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The Papovaviridae Volume 2 TIlE PAPILLOMA 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
THE ADENOVIRUSES Edited by Harold S. Ginsberg
THE HERPESVIRUSES Volumes 1-3 • Edited by Bernard Roizman Volume 4 • Edited by Bernard Roizman and Carlos Lopez
THEPAPOVAVIRIDAE Volume 1 • Edited by Norman P. Salzman Volume 2 • Edited by Norman P. Salzman and Peter M. Howley
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
THE REOVIRIDAE Edited by Wolfgang K. Joklik
THE RHABDOVIRUSES Edited by Robert R. Wagner
THE TOGA VIRIDAE AND FLAVIVIRIDAE Edited by Sondra Schlesinger and Milton J. Schlesinger
THE VIROIDS Edited by T. o. Diener
The Papovaviridae Volume 2 THE PAPILLOMAVIRUSES
Edited by
NORMAN P. SALZMAN and
PETER M. HOWLEY National Institutes of Health Bethesda, Maryland
PLENUM PRESS • NEW YORK AND LONDON
Library of Congress Cataloging in Publication Data
(Revised for volume 2)
The papovaviridae.
(The Viruses) Volume 2 edited by Norman P. Salzman and Peter M. Howley. Includes bibliographies and index. Contents: v. 1. The polyomaviruses- The papillomaviruses. 1. Papovaviruses-Collected works. I. Salzman, Norman P. II. Howley, Peter M.
III. Series: The Viruses. [DNLM: 1. Papovaviridae. QW 165.5.P2 P218j QR406.P36 1986 576'.6484 86-15160 ISBN 978-1-4757-0586-7 ISBN 978-1-4757-0584-3 (eBook) DOI 10.1007/978-1-4757-0584-3
© 1987 Plenum Press, New York Softcover reprint of the hardcover 1st edition 1987
A Division of Plenum Publishing Corporation 233 Spring Street, New York, N.Y. 10013
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
Contributors
Eberhard Amtmann, Institute for Virus Research, German Cancer Research Center, 6900 Heidelberg, Federal Republic of Germany
Harri Ahola, Department of Medical Genetics, University of Uppsala, Biomedical Center, S-751 23 Uppsala, Sweden
Carl C. Baker, Laboratory of Tumor Virus Biology, National Cancer Institute, Bethesda, Maryland 20892
Daniel DiMaio, Department of Human Genetics, Yale University School of Medicine, New Haven, Connecticut 06510
Peter M. Howley, Laboratory of Tumor Virus Biology, National Cancer Institute, Bethesda, Maryland 20892
Robert F. LaPorta, Department of Oral Biology and Pathology, School of Dental Medicine, State University of New York, Stony Brook, New York 11794
Jorge Moreno-Lopez, Department of Veterinary Microbiology (Virology), Swedish University of Agricultural Sciences, Biomedical Center, S-751 23 Uppsala, Sweden
Carl Olson, Department of Veterinary Science, University of Wisconsin, Madison, Wisconsin 53706
Gerard Orth, Unite des Papillomavirus, Unite INSERM 190, Institut Pasteur, 75724 Paris Cedex IS, France
Ulf Pettersson, Department of Medical Genetics, University of Uppsala, Biomedical Center, S-751 23 Uppsala, Sweden
Herbert Pfister, Institut fiir Klinische Virologie, Universitiit ErlangenNiirnberg, 0-8520 Erlangen, Federal Republic of Germany
Richard Schlegel, Laboratory of Tumor Virus Biology, National Cancer Institute, Bethesda, Maryland 20892
Achim Schneider, Sektion fiir Gyniikologische Zytologie und Histologie, Universitiits-Frauenklinik, 7900 Ulm, Federal Republic of Germany
Bettie M. Steinberg, Department of Otolaryngology, Long Island Jewish Medical Center, New Hyde Park, New York 11042
v
vi CONTRIBUTORS
Arne Stenlund, Department of Medical Genetics, University of Uppsala, Biomedical Center, S-751 23 Uppsala, Sweden; Present address: Department of Molecular Biology, University of California, Berkeley, California 94720
Lorne B. Taichman, Department of Oral Biology and Pathology, School of Dental Medicine, State University of New York, Stony Brook, New York 11794
Klaus Wayss, Institute for Pathology, German Cancer Research Center, 6900 Heidelberg, Federal Republic of Germany
Felix o. Wettstein, Department of Microbiology and Immunology, UCLA School of Medicine, Los Angeles, California 90024
Harald zur Hausen, Deutsches Krebsforschungszentrum, 6900 Heidelberg, Federal Republic of Germany
Preface
In recent years there has been an explosion in research on the papillomaviruses. The viral nature of human warts was first suggested 80 years ago by Ciuffo, who demonstrated transmission using cell-free filtrates. Shope described the first papillomavirus over 50 years ago as the etiologic agent in infectious papillomatosis in rabbits. Subsequent studies by Rous established that benign rabbit papillomas induced by this virus could progress to carcinomas when treated with specific nonviral cofactors. Despite these rich beginnings, the papillomavirus field lay virtually dormant until the late 1970s because no one was able to propagate these viruses in culture successfully. In the late 1970s the molecular cloning of the papillomavirus genomes permitted investigators to partially circumvent this obstacle to their progress. The cloning of the viral genomes permitted the standardization of viral reagents and provided sufficient material to begin a systematic evaluation of the biology of this group of viruses.
This volume contains a series of chapters designed to provide a historical perspective on this field and to review the current state of research involving the papillomaviruses. Chapters 1 and 2 serve as introductory chapters, providing a general description and overview of the papillomaviruses and a historical perspective. Chapters 3-5 cover the molecular biology and cell biology of the papillomaviruses, focusing on transcription, gene expression in epithelial cells, and viral transformation. The rabbit and the Mastomys natalensis provide animal models for studying carcinogenic progression and are covered in Chapters 6 and 7. In humans, HPVs have been associated with cutaneous carcinomas in patients with epidermodysplasia verruciformis and with anogenital carcinomas; these topics are reviewed in Chapters 8 and 9. Laryngeal papillomas are associated with the same HPV types seen in the genital tract and are also capable of malignant progression, as discussed in Chapter 10. The viral DNA of the bovine papillomavirus type 1 has been developed into a
vii
viii PREFACE
mammalian cell cloning vector based on its ability to remain as a stable plasmid in transformed cells, and its use as a cloning vector is reviewed in Chapter 11. Finally, an appendix is included presenting an analysis of sequences of the viral genomes available at the time this book was completed.
Norman P. Salzman Peter M. Howley
Contents
Chapter 1
Papillomaviruses: General Description, Taxonomy, and Classification
Herbert Pfister
I. Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1 II. Properties of the Virions . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
A. Particle Structure and Protein Composition ............ 2 B. Nucleic Acid. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
III. Biological Properties. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5 A. Characteristics of the Benign Tumors . . . . . . . . . . . . . . . . . . 5 B. Immune Response of the Host. . . . . . . . . . . . . . . . . . . . . . . . . 9 C. Malignant Conversion-General Aspects . . . . . . . . . . . . . . . 10
IV. Classification and Nomenclature. . . . . . . . . . . . . . . . . . . . . . . . . . 13 V. Description of Virus Types. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15
A. Human Papillomaviruses. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15 B. Bovine Papillomaviruses ..... . . . . . . . . . . . . . . . . . . . . . . . . . 22 C. Other Papillomaviruses from Animals ................. 25
VI. Evolutionary Aspects. . . . . . . . . . . . .. . . . . . . .. . . . . . . . ... . . . . . 27 A. Papillomaviruses and Miopapovaviruses . . . . . . . . . . . . . . . . 27 B. Evolution of Papillomaviruses . . . . . . . . . . . . . . . . . . . . . . . . . 27 C. Papillomaviruses from Epidermodysplasia Verruciformis 28 D. Reservoir of Papillomaviruses ......................... 29
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 30
ix
x CONTENTS
Chapter 2
Animal Papillomas: Historical Perspectives
Carl Olson
I. Canine Oral Papillomatosis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 39 II. Shope Rabbit Papilloma .................................. 42
III. Equine Papillomatosis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 44 IV. Bovine Papillomatosis. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 45 V. Papillomatosis in Sheep .................................. 58
VI. Oral Papillomatosis of Rabbits ............................ 58 VII. Deer Fibromatosis. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 58
VIII. European Elk Papillomaviruses . . . . . . . . . . . . . . . . . . . . . . . . . . . . 59 IX. Viral Papillomatosis in Other Animals. . . . . . . . . . . . . . . . . . . . . 60
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 62
Chapter 3
Organization and Expression of Papillomavirus Genomes
Ull Pettersson, Harri Ahola, Arne Stenlund, and Jorge Moreno-Lopez
I. Introduction............................................. 67 II. General Properties of Papillomavirus Genomes . . . . . . . . . . . . . 68
III. The Genome of BPV-l ........... '" .... . .. . ..... . .. . . ... . 71 A. Organization of the E (Transforming)-Region ........... 71 B. Organization of the L-Region . . . . . . . . . . . . . . . . . . . . . . . . . . 72 C. The Noncoding Region ............................... 72 D. Transcriptional Signals. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 72 E. The Origin of DNA Replication ....................... 74
IV. Comparative Anatomy of Papillomavirus Genomes ......... 75 A. Features of the HPV-la Genome.... ... . .... . ... . .... .. 75 B. Features of the HPV-6b Genome.... .... .... . .... ...... 77 C. Features of the CRPV Genome ........................ 79 D. Features of Other Papillomavirus Genomes.... ..... .... 81
V. Predicted Functions of Papillomavirus Proteins. . . . . . . . . . . . . 82 VI. Transcriptional Organization of Papillomavirus Genomes . . . 84
A. RNAs Expressed in Rodent Cells Transformed by BPV-l 85 B. mRNAs Transcribed in Virus-Producing BPV-I-Induced
Fibropapillomas ...................................... 93 C. The Transcription Map of the BPV -1 Genome Is
Incomplete. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 95 D. Transcriptional Organization of the CRPV Genome. . ... 97
CONTENTS xi
E. Common Features between BPV-1 and CRPV Transcription ........................................ 100
F. Transcriptional Organization of Human Papillomavirus Genomes ............................................ 102
VII. Conclusions............................................. 102 References .................................................. " 103
Chapter 4
The Expression of Papillomaviruses in Epithelial Cells
Lorne B. Taichman and Robert F. LaPorta
I. Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 109 II. Epithelium and Keratinization ............................ 110
III. Behavioral Classification of PVs . . . . . . . . . . . . . . . . . . . . . . . . . .. 116 IV. Histological Characteristics of PV-Induced Lesions ......... 117 V. Transformation.......................................... 121
VI. Vegetative Viral Growth. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 123 VII. Cytopathic Effects. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 127
VIII. Expression of PV in Cultured Keratinocytes . . . . . . . . . . . . . . .. 128 IX. Expression of PV in Malignant Cells. . . . . . . . . . . . . . . . . . . . . .. 131 X. Conclusions ............................................. 133
References. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 134
Chapter 5
Papilloma virus Transformation
Peter M. Howley and Richard Schlegel
I. Introduction............................................. 141 II. Transformation Biology. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 143
III. Transcription............................................ 147 IV. Transforming Functions. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 150
A. Genetics............................................. 150 B. E6 Gene .......................... ,. . .. . . . . . . .... . . .. 153 C. E5 Gene.. . . . . . . . . . .. . . . . .. . .... . . . . .. . .. . . . . . . . ... .. 154 D. E2 Gene. . . . . . . . . . . .. . . . ... . . ... . . . . .. . ... . . . . . . . .. .. 156
V. Plasmid Maintenance Functions. . . . . . . . . . . . . . . . . . . . . . . . . .. 158 A. Cis Elements. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 159 B. Trans Functions. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 160
VI. Summary and Conclusions. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 160 References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 163
xii CONTENTS
Chapter 6
Papillomaviruses and Carcinogenic Progression I: Cottontail Rabbit (Shope) Papillomavirus
Felix O. Wettstein
I. Introduction............................................. 167 II. Biological Properties of the System. . . . . . . . . . . . . . . . . . . . . . .. 168
A. Virus Multiplication and Tumor Induction . . . . . . . . . . . .. 168 B. Tumor Progression ................................... 170 C. Interaction with Chemical Carcinogens . . . . . . . . . . . . . . .. 171 D. Regression . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 172
III. Molecular Aspects of the System. . . . . . . . . . . . . . . . . . . . . . . . .. 173 A. The Virus and Viral Genome . . . . . . . . . . . . . . . . . . . . . . . . .. 173 B. State of the Viral DNA ............................... 176 C. Viral Transcripts ..................................... 179 D. Conclusion .......................................... 180
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 183
Chapter 7
Papillomaviruses and Carcinogenic Progression II: The Mastomys natalensis Papillomavirus
Eberhard Amtmann and Klaus Wayss
I. The Animal System...... ...... ......... ..... ..... ..... .. 187 II. Epithelial Skin Tumors in Mastomys .............. '" . .... 188
III. Identification of MnPV as the Causative Agent of Skin Tumors ................................................. 189
IV. Characterization of the Virus ............................. 190 V. Persistence of MnPV Genomes in Normal Tissu~s... ....... 191
VI. Accumulation of Viral DNA in Normal Skin during Aging.. 192 VII. MnPV Genome Expression in Tumors.. ... .. ... ..... . ..... 194
VIII. Induction of MnPV by Tumor Promoters ................. " 195 IX. Induction of Antibodies against MnPV in Mastomys ........ 197
References .................................................... 198
Chapter 8
Epidermodysplasia Verruciformis
Gerard Orth
I. Introduction............................................. 199 II. Clinical Aspects of EV . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 201
CONTENTS xiii
A. Clinical Course of the Disease. . . . . . . . . . . . . . . . . . . . . . . .. 201 B. Morphology of EV Lesions ............................ 202
III. Histology and Ultrastructure of EV Lesions ................ 204 A. Benign Lesions . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 204 B. Malignant Lesions . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 206
IV. Genetic Factors in EV ....... . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 207 V. Immunological Factors in EV ............................. 209
A. Humoral Immunity ................................... 210 B. Cell-Mediated Immunity. . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 211 C. Regression of EV ..................................... 212
VI. HPV Types Associated with EV ........................... 212 A. Multiplicity and Specificity. . . . . . . . . . . . . . . . . . . . . . . . . .. 212 B. Biochemical Properties. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 214 C. HPV Infection in EV Patients. . . . . . . . . . . . . . . . . . . . . . . . .. 223
VII. Role of HPVs in EV Carcinomas. . . . . . . . . . . . . . . . . . . . . . . . . .. 224 A. HPV Types Found in EV Cancers . . . . . . . . . . . . . . . . . . . . .. 224 B. Physical State of HPV Genomes . . . . . . . . . . . . . . . . . . . . . .. 225 C. Expression of HPV Genomes ..... . . . . . . . . . . . . . . . . . . . .. 227
VIII. HPVs and Skin Neoplasia in Non-EV Patients. . . . . . . . . . . . .. 227 A. Immunodepression or Immunosuppression . . . . . . . . . . . .. 227 B. General Population . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 229
IX. In Vitro Studies on EV HPVs . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 231 X. Questions Raised by EV .................................. 231
A. Plurality of HPVs. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 231 B. Control of HPV Infection ............................. 232 C. Cofactors in HPV Oncogenesis ........................ 233 D. Mechanisms of Tumor Progression .................... 234
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 235
Chapter 9
The Role of Papillomaviruses in Human Anogenital Cancer
Harald zur Hausen and Achim Schneider
I. Introduction and Historical Aspects ....................... 245 II. Isolation of Genital Papillomaviruses and Their
Characterization ......................................... 248 III. Biology of Genital Papillomavirus Infections ............... 250 IV. HPV DNA Sequences in Cell Lines ........................ 255 V. State of Viral DNA in Benign and Malignant Lesions ....... 256
VI. Interaction of Papillomavirus Infections with Initiators in Malignant Conversion . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 257
VII. Association of Malignant Tumors with Genital Papillomavirus Infections at Extragenital Sites ............. 258
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 259
xiv CONTENTS
Chapter 10
Laryngeal Papillomas: Clinical Aspects and in Vitro Studies
Bettie M. Steinberg
I. Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 265 II. Clinical Description. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 266
III. Treatment ............................................... 269 A. Physical Treatment. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 270 B. Surgical Treatment. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 270 C. Medical Treatment . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 270 D. Immunotherapy ...................................... 271
IV. Pathology................................................ 272 V. Molecular and Cellular Biology. . . . . . . . . . . . . . . . . . . . . . . . . . .. 275
A. Etiology ............................................. 275 B. Tissue Culture Studies. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 283
VI. Laryngeal Papillomas and Laryngeal Cancer . . . . . . . . . . . . . . .. 286 VII. Major Unanswered Questions. " ........... , .............. 287 References .................................................... 289
Chapter 11
Papillomavirus Cloning Vectors
Daniel DiMaio
I. Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 293 II. Transformation by Bovine Papillomavirus . . . . . . . . . . . . . . . . .. 294
III. Development of BPV-1 Vectors ............................ 297 IV. Expression of Genes Cloned on BPV-1 ..................... 301
A. Human I3-Globin Gene .............................. " 302 B. Metallothioneins . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 303 C. Hepatitis B Surface Antigen . . . . . . . . . . . . . . . . . . . . . . . . . .. 304 D. Human I3-Interferon Gene ........................... " 305 E. Mouse Mammary Tumor Virus. . . . . . . . . . . . . . . . . . . . . . .. 308 F. Polypeptide Hormones ................................ 309
G. Other Genes ......................................... 309 V. Stability of BPV-1 Recombinants ........................ " 310
VI. FutureDirections ........................................ 312 References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 315
Appendix
Sequence Analysis of Papillomavirus Genomes
Carl C. Baker
I. Papillomavirus Sequences. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 321 II. Restriction Site Analysis. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 352
CONTENTS xv
III. Genome Organization . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 366 IV. Protein Comparisons. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 367
A. E1 ORF . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 371 B. E20RF .............................................. 371 C. E40RF .............................................. 372 D. ES ORF . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 377 E. E60RF .............................................. 377 F. E70RF.............................................. 377
G. Ll ORF .............................................. 379 H. L20RF .............................................. 379
References .................................................... 384
Index ......................................................... 387