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SV40

sv40 virus, sv40 polio vaccine
SV40 is an abbreviation for Simian vacuolating virus 40 or Simian virus 40, a polyomavirus that is found in both monkeys and humans It was named for the effect it produced on infected green monkey cells, which developed an unusual number of vacuoles Like other polyomaviruses, SV40 is a DNA virus that has the potential to cause tumors in animals, but most often persists as a latent infection

The discovery of SV40 revealed that between 1955 and 1963 around 90% of children and 60% of adults in USA were inoculated with SV40-contaminated polio vaccines1

Contents

  • 1 Historical background
  • 2 Virology
  • 3 Multiplicity reactivation
  • 4 Transcription
  • 5 SV40 in animals
  • 6 Hypothesized role in human disease
    • 61 p53 damage and carcinogenicity
    • 62 Polio vaccine contamination
  • 7 See also
  • 8 References
  • 9 External links
    • 91 CDC FAQ
    • 92 NIH 1997 Conference on SV40
    • 93 Other

Historical backgroundedit

SV40 was first identified by Ben Sweet and Maurice Hilleman in 1960 when they found that between 10-30% of polio vaccines in the USA were contaminated with SV402 In 1962, Bernice Eddy described the SV40 oncogenic function inducing sarcoma and ependymomas in hamsters inoculated with monkeys cells infected with SV403 The complete viral genome was sequenced by Fiers and his team at the University of Ghent Belgium in 19784

Virologyedit

SV40 consists of an unenveloped icosahedral virion with a closed circular dsDNA genome5 of 52 kb6 The virion adheres to cell surface receptors of MHC class I by the virion glycoprotein VP1 Penetration into the cell is through a caveolin vesicle Inside the cell nucleus, the cellular RNA polymerase II acts to promote early gene expression This results in an mRNA that is spliced into two segments The small and large T antigens result from this The large T antigen has two functions: 5% goes to the plasma cell membrane and 95% returns to the nucleus Once in the nucleus the large T antigen binds three viral DNA sites, I, II and III Binding of sites I and II autoregulates early RNA synthesis Binding to site II takes place in each cell cycle Binding site I initiates DNA replication at the origin of replication Early transcription gives two spliced RNAs that are both 19s Late transcription gives both a longer 16s, which synthesizes the major viral capsid protein VP1; and the smaller 19s, which gives VP2 and VP3 through leaky scanning All of the proteins, besides the 5% of large T, return to the nucleus because assembly of the viral particle happens there Eventual release of the viral particles is cytolytic and results in cell deathcitation needed

Multiplicity reactivationedit

SV40 is capable of multiplicity reactivation MR78 MR is the process by which two or more virus genomes containing otherwise lethal damage interact within an infected cell to form a viable virus genome Yamamato and Shimojo observed MR when SV40 virions were irradiated with UV light and allowed to undergo multiple infection of host cells7 Hall studied MR when SV 40 virions were exposed to the DNA crosslinking agent 4, 5’, 8-trimethylpsoralen8 Under conditions in which only a single virus particle entered each host cell, approximately one DNA cross-link was lethal to the virus and could not be repaired In contrast, when multiple viral genomes infected a host cell, psoralen-induced DNA cross-links were repaired; that is, MR occurred Hall suggested that the virions with cross-linked DNA were repaired by recombinational repair8 Michod et al reviewed numerous examples of MR in different viruses and suggested that MR is a common form of sexual interaction that provides the advantage of recombinational repair of genome damages9

Transcriptionedit

The early promoter for SV40 contains three elements The TATA box is located approximately 20 base-pairs upstream from the transcriptional start site The 21 base-pair repeats contain six GC boxes and are the site that determines the direction of transcription Also, the 72 base-pair repeats are transcriptional enhancers When the SP1 protein interacts with the 21 bp repeats it binds either the first or the last three GC boxes Binding the first three initiates early expression and binding the last three initiates late expression The function of the 72 bp repeats is to enhance the amount of stable RNA and increase the rate of synthesis This is done by binding dimerization with the AP1 activator protein 1 to give a primary transcript that is 3' polyadenylated and 5' cappedcitation needed

SV40 in animalsedit

SV40 is dormant and is asymptomatic in rhesus monkeys The virus has been found in many macaque populations in the wild, where it rarely causes disease However, in monkeys that are immunodeficient—due to, for example, infection with Simian immunodeficiency virus—SV40 acts much like the human JC and BK polyomaviruses, producing kidney disease and sometimes a demyelinating disease similar to PML In other species, particularly hamsters, SV40 causes a variety of tumors, generally sarcomas In rats, the oncogenic SV40 large T-antigen was used to establish a brain tumor model for PNETs and medulloblastomas10

The molecular mechanisms by which the virus reproduces and alters cell function were previously unknown, and research into SV40 vastly increased biologists' understanding of gene expression and the regulation of cell growthcitation needed

Hypothesized role in human diseaseedit

The hypothesis that SV40 might cause cancer in humans has been a particularly controversial area of research11 Several methods have detected SV40 in a variety of human cancers, although how reliable these detection methods are, and whether SV40 has any role in causing these tumors, remains unclear12 As a result of these uncertainties, academic opinion remains divided, with some arguing that this hypothesis is not supported by the data13 and others arguing that some cancers may involve SV401415 The US National Cancer Institute announced in 2004 that although SV40 does cause cancer in some animal models, "substantial epidemiological evidence has accumulated to indicate that SV40 likely does not cause cancer in humans"16 This announcement was based on two studies1718 This 2004 announcement is in contrast to a 2002 study performed by The National Academy of Sciences Immunization Safety Review committee that stated, "The committee concludes that the biological evidence is moderate that SV40 exposure could lead to cancer in humans under natural conditions”19 However, Namika, Goodison,and Rosser found that the SV40 large t-antigen, in combination with mycoplasma, often a contaminate of vaccines, can cause prostate cells to turn cancerous Whether or not this is true for other human cells is debated20

p53 damage and carcinogenicityedit

SV40 is believed to suppress the transcriptional properties of the tumor-suppressing p53 in humans through the SV40 large T-antigen and SV40 small T-antigen p53 is responsible for initiating regulated cell death "apoptosis", or cell cycle arrest when a cell is damaged A mutated p53 gene may contribute to uncontrolled cellular proliferation, leading to a tumor

SV40 may act as a co-carcinogen with crocidolite asbestos to cause both Peritoneal and Pleural Mesothelioma2122

When SV40 infects nonpermissive cells, such as 3T3 mouse cells, the dsDNA of SV40 becomes covalently integrated In nonpermissive cells only early gene expression occurs and this leads to transformation, or oncogenesis The nonpermissive host needs the large T-antigen and the small t-antigen in order to function The small T-antigen interacts with and integrates with the cellular phosphatase pp2A This causes the cell to lose the ability to initiate transcriptioncitation needed

Polio vaccine contaminationedit

Soon after its discovery, SV40 was identified in the oral form of the polio vaccine produced between 1955 and 1961 by American Home Products dba Lederle This is believed to be due to two sources: 1 SV40 contamination of the original seed strain coded SOM; 2 contamination of the substrate—primary kidney cells from infected monkeys used to grow the vaccine virus during production Both the Sabin vaccine oral, live virus and the Salk vaccine injectable, killed virus were affected; the technique used to inactivate the polio virus in the Salk vaccine, by means of formaldehyde, did not reliably kill SV40 The contaminated vaccine continued to be distributed to the public through 19632324

It was difficult to detect small quantities of virus until the advent of PCR; since then, stored samples of vaccine made after 1962 have tested negative for SV40 In 1997, Herbert Ratner of Oak Park, Illinois, gave some vials of 1955 Salk vaccine to researcher Michele Carbone25 Ratner, the Health Commissioner of Oak Park at the time the Salk vaccine was introduced, had kept these vials of vaccine in a refrigerator for over forty years2627 Upon testing this vaccine, Carbone discovered that it contained not only the SV40 strain already known to have been in the Salk vaccine containing two 72-bp enhancers but also the same slow-growing SV40 strain currently found in some malignant tumors and lymphomas containing one 72-bp enhancers28 It is unknown how widespread the virus was among humans before the 1950s, though one study found that 12% of a sample of German medical students in 1952 had SV40 antibodies29

An analysis presented at the Vaccine Cell Substrate Conference in 200430 suggested that vaccines used in the former Soviet bloc countries, China, Japan, and Africa, could have been contaminated up to 1980, meaning that hundreds of millions more could have been exposed to the virus unknowingly

Population level studies show no evidence of any increase in cancer incidence as a result of exposure,31 though SV40 has been extensively studied32 A thirty-five year followup found no excess of the cancers putatively associated with SV4033

See alsoedit

  • Viruses portal
  • Mason-Pfizer monkey virus packaging signal
  • SV40 Cancer Foundation

Referencesedit

  1. ^ Shah, K; Nathanson, N January 1976 "Human exposure to SV40: Review and comment" American Journal of Epidemiology 103 1: 1–12 PMID 174424 
  2. ^ Sweet, B H; Hilleman, M R November 1960 "The vacuolating virus, SV 40" Proceedings of the Society for Experimental Biology and Medicine Society for Experimental Biology and Medicine 105 2: 420–427 PMID 13774265 doi:103181/00379727-105-26128 
  3. ^ Eddy, B E; Borman, G S; Grubbs, G E; Young, R D May 1962 "Identification of the oncogenic substance in rhesus monkey kidney cell culture as simian virus 40" Virology 17: 65–75 PMID 13889129 doi:101016/0042-68226290082-x 
  4. ^ Fiers, W; Contreras, R; Haegemann, G; Rogiers, R; Van De Voorde, A; Van Heuverswyn, H; Van Herreweghe, J; Volckaert, G; Ysebaert, M May 1978 "Complete nucleotide sequence of SV40 DNA" Nature 273 5658: 113–20 Bibcode:1978Natur273113F PMID 205802 doi:101038/273113a0 
  5. ^ Fanning, E; Zhao, K 2009 "SV40 DNA replication: from the A gene to a nanomachine" Virology 384 2: 352–359 PMC 2718763  PMID 19101707 doi:101016/jvirol200811038 
  6. ^ Sowd, GA; Fanning, E 2012 "A wolf in sheep's clothing: SV40 co-opts host genome maintenance proteins to replicate viral DNA" PLoS Pathogens 8 11: e1002994 PMC 3493471  PMID 23144614 doi:101371/journalppat1002994 
  7. ^ a b Yamamoto, Hiroshi; Shimojo, H August 1971 "Multiplicity reactivation of human adenovirus type 12 and simian virus 40 irradiated by ultraviolet light" Virology 45 2: 529–31 PMID 4328814 doi:101016/0042-68227190355-2 
  8. ^ a b c Hall, J D 1982 "Repair of psoralen-induced crosslinks in cells multiply infected with SV40" Molecular & General Genetics 188 1: 135–8 PMID 6294477 doi:101007/bf00333007 
  9. ^ Michod, Richard E; Bernstein, Harris; Nedelcu, Aurora M 2008 "Adaptive value of sex in microbial pathogens" Infection, Genetics and Evolution 8 3: 267–85 PMID 18295550 doi:101016/jmeegid200801002 
  10. ^ Eibl, R H; Kleihues, P; Jat, P S; Wiestler, O D 1994 "A model for primitive neuroectodermal tumors in transgenic neural transplants harboring the SV40 large T antigen" The American Journal of Pathology 144 3: 556–64 PMC 1887088  PMID 8129041 
  11. ^ Poulin, D L; Decaprio, J A 2006 "Is There a Role for SV40 in Human Cancer" Journal of Clinical Oncology 24 26: 4356–65 PMID 16963733 doi:101200/JCO2005037101 
  12. ^ Lowe, D B; Shearer, M H; Jumper, C A; Kennedy, R C 2007 "SV40 association with human malignancies and mechanisms of tumor immunity by large tumor antigen" Cellular and Molecular Life Sciences 64 7–8: 803–14 PMID 17260087 doi:101007/s00018-007-6414-6 
  13. ^ Shah, K V 2007 "SV40 and human cancer: A review of recent data" International Journal of Cancer Journal International Du Cancer 120 2: 215–23 PMID 17131333 doi:101002/ijc22425 
  14. ^ Moens, U; Van Ghelue, M; Johannessen, M 2007 "Oncogenic potentials of the human polyomavirus regulatory proteins" Cellular and molecular life sciences : CMLS 64 13: 1656–78 PMID 17483871 doi:101007/s00018-007-7020-3 
  15. ^ Barbanti-Brodano, G; Sabbioni, S; Martini, F; Negrini, M; Corallini, A; Tognon, M 2004 "Simian virus 40 infection in humans and association with human diseases: results and hypotheses" Virology 318 1: 1–9 PMID 15015494 doi:101016/jvirol200309004 
  16. ^ "Studies Find No Evidence That SV40 is Related to Human Cancer" National Cancer Institute, National Institutes of Health 23 August 2004 
  17. ^ Engels, E A; Chen, J; Hartge, P; Cerhan, J R; Davis, S; Severson, R K; Cozen, W; Viscidi, R P 2005 "Antibody Responses to Simian Virus 40 T Antigen: A Case-Control Study of Non-Hodgkin Lymphoma" Cancer Epidemiology Biomarkers & Prevention 14 2: 521–4 PMID 15734981 doi:101158/1055-9965epi-04-0441 
  18. ^ Engels, Eric A; Katki, Hormuzd A; Nielsen, Nete M; Winther, Jeanette F; Hjalgrim, Henrik; Gjerris, Flemming; Rosenberg, Philip S; Frisch, Morten 2003 "Cancer Incidence in Denmark Following Exposure to Poliovirus Vaccine Contaminated with Simian Virus 40" JNCI Journal of the National Cancer Institute 95 7: 532–9 PMID 12671021 doi:101093/jnci/957532 
  19. ^ Stratton, Kathleen; Almario, Donna A; McCormick, Marie C, eds 2002 "Immunization Safety Review: SV40 Contamination of Polio Vaccine and Cancer" Immunization Safety Review: SV40 Contamination of Polio Vaccine and Cancer The National Academy of Sciences pp 19–84 ISBN 978-0-309-08610-3 
  20. ^ Namiki, K; Goodison, S; Porvasnik, S; Allan, R W; Iczkowski, K A; Urbanek, C; Reyes, L; Sakamoto, N; Rosser, C J 1 September 2009 "Persistent exposure to Mycoplasma induces malignant transformation of human prostate cells" PLoS ONE 4 9: e6872 Bibcode:2009PLoSO46872N PMC 2730529  PMID 19721714 doi:101371/journalpone0006872 
  21. ^ Kroczynska, B; Cutrone, R; Bocchetta, M; Yang, H; Elmishad, A G; Vacek, P; Ramos-Nino, M; Mossman, B T; Pass, H I; Carbone, M 2006 "Crocidolite asbestos and SV40 are cocarcinogens in human mesothelial cells and in causing mesothelioma in hamsters" Proceedings of the National Academy of Sciences of the United States of America 103 38: 14128–33 Bibcode:2006PNAS10314128K PMC 1599923  PMID 16966607 doi:101073/pnas0604544103 
  22. ^ Pershouse, M A; Heivly, S; Girtsman, T 2006 "The role of SV40 in malignant mesothelioma and other human malignancies" Inhalation toxicology 18 12: 995–1000 PMID 16920674 doi:101080/08958370600835377 
  23. ^ CDC website http://wwwcdcgov/vaccinesafety/concerns/concerns-historyhtml Accessed Nov 1, 2016
  24. ^ Study Is Unsure on Tainted Polio Vaccine's Cancer Role Denise Grady NY Times Oct 23, 2002 http://wwwnytimescom/2002/10/23/us/study-is-unsure-on-tainted-polio-vaccine-s-cancer-rolehtml Accessed Nov 1, 2016
  25. ^ Ferber, Dan 2002 "Virology Monkey virus link to cancer grows stronger" Science 296 5570: 1012–1015 PMID 12004103 doi:101126/science29655701012 Archived from the original on May 10, 2002 
  26. ^ Baggott, Mary Tim, ed 2007 Nature, the Physician, and the Family: Selected Writings of Herbert Ratner, MD AuthorHouse pp xv–xvii ISBN 978-1-4184-7510-9 self-published source
  27. ^ Bookchin, Debbie; Schumacher, Jim 2004 The Virus and the Vaccine St Martin's Press pp 226–28 ISBN 0-312-27872-1 
  28. ^ Rizzo, Paola; Di Resta, Ilaria; Powers, Amy; Ratner, Herbert; Carbone, Michele 1999 "Unique Strains of SV40 in Commercial Poliovaccines from 1955 Not Readily Identifiable with Current Testing for SV40 Infection" Cancer Research 59 24: 6103–8 PMID 10626798 
  29. ^ Martini, F; Corallini, A; Balatti, V; Sabbioni, S; Pancaldi, C; Tognon, M 9 July 2007 "Simian virus 40 in humans" Infectious agents and cancer 2: 13 PMC 1941725  PMID 17620119 doi:101186/1750-9378-2-13 
  30. ^ Bookchin, Debbie 7 July 2004 "Vaccine scandal revives cancer fear" New Scientist 
  31. ^ NIH/National Cancer Institute 2004-08-25 "Studies Find No Evidence That Simian Virus 40 Is Related To Human Cancer" Science Daily 
  32. ^ Hilleman MR 1998 "Discovery of simian virus 40 SV40 and its relationship to poliomyelitis virus vaccines" Dev Biol Stand 94: 183–90 PMID 9776239 
  33. ^ Carroll-Pankhurst, C; Engels, EA; Strickler, HD; Goedert, JJ; Wagner, J; Mortimer EA Jr Nov 2001 "Thirty-five year mortality following receipt of SV40- contaminated polio vaccine during the neonatal period" Br J Cancer 85 9: 1295–7 PMC 2375249  PMID 11720463 doi:101054/bjoc20012065 

External linksedit

CDC FAQedit

  • Frequently Asked Questions about Cancer, Simian Virus 40 SV40, and Polio Vaccine, Science Coordination and Innovation, United States Centers for Disease Control

NIH 1997 Conference on SV40edit

  • Simian Virus 40 SV40: A Possible Human Polyomavirus Workshop Monday January 27, 1997, Morning Session, transcript of 1997 National Institutes of Health conference on SV40 in humans, part 1 of 3, United States Food and Drug Administration FDA
  • Simian Virus 40 SV40: A Possible Human Polyomavirus Workshop Monday January 27, 1997 Afternoon Session, transcript of 1997 National Institutes of Health conference on SV40 in humans part 2 of 3, United States Food and Drug Administration FDA
  • Simian Virus 40 SV40: A Possible Human Polyomavirus Workshop, Tuesday, 28 January 1997, transcript of 1997 National Institutes of Health conference on SV40 in humans part 3 of 3 , United States Food and Drug Administration FDA

Otheredit

  • Simian virus 40 at the US National Library of Medicine Medical Subject Headings MeSH
  • SV40 entry in the NCBI Taxonomy database
  • SV40 entry in the NCBI Genome database

sv40, sv40 cancer, sv40 genome, sv40 ori, sv40 polio vaccine, sv40 polya sequence, sv40 polyadenylation signal, sv40 promoter, sv40 t antigen, sv40 virus


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