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Basic

IMMUNOLOGY

FUNCTIONS AND DISORDERS OF THE IMMUNE SYSTEM

Abul K. Abbas, MBBS

Professor Emeritus, Pathology

University of California San Francisco

San Francisco, California

Andrew H. Lichtman, MD, PhD

Professor of Pathology

Harvard Medical School

Brigham and Women’s Hospital

Boston, Massachusetts

Shiv Pillai, MBBS, PhD

Professor of Medicine and Health Sciences

and Technology

Harvard Medical School

Massachusetts General Hospital

Boston, Massachusetts

Illustrations by David L. Baker, MA

DNA Illustrations, Inc.

SIXTH EDITION

BASIC IMMUNOLOGY: FUNCTIONS AND DISORDERS OF

THE IMMUNE SYSTEM, SIXTH EDITION ISBN: 978-0-323-54943-1

Copyright © 2020 by Elsevier Inc. All rights reserved.

No part of this publication may be reproduced or transmitted in any form or by any means, electronic or mechanical, including photocopying, recording, or any information storage and retrieval system, without permission in writing from the publisher. Details on how to seek permission, further information about the Publisher’s

permissions policies, and our arrangements with organizations such as the Copyright Clearance Center and the

Copyright Licensing Agency can be found at our website: www.elsevier.com/permissions.

This book and the individual contributions contained in it are protected under copyright by the Publisher (other

than as may be noted herein).

Notice

Practitioners and researchers must always rely on their own experience and knowledge in evaluating and

using any information, methods, compounds or experiments described herein. Because of rapid advances in

the medical sciences, in particular, independent verification of diagnoses and drug dosages should be made.

To the fullest extent of the law, no responsibility is assumed by Elsevier, authors, editors or contributors for

any injury and/or damage to persons or property as a matter of products liability, negligence or otherwise, or

from any use or operation of any methods, products, instructions, or ideas contained in the material herein.

Previous editions copyrighted 2016, 2014, 2011, 2009, 2006, 2004, and 2001.

International Standard Book Number: 978-0-323-54943-1

Executive Content Strategist: James Merritt

Senior Content Strategist: Jeremy Bowes

Director, Content Development: Rebecca Gruliow

Publishing Services Manager: Julie Eddy

Book Production Specialist: Clay S. Broeker

Design Direction: Brian Salisbury

Printed in China

Last digit is the print number: 9 8 7 6 5 4 3 2 1

1600 John F. Kennedy Blvd.

Ste. 1600

Philadelphia, PA 19103-2899

To our students

vi

The sixth edition of Basic Immunology has been revised

to include recent important advances in our knowledge

of the immune system. The original goals of this book,

from the earliest edition, were to present current concepts in immunology cogently and also in sufficient

detail that they would be understood by students of

the discipline, as well as to emphasize clinical aspects,

including disease pathogenesis and the development of

novel therapies based on the basic science of immunology. These are the goals that we continue to strive for.

With improving understanding of the normal immune

response, we believe it is possible to present the fundamental knowledge in a concise way. In addition, there

has been exciting progress in applying basic principles

to understanding and treating human diseases, a topic

that is of paramount interest for students of medicine

and allied health sciences. Foremost among these recent

advances is the development of cancer immunotherapy,

which dramatically illustrates how foundational science

can be translated into clinical practice.

More specifically, we have focused on the following

objectives. First, we have presented the most important

principles governing the function of the immune system by synthesizing key concepts from the vast amount

of experimental data that have emerged in the field of

immunology. Our judgment of what is most important

is based largely on what is most clearly established by

scientific investigation and is essential for understanding the major functions of the immune system. We

have also prioritized content that is relevant to human

health and disease. We have realized that in any concise discussion of complex phenomena, it is inevitable

that exceptions and caveats cannot be considered in

detail, so these have largely been omitted. Second, we

have focused on immune responses against infectious

microbes, and most of our discussions of the immune

system are in this context. Third, we have made liberal

use of illustrations to highlight important principles,

but we have reduced factual details that may be found

in more comprehensive textbooks. Fourth, we have

also discussed immunologic diseases from the perspective of principles, emphasizing their relation to normal

immune responses and avoiding details of clinical syndromes and treatments. We have included selected clinical cases in an appendix to illustrate how the principles

of immunology may be applied to common human diseases. Finally, in order to make each chapter readable on

its own, we have repeated key ideas in different places in

the book. We feel such repetition will help students to

grasp the most important concepts.

We hope that students will find this new edition of

Basic Immunology clear, cogent, manageable, and enjoyable to read. We hope the book will convey our sense

of excitement about how the field has evolved and how

it continues to grow in relevance to human health and

disease. Finally, although we were spurred to tackle this

project because of our associations with medical school

courses, we hope the book will be valued by students of

allied health and biology as well. We will have succeeded

if the book can answer many of the questions these students have about the immune system and, at the same

time, encourage them to delve even more deeply into

immunology.

Several individuals played key roles in the writing of

this book. Our editor, James Merritt, has been an enthusiastic source of encouragement and advice. Our talented illustrator, David Baker, continues to effectively

convert our ideas into pictures that are informative and

aesthetically pleasing. Our development editor, Rebecca

Gruliow, has kept the project organized and on track

despite pressures of time and logistics. Clay Broeker

has moved the book through the production process