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The Reoviridae THE VIRUSES Series Editors HEINZ FRAENKEL-CONRAT, University of California Berkeley, California ROBERT R. WAGNER, University of Virginia School of Medicine Charlottesville, Virginia THE HERPESVIRUSES, Volumes I, 2, 3, and 4 Edited by Bernard Roizman THE REOVIRIDAE Edited by Wolfgang K. Joklik THE PARVOVIRUSES Edited by Kenneth I. Berns The Reoviridae Edited by WOLFGANG K. JOKLIK Duke University Medical Center Durham, North Carolina Springer Science+Business Media, LLC Library of Congress Cataloging in Publication Data Main entry under title: The Reoviridae. (Viruses) Includes bibliographical references and index. 1. Reoviruses. L Joklik, Wolfgang K. II. Series. QR414.R46 1983 576/.64 83-6276 ISBN 978-1-4899-0582-6 ISBN 978-1-4899-0582-6 ISBN 978-1-4899-0580-2 (eBook) DOI 10.1007/978-1-4899-0580-2 © Springer Science+Business Media New York 1983 Originally published by Plenum Press, New York in 1983 Softcover reprint of the hardcover 1st edition 1983 All rights reserved No part of this book may be reproduced, stored in a retrieval system, or transmitted in any form or by any means, electronic, mechanical, photocopying, microfilming, recording, or otherwise, without written permission from the Publisher Con tribu tors Guido 'Boccardo, Istituto di Fitovirologia Applicata del C.N.R., 10135 To­ rino, Italy Bernard N. Fields, Department of Microbiology and Molecular Genetics, Harvard Medical School, Boston, Massachusetts 02115; and Depart­ ment of Medicine, Brigham and Women's Hospital, Boston, Massa­ chusetts 02115 R. I. B. Fran'cki, Department of Plant Pathology, Waite Agricultural Re­ search Institute, The University of Adelaide, Adelaide 5064, South Australia Barry M. Gorman, Queensland Institute of Medical Research, Bramston Terrace, Brisbane 4006, Australia Ian H. Holmes, Department of Microbiology, University of Melbourne, Parkville, Victoria 3052, Australia Wolfgang K. Joklik, Department of Microbiology and Immunology, Duke University Medical Center, Durham, North Carolina 27710 Marilyn Kozak, Department of Biological Sciences, University of Pitts­ burgh, Pittsburgh, Pennsylvania 15260 Peter P. C. Mertens, Animal Virus Research Institute, Pirbright, Woking, Surrey GU24 ONF, United Kingdom Stewart Millward, Department of Biochemistry, McGill University, Mon­ treal, Quebec, Canada H3G 1Y6 Christopher C. Payne, Glasshouse Crops Research Institute, Littlehamp­ ton, West Sussex BN16 3PU, United Kingdom Robert F. Ramig, Department of Virology and Epidemiology, Baylor Col­ lege of Medicine, Houston, Texas 77030 Arlene H. Sharpe, Department of Microbiology and Molecular Genetics, Harvard Medical School, Boston, Massachusetts 02115 Aaron J. Shatkin, Department of Cell Biology, Roche Institute of Mole­ Biology, Nutley, New Jersey 07110 v vi CONTRIBUTORS Jill Taylor, Queensland Institute of Medical Research, Bramston Terrace, Brisbane 4006, Australia Peter J. Walker, Queensland Institute of Medical Research, Bramston Ter­ race, Brisbane 4006, Australia Helmut Zarbl, Department of Biochemistry, McGill University, Mon­ treal, Quebec, Canada H3G 1Y6 Preface It is now just 20 years since Gomatos and his co-workers at the Rocke­ feller University showed that the nucleic acid in reovirus particles is double-stranded RNA (dsRNA). This discovery created great excitement, for dsRNA was at that time under intense investigation as the replicative form of viral genomes consisting of single-stranded RNA. An equally interesting and important finding followed soon after: it was found that the reovirus genome consists, not of a single nucleic acid molecule, but of 10 discrete "segments," each with its specific sequence content and each transcribed into its own messenger RNA. It is clear now that these segments are genes. Not surprisingly, the availability of a viral genome consisting of 10 unlinked genes has permitted some unique lines of in­ vestigation in molecular biology. Mammalian and avian reoviruses proved to be but the first of several viruses recognized as sharing Similarity in size and morphology and ge­ nomes consisting of 10, II, or 12 separate genes. These viruses are dis­ tributed throughout living organisms; among the natural hosts of mem­ bers of this virus family are vertebrates, Insects, and plants. Members of the Reoviridae family differ widely in the virulence that they exhibit toward their hosts . .For example, the first discovered mam­ malian reovirus literally is, as the name signifies, a "respiratory enteric orphan" virus, that is, a virus unassociated with disease. In contrast, the most recently discovered members of this family, the rotaviruses, which were recognized as comprising a separate genus not much more than a decade ago, cause a disease, infantile gastroenteritis, that, while not se­ vere in communities where health care is good, is one of the most im­ portant, if not the most important, cause of death in infants under the age of 2 years in less developed countries. Studies of this family of viruses from molecular, biological, and medical perspectives complement each other to an unusual degree because of the unique nature of the genomes of these viruses. The ten chapters in this book describe what is currently known concerning the molecular biology of members of this virus family on the vii viii PREFACE one hand and the pathogenesis that they exhibit toward their ·hosts on the other. About half the chapters are devoted to the mammalian reo­ viruses, which are the Reoviridae that have been studied most intensively in molecular and genetic terms; there are chapters that deal with the nature of reovirus particles and of the reovirus genome, its transcription and replication, the reovirus multiplication cycle, and reovirus genetics. There is also a chapter that describes reovirus pathogenesis in terms of the effects that various individual reovirus gene products exert on host cells. Other chapters deal with the orbiviruses, which include some im­ portant animal pathogens such as bluetongue virus; the rotaviruses, im­ portant human pathogens; the cytopla~mic polyhedrosis viruses of in­ sects; and the plant reoviruses, which include some important plant pathogens. Wolfgang K. Joklik Contents Chapter 1 The Members of the Family Reoviridae Wolfgang K. Joklik I. Introduction ........................................................................... 1 n. Properties of Members of the Family Reoviridae ................... 2 A. Possession of Double-Stranded RNA ................................. 2 B. Structure ........................................................................... 2 C. Nature of the Genome ...................................................... 3 D. Lack of Complete Uncoating ........ .......... ... ... .... ......... ....... 4 E. Possession of Enzymes That Transcribe the Double- Stranded RNA into Messenger RNA .. ... ................ ............ 4 III. Unclassified Members of the Family Reoviridae .................... 5 A. Drosophila Virus F and Ceratitis Virus I ... .... ... .... ...... ...... 5 B. Housefly Virus .................................................................. 5 C. Chum Salmon Virus ..... ...... .... ................ .......... ...... ...... .... 6 IV. Other Viruses That Contain Double-Stranded RNA ....... ... .... 6 References ...................................................................................... 6 Chapter 2 The Reovirus Particle Wolfgang K. Joklik I. Introduction ........................................................................... 9 n. Morphology ... .......... .......................... ...... .......... .......... ........... 12 A. Reovirus Particles ............................................................. 12 B. Cores ................................................................................. 14 C. Top-Component Particles ................................................. 16 m. Purification ............................................................................ 17 IV. Measurement ......................................................................... 17 A. Plaque Formation .............................................................. 17 B. Hemagglutination ............................................................. 18 ix x CONTENTS C. Enumeration of Virus Particles by Measurement of Optical Density................................................................ 18 V. Reovirus Genome .................................................................. 19 VI. Proteins of Reovirus ............................................................... 22 A. Outer Capsid Shell ............................................................ 25 B. Inner Capsid Shell: The Core ............................................ 28 C. Nonstructural Reovirus Proteins ...................................... 30 D. Modification of Reovirus Proteins ..................................... 31 E. Mechanisms of Chemical and Physical Inactivation of Reovirus Particles ............................................... ;............. 32 VII. Information Content of the Ten Reovirus Genes ................... 33 VIII. Genetic Relatedness among the Three Serotypes of Reovirus ................................................................................. 35 IX. Nature of the Antibodies Elicited by Reovirus-Specified Proteins .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. 40 X. Sequence of Reovirus Genes and Messenger RNAs ............... 44 A. Sequence Identity of the Plus Strands of Reovirus Genes and of Their Transcripts

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