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Fenner and White's Medical Virology Fenner and White’s Medical Virology Fenner and White’s Medical Virology Fifth Edition Christopher J. Burrell School of Molecular and Biomedical Science, University of Adelaide, Adelaide, South Australia, Australia Colin R. Howard College of Medicine and Dentistry, University of Birmingham, United Kingdom Frederick A. Murphy University of Texas Medical Branch, Galveston, Texas, USA AMSTERDAM • BOSTON • HEIDELBERG • LONDON NEW YORK • OXFORD • PARIS • SAN DIEGO SAN FRANCISCO • SINGAPORE • SYDNEY • TOKYO Academic Press is an imprint of Elsevier Academic Press is an imprint of Elsevier 125 London Wall, London EC2Y 5AS, United Kingdom 525 B Street, Suite 1800, San Diego, CA 92101-4495, United States 50 Hampshire Street, 5th Floor, Cambridge, MA 02139, United States The Boulevard, Langford Lane, Kidlington, Oxford OX5 1GB, United Kingdom Copyright © 2017, 2011, 1999 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). Notices Knowledge and best practice in this field are constantly changing. As new research and experience broaden our understanding, changes in research methods, professional practices, or medical treatment may become necessary. Practitioners and researchers must always rely on their own experience and knowledge in evaluating and using any information, methods, compounds, or experiments described herein. In using such information or methods they should be mindful of their own safety and the safety of others, including parties for whom they have a professional responsibility. To the fullest extent of the law, neither the Publisher nor the authors, contributors, or editors, assume any liability 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. British Library Cataloguing-in-Publication Data A catalogue record for this book is available from the British Library Library of Congress Cataloging-in-Publication Data A catalog record for this book is available from the Library of Congress ISBN: 978-0-12-375156-0 For Information on all Academic Press publications visit our website at https://www.elsevier.com Publisher: Sara Tenney Acquisition Editor: Jill Leonard Editorial Project Manager: Pat Gonzalez Production Project Manager: Julia Haynes Designer: Matt Limbert Typeset by MPS Limited, Chennai, India Dedication This book is dedicated to our friends Frank J. Fenner (1914– an independent scientific and medical discipline and a 2010) and David O. White (1931–2004), the founders of the significant contributor to human, animal, and environmental series of books now spanning five editions of MEDICAL well-being. We hope our students will come to understand VIROLOGY and five editions of VETERINARY VIROLOGY. the “big picture” of medical virology as well as Frank and They set a standard of scholarship that is impossible to David achieved throughout their outstanding careers. match and a joie de vivre that made the writing and editing We would also like to dedicate this book to our wives almost fun. and families, our teachers and mentors, together with our They taught us that the subject of virology must be seen students, all of whom have shaped our thinking and provided within the context of society as a whole as well as within the us inspiration over the years in so many different ways. context of science. They envisioned virology as extending broadly, from its roots in the science of microbiology and Christopher J. Burrell the practice of infectious disease, through its key roles in Colin R. Howard the development of molecular and cell biology, to become Frederick A. Murphy Foreword To say that this new version of the classic Medical Virology institutional structures that they established, or the insights text by Frank Fenner (1914–2010) and David White (1931– they developed. My current office is in the Department 2004) is timely is, to say the least, an understatement. It’s of Microbiology and Immunology at the University of been 21 years since the fourth edition, with authors Chris Melbourne, which was headed for many years by David Burrell, Colin Howard, and Fred Murphy building on that White. Perhaps, as Isaac Newton (1642–1726) said “we stand much respected foundation to produce a greatly refreshed on the shoulders of giants” though, when I’ve met the “giants” fifth edition. The basic philosophy and structure that made of science, they’ve pretty much turned out to be hard working, this such a useful volume for many generations of advanced smart, and dedicated people who quietly and systematically medical students, clinicians, and pathologists remain but, as built a body of knowledge and a tradition of scholarship. That all who have been even peripherally involved in molecular very much describes the discipline of virology. science over the past two decades will realize, there was an Thus, while reflecting on the up-to-date understanding enormous amount of new information that had to be read, of the authors, and of other researchers who have provided sifted, and selected for relevance to the information needs key insights concerning each particular pathogen, Fenner of this target audience. and White’s Medical Virology reflects a cumulated wisdom Basic virology has, of course, been well served by the that goes back to the early days of 20th century virology. many revisions of Bernie Fields’ (1938–1995) exhaustive Back then, the thinking and technical range of mammalian text and, following the example set by Field’s Virology, the virologists was very much focused on pathogenesis and fifth edition will appear as Fenner and White’s Medical disease process. Now, after decades of illuminating the Virology. It is a fitting tribute. Taken together, the original molecular characteristics of viruses and infected cells, it authors, and those responsible for this latest version, have is gratifying to see these precise investigative tools being variously been active and publishing on one or the other applied to the understanding of what actually happens in aspect of virus-induced disease and/or pathology since an infected individual. This is a great time for any medical 1948: and the virology lineage goes back even further! researcher who is fascinated by the linked issues of disease Frank Fenner was the student of MacFarlane Burnet process, pathology, and treatment. (1899–1985) who, applying technical and conceptual This fifth edition provides an accessible and informative approaches learned from his early studies with bacteriophage, account of medical virology, based on both contemporary pioneered quantitative mammalian virology, and genetics. science and what went before. It constitutes an excellent David White began his career as a virologist in the university resource for both the physician and the research investigator. microbiology department headed by Frank Fenner. Both Fred New discoveries are constantly being made and, while Murphy and I worked for a time and Chris Burrell completed much remains unclear, the rate of advance in understanding his PhD in that same department, where Rolf Zinkernagel and is extraordinary. Hopefully, we will not need to wait another I did the experiments in viral immunity that led to a Nobel 20 years for the next edition of this seminal text. Prize. Reflecting on lineages and the solid foundations that enable the work of subsequent generations, I realize that I’ve Peter C. Doherty met all the people named above in one or other professional University of Melbourne or personal context and have benefitted from either the December 10, 2015 xvii Preface Enormous strides have taken place since the fourth edition of responsible. Much of the approach adopted has been shaped this book in regard to our understanding of virus properties by the authors’ experience in teaching virology to science and how viral diseases can be controlled, treated, and and medical students and graduates—which has itself been prevented. Considerable advances have been made across shaped by earlier editions of this book. the whole spectrum of virology, particularly in relation to This edition is designed to meet the needs of advanced virus structure and replication, “point-of-care” diagnostics medical students, clinicians and pathologists, university and the wide use of antiviral therapies. This progress has teachers, researchers, and public health workers who are done much to contribute to a substantial improvement in seeking a single accessible source of key information about public health across the globe. the range of major human viruses and the options for treatment At the same time, we have seen the emergence of new and control. We have included historical perspectives, so that viral agents, and older agents in new guises, sometimes readers gain some insight into the paths and the personalities creating a need for radically
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