Fenner’s Veterinary Virology

570 Pages·2016·83.01 MB·English

by  N. James Maclachlan

Fenner’s Veterinary Virology
Fenner’s Veterinary Virology
Fifth Edition
Edited by Associate Editors:
N. James MacLachlan, Stephen W. Barthold, DVM, PhD,
BVSc, PhD, Dip ACVP Dip ACVP
Distinguished Professor Distinguished Professor Emeritus
Department of Pathology, Department of Pathology, Microbiology and
Microbiology and Immunology Immunology
School of Veterinary Medicine School of Veterinary Medicine
University of California,
University of California
Davis, California, USA
Davis, California, USA
and David E. Swayne, DVM, PhD,
Dip ACVP, Dip ACPV
Extraordinary Professor
Department of Veterinary Tropical Diseases
Center Director
Faculty of Veterinary Science
USDA/Agricultural Research Services
University of Pretoria
Exotic and Emerging Avian Viral Diseases
Onderstepoort, Republic of South Africa
Research Unit
Southeast Poultry Research Laboratory
Edward J. Dubovi,
Athens, Georgia, USA
MS, PhD, Dip ACVM (Hon)
James R. Winton, BA, PhD
Director, Virology Section
Animal Health Diagnostic Center Chief, Fish Health Section
Department of Population Medicine United States Geological Survey
and Diagnostic Sciences Western Fisheries Research Center
College of Veterinary Medicine Seattle, Washington, USA
Cornell University
Ithaca, New York, USA
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Dedication
We recognize the remarkable accomplishment that was the global eradication
of rinderpest, long a scourge of animal production systems in much of the
world. The global eradication of rinderpest in 2011 represents only the second
time a pathogen has been eradicated from the planet through concerted human
effort, and serves as an inspirational example of the constructive potential
benefits of science to the entire global community. The collaborative efforts of
academic institutions and governments along with animal health experts and
vaccine delivery teams will hopefully provide the model for future disease
eradication efforts.
List of Contributors
Udeni B.R. Balasuriya, BVSc, MS, PhD, Professor of Ruben O. Donis, MV, PhD, Deputy Director, Influenza
Virology, Maxwell H. Gluck Equine Research Center, Division, Biomedical Advanced Research and
Department of Veterinary Science, University of Development Authority (BARDA), Assistant Secretary
Kentucky,Lexington,Kentucky,USA for Preparedness and Response, Department of Health
ArteriviridaeandRoniviridae;Togaviridae;Flaviviridae andHumanServices,WashingtonDC,USA
Orthomyxoviridae
Simon Barratt-Boyes, BVSc, PhD, Professor,
Department of Infectious Disease & Microbiology, Ian Gardner, BVSc, MPVM, PhD, Professor and
Center for Vaccine Research, University of Pittsburgh, Canada Research Excellence Chair, Department of
Pittsburgh,Pennsylvania,USA Health Management, Atlantic Veterinary College,
AntiviralImmunityandVirusVaccines University of Prince Edward Island, Charlottetown,
PrinceEdwardIsland,Canada
Martin Beer, PhD, Institute of Diagnostic Virology,
EpidemiologyandControlofViralDiseases
Friedrich-Loeffler-Institut, Federal Research Institute
forAnimalHealth,Su¨dufer,Greifswald,Germany James Gilkerson, BVSc, BSc(Vet), PhD, Professor,
Bunyaviridae Faculty of Veterinary and Agricultural Science, The
UniversityofMelbourne,Melbourne,Victoria,Australia
Brian Bird, DVM, MSPH, PhD, Veterinary Medical Picornaviridae
Officer, Viral Special Pathogens Branch, Centers for
DiseaseControlandPrevention,Atlanta,Georgia,USA William T. Golde, PhD, Senior Scientist, Department of
Arenaviridae Immunology, Plum Island Animal Disease Center,
Agricultural Research Service, USDA, Orient Point,
JoeBrownlie,BVSc,PhD,FRCVS,DipECVP, Emeritus USA
Professor of Veterinary Pathology, Pathology and AntiviralImmunityandVirusVaccines
Pathogen Biology, Royal Veterinary College, London,
Carol Hartley, BSc(Hons), PhD, Senior Research
Hertfordshire,UK
Fellow, Faculty of Veterinary and Agricultural Science,
Coronaviridae
The University of Melbourne, Melbourne, Victoria,
LarkL.Coffey,PhD, AssistantProfessor,Departmentof Australia
Pathology, Microbiology and Immunology, School of Picornaviridae
Veterinary Medicine, University of California, Davis,
Hans Heidner, BS, MS, PhD, Professor, Department of
California,USA
Biology, University of Texas at San Antonio, San
TheNatureofViruses
Antonio,Texas,USA
John M. Cullen, VMD, PhD, Dip ACVP, Professor, VirusReplication
Population Health and Pathobiology, North Carolina
State University College of Veterinary Medicine, Christine Herden, DVM, PhD, Dipl ECVP, Professor,
Raleigh,NorthCarolina,USA Institute for Veterinary Pathology, Justus-Liebig-
OtherViruses:Hepeviridae,Hepadnaviridae, Universita¨tGiessen,Giessen,Germany
Delataviruses,Nodaviridae,andUnclassifiedViruses Bornaviridae
Gustavo A. Delhon, DVM, MS, PhD, Associate Peter Kirkland, BVSc, PhD, Senior Principal Research
Professor,SchoolofVeterinaryMedicine&Biomedical Scientist, Virology Laboratory, Elizabeth Macarthur
Science, University of Nebraska-Lincoln, Lincoln, Agriculture Institute, Menangle, New South Wales,
Nebraska,USA Australia
Poxviridae Bunyaviridae
xvii
xviii ListofContributors
Donald P. Knowles, DVM, PhD, Dip ACVP, Research John Parker, BVMS, PhD, Associate Professor, Baker
Leader, USDA/Agricultural Research Services, Animal InstituteforAnimalHealth,DepartmentofMicrobiology
Diseases Research Unit, Professor, Department of and Immunology, College of Veterinary Medicine,
Veterinary Microbiology and Pathology, College of CornellUniversity,Ithaca,NewYork,USA
Veterinary Medicine, Washington State University, CaliciviridaeandAstroviridae
Pullman,Washington,USA
Rhabdoviridae Colin R. Parrish, PhD, Professor of Virology, Baker
InstituteforAnimalHealth,DepartmentofMicrobiology
Xiang-Jin Meng, MD, MS, PhD, University
and Immunology, College of Veterinary Medicine,
Distinguished Professor, Department of Biomedical
CornellUniversity,Ithaca,NewYork,USA
Sciences and Pathobiology, Virginia-Maryland College Parvoviridae
ofVeterinaryMedicine,Blacksburg,Virginia,USA
CircoviridaeandAnelloviridae;OtherViruses:
Patricia Pesavento, DVM, PhD, Dip ACVP, Professor,
Hepeviridae,Hepadnaviridae,Delataviruses,
Department of Pathology, Microbiology and
Nodaviridae,andUnclassifiedViruses
Immunology, School of Veterinary Medicine,
John Munday, BVSc, PhD, Dip ACVP, Associate UniversityofCalifornia,Davis,California,USA
Professor, Department of Pathobiology, Institute of PapillomaviridaeandPolyomaviridae;Caliciviridae
Veterinary, Animal and Biomedical Sciences, Massey andAstroviridae
University,PalmerstonNorth,NewZealand
PapillomaviridaeandPolyomaviridae William Reisen, BS, MS, PhD, Emeritus Professor,
Center for Vectorborne Diseases, Department of
Brian Murphy, DVM, PhD, Dip ACVP, Associate Pathology, Microbiology and Immunology, School of
Professor, Department of Pathology, Microbiology and Veterinary Medicine, University of California, Davis,
Immunology,SchoolofVeterinaryMedicine,University California,USA
ofCalifornia,Davis,California,USA Togaviridae;Flaviviridae
Retroviridae
Juergen A. Richt, DVM, PhD, Regents Distinguished
Stefan Niewiesk, DVM, PhD, Dip ACVP, Professor,
Professor, Department of Diagnostic Medicine/
Department of Veterinary Biosciences, The Ohio State
Pathobiology, College of Veterinary Medicine, Kansas
University,Columbus,Ohio,USA
StateUniversity,Manhattan,Kansas,USA
PathogenesisofViralInfectionsandDiseases
Bornaviridae
Michael Oglesbee, DVM, PhD, Dip ACVP, Professor
Christina J. Sigurdson, DVM, PhD, Dip
and Chair, Department of Veterinary Biosciences, The
ACVP, Associate Professor, Department of Pathology,
OhioStateUniversity,Columbus,Ohio,USA
University of California, San Diego, School of
PathogenesisofViralInfectionsandDiseases
Medicine,LaJolla,California,USA
Prions:AgentsofTransmissibleSpongiform
Klaus Osterrieder, PhD, Professor, Managing Director,
Encephalopathies
Department of Veterinary Medicine, Institute of
Virology,FreieUniversita¨tBerlin,Berlin,Germany
Jonathan Towner, PhD, Lead, Virus Host Ecology
Herpesvirales
Section, Viral Special Pathogens Branch, Centers for
DiseaseControlandPrevention,Atlanta,Georgia,USA
Christopher Oura, BVetMed, MSc, PhD, Professor of
Filoviridae
Veterinary Virology, Department of Basic Veterinary
Sciences, The School of Veterinary Medicine, The
Veronika von Messling, DVM, PhD, Director and
University of the West Indies, St Augustine, Trinidad
andTobago Professor, Veterinary Medicine Division, Paul-Ehrlich-
AsfarviridaeandIridoviridae Institut,Langen,Germany
ParamyxoviridaeandPneumoviridae
Massimo Palmarini, DVM, PhD, Professor, MRC-
University of Glasgow Centre for Virus Research, Gary Whittaker, BSc, PhD, Professor, Department of
Institute of Infection, Immunity and Inflammation, Microbiology and Immunology, College of Veterinary
Glasgow,UK Medicine,CornellUniversity,Ithaca,NewYork,USA
Reoviridae Coronaviridae
Acknowledgements
Murphy. The goal of this fifth edition of Veterinary
Virology is to continue the agenda we began in the
fourth edition, specifically to bring all aspects of
veterinary and zoonotic virus diseases under a common
umbrella. To that end, we gratefully acknowledge the
Associate Editors of this text who contributed their spe-
cialist expertise on virus diseases of laboratory animals,
fish and aquatic species, and birds. We also gratefully
acknowledgethe34contributorswhoacceptedtheonerous
task of updating individual chapters. Lastly, we are espe-
cially grateful to those individuals who, without recogni-
In 1985, Frank Fenner (1914(cid:1)2010) had the idea to
tion,agreedtoproofvariouschaptersandsectionsthereof,
writeaveterinaryfocusedcompaniontothebook,Medical
notably Drs.Kimberly Dodd,Kirsten Murphy, Linda Saif,
Virology,whichheandDavidO.White(1931(cid:1)2004)had
andDonalO’Toole,alongwiththosewhoprovidedfigures
co-authoredin1970and1976.Thus,in1987,thefirstedi-
includingDr.KevinKeelwhocontributedthepictureused
tion of Veterinary Virology was published, with subse-
onthecover.
quent editionsin1993 and 1999.Other contributorstothe
Asinprioreditions,weacknowledgeourfamilies,tea-
firstthreeeditionsofthetextincludedPeterA.Bachmann
chers, mentors and students for their inspiration and
(1939(cid:1)1985),RudolfRott(1926(cid:1)2003),E.PaulJ.Gibbs,
direction.
Michael J.Studdert,MarianC.Horzinek andFrederickA.
xix
Chapter 1
The Nature of Viruses
ChapterOutline
Introduction:ABriefHistoryofAnimalVirology 3 ViralMembraneLipids 10
CharacteristicsofViruses 7 ViralMorphology 10
ChemicalCompositionoftheVirion 8 ViralTaxonomy 13
ViralNucleicAcidsintheVirion 8 PhylogeneticComparisonofVirusSequences 16
ViralProteinsintheVirion 9
INTRODUCTION: A BRIEF HISTORY OF early studies provided the essential operational definition
of viruses as filterable agents. Chemical and physical
ANIMAL VIROLOGY
studies revealed the structural basis of viruses nearly 40
The history of human development has been shaped by at yearslater.
least three major recurring elements: (1) environmental In the early 20th century, use of the filtration criteria
changes; (2) human conflicts; (3) infectious diseases. led to the association of many acute animal diseases with
Infectious diseases have impacted both humans and our what were later defined as viral infections: African horse
food supply. The origins of veterinary medicine are sickness, fowl plague (high pathogenicity avian influenza),
rooted in efforts to maintain the health of animals for rabies, canine distemper, equine infectious anemia, rinder-
foodandfiberproduction,andanimals essential forwork- pest,andclassicalswinefever(hogcholera)(Table1.1).In
related activities. Control ofanimal disease outbreaks was 1911, Rous discovered the first virus that could produce
not possible until the pioneering work of the late 19th neoplasia (tumors), and for this discovery he was awarded
century that linked microbes to specific diseases of plants a Nobel Prize. This early phase of virology was fraught
and animals (see Murphy, F.A., 2012. The Foundations of with skepticism and uncertainty because of the limited
Virology, for detailed discussion). Many attribute the tools available to study and characterize the filterable
beginning of virology with the work of Ivanofsky and agents. Experiments using filters with varying retention
Beijerinck on the transmission of tobacco mosaic virus. parameters demonstrated the existence of filterable agents
Both scientists were able to demonstrate transmission of of different sizes. Some agents were inactivated with
the agent causing disease in tobacco plants through filters organic solvents, whereas others were resistant. For equine
that retained bacteria. Beijerinck also noted that the filter- infectious anemia, the acute and chronic forms of the dis-
able agent could regain its “strength” from diluted mate- easewere perplexingand anunresolved conundrum.These
rial, but only if it were put back into the tobacco plants. types of apparent inconsistencies made it difficult to estab-
The concept of a replicating entity rather than a chemical lish a unifying conceptual description of the filterable
or toxin had its genesis with these astute observations. agents. For research on animal diseases, early workers
The era of veterinary virology had its beginning virtually were restricted to using animal inoculation in order to
at the same time as Beijerinck was characterizing tobacco assess the impact of a treatment on any putative disease-
mosaic virus transmission. Loeffler and Frosch applied causing agent. The logistics could be especially daunting
the filtration criteria to a disease in cattle that later would forstudiesincattleandhorses.Helpinprovidingdefinition
be known as foot and mouth disease. Repeated passage of tofilterable agents came fromthe discovery ofviruses that
the filtrate into susceptible animals with the reproduction infectedbacteria.Twortin1915detectedtheexistenceofa
of acute disease firmly established the “contagious” filterable agent that could kill bacteria. Like its plant and
nature of the filtrate and provided more evidence for a animal counterparts, the strength of a dilute solution of the
process that was inconsistent with toxic substances. These bacterial virus could be regained by inoculating new
Fenner’sVeterinaryVirology.DOI:http://dx.doi.org/10.1016/B978-0-12-800946-8.00001-5
©2017ElsevierInc.Allrightsreserved. 3
4 PART | I ThePrinciplesofVeterinaryandZoonoticVirology
TABLE1.1 SelectedMomentsintheHistoryofVirology
Year Investigator(s) Event Year Investigator(s) Event
1892 Ivanofsky Identificationoftobacco 1939 EllisandDelbruck One-stepgrowthcurve—
mosaicvirusasfilterable bacteriophage
agent
1946 Olafson,MacCallum,andFox Bovineviraldiarrheavirus
1898 LoefflerandFrosch Foot-and-mouthdisease
1948 Sanford,Earle,andLikely Cultureofisolated
causedbyfilterableagent
mammaliancells
1898 Sanarelli Myxomavirus
1952 DulbeccoandVogt Plaqueassayforfirstanimal
1900 Reed Yellowfevervirus virus—poliovirus
1900 McfadyeanandTheiler Africanhorsesicknessvirus 1956 Madin,York,andMcKercher Isolationofbovine
herpesvirus1
1901 Centanni,Lode,andGruber Fowlplaguevirus(avian
influenzavirus) 1957 IsaacsandLindemann Discoveryofinterferon
1902 NicolleandAdil-Bey Rinderpestvirus 1958 HorneandBrenner Developmentofnegative-
stainelectronmicroscopy
1902 SpruellandTheiler Bluetonguevirus
1961 Becker Firstisolationofavian
1902 Aujeszky Pseudorabiesvirus
influenzavirusfromwild
1903 RemlingerandRiffat-Bay Rabiesvirus birdreservoir
1903 DeSchweinitzandDorset Hogcholeravirus(classical 1963 PlummerandWaterson Equineabortionvirus 5
swinefevervirus) herpesvirus
1904 Carre´andValle´e Equineinfectiousanemia 1970 TeminandBaltimore Discoveryofreverse
virus transcriptase
1905 Spreull Insecttransmissionof 1978 Carmichael,Appel,andScott Canineparvovirus2
bluetonguevirus
1979 WorldHealthOrganization WHOdeclaressmallpox
1905 Carre´ Caninedistempervirus eradicated
1908 EllermannandBang Avianleukemiavirus 1981 Pedersen Felinecoronavirus
1909 LandsteinerandPopper Poliovirus 1981 Baltimore Firstinfectiouscloneofan
RNAvirus
1911 Rous Roussarcomavirus—first
tumorvirus 1983 Montagnier,Barre-Sinoussi, Discoveryofhuman
andGallo immunodeficiencyvirus
1915 Twortandd’Herelle Bacterialviruses
1987 Pedersen Felineimmunodeficiency
1917 d’Herelle Developmentoftheplaque virus
assay
1991 WensvoortandTerpstra Isolationofporcine
1927 Doyle Newcastlediseasevirus reproductiveandrespiratory
syndromevirus(PRRSV)
1928 VergeandChristofornoni Felineparvovirus(feline
SeifriedandKrembs panleukopeniavirus) 1994 Murray Hendravirusisolated
1930 Green Foxencephalitis(canine 1999 WestNilevirusentersNorth
adenovirus1) America
1931 Shope Swineinfluenzavirus 2002 Severeacuterespiratory
syndromeoutbreak
1931 WoodruffandGoodpasture Embryonatedeggsforvirus
propagation 2005 Palase,Garcia-Sastre, Reconstructionofthe1918
Tumpey,andTaubenberger pandemicinfluenzavirus
1933 DimmockandEdwards Viraletiologyforequine
abortions 2008 Developmentofmolecular
toolsandcomputersoftware
1933 Andrewes,Laidlaw,and Firstisolationofhuman
for“nextgeneration
Smith influenzavirus
sequencing”and
1933 Shope Swinenaturalhostof metagenomicanalyses
pseudorabies
2011 WorldOrganisationof Declarationoftheglobal
1933 BushnellandBrandly Avianbronchitisvirus AnimalHealth(OIE) eradicationofrinderpest
1935 Stanley Tobaccomosaicvirus(TMV) 2012 RecognitionofMiddleEast
crystallized;proteinnature respiratorysyndrome
ofvirusesconfirmed
2014 ReemergenceofEbolain
1938 Kausche,Ankuch,andRuska Firstelectronmicroscopy WestAfrica
pictures—TMV
TheNatureofViruses Chapter | 1 5
cultures of bacteria. Felix d’Herelle also noted the killing a human cell line was developed, and growth of poliovi-
of bacteria by an agent that he called “bacteriophage.” He rus in a nonneuronal cell was demonstrated. These
defined the plaque assay for quantitating bacteriophage, a advances all permitted the development of a plaque assay
techniquetoenumeratevirusparticlesbasedontheirability for poliovirus 35 years after the concept was defined for
to kill cultured cells and therefore produce holes, or bacteriophage. Basic studies on animal viruses that were
plaques in the cell layer that became a keystone for defin- hindered by the necessity to work in animal systems were
ingthepropertiesofviruses. now possible in vitro, and the principles established for
The initial studies on tobacco mosaic virus led to fur- bacteriophage could be explored for animal viruses. The
ther understanding of“filterable agents”—namely viruses. cellcultureeraofanimal virologyhadbegun.
Specifically, the high concentration of virus produced in The advances in virology driven by human disease
infected tobacco plants permitted the chemical and physi- control efforts were directly applicable to animal virol-
cal characterization of the infectious material. By the ogy. Bovine viral diarrhea virus was identified as a new
early 1930s, there was evidence that the agent infecting disease-causing agent in cattle in 1946 and by the late
tobacco plants was composed of protein, and that antibo- 1950s was considered the most economically important
dies produced in rabbits could neutralize the virus. The disease of cattle in the United States. Cell culture proce-
tobacco mosaic virus was crystallized in 1935, and in dures permitted isolation of the virus and the production
1939 was the first virus to be viewed using an electron of a vaccine by the early 1960s. Influenza virus was
microscope. The particulate nature of viruses was now an detectedfor the first time inwild birdsin1961, which led
established fact. A further advance in animal virology to the identification of water fowl and shore birds as the
was the use of embryonated eggs for culturing virus in natural reservoir of influenza A viruses. An apparent
1931. In the same year, Shope identified influenza virus cross-species incursion of a feline parvovirus variant pro-
in swine; in 1933, influenza virus was isolated from duced theworldwideepizootic ofcanine parvovirusinthe
human cases. The identification of the strain H1N1 of late 1970s. Again, standard in vitro cell culture proce-
influenza virus in swine might be considered the first dures identified the new agent and soon enabled the pro-
comprehensive description of an “emerging” disease in duction of an effective vaccine. The entire arterivirus
animals—that is, a virus crossing a species barrier and family (Arteriviridae) was identified in the cell culture
maintaining itself as an agent of disease in the new spe- era of virology—specifically, equine arteritis virus
cies. In an attempt to move away from large-animal (1953),lactatedehydrogenase-elevatingvirus(1960),sim-
experimentation, andtoprovidemodelsystemsforhuman ian hemorrhagic fever virus (1964), porcine reproductive
diseases such as influenza, mice and rats became impor- and respiratory syndrome virus (1991), and most recently,
tant tools for studying animal viruses. These advances wobbly possum virus (2012). The discovery of human
spawned the birth of laboratory animal medicine pro- immunodeficiency virus (HIV) in 1983 attracted global
gramsthathavebecomeanessentialbackboneofbiomed- attention, but the identification of simian immunodefi-
icalresearch. ciency virus (SIV) shortly thereafter may ultimately be of
The decade 1938(cid:1)48 saw major advances by Ellis, equal importance to the eventual control of human HIV
Delbruck, and Luria in the use of bacteriophage to probe infection. The primate system has provided the animal
the mechanism ofinheritance of phenotypic traits of these models for studies of pathogenesis and vaccine develop-
bacterial viruses. Advances in understanding the proper- ment. Genetic analyses established that HIV-1 and HIV-2
ties of viruses progressed much more rapidly with bacte- were closely relatedtothe SIVspresent inOld World pri-
rial viruses, because the work could be done in artificial mates, and that they were independently derived via
media, without any requirement for laborious and time- cross-speciestransmissionofthesesimianviruses.
consuming propagation of viruses in either animals or The beginnings of the molecular era of virology date
plants. A key concept in virus replication, namely the to the late 1970s and early 1980s. Although not specifi-
latent period, was defined using one-step growth curve cally designed for viruses, the development of the poly-
experiments with bacteriophage (see Chapter 2: Virus merase chain reaction (PCR) in 1983 had a profound
Replication). This observation of the loss of infectivity impact on virus research. Cloning of nucleic acid
for a period after the initiation of the infection directed sequences led to the first infectious molecular clone of a
research to define the mode of replication of viruses as virus (poliovirus) in 1981. The impact of molecular tech-
totallydistinctfromthatofallotherreplicatingentities. niques on virus detection and diagnostics was demon-
Animal virus studies made a dramatic shift in empha- strated with the identification of hepatitis C virus by
sis with the development of reliable in vitro animal cell molecular means without isolation and in vitro propaga-
cultures (1948(cid:1)55). As a result of intensive efforts to tion of the virus in cell culture. Viruses that could not be
control poliovirus infections, single cell culture proce- easily cultured in vitro—such as papillomaviruses, noro-
dures were defined, cell culture media were standardized, viruses, rotaviruses, and certain nidoviruses amongst