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B W J Mahy M H V van Regenmortel Encyclopedia of
Virology Five Volume Set Volume 1 5 Third Edition 2008
B.W.J. Mahy Digital Instant Download
Author(s): B.W.J. Mahy, M. H.V. van Regenmortel - Encyclopedia of
Virology, Five-Volume Set, Volume 1-5, Third Edition (2008)
ISBN(s): 9781865843834, 0123739357
Edition: 3
File Details: PDF, 80.42 MB
Year: 2008
Language: english
Encyclopedia of
VIROLOGY
THIRD EDITION
Encyclopedia of
VIROLOGY
THIRD EDITION
EDITORS-IN-CHIEF

Dr BRIAN W J MAHY
and
Dr MARC H V VAN REGENMORTEL
Academic Press is an imprint of Elsevier
Linacre House, Jordan Hill, Oxford, OX2 8DP, UK
525 B Street, Suite 1900, San Diego, CA 92101-4495, USA

Copyright ã 2008 Elsevier Inc. All rights reserved

The following articles are US government works in the public domain and are not subject to copyright:
Bovine Viral Diarrhea Virus, Coxsackieviruses, Prions of Yeast and Fungi, Human Respiratory Syncytial Virus,
Fish Rhabdoviruses, Varicella-Zoster Virus: General Features, Viruses and Bioterrorism, Bean Common Mosaic
Virus and Bean Common Mosaic Necrosis Virus, Metaviruses, Crimean-Congo Hemorrhagic Fever Virus and Other
Nairoviruses, AIDS: Global Epidemiology, Papaya Ringspot Virus, Transcriptional Regulation in Bacteriophage.

Nepovirus, Canadian Crown Copyright 2008

No part of this publication may be reproduced, stored in a retrieval system or transmitted in any form or by
any means electronic, mechanical, photocopying, recording or otherwise without the prior written
permission of the publisher

Permissions may be sought directly from Elsevier’s Science & Technology Rights Department in Oxford,
UK: phone (+44) (0) 1865 843830; fax (+44) (0) 1865 853333; email: [email protected]. Alternatively
you can submit your request online by visiting the Elsevier web site at (http://elsevier.com/locate/permission),
and selecting Obtaining permission to use Elsevier material

Notice
No responsibility is assumed by the publisher for any injury and/or damage to persons or property as a
matter of products liability, negligence or otherwise, or from any use or operation of any methods, products,
instructions or ideas contained in the material herein. Because of rapid advances in the medical sciences,
in particular, independent verification of diagnoses and drug dosages should be made

British Library Cataloguing in Publication Data

A catalogue record for this book is available from the British Library

Library of Congress Catalog Number: 200892260

ISBN: 978-0-12-373935-3

For information on all Elsevier publications

visit our website at books.elsevier.com

PRINTED AND BOUND IN SLOVENIA

08 09 10 11 10 9 8 7 6 5 4 3 2 1
EDITORS-IN-CHIEF

Brian W J Mahy MA PhD ScD DSc

Senior Scientific Advisor,
Division of Emerging Infections and Surveillance Services,
Centers for Disease Control and Prevention,
Atlanta GA, USA

Marc H V Van Regenmortel PhD

Emeritus Director at the CNRS,
French National Center for Scientific Research,
Biotechnology School of the University of Strasbourg,
Illkirch, France

v
ASSOCIATE EDITORS

Dennis H Bamford, Ph.D. Robert A. Lamb, Ph.D., Sc.D.

Department of Biological and Environmental Sciences Department of Biochemistry,
and Institute of Biotechnology, Biocenter 2, Molecular Biology and Cell Biology
P.O. Box 56 (Viikinkaari 5), Howard Hughes Medical Institute
00014 University of Helsinki, Northwestern University
Finland 2205 Tech Dr.
Evanston
Charles Calisher, B.S., M.S., Ph.D.
IL 60208-3500
Arthropod-borne and Infectious Diseases Laboratory
USA
Department of Microbiology, Immunology and Pathology
College of Veterinary Medicine and Biomedical Sciences Olivier Le Gall
Colorado State University IPV, UMR GDPP, IBVM,
Fort Collins INRA Bordeaux-Aquitaine, BP 81,
CO 80523 F-33883 Villenave d’Ornon Cedex
USA FRANCE
Andrew J Davison, M.A., Ph.D. Vincent Racaniello, Ph.D.
MRC Virology Unit Department of Microbiology
Institute of Virology Columbia University
University of Glasgow New York, NY 10032
Church Street USA
Glasgow G11 5JR
UK David A. Theilmann, Ph.D., B.Sc., M.Sc
Pacific Agri-Food Research Centre
Claude Fauquet
Agriculture and Agri-Food Canada
ILTAB/Donald Danforth Plant Science Center
Box 5000, 4200 Highway 97
975 North Warson Road
Summerland
St. Louis, MO 63132
BC V0H 1Z0
Said Ghabrial, B.S., M.S., Ph.D. Canada
Plant Pathology Department
University of Kentucky H. Josef Vetten, Ph.D.
201F Plant Science Building Julius Kuehn Institute, Federal Research Centre for
1405 Veterans Drive Cultivated Plants (JKI)
Lexington Messeweg 11-12
KY 4050546-0312 38104 Braunschweig
USA Germany

Eric Hunter, B.Sc., Ph.D. Peter J Walker, B.Sc., Ph.D.

Department of Pathology and Laboratory Medicine, and CSIRO Livestock Industries
Emory Vaccine Center Australian Animal Health Laboratory (AAHL)
Emory University Private Bag 24
954 Gatewood Road NE Geelong
Atlanta Georgia 30329 VIC 3220
USA Australia

vii
PREFACE

This third edition of the Encyclopedia of Virology is being published nine years after the second edition, a period which has
seen enormous growth both in our understanding of virology and in our recognition of the viruses themselves, many of
which were unknown when the second edition was prepared. Considering viruses affecting human hosts alone, the
worldwide epidemic of severe acute respiratory syndrome (SARS), caused by a previously unknown coronavirus, led to
the discovery of other human coronaviruses such as HKU1 and NL63. As many as seven chapters are devoted to the
AIDS epidemic and to human immunodeficiency viruses. In addition, the development of new molecular technologies
led to the discovery of viruses with no obvious disease associations, such as torque-teno virus (one of the most ubiquitous
viruses in the human population), human bocavirus, human metapneumovirus, and three new human polyomaviruses.
Other new developments of importance to human virology have included the introduction of a virulent strain of West
Nile virus from Israel to North America in 1999. Since that time the virus has become established in mosquito, bird and
horse populations throughout the USA, the Caribbean and Mexico as well as the southern regions of Canada.
As in the two previous editions, we have tried to include information about all known species of virus infecting
bacteria, fungi, invertebrates, plants and vertebrates, as well as descriptions of related topics in virology such as antiviral
drug development, cell- and antibody-mediated immunity, vaccine development, electron microscopy and molecular
methods for virus characterization and identification. Many chapters are devoted to the considerable economic
importance of virus diseases of cereals, legumes, vegetable crops, fruit trees and ornamentals, and new approaches to
control these diseases are reviewed.
General issues such as the origin, evolution and phylogeny of viruses are also discussed as well as the history of the
different groups of viruses.

To cover all these subjects and new developments, we have had to increase the size of the Encyclopedia from three to five
volumes.
Throughout this work we have relied upon the 8th Report of the International Committee on Taxonomy of Viruses
published in 2005, which lists more than 6000 viruses classified into some 2000 virus species distributed among more
than 390 different genera and families. In recent years the criteria for placing viruses in different taxa have shifted away
from traditional serological methods and increasingly rely upon molecular techniques, particularly the nucleotide
sequence of the virus genome. This has changed many of the previous groupings of viruses, and is reflected in this
third edition.
Needless to say, a work of this magnitude has involved many expert scientists, who have given generously of their time
to bring it to fruition. We extend our grateful thanks to all contributors and associate editors for their excellent and
timely contributions.

Brian W J Mahy
Marc H V van Regenmortel

ix
HOW TO USE THE ENCYCLOPEDIA

Structure of the Encyclopedia

The major topics discussed in detail in the text are presented in alphabetical order (see the Alphabetical Contents list
which appears in all five volumes).

Finding Specific Information

Information on specific viruses, virus diseases and other matters can be located by consulting the General Index at the
end of Volume 5.

Taxonomic Groups of Viruses

For locating detailed information on the major taxonomic groups of viruses, namely virus genera, families and orders, the
Taxonomic Index in Volume 5 (page. . .) should be consulted.

Cross referencing between articles

The ‘‘See also’’ section at the end of each article directs the reader to other entries on related topics. For example. The
entry Lassa, Junin, Machupo and Guanarito Viruses includes the following cross-references:

D Gallitelli M-A Grandbastien

Università degli Studi and Istituto di Virologia Vegetale del INRA, Versailles, France
CNR, Bari, Italy
R Grassmann
F Garcı́a-Arenal University of Erlangen – Nürnberg, Erlangen, Germany
Universidad Politécnica de Madrid, Madrid, Spain
M Gravell
J A Garcı́a National Institutes of Health, Bethesda, MD, USA
Centro Nacional de Biotecnologı́a (CNB), CSIC, Madrid, M V Graves
Spain University of Massachusetts–Lowell, Lowell, MA, USA
R A Garrett K Y Green
Copenhagen University, Copenhagen, Denmark National Institutes of Health, Bethesda, MD, USA
S Gaumer H B Greenberg
Université Versailles St-Quentin, CNRS, Versailles, Stanford University School of Medicine and
France Veterans Affairs Palo Alto Health Care System, Palo Alto,
CA, USA
R J Geijskes
Queensland University of Technology, Brisbane, QLD, B M Greenberg
Australia Johns Hopkins School of Medicine, Baltimore, MD, USA
T W Geisbert I Greiser-Wilke
National Emerging Infectious Diseases Laboratories, School of Veterinary Medicine, Hanover, Germany
Boston, MA, USA
D E Griffin
E Gellermann Johns Hopkins Bloomberg School of Public Health,
Hannover Medical School, Hannover, Germany Baltimore, MD, USA
A Gessain T S Gritsun
Pasteur Institute, CNRS URA 3015, Paris, France University of Reading, Reading, UK
S A Ghabrial R J de Groot
University of Kentucky, Lexington, KY, USA Utrecht University, Utrecht, The Netherlands
Contributors xix

A J Gubala S Hertzler
CSIRO Livestock Industries, Geelong, VIC, Australia University of Illinois at Chicago, Chicago, IL, USA
D J Gubler F van Heuverswyn
John A. Burns School of Medicine, Honolulu, HI, USA University of Montpellier 1, Montpellier, France
A-L Haenni J Hilliard
Institut Jacques Monod, Paris, France Georgia State University, Atlanta, GA, USA
D Haig B I Hillman
Nottingham University, Nottingham, UK Rutgers University, New Brunswick, NJ, USA
F J Haines S Hilton
Oxford Brookes University, Oxford, UK University of Warwick, Warwick, UK
J Hamacher D M Hinton
INRES, University of Bonn, Bonn, Germany National Institutes of Health, Bethesda, MD, USA
J Hammond A Hinz
USDA-ARS, Beltsville, MD, USA UMR 5233 UJF-EMBL-CNRS, Grenoble, France
R M Harding A E Hoet
Queensland University of Technology, Brisbane, QLD, The Ohio State University, Columbus, OH, USA
Australia
S A Hogenhout
J M Hardwick The John Innes Centre, Norwich, UK
Johns Hopkins University Schools of Public Health and
T Hohn
Medicine, Baltimore, MD, USA
Basel university, Institute of Botany, Basel,
D Hariri Switzerland
INRA – Département Santé des Plantes et
J S Hong
Environnement, Versailles, France
Seoul Women’s University, Seoul, South Korea
B Harrach
M C Horzinek
Veterinary Medical Research Institute, Budapest,
Utrecht University, Utrecht, The Netherlands
Hungary
T Hovi
P A Harries
National Public Health Institute (KTL), Helsinki, Finland
Samuel Roberts Noble Foundation, Inc., Ardmore, OK,
USA A M Huger
Institute for Biological Control, Darmstadt, Germany
L E Harrington
University of Alabama at Birmingham, Birmingham, AL, L E Hughes
USA University of St. Andrews, St. Andrews, UK
T J Harrison R Hull
University College London, London, UK John Innes Centre, Colney, UK
T Hatziioannou E Hunter
Aaron Diamond AIDS Research Center, The Rockefeller Emory University Vaccine Center, Atlanta, GA, USA
University, New York, NY, USA
A D Hyatt
J Hay Australian Animal Health Laboratory, Geelong, VIC,
The State University of New York, Buffalo, NY, USA Australia
G S Hayward T Hyypiä
Johns Hopkins School of Medicine, Baltimore, MD, USA University of Turku, Turku, Finland
E Hébrard T Iwanami
IRD, Montpellier, France National Institute of Fruit Tree Science, Tsukuba, Japan
R W Hendrix A O Jackson
University of Pittsburgh, Pittsburgh, PA, USA University of California, Berkeley, CA, USA
L E Hensley P Jardine
USAMRIID, Fort Detrick, MD, USA University of Minnesota, Minneapolis, MN, USA
M de las Heras J A Jehle
University of Glasgow Veterinary School, Glasgow, UK DLR Rheinpfalz, Neustadt, Germany
xx Contributors

A R Jilbert R G Kleespies
Institute of Medical and Veterinary Science, Adelaide, SA, Institute for Biological Control, Darmstadt, Germany
Australia
D F Klessig
P John Boyce Thompson Institute for Plant Research, Ithaca, NY,
Indian Agricultural Research Institute, New Delhi, India USA
J E Johnson W B Klimstra
The Scripps Research Institute, La Jolla, CA, USA Louisiana State University Health Sciences Center at
R T Johnson Shreveport, Shreveport, LA, USA
Johns Hopkins School of Medicine, Baltimore, MD, USA V Klimyuk
W E Johnson Icon Genetics GmbH, Weinbergweg, Germany
New England Primate Research Center, Southborough, N Knowles
MA, USA Institute for Animal Health, Pirbright, UK
S L Johnston
R Koenig
Imperial College London, London, UK
Biologische Bundesanstalt für Land- und Forstwirtschaft,
A T Jones Brunswick, Germany
Scottish Crop Research Institute, Dundee, UK
R Koenig
R Jordan Institut für Pflanzenvirologie, Mikrobiologie und
USDA-ARS, Beltsville, MD, USA biologische Sicherheit, Brunswick, Germany
Y Kapustin G Konaté
National Institutes of Health, Bethesda, MD, USA INERA, Ouagadougou, Burkina Faso
P Karayiannis C N Kotton
Imperial College London, London, UK Massachusetts General Hospital, Boston, MA, USA
P Kazmierczak L D Kramer
University of California, Davis, CA, USA Wadsworth Center, New York State Department of
K M Keene Health, Albany, NY, USA
Colorado State University, Fort Collins, CO, USA P J Krell
C Kerlan University of Guelph, Guelph, ON, Canada
Institut National de la Recherche Agronomique (INRA), Le
J Kreuze
Rheu, France
International Potato Center (CIP), Lima, Peru
K Khalili
M J Kuehnert
Temple University School of Medicine, Philadelphia, PA,
Centers for Disease Control and Prevention, Atlanta, GA,
USA
USA
P H Kilmarx
Centers for Disease Control and Prevention, Atlanta, GA, R J Kuhn
USA Purdue University, West Lafayette, IN, USA

L A King G Kurath
Oxford Brookes University, Oxford, UK Western Fisheries Research Center, Seattle, WA, USA

P D Kirkland I Kusters
Elizabeth Macarthur Agricultural Institute, Menangle, sanofi pasteur, Lyon, France
NSW, Australia I V Kuzmin
C D Kirkwood Centers for Disease Control and Prevention, Atlanta,
Murdoch Childrens Research Institute Royal Children’s GA, USA
Hospital, Melbourne, VIC, Australia M E Laird
R P Kitching New England Primate Research Center, Southborough,
Canadian Food Inspection Agency, Winnipeg, MB, MA, USA
Canada
R A Lamb
P J Klasse Howard Hughes Medical Institute at Northwestern
Cornell University, New York, NY, USA University, Evanston, IL, USA
N R Klatt P F Lambert
University of Pennsylvania School of Medicine, University of Wisconsin School of Medicine
Philadelphia, PA, USA and Public Health, Madison, WI, USA
Contributors xxi

A S Lang M L Linial
Memorial University of Newfoundland, St. John’s, NL, Fred Hutchinson Cancer Research Center, Seattle, WA,
Canada USA
H D Lapierre D C Liotta
INRA – Département Santé des Plantes et Emory University, Atlanta, GA, USA
Environnement, Versailles, France
W Ian Lipkin
G Lawrence Columbia University, New York, NY, USA
The Children’s Hospital at Westmead, Westmead, NSW,
Australia and H L Lipton
University of Sydney, Westmead, NSW, Australia University of Illinois at Chicago, Chicago, IL, USA

H Lecoq A S Liss
Institut National de la Recherche Agronomique (INRA), University of Texas at Austin, Austin, TX, USA
Station de Pathologie Végétale, Montfavet, France
J J López-Moya
B Y Lee Instituto de Biologı́a Molecular de Barcelona (IBMB),
Seoul Women’s University, Seoul, South Korea CSIC, Barcelona, Spain
E J Lefkowitz G Loebenstein
University of Alabama at Birmingham, Birmingham, AL, Agricultural Research Organization, Bet Dagan, Israel
USA
C-F Lo
J P Legg National Taiwan University, Taipei, Republic of China
International Institute of Tropical Agriculture, Dar es
Salaam, Tanzania, S A Lommel
UK and North Carolina State University, Raleigh, NC, USA
Natural Resources Institute, Chatham Maritime, UK
G P Lomonossoff
P Leinikki John Innes Centre, Norwich, UK
National Public Health Institute, Helsinki, Finland
M Luo
J Lenard University of Alabama at Birmingham, Birmingham, AL,
University of Medicine and Dentistry of New Jersey USA
(UMDNJ), Piscataway, NJ, USA
S A MacFarlane
J C Leong Scottish Crop Research Institute, Dundee, UK
University of Hawaii at Manoa, Honolulu, HI, USA
J S Mackenzie
K N Leppard Curtin University of Technology, Shenton Park, WA,
University of Warwick, Coventry, UK Australia
A Lescoute R Mahieux
Université Louis Pasteur, Strasbourg, France Pasteur Institute, CNRS URA 3015, Paris, France
D-E Lesemann B W J Mahy
Biologische Bundesanstalt für Land- und Forstwirtschaft, Centers for Disease Control and Prevention, Atlanta, GA,
Brunswick, Germany USA
J-H Leu E Maiss
National Taiwan University, Taipei, Republic of China Institute of Plant Diseases and Plant Protection,
H L Levin Hannover, Germany
National Institutes of Health, Bethesda, MD, USA
E O Major
D J Lewandowski National Institutes of Health, Bethesda, MD, USA
The Ohio State University, Columbus, OH, USA
V G Malathi
H-S Lim Indian Agricultural Research Institute, New Delhi,
University of California, Berkeley, Berkeley, CA, USA India
M D A Lindsay A Mankertz
Western Australian Department of Health, Mount Robert Koch-Institut, Berlin, Germany
Claremont, WA, Australia
S Mansoor
R Ling National Institute for Biotechnology and Genetic
University of Warwick, Coventry, UK Engineering, Faisalabad, Pakistan
Contributors xxiii

L Moser M S Oberste
University of Wisconsin – Madison, Madison, WI, USA Centers for Disease Control and Prevention, Atlanta, GA,
USA
B Moury
INRA – Station de Pathologie Végétale, Montfavet, France W A O’Brien
University of Texas Medical Branch – Galveston,
J W Moyer Galveston, TX, USA
North Carolina State University, Raleigh, NC, USA
D J O’Callaghan
R W Moyer Louisiana State University Health Sciences Center,
University of Florida, Gainesville, FL, USA Shreveport, LA, USA
E Muller W F Ochoa
CIRAD/UMR BGPI, Montpellier, France University of California, San Diego, La Jolla, CA, USA
F A Murphy M R Odom
University of Texas Medical Branch, Galveston, University of Alabama at Birmingham, Birmingham, AL,
TX, USA USA
A Müllbacher M M van Oers
Australian National University, Canberra, ACT, Australia Wageningen University, Wageningen, The Netherlands
K Nagasaki M B A Oldstone
Fisheries Research Agency, Hiroshima, Japan The Scripps Research Institute, La Jolla, CA, USA
T Nakayashiki G Olinger
National Institutes of Health, Bethesda, MD, USA USAMRIID, Fort Detrick, MD, USA
A A Nash K E Olson
University of Edinburgh, Edinburgh, UK Colorado State University, Fort Collins, CO, USA
N Nathanson A Olspert
University of Pennsylvania, Philadelphia, PA, USA Tallinn University of Technology, Tallinn, Estonia
C K Navaratnarajah G Orth
Purdue University, West Lafayette, IN, USA Institut Pasteur, Paris, France
M S Nawaz-ul-Rehman J E Osorio
Danforth Plant Science Center, St. Louis, MO, USA University of Wisconsin, Madison, WI, USA
J C Neil N Osterrieder
University of Glasgow, Glasgow, UK Cornell University, Ithaca, NY, USA
R S Nelson S A Overman
Samuel Roberts Noble Foundation, Inc., Ardmore, University of Missouri – Kansas City, Kansas City,
OK, USA MO, USA
P Nettleton R A Owens
Moredun Research Institute, Edinburgh, UK Beltsville Agricultural Research Center, Beltsville, MD,
USA
A W Neuman
Emory University, Atlanta, GA, USA M S Padmanabhan
Yale University, New Haven, CT, USA
A R Neurath
Virotech, New York, NY, USA S Paessler
University of Texas Medical Branch, Galveston, TX, USA
M L Nibert
Harvard Medical School, Boston, MA, USA P Palese
Mount Sinai School of Medicine, New York, NY, USA
L Nicoletti
Istituto Superiore di Sanità, Rome, Italy M A Pallansch
Centers for Disease Control and Prevention, Atlanta, GA,
N Noah
USA
London School of Hygiene and Tropical Medicine,
London, UK M Palmarini
University of Glasgow Veterinary School, Glasgow, UK
D L Nuss
University of Maryland Biotechnology Institute, Rockville, P Palukaitis
MD, USA Scottish Crop Research Institute, Invergowrie, Dundee, UK
xxii Contributors

A A Marfin T C Mettenleiter
Centers for Disease Control and Prevention, Atlanta, Friedrich-Loeffler-Institut, Greifswald-Insel Riems, Germany
GA, USA
H Meyer
S Marillonnet Bundeswehr Institute of Microbiology, Munich, Germany
Icon Genetics GmbH, Weinbergweg, Germany
R F Meyer
G P Martelli Centers for Disease Control and Prevention, Atlanta, GA,
Università degli Studi and Istituto di Virologia vegetale USA
CNR, Bari, Italy
P de Micco
M Marthas Etablissement Français du Sang Alpes-Méditerranée,
University of California, Davis, Davis, CA, USA Marseilles, France
D P Martin B R Miller
University of Cape Town, Cape Town, South Africa Centers for Disease Control and Prevention (CDC), Fort
Collins, CO, USA
P A Marx
Tulane University, Covington, LA, USA C J Miller
University of California, Davis, Davis, CA, USA
W S Mason
Fox Chase Cancer Center, Philadelphia, PA, USA R G Milne
Istituto di Virologia Vegetale CNR, Torino, Italy
T D Mastro
Centers for Disease Control and Prevention, Atlanta, P D Minor
GA, USA NIBSC, Potters Bar, UK

A A McBride S Mjaaland
National Institutes of Health, Bethesda, MD, USA Norwegian School of Veterinary Science, Oslo, Norway

L McCann E S Mocarski
National Institutes of Health, Bethesda, MD, USA Emory University School of Medicine, Atlanta, GA, USA

M McChesney E S Mocarski, Jr.

University of California, Davis, Davis, CA, USA Emory University School of Medicine, Emory, GA, USA

J B McCormick V Moennig
University of Texas, School of Public Health, Brownsville, School of Veterinary Medicine, Hanover, Germany
TX, USA
P Moffett
G McFadden Boyce Thompson Institute for Plant Research, Ithaca, NY,
University of Florida, Gainesville, FL, USA USA

G McFadden T P Monath
The University of Western Ontario, London, ON, Kleiner Perkins Caufield and Byers, Menlo Park, CA, USA
Canada
R C Montelaro
D B McGavern University of Pittsburgh School of Medicine, Pittsburgh,
The Scripps Research Institute, La Jolla, CA, USA PA, USA

A L McNees P S Moore
Baylor College of Medicine, Houston, TX, USA University of Pittsburgh Cancer Institute, Pittsburgh,
PA, USA
M Meier
Tallinn University of Technology, Tallinn, Estonia F J Morales
International Center for Tropical Agriculture, Cali,
P S Mellor Colombia
Institute for Animal Health, Woking, UK
H Moriyama
X J Meng Tokyo University of Agriculture and Technology, Fuchu,
Virginia Polytechnic Institute and State University, Japan
Blacksburg, VA, USA
T J Morris
A A Mercer University of Nebraska, Lincoln, NE, USA
University of Otago, Dunedin, New Zealand
S A Morse
P P C Mertens Centers for Disease Control and Prevention, Atlanta, GA,
Institute for Animal Health, Woking, UK USA
xxiv Contributors

I Pandrea R T M Poulter
Tulane National Primate Research Center, Covington, LA, University of Otago, Dunedin, New Zealand
USA
A M Powers
O Papadopoulos Centers for Disease Control and Prevention, Fort Collins,
Aristotle University, Thessaloniki, Greece CO, USA
H R Pappu D Prangishvili
Washington State University, Pullman, WA, USA Institut Pasteur, Paris, France
S Parker C M Preston
Saint Louis University School of Medicine, St. Louis, MO, Medical Research Council Virology Unit, Glasgow, UK
USA
S L Quackenbush
C R Parrish Colorado State University, Fort Collins, CO, USA
Cornell University, Ithaca, NY, USA
F Qu
R F Pass University of Nebraska, Lincoln, NE, USA
University of Alabama School of Medicine, Birmingham,
AL, USA B C Ramirez
CNRS, Paris, France
J L Patterson
Southwest Foundation for Biomedical Research, San A Rapose
Antonio, TX, USA University of Texas Medical Branch – Galveston,
Galveston, TX, USA
T A Paul
Cornell University, Ithaca, NY, USA D V R Reddy
Hyderabad, India
A E Peaston
The Jackson Laboratory, Bar Harbor, ME, USA A J Redwood
The University of Western Australia,
M Peeters Crawley, WA, Australia
University of Montpellier 1, Montpellier, France
M Regner
J S M Peiris Australian National University, Canberra, ACT, Australia
The University of Hong Kong, Hong Kong, People’s
Republic of China W K Reisen
University of California, Davis, CA, USA
P J Peters
Centers for Disease Control and Prevention, Atlanta, GA, T Renault
USA IFREMER, La Tremblade, France

M Pfeffer P A Revill
Bundeswehr Institute of Microbiology, Munich, Germany Victorian Infectious Diseases Reference Laboratory,
Melbourne, VIC, Australia
H Pfister
University of Köln, Cologne, Germany A Rezaian
University of Adelaide, Adelaide, SA, Australia
O Planz
Federal Research Institute for Animal Health, Tuebingen, J F Ridpath
Gemany USDA, Ames, IA, USA

L L M Poon B K Rima
The University of Hong Kong, Hong Kong, People’s The Queen’s University of Belfast, Belfast, UK
Republic of China E Rimstad
M M Poranen Norwegian School of Veterinary Science, Oslo,
University of Helsinki, Helsinki, Finland Norway

K Porter F J Rixon
The University of Melbourne, Parkville, VIC, Australia MRC Virology Unit, Glasgow, UK

A Portner Y-T Ro
St. Jude Children’s Research Hospital, Memphis, TN, USA Konkuk University, Seoul, South Korea
R D Possee C M Robinson
NERC Institute of Virology and Environmental University of Oklahoma Health Sciences Center,
Microbiology, Oxford, UK Oklahoma City, OK, USA
Contributors xxv

G F Rohrmann J T Sample
Oregon State University, Corvallis, OR, USA The Pennsylvania State University College of Medicine,
Hershey, PA, USA
M Roivainen
National Public Health Institute (KTL), Helsinki, C E Sample
Finland The Pennsylvania State University College of Medicine,
Hershey, PA, USA
L Roux
University of Geneva Medical School, Geneva, R M Sandri-Goldin
Switzerland University of California, Irvine, Irvine, CA, USA
J Rovnak H Sanfaçon
Colorado State University, Fort Collins, CO, USA Pacific Agri-Food Research Centre, Summerland, BC,
Canada
D J Rowlands
University of Leeds, Leeds, UK R Sanjuán
Instituto de Biologı́a Molecular y Cellular de Plantas,
P Roy
CSIC-UPV, Valencia, Spain
London School of Hygiene and Tropical Medicine,
London, UK N Santi
Norwegian School of Veterinary Science, Oslo, Norway
L Rubino
Istituto di Virologia Vegetale del CNR, Bari, Italy C Sarmiento
Tallinn University of Technology, Tallinn, Estonia
R W H Ruigrok
CNRS, Grenoble, France T Sasaya
National Agricultural Research Center, Ibaraki, Japan
C E Rupprecht
Centers for Disease Control and Prevention, Atlanta, Q J Sattentau
GA, USA University of Oxford, Oxford, UK
R J Russell C Savolainen-Kopra
University of St. Andrews, St. Andrews, UK National Public Health Institute (KTL), Helsinki, Finland
B E Russ B Schaffhausen
The University of Melbourne, Parkville, VIC, Australia Tufts University School of Medicine, Boston, MA, USA
W T Ruyechan K Scheets
The State University of New York, Buffalo, NY, USA Oklahoma State University, Stillwater, OK, USA
E Ryabov M J Schmitt
University of Warwick, Warwick, UK University of the Saarland, Saarbrücken, Germany
M D Ryan A Schneemann
University of St. Andrews, St. Andrews, UK The Scripps Research Institute, La Jolla, CA, USA
E P Rybicki G Schoehn
University of Cape Town, Cape Town, South Africa CNRS, Grenoble, France
K D Ryman J E Schoelz
Louisiana State University Health Sciences Center at University of Missouri, Columbia, MO, USA
Shreveport, Shreveport, LA, USA
L B Schonberger
K D Ryman Centers for Disease Control and Prevention, Atlanta, GA,
Louisiana State University Health Sciences Center, USA
Shreveport, LA, USA
U Schubert
K H Ryu Klinikum der Universität Erlangen-Nürnberg, Erlangen,
Seoul Women’s University, Seoul, South Korea Germany
M Safak D A Schultz
Temple University School of Medicine, Philadelphia, PA, Johns Hopkins University School of Medicine, Baltimore,
USA MD, USA
M Salas S Schultz-Cherry
Universidad Autónoma, Madrid, Spain University of Wisconsin – Madison, Madison, WI, USA
S K Samal T F Schulz
University of Maryland, College Park, MD, USA Hannover Medical School, Hannover, Germany
xxvi Contributors

P D Scotti P Sreenivasulu
Waiatarua, New Zealand Sri Venkateswara University, Tirupati, India
B L Semler J Stanley
University of California, Irvine, CA, USA John Innes Centre, Colney, UK

J M Sharp K M Stedman
Veterinary Laboratories Agency, Penicuik, UK Portland State University, Portland, OR, USA

M L Shaw D Stephan
Mount Sinai School of Medicine, New York, NY, USA Institute of Plant Diseases and Plant Protection,
Hannover, Germany
G R Shellam
The University of Western Australia, C C M M Stijger
Crawley, WA, Australia Wageningen University and Research Centre, Naaldwijk,
The Netherlands
D N Shepherd
University of Cape Town, Cape Town, South Africa L Stitz
Federal Research Institute for Animal Health, Tuebingen,
N C Sheppard Gemany
University of Oxford, Oxford, UK
P G Stockley
F Shewmaker University of Leeds, Leeds, UK
National Institutes of Health, Bethesda, MD, USA
M R Strand
P A Signoret University of Georgia, Athens, GA, USA
Montpellier SupAgro, Montpellier, France
M J Studdert
A Silaghi The University of Melbourne, Parkville, VIC, Australia
University of Manitoba, Winnipeg, MB, Canada
C A Suttle
G Silvestri University of British Columbia, Vancouver, BC,
University of Pennsylvania, Philadelphia, PA, USA Canada
T L Sit N Suzuki
North Carolina State University, Raleigh, NC, USA Okayama University, Okayama, Japan
N Sittidilokratna J Y Suzuki
Centex Shrimp and Center for Genetic Engineering and USDA, Pacific Basin Agricultural Research Center, Hilo,
Biotechnology, Bangkok, Thailand HI, USA

M A Skinner R Swanepoel
Imperial College London, London, UK National Institute for Communicable Diseases,
Sandringham, South Africa
D W Smith
PathWest Laboratory Medicine WA, Nedlands, WA, S J Symes
Australia The University of Melbourne, Parkville, VIC, Australia

G L Smith G Szittya
Imperial College London, London, UK Agricultural Biotechnology Center, Godollo, Hungary

L M Smith M Taliansky
The University of Western Australia, Scottish Crop Research Institute, Dundee, UK
Crawley, WA, Australia P Tattersall
E J Snijder Yale University Medical School, New Haven, CT, USA
Leiden University Medical Center, Leiden, The T Tatusova
Netherlands National Institutes of Health, Bethesda, MD, USA
M Sova S Tavantzis
University of Texas Medical Branch – Galveston, University of Maine, Orono, ME, USA
Galveston, TX, USA
J M Taylor
J A Speir Fox Chase Cancer Center, Philadelphia, PA, USA
The Scripps Research Institute, La Jolla, CA, USA
D A Theilmann
T E Spencer Agriculture and Agri-Food Canada, Summerland, BC,
Texas A&M University, College Station, TX, USA Canada
Contributors xxvii

F C Thomas Allnutt R Vainionpää

National Science Foundation, Arlington, VA, USA University of Turku, Turku, Finland

G J Thomas Jr. A M Vaira

University of Missouri – Kansas City, Kansas City, MO, Istituto di Virologia Vegetale, CNR, Turin, Italy
USA
N K Van Alfen
J E Thomas University of California, Davis, CA, USA
Department of Primary Industries and Fisheries,
Indooroopilly, QLD, Australia R A A Van der Vlugt
Wageningen University and Research Centre,
H C Thomas Wageningen, The Netherlands
Imperial College London, London, UK
M H V Van Regenmortel
A N Thorburn CNRS, Illkirch, France
The University of Melbourne, Parkville, VIC, Australia
P A Venter
P Tijssen The Scripps Research Institute, La Jolla, CA, USA
Université du Québec, Laval, QC, Canada
J Verchot-Lubicz
S A Tolin Oklahoma State University, Stillwater, OK, USA
Virginia Polytechnic Institute and State University,
Blacksburg, VA, USA R A Vere Hodge
Vere Hodge Antivirals Ltd., Reigate, UK
L Torrance
Scottish Crop Research Institute, Invergowrie, UK H J Vetten
Federal Research Centre for Agriculture and Forestry
S Trapp
(BBA), Brunswick, Germany
Cornell University, Ithaca, NY, USA
L P Villarreal
S Tripathi
University of California, Irvine, Irvine, CA, USA
USDA, Pacific Basin Agricultural Research Center, Hilo,
HI, USA J M Vlak
Wageningen University, Wageningen, The Netherlands
E Truve
Tallinn University of Technology, Tallinn, Estonia P K Vogt
J-M Tsai The Scripps Research Institute, La Jolla, CA, USA
National Taiwan University, Taipei, Republic of China L E Volkman
M Tsompana University of California, Berkeley, Berkeley, CA, USA
North Carolina State University, Raleigh, NC, USA J Votteler
Klinikum der Universität Erlangen-Nürnberg, Erlangen,
R Tuma
Germany
University of Helsinki, Helsinki, Finland
A S Turnell D F Voytas
The University of Birmingham, Birmingham, UK Iowa State University, Ames, IA, USA

K L Tyler J D F Wadsworth
University of Colorado Health Sciences, Denver, CO, USA University College London, London, UK

A Uchiyama E K Wagner
Cornell University, Ithaca, NY, USA University of California, Irvine, Irvine, CA, USA

C Upton P J Walker
University of Victoria, Victoria, BC, Canada CSIRO Australian Animal Health Laboratory, Geelong,
VIC, Australia
A Urisman
University of California, San Francisco, A L Wang
San Francisco, CA, USA University of California, San Francisco, CA, USA

J K Uyemoto X Wang
University of California, Davis, CA, USA University of Wisconsin – Madison, Madison, WI, USA

A Vaheri C C Wang
University of Helsinki, Helsinki, Finland University of California, San Francisco, CA, USA
xxviii Contributors

L-F Wang R G Will

Australian Animal Health Laboratory, Geelong, VIC, Western General Hospital, Edinburgh, UK
Australia
T Williams
R Warrier Instituto de Ecologı́a A.C., Xalapa, Mexico
Purdue University, West Lafayette, IN, USA
K Willoughby
S C Weaver Moredun Research Institute, Edinburgh, UK
University of Texas Medical Branch, Galveston, TX, USA
S Winter
B A Webb Deutsche Sammlung für Mikroorganismen und
University of Kentucky, Lexington, KY, USA Zellkulturen, Brunswick, Germany
F Weber J Winton
University of Freiburg, Freiburg, Germany Western Fisheries Research Center, Seattle, WA, USA
R P Weir J K Yamamoto
Berrimah Research Farm, Darwin, NT, Australia University of Florida, Gainesville, FL, USA
R A Weisberg M Yoshida
National Institutes of Health, Bethesda, MD, USA University of Tokyo, Chiba, Japan
W Weissenhorn N Yoshikawa
UMR 5233 UJF-EMBL-CNRS, Grenoble, France Iwate University, Ueda, Japan
R M Welsh L S Young
University of Massachusetts Medical School, Worcester, University of Birmingham, Birmingham, UK
MA, USA
R F Young,III
J T West Texas A&M University, College Station, TX, USA
University of Oklahoma Health Sciences Center,
T M Yuill
Oklahoma City, OK, USA
University of Wisconsin, Madison, WI, USA
E Westhof
A J Zajac
Université Louis Pasteur, Strasbourg, France
University of Alabama at Birmingham, Birmingham, AL,
S P J Whelan USA
Harvard Medical School, Boston, MA, USA
S K Zavriev
R L White Shemyakin and Ovchinnikov Institute of Bioorganic
Texas A&M University, College Station, TX, USA Chemistry, Russian Academy of Sciences,
Moscow, Russia
C A Whitehouse
United States Army Medical Research Institute of J Ziebuhr
Infectious Diseases, Frederick, MD, USA The Queen’s University of Belfast, Belfast, UK
R B Wickner E I Zuniga
National Institutes of Health, Bethesda, MD, USA The Scripps Research Institute, La Jolla, CA, USA
CONTENTS

Editors-in-Chief v
Associate Editors vii
Preface ix
How to Use the Encyclopedia xi
Contributors xiii

VOLUME 1
A
Adenoviruses: General Features B Harrach 1
Adenoviruses: Malignant Transformation and Oncology A S Turnell 9
Adenoviruses: Molecular Biology K N Leppard 17
Adenoviruses: Pathogenesis M Benkő 24
African Cassava Mosaic Disease J P Legg 30
African Horse Sickness Viruses P S Mellor and P P C Mertens 37
African Swine Fever Virus L K Dixon and D Chapman 43
AIDS: Disease Manifestation A Rapose, J East, M Sova and W A O’Brien 51
AIDS: Global Epidemiology P J Peters, P H Kilmarx and T D Mastro 58
AIDS: Vaccine Development N C Sheppard and Q J Sattentau 69
Akabane Virus P S Mellor and P D Kirkland 76
Alfalfa Mosaic Virus J F Bol 81
Algal Viruses K Nagasaki and C P D Brussaard 87
Allexivirus S K Zavriev 96
Alphacryptovirus and Betacryptovirus R Blawid, D Stephan and E Maiss 98
Anellovirus P Biagini and P de Micco 104
Animal Rhabdoviruses H Bourhy, A J Gubala, R P Weir and D B Boyle 111
Antigen Presentation E I Zuniga, D B McGavern and M B A Oldstone 121
Antigenic Variation G M Air and J T West 127
Antigenicity and Immunogenicity of Viral Proteins M H V Van Regenmortel 137

xxix
XXX Contents

Antiviral Agents H J Field and R A Vere Hodge 142

Apoptosis and Virus Infection J R Clayton and J M Hardwick 154
Aquareoviruses M St J Crane and G Carlile 163
Arboviruses B R Miller 170
Arteriviruses M A Brinton and E J Snijder 176
Ascoviruses B A Federici and Y Bigot 186
Assembly of Viruses: Enveloped Particles C K Navaratnarajah, R Warrier and R J Kuhn 193
Assembly of Viruses: Nonenveloped Particles M Luo 200
Astroviruses L Moser and S Schultz-Cherry 204

B
Baculoviruses: Molecular Biology of Granuloviruses S Hilton 211
Baculoviruses: Molecular Biology of Mosquito Baculoviruses J J Becnel and C L Afonso 219
Baculoviruses: Molecular Biology of Sawfly Baculoviruses B M Arif 225
Baculoviruses: Apoptosis Inhibitors R J Clem 231
Baculoviruses: Expression Vector F J Haines, R D Possee and L A King 237
Baculoviruses: General Features P J Krell 247
Baculoviruses: Molecular Biology of Nucleopolyhedroviruses D A Theilmann and G W Blissard 254
Baculoviruses: Pathogenesis L E Volkman 265
Banana Bunchy Top Virus J E Thomas 272
Barley Yellow Dwarf Viruses L L Domier 279
Barnaviruses P A Revill 286
Bean Common Mosaic Virus and Bean Common Mosaic Necrosis Virus R Jordan and J Hammond 288
Bean Golden Mosaic Virus F J Morales 295
Beet Curly Top Virus J Stanley 301
Benyvirus R Koenig 308
Beta ssDNA Satellites R W Briddon and S Mansoor 314
Birnaviruses B Delmas 321
Bluetongue Viruses P Roy 328
Border Disease Virus P Nettleton and K Willoughby 335
Bornaviruses L Stitz, O Planz and W Ian Lipkin 341
Bovine and Feline Immunodeficiency Viruses J K Yamamoto 347
Bovine Ephemeral Fever Virus P J Walker 354
Bovine Herpesviruses M J Studdert 362
Bovine Spongiform Encephalopathy R G Will 368
Bovine Viral Diarrhea Virus J F Ridpath 374
Brome Mosaic Virus X Wang and P Ahlquist 381
Bromoviruses J J Bujarski 386
Contents XXXI

Bunyaviruses: General Features R M Elliott 390

Bunyaviruses: Unassigned C H Calisher 399

C
Cacao Swollen Shoot Virus E Muller 403
Caliciviruses M J Studdert and S J Symes 410
Capillovirus, Foveavirus, Trichovirus, Vitivirus N Yoshikawa 419
Capripoxviruses R P Kitching 427
Capsid Assembly: Bacterial Virus Structure and Assembly S Casjens 432
Cardioviruses C Billinis and O Papadopoulos 440
Carlavirus K H Ryu and B Y Lee 448
Carmovirus F Qu and T J Morris 453
Caulimoviruses: General Features J E Schoelz 457
Caulimoviruses: Molecular Biology T Hohn 464
Central Nervous System Viral Diseases R T Johnson and B M Greenberg 469
Cereal Viruses: Maize/Corn P A Signoret 475
Cereal Viruses: Rice F Morales 482
Cereal Viruses: Wheat and Barley H D Lapierre and D Hariri 490
Chandipura Virus S Basak and D Chattopadhyay 497
Chrysoviruses S A Ghabrial 503
Circoviruses A Mankertz 513
Citrus Tristeza Virus M Bar-Joseph and W O Dawson 520
Classical Swine Fever Virus V Moennig and I Greiser-Wilke 525
Coltiviruses H Attoui and X de Lamballerie 533
Common Cold Viruses S Dreschers and C Adams 541
Coronaviruses: General Features D Cavanagh and P Britton 549
Coronaviruses: Molecular Biology S C Baker 554
Cotton Leaf Curl Disease S Mansoor, I Amin and R W Briddon 563
Cowpea Mosaic Virus G P Lomonossoff 569
Cowpox Virus M Bennett, G L Smith and D Baxby 574
Coxsackieviruses M S Oberste and M A Pallansch 580
Crenarchaeal Viruses: Morphotypes and Genomes D Prangishvili, T Basta and R A Garrett 587
Crimean–Congo Hemorrhagic Fever Virus and Other Nairoviruses C A Whitehouse 596
Cryo-Electron Microscopy W Chiu, J T Chang and F J Rixon 603
Cucumber Mosaic Virus F Garcı́a-Arenal and P Palukaitis 614
Cytokines and Chemokines D E Griffin 620
Cytomegaloviruses: Murine and Other Nonprimate Cytomegaloviruses A J Redwood, L M Smith and
G R Shellam 624
Cytomegaloviruses: Simian Cytomegaloviruses D J Alcendor and G S Hayward 634
XXXII Contents

VOLUME 2
D
Defective-Interfering Viruses L Roux 1
Dengue Viruses D J Gubler 5
Diagnostic Techniques: Microarrays K Fischer, A Urisman and J DeRisi 14
Diagnostic Techniques: Plant Viruses R Koenig, D-E Lesemann, G Adam and S Winter 18
Diagnostic Techniques: Serological and Molecular Approaches R Vainionpää and P Leinikki 29
Dicistroviruses P D Christian and P D Scotti 37
Disease Surveillance N Noah 44
DNA Vaccines S Babiuk and L A Babiuk 51

E
Ebolavirus K S Brown, A Silaghi and H Feldmann 57
Echoviruses T Hyypiä 65
Ecology of Viruses Infecting Bacteria S T Abedon 71
Electron Microscopy of Viruses G Schoehn and R W H Ruigrok 78
Emerging and Reemerging Virus Diseases of Plants G P Martelli and D Gallitelli 86
Emerging and Reemerging Virus Diseases of Vertebrates B W J Mahy 93
Emerging Geminiviruses C M Fauquet and M S Nawaz-ul-Rehman 97
Endogenous Retroviruses W E Johnson 105
Endornavirus T Fukuhara and H Moriyama 109
Enteric Viruses R F Bishop and C D Kirkwood 116
Enteroviruses of Animals L E Hughes and M D Ryan 123
Enteroviruses: Human Enteroviruses Numbered 68 and Beyond T Hovi, S Blomqvist,
C Savolainen-Kopra and M Roivainen 130
Entomopoxviruses M N Becker and R W Moyer 136
Epidemiology of Human and Animal Viral Diseases F A Murphy 140
Epstein–Barr Virus: General Features L S Young 148
Epstein–Barr Virus: Molecular Biology J T Sample and C E Sample 157
Equine Infectious Anemia Virus J K Craigo and R C Montelaro 167
Evolution of Viruses L P Villarreal 174

F
Feline Leukemia and Sarcoma Viruses J C Neil 185
Filamentous ssDNA Bacterial Viruses S A Overman and G J Thomas Jr. 190
Filoviruses G Olinger, T W Geisbert and L E Hensley 198
Fish and Amphibian Herpesviruses A J Davison 205
Fish Retroviruses T A Paul, R N Casey, P R Bowser, J W Casey, J Rovnak and S L Quackenbush 212
Contents XXXIII

Fish Rhabdoviruses G Kurath and J Winton 221

Fish Viruses J C Leong 227
Flaviviruses of Veterinary Importance R Swanepoel and F J Burt 234
Flaviviruses: General Features T J Chambers 241
Flexiviruses M J Adams 253
Foamy Viruses M L Linial 259
Foot and Mouth Disease Viruses D J Rowlands 265
Fowlpox Virus and Other Avipoxviruses M A Skinner 274
Fungal Viruses S A Ghabrial and N Suzuki 284
Furovirus R Koenig 291
Fuselloviruses of Archaea K M Stedman 296

G
Gene Therapy: Use of Viruses as Vectors K I Berns and T R Flotte 301
Genome Packaging in Bacterial Viruses P Jardine 306
Giardiaviruses A L Wang and C C Wang 312

H
Hantaviruses A Vaheri 317
Henipaviruses B T Eaton and L-F Wang 321
Hepadnaviruses of Birds A R Jilbert and W S Mason 327
Hepadnaviruses: General Features T J Harrison 335
Hepatitis A Virus A Dotzauer 343
Hepatitis B Virus: General Features P Karayiannis and H C Thomas 350
Hepatitis B Virus: Molecular Biology T J Harrison 360
Hepatitis C Virus R Bartenschlager and S Bühler 367
Hepatitis Delta Virus J M Taylor 375
Hepatitis E Virus X J Meng 377
Herpes Simplex Viruses: General Features L Aurelian 383
Herpes Simplex Viruses: Molecular Biology E K Wagner and R M Sandri-Goldin 397
Herpesviruses of Birds S Trapp and N Osterrieder 405
Herpesviruses of Horses D J O’Callaghan and N Osterrieder 411
Herpesviruses: Discovery B Ehlers 420
Herpesviruses: General Features A J Davison 430
Herpesviruses: Latency C M Preston 436
History of Virology: Bacteriophages H-W Ackermann 442
History of Virology: Plant Viruses R Hull 450
History of Virology: Vertebrate Viruses F J Fenner 455
XXXIV Contents

Hordeivirus J N Bragg, H-S Lim and A O Jackson 459

Host Resistance to Retroviruses T Hatziioannou and P D Bieniasz 467
Human Cytomegalovirus: General Features E S Mocarski Jr. and R F Pass 474
Human Cytomegalovirus: Molecular Biology W Gibson 485
Human Eye Infections J Chodosh, A V Chintakuntlawar and C M Robinson 491
Human Herpesviruses 6 and 7 U A Gompels 498
Human Immunodeficiency Viruses: Antiretroviral Agents A W Neuman and D C Liotta 505
Human Immunodeficiency Viruses: Molecular Biology J Votteler and U Schubert 517
Human Immunodeficiency Viruses: Origin F van Heuverswyn and M Peeters 525
Human Immunodeficiency Viruses: Pathogenesis N R Klatt, A Chahroudi and G Silvestri 534
Human Respiratory Syncytial Virus P L Collins 542
Human Respiratory Viruses J E Crowe Jr. 551
Human T-Cell Leukemia Viruses: General Features M Yoshida 558
Human T-Cell Leukemia Viruses: Human Disease R Mahieux and A Gessain 564
Hypovirulence N K Van Alfen and P Kazmierczak 574
Hypoviruses D L Nuss 580

VOLUME 3
I
Icosahedral dsDNA Bacterial Viruses with an Internal Membrane J K H Bamford and S J Butcher 1
Icosahedral Enveloped dsRNA Bacterial Viruses R Tuma 6
Icosahedral ssDNA Bacterial Viruses B A Fane, M Chen, J E Cherwa and A Uchiyama 13
Icosahedral ssRNA Bacterial Viruses P G Stockley 21
Icosahedral Tailed dsDNA Bacterial Viruses R L Duda 30
Idaeovirus A T Jones and H Barker 37
Iflavirus M M van Oers 42
Ilarvirus K C Eastwell 46
Immune Response to Viruses: Antibody-Mediated Immunity A R Neurath 56
Immune Response to Viruses: Cell-Mediated Immunity A J Zajac and L E Harrington 70
Immunopathology M B A Oldstone and R S Fujinami 78
Infectious Pancreatic Necrosis Virus Ø Evensen and N Santi 83
Infectious Salmon Anemia Virus B H Dannevig, S Mjaaland and E Rimstad 89
Influenza R A Lamb 95
Innate Immunity: Defeating C F Basler 104
Innate Immunity: Introduction F Weber 111
Inoviruses L A Day 117
Insect Pest Control by Viruses M Erlandson 125
Insect Reoviruses P P C Mertens and H Attoui 133
Insect Viruses: Nonoccluded J P Burand 144
Contents XXXV

Interfering RNAs K E Olson, K M Keene and C D Blair 148

Iridoviruses of Vertebrates A D Hyatt and V G Chinchar 155
Iridoviruses of Invertebrates T Williams and A D Hyatt 161
Iridoviruses: General Features V G Chinchar and A D Hyatt 167

J
Jaagsiekte Sheep Retrovirus J M Sharp, M de las Heras, T E Spencer and M Palmarini 175
Japanese Encephalitis Virus A D T Barrett 182

K
Kaposi’s Sarcoma-Associated Herpesvirus: General Features Y Chang and P S Moore 189
Kaposi’s Sarcoma-Associated Herpesvirus: Molecular Biology E Gellermann and T F Schulz 195

L
Lassa, Junin, Machupo and Guanarito Viruses J B McCormick 203
Legume Viruses L Bos 212
Leishmaniaviruses R Carrion Jr, Y-T Ro and J L Patterson 220
Leporipoviruses and Suipoxviruses G McFadden 225
Luteoviruses L L Domier and C J D’Arcy 231
Lymphocytic Choriomeningitis Virus: General Features R M Welsh 238
Lymphocytic Choriomeningitis Virus: Molecular Biology J C de la Torre 243
Lysis of the Host by Bacteriophage R F Young III and R L White 248

M
Machlomovirus K Scheets 259
Maize Streak Virus D P Martin, D N Shepherd and E P Rybicki 263
Marburg Virus D Falzarano and H Feldmann 272
Marnaviruses A S Lang and C A Suttle 280
Measles Virus R Cattaneo and M McChesney 285
Membrane Fusion A Hinz and W Weissenhorn 292
Metaviruses H L Levin 301
Mimivirus J-M Claverie 311
Molluscum Contagiosum Virus J J Bugert 319
Mononegavirales A J Easton and R Ling 324
Mouse Mammary Tumor Virus J P Dudley 334
Mousepox and Rabbitpox Viruses M Regner, F Fenner and A Müllbacher 342
Movement of Viruses in Plants P A Harries and R S Nelson 348
XXXVI Contents

Mumps Virus B K Rima and W P Duprex 356

Mungbean Yellow Mosaic Viruses V G Malathi and P John 364
Murine Gammaherpesvirus 68 A A Nash and B M Dutia 372
Mycoreoviruses B I Hillman 378

N
Nanoviruses H J Vetten 385
Narnaviruses R Esteban and T Fujimura 392
Nature of Viruses M H V Van Regenmortel 398
Necrovirus L Rubino and G P Martelli 403
Nepovirus H Sanfaçon 405
Neutralization of Infectivity P J Klasse 413
Nidovirales L Enjuanes, A E Gorbalenya, R J de Groot, J A Cowley, J Ziebuhr and E J Snijder 419
Nodaviruses P A Venter and A Schneemann 430
Noroviruses and Sapoviruses K Y Green 438

O
Ophiovirus A M Vaira and R G Milne 447
Orbiviruses P P C Mertens, H Attoui and P S Mellor 454
Organ Transplantation, Risks C N Kotton, M J Kuehnert and J A Fishman 466
Origin of Viruses P Forterre 472
Orthobunyaviruses C H Calisher 479
Orthomyxoviruses: Molecular Biology M L Shaw and P Palese 483
Orthomyxoviruses: Structure of Antigens R J Russell 489
Oryctes Rhinoceros Virus J M Vlak, A M Huger, J A Jehle and R G Kleespies 495
Ourmiavirus G P Accotto and R G Milne 500

VOLUME 4
P
Papaya Ringspot Virus D Gonsalves, J Y Suzuki, S Tripathi and S A Ferreira 1
Papillomaviruses: General Features of Human Viruses G Orth 8
Papillomaviruses: Molecular Biology of Human Viruses P F Lambert and A Collins 18
Papillomaviruses of Animals A A McBride 26
Papillomaviruses: General Features H U Bernard 34
Paramyxoviruses of Animals S K Samal 40
Parainfluenza Viruses of Humans E Adderson and A Portner 47
Paramyxoviruses R E Dutch 52
Parapoxviruses D Haig and A A Mercer 57
Contents XXXVII

Partitiviruses of Fungi S Tavantzis 63

Partitiviruses: General Features S A Ghabrial, W F Ochoa, T S Baker and M L Nibert 68
Parvoviruses of Arthropods M Bergoin and P Tijssen 76
Parvoviruses of Vertebrates C R Parrish 85
Parvoviruses: General Features P Tattersall 90
Pecluvirus D V R Reddy, C Bragard, P Sreenivasulu and P Delfosse 97
Pepino Mosaic Virus R A A Van der Vlugt and C C M M Stijger 103
Persistent and Latent Viral Infection E S Mocarski and A Grakoui 108
Phycodnaviruses J L Van Etten and M V Graves 116
Phylogeny of Viruses A E Gorbalenya 125
Picornaviruses: Molecular Biology B L Semler and K J Ertel 129
Plant Antiviral Defense: Gene Silencing Pathway G Szittya, T Dalmay and J Burgyan 141
Plant Reoviruses R J Geijskes and R M Harding 149
Plant Resistance to Viruses: Engineered Resistance M Fuchs 156
Plant Resistance to Viruses: Geminiviruses J K Brown 164
Plant Resistance to Viruses: Natural Resistance Associated with Dominant Genes
P Moffett and D F Klessig 170
Plant Resistance to Viruses: Natural Resistance Associated with Recessive Genes
C Caranta and C Dogimont 177
Plant Rhabdoviruses A O Jackson, R G Dietzgen, R-X Fang, M M Goodin, S A Hogenhout,
M Deng and J N Bragg 187
Plant Virus Diseases: Economic Aspects G Loebenstein 197
Plant Virus Diseases: Fruit Trees and Grapevine G P Martelli and J K Uyemoto 201
Plant Virus Diseases: Ornamental Plants J Engelmann and J Hamacher 207
Plant Virus Vectors (Gene Expression Systems) Y Gleba, S Marillonnet and V Klimyuk 229
Plum Pox Virus M Glasa and T Candresse 238
Poliomyelitis P D Minor 242
Polydnaviruses: Abrogation of Invertebrate Immune Systems M R Strand 250
Polydnaviruses: General Features A Fath-Goodin and B A Webb 256
Polyomaviruses of Humans M Safak and K Khalili 261
Polyomaviruses of Mice B Schaffhausen 271
Polyomaviruses M Gravell and E O Major 277
Pomovirus L Torrance 282
Potato Virus Y C Kerlan and B Moury 287
Potato Viruses C Kerlan 296
Potexvirus K H Ryu and J S Hong 310
Potyviruses J J López-Moya and J A Garcı́a 313
Poxviruses G L Smith, P Beard and M A Skinner 322
Prions of Vertebrates J D F Wadsworth and J Collinge 330
Prions of Yeast and Fungi R B Wickner, H Edskes, T Nakayashiki, F Shewmaker, L McCann,
A Engel and D Kryndushkin 336
XXXVIII Contents

Pseudorabies Virus T C Mettenleiter 341

Pseudoviruses D F Voytas 352

Q
Quasispecies R Sanjuán 359

R
Rabies Virus I V Kuzmin and C E Rupprecht 367
Recombination J J Bujarski 374
Reoviruses: General Features P Clarke and K L Tyler 382
Reoviruses: Molecular Biology K M Coombs 390
Replication of Bacterial Viruses M Salas and M de Vega 399
Replication of Viruses A J Cann 406
Reticuloendotheliosis Viruses A S Liss and H R Bose Jr. 412
Retrotransposons of Fungi T J D Goodwin, M I Butler and R T M Poulter 419
Retrotransposons of Plants M-A Grandbastien 428
Retrotransposons of Vertebrates A E Peaston 436
Retroviral Oncogenes P K Vogt and A G Bader 445
Retroviruses of Insects G F Rohrmann 451
Retroviruses of Birds K L Beemon 455
Retroviruses: General Features E Hunter 459
Rhinoviruses N W Bartlett and S L Johnston 467
Ribozymes E Westhof and A Lescoute 475
Rice Tungro Disease R Hull 481
Rice Yellow Mottle Virus E Hébrard and D Fargette 485
Rift Valley Fever and Other Phleboviruses L Nicoletti and M G Ciufolini 490
Rinderpest and Distemper Viruses T Barrett 497
Rotaviruses J Angel, M A Franco and H B Greenberg 507
Rubella Virus T K Frey 514

S
Sadwavirus T Iwanami 523
Satellite Nucleic Acids and Viruses P Palukaitis, A Rezaian and F Garcı́a-Arenal 526
Seadornaviruses H Attoui and P P C Mertens 535
Sequiviruses I-R Choi 546
Severe Acute Respiratory Syndrome (SARS) J S M Peiris and L L M Poon 552
Shellfish Viruses T Renault 560
Shrimp Viruses J-R Bonami 567
Contents XXXIX

Sigma Rhabdoviruses D Contamine and S Gaumer 576

Simian Alphaherpesviruses J Hilliard 581
Simian Gammaherpesviruses A Ensser 585
Simian Immunodeficiency Virus: Animal Models of Disease C J Miller and M Marthas 594
Simian Immunodeficiency Virus: General Features M E Laird and R C Desrosiers 603
Simian Immunodeficiency Virus: Natural Infection I Pandrea, G Silvestri and C Apetrei 611
Simian Retrovirus D P A Marx 623
Simian Virus 40 A L McNees and J S Butel 630
Smallpox and Monkeypox Viruses S Parker, D A Schultz, H Meyer and R M Buller 639
Sobemovirus M Meier, A Olspert, C Sarmiento and E Truve 644
St. Louis Encephalitis W K Reisen 652
Sweetpotato Viruses J Kreuze and S Fuentes 659

VOLUME 5
T
Taura Syndrome Virus A K Dhar and F C T Allnutt 1
Taxonomy, Classification and Nomenclature of Viruses C M Fauquet 9
Tenuivirus B C Ramirez 24
Tetraviruses J A Speir and J E Johnson 27
Theiler’s Virus H L Lipton, S Hertzler and N Knowles 37
Tick-Borne Encephalitis Viruses T S Gritsun and E A Gould 45
Tobacco Mosaic Virus M H V Van Regenmortel 54
Tobacco Viruses S A Tolin 60
Tobamovirus D J Lewandowski 68
Tobravirus S A MacFarlane 72
Togaviruses Causing Encephalitis S Paessler and M Pfeffer 76
Togaviruses Causing Rash and Fever D W Smith, J S Mackenzie and M D A Lindsay 83
Togaviruses Not Associated with Human Disease L L Coffey, 91
Togaviruses: Alphaviruses A M Powers 96
Togaviruses: Equine Encephalitic Viruses D E Griffin 101
Togaviruses: General Features S C Weaver, W B Klimstra and K D Ryman 107
Togaviruses: Molecular Biology K D Ryman, W B Klimstra and S C Weaver 116
Tomato Leaf Curl Viruses from India S Chakraborty 124
Tomato Spotted Wilt Virus H R Pappu 133
Tomato Yellow Leaf Curl Virus H Czosnek 138
Tombusviruses S A Lommel and T L Sit 145
Torovirus A E Hoet and M C Horzinek 151
Tospovirus M Tsompana and J W Moyer 157
Totiviruses S A Ghabrial 163
XL Contents

Transcriptional Regulation in Bacteriophage R A Weisberg, D M Hinton and S Adhya 174

Transmissible Spongiform Encephalopathies E D Belay and L B Schonberger 186
Tumor Viruses: Human R Grassmann, B Fleckenstein and H Pfister 193
Tymoviruses A-L Haenni and T W Dreher 199

U
Umbravirus M Taliansky and E Ryabov 209
Ustilago Maydis Viruses J Bruenn 214

V
Vaccine Production in Plants E P Rybicki 221
Vaccine Safety C J Clements and G Lawrence 226
Vaccine Strategies I Kusters and J W Almond 235
Vaccinia Virus G L Smith 243
Varicella-Zoster Virus: General Features J I Cohen 250
Varicella-Zoster Virus: Molecular Biology W T Ruyechan and J Hay 256
Varicosavirus T Sasaya 263
Vector Transmission of Animal Viruses W K Reisen 268
Vector Transmission of Plant Viruses S Blanc 274
Vegetable Viruses P Caciagli 282
Vesicular Stomatitis Virus S P J Whelan 291
Viral Killer Toxins M J Schmitt 299
Viral Membranes J Lenard 308
Viral Pathogenesis N Nathanson 314
Viral Receptors D J Evans 319
Viral Suppressors of Gene Silencing J Verchot-Lubicz and J P Carr 325
Viroids R Flores and R A Owens 332
Virus Classification by Pairwise Sequence Comparison (PASC) Y Bao, Y Kapustin and T Tatusova 342
Virus Databases E J Lefkowitz, M R Odom and C Upton 348
Virus Entry to Bacterial Cells M M Poranen and A Domanska 365
Virus Evolution: Bacterial Viruses R W Hendrix 370
Virus-Induced Gene Silencing (VIGS) M S Padmanabhan and S P Dinesh-Kumar 375
Virus Particle Structure: Nonenveloped Viruses J A Speir and J E Johnson 380
Virus Particle Structure: Principles J E Johnson and J A Speir 393
Virus Species M H V van Regenmortel 401
Viruses and Bioterrorism R F Meyer and S A Morse 406
Viruses Infecting Euryarchaea K Porter, B E Russ, A N Thorburn and M L Dyall-Smith 411
Visna-Maedi Viruses B A Blacklaws 423
Contents XLI

W
Watermelon Mosaic Virus and Zucchini Yellow Mosaic Virus H Lecoq and C Desbiez 433
West Nile Virus L D Kramer 440
White Spot Syndrome Virus J-H Leu, J-M Tsai and C-F Lo 450

Y
Yatapoxviruses J W Barrett and G McFadden 461
Yeast L-A Virus R B Wickner, T Fujimura and R Esteban 465
Yellow Fever Virus A A Marfin and T P Monath 469
Yellow Head Virus P J Walker and N Sittidilokratna 476

Z
Zoonoses J E Osorio and T M Yuill 485

Taxonomic Index 497

Introduction respectively. The other two genera (Atadenovirus and Siade-

novirus) have a broader range of hosts. Atadenoviruses were
Adenoviruses are middle-sized, nonenveloped, icosahe- named after a bias toward high AþTcontent in the genomes
dral, double-stranded DNA viruses of animals. The prefix of the initial representatives, which infect various ruminant
adeno comes from the Greek word a dZn (gland), reflect- and avian hosts, as well as a marsupial. Every known reptil-
ing the first isolation of a virus of this type from human ian adenovirus also belongs to the atadenoviruses, although
adenoid tissue half a century ago. Adenoviruses have since their genomes do not show the same bias toward high AþT
been isolated from a large variety of hosts, including content. The very few known siadenoviruses were isolated
representatives of every major vertebrate class from fish from or detected by PCR in birds and a frog. This genus was
to mammals. Using polymerase chain reaction (PCR) named in recognition of the presence of a gene encoding a
technology, a large variety of putative novel adenoviru- potential sialidase in the viruses concerned. The single
ses have been detected, but isolation of such viruses and confirmed fish adenovirus falls into a separate group that
in vitro propagation is hampered in most cases by the lack may eventually found a fifth genus; adenovirus-like parti-
of appropriate permissive cell cultures. Some human and cles have been described in additional fish species.
animal adenoviruses can cause diseases or even death, but Within each genus, the viruses are grouped into species,
most are not pathogenic in non-immuno-compromised, which are named according to the host first described and
healthy individuals. Adenoviruses have been used as model supplemented with letters of the alphabet (Table 1). Host
organisms in molecular biology, and important findings of origin is only one of several criteria that are used to demar-
general relevance have emerged from such studies, includ- cate the species. Phylogenetic distance is the most signifi-
ing splicing in eukaryotes. Adenoviruses have become one cant criterion, with species defined as separated by more
of the most popular vector systems for virus-based gene than 5–10% amino acid sequence divergence of hexon and
therapy and vaccination and have potential as antitumor DNA polymerase (pol), respectively. Further important
tools. Wide prevalence in diverse host species and a characteristics come into play, especially if DNA sequence
substantially conserved genome organization make ade- data are not available: DNA hybridization, restriction frag-
noviruses an ideal model for studying virus evolution. ment length polymorphism, nucleotide composition, onco-
genicity in rodents, growth characteristics, host range,
cross-neutralization, ability to recombine, number of
Taxonomy virus-associated (VA) RNA genes, hemagglutination
properties, and organization of the genome. However, all
Adenoviruses belong to the family Adenoviridae. No higher of these ancillary data are expected to accord with the
taxonomical level has yet been established, despite the fact results of phylogenetic calculations. Thus, for example,
that certain bacteriophages (Tectiviridae), the green alga chimpanzee adenoviruses are classified into human ade-
virus Paramecium bursaria Chlorella virus 1 (Phycodnaviridae), novirus species. Adenoviruses of humans have been stud-
and a virus of Archaea living in hot springs (sulfolobus ied far more intensively than those of other animals, and
turreted icosahedral virus) seem to have common evolu- the six species (Human adenovirus A to Human adenovirus F;
tionary roots with adenoviruses. abbreviated informally to HAdV-A to HAdV-F) corre-
There are four official genera in the family. Two genera spond to substantial ‘groups’ or ‘subgenera’ defined previ-
(Mastadenovirus and Aviadenovirus) comprise adenoviruses ously. Each human adenovirus serotype is abbreviated to
that have probably co-evolved with mammals and birds, HAdV hyphenated to a number.

1
2 Adenoviruses: General Features

Table 1 The taxonomy of family Adenoviridaea

Genus/species Serotype Strain Genome

Mastadenovirus
Bovine adenovirus A BAdV-1 G
Bovine adenovirus B BAdV-3 G
Bovine adenovirus C BAdV-10
Canine adenovirus CAdV-1, 2 G
Equine adenovirus A EAdV-1
Equine adenovirus B EAdV-2
Human adenovirus A HAdV-12, 18, 31 12
Human adenovirus B HAdV-3, 7, 11, 14, 16, 21, 34, 35, 50 G
Simian adenovirus 21 (SAdV-21) G
Human adenovirus C BAdV-9, HAdV-1, 2, 5, 6 1, 2, 5
Human adenovirus D HAdV-8–10, 13, 15, 17, 19, 20, 9, 17
22–30, 32, 33, 36–39, 42–49, 51 26, 46, 48, 49
Human adenovirus E HAdV-4, SAdV-22–25 G
Human adenovirus F HAdV-40, 41 G
Murine adenovirus A MAdV-1 G
Ovine adenovirus A BAdV-2, OAdV-2–5 BAdV-2
Ovine adenovirus B OAdV-1
Porcine adenovirus A PAdV-1, 2, 3 3
Porcine adenovirus B PAdV-4
Porcine adenovirus C PAdV-5 G
Tree shrew adenovirus TSAdV-1 G
Goat adenovirus (ts) GAdV-2
Guinea pig adenovirus (ts) GPAdV-1
Murine adenovirus B (ts) MAdV-2
Ovine adenovirus C (ts) OAdV-6
Simian adenovirus A (ts) SAdV-3 G
Squirrel adenovirus (ts) SqAdV-1
? HAdV-52 G
? SAdV-1–2, 4–20 1, 6, 7, 20
Aviadenovirus
Fowl adenovirus A FAdV-1 CELO G
Fowl adenovirus B FAdV-5 340
Fowl adenovirus C FAdV-4, 10 KR95, CFA20
Fowl adenovirus D FAdV-2, 3, 9, 11 P7-A, 75, A2-A, 380 9
Fowl adenovirus E FAdV-6, 7, 8a, 8b CR119, YR36, TR59, 764
Goose adenovirus GoAdV-1–3
Duck adenovirus B (ts) DAdV-2
Pigeon adenovirus (ts) PiAdV-1
Turkey adenovirus B (ts) TAdV-1, 2
Psittacine adenovirus? Psittacine adenovirus 1
Falcon adenovirus? Falcon adenovirus 1
Atadenovirus
Bovine adenovirus D BAdV-4, 5, 8, strain Rus 4
Duck adenovirus A DAdV-1 G
Ovine adenovirus D GAdV-1, OAdV-7 7
Possum adenovirus PoAdV-1
Bearded dragon adenovirus (ts) BDAdV-1
Bovine adenovirus E (ts) BAdV-6
Bovine adenovirus F (ts) BAdV-7
Cervine adenovirus (ts) Odocoileus adenovirus 1 (OdAdV-1)
Chameleon adenovirus (ts) ChAdV-1
Gecko adenovirus (ts) GeAdV-1 Fat-tailed gecko
Snake adenovirus (ts) SnAdV-1 Corn snake, python G
Gekkonid adenovirus(?) Tokay gecko adenovirus
Helodermatid adenovirus(?) Gila monster adenovirus
Scincid adenovirus(?) Blue-tongued skink adenovirus
Genus Siadenovirus
Frog adenovirus FrAdV-1 G
Turkey adenovirus A TAdV-3 G
? Raptor adenovirus 1 Harris hawk

Continued
Adenoviruses: General Features 3

Table 1 Continued

Genus/species Serotype Strain Genome

Unassigned Viruses in the Family

? White sturgeon adenovirus 1 (WSAdV-1)
? Crocodile adenovirus
a
Official genus and species names as published in the Eighth Report of the ICTV are in italics, and tentative species (ts), proposed
species (marked by a query) or single isolates are not. Because of confusion in the serotype numbering in some cases (e.g. among fowl
adenoviruses), certain characteristic strains are shown for easier identification. Available full genome sequences are noted by G or by
the number of the sequenced serotype(s) if those listed are not all available.

To illustrate the need to proceed carefully in developing present in the members of any other genus. Two mono-
adenovirus taxonomy, the case of the newest adenovirus mers of polypeptide IIIa penetrate the hexon shell at the
isolated from human samples (HAdV-52) is illuminating. edge of each facet, and multiple copies of polypeptide VI
This virus seems to be sufficiently different from other form a ring underneath the peripentonal hexons. Penton
human adenoviruses to merit the erection of a new species. bases are formed at the vertices by the interaction of five
However, it is very similar to some previously characterized copies of polypeptide III, and are tightly associated with
Old World monkey adenoviruses (simian adenoviruses 1 and one (or, in the aviadenoviruses, two) fibers, each consist-
7 (SAdV-1, SAdV-7) plus others). One taxonomical proposal ing of three copies of polypeptide IV in the form of a shaft
would be to establish a new species, Human adenovirus G, of characteristic length with a distal knob. Polypeptide
containing HAdV-52 and the related monkey adenoviruses. VIII is situated at the inner surface of the hexon shell.
Clearly, this would depend on epidemiological data demon- Polypeptides VI and VIII appear to link the capsid to the
strating that HAdV-52 is properly a human virus and not an virion core, which consists of a single copy of the DNA
occasional, opportunistic transfer from monkeys. For similar genome complexed with four polypeptides (V, VII, X,
reasons and others, classification of many nonhuman adeno- TP). Polypeptide V exists only in mastadenoviruses.
viruses into species is not yet resolved. Adenoviruses are stable on storage in the frozen state.
They are stable to mild acid and insensitive to lipid
solvents. Heat sensitivity varies among the genera.
Virion Morphology and Properties

The icosahedral capsid is 70–90 nm in diameter and con- Nucleic Acid, Genome Organization, and
sists of 240 nonvertex capsomers (called hexons), each Replication
8–10 nm in diameter, and 12 vertex capsomers (pentons),
each with a protruding fiber 9–77.5 nm in length. The The adenovirus genome is a linear molecule of double-
members of genus Aviadenovirus that have been studied stranded DNA (26 163–45 063 bp) containing an inverted
have two fiber proteins per vertex. Fowl adenovirus 1 terminal repeat (ITR) of 36–368 bp at its termini, with the 50
(FAdV-1) even has two, tandem fiber genes of different ends of the genome linked covalently to a terminal protein
lengths, resulting in two fibers of different sizes at each (TP). The nucleotide composition is 33.7–63.8% GþC.
vertex. Members of species Human adenovirus F (and The genetic organization of the central part of the genome
HAdV-52 and the related monkey viruses) also have two is conserved throughout the family, whereas the terminal
fiber genes of different lengths, but the fibers are parts show large variations in length and gene content
distributed in single copies alternately on the vertices. (Figure 1). Splicing was first discovered in adenoviruses,
The main capsomers (hexons) are formed by the interaction and is a common means of expressing mRNAs in this virus
of three identical polypeptides (designated hexon, and also family. In the conserved region, most late genes are
as polypeptide II, after a Roman numeral system based on expressed by splicing from the rightward-oriented major
the relative mobilities of structural proteins under reducing late promoter located in the pol gene. The early genes
conditions in sodium dodecyl sulfate-polyacrylamide gel encoding pol, the precursor of TP (pTP), and the DNA-
electrophoresis). Each hexon has two characteristic parts: a binding protein (DBP) are spliced from leftward-oriented
triangular top with three ‘towers’, and a pseudohexagonal promoters. Where it has been examined, splicing is also a
base with a central cavity. Hexons, or more exactly their common feature of genes in the nonconserved regions.
bases, are packed tightly to form a protein shell that Replication of various human adenoviruses has been
protects the inner components of the virion. studied in detail, in particular with HAdV-2. Virus entry
In members of the genus Mastadenovirus 12 copies of takes place via interactions of the fiber knob with specific
polypeptide IX are found between the nine hexons in the receptors on the surface of a susceptible cell followed by
center of each facet. However, polypeptide IX is not internalization via interactions between the penton base
Exploring the Variety of Random
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"Yes, I do," I whispered.
"I am going to inject some serum." He was busy tying a tourniquet of
rubber tubing above my elbow to bring out the veins. I felt the needle
probing for the collapsed tissues and later the pressure as he pulled it
out and stopped the bleeding. He jabbed me again in the biceps.
"You have had your anti-serum and a sedative," he said, leaning
close to be sure I heard. "Now you must relax and concentrate on
getting well."
With that thought in my mind I went to sleep.

Three days later I was over the worst of it. I had bled again from the
kidneys but fortunately the disease had not been severe enough to
cause a massive internal hemorrhage that would have choked their
filtering mechanism and killed me.
"How do I look?" I said to Anders that morning as he examined me
where I lay, in the dappled shade of the clearing.
"Your eyes are very red, of course," he smiled, "and you have
purpuric spots ... what your laymen call bruising, isn't it ... in the
creases of your elbows and thighs, but I think you have been
fortunate."
"I agree with that statement, Doctor," I said as I looked over at his
other patients, lying there so quietly beside me. A horse fly lit on
Kang's nose. Feebly his face twitched, trying to dislodge it. He lifted
his right hand, bending the arm from the elbow. It stayed there, too
weak to go farther. Anders shooed the fly. Kang's hand, poised
uncertainly for a time, slowly fell back to his side. To all appearances
he was lifeless.
"They're in bad shape, aren't they?" I asked.
"Yes, but they should recover. You saved their lives, you know."
"I did? How?"
"By using that cold water. When I checked them, their temperatures
were very low, especially Kang. You might say you had put them into
artificial hibernation. They were both in shock but, with the low body
temperatures reducing their metabolism during the crucial stage, I am
sure they have a much better chance of returning to normal. I
maintained their low temperatures with one of our new hypothermic
drugs for the first two days. Now they have returned to a more normal
state except that they are still asleep."
"They look more dead than asleep," I said and raised myself up to sit.
Even that was an effort as my swimming head and pounding heart
warned me. In a moment or two I felt better. I inched over to a tree
and used it as a back rest. Soaking in the friendly warmth of the sun
like a cat on a garden wall, I dozed off.
"Take this, Colonel." Anders' face was close to mine as he woke me
gently and held out a bowl of warm rice. The sparse light-colored
stubble on his unshaven chin stood out like the tattered wheat stalks
on a dustbowl farm. Gaunt with fatigue, bleary-eyed and scruffy
though he was, his red-rimmed eyes shone with a fierce
determination to pull us through and cheat his former masters of at
least three victims. I ate and watched as he gently spooned a thin
paste of rice into the cracked and crusted mouths of his patients. As it
touched their tongues, they swallowed automatically like patients
under anaesthesia, which, in a way, I suppose they were.
"Have you had any sleep at all?" I said, watching him.
"Not much. An hour here and there. I was afraid to sleep."
"Then why don't you sleep now while I watch. I can wake you easily if
you lie down here."
"Thank you. I will do that. I am very tired."
I let him sleep six hours. The sun was low over the ridges and Kim
and his men were preparing the evening meal when he awoke.
Renewed vigor showed in all his actions as he moved about lighting a
fire and preparing our rice gruel. This time I crawled over to help him
with the patients. As we dripped the thick rice soup into those
impassive faces and later washed the dry drum-tight skin stretched
over bare bones, I asked about our plans.
"Yesterday," he said, "Lieutenant Pak On took a small party down to
the coast, to Wongpo. They are to find Lee Sung and tell him that
there will be a delay until you are well enough to travel. They also
must try to get more food. It will be at least three days before they are
back. I have vaccinated all your men against the bleeding death and
we must wait until you are no longer infectious and I am sure they are
immune before we escape. We cannot risk spreading the disease in
the western democracies."
"Do you think it got spread when the tank cars were blown up?" I
asked.
"It is quite possible. The concentration of virus in that river must have
been very high. Unfortunately there are still villages down its course
and along the Imjin where people live, and they may get it. For that
reason we must move as soon as it is safe. If disease breaks out
near the coast we will never be able to get a boat to take us off."
The thought worried me. Suppose Lee Sung died? Only he could
make the contacts to get us away, I supposed, by small fishing boat
out to sea where a submarine, or perhaps a destroyer, could pick us
up outside the territorial limits. We had to avoid the coastal patrols too
and only Sung could help us there.
By the end of that week, May was two thirds gone and we were all
recovering slowly. Pak came back and the news was bad. Lee Sung
and Blackie had come with him. I met them as I strolled along the trail
and went back with them to our camp for a conference. Before it
began, Anders got out his syringe and inoculated Lee Sung.
"We can't afford to lose our only contact with freedom," he said.
"I appreciate your kindness, even if it is somewhat self-centered," Lee
replied, with a disarming smile.

"Let's have it," I said to Pak when all the officers were gathered in the
glade. Makstutis, too weak as yet to participate actively, was lying
quietly taking in all that was said.
"We got into Wongpo without too much trouble," he began. "We kept
away from any signs of people on the way. It wasn't too hard to locate
Lee Sung either when I walked into the little town; several people
knew him and I pretended the North Korean Army had business with
him. I found him down at the wharf where his boat is moored and he
took me aboard. I was alone of course," he said, as an afterthought.
"The men stayed back in the hills."
"Is that the boat you mentioned when we first met?" I interrupted to
ask Lee.
"Yes. Actually it belongs to the United States," Lee Sung said. "It is
fitted as a deep sea fishing or trading junk. It has souped up engines
that look ordinary and a false bottom where I hide guns or radio or
anything we need to smuggle into or out of North Korea. The boat is
registered in my name of course. I'm supposed to be a part-time
fisherman and local cargo carrier, as well as a merchant. I have a
small store in Wongpo. The Reds used to wink their eyes at my
activities because I smuggled things they wanted from Hong Kong or
South Korea."
"Sorry to interrupt," I said to Pak. "I wanted to get the background
straight."
"That's all right, sir," he replied. "Now, where was I?"
"You'd got to Lee's boat," I said.
"Oh yes. We had to get some food so that's the first thing we talked
about. Lee had bags of rice in his store so we went there and loaded
up a mule cart he borrowed. We were going to drive it as far as we
could to where our men could get the rice and pack the sacks on A-
frames back over the trail to the farmhouse. We thought it would be
less noticeable if we did it that night. In the meantime Lee went out to
get some vegetables and see if he could scrounge any meat. That's
when the trouble started." He turned to Lee Sung. "Maybe you'd
better tell the rest of it."
"I went to the house of a farmer, an old friend of mine, who lives on
the edge of the village, to bargain for some vegetables and perhaps a
pig," Lee said. "I was still there, drinking tea to conclude the
transaction, as is our custom, when a detachment of about fifty North
Korean soldiers in three trucks rolled along the coast road into the
village. I finished my business as rapidly as possible, and, with the
help of the farmers' sons, brought the food down to my store. Then I
walked out around the village seeking information. Lieutenant Pak
stayed with the supplies. I was afraid someone would have told the
detachment commander of the presence of another North Korean
officer but fortunately he was so busy and the people so frightened
that no one remembered Lieutenant Pak."
"That evening the commander called an open meeting in the village
and announced there was to be curfew for everyone beginning that
night. Anyone who disobeyed would be shot. He also announced that
nobody could enter or leave the village by land or sea and he has
seized the fishing vessels, including mine."
"Oh, my God!" I said. "Why did he do that?"
"Apparently a new epidemic of some sort has broken out at several
villages along the Imjin River."
I looked across at Anders and shook my head. He lifted his shoulders
in a shrug of resignation.

Lee Sung continued. "There is a great search being made for

American bandits of oriental descent masquerading as soldiers of the
Peoples Army who attacked the virus factory. You will be interested to
know that it was a large and heavily armed force of capitalist
reactionaries which was driven off with very heavy losses."
"Losses to whom?" Blackie asked with a grin.
"To the Americans of course. The virus factory was not damaged."
"At least that's the truth; we got the powerhouse," Pak laughed.
"The moment he made the announcement about American soldiers I
left the meeting quietly and went back to my store. Somebody was
going to wake up, perhaps soon. I told Pak and we decided to risk it
as the soldiers were still moving into the police barracks and getting
set up. We drove the mule cart quickly out the back end of the village
and got away without being challenged. In fact we didn't see anyone
at all until we met our own people."
"It looks like we've stirred up the whole country," I said. "They
certainly seem frightened."
"I'm not surprised," Anders broke in. "When I left the factory, besides
the virus cultures, my rucksack was full of bottles of vaccine against
the bleeding death and as much of the anti-serum as I could carry. I
expected we might need it. I destroyed all I could of what I had to
leave behind and the papers too. The Communists have very little
left."
"A fine piece of work, Doctor," I said. "You saved our lives and
deprived the Reds of their protection, all at the same time." I turned
back to Lee Sung. "I wonder how they figured we were Americans. All
our papers were in order. There wasn't a thing to show we weren't
native guerrillas, admittedly in the service of an unfriendly power.
Why not think we were from South Korea?"
"Possibly some of the men we left in the factory lived long enough to
talk," Blackie said, "put I doubt it very much. Of course seeing the
white officers would give them grounds for suspicion."
"Suspicion, yes, but not fact," I said.
"The radio has been talking mysterious explosions and guerrilla
warfare in Siberia and parts of China recently," Lee Sung reported.
"That's it, by Golly!" Blackie burst out. "The raids on the nerve gas
centers must have started. It could be somebody has been captured
and brainwashed."
"Could be," I said, "and if so, we'd better get home. If the Reds can
suppress news of how successful the raids are, they may still bluff the
democracies, with threats of nerve gas and CBR warfare, into giving
them more food and a good settlement of the war, but if we get home
with our story then they'll realize they are licked and maybe quit."
The following day we set out to do the last few miles to the
farmhouse. The Reds didn't have enough men to search the hills and
the wilder the area the safer we'd be. Our trouble would be to break
through the barrier at the coast. With one wounded man and two sick
ones on litters we were heavily loaded and could make only slow
time. I had all I could do to carry my own weight and when we got to
the house late that night I collapsed on a pile of straw and stayed
there for the whole of the next day.

CHAPTER 16
We stayed in the village for three weeks. Each day Makstutis and
Kang were a little better.
"We have to get out of here," I said to Anders one day in the last
week, after we had examined our patients. "The A-bomb carrier is
probably on its way right now."
"They can't march all the way to the coast," Anders said dubiously. "If
we must go, we shall have to carry them."
Blackie and Kim had been watching us with interest. Now Kim spoke
up.
"We've got some real husky boys in the unit, Doc. How about fixing
up seats on a couple of A-frames. Then we could chogi them up the
hills and they could maybe make it down the other side."
"It's a good idea, sir," Blackie agreed. "Those back trails are too
narrow for litters. We can change chogi bearers frequently."
"What about Yip Kee?" Kim said. "Can he travel the same way?"
I looked at Anders. "What do you say, Doctor? It's a month since his
leg was fractured. I think we could take a chance on it provided he is
carried all the way."
"I see no alternative," Anders agreed.
We borrowed the A-frames from the farmhouse and Pak wove basket
seats across the carrying prongs. With wider shoulder straps and
some padding our men could carry the patients quite well, changing
frequently. We assigned two bearers to each A-frame; it was all we
could spare. The first time we tried it, Makstutis, irrepressible as ever,
cracked, "This'll be the first time I ever went into action sitting on my
ass. I feel like a damn tanker." The name stuck; from then on they
were called the tank section.
In the first part of June, Lieutenant Pak and Lee Sung made a
reconnaissance and came back with an encouraging report. The furor
over the raid on the virus factory had died down. Work on the power
house had started but in a half-hearted fashion, either from a sense
of defeat or perhaps a shortage of supplies and workmen. The
bleeding death had hit hard along the Imjin and spread over the
watershed to the coastal villages. It continued to spread as the panic-
stricken natives, completely out of control after two terrible epidemics,
fled from the disease and disseminated it wherever they went. Most
of the enemy troops were being used to try to halt the crazy rush
away from the death zone but some of them had also become
infected, either by contact with refugees or perhaps in the age-old
fashion by consorting with prostitutes in the towns. The result was
disorganization and a very low morale.

The garrison at Wongpo, still kept at fifty men, was in good health as
they had commandeered plentiful food supplies and driven out or
killed most of the villagers who had not already died. They held the
harbor and the three boats tied up there. One was Lee Sung's, the
other two were much smaller fishing boats.
Counting Anders and the three convalescents, we were down to a
total strength of twenty-three. It would not be easy to capture the boat
unless we could catch the North Koreans by surprise, but we had to
try it. We set off over the wildest part of the country, avoiding all
villages or farmland that might still be inhabited. By the evening of the
third day we lay on a ridge overlooking Wongpo. Shortly after dark,
Lee Sung and Pak went down to see what the situation was. The day
had been warm but a cool breeze began blowing towards the sea as
the land cooled off. I fell asleep, lulled by the quiet murmur of the
distant breakers and the rustle of leaves in the steady wind.
"Doc, Doc, wake up! Wake up, Doc," the insistent whisperer was Kim.
"Yeah, what, what's that?" I struggled confusedly back to
consciousness. Obviously something was wrong the way Kim was
still shaking me. "OK Kim ... lay off ... I'm awake," I said crossly. I was
still fagged out and hated to come back to reality.
"Sorry sir, there's trouble. Lee Sung has been captured."
"Captured!" I echoed. "How do you know?"
"Pak just came back. He says Lee Sung left him hidden near his store
while he tried to sneak back on board the junk. He could see Lee go
aboard but he never came on deck again. A few minutes later a North
Korean soldier came off the boat and went to the police barracks.
Two officers came back with him and went aboard. Then Pak figured
he'd need help and high-tailed it up here."
"OK, get everybody up. We move out right now," I growled and
started to put my equipment on. "Where's Blackie?"
"Talking to Pak, Colonel," he said as he moved away.
While the men got ready, the officers gathered around me.
"What's the plan, sir?" Blackie said.
"I'm not absolutely sure," I said, "but I do know this, if we don't rescue
Lee Sung and his junk you might as well figure on walking back to the
States via Siberia." I thought over the plan of the village for a
moment. "The boat is lying alongside the jetty about five hundred
yards north of the police barracks. The houses there thin out along
the coast road. You, Blackie, take four men. Swing north from here
and come in at the jetty as quietly as you can. If it isn't well guarded
maybe you can get aboard before they suspect. Then wait for us.
Better take Pak with you ... he knows exactly where the boat is in this
darkness. Kim, you take five men and surround the police barracks
as well as you can. At least try to cover the way north to the boat.
Don't do a thing unless you hit trouble or Blackie gets into a fight." I
paused, there was some detail I wanted to be sure of. "Oh yes, locate
all power and telephone lines you can and cut them the moment
shooting starts. Then hold off the Commies as long as possible and
withdraw towards the junk. I'll take the tank section and Dr. Anders.
We'll go along with Kim as far as seems safe and then make for the
boat by way of the beach while he tackles the barracks. The
challenge is Pusan ... the answer, Tokyo. One thing more. I don't want
to leave without Lee Sung for two reasons. One, we owe our lives to
him, and two, without him we'll have one hell of a time running that
boat and contacting help. I'm betting he is still held on the junk but
there's no guarantee of that. I wouldn't be a damn bit surprised if the
Reds were holding him for bait to catch us, so watch yourselves
every minute. Is that clear?"

It was. Blackie and his boys moved off first, going north over a trail
that would gradually lead them down the slope and north of the
village. A few minutes later we followed. Makstutis and Kang were to
walk downhill although they were exhausted from the day's march,
but Yip Kee had to ride. There was not quite a half moon, enough to
see the trail but not enough to make us conspicuous. I looked down
to the village. There were no lights. Even in the police barracks there
was blackout, either in fear of guerrilla sniping or perhaps waiting for
us. In thirty minutes we were on level ground with the beach a quarter
of a mile away. There was little wind now and the waves must have
been small. I couldn't hear anything but my own breathing and the
scuffing of our feet. There were no dogs and I wondered if there ever
had been; dog is a tasty meal to some of these people. Tonight
certainly it was a blessing.
Kim came back to me quietly, a short strong silhouette against the low
moon, and stuck his mouth up close to my ear.
"We go straight from here, Doc. The Police barracks is on the coast
road dead ahead. You'd better cut north a bit before you get on the
beach. Watch it crossing that road. In this light you'll stand out like a
neon sign."
I nodded to him. He and his men moved away, shadows that merely
faded until I was not sure they were there at all. I waited a little longer,
then I took the lead with Anders next, followed by the three tanks, all
the patients now riding, and the three spare men acting as a
bodyguard. We walked in a wide arc, going north and finally swinging
down a narrow mud lane between thatched houses to come to the
coast road. In the moonlight I could see no sign of life, so, one at a
time, we skimmed across it as quietly as we could, dodged the fishing
net racks that cluttered the soft sand and got out on the tidal area of
the beach. I saw at once that I had made a mistake. Kim was right.
We were far too conspicuous out there on the hard sand. I led them
back close to the nets and we stumbled on, tripping over the rocks
and loose stones that thrust up through the sandy patches, tiring
ourselves out in that loose shifting footing.
I stopped, trying to breathe silently. Faintly ahead I saw the outlines of
the jetty, the masts of the three boats silhouetted above it.
"Let's get back beside the road," I whispered to Anders. "We have to
get on it soon anyway, to approach that pier where the boats are. The
chogi bearers are about all in, trying to carry the patients in this soft
sand."

We had just stumbled and crawled back over the rocks and debris to
the side of the road when a light machine gun chittered angrily to the
south. I heard the yelling of commands cut short by the quick blasting
crack of a hand grenade. For a moment the flame burned a pattern
on my retina so I couldn't see clearly. I thought there were figures
moving down that way but I wasn't sure. Ahead, where the masts of
the junks jousted at the stars, there was a flash of light as a door
opened. A man's harsh scream followed it as thunder follows lightning
and then there was a splash, shouts, and running feet on the planks
of the wharf. The firing at the police barracks was heavier now and I
could see rifle flashes that appeared to come from a second floor
window. One of our boys must have marked it down too. The rifle
cracked once more, followed instantly by a grenade explosion inside
the room. It lit up the outline of the window like a furnace door opened
in a dark cellar. Something fell out. After that there was a lull behind
us. Ahead, sporadic shooting rattled back and forth from boat to jetty
to shore, the flashes jerking about like fireflies playing tag.
I could make no sense out of that battle so I gave orders in a low
voice.
"Let's stay here right now. Tanks, dismount and cover the rear.
Anders, you and the three guards move north twenty-five yards and
cover both sides of the road. Stay hidden, halt everybody, and don't
forget to give the challenge before you shoot." I returned to Makstutis.
"I'll drop back south a bit," I said, "and outflank anybody you stop."
I walked away and hid behind poles supporting one end of a large
fishing net that was hung on the long racks to dry. The shadows
broke my silhouette but I could see well through the net.
Across the road the low thatched roofs of the houses formed an
almost unbroken bar of shadow against the faint light of the moon. I
had been looking at it for a long time. I stared at it once more and
thought I saw slight movements in the blackness. I looked away and
tried the old trick of not staring straight at where I wanted to see, to
give my night vision a better chance. There it was again!
"Halt!" Makstutis gave the order in Korean. There was neither
movement nor sound now.
"Pusan!" He hissed the word explosively. Still no answer!

The light from his grenade was an instant before the roar. Crouched
along the walls of the houses across from me was a group of men,
more than five, maybe ten, spread apart for safety. He had caught the
first two with the explosion, the grenade right between them. The
others opened up, firing generally north and across the road, hoping
to catch their assailant.
"Makstutis is smart," I was thinking. "I'd probably have used my gun
and given my position away first thing." I marked the approximate
area of the flashes and, from my knees, covered it with one swinging
burst and then dropped behind the poles. The answering fire went
over my head and now our tanks really let go, all six of the men
blasting at the black shadow. It was enough. There was no answer.
Except for an occasional moan and some dragging and scrabbling in
the dirt, I heard only the ringing in my ears. Five minutes later I
decided it was safe to go back to my men.
The battle south of us stuttered and chattered as the burp guns spat
at each other. Northward it was quiet, too much so. An hour went by.
Then I heard the challenge again.
"Halt! Pusan!"... "Tokyo!" came the answer. It was Kim.
"What the Hell's going on?" he asked.
"I don't know," I said. "I think Blackie has the bear by the tail and can't
let go. There hasn't been a sound for a long time."
"We've got the Reds bottled up in the barracks," Kim said cheerfully.
"All except a patrol of ten that got out and went north. I figured you
could hold them while we took care of the rest."
"We cleaned them up. They're lying over there across the road."
"That's real neat work, Doc." I imagined him smiling in the dim light.
"Now what do we do?"
"Damned if I know," I admitted. "We'd better try to find Blackie, I
guess."
We found him lying behind the heavy timbers of the jetty where it
joined the road. He was boiling over with anger and frustration.
"The so-and-so's went back down inside the boat when the fighting
started and I don't dare go after them. They've still got Lee Sung
there and threaten to kill him if we attack. I told them we'd show them
real torture if they hurt him and promised to let them go free if they
surrendered but I guess they're counting on being rescued."
"Any of your men ever do any sailing?" I asked.
"I have a couple who know how," said Kim. "What's in your mind?"
"We can't get to the engines of the junk and I'm afraid to stay here." I
turned to Kim. "Did you ever get those power and phone lines, by the
way?"
"Yeah, I got them; but they could call for help if they have a battery-
powered transmitter."
"That's what's worrying me," I said. "The only alternative I can think of
is to get on that junk and try to sail the damn thing away. Maybe,
when they find out they are at sea, the gooks will surrender. Kim, you
go back and hold the fort while we try to get on the boat."
"But sir," Blackie spoke up, "when we go we'll have to take all three
boats or the garrison might take after us."
"I forgot about that, Blackie," I said ruefully, and then, as the thought
struck me, "Say, maybe one of them has an auxiliary motor and we
could tow the whole lot out to sea. Is there anyone on the little
boats?"
"I don't think so. I watched for a while when the fight started and didn't
see anybody. But they're small," he concluded doubtfully. "I wouldn't
bet on any engines."
"Engines or no engines," I decided, "we take all three boats. Can you
get aboard them safely?"
"We'll have to rush the big one," Blackie replied. "There's a couple of
ports they can fire through that cover most of the wharf except out
towards the bow."
"Let's see if we can find a rowboat first, or make a raft from those fish
racks," I said. "Then you can row out to the end of the wharf with
three or four men and approach the junk head on. While you're about
it, check the fishing boats for engines but don't start them up if they
have any. Tie all three boats together and find something we can use
for paddles or oars too. We'll try to float away with the tide. It seems
to be going out now."
Down by the nets we found a long flat-bottomed rowboat that seemed
serviceable. It was a struggle to get it to the water but we managed
with the help of some choice swearing and rude remarks about
Korean fishermen and Marine operations in general. The long sweep
oars were stacked by the nets and, in a short time, Blackie and his
amateur crew splashed out into the darkness. Some time later he
was back with one man.
"We got aboard," he related proudly, "and found the two fishing boats
have small motors that might be enough to pull the big junk along for
a while. We've got them all tied together and I left three men on the
deck of Lee Sung's boat. They can make sure the gooks keep their
heads inside but we'll have to ferry everybody out to the small boats
first. We still can't risk crossing that wharf."
"OK," I agreed. "Start ferrying the tanks. I'll go and get Kim and his
gang."
There was no more firing around the barracks. Either the Reds were
waiting for daylight or perhaps for help. The moon had set and in the
blackness finally I found Kim and explained the situation. We sent off
all the men and together we sat and watched for a surprise sortie
from the building. It must have been about two in the morning when
Blackie sent back for us. We were the last to leave and, as I passed
the racks, I pulled off a fishing net.
"Give me a hand with this, Blackie, I want to take it along with me."
"What on earth for, Colonel?"
"We can't get below decks on the junk. Our food is low. We can try for
fish with this. What about water?"
"Everybody filled their canteens with water before starting.
"It's not enough," I complained, "but we can't wait now."
By this time half our men were on the deck of the junk. The sick men
stayed in the smaller craft in case of trouble. We filled the rowboat
with six of the strongest men and cut loose from shore. With the
ebbing tide to help, the rowboat crew pulled slowly away from the
wharf, aided by others paddling in the fishing craft. Our prisoners
made no noise and we could hear no sounds of pursuit. An hour later
we started the small boats' motors.
My first impulse had been to run for the open sea, beyond the
territorial limits of North Korea but I reconsidered. The Soviets, if they
were looking for us, wouldn't bother about the niceties of international
law. We were fair game until picked up. So we putted along the coast,
running towards the thirty-eighth parallel. Shortly before dawn we
sailed close in to the rugged shoreline and anchored. We loosed the
small boats and ran them in to shore behind a rocky headland.
Perhaps a reconnaissance plane would miss us in the shelter of the
cliffs. We would have to chance the wind and weather in our rather
insecure hideout.

Sitting on the pebbly beach beside Anders, I was wondering what to

do next when he broke the silence.
"I believe I have a solution, Doctor," he said in his precise manner, "if
you will give me permission to try."
"What can we lose?" I said.
Approaching Lee Sung's vessel from dead ahead, we climbed over
the bow. Anders leaned over the side and yelled in fluent Korean for
the senior officer of the Communist soldiers. After a short silence
there was a rough shout from the forward port.
"What do you want with me?"
Anders talked slowly and clearly. "I am Dr. Anders from the virus
factory. You know I escaped and that I cannot go back if I want to live.
Therefore, if this boat is found by your comrades I will kill you before I
die myself."
"You cannot touch us and you cannot sink the boat. Your threat does
not scare me." The Red officer did not attempt to conceal his scorn.
"I promise you that if you free Lee Sung we will set you ashore and let
you go unharmed."
"I do not trust traitors," yelled the Korean. "We will not surrender. You
will be caught soon by our patrols."
"That will do you no good," countered Anders. "Listen to me! I have
vials of the bleeding death with me. All of us, including your prisoner,
are protected against it. If you do not surrender now I will break the
vials and spread the disease through the ship. Even if you are
rescued you will still die."
We could hear the angry arguments below deck. All of them had seen
death from hemorrhagic disease in its new virulent form. It was a
horrible sight even to a physician, and, to the uneducated soldiers,
the thought of those purple mottled bodies with blood red eyes,
retching and vomiting their lives away, must have been terrifying. The
wrangling stopped and the senior officer called out.
"How do you plan to do this if we consent?"
"How many men have you?" Anders asked.
"We are twelve altogether."
"Then send up four men, including the other officer, unarmed. We will
put them ashore where you can see them. The second time four
more will go. The last time, you will come up and bring Lee Sung. If
he is in reasonable condition you too will go. Otherwise you die."
"It is agreed. We come now."
There was no further trouble. Lee Sung had been beaten in the usual
Korean fashion but he was so glad to be free he claimed he felt fine.
The North Koreans disappeared quickly along the beach as if afraid
we might shoot.
"We'd better get out of here right now," said Lee Sung. "They can
reach a good sized village north of here in an hour and give the
alarm."
"All right," I agreed. "You take over."
He led us below and, after shifting some cargo, opened up a small
space under the false deck, forward of the engine room. In it he had a
powerful radio transmitter, a case containing two heavy machine guns
with ammunition, and a few boxes of burp guns and grenades.
"I used to run guns to the guerrillas," he explained. "These may be
very useful."

We set up the machine guns on deck and I felt better. By now we

were running south at the full speed of the powerful engines, the two
small junks towing behind, still manned and helping with their own
engines. Sung had said we might need them when I suggested
sinking them before we hauled up our anchor. I steered the course
while he worked his radio, trying to raise his contacts and get help to
us. It took some time but finally he came on deck smiling.
"I got them," he said. "We rendezvous with a destroyer off the coast
tonight. It will escort us to Japan."
"What do we do in the meantime?"
"There is danger that the Communists picked up my signals and got a
bearing. If the coast patrol or the jets don't see us we will be OK."
"What about our own jets? Can't we get fighter cover?"
"Only as a last resort ... and it would probably be too late. The Air
Force has been warned to avoid all incidents and they do not wish to
fly close to the coast."

The sun was almost gone behind the hills of the steep Korean coast
when the Red jets found us. They came out of the sun, as
experienced fighters do, and the high whistle was already over us
before we saw them.
"Migs!" I yelled and ran to the stern. "Cut loose! Spread out and head
in to shore."
The little boats swung to starboard almost at once and wavered off
like water beetles trying to dodge a dragon fly. Lee Sung was at the
wheel again. He spun it sharply and the bow swung towards the
shore. There was shelter in a narrow cleft between a rocky pinnacle
and the cliffs of the mainland if we could reach it. We would have to
chance the depth of the water. By now the jets were around again
and peeling off for the attack. They were coming in low from the
northwest this time as we were getting some protection from the
shoreline. I watched them come, feeling helpless without a weapon,
ready to drop behind the mainmast when I saw the angle of flight. At
the stern, Makstutis was lying flat, his helmet back on his head and
his teeth bared as he squinted over the barrel of the heavy machine
gun into the bright light. Beside him Kang was feeding the belts.
Propped up against the side, Yip Kee braced his automatic rifle on
the wooden rail and waited calmly. I swivelled around. Blackie and
two others had the forward gun aimed and waiting. In the little junks,
dropping rapidly astern, I saw that Kim and his men were already
fighting. Their puny burp guns popped bravely at the two jets which,
ignoring them as too small, were concentrating all their attention on
us.
The leading jet grew larger, filling the sky with its round open face and
stubby wings. The tracers from Makstutis' gun floated lazily upwards
and then seemed to snap past, below the airplane, and wink out. Too
low, I thought, and dropped flat as the Mig hit out at us. The ship
heeled over, sliding like a runner for base as Sung clawed at the
wheel. The screaming roar of the jet and the impact of cannon shells
and bullets on wood went by me once, and then again as the second
Mig swept overhead. I looked back. Makstutis was unhurt but Kang
was rolling around clutching at his legs. I got up and ran to him. That
last minute swerve and the sight of the tracers coming up had been
enough to divert the pilot's aim. Perhaps the bumpy air currents of the
cliffs helped. I thanked God as I ran that there were none of the new
guided missile planes around. The shells had ploughed through the
stern rail and ripped up the port side of the deck, missing the machine
gun but catching Kang's legs.
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Virology Five Volume Set Volume 1 5 Third Edition 2008 B.W.J. Mahy

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Copyright ã 2008 Elsevier Inc. All rights reserved

Nepovirus, Canadian Crown Copyright 2008

British Library Cataloguing in Publication Data

Library of Congress Catalog Number: 200892260

For information on all Elsevier publications

PRINTED AND BOUND IN SLOVENIA

Brian W J Mahy MA PhD ScD DSc

Marc H V Van Regenmortel PhD

Dennis H Bamford, Ph.D. Robert A. Lamb, Ph.D., Sc.D.

Eric Hunter, B.Sc., Ph.D. Peter J Walker, B.Sc., Ph.D.

Structure of the Encyclopedia

Finding Specific Information

Taxonomic Groups of Viruses

Further Reading sections

Cross referencing between articles

See also: Lymphocytic Choriomeningitis Virus: General Features.

D Gallitelli M-A Grandbastien

M McChesney E S Mocarski, Jr.

F C Thomas Allnutt R Vainionpää

G J Thomas Jr. A M Vaira

L-F Wang R G Will

Antiviral Agents H J Field and R A Vere Hodge 142

Bunyaviruses: General Features R M Elliott 390

Fish Rhabdoviruses G Kurath and J Winton 221

Hordeivirus J N Bragg, H-S Lim and A O Jackson 459

Interfering RNAs K E Olson, K M Keene and C D Blair 148

Mumps Virus B K Rima and W P Duprex 356

Partitiviruses of Fungi S Tavantzis 63

Pseudorabies Virus T C Mettenleiter 341

Sigma Rhabdoviruses D Contamine and S Gaumer 576

Transcriptional Regulation in Bacteriophage R A Weisberg, D M Hinton and S Adhya 174

Taxonomic Index 497

Introduction respectively. The other two genera (Atadenovirus and Siade-

Table 1 The taxonomy of family Adenoviridaea

Genus/species Serotype Strain Genome

Genus/species Serotype Strain Genome

Unassigned Viruses in the Family

Lee Sung continued. "There is a great search being made for

Sitting on the pebbly beach beside Anders, I was wondering what to

We set up the machine guns on deck and I felt better. By now we

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