The Insect Viruses THE VIRUSES Series Editors HEINZ FRAENKEL-CONRAT, University of California Berkeley, California ROBERT R. WAGNER, University of Virginia School of Medicine Charlottesville, Virginia THE VIRUSES: Catalogue, Characterization, and Classification Heinz Fraenkel-Conrat Other volumes in the series: THE BACULOVIRUSES Edited by Lois K. Miller THE BUNYAVIRIDAE Edited by Richard M. Elliott THE CORONAV IRIDAE Edited by Stuart G. Siddell THE INSECT VIRUSES Edited by Lois K. Miller and L. Andrew Ball THE PA POVAV IRIDAE Volume 1 • Edited by Norman P. Salzman Volume 2· Edited by Norman P. Salzman and Peter M. Howley THE PARAMYXOVIRUSES Edited by David W. Kingsbury THE PARVOVIRUSES Edited by Kenneth I. Berns THE PLANT VIRUSES Volume 1 • Edited by R. I. B. Francki Volume 2· Edited by M. H. V. Van Regenmortel and Heinz Fraenkel-Conrat Volume 3 • Edited by Renate Koenig Volume 4· Edited by R. G. Milne Volume 5 • Edited by B. D. Harrison and A. F. Murant THE REOVIRIDAE Edited by Wolfgang K. Joklik THE RETROVIRIDAE Volumes 1-4 • Edited by Jay A. Levy THE RHABDOVIRUSES Edited by Robert R. Wagner THE TOGAV IRIDAE AND FLAV IVIRIDAE Edited by Sondra Schlesinger and Milton J. Schlesinger THE VIROIDS Edited by T. O. Diener A complete listing of volumes in this series appears at the back of this volume. The Insect Viruses Edited by LOIS K. MILLER The University of Georgia Athens, Georgia and L. ANDREW BALL University of Alabama at Birmingham Brrmingham,iUabama SPRINGER SCIENCE+BUSINESS MEDIA, LLC Llbrarv of Congre •• C8taloglng-ln-Publlcatlon Data Tha Insact vlrusas I adlted by Lols K. Mlller and L. Andraw BaII. p. cm. -- (Vlrusas) Includas blbllographlcal rafarancas and Index. ISBN 978-1-4613-7437-4 ISBN 978-1-4615-5341-0 (eBook) DOI 978-1-4615-5341-0 1. Insacts--Vlrusas. 1. Mlllar, Lols K. II. BaII, L. Andraw (Lauranca Andrew), 1944- III. Sarlas. CR327.I57 1998 579.2·3--dc~1 98-27923 CIP ISBN 978-1-4613 -7437-4 ©1998 Springer Science+Business Media New York Original1y published by Plenum Press, New York in 1998 lO987654321 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, mechanica1, photocopying, microfilming, recording, or otherwise, without written permission from the Publisher Contributors 1. Andrew Ball, Department of Microbiology, University of Alabama at Birmingham, Birmingham, Alabama 35294 Barry Beaty, Arthropod-borne and Infectious Diseases Laboratory, Department of Microbiology, Colorado State University, Fort Collins, Colorado 80523-1682 Serge Belloncik, Centre de Recherche en Virologie, Institut Armand-Frappier, Laval H7N 4Z3, Canada Max Bergoin, Station de Recherches de Pathologie Comparee, INRA-UA CNRS 1184, 30380 Saint Christol-Iez-Ales, France John P. Burand, Departments of Entomology and Microbiology, University of Massachusetts-Amherst, Amherst, Massachusetts 01003 Peter D. Christian, CSIRO Division of Entomology, Canberra, ACT 2601, Australia Karl H.J. Gordon, CSIRO Division of Entomology, Canberra, ACT 2601, Australia Terry N. Hanzlik, CSIRO Division of Entomology, Canberra, ACT 2601, Australia Stephen Higgs, Arthropod-borne and Infectious Diseases Laboratory, Depart- ment of Microbiology, Colorado State University, Fort Collins, Colorado 80523-1682 John E. Johnson, Department of Molecular Biology, The Scripps Research Institute, La Jolla, California 92117 Kyle L. Johnson, Department of Microbiology, University of Alabama at Birmingham, Birmingham, Alabama 35294 Linda A. King, School of Biological and Molecular Sciences, Oxford Brookes University, Gipsy Lane Campus, Oxford OX3 OBP, United Kingdom Susan A. Marlow, School of Biological and Molecular Sciences, Oxford Brookes University, Gipsy Lane Campus, Oxford OX3 OBP, United Kingdom Davin P. Miller, School of Biological and Molecular Sciences, Oxford Brookes University, Gipsy Lane Campus, Oxford OX3 OBP, United Kingdom Lois K. Miller, Departments of Entomology and Genetics, The University of Georgia, Athens, Georgia 30602 Hajime Mori, Kyoto Institute of Technology, Kyoto, Japan v vi CONTRIBUTORS Kenneth Olson, Arthropod-borne and Infectious Diseases Laboratory, Depart- ment of Microbiology, Colorado State University, Fort Collins, Colorado 80523-1682 Vijay Reddy, Department of Molecular Biology, The Scripps Research Institute, La Jolla, California 92117 Paul D. Scotti, The Horticulture and Food Research Institute of New Zealand Ltd., Mt. Albert Research Centre, Auckland, New Zealand Peter Tijssen, Centre de Recherche en Virologie, Institut Amand-Frappier, Universite du Quebec, Laval-des-Rapides, Quebec City, H7N 4Z3, Canada Bruce A. Webb, Department of Entomology, University of Kentucky, Lexing- ton, Kentucky 40546 Nicola Wilkinson, School of Biological and Molecular Sciences, Oxford Brookes University, Gipsy Lane Campus, Oxford OX3 OBP, United Kingdom Trevor Williams, ECOSUR, Tapachula 30700, Chiapas, Mexico Preface There are over a million different species of insects, and individually they outnumber humans by more than 108 to 1. Moreover, some insects live in close association with both plants and higher animals and naturally exchange viruses with them. It has even been speculated that viruses in general may have radiated through the plant and animal kingdoms from common insect origins. Be that as it may, since insects play pivotal roles in the biosphere, both to the benefit and detriment of mankind, they and the viruses that infect them are important subjects for study. Insects are infected by a diverse medley of viruses, and this volume focuses on those insect virus families that are found primarily or exclusively in insects. All major families of insect-selective viruses are covered except for the baculo- viruses, which were described in a separate volume of The Viruses series. Included in this volume are the established families of insect viruses, the newly recognized ascovirus family, and the nudiviruses, which probably represent a separate family but currently remain unclassified. The coverage of arboviruses is limited to a single chapter that focuses on their potential utility as vectors and in insect control. Omitted for practical reasons are several individual insect viruses that have been described sporadically in the literature and probably represent novel virus families. Insect viruses provide novel insights into virus evolution. The polydna- viruses, for example, appear to have a symbiotic relationship with their insect hosts, parasitic wasps. The viruses are vertically transmitted and provide a survival advantage to the wasp by abrogating the protective response of its host. The nudivinises and ascoviruses may also have a complex mutualistic relation- ship with their hosts. The entomopoxviruses and insect iridoviruses are dis- tinctive permutations of their mammalian counterparts, while the tetraviruses and nodaviruses provide novel variations on the positive-strand ribovirus strat- egy. Understanding how all these viruses have evolved in their arthropod hosts is likely to provide important links in understanding the evolution of mam- malian viruses. Although most of the viruses covered in this volume have little direct relevance to human health, they nevertheless represent important viruses from vii viii PREFACE an environmental perspective. Viral diseases in beneficial insects can lead to ecological problems, while diseases of insect pests are often welcomed as a means of reducing economic and medical problems. Furthermore, like the baculoviruses, some of the viruses described in this volume are likely to be- come valuable tools in the escalating human efforts to control and manipulate the ecosystem. The large DNA viruses are described first, beginning with the entomopox- viruses and iridoviruses, followed by those that rely more heavily on insect vec- tors for their propagation, and then the small DNA densoviruses. These are fol- lowed by the RNA viruses, beginning with an account of the three-dimensional structures of small RNA viruses, followed by chapters describing each of the major families of insect RNA viruses, and ending with a review of the develop- ment of arbovirus expression systems and how they may be employed in the future. Lois K. Miller L. Andrew Ball Contents Chapter 1 Entomopoxviruses Linda A. King, Nicola Wilkinson, Davin P. Miller, and Susan A. Marlow I. Introduction ................................................. 1 ll. Classification . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 III. Virus Structure .............................................. 3 A. Morphology .............................................. 3 B. Structural Proteins .................. , . .. . .. . . . . . . . . . . . . . . . 3 C. Virion-Associated Enzyme Activities ....................... 4 D. Spheroids ................................................ 5 E. Spindles.................................................. 5 IV. Virus Replication ............................................ 6 A. Pathology ................................................ 6 B. Replication Cycle in Vivo ................................. 7 C. Replication Cycle in Vitro . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10 V. Entomopoxvirus Genome Structure ........................... 16 A. Entomopoxvirus Genes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17 B. Entomopoxvirus Promoters ........ . . . . . . . . . . . . . . . . . . . . . . . . 22 C. Genetic Modification of the Entomopoxvirus Genome . . . . . . . 23 VI. Biological Control ..................................... , ... " . 24 References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 25 Chapter 2 Invertebrate Iridescent Viroses Trevor Williams I. Introduction ................................................. 31 ll. Classification .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 32 ix x CONTENTS A. Problems with the Established Classification of Iridescent "Viruses ................................................... 32 B. New Changes in the Classification of Iridescent Viruses. .. . . 33 C. The Nomenclature Problem ............................... 34 III. Structure and Composition ................................... 35 A. Capsid ........." .......................................... 35 B. Lipid Layer . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . • . . . . . . 36 C. Core ............................................. "........ 37 D. Physicochemical Characteristics and Composition".......... 37 IV. Replication and Molecular Biology ............................ 38 A. Penetration and Shutdown of Macromolecular Synthesis . . . . . 39 B. DNA Replication ......................................... 39 C. Transcription, Translation, and IV Genes ................... 40 D. Particle Assembly" . . . . . . . . . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . 42 V. Signs and Characteristics of Patent Disease .................... 43 A. Pathology ................................................ 43 B. IV Production and Quantification .......................... 44 C. Dose-Response Relationships and Survival Time ........... 45 VI. Relationships among IVs ..................................... 46 A. Serology ......................................... ,....... 46 B. Genetic .................................................. 47 C. DNA Sequence Analysis .................................. 50 D. Biochemical .............................................. 52 VII. Ecology ..................................................... 52 A. Transmission .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 52 B. Persistence ............................................... 54 C. Host Range. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 57 VIII. Use of Iridescent Viruses in Biocontrol ........................ 58 References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 60 Chapter 3 Nudiviruses John P. Burand I. Introduction................................................. 69 II. Oryctes Virus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 70 A. Physical and Biological Properties of the Oryctes Virus ...... 70 B. Physical Map of the Oryctes Virus Genome ................ 71 C. Replication of Oryctes Virus in Cell Culture . . . . . . . . . . . . . . . . 71 D. Oryctes Virus Structural and Intracellular Proteins .......... 73 m. Hz-1 Virus .................................................. 73 A. Discovery and Isolation of Hz-1 V .......................... 73 B. Biochemical Properties of Hz-1 V ........................... 74 C. Persistence of Hz-! V ...................................... 75