Simian Virology Simian Virology Alexander F. Voevodin, MD, PhD, DSc, FRCPath Professor of Virology Vir&Gen, Toronto, Canada Preston A. Marx, Jr., PhD Professor of Tropical Medicine Tulane National Primate Research Center Covington, Louisiana, USA A John Wiley & Sons, Ltd., Publication Edition first published 2009 C 2009 Alexander F. Voevodin and Preston A. Marx, Jr. Blackwell Publishing was acquired by John Wiley & Sons in February 2007. Blackwell’s publishing program has been merged with Wiley’s global Scientific, Technical, and Medical business to form Wiley-Blackwell. Editorial Office 2121 State Avenue, Ames, Iowa 50014-8300, USA For details of our global editorial offices, for customer services, and for information about how to apply for permission to reuse the copyright material in this book, please see our website at www.wiley.com/wiley-blackwell. 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No warranty may be created or extended by any promotional statements for this work. Neither the publisher nor the author shall be liable for any damages arising herefrom. 1 2009 To Boris A. Lapin, the Dean of the Medical Primatology Corps, with gratitude A.F.V. To Murray B. E. Gardner, an outstanding colleague, mentor, and friend. P.A.M. Contents Preface ix Acknowledgments xi Part I: Introduction to Primatology and Virology 1 Classification of Nonhuman Primates 3 2 Principles of Virology 39 Part II: Simian Viruses and Nonhuman Primate Models of Viral Infections Introduction to Part II 63 Section 1: RNA Viruses Section 1.1: Simian Retroviruses Introduction to Retroviruses 69 3 Lentiviruses in Their Natural Hosts 77 4 Lentivirus AIDS Models 119 5 Origins of Epidemic Forms HIV-1 and HIV-2 155 6 Betaretroviruses 163 7 Gammaretroviruses 183 8 Deltaretroviruses 193 9 Spumaviruses 217 Section 1.2: Other Simian RNA Viruses 10 Picornaviruses 237 11 Arteriviruses 247 Section 2: DNA Viruses Section 2.1: Simian Herpesviruses Introduction to Herpesviruses 259 12 Simplexviruses 267 vii viii Contents 13 Varicelloviruses 295 14 Cytomegaloviruses 309 15 Lymphocryptoviruses 323 16 Rhadinoviruses 347 Section 2.2: Other Simian DNA Viruses 17 Parvoviruses 371 18 Polyomaviruses 377 19 Papillomaviruses 389 20 Hepadnaviruses 397 21 Adenoviruses 407 Section 3: Miscellaneous RNA and DNA Viruses 22 Miscellaneous Viruses 419 Section 4: Experimental and Natural Infection of Nonhuman Primates with Nonsimian Viruses 23 Experimental Infection of Nonhuman Primates with Viruses of Medical Importance 433 24 Natural Infection of Nonhuman Primates with Nonsimian Viruses 469 Index 475 Preface Simian virology became a “centenarian” last year. It was grate this huge body of information into a single book. conceived in 1908 by Carl Landsteiner and Erwin Pop- This book is an attempt to fill this gap by providing an per who demonstrated that poliomyelitis was a viral dis- up-to-date compendium for the readership ranging from ease transmissible to monkeys. Their work laid a foun- experienced professionals interested in simian viruses dation for a spectacular victory in the fight against po- and NHP models of viral diseases to advanced under- liomyelitis. During the next 40 years progress in simian graduate students who may use the book as a stepping- virology was intermittent and generally slow. Never- stone to infectious disease research in NHPs. theless, some important achievements were made such To accommodate readers with different backgrounds as proving the viral etiology of yellow fever (Adrian the book is divided into two parts. Part I (Chapters 1 Stokes, 1927; see Chapter 23) and identification of and 2) presents a minimalist approach to primate tax- “herpes B” virus, a simian virus lethal for humans (Al- onomy and basic virology, a kind of primer for under- bert Sabin and Arthur Wright, 1934; see Chapter 12). graduate and graduate students. Scientific and medical The first wave of explosive growth of simian virology professionals who lack primatological and virological was sparked in the early 1950s by the development of background or need a brief refresher may also benefit vaccines against poliomyelitis. Evaluation of the effi- from reading Chapters 1 and 2. cacy of the vaccine, large-scale propagation of the vac- Part II (Chapters 3–24) is mainly devoted to specific cine strains, and safety testing would have been impos- viral families that contain simian viruses. It is assumed sible without the use of nonhuman primates (NHPs). As that a reader of these chapters has a core knowledge of a by-product of polio vaccine development, a number of primatology and virology, at least at the level of Part I. simian viruses were discovered and characterized in the Our aim in Part II is not simply the “cataloguing” of 1950s and 1960s. data relevant to the individual simian viruses, but rather The advent of AIDS in 1981 ushered in the mod- we want to present a structured and critically assessed ern era of simian virology. Human immunodeficiency “state-of-the art” knowledge in these fields formatted viruses type 1 (HIV-1) and type 2 (HIV-2) were dis- to be useful for both professionals and students. We covered in 1982 and 1986, respectively (see Chapters 3 do not pretend to be comprehensive in each and every and 5). Both HIVs originated from simian ancestors, the chapter of Part II. The specialists may find some omis- simian immunodeficiency viruses (SIVs). These discov- sions and we invite suggestions if the reader feels that eries focused simian virology mainly on AIDS-related the omissions were of central importance to the field. themes. Although other areas of simian virology moved Several chapters in Part II stand apart. Chapter 5 con- at a slower pace compared to AIDS-related research, tains a critical analysis of data and hypotheses relevant knowledge of simian viruses other than SIV advanced, to the origin of HIV/AIDS pandemic. This chapter may particularly during last 10 years. be of interest not only to professionals, but also to stu- There are more than 20,000 simian virology-related dents and even to lay readers. Chapter 23, in a very publications. However, surprisingly, during past 40 concise form, covers NHP models of human and ani- years there has been no attempt to structure and inte- mal diseases produced by the inoculation of NHPs with ix x Preface nonsimian viruses. Chapter 24 contains examples of nat- knowledge of simian viruses, far more than we expected. ural (nonexperimental) infections of NHPs with non- We hope that our book will spur new investigations to simian viruses, including human viruses. close these gaps. Writing this book was a challenging and enriching experience.