Yechiel Becker, Series Editor Julia Hadar, Managing Editor

Transcription

1 A VIAN LEUKOSIS

2 DEVELOPMENTS IN VETERINARY VIROLOGY Yechiel Becker, Series Editor Julia Hadar, Managing Editor Payne, L.N. (ed.) Marek's Disease (1985) Burny, A. and Mammerickx, M. (eds.) Enzootic Bovine Leukosis and Bovine Leukemia Virus (1987) Becker, Y. (ed.) African Swine Fever (1987) De Boer, O.F. (ed.) Avian Leukosis (1987)

3 AVIAN LEUKOSIS edited by G.F. De Boer Central Veterinary Institute Lelystad, The Netherlands UI " Martinus Nijhoff Publishing a member of the Kluwer Academic Publishers Group Boston/DordrechtiLancaster

4 Distributors for North America: Kluwer Academic Publishers 101 Philip Drive Assinippi Park Norwell, Massachusetts 02061, USA Distributors for the UK and Ireland: Kluwer Academic Publishers MTP Press Limited Falcon House, Queen Square Lancaster, LAI IRN, UNITED KINGDOM Distributors for all other countries: Kluwer Academic Publishers Group Distribution Centre Post Office Box AH Dordrecht, THE NETHERLANDS Library of Congress Cataloging-in-Publication Data Boer, G. F. de. Avian leukosis. (Developments in veterinary virology) Includes bibliographies and index. I. Avian leukosis. I. Title. II. Series. SF995.6.L4B '089' ISBN-13: e-isbn-13: DOl: / Copyright 1987 by Martinus Nijhoff Publishing, Boston Softcover reprint of the hardcover 1st edition 1987 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, mechanical, photocopying, recording, or otherwise, without the prior written permission of the publisher, Martinus Nijhoff Publishing, 101 Philip Drive, Assinippi Park, Norwell, Massachusetts

5 CONTENTS LIST OF CONTRIBUTORS PREFACE viii ix 1. A historical review of avian retrovirus research 1 R.M. Dougherty 1. The origins of avian retrovirus research Avian retrovirus research during the period Contemporary research with avian retroviruses Concluding remarks Structure and virus-associated oncogenes of avian sarcoma 29 and leukemia viruses P.J. Enrietto and M.J. Hayman 1. Introduction Defective leukemia viruses Avian sarcoma viruses Concluding remarks Epizootiology of avian leukosis virus infections 47 L.N. Payne 1. Introduction Virus classi fication Incidence and prevalence Modes of natural transmission Infection patterns within flocks Viral factors influencing infection and disease Host genetic factors influencing infection and disease Other host factors influencing infection and disease Environmental factors influencing infection and disease Molecular basis of oncogenesis by non-acute avian retro\"iruses 77 H.-J. Kung and N.J. Maihle 1. Introduction Structure and replication of non-acute retroviruses The structure of the long terminal repeat (L TR) Possible schemes for host oncogene activation by non-acute retroviruses B-lymphoma induction by ALV: c-myc activation B-lymphoma induction by REV: c-myc activation Erythroblastosis induction by AL V: c-erbb activation RPV-induced neoplasms MAV-induced neoplasms Concluding remarks Endogenous avian leukemia viruses 101 E.J. Smith 1. Introduction Evidence that normal chickens harbor avian retroviral genes The endogenous AL V genome Expression of ev genes Transcriptionalcontrol elements within the long terminal repeat Chromosomal distribution of ev genes 110

6 vi 7. Significance of ev genes among avian species 8. Genetic diversifyamong avian retroviruses 9. Biological influences of endogenous proviruses 6. Immunology of avian leukosis virus infections R.E. Smith 1. Introduction 2. Neoplastic transformation of lymphoid cells 3. Immune response to AL V infection 4. Immunosuppression III Avian sarcomas: immune responsiveness and pathology 131 M.A. Wainberg and M.S. Halpern 1. Introduction Cellular transformation by avian sarcoma viruses Viral-associated antigens ASV-induced pathogenesis and immunosuppression Development of anti-tumor immunity in tumor-bearing hosts The immunogenicity of retroviral envelope glycoprotein Characteristics of tumor cell growth in vh"o and in vitro Virus-cell interactions of avian sarcoma and defective leukemia viruses 153 C. Moscovici and L. Gazzolo 1. Introduction Avian sarcoma viruses Pathogenic effects of avian sarcoma viruses in vivo Transforming effects of avian sarcoma viruses in vitro Defective avian leukemia viruses Conclusions and future trends The pathogenesis and pathology of neoplasms caused by avian leukosis viruses H.G. Purchase 1. Introduction 2. Virus classification 3. Factors affecting the pathogenic response 4. Classification of leukemias and leukoses 5. Other tumors and tumorous conditions 6. Non-neoplastic conditions 7. Transplantability and development of cell lines 8. Lymphoid leukosis 9. Erythroblastosis 10. Myeloblastosis 11. Nephroblastomatosis 12. Osteopetrosis 13. Concluding remarks 10. Differential diagnosis of lymphoid leukosis A.M. Fadly 1. Introduction 2. Clinical signs 3. Etiology 4. Pathogenesis 5. Gross pathology

7 vii 6. Microscopic pathology 7 Serological and immunological techniques Laboratory diagnostic procedures for detecting avian leukosis virus infections 213 J.L. Spencer 1. Introduction Collection and handling of specimens In vivo assay of RSV and AL V In vitro assay of RSV and AL V Production of virus stocks and antiserum Tests based on interaction between AL V and RSV Tests for AL V based on detection of group-specific antigens Detection and distribution of viral antigens in host tissues Detection of antibody against subgroup- and group-specific antigens Concluding remarks Influences of avian leukosis virus infection on production and mortality and the role of genetic selection in the control of lymphoid leukosis 241 J.S. Gavora 1. Introduction Frequency of AL V infection Effects of AL V infection on production and mortality Prospects for the future Approaches to control avian lymphoid leukosis 261 G.F. de Boer 1. Introduction Detection of congenital ALV transmission Prevention of horizontal virus exposure LL control by elimination of ALV shedding hens Manipulation of the immune response Genetic resistance Concluding remarks 282 INDEX 287

8 viii LIST OF CONTRIBUTORS 1. R.M. DOUGHERTY M.S. HALPERN Department of Microbiology Wistar Institute of Anatomy and State University of New York Biology Upstate Medical Center 36th Street at Spruce 766 Irving Avenue, Syracuse Philadelphia, Pennsylvania 19104, USA New York 13210, USA 8. C. MOSCOVICI 2. P.J. ENRIETTO Tumor Virology Laboratory Department of Microbiology Research Service, VAMC, and State University of New York Department of Pathology at Stony Brook College of Medicine Stony Brook, New York 11794, USA University of Florida Gainesville, Florida 32602, USA M.J. HAYMAN Department of Microbiology L. GAZZOLO State University of New York Laboratoire d'epidemiologie et Stony Brook, New York 11794, USA d'immunovirologie des Tumeurs CNRS, Faculte de Medecine A. Carrel 3. L.N. PAYNE Rue G. Para din Houghton Poultry Research Station Lyon, France Houghton, Huntingdon Cambs PE17 2DA, England 9. H.G. PURCHASE US Department of Agriculture 4. H.-J. KUNG Agricultural Research Service Department of Molecular Biology Beltsville Area and Microbiology Beltsville, Maryland 20705, USA Case Western Reserve University School of Medicine 10. A.M. FADLY Cleveland, Ohio 44106, USA Regional Poultry Research Laboratory 3606 East Mount Hope Road N.J. MAIHLE East Lansing, Michigan 48823, USA Department of Molecular Biology and Microbiology 11. J.L. SPENCER Case Western Reserve University Animal Diseases Research Institute School of Medicine P.O. Box Cleveland, Ohio 44106, USA Station H, Nepean Ontario K2H 8P9, Canada 5. E.J. SMITH Regional Poultry Research Laboratory 3606 East Mount Hope Road 12. J.S. GAVORA East Lansing, Michigan 48823, USA Animal Research Centre Agriculture Canada, Building R.E. SMITH Central Experimental Farm Department of Microbiology and Ottawa, Ontario KIA OC6, Canada Environmental Health Colorado State University 13. G.F. DE BOER Fort Collins, Colorado 80523, USA Central Veterinary Institute Department of Virology 7. M.A. WAINBERG P.O. Box 365 Lady Davis Institute for Medical 8200 AJ Lelystad, The Netherlands Research, Sir Mortimer B. Davis - Jewish General Hospital 3755 Cote St. Catherine Road Montreal, Quebec H3T le2, Canada

9 PREFACE The intention of the series Developments in Veterinary Virology is to provide monographs dealing with the major animal viral diseases. Each volume will include the latest achievements in fundamental research and practical applications and should be readable for people from various disciplines and different backgrounds. The multi-author approach provides the best opportunity to keep each chapter at the highest level and makes the composition of the volumes manageable to the editors. This monograph on Avian Leukosis presents comprehensive reviews on the recent history of avian retrovirus research, on epizootiological, virological, pathological aspects, on tumor induction, the immune response to avian retroviruses, virus-cell interactions and on techniques for diagnosis. The volume deals mainly with exogenous avian leukosis virus (ALV) infections, but one chapter is entirely devoted to endogenous avian leukemia virus. Molecular biology aspects are confined to various oncogenes and to lymphoma induction since retroviruses, including those specific for avian species, have recently been described in detail in the Cold Spring Harbor Laboratory series "Molecular Biology of Tumor Viruses". Two chapters are devoted to the practical application of insights obtained from avian leukosis research: influences of AL V infection on production performance and eradication procedures. The study of avian retrovirus-induced neoplasia led to important developments in the understanding of the nature and development of various tumors in avian species and has also provided, as a result of comparative research, insight into the structure and biology of retroviruses, virus-cell interactions, oncogenes and neoplasia in general. Current knowledge of the viral etiology of tumors, oncogenes, in vitro transformation, defective genomes, helper viruses and endogenous viruses (to name only a few) is for a considerable part based on work performed in the avian system. The genetically defined resistance (or susceptibility) at the cellular level to the various viral subgroups has complicated avian leukemia research, but has at the same time made the research projects of greater scientific interest. Seventeen authors, whose names are listed on a previous page, have contributed to this monograph. I gratefully acknowledge the willingness of all sixteen experts in the various subdisciplines to add a chapter to this volume. Their

10 x pre-eminent cooperation in the joint effort was very encouraging. The final product attained a quality that probably no single one of us could have generated for the whole field on his own. The communication between various authors was of great help in expressed. avoiding overlaps between chapters or conflicting opinions The illustrations were kindly prepared by Johan IJzerman, an amateur artist working in our laboratory as a technician. The cover design symbolizes the crucial moment in the epizootiology of ALV: congenital infection may have been established and horizontal spread of virus may start with great intensity. Finally, it is my privilege to thank, on behalf of all the authors, all colleagues who assisted in scrutinizing the various chapters of this monograph. One or more manuscripts were kindly reviewed by Drs. T.J. Bagust, K. McCullough, A.J. van der Eb, J. Hadar, J. Ignjatovic, J.H. van Middelkoop, J.M. Phillips, H.G. Purchase and M.A. Wainberg, and, from my own laboratory, J.G. van Bekkum, R. de Bruin, G. Koch, R.J.M. Moormann and P.W. van DIm. The figures in chapter 2 were kindly computer-designed by Dr. H. Vreeswijk. The photographs in a number of chapters were remade by Mr. K. Dekker and Mr. A.P.M. van Alphen. Above all it is my pleasure to thank Mrs. Ria de Kok-Heuckeroth and Mrs. Jacqueline C. Hoogeveen-Hilhorst who mastered the word processor. This new administrative gadget created conditions in which several versions of all chapters could be typed by our administrative staff with increasing interest in the editing activities, and at the same time they provided significant editorial assistance. r hope that the comprehensive information on avian leukosis virus infection in this monograph will be of value to research workers in various disciplines, to teachers and students, and also to veterinarians, poultrymen and vaccine manufacturers who need this information for practical applications. G.F. de Boer.

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