V IROLOGY V IROLOGY PRINCIPLES AND APPLICATIONS John B. Carter and Venetia A. Saunders School of Biomolecular Sciences, Liverpool John Moores University, UK Copyright2007 JohnWiley&SonsLtd,TheAtrium,SouthernGate,Chichester, WestSussexPO198SQ,England Telephone(+44)1243779777 Email(forordersandcustomerserviceenquiries):[email protected] VisitourHomePageonwww.wileyeurope.comorwww.wiley.com AllRightsReserved.Nopartofthispublicationmaybereproduced,storedinaretrievalsystemortransmittedinanyformorby anymeans,electronic,mechanical,photocopying,recording,scanningorotherwise,exceptunderthetermsoftheCopyright, DesignsandPatentsAct1988orunderthetermsofalicenceissuedbytheCopyrightLicensingAgencyLtd,90TottenhamCourt Road,LondonW1T4LP,UK,withoutthepermissioninwritingofthePublisher.RequeststothePublishershouldbeaddressed tothePermissionsDepartment,JohnWiley&SonsLtd,TheAtrium,SouthernGate,Chichester, WestSussexPO198SQ,England,[email protected],orfaxedto(+44)1243770620. 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To Myra, Robert, Jon and Mark Contents Preface xv Abbreviations used in this book xvii Greek letters used in this book xxi Colour coding for molecules xxiii 1 Viruses and their importance 1 Ataglance 1 1.1 VirusesareubiquitousonEarth 3 1.2 Reasonsforstudyingviruses 3 1.3 Thenatureofviruses 4 1.4 Theremainderofthebook 7 Learningoutcomes 7 Sourcesoffurtherinformation 7 2 Methods used in virology 9 Ataglance 9 2.1 Introductiontomethodsusedinvirology 11 2.2 Cultivationofviruses 12 2.3 Isolationofviruses 13 2.4 Centrifugation 13 2.5 Structuralinvestigationsofcellsandvirions 17 2.6 Electrophoretictechniques 17 2.7 Detectionofvirusesandviruscomponents 18 2.8 Infectivityassays 22 2.9 Virusgenetics 26 Learningoutcomes 28 Sourcesoffurtherinformation 28 3 Virus structure 31 Ataglance 31 3.1 Introductiontovirusstructure 32 3.2 Virusgenomes 32 3.3 Virusproteins 37 3.4 Capsids 39 3.5 Virionmembranes 45 3.6 Occlusionbodies 47 viii CONTENTS 3.7 Othervirioncomponents 47 Learningoutcomes 48 Sourcesoffurtherinformation 48 4 Virus transmission 49 Ataglance 49 4.1 Introductiontovirustransmission 50 4.2 Transmissionofplantviruses 51 4.3 Transmissionofvertebrateviruses 54 4.4 Transmissionofinvertebrateviruses 56 4.5 Permissivecells 57 Learningoutcomes 58 Sourcesoffurtherinformation 58 5 Attachment and entry of viruses into cells 59 Ataglance 59 5.1 Overviewofvirusreplication 60 5.2 Animalviruses 61 5.3 Bacteriophages 68 Learningoutcomes 68 Sourcesoffurtherinformation 68 6 Transcription, translation and transport 69 Ataglance 69 6.1 Introductiontotranscription,translationandtransport 70 6.2 Transcriptionofvirusgenomes 70 6.3 Transcriptionineukaryotes 72 6.4 Translationineukaryotes 77 6.5 Transportineukaryoticcells 81 6.6 Transcriptionandtranslationinbacteria 83 Learningoutcomes 84 Sourcesoffurtherinformation 84 7 Virus genome replication 85 Ataglance 85 7.1 Overviewofvirusgenomereplication 86 7.2 Locationsofvirusgenomereplicationineukaryoticcells 88 7.3 Initiationofgenomereplication 88 7.4 Polymerases 89 7.5 DNAreplication 90 7.6 Double-strandedRNAreplication 91 7.7 Single-strandedRNAreplication 92 7.8 Reversetranscription 92 Learningoutcomes 93 Sourcesoffurtherinformation 93 CONTENTS ix 8 Assembly and exit of virions from cells 95 Ataglance 95 8.1 Introductiontoassemblyandexitofvirionsfromcells 96 8.2 Nucleocapsidassembly 96 8.3 Formationofvirionmembranes 98 8.4 Virionexitfromtheinfectedcell 101 Learningoutcomes 101 Sourcesoffurtherinformation 102 9 Outcomes of infection for the host 103 Ataglance 103 9.1 Introductiontooutcomesofinfectionforthehost 105 9.2 Factorsaffectingoutcomesofinfection 105 9.3 Non-productiveinfections 110 9.4 Productiveinfections 112 Learningoutcomes 114 Sourcesoffurtherinformation 114 10 Classification and nomenclature of viruses 115 Ataglance 115 10.1 Historyofvirusclassificationandnomenclature 116 10.2 Modernvirusclassificationandnomenclature 116 10.3 Baltimoreclassificationofviruses 119 Learningoutcomes 119 Sourcesoffurtherinformation 120 11 Herpesviruses (and other dsDNA viruses) 121 Ataglance 121 11.1 Introductiontoherpesviruses 122 11.2 Thehumanherpesviruses 122 11.3 Theherpesvirusvirion 123 11.4 HSV-1genomeorganization 124 11.5 HSV-1replication 124 11.6 Latentherpesvirusinfection 130 11.7 OtherdsDNAviruses 132 Learningoutcomes 135 Sourcesoffurtherinformation 135 12 Parvoviruses (and other ssDNA viruses) 137 Ataglance 137 12.1 Introductiontoparvoviruses 138 12.2 Examplesofparvoviruses 138 12.3 Parvovirusvirion 139 12.4 Parvovirusreplication 141