2013 complicate surveillance tinuing process of evolution of these viruses. due to the lack of poultry mortality as a driver of diagnos- Cross-species infections occur sporadically between tic activities. Domestic swine are considered an important birds and mammals, including swine, horses, dogs, mink, intermediate (“bridge”) host in those areas of the world marine mammals, and humans (Fig. 21.1). Incursions of where there is frequent contact between poultry and TABLE 21.1 Hemagglutinin Subtype Distributiona Between Different Birds (Class: Aves) and Mammals (Class: Mammalia) HA Host of Origin Subtypec Mammalia Aves Humans Swine Equine Anseriformes Charadriiformes and Galliformes (eg, dabbling ducks) Procellariiformes (eg, shorebirds, (Domestic gulls, seabirds) Poultry) H1 11 11 1 1 11 e H2 (11 )b 61 1 1 H3 11 11 11 11 1111e H4 6111 1 H5 66 1 1 11b H6 61111 H7 66(11 )b 11 11b H8 66 H9 66 1 11 11 H10 6111 H11 1111 H12 11 6 H13 1111 H14d 6 H15d 66 H16 1 a 6, sporadic; 1, multiple reports;11, most common. b(), Previously common but now not reported. cBoth LP and HP viruses. dRare subtypes. ePrimarily swine influenza virus infections of domestic turkeys. From Swayne, D.E. (Ed.). Animal Influenza. Copyright r John Wiley and Sons (2009), with permission. Orthomyxoviridae Chapter | 21 391 FIGURE 21.1 Interspecies transmission of influenza A viruses. Diagrammatic representation of the source and movement of influenza A viruses or their genes within avian and mammalian ecological and epidemiological situations. H, hemagglutinin subtype; those in ( ) were previously common but no longer are in circulation. From Swayne, D.E. (Ed.), Avian Influenza, p. 62. Copyright r John Wiley & Sons (2009), with permission. swine, although the premise that swine are an essential and reassortment can complicate the interpretation of phy- intermediate host for the development of pandemic influ- logenetic trees. In contrast, analyses of the matrix protein enza virus strains has not been substantiated. Whole- gene (M) of field strains of influenza A virus show two genome sequencing of influenza viruses is now widely major avian lineages (North American and Eurasian), two used to reconstruct the phylogeny of each of the eight equine lineages, two gull lineages (North American and viral gene segments. For instance, comparison of the hem- European), two swine lineages (North American and agglutinin (HA) and neuraminidase (NA) genes shows Eurasian) and a human lineage. Analysis of the PB1 gene divergence into multiple subtypes (H1 through H16 and segregates human viruses between the North American N1 through N9, respectively) and host-specific lineages swine and Eurasian avian groups. Predictably, there are within some subtypes. However, interspecies transmission exceptions such as an outbreak of equine influenza in 392 PART | II Veterinary and Zoonotic Viruses China in 1989 that was caused by an H3N8 virus from a PROPERTIES OF ORTHOMYXOVIRUSES contemporary Eurasian avian source, whereas the classi- cal H3N8 equine lineage has its origin from a North Classification American avian lineage virus. Influenza A viruses from The family Orthomyxoviridae comprises the genera wild birds are very diverse, comprising the majority of Influenzavirus A, Influenzavirus B, Influenzavirus C, the viral gene pool, and all gene segments evolve con- Thogotovirus, Quaranjavirus, and Isavirus. The name of stantly. However, the virion surface genes accumulate the family is derived from the Greek myxa, meaning amino acid changes most frequently. Evolution is detected mucus, and orthos, meaning correct or right. The name not only in mammalian (eg, equine) and domestic poultry was intended to distinguish the orthomyxoviruses from viruses, but also in viruses from wild birds. the paramyxoviruses. Influenza is the Italian form of With renewed appreciation of the natural history of Latin, from influentia, “influence,” so used because epi- influenza viruses as “species jumpers,” prevention efforts demics were believed to be caused by astrological or logically will continue to focus on those situations in other occult influences. Influenza A viruses are common which high densities of birds and mammals are main- pathogens of horses, swine, humans, and domestic poultry tained in close proximity and in which there are rapid throughout much of the world, but they also are the cause turnover in populations, such as live-animal markets. The of sporadic or geographically limited
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