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This ninetieth volume of the IARC Monographs on the Evaluation of Carcinogenic Risks to Humansconsiders human papillomaviruses (HPVs) which were evaluated by a previous Working Group (IARC, 1995). The monograph in the present volume incorporates new data that have become available during the past decade. HPVs represent the most common infectious agents that are transmitted sexually throughout the world; the major risk factors are behaviors associated with sexual activity. Although most infections are asymptomatic and are cleared within a period of 2 years, genital HPV infection can lead to clinical disease, including anogenital warts, cervical neoplasia, cervical cancer and other anogenital cancers. The risk for persistence of infection and progression of the more than 40 genital HPV types to grade 3 cervical intraepithelial neoplasia (CIN3) and cancer differs widely. Persistent infection with carcinogenic HPVs occurs in virtually all cases of cervical cancer. Previous evaluations of HPVs have classified types 16 and 18 as carcinogenic to humans (group 1), types 31 and 33 as probably carcinogenic to humans (Group 2A) and some types other than 16, 18, 31 and 33 as possibly carcinogenic to humans (Group 2B). At that time, the evaluation of types 16 and 18 was based on the strong association between infection with these HPVs and cervical cancer. For types 31 and 33, the association was less strong. The new epidemiological data reviewed in the present volume strongly support and further confirm the previous evaluation of types 16 and 18, and provide new evidence for other HPVs. This information, which includes strong evidence of carcinogenicity at sites other than the cervix, supports new evaluations for several other HPV types in addition to those mentioned above. Since the Working Group was convened in 2005, important innovations in HPV prophylaxis have occurred and these needed to be included in this volume. To date two prophylactic vaccines have been developed and used in large multicentric trials. This prophylactic vaccination is expected to reduce the incidence of HPV-related genital diseases. However, the benefits of prophylactic vaccines in a broad public health perspective will be achieved only if such vaccines can be provided to those groups of women for whom access to cervical cancer screening services is most problematic. Therefore, the development of second-generation vaccines that are expected to be cheaper, easier to deliver and to provide T-cell response against pre-existing HPV infections is highly desirable.
It has been more than twenty years since the isolation of polyoma virus and SV40, and the reports that they could produce tumors in animals and transformation of cells in culture. What was startling was that these biologic properties are associated with viruses that contain genetic in formation that is able to code for only five or six proteins. Since that time, investigations with these viruses have been in four principal areas. One major area of study has been on cells transformed by viruses that show altered growth properties and specify new viral and cellular proteins. Transformation studies have focused on the tumor (T) antigens that are specified by the virus and are required to initiate and to maintain the transformed state. Current studies on transformation are summarized in Chapter 4. The second broad area of investigation concerns replication of viruses during a lytic cycle of infection. T-antigens that are the hallmark of transformed cells are also expressed in cells that are lytically infected and are required for viral DNA replication and also function to alter rates of transcription of the early and late viral genes. Except for T-antigen, virus replication depends on the cellular enzymatic machinery and so the description of viral macromolecular synthesis has provided valuable insights into the cellular biosynthetic pathways. These studies are described in Chapters 1-3. The studies that have medical relevance concern JC and BK viruses and there is evidence of widespread exposure of human populations to these agents.
This book explains the ecology of viruses by examining their interactive dynamics with their hosting species (in this volume, in animals), including the types of transmission cycles that viruses have evolved encompassing principal and alternate hosts, vehicles and vectoring species. Examining virology from an organismal biology approach and focusing on the concept that viral infections represent areas of overlap in the ecologies of the involved species, Viral Ecology is essential for students and professionals who either may be non-virologists or virologists whose previous familiarity has been very specialized.
The past decade has witnessed an explosion of information on the molecular biology of insect viruses and a frenzy of activity in applying this information to medicine and agriculture. Genetically engineered baculoviruses are presently being tested for commercial use as pesticides, and the study of such viruses is also revealing remarkable insights into basic cellular processes such as apoptosis. This comprehensive volume provides readers with knowledge of basic and applied baculovirology so that current literature in the field can be appreciated.
In this volume, a distinguished international group of contributors present the latest molecular, organismal, and epidemiological research on arenaviruses. Their work will broaden both the clinician's and the researcher's knowledge of basic mechanisms of immunological tolerance, viral immunosuppression, the nature of protective immune responses to vaccination, and viral effects on cell functions.
The viruses of the family Rhabdoviridae have an exceedingly broad host range and are widely distributed throughout the animal and plant king doms. Animal rhabdoviruses infect and often cause disease in insects, fish, and mammals, including man. The prototype rhabdovirus, vesicular stomatitis virus !VSV), has been extensively studied and provides perhaps the best model system for studying negative-strand viruses. The popularity of VSV as a model system is to a considerable extent due to its relative simplicity and to its rapid growth, generally to high titer, in many cell types ranging from yeast to human. The nucleocapsids of these viruses also carry transcriptional and replicative functions that are expressed in cell-free systems. The first RNA-dependent RNA poly merase was described in VSV and its G protein provided an early model system for studying the synthesis, processing, and membrane insertion of mammalian glycoproteins. VSV is also highly cytopathogenic and has been studied quite extensively for its capacity to kill cells and to shut off cellular macromolecular synthesis. Even earlier, VSV was discovered to be highly susceptible to the action of interferons and has served ever since as a means for quantitating the activity of interferons. To my way of thinking, the spark that ignited the explosion of re search in this field was struck at the First International Colloquium on Rhabdoviruses, attended by 30 or so participants in Roscoff, France, in June 1972.
More than seven years have passed since the first monograph on viroids was published. At that time, the existence of viroids as a novel type of pathogen far smaller than viruses had been amply demonstrated and some of their unusual molecular properties had been elucidated, but the entry of molecular biology into viroid research was still in its infancy. Since that time, our knowledge of the molecular properties of viroids has increased exponentially and viroids have become even more fasci nating than was the case seven years ago. Today, aside from transfer RNA, viroids are probably the best known type of RNA-at least from a struc tural standpoint. Much less is known of the mechanisms of viroid func tion, such as the exact pathway and enzymology of viroid replication and the biochemistry of viroid pathogenesis. Recently, however, emphasis in viroid research has shifted from structural to functional themes and im portant beginnings have been made in the elucidation of viroid struc ture-function relationships. With the discovery of viroidlike RNAs within the capsids of certain plant viruses and the finding of surprising structural similarities between viroids and plant satellite RNAs, the conceptual gap between viroids and conventional viruses has significantly narrowed. Even beyond virology, connecting links with cellular RNAs have come to light and the long isolation of viroids land "viroidologists"J has come to an end.
What justifies the size of this compendium of reviews on the paramyxoviruses? As intracellular parasites that reproduce with almost complete indifference to nuclear activities, paramyxoviruses have not been providing insights about genes that regulate cellular activities and development, topics that account for much of the excitement in modem biology. For contributions of virus research to those topics, we must look to the retroviruses, which have the propensity to steal developmentally important genes and subvert them to malignant pur poses, and to the nuclear DNA viruses, whose gene expression depends heavily upon cellular transcription machinery, making them exceptionally useful tools for identifying and characterizing components of that machinery. From this perspective, it may appear that purely lytic viruses like the paramyxoviruses are sitting on the sidelines of contemporary biology. But there is plenty of action on the sidelines. Paramyxoviruses remain unconquered, devastating agents of disease. Human deaths attributable to paramyxoviruses worldwide, especially in children, are numbered in the mil lions annually. There are many pathogenic paramyxoviruses and too few effec tive vaccines, and those vaccines (against measles and mumps) are affordable only by relatively affluent nations. Moreover, the paramyxoviruses are intrin sically interesting organisms, presenting the challenge of understanding the self-replication of RNA and many other challenges peculiar to the structures and functions of their proteins, not only as individual entities, but also as they act in concert during virus reproduction and interact with vital functions of the cells they infect and often (but not always) destroy.
Providing both historical background and recent advances, this series reviews in-depth the biologic, molecular, immunologic, and patholic features of this facinating virus family. The current volume focuses on the avian and murine species which have generated novel insights into cancer, and the evolution of the retroviridae.