Filoviruses: A Compendium of 40 Years of Epidemiological, Clinical, and Laboratory Studies

410 Pages·7.271 MB·English

by  Jens H. Kuhn MD, ScD, MS (auth.)

W
Archives of Virology
Supplement 20
Edited by
Charles H. Calisher
With advice and contributions by
Aleksandr P. Agafonov, M. Javad Aman, Kevin Anderson,
Daniel G. Bausch, Sina Bavari, Yevgenii F. Belanov,
Matthias Borchert, Joel G. Breman, Alexander A. Bukreyev,
Kartik Chandran, Robert Colebunders,
Heinz Feldmann, Claude Fauquet,
Thomas W. Geisbert, Jean-Paul Gonzalez, Mark Gorwitz,
Barry S. Hewlett, David L. Heymann, Frank Hufert,
M. Sofi Ibrahim, Tetsuro Ikegami,
Peter B. Jahrling, Barbara Johnson, Karl M. Johnson,
Alla V. Kachko, Philip J. Kranzusch,
Elliot J. Lefkowitz, Eric M. Leroy, Loreen L. Lofts,
Shigeru Morikawa, Elke Mühlberger, Frederick A. Murphy,
Jean-Jacques Muyembe-Tamfum, Sergei V. Netyosov,
Gene G. Olinger, Jr., Sheli R. Radoshitzky, Yurii N. Rassadkin,
Yelena I. Ryabchikova, Jean-François Saluzzo,
Aleksandr N. Sergeyev, Aleksandr M. Shestopalov,
Werner Slenczka, Aleksandr V. Sorokin, Ute Ströher,
Ayato Takada, Vladimir A. Ternovoi,
Aleksandr S. Vladyko, Viktor Ye. Volchkov,
Guido van der Groen, Manfred Weidmann
Jens H. Kuhn
Filoviruses
A Compendium of 40 Years
of Epidemiological, Clinical,
and Laboratory Studies
SpringerWienNewYork
Jens H. Kuhn, MD, ScD, MS
New England Primate Research Center
Harvard Medical School
Southborough, MA, USA
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Foreword
Thefiloviruses(LakeVictoriamarburgvirusandthe Foreaseofretrievalbythereader,allreferencesare
ebolaviruses) are etiological agents of severe hem- cited with original titles in the original spelling,
orrhagic fevers with extraordinary high case- first author names, full journal titles, and issue
fatality rates for humans, chimpanzees, gorillas, numbers where possible. The text of this book
and probably other animals. Natural reservoirs and and the accompanying reference resource will be
true geographic distributions of these viruses are updatedinthefuture,andthuswillprovidearobust
unknown. Vaccines for the prevention of filovirus source of information for filovirologists, other vir-
infections and antiviral therapies for the treatment ologists and scientists in general, clinicians, stu-
of patients or of infected animals are not yet avail- dents, journalists, and biodefense professionals.
able. Lake Victoria marburgvirus and ebolavirus This review is a work in progress. It is ex-
research requires maximum-containment laboratory pected that it will be expanded, updated, and cor-
facilities, of which there are only a few worldwide. rected at regular intervals. To further improve
In the U.S., the filoviruses are classified as Select this review, the reader is strongly encouraged to
Agents and National Institute of Allergy and In- contact the author (fi[email protected]),
fectious Diseases (NIAID) Category A Priority editor ([email protected]), or Springer
Pathogens, and are handled under biosafety-level- (fi[email protected]) to point out errors or
4 containment to protect laboratory workers from to provide missing or ill-cited references, personal
infection and to protect the civilian population. thoughts, materials, or observations relevant to
Many institutions and experts consider filoviruses filovirology, any or all of which would be most
as potential threats to humans because they could welcome.
be used as biological weapons. In a work this large and inclusive, surely many
This review summarizes in detail the contempo- errors remain, for which we apologize. Any errors
rary knowledge of filoviruses and the diseases they in this book are the responsibility of the author and
cause. The entirety of the open literature on filo- editor. Co-editors and reviewers provided guidance.
viruses, covering all major scientific and clinical Some assessed particular chapters covering only
fields, is referenced and summarized in this text, their areas of expertise, whereas others evaluated
including most of the conference abstracts, book the entire manuscript. Occasionally, one reviewer’s
chapters, dissertations, government reports, patents, pointofviewcontradictedthatofanotherreviewer’s
theses, and journal publications in many languages. orthatoftheauthorandeditor.Theauthorandeditor
In total, almost 4,500 references ((cid:1)3,350 citations decided which suggestions to follow. Theviewsand
and (cid:1)1,150 subcitations, such as abstracts, reprints, opinions expressed in this book do not necessarily
or translations) are provided on a CD-ROM as an stateorreflectthoseoftheinstitutionsaffiliatedwith
accompanying reference source for the book – the author, editor, or any reviewer or co-editor.
roughly three times the number of references that Jens H. Kuhn
can be recovered using PubMed searches ((cid:1)1,450). Charles H. Calisher
1 Preface
This review originated from my own need for an gence of Lake Victoria marburgvirus is described.
extensive review of filovirus (Lake Victoria mar- However, these translations frequently appeared
burgvirus and ebolavirus) research during my par- years later than the original publications, or, even
ticipation in the vaccine-development program at worse, are internal translations of specialized insti-
the United States Army Medical Research Institute tutes and are not easily accessible to the scientific
of Infectious Diseases, Frederick, Maryland from community or the public. Copying of titles in lan-
April1998toApril1999.Iwassurprisedtodiscov- guages other than English with the typical ac-
eronlyspecializedarticles – acomprehensivework cumulation of typing errors, and the translation of
covering all aspects of filovirus research did not non-English journal titles into English often result-
exist. edinalmostindecipherablereferencing – totheex-
Lake Victoria marburgvirus was discovered in tentthatevenprofessionallibrarianswereunableto
1967 in Germany and Yugoslavia. At the time, it locate these articles. The bad habit of ‘‘correcting’’
wasnotunusualforresearcherstopublishscientific thosespelling errors considered to be ‘‘obvious’’ in
articles in their respective language followed by author lists or titles of journal articles has contrib-
articles in English. Thus, a large body of literature uted to the confusion.
in non-English languages accumulated, and, to a Thewidespreadinterestinfilovirusesbythepub-
lesser extent, still accumulates today. In fact, most lic and the scientific community, and the above-
of the important Soviet and Russian contributions mentioned flaws of published articles, stimulated
tofilovirusresearchwerepublishedinRussian,and the idea to write this review. Its purpose is to pro-
many of them have not been indexed or translated, vide an extensive bibliographyoffilovirus-research
let alone summarized in review articles. I came to publications with emphasis on the African, Dutch,
realize that most of the non-English publications French, German, Japanese, and Russian literature,
are not cited in the filovirus literature. One reason andtoreviewtheircontents.Throughthereferences,
for this might be that only a few dedicated re- credit is given to most researchers who, in the past
searchers would undertake the effort of obtaining four decades, contributed to today’s knowledge of
and translating these articles. An even more impor- filoviruses. The references are designed to provide
tant reason might be that many of the non-English accurate and detailed information to easily locate
articles are not indexed in the common computer- and retrieve the respective publications. Additional
ized databases such as PubMed or Ingenta. Many information regarding the articles is also provided,
researchers may not be fully aware of all publi- e.g. whether translations of articles are available,
cations, which might lead to redundant research. whetheranarticlehasbeenpublishedwithabstracts
Another major problem I have encountered is that in different languages, whether reprints of articles
many references are cited incorrectly or have ty- are available, and much more. The full names of
pographical errors. For example, translations of researchers are provided if they appeared in a cited
Germanarticlesareoftencitedwhenthefirstemer- publication; the full journal names and the issue
2 Preface
numbers of references are provided for easier Biotechnology ‘‘Vector’’ in Koltsovo, Novosibirsk
retrieval by the reader. Region, Russia (in 2001), allowed me to tap into
The collection and analysis of the filovirus re- theknowledgeofworld-renownedfilovirusexperts.
search-related documents cited in this review took Many of them reviewed this manuscript and con-
almost nine years. Often, articles could only be tributed material, personal thoughts, and organiza-
retrieved by contacting individual scientists direct- tional help. These experts are listed by name in
ly, or by visiting local libraries in many countries. alphabetical order at the beginning of the book. I
Laboratory rotations at the United States Army am eternally grateful for their intellectual input
Medical Research Institute of Infectious Diseases and their enthusiastic support. This review would
(1998–1999),at the U.S.Centers for DiseaseCon- not have been possible without their help and that
trol and Prevention in Atlanta, Georgia (in 2000), of many kind and incredibly patient librarians.
and at the State Research Center for Virology and Jens H. Kuhn, September 4, 2007
2 Contents
3 LIST OFABBREVIATIONS. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9
4 INTRODUCTION . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13
4.1 Viral hemorrhagic fevers . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19
4.2 Filovirus taxonomy, evolution, and phylogeny. . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . 23
4.3 Biosafety concerns in filovirus research . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 37
4.4 Biosecurity concerns in filovirus research. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 50
5 HISTORY OF FILOVIRAL DISEASE OUTBREAKS . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 59
5.1 Lake Victoria marburgvirus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 64
5.1.1 Germany and Yugoslavia, 1967, and Uganda, 1967 (speculative) . . . . . .. . . . . . 64
5.1.2 South Africa, 1973 (speculative). . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . 68
5.1.3 Rhodesia, 1975. . . . . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . 68
5.1.4 Kenya, 1980. . . . . . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . 70
5.1.5 Zimbabwe=South Africa, 1982 (speculative). . . . . . . . . . . . . . . . . . . . . . . . . . . 70
5.1.6 Togo, 1985 (speculative) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 70
5.1.7 Kenya, 1987. . . . . . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . 71
5.1.8 Laboratory accident(s), U.S.S.R., 1988 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 71
5.1.9 Kenya=Sweden, 1990 (speculative) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 71
5.1.10 Laboratory accident, U.S.S.R., 1990 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 72
5.1.11 Kenya=Germany, 1993 (speculative) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 72
5.1.12 Zaire, 1987–1996=Democratic Republic of the Congo, 1997–2000. . . . . .. . . . . 73
5.1.13 Angola, 2004–2005. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 75
5.1.14 Uganda, 2007. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 75
5.2 Zaire ebolavirus . . . . . . . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . . 75
5.2.1 Zaire, 1976 . . . . . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . 75
5.2.2 Zaire, 1977 . . . . . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . 78
5.2.3 Kenya, 1980 (speculative) . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . 79
5.2.4 Zaire, 1980–1985 (speculative). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 79
5.2.5 Five independent outbreaks, Gabon, 1994–1995. . . . . . . . . . . . . . . . . . . . . . . . 79
5.2.6 Zaire, 1995 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 80
5.2.7 Laboratory accident, Russia, 1996. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 82
5.2.8 Gabon, 1996. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 82
5.2.9 Gabon=South Africa, 1996–1997 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 82
5.2.10 EightindependentoutbreaksinGabonandCongo(Brazzaville),2001–2002 ..... 83
5.2.11 Congo (Brazzaville) and Gabon, 2002. . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . 85
5.2.12 Three independent outbreaks, Congo (Brazzaville), 2002–2003 . . . . . .. . . . . . . 85
5.2.13 Congo (Brazzaville), 2003–2004. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 85
4 Contents
5.2.14 Laboratory accident, U.S.A., 2004 (speculative) . . . . . . . . . . . . . . . . . . . . . . . . 86
5.2.15 Laboratory accident, Russia, 2004. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 86
5.2.16 Congo (Brazzaville), 2005 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 86
5.3 Sudan ebolavirus. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 86
5.3.1 Sudan, 1976. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 86
5.3.2 Laboratory accident, U.K., 1976. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 88
5.3.3 Sudan, 1979. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 88
5.3.4 Uganda, 2000–2001 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 88
5.3.5 Sudan, 2004. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 90
5.4 Reston ebolavirus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 90
5.4.1 The Philippines=U.S.A., 1989–1990 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 90
5.4.2 The Philippines=Italy, 1992 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 93
5.4.3 The Philippines=U.S.A., 1996 . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . 93
5.5 C^oote d’Ivoire ebolavirus. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 94
5.5.1 C^oote d’Ivoire, 1992 (speculative) and C^oote d’Ivoire=Switzerland 1994 . . . .. . . . . 94
5.5.2 Liberia and C^oote d’Ivoire, 1995 (speculative). . . . . . . . . . . . . .. . . . . . . . . . . . . 95
5.6 Speculations regarding the occurrence of additional filovirus infections . . . . . . . . . . . . . . 95
5.7 Conclusions . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 96
6 CLINICAL PRESENTATION OF FILOVIRAL DISEASE . . . . . . . . . . . . ... . . . . . . . . . . . . 99
6.1 Lake Victoria marburgvirus infections . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . 99
6.1.1 In humans . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 99
6.1.2 In nonhuman primates. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 104
6.1.3 In guinea pigs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 105
6.1.4 In other animals. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 105
6.2 Zaire ebolavirus infections . . . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . .106
6.2.1 In humans . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 106
6.2.2 In nonhuman primates. . . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . 110
6.2.3 In guinea pigs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 113
6.2.4 In mice . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 113
6.2.5 In other animals. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 115
6.3 Sudan ebolavirus infections. . . . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . .115
6.3.1 In humans . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 115
6.3.2 In nonhuman primates. . . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . 116
6.3.3 In guinea pigs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 116
6.3.4 In mice . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 117
6.4 Reston ebolavirus infections . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .117
6.5 C^oote d’Ivoire ebolavirus infections . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .118
6.6 Conclusions . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .118
7 FILOVIRAL DISEASE PATHOLOGY. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 121
7.1 Marburgvirus disease pathology . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .121
7.1.1 In humans . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 121
7.1.2 In nonhuman primates. . . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . 123
7.1.3 In guinea pigs, hamsters, and mice. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 124
7.2 Zaire ebolavirus disease pathology. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .128
7.2.1 In humans . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 128
7.2.2 In nonhuman primates. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 130