Comprehensive Urology 5 Comprehensive Urology Edited by Heinz Fraenkel-Conrat University of California at Berkeley and Robert R. Wagner University of Virginia Editorial Board Purnell W. Choppin, Rockefeller University Harrison Echols, University of California at Berkeley Harold S. Ginsberg, Columbia University Leon Hirth, University of Strasbourg Peter H. Hofschneider, Max Planck Institut fiir Biochemie E. M. J. Jaspars, University of Leiden R. E. F. Matthews, University of Auckland Yoshimi Okada, University of Tokyo Lennart Philipson, University of Uppsala Aaron J. Shatkin, Roche Institute of Molecular Biology Robert L. Sinsheimer, California Institute of Technology Peter K. Vogt, University of Southern California Robley C. Williams, University of California at Berkeley William B. Wood, California Institute of Technology Volume 1: Descriptive Catalogue of Viruses - by Heinz Fraenkel-Conrat Reproduction Volume 2: Small and Intermediate RNA Viruses - Contributors: J.T. August, L. Eoyang, A. Siegel, V. Hariharasubramanian, L. Levintow, E. R. Pfefferkorn, D. Shapiro, and W. K. Joklik Volume 3: DNA Animal Viruses - Contributors: L. Philipson, U. lindberg, J.A. Rose, N.P. Salzman, G. Khoury, B. Moss, B. Roizman, and D. Furlong Volume 4: Large RNA Viruses - Contributors: P.W. Choppin, RW. Compans, R.R. Wagner, and J.P. Bader In preparation: DNA Bacteriophages - Contributors: D. T. Denhardt, C.K. Mathews, and RS. Ray Structure and Assembly Volume 5: Virions, Pseudo virions, and Intraviral Nucleic Acids - Contributors: T.I. Tikchonenko, John T. Finch, lionel V. Crawford, and H. Vasken Aposhian Volume 6: Assembly of Small RNA Viruses - Contributors: L. Hirth, P.P. Hung, T. Ohno, Y. Okada, K.E. Richards, RR. Rueckert, and R.C. Williams Other W. Bauer, D.L.D. Caspar, P.W. Choppin, RW. Compans, F.A. Eiserling, contributors W. Fiers, RM. Franklin, H.S. Ginsberg, K.A. Harrap, J. King, T.W. Tinsley, include: J. Vinograd, and W.B. Wood Regulation and Genetics Volume 7: Bacteriophages - Contributors: R. Calendar, E. P. Geiduschek, M.E. Gottesman, S. Gottesman, and R. Weisberg Other J. Atabekov, D. Baltimore, D. Bishop, M.A. Bratt, A. Campbell, P. Cooper, con tribu tors W. Doerfler,W. Eckhart, B. Fields, H. Ginsberg, H. Hanafusa, A. Huang, include: E.M.J. Jaspars, J. Kates, A. Lewis, D. Nathans, E.R. Pfefferkorn, C. Pringle, J.R. Roberts, H.L. Sanger, A.J. Shatkin, F. W. Stahl, J.H. Subak-Sharpe, W. Szybalski, L. Van Vloten-Doting, and P. Vogt Interaction of Viruses and Their Hosts Effects of Physical and Chemical Agents omprehensive Edited by Heinz Fraenkel-Conrat Department 0/ Molecular Biology and Virus Laboratory University o/Cali/omiD, Berkeley, Cali/omiD and Robert R.Wagner Department 0/M icrobiology University 0/ VirginiD, Charlottesville, Virginia irology 5 Structure and Assembly Virions, Pseudovirions, and Intraviral Nucleic Acids PLENUM PRESS. NEW YORK AND LONDON Library of Congress Cataloging in Publication Data Fraenkel-Conrat, Heinz, 1910- Comprehensive virology. Includes bibliographies. 1. Virology-Collected works. I. Wagner, Robert R., 1923- joint author. II. Title. [DNLM: 1. Virus diseases. 2. Viruses. QW160 F799cal QR357.F72 576'.64'08 74-5494 ISBN 978-1-4684-2711-0 ISBN 978-1-4684-2709-7 (eBook) DOl 10.1007/978-1-4684-2709-7 © 1975 Plenum Press, New York Softcover reprint of the hardcover 1st edition 1975 A Division of Plenum Publishing Corporation 227 West 17th Street, New York, N.Y. 10011 United Kingdom edition published by Plenum Press, London A Division of Plenum Publishing Company, Ltd. Davis House (4th Floor), 8 Scrubs Lane, Harlesden, London, NWI0 6SE, England 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 Foreword The time seems ripe for a critical compendium of that segment of the biological universe we call viruses. Virology, as a science, having passed only recently through its descriptive phase of naming and num hering, has probably reached that stage at which relatively few new-truly new-viruses will be discovered. Triggered by the in tellectual probes and techniques of molecular biology, genetics, biochemical cytology, and high-resolution microscopy and spectroscopy, the field has experienced a genuine information explo sion. Few serious attempts have been made to chronicle these events. This comprehensive series, which will comprise some 6000 pages in a total of about 22 volumes, represents a commitment by a large group of active investigators to analyze, digest, and expostulate on the great mass of data relating to viruses, much of which is now amorphous and disjointed, and scattered throughout a wide literature. In this way, we hope to place the entire field in perspective, and to develop an in valuable reference a·nd sourcebook for researchers and students at all levels. This series is designed as a continuum that can be entered anywhere, but which also provides a logical progression of developing facts and integrated concepts. Volume 1 contains an alphabetical catalogue of almost all viruses of vertebrates, insects, plants, and protists, describing them in general terms. Volumes 2-4 deal primarily, but not exclusively, with the processes of infection and reproduction of the major groups of viruses in their hosts. Volume 2 deals with the simple RNA viruses of bacteria, plants, and animals; the togaviruses (formerly called arboviruses), which share with these only the feature that the virion's RNA is able to act as messenger RNA in the host cell; and the reoviruses of animals and plants, which all share several structurally singular features, the most important being the double-strandedness of their multiple RNA vii viii Foreword molecules. This grouping, of course, has only slightly more in its favor than others that could have been, or indeed were, considered. Volume 3 addresses itself to the reproduction of all DNA-contain ing viruses of vertebrates, a seemingly simple act of classification, even though the field encompasses the smallest and the largest viruses known. (The reproduction of the DNA-bacteriophages will be dealt with later, probably in Volume 8.) The reproduction of the larger and more complex RNA viruses is the subject matter of Volume 4. These viruses share the property of lipid-rich envelopes with the togaviruses included in Volume 2. They share as a group, and with the reoviruses, the presence of enzymes in their virions and the need for their RNA to become transcribed before it can serve messenger functions. Volumes 5 and 6 represent the first in a series that focuses primarily on the structure and assembly of virus particles. Other volumes nearing completion will deal with animal virus genetics, multicomponent plant viruses, regulation of viral replication, virus-host relationships, etc. The completed series will endeavor to encompass all aspects of the molecular biology and the behavior of viruses. We hope to keep this series up to date at all times by prompt and rapid publication of all contributions, and by encouraging the authors to update their chapters by additions or corrections whenever a volume is reprinted. Contents Chapter 1 Structure of Viral Nucleic Acids in Situ T. I. Tikchonenko 1. Viruses with Single-Stranded Nucleic Acids. . . . . . . . . . . 2 1.1. Rodlike and Filamentous RNA-Containing Viruses. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 1.2. Filamentous DNA-Containing Phages .......... 25 1.3. Isometric RNA-Containing Viruses. . . . . . . . . . . . . 30 1.4. Isometric DNA-Containing Viruses. . . . . . . . . . . . . 53 2. Viruses with Double-Stranded DNA. . . . . . . . . . . . . . . . . 54 2.1. The Tertiary Structure of DNA. . . . . . . . . . . . . . . . 54 2.2. The Secondary Structure of DNA. . . . . . . . . . . . . . 66 3. Concluding Remarks . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 98 . 4. References....................................... 99 Chapter 2 Structure of Small DNA-Containing Animal Viruses John T. Finch and Lionel V. Crawford 1. General Principles of Virus Structure ................ 119 2. Papilloma Viruses ................................ 120 2.1. Biology of the Viruses ........................ 120 2.2. Composition................................ 121 2.3. Structure................................... 123 3. SV40 and Polyoma Virus .......................... 139 3.1. Biology of the Viruses ........................ 139 3.2. Composition................................ 140 ix x Contents 3.3. Structure................................... 145 3.4. Related Human Viruses ...................... 149 4. References....................................... 151 Chapter 3 Pseudovirions in Animals, Plants, and Bacteria H. Vas ken Aposhian 1. Introduction..................................... 155 1.1. Definitions.................................. 155 1.2. Scope of the Chapter. . . . . . . . . . . . . . . . . . . . . . . .. 156 2. Animal Cell Pseudovirions ......................... 157 2.1. Polyoma Pseudovirions . . . . . . . . . . . . . . . . . . . . . .. 157 2.2. SV40 Pseudovirions . . . . . . . . . . . . . . . . . . . . . . . . .. 186 2.3. Other Animal Virus Systems Studied for Pseudovirions ............................. 196 3. Plant Cell Pseudovirions (Tobacco Mosaic Pseudovirions) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 197 3.1. Separation.................................. 197 3.2. Characterization............................. 198 4. Bacterial Cell Pseudovirions(Generalized Transducing Phages) ....................................... 199 5. Orphan Pseudo virions ............................. 201 5.1. From Bacillus subtilis ........................ 201 5.2. From Bacillus licheniformis ................... 210 6. Possible Biological Roles of Pseudovirions . . . . . . . . . . .. 211 7. Summary........................................ 212 8. References....................................... 213 Index ..... .................................... "" ...... , 219