,VeterinaryI
mmunology
l
PrinciplesandPractice l l
Michael J. Day l l
BScl BVMS(Hons)l PhDl DScl DiplECVPl FASMl FRCPathl FRCVS
ProfessorloflVeterinarylPathology
DivisionloflVeterinarylPathology,lInfectionlandlImmunitylSchooll
oflVeterinarylScience
UniversityloflBristollLangford
,lBristol,lUK
inl collaborationl with
Ronald D. Schultz
l l
BSlMSlPhDlDACVM
ProfessorlandlChairlDepartment
loflPathobiologicallSciences
SchoolloflVeterinarylMedicinelUniversity
loflWisconsin-
MadisonlMadison,lWisconsin,lUSA
MANSON PUBLISHING/THE VETERINARY PRESS
l l l
,Copyrightl©l2011lMansonlPublishinglLtdlISBN:
l978-1-84076-143-6
Alllrightslreserved.lNolpartloflthislpublicationlmaylbelreproduced,lstoredlinlalretrievallsystemlorltransmittedlinlanylf
ormlorlbylanylmeanslwithoutlthelwrittenlpermissionloflthelcopyrightlholderlorlinlaccordancelwithlthelprovisionsloflt
helCopyrightlActl1956l(aslamended),lorlunderltheltermsloflanyllicencelpermittingllimitedlcopyinglissuedl byl thel Cop
yrightl Licensingl Agency,l 33–34l Alfredl Place,l Londonl WC1El 7DP,l UK.
Anylpersonlwholdoeslanylunauthorizedlactlinlrelationltolthislpublicationlmaylbelliableltolcriminallprosecutionlandlci
villclaimslforldamages.
Al CIPl cataloguel recordl forl thisl bookl isl availablel froml thel Britishl Library.
ForlfullldetailsloflalllMansonlPublishinglLtdltitleslpleaselwritelto:lMansonl
PublishinglLtd,l73lCorringhamlRoad,lLondonlNW11l7DL,lUK.lTel:l+44(0)2
0l8905l5150
Fax:l +44(0)20l 8201l 9233
Email:l
ite:l www.mansonpublishing.com
Commissioningleditor:lJilllNorthcottlP
rojectl manager:l Katel Nardonil Copyl
editor:lPeterlBeynonlIllustration:lcact
usdesign.co.uk
Colourlreproduction:lTenonl&lPolertlColourlScanninglLtd,lHonglKonglPri
ntedlin:lThelCzechlRepublic
, Foreword 5 Antigenl uptake 74
Abbreviations 6 Antigenlprocessinglandlpresentation 77
AntigenlpresentationlbylCD1 79
Preface 8 AntigenlpresentationlbylBlcellsland
KeyltolSymbols 8 nonprofessionall APCs 79
Cytokines 80
1 AnlOverviewloflthelImmunelSystem:lInnatelandlAdapti Keyl points 82
velImmunitylandlthelInflammatorylResponse 9 8 ThelBiologyloflTlLymphocytes 83
Introduction 9 Introduction 83
Historyl ofl immunology 9 Thel T-celll receptor 83
Thelimmunelsystem:lanloverview 10 T-cellldevelopmentlandlmaturation 86
Thel evolutionl ofl thel immunel system 16 T-celllactivation 86
Keyl points 18 Thel rolel ofl CD4+l Tl lymphocytes 88
2 AntigenslandlAntibodies 19 Cytotoxicity 90
Introduction 19 Memoryl cells 94
Antigens 19 llymphocytes 95
Antibodies 22 NaturallkillerlTlcells 95
Immunoglobulinl classes 22 Keyl points 96
Antigen–antibodyl interaction 28 9 ThelBiologyloflBlLymphocytes 97
Keyl points 29 Introduction 97
3 ThelComplementlSystem 30 Antigenl recognitionl byl Bl cells 97
Introduction 30 Developmentl andl maturationl ofl Bl lymphocytes 98
Thel classicall andl lectinl pathways 31 Activationl ofl thel Bl lymphocyte 98
Thelalternativelpathway 32 B-celll receptorl diversity 100
Thel terminall pathway 34 Thel immunoglobulinl classl switch 102
Biologicall consequencesl ofl complement Affinityl maturation 102
activation 34 Kineticsl ofl thel humorall immunel response 103
Testsloflcomplementlfunction 37 B1l andl B2l lymphocytes 103
Keyl points 38 Memoryl Bl lymphocytes 104
Monoclonall antibodies 104
4 SerologicallTesting 39 Keyl points 106
Introduction 39
Serologicall tests 40 10 TestingloflCellularlImmunelFunction 107
Keyl points 48 Introduction 107
Lymphocytelstimulationltests 108
5 CellslandlTissuesloflthelImmunelSystem 49 Testsl ofl cytotoxicl function 109
Introduction 49 Testsloflphagocyticl celllfunction 110
Primaryllymphoidltissue 49 Keyl points 111
Secondaryllymphoidltissue 51
Lymphocytes 54 11 Immunel Suppression 112
Distinguishingl betweenl immunel cells 56 Introduction 112
Lymphocytelrecirculation 60 Antigenl usage 112
Thel commonl mucosall system 64 Antibody-mediatedlsuppression 113
Keyl points 64 Thel neuroendocrine–immunologicall loop 114
Regulatoryl (suppressor)l Tl lymphocytes 115
6 ThelMajorlHistocompatibilitylComplex 65 Keyl points 119
Introduction 65
Graftl rejection 65 12 Hypersensitivityl Mechanisms 120
Majorl histocompatibilityl antigens 66 Introduction 120
Thel majorl histocompatibilityl complex 67 ThelGelllandlCoombslclassificationlofl hypersensitivity
Thel inheritancel ofl MHC 68 120
MHCl diseasel associations 69 Typel Il hypersensitivity 121
Otherl applicationsl ofl tissuel typing 70 Typel IIl hypersensitivity 123
Keyl points 71 Typel IIIl hypersensitivity 125
Typel IVl hypersensitivity 127
7 AntigenlPresentationlandlCytokines 72 Keyl points 130
Introduction 72
Antigenl presentingl cells 72
mmunology
l
PrinciplesandPractice l l
Michael J. Day l l
BScl BVMS(Hons)l PhDl DScl DiplECVPl FASMl FRCPathl FRCVS
ProfessorloflVeterinarylPathology
DivisionloflVeterinarylPathology,lInfectionlandlImmunitylSchooll
oflVeterinarylScience
UniversityloflBristollLangford
,lBristol,lUK
inl collaborationl with
Ronald D. Schultz
l l
BSlMSlPhDlDACVM
ProfessorlandlChairlDepartment
loflPathobiologicallSciences
SchoolloflVeterinarylMedicinelUniversity
loflWisconsin-
MadisonlMadison,lWisconsin,lUSA
MANSON PUBLISHING/THE VETERINARY PRESS
l l l
,Copyrightl©l2011lMansonlPublishinglLtdlISBN:
l978-1-84076-143-6
Alllrightslreserved.lNolpartloflthislpublicationlmaylbelreproduced,lstoredlinlalretrievallsystemlorltransmittedlinlanylf
ormlorlbylanylmeanslwithoutlthelwrittenlpermissionloflthelcopyrightlholderlorlinlaccordancelwithlthelprovisionsloflt
helCopyrightlActl1956l(aslamended),lorlunderltheltermsloflanyllicencelpermittingllimitedlcopyinglissuedl byl thel Cop
yrightl Licensingl Agency,l 33–34l Alfredl Place,l Londonl WC1El 7DP,l UK.
Anylpersonlwholdoeslanylunauthorizedlactlinlrelationltolthislpublicationlmaylbelliableltolcriminallprosecutionlandlci
villclaimslforldamages.
Al CIPl cataloguel recordl forl thisl bookl isl availablel froml thel Britishl Library.
ForlfullldetailsloflalllMansonlPublishinglLtdltitleslpleaselwritelto:lMansonl
PublishinglLtd,l73lCorringhamlRoad,lLondonlNW11l7DL,lUK.lTel:l+44(0)2
0l8905l5150
Fax:l +44(0)20l 8201l 9233
Email:l
ite:l www.mansonpublishing.com
Commissioningleditor:lJilllNorthcottlP
rojectl manager:l Katel Nardonil Copyl
editor:lPeterlBeynonlIllustration:lcact
usdesign.co.uk
Colourlreproduction:lTenonl&lPolertlColourlScanninglLtd,lHonglKonglPri
ntedlin:lThelCzechlRepublic
, Foreword 5 Antigenl uptake 74
Abbreviations 6 Antigenlprocessinglandlpresentation 77
AntigenlpresentationlbylCD1 79
Preface 8 AntigenlpresentationlbylBlcellsland
KeyltolSymbols 8 nonprofessionall APCs 79
Cytokines 80
1 AnlOverviewloflthelImmunelSystem:lInnatelandlAdapti Keyl points 82
velImmunitylandlthelInflammatorylResponse 9 8 ThelBiologyloflTlLymphocytes 83
Introduction 9 Introduction 83
Historyl ofl immunology 9 Thel T-celll receptor 83
Thelimmunelsystem:lanloverview 10 T-cellldevelopmentlandlmaturation 86
Thel evolutionl ofl thel immunel system 16 T-celllactivation 86
Keyl points 18 Thel rolel ofl CD4+l Tl lymphocytes 88
2 AntigenslandlAntibodies 19 Cytotoxicity 90
Introduction 19 Memoryl cells 94
Antigens 19 llymphocytes 95
Antibodies 22 NaturallkillerlTlcells 95
Immunoglobulinl classes 22 Keyl points 96
Antigen–antibodyl interaction 28 9 ThelBiologyloflBlLymphocytes 97
Keyl points 29 Introduction 97
3 ThelComplementlSystem 30 Antigenl recognitionl byl Bl cells 97
Introduction 30 Developmentl andl maturationl ofl Bl lymphocytes 98
Thel classicall andl lectinl pathways 31 Activationl ofl thel Bl lymphocyte 98
Thelalternativelpathway 32 B-celll receptorl diversity 100
Thel terminall pathway 34 Thel immunoglobulinl classl switch 102
Biologicall consequencesl ofl complement Affinityl maturation 102
activation 34 Kineticsl ofl thel humorall immunel response 103
Testsloflcomplementlfunction 37 B1l andl B2l lymphocytes 103
Keyl points 38 Memoryl Bl lymphocytes 104
Monoclonall antibodies 104
4 SerologicallTesting 39 Keyl points 106
Introduction 39
Serologicall tests 40 10 TestingloflCellularlImmunelFunction 107
Keyl points 48 Introduction 107
Lymphocytelstimulationltests 108
5 CellslandlTissuesloflthelImmunelSystem 49 Testsl ofl cytotoxicl function 109
Introduction 49 Testsloflphagocyticl celllfunction 110
Primaryllymphoidltissue 49 Keyl points 111
Secondaryllymphoidltissue 51
Lymphocytes 54 11 Immunel Suppression 112
Distinguishingl betweenl immunel cells 56 Introduction 112
Lymphocytelrecirculation 60 Antigenl usage 112
Thel commonl mucosall system 64 Antibody-mediatedlsuppression 113
Keyl points 64 Thel neuroendocrine–immunologicall loop 114
Regulatoryl (suppressor)l Tl lymphocytes 115
6 ThelMajorlHistocompatibilitylComplex 65 Keyl points 119
Introduction 65
Graftl rejection 65 12 Hypersensitivityl Mechanisms 120
Majorl histocompatibilityl antigens 66 Introduction 120
Thel majorl histocompatibilityl complex 67 ThelGelllandlCoombslclassificationlofl hypersensitivity
Thel inheritancel ofl MHC 68 120
MHCl diseasel associations 69 Typel Il hypersensitivity 121
Otherl applicationsl ofl tissuel typing 70 Typel IIl hypersensitivity 123
Keyl points 71 Typel IIIl hypersensitivity 125
Typel IVl hypersensitivity 127
7 AntigenlPresentationlandlCytokines 72 Keyl points 130
Introduction 72
Antigenl presentingl cells 72
