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IMMUNOLOGY A Short Course To Lisa, Jonathan, and Jennifer R.C. To Ilene, Caroline, Alex, and Pearl G.S. This title is also available as an e-book. For more details, please see www.wiley.com/buy/9781118396919 or scan this QR code: IMMUNOLOGY A Short Course SEVENTH EDITION Richard Coico SUNY Downstate College of Medicine, Brooklyn, New York Geoffrey Sunshine Heath Effects Institute, and Tufts University School of Medicine, Boston, Massachusetts This edition first published 2015 © 2015 by John Wiley & Sons Ltd Registered office: John Wiley & Sons, Ltd, The Atrium, Southern Gate, Chichester, West Sussex, PO19 8SQ, UK Editorial offices: 9600 Garsington Road, Oxford, OX4 2DQ, UK The Atrium, Southern Gate, Chichester, West Sussex, PO19 8SQ, UK 111 River Street, Hoboken, NJ 07030-5774, USA For details of our global editorial offices, for customer services, and for information about how to apply for permission to reuse the copyright material in this book, please see our website at www.wiley.com/wiley-blackwell. The right of the author to be identified as the author of this work has been asserted in accordance with the UK Copyright, Designs and Patents Act 1988. All rights reserved. No part of this publication may be reproduced, stored in a retrieval system, 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 Act 1988, without the prior permission of the publisher. Designations used by companies to distinguish their products are often claimed as trademarks. All brand names and product names used in this book are trade names, service marks, trademarks, or registered trademarks of their respective owners. The publisher is not associated with any product or vendor mentioned in this book. It is sold on the understanding that the publisher is not engaged in rendering professional services. If professional advice or other expert assistance is required, the services of a competent 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 recommending or promoting a specific method, diagnosis, or treatment by health science practitioners for any particular patient. The publisher and the author make no representations or warranties with respect to the accuracy or completeness of the contents of this work and specifically disclaim all warranties, including without limitation any implied warranties of fitness for a particular purpose. In view of ongoing research, equipment modifications, changes in governmental regulations, and the constant flow of information relating to the use of medicines, equipment, and devices, the reader is urged to review and evaluate the information provided 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 added warnings and precautions. Readers should consult with a specialist where appropriate. The fact that an organization or website is referred to in this work as a citation and/or a potential source of further information does not mean that the author or the publisher endorses the information the organization or website may provide or recommendations it may make. Further, readers should be aware that Internet websites listed in this work may have changed or disappeared between the time when this work was written and when it is read. No warranty may be created or extended by any promotional statements for this work. Neither the publisher nor the author shall be liable for any damages arising herefrom. Library of Congress Cataloging-in-Publication Data Coico, Richard, author. Immunology : a short course / Richard Coico, Geoffrey Sunshine. – Seventh edition. p. ; cm. Includes bibliographical references and index. ISBN 978-1-118-39691-9 (pbk.) I. Sunshine, Geoffrey, author. II. Title. [DNLM: 1. Allergy and Immunology. 2. Immune System Diseases. 3. Immune System Processes. 4. Immunity. QW 504] QR181 616.07′9–dc23 2014023101 A catalogue record for this book is available from the British Library. Wiley also publishes its books in a variety of electronic formats. Some content that appears in print may not be available in electronic books. Cover image: Russell Kightley Set in 10/12 pt Times by Toppan Best-set Premedia Limited 1 2015 CONTENTS IN BRIEF 1 11 OVERVIEW OF THE IMMUNE SYSTEM, 1 ACTIVATION AND FUNCTION OF T CELLS, 153 2 12 INNATE IMMUNITY, 11 CYTOKINES, 176 3 13 ADAPTIVE IMMUNITY, 26 TOLERANCE AND AUTOIMMUNITY, 194 4 14 IMMUNOGENS AND ANTIGENS, 35 COMPLEMENT, 217 15 5 HYPERSENSITIVITY: TYPE I, 233 ANTIBODY STRUCTURE AND FUNCTION, 47 16 6 HYPERSENSITIVITY: TYPES II AND III, 249 ANTIGEN–ANTIBODY INTERACTIONS, IMMUNE ASSAYS, AND EXPERIMENTAL SYSTEMS, 67 17 HYPERSENSITIVITY: TYPE IV, 259 7 THE GENETIC BASIS OF ANTIBODY 18 STRUCTURE, 88 IMMUNODEFICIENCY DISORDERS AND NEOPLASIAS OF THE LYMPHOID SYSTEM, 268 8 BIOLOGY OF THE B LYMPHOCYTE, 100 19 TRANSPLANTATION, 298 9 HOW T CELLS RECOGNIZE ANTIGEN: 20 THE ROLE OF THE MAJOR HISTOCOMPATIBILITY TUMOR IMMUNOLOGY, 312 COMPLEX, 117 21 RESISTANCE AND IMMUNIZATION TO 10 BIOLOGY OF THE T LYMPHOCYTE, 137 INFECTIOUS DISEASES, 328 v CONTENTS About the Authors, xv Inflammation, 20 Hallmark Signs of Inflammation, 20 Contributors, xvi Localized Inflammatory Responses, 21 Preface and Acknowledgments, xvii Chronic Inflammation, 23 How to Use Your Textbook, xix Fever, 23 References and Bibliography, 24 About the Companion Website, xxiii Review Questions, 24 Answers to Review Questions, 25 1 OVERVIEW OF THE IMMUNE SYSTEM, 1 3 Introduction, 1 ADAPTIVE IMMUNITY, 26 Innate and Adaptive Immunity, 2 Cells and Organs Involved in Adaptive Immunity, 26 Innate Immunity, 2 The Lymphatic Organs, 27 Adaptive Immunity, 2 Lymphocyte Migration and Recirculation, 29 Clonal Selection Theory, 3 The Fate of Antigen after Penetration, 31 Active, Passive, and Adoptive Immunization, 5 Frequency of Antigen-Specific Naïve Major Characteristics of the Adaptive Immune Lymphocytes, 32 Response, 5 Interrelationship between Innate and Adaptive Cells Involved in the Adaptive Immunity, 33 Immune Response, 5 Review Questions, 34 Humoral and Cellular Immunity, 6 Answers to Review Questions, 34 Humoral Immunity, 6 Cell-Mediated Immunity, 7 4 IMMUNOGENS AND ANTIGENS, 35 Generation of Diversity in the Immune Response, 8 Introduction, 35 Benefits of Immunology, 8 Requirements for Immunogenicity, 35 Damaging Effects of the Foreignness, 35 Immune Response, 9 High Molecular Weight, 36 The Future of Immunology, 9 Chemical Complexity, 36 The Short Course Begins Here, 10 Degradability, 36 References and Bibliography 10 Haptens, 36 Further Requirements for Immunogenicity, 37 2 INNATE IMMUNITY, 11 Primary and Secondary Responses, 38 Introduction, 11 Antigenicity and Antigen-Binding Site, 38 Physical and Chemical Barriers of Innate Immunity, 11 Epitopes Recognized by B Cells and T Cells, 39 Origin, Differentiation, and Characterization of Cells Major Classes of Antigens, 40 of the Innate Immune System, 12 Binding of Antigen with Antigen-Specific Antibodies or Pattern Recognition: The Hallmark of Innate Immune T Cells, 41 Responses, 15 Cross-Reactivity, 41 Pattern Recognition Receptors, 15 Adjuvants, 42 Complement, 18 References and Bibliography, 44 Intracellular and Extracellular Killing of Review Questions, 45 Microorganisms, 19 Answers to Review Questions, 45 vii viii CONTENTS 5 Primary Interactions between Antibody and Antigen, 68 ANTIBODY STRUCTURE AND FUNCTION, 47 Association Constant, 68 Introduction, 47 Affinity and Avidity, 70 Isolation and Characterization of Immunoglobulins, 48 Secondary Interactions between Antibody and Structure of Light and Heavy Chains, 48 Antigen, 70 Domains, 51 Agglutination Reactions, 70 Hinge Region, 51 Precipitation Reactions, 72 Variable Region, 51 Immunoassays, 74 Immunoglobulin Variants, 53 Direct-Binding Immunoassays, 74 Isotypes, 53 Solid-Phase Immunoassays, 75 Allotypes, 54 Immunofluorescence, 76 Idiotypes, 54 Direct Immunofluorescence, 76 Structural Features of IgG, 55 Indirect Immunofluorescence, 76 Biologic Properties of IgG, 55 Flow Cytometry, 76 Agglutination and Formation of Precipitate, 56 Immunoabsorption and Immunoadsorption, 78 Passage through the Placenta and Absorption in Cellular Assays, 78 Neonates, 56 Assays of Lymphocyte Function, 78 Opsonization, 57 B-Cell and T-Cell Proliferation Assays, 78 Antibody-Dependent Cell-Mediated Cytotoxicity, 58 Antibody Production by B Cells, 78 Activation of Complement, 58 Effector Cell Assays for T Cells and Natural Killer Neutralization of Toxins, 58 Cells, 79 Immobilization of Bacteria, 58 Cell Culture, 79 Neutralization of Viruses, 58 Primary Cell Cultures and Cloned Lymphoid Cell Structural Features of IgM, 59 Lines, 79 Biologic Properties of IgM, 59 B-Cell Hybridomas and Monoclonal Antibodies, 80 Complement Fixation, 59 T-Cell Hybridomas, 80 Neonatal Immunity and First Line of Humoral Genetically Engineered Molecules and Defense, 59 Receptors, 80 Agglutination, 60 Experimental Animal Models, 81 Isohemagglutinins, 60 Inbred Strains, 81 Structural and Biologic Properties of IgA, 60 Adoptive Transfer, 82 Biologic Properties of IgA, 60 SCID Mice, 82 Role in Mucosal Infections, 60 Thymectomized and Congenitally Athymic (Nude) Bactericidal Activity, 61 Mice, 82 Antiviral Activity, 61 Transgenic Mice and Gene Targeting, 82 Structural and Biologic Properties of IgD, 61 Transgenic Mice, 82 Structural and Biologic Properties of IgE, 62 Knockout and Knock-in Mice, 83 Importance of IgE in Parasitic Infections and Analysis of Gene Expression, 83 Hypersensitivity Reactions, 62 Microarrays to Assess Gene Expression, 83 Kinetics of the Antibody Response Following References and Bibliography, 85 Immunization, 62 Review Questions, 86 Primary Response, 62 Answers to Review Questions, 86 Secondary Response, 62 The Immunoglobulin Superfamily, 63 References and Bibliography, 65 7 THE GENETIC BASIS OF ANTIBODY Review Questions, 65 STRUCTURE, 88 Answers to Review Questions, 66 Introduction, 88 A Brief Review of Nonimmunoglobulin Gene Structure 6 ANTIGEN–ANTIBODY INTERACTIONS, IMMUNE and Gene Expression, 88 ASSAYS, AND EXPERIMENTAL SYSTEMS, 67 Genetic Events in Synthesis of Ig Chains, 90 Introduction, 67 Organization and Rearrangement of Antigen–Antibody Interactions, 67 Light-Chain Genes, 90

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