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Roitt’sEssentialImmunology

Peter J. DelvesProfessor Delves obtained his PhD from the University ofLondon in 1986 and is a Professor of Immunology at UCL(University College London). His research has focused onmolecular aspects of antigen recognition. He has authored andedited a number of immunology books, and teaches the subjectat a broad range of levels.Seamus J. MartinProfessor Martin received his PhD from The National Universityof Ireland in 1990 and trained as a post-doctoral fellow atUniversity College London (with Ivan Roitt) and The La JollaInstitute for Allergy and Immunology, California, USA (withDoug Green). Since 1999, he is the holder of the Smurfit Chairof Medical Genetics at Trinity College Dublin and is also aScience Foundation Ireland Principal Investigator. His research isfocused on various aspects of programmed cell death (apoptosis)in the immune system and in cancer and he has received severalawards for his work in this area. He has previously edited twobooks on apoptosis and was elected as a Member of The RoyalIrish Academy in 2006 and as a member of The EuropeanMolecular Biology Organisation (EMBO) in 2009.Dennis R. BurtonProfessor Burton obtained his BA in Chemistry from theUniversity of Oxford in 1974 and his PhD in PhysicalBiochemistry from the University of Lund in Sweden in 1978.After a period at the University of Sheffield, he moved to theScripps Research Institute in La Jolla, California in 1989 wherehe is Professor of Immunology and Molecular Biology. Hisresearch interests include antibodies, antibody responses topathogens and vaccine design, particularly in relation to HIV.Ivan M. RoittProfessor Roitt was born in 1927 and educated at King Edward’sSchool, Birmingham and Balliol College, Oxford. In 1956,together with Deborah Doniach and Peter Campbell, he madethe classic discovery of thyroglobulin autoantibodies inHashimoto’s thyroiditis which helped to open the whole conceptof a relationship between autoimmunity and human disease. Thework was extended to an intensive study of autoimmunephenomena in pernicious anaemia and primary biliary cirrhosis.In 1983 he was elected a Fellow of The Royal Society, and hasbeen elected to Honorary Membership of the Royal College ofPhysicians and appointed Honorary Fellow of The Royal Societyof Medicine.

TWELF TH EDITIONRoitt’sEssentialImmunologyPeter J. DelvesPhDDivision of Infection and ImmunityUCLLondon, UKSeamus J. MartinPhD, FTCD, MRIAThe Smurfit Institute of GeneticsTrinity CollegeDublin, IrelandDennis R. BurtonPhDDepartment of Immunology and Molecular BiologyThe Scripps Research InstituteLa Jolla, California, USAIvan M. RoittMA, DSc(Oxon), FRCPath, Hon FRCP (Lond), FRSCentre for Investigative and Diagnostic OncologyMiddlesex UniversityLondon, UKA John Wiley & Sons, Ltd., Publication

This edition first published 2011 1971, 1974, 1977, 1980, 1984, 1988, 1991, 1994,1997, 2001, 2006 by Peter J Delves, Seamus J. Martin, Dennis R. Burton, Ivan M. RoittBlackwell Publishing was acquired by John Wiley & Sons in February 2007. Blackwell’spublishing program has been merged with Wiley’s global Scientific, Technical and Medicalbusiness to form Wiley-Blackwell.Registered office: John Wiley & Sons, Ltd, The Atrium, Southern Gate, Chichester, WestSussex, PO19 8SQ, UKEditorial offices: 9600 Garsington Road, Oxford, OX4 2DQ, UKThe Atrium, Southern Gate, Chichester, West Sussex, PO19 8SQ, UK111 River Street, Hoboken, NJ 07030-5774, USAFor details of our global editorial offices, for customer services and for information abouthow to apply for permission to reuse the copyright material in this book please see ourwebsite at www.wiley.com/wiley-blackwellThe right of the author to be identified as the author of this work has been asserted inaccordance with the UK Copyright, Designs and Patents Act 1988.All rights reserved. No part of this publication may be reproduced, stored in a retrievalsystem, or transmitted, in any form or by any means, electronic, mechanical, photocopying,recording or otherwise, except as permitted by the UK Copyright, Designs and Patents Act1988, without the prior permission of the publisher.Designations used by companies to distinguish their products are often claimed astrademarks. All brand names and product names used in this book are trade names, servicemarks, trademarks or registered trademarks of their respective owners. The publisher is notassociated with any product or vendor mentioned in this book. This publication is designedto provide accurate and authoritative information in regard to the subject matter covered. Itis sold on the understanding that the publisher is not engaged in rendering professionalservices. If professional advice or other expert assistance is required, the services of acompetent professional should be sought.The contents of this work are intended to further general scientific research, understanding,and discussion only and are not intended and should not be relied upon as recommendingor promoting a specific method, diagnosis, or treatment by physicians for any particularpatient. The publisher and the author make no representations or warranties with respect tothe accuracy or completeness of the contents of this work and specifically disclaim allwarranties, including without limitation any implied warranties of fitness for a particularpurpose. In view of ongoing research, equipment modifications, changes in governmentalregulations, and the constant flow of information relating to the use of medicines,equipment, and devices, the reader is urged to review and evaluate the informationprovided in the package insert or instructions for each medicine, equipment, or device for,among other things, any changes in the instructions or indication of usage and for addedwarnings and precautions. Readers should consult with a specialist where appropriate. Thefact that an organization or Website is referred to in this work as a citation and/or apotential source of further information does not mean that the author or the publisherendorses the information the organization or Website may provide or recommendations itmay make. Further, readers should be aware that Internet Websites listed in this work mayhave changed or disappeared between when this work was written and when it is read. Nowarranty may be created or extended by any promotional statements for this work. Neitherthe publisher nor the author shall be liable for any damages arising herefrom.Library of Congress Cataloging-in-Publication DataRoitt’s essential immunology / Peter J. Delves . [et al.]. – 12th ed.p. cm. – (Essentials ; 16)Rev. ed. of: Roitt’s essential immunology / Ivan M. Roitt, Peter J. Delves.Includes bibliographical references and index.ISBN 978-1-4051-9683-3 (pbk.)1. Immunology. I. Delves, Peter J. II. Roitt, Ivan M. (Ivan Maurice) Roitt’s essentialimmunology. III. Title: Essential immunology.QR181.R57 2011616.07'9–dc222010047392A catalogue record for this book is available from the British Library.Set in 10/12 pt Adobe Garamond Pro by Toppan Best-set Premedia Limited12011

ContentsAcknowledgmentsPrefaceAbbreviationsHow to get the best out of your textbookviviiviiixiiPART 1: Fundamentals of Immunology1. Innate immunity32. Specific acquired immunity353. Antibodies534. Membrane receptors for antigen795. The primary interaction with antigen1136. Immunological methods and applications1417. The anatomy of the immune response1888. Lymphocyte activation2059. The production of effectors22610. Control mechanisms26311. Ontogeny and phylogeny283PART 2: Applied Immunology12. Adversarial strategies during infection31313. Vaccines34514. Immunodeficiency36915. Allergy and other hypersensitivities39416. Transplantation42317. Tumor immunology44518. Autoimmune diseases475GlossaryIndex511522Companion website www.roitt.com

AcknowledgmentsThe input of the editorial team of Elizabeth Johnston, Laura Murphy and Cathryn Gates at Wiley-Blackwell and the projectmanagement of Ruth Swan is warmly acknowledged. We are much indebted to the co-editors of Immunology, J. Brostoff, D.Roth and D. Male, together with the publishers, Mosby, and the following individuals for permission to utilize or modify theirfigures: J. Brostoff and A. Hall for figure 15.11; J. Horton for figure 11.19; and J. Taverne for figures 12.23 and 12.24.IMR would like to acknowledge the indefatigable secretarial assistance of Christine Griffin. DRB wishes to particularlyacknowledge the invaluable contributions of Amandeep Gakhal, Erin Scherer, Rena Astronomo and Wendelien Oswald. He isgrateful to Jenny Woof, Ann Feeney, Beatrice Hahn, Jim Marks, Don Mosier, Paul Sharp, Robyn Stanfield, James Stevens andMario Stevenson for many very helpful comments. PJD would particularly like to thank Per Brandtzaeg, Volker Brinkmann,Greg Campbell, Peter Lydyard, Rand Swenson and Ulrich Wahn. SJM is indebted to Ed Lavelle, Sean Cullen, CristinaMunoz-Pinedo and all of the members of his laboratory for comments, suggestions and support. He would also like to thankMia, Madeleine and Jamie for their support and indulgence.Every effort has been made by the authors and the publisher to contact all the copyright holders to obtain their permissionto reproduce copyright material. However, if any have been inadvertently overlooked, the publisher will be pleased to make thenecessary arrangements at the first opportunity.A number of scientists very generously provided illustrations for inclusion in this edition, and we have acknowledged ourgratitude to them in the relevant figure legends.Companion websiteThis book is accompanied by a companion website:www.roitt.comThe website includes: Interactive MCQs and SBA questions for each chapter, with feedback on all answers selected Animations and videos showing key concepts Fully downloadable figures and illustrations, further reading and useful links Extracts from the Encyclopaedia of Life Sciences Podcasts to reinforce the key principles explained in the text: ideal for revision ‘on the go’

PrefaceWelcome to this new edition! When Ivan wrote the first edition some 40 years ago, he wanted to feel that he was chatting tothe reader almost informally, rather than preaching, and it has been our intention to maintain this style. As a subject, immunology is exciting and dynamic and to persuade you that it is absolutely worthwhile for you to tackle this new edition we havemade very extensive changes to update the previous edition. Accordingly, apart from the introduction of numerous new illustrations, we have: Expanded discussion of pathogen- and danger-associated molecular patterns (PAMPs & DAMPs)Introduced a new section on dendritic cells and their role in antigen processing including cross-presentationUpdated sections on B-cell and NK receptorsEnhanced discussion of lymphocyte traffickingIncorporated the latest findings on T-cell subsets, particularly Th17 and the diversity of regulatory T-cellsRecorded newer information on NK and cytotoxic T-cell killing mechanismsGiven more insight into the effects of aging on immune responsesCarried out a major rewrite of the vaccine chapter with new emphasis on mechanisms of action of conventional and carbohydrate vaccines, and new approaches to vaccine development including reverse vaccinology, together with progress in malariavaccines and adjuvant actionProvided new information on novel genetic immunodeficiency defects, on the origin of AIDS and the ever-expanding plethoraof AIDS drugs plus results from the latest HIV vaccine trialsClarified the recent findings on the cellular transformations leading to cancer, the manipulation of the immune system bytumors and the links between infection, inflammation and cancerSubstantially rewritten the chapter on autoimmune diseases.It is our fond expectation that you will enjoy and benefit from a reading of our offering.Peter J. DelvesSeamus J. MartinDennis R. BurtonIvan M. Roitt

CFAcGMPChIPCHIPCH(L)CLACLIPCMICMLadeno-associated virusantibodyacetylcholine receptoradoptive cell transferadrenocorticotropic hormoneadenosine deaminaseantibody-dependent cellular cytotoxicityasparagine endopeptidaseantigenactivation-induced cytidine deaminaseacquired immunodeficiency syndromeautoimmune regulatoraddressable laser bead assayantineutrophil cytoplasmic antibodiesantigen-presenting cellantigen receptor response element-1antigen receptor response element-2antiretroviral therapyAfrican swine fever viruszidovudine (3′-azido-3′-deoxythymidine)B-cell-activating factor of the tumornecrosis factor familylymphocyte which matures in bonemarrowbacille Calmette–Guérin attenuated formof tuberculosisB-cell receptorbone marrowbovine serum albuminbovine spongiform encephalopathyBruton’s tyrosine kinasebromodeoxyuridinecomplementconstant part of TCR α(β/γ/δ) chaincommon acute lymphoblastic leukemiaantigencyclic adenosine monophosphatecomplement control protein repeatcluster of differentiationcomplementarity determining regions of Igor TCR variable portioncarcinoembryonic antigencomplete Freund’s adjuvantcyclic guanosine monophosphatechromatin immunoprecipitationchemotaxis inhibitory proteinconstant part of Ig heavy (light) chaincutaneous lymphocyte antigenclass II-associated invariant chain peptidecell-mediated immunitycell-mediated lympholysisCMVCnCniCnCnaCpGCR(n)CRPCSFCSRCTLRD GADSg.b.m.G-CSFcytomegaloviruscomplement component “n”activated complement component “n”inactivated complement component “n”small peptide derived by proteolyticactivation of Cncytosine phosphate-guanosine dinucleotidemotifcomplement receptor “n”C-reactive proteincerebrospinal fluidclass switch recombinationC-type lectin receptordiversity minigene joining V and Jsegments to form variable regiondecay accelerating factordiacylglyceroldanger-associated molecular patterndendritic cellsdisease-modifying antirheumatic drugdinitrophenyldelayed-type hypersensitivitydiphtheria, tetanus, pertussis triplevaccineexperimental autoimmune (allergic)encephalomyelitisEpstein–Barr virusenzyme-linked immunosorbent assayelectron microscopeeosinophilerythropoietinendoplasmic reticulumembryonic stem (cell)exfoliative toxinsfactor (B, etc.)monovalent Ig antigen-binding fragmentafter papain digestiondivalent antigen-binding fragment afterpepsin digestionFas-ligandfluorescence-activated cell sorterIg crystallisable-fragment originally; nownon-Fab part of Igreceptor for IgG Fc fragmentfollicular dendritic cellflk-2 ligand(single chain) VH–VL antigen bindingfragmentGRB2-related adapter proteinglomerular basement membranegranulocyte colony-stimulating factor

Abbreviations / RHRFHSAHSChsp5HTHTLVH-YIBDICAM-1Id IL-1iNOSIP3ISCOMITAMguanine-nucleotide exchange factorsgranulocyte–macrophage colonystimulating factorn kDa glycoproteingrowth factor receptor-binding protein 2glycogen synthase kinase 3graft versus hostthe mouse major histocompatibilitycomplexmain loci for classical class I (class II)murine MHC moleculeshuman antimouse antibodieshuman anti-toxin antibodyhepatitis B surface antigenhuman chorionic gonadotropinhuman cytomegalovirushen egg lysozymehigh-walled endothelium of post capillaryvenulehuman immunodeficiency virusthe human major histocompatibilitycomplexmain loci for classical class I (class II)human MHC moleculeshigh mobility grouphypersensitive responsehomologous restriction factorheat-stable antigenhematopoietic stem cellheat-shock protein5-hydroxytryptaminehuman T-cell leukemia virusmale transplantation antigeninflammatory bowel diseaseintercellular adhesion molecule-1idiotype (anti-idiotype)interdigitating dendritic cellsinsulin-dependent diabetes mellitusindoleamine 2,3-dioxygenaseintraepithelial lymphocyteα-interferon (also IFNβ, IFNγ)interferon-regulated factorimmunoglobulinimmunoglobulin G (also IgM, IgA, IgD,IgE)surface immunoglobulinmembrane peptide chains associated withsIg B-cell receptorimmunoglobulin superfamilyinterleukin-1 (also IL-2, IL-3, etc.)inducible nitric oxide synthaseinositol triphosphateimmunostimulating compleximmunoreceptor tyrosine-based activationmotifITIMITPIVIgJAKJ chainJ LGLLHRHLIFLRRLT(B)LPSMφmAbMACMAdCAMMALTMAMMAP IICMLAMLRMMTVMRSAMSMSCimmunoreceptor tyrosine-based inhibitorymotifidiopathic thrombocytopenic purpuraintravenous immunoglobulinJanus kinasespolypeptide chain in IgA dimer and IgMjoining gene linking V or D segment toconstant regionassociation (dissociation) affinity constant(usually Ag–Ab reactions)units of molecular mass in kilo Daltonskiller immunoglobulin-like receptorskeyhole limpet hemocyaninlymphokine-activated killer celllysosomal-associated membrane proteinslinker for activation of T cellslong-acting thyroid stimulatorLPS binding proteinlymphocytic choriomeningitis virusLewisa/b/x blood group antigenslymphocyte functional antigen-1large granular lymphocyteluteinizing hormone releasing hormoneleukemia inhibiting factorleucine-rich repeatleukotriene (B etc.)lipopolysaccharide (endotoxin)macrophagemonoclonal antibodymembrane attack complexmucosal addressin cell adhesion moleculemucosa-associated lymphoid tissueMycoplasma arthritidis mitogenmitogen-activated protein kinasemitogen-associated protein kinase kinasekinasemannose binding lectinmajor basic protein of eosinophils (alsomyelin basic protein)membrane cofactor protein (complementregulation)monocyte chemotactic protein-1macrophage colony-stimulating factormuramyl dipeptidemajor histocompatibility complexMHC class I chain-related A chainmetal ion-dependent adhesion sitemacrophage migration inhibitory factorMHC class II-enriched compartmentsmonophosphoryl lipid Amixed lymphocyte reactionmouse mammary tumor virusmethicillin-resistant Staphylococcus aureusmultiple sclerosismesenchymal stem cell

x / NO·NODNZBNZB W O2 RRPTFEPTKPWMRARANTESRASTRFmelanocyte stimulating hormonemicrosomal triglyceride-transfer proteinmurine leukemia virusnicotinamide adenine dinucleotidephosphateneutrophil activating peptidenitroblue tetrazoliumneutrophil chemotactic factornuclear factor of activated T-cellsnuclear transcription factornatural killer cellnod-like receptornitric oxideNonobese diabetic mouseNew Zealand Black mouseNew Zealand Black mouse NZ WhiteF1 hybridsuperoxide anionoptical densityopen reading frameobese strain chickenovalbuminplatelet activating factor (-receptor)polyacrylamide gel electrophoresispathogen-associated molecular patternperipheral blood stem cellspassive cutaneous anaphylaxispolymerase chain reactionporcine endogenous retrovirusesprostaglandin (E etc.)phytohemagglutininphagocyte oxidasephosphatidylinositol 3-kinaseprotein inhibitor of activated STATpoly-Ig receptorphosphatidylinositol diphosphateprotein kinase CRNA-dependent protein kinasephospholipase Cphospholipase Cγ2polymorphonuclear neutrophilphotomultiplier tubeparoxysmal nocturnal hemoglobinuriaperoxisome proliferator-activated receptorpurified protein derivative fromMycobacterium tuberculosispattern recognition receptorspolytetrafluoroethyleneprotein tyrosine kinasepokeweed mitogenrheumatoid arthritisregulated upon activation normal T-cellexpressed and secreted chemokineradioallergosorbent testrheumatoid SCFscFvSCGSCIDSDFSDSSDS-PAGESEA(B CITAPT-ALLTBTcT-cellTCFTCR1(2)TdTTG-A-LTGFβrhesus blood group (D)rat insulin promoterRIG-like helicase re

Roitt’s Essential Immunology Peter J. Delves PhD Division of Infection and Immunity UCL London, UK Seamus J. Martin PhD, FTCD, MRIA Th e Smurfi t Institute of Genetics Trinity College Dublin, Ireland Dennis R. Burton PhD Department of Immunology and Molecular Biology Th e Scripps Research Institute La Jolla, California, USA Ivan M. Roitt

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Roitt IM. Roitt’s Essential Immunology 10th ed. Blackwell Science 2001 3. Hochburg et al. Rheumatology 3rd ed. Mosby 2003 4. UpToDate 12.3 5. Kalla AA. Rheumatology Handbook. Rheumatic Diseases Unit Univrersity of Cape Town. 2003 References (cont) 6. Parkin J, Cohen B. An overview of the immune system. Lancet 2001;357: 1777-1789. 7. Mackay IR .

5. Roitt’s Essential Immunology 11th ed– Delves P, Martin S, Burton D, Roitt I. Blackwell Publishing 6. Immunology 2ed – C.V.Rao. Narosa Publication 7. Molecular Biology of the Cell – Alberts et al. Garland Science 8. Molecular cell Biology – Lodish et al. Freeman Publishers 9.

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