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Review Article

Human endogenous retrovirus-K (HML-2): a comprehensive review

, , &
Pages 715-738 | Received 03 Jan 2018, Accepted 02 Jul 2018, Published online: 14 Oct 2018

References

  • Aaronson SA, Todaro GJ, Scolnick EM. 1971. Induction of murine C-type viruses from clonal lines of virus-free BALB-3T3 cells. Science. 174(4005):157–159.
  • Ade C, Roy-Engel AM, Deininger PL. 2013. Alu elements: an intrinsic source of human genome instability. Curr Opin Virol. 3(6):639–645.
  • Agoni L, Guha C, Lenz J. 2013. Detection of human endogenous retrovirus k (HERV-K) transcripts in human prostate cancer cell lines. Front Oncol. 3:180.
  • Alfahad T, Nath A. 2013. Retroviruses and amyotrophic lateral sclerosis. Antiviral Res. 99(2):180–187.
  • Andersson ML, Lindeskog M, Medstrand P, Westley B, May F, Blomberg J. 1999. Diversity of human endogenous retrovirus class II-like sequences. J Gen Virol. 80(Pt 1):255–260.
  • Andersson ML, Medstrand P, Yin H, Blomberg J. 1996. Differential expression of human endogenous retroviral sequences similar to mouse mammary tumor virus in normal peripheral blood mononuclear cells. AIDS Res Hum Retroviruses. 12(9):833–840.
  • Argaw-Denboba A, Balestrieri E, Serafino A, Cipriani C, Bucci I, Sorrentino R, Sciamanna I, Gambacurta A, Sinibaldi-Vallebona P, Matteucci C. 2017. HERV-K activation is strictly required to sustain CD133 + melanoma cells with stemness features. J Exp Clin Canc Res. 36:20.
  • Ariza ME, Williams MV. 2011. A human endogenous retrovirus K dUTPase triggers a TH1, TH17 cytokine response: does it have a role in psoriasis? J Invest Dermatol. 131(12):2419–2427.
  • Armbruester V, Sauter M, Krautkraemer E, Meese E, Kleiman A, Best B, Roemer K, Mueller-Lantzsch N. 2002. A novel gene from the human endogenous retrovirus K expressed in transformed cells. Clin Cancer Res. 8(6):1800–1807.
  • Armbruester V, Sauter M, Roemer K, Best B, Hahn S, Nty A, Schmid A, Philipp S, Mueller A, Mueller-Lantzsch N. 2004. Np9 protein of human endogenous retrovirus K interacts with ligand of numb protein X. J Virol. 78(19):10310–10319.
  • Aydin H, Cook JD, Lee JE. 2014. Crystal structures of beta- and gammaretrovirus fusion proteins reveal a role for electrostatic stapling in viral entry. J Virol. 88(1):143–153.
  • Babaian A, Mager DL. 2016. Endogenous retroviral promoter exaptation in human cancer. Mob DNA. 7:24.
  • Bannert N, Kurth R. 2004. Retroelements and the human genome: new perspectives on an old relation. Proc Natl Acad Sci U S A. 101(Suppl 2):14572–14579.
  • Barazandeh M, Lambert SA, Albu M, Hughes TR. 2018. Comparison of ChIP-Seq data and a reference motif set for human KRAB C2H2 zinc finger proteins. G3 (Bethesda). 8(1):219–229.
  • Barbulescu M, Turner G, Seaman MI, Deinard AS, Kidd KK, Lenz J. 1999. Many human endogenous retrovirus K (HERV-K) proviruses are unique to humans. Curr Biol. 9(16):861–868.
  • Barmak K, Harhaj E, Grant C, Alefantis T, Wigdahl B. 2003. Human T cell leukemia virus type I-induced disease: pathways to cancer and neurodegeneration. Virology. 308(1):1–12.
  • Belshaw R, Dawson AL, Woolven-Allen J, Redding J, Burt A, Tristem M. 2005. Genomewide screening reveals high levels of insertional polymorphism in the human endogenous retrovirus family HERV-K(HML2): implications for present-day activity. J Virol. 79(19):12507–12514.
  • Bentvelzen P, Daams JH, Hageman P, Calafat J. 1970. Genetic transmission of viruses that incite mammary tumor in mice. Proc Natl Acad Sci U S A. 67(1):377.
  • Berkhout B, Jebbink M, Zsiros J. 1999. Identification of an active reverse transcriptase enzyme encoded by a human endogenous HERV-K retrovirus. J Virol. 73(3):2365–2375.
  • Best S, Le Tissier PR, Stoye JP. 1997. Endogenous retroviruses and the evolution of resistance to retroviral infection. Trends Microbiol. 5(8):313–318.
  • Bhardwaj N, Maldarelli F, Mellors J, Coffin JM. 2014. HIV-1 infection leads to increased transcription of human endogenous retrovirus HERV-K (HML-2) proviruses in vivo but not to increased virion production. J Virol. 88(19):11108–11120.
  • Bhardwaj N, Montesion M, Roy F, Coffin JM. 2015. Differential expression of HERV-K (HML-2) proviruses in cells and virions of the teratocarcinoma cell line Tera-1. Viruses. 7(3):939–968.
  • Bhat RK, Rudnick W, Antony JM, Maingat F, Ellestad KK, Wheatley BM, Tonjes RR, Power C. 2014. Human endogenous retrovirus-K(II) envelope induction protects neurons during HIV/AIDS. PLoS One. 9(7):e97984.
  • Blaise S, de Parseval N, Benit L, Heidmann T. 2003. Genomewide screening for fusogenic human endogenous retrovirus envelopes identifies syncytin 2, a gene conserved on primate evolution. Proc Natl Acad Sci U S A. 100(22):13013–13018.
  • Blikstad V, Benachenhou F, Sperber GO, Blomberg J. 2008. Evolution of human endogenous retroviral sequences: a conceptual account. Cell Mol Life Sci. 65(21):3348–3365.
  • Blissenbach M, Grewe B, Hoffmann B, Brandt S, Uberla K. 2010. Nuclear RNA export and packaging functions of HIV-1 Rev revisited. J Virol. 84(13):6598–6604.
  • Blomberg J, Benachenhou F, Blikstad V, Sperber G, Mayer J. 2009. Classification and nomenclature of endogenous retroviral sequences (ERVs): problems and recommendations. Gene. 448(2):115–123.
  • Blond JL, Lavillette D, Cheynet V, Bouton O, Oriol G, Chapel-Fernandes S, Mandrand B, Mallet F, Cosset FL. 2000. An envelope glycoprotein of the human endogenous retrovirus HERV-W is expressed in the human placenta and fuses cells expressing the type D mammalian retrovirus receptor. J Virol. 74(7):3321–3329.
  • Boller K, Janssen O, Schuldes H, Tonjes RR, Kurth R. 1997. Characterization of the antibody response specific for the human endogenous retrovirus HTDV/HERV-K. J Virol. 71(6):4581–4588.
  • Boller K, Konig H, Sauter M, Muellerlantzsch N, Lower R, Lower J, Kurth R. 1993. Evidence that Herv-K is the endogenous retrovirus sequence that codes for the human teratocarcinoma-derived retrovirus Htdv. Virology. 196(1):349–353.
  • Bowen LN, Tyagi R, Li W, Alfahad T, Smith B, Wright M, Singer EJ, Nath A. 2016. HIV-associated motor neuron disease: HERV-K activation and response to antiretroviral therapy. Neurology. 87(17):1756–1762.
  • Brady T, Lee YN, Ronen K, Malani N, Berry CC, Bieniasz PD, Bushman FD. 2009. Integration target site selection by a resurrected human endogenous retrovirus. Genes Dev. 23(5):633–642.
  • Bray S, Turnbull M, Hebert S, Douville RN. 2016. Insight into the ERVK integrase – propensity for DNA damage. Front Microbiol. 7:1941.
  • Brinzevich D, Young GR, Sebra R, Ayllon J, Maio SM, Deikus G, Chen BK, Fernandez-Sesma A, Simon V, Mulder LC. 2014. HIV-1 interacts with human endogenous retrovirus K (HML-2) envelopes derived from human primary lymphocytes. J Virol. 88(11):6213–6223.
  • Buscher K, Hahn S, Hofmann M, Trefzer U, Ozel M, Sterry W, Lower J, Lower R, Kurth R, Denner J. 2006. Expression of the human endogenous retrovirus-K transmembrane envelope, Rec and Np9 proteins in melanomas and melanoma cell lines. Melanoma Res. 16(3):223–234.
  • Buscher K, Trefzer U, Hofmann M, Sterry W, Kurth R, Denner J. 2005. Expression of human endogenous retrovirus K in melanomas and melanoma cell lines. Cancer Res. 65(10):4172–4180.
  • Buzdin A, Kovalskaya-Alexandrova E, Gogvadze E, Sverdlov E. 2006. At least 50% of human-specific HERV-K (HML-2) long terminal repeats serve in vivo as active promoters for host nonrepetitive DNA transcription. J Virol. 80(21):10752–10762.
  • Buzdin A, Ustyugova S, Khodosevich K, Mamedov I, Lebedev Y, Hunsmann G, Sverdlov E. 2003. Human-specific subfamilies of HERV-K (HML-2) long terminal repeats: three master genes were active simultaneously during branching of hominoid lineages. Genomics. 81(2):149–156.
  • Cai BH, Chen JY, Lu MH, Chang LT, Lin HC, Chang YM, Chao CF. 2009. Functional four-base A/T gap core sequence CATTAG of P53 response elements specifically bound tetrameric P53 differently than two-base A/T gap core sequence CATG bound both dimeric and tetrameric P53. Nucleic Acids Res. 37(6):1984–1990.
  • Callahan R, Chiu IM, Wong JF, Tronick SR, Roe BA, Aaronson SA, Schlom J. 1985. A new class of endogenous human retroviral genomes. Science. 228(4704):1208–1211.
  • Cegolon L, Salata C, Weiderpass E, Vineis P, Palu G, Mastrangelo G. 2013. Human endogenous retroviruses and cancer prevention: evidence and prospects. BMC Cancer. 13:4.
  • Chaffanet M, Popovici C, Leroux D, Jacrot M, Adelaide J, Dastugue N, Gregoire MJ, Hagemeijer A, Lafage-Pochitaloff M, Birnbaum D et al. 1998. t(6;8), t(8;9) and t(8;13) translocations associated with stem cell myeloproliferative disorders have close or identical breakpoints in chromosome region 8p11-12. Oncogene. 16(7):945–949.
  • Chen T, Meng Z, Gan Y, Wang X, Xu F, Gu Y, Xu X, Tang J, Zhou H, Zhang X et al. 2013. The viral oncogene Np9 acts as a critical molecular switch for co-activating beta-catenin, ERK, Akt and Notch1 and promoting the growth of human leukemia stem/progenitor cells. Leukemia. 27(7):1469–-1478. English.
  • Cohen M, Larsson E. 1988. Human endogenous retroviruses. Bioessays. 9(6):191–196.
  • Contreras-Galindo R, Almodovar-Camacho S, Gonzalez-Ramirez S, Lorenzo E, Yamamura Y. 2007. Comparative longitudinal studies of HERV-K and HIV-1 RNA titers in HIV-1-infected patients receiving successful versus unsuccessful highly active antiretroviral therapy. AIDS Res Hum Retroviruses. 23(9):1083–1086.
  • Contreras-Galindo R, Contreras-Galindo A, Lorenzo E, Yamamura Y. 2006. Evidence for replication of human endogenous retroviruses type-K (HERV-K) in HIV-1 positive patients. Retrovirology. 3:S33.
  • Contreras-Galindo R, Dube D, Fujinaga K, Kaplan MH, Markovitz DM. 2017. Susceptibility of human endogenous retrovirus type K to reverse transcriptase inhibitors. J Virol. 91(23):e01309-17.
  • Contreras-Galindo R, Kaplan MH, Contreras-Galindo AC, Gonzalez-Hernandez MJ, Ferlenghi I, Giusti F, Lorenzo E, Gitlin SD, Dosik MH, Yamamura Y et al. 2012. Characterization of human endogenous retroviral elements in the blood of HIV-1-infected individuals. J Virol. 86(1):262–276.
  • Contreras-Galindo R, Kaplan MH, Dube D, Gonzalez-Hernandez MJ, Chan S, Meng F, Dai M, Omenn GS, Gitlin SD, Markovitz DM. 2015. Human endogenous retrovirus type K (HERV-K) particles package and transmit HERV-K-related sequences. J Virol. 89(14):7187–7201.
  • Contreras-Galindo R, Lopez P, Velez R, Yamamura Y. 2007. HIV-1 infection increases the expression of human endogenous retroviruses type K (HERV-K) in vitro. AIDS Res Hum Retroviruses. 23(1):116–122.
  • Costas J. 2001. Evolutionary dynamics of the human endogenous retrovirus family HERV-K inferred from full-length proviral genomes. J Mol Evol. 53(3):237–243.
  • de Parseval N, Lazar V, Casella JF, Benit L, Heidmann T. 2003. Survey of human genes of retroviral origin: identification and transcriptome of the genes with coding capacity for complete envelope proteins. J Virol. 77(19):10414–10422.
  • Delelis O, Saib A, Sonigo P. 2003. Biphasic DNA synthesis in spumaviruses. J Virol. 77(14):8141–8146.
  • Denne M, Sauter M, Armbruester V, Licht JD, Roemer K, Mueller-Lantzsch N. 2007. Physical and functional interactions of human endogenous retrovirus proteins Np9 and rec with the promyelocytic leukemia zinc finger protein. J Virol. 81(11):5607–5616.
  • Denner J. 1998. Immunosuppression by retroviruses: implications for xenotransplantation. Ann N Y Acad Sci. 862:75–86.
  • Denner J. 2000. How does HIV induce AIDS? The virus protein hypothesis. J Human Virol. 3(2):81–82.
  • Dewannieux M, Blaise S, Heidmann T. 2005. Identification of a functional envelope protein from the HERV-K family of human endogenous retroviruses. J Virol. 79(24):15573–15577.
  • Dewannieux M, Harper F, Richaud A, Letzelter C, Ribet D, Pierron G, Heidmann T. 2006. Identification of an infectious progenitor for the multiple-copy HERV-K human endogenous retroelements. Genome Res. 16(12):1548–1556.
  • Dickerson F, Rubalcaba E, Viscidi R, Yang S, Stallings C, Sullens A, Origoni A, Leister F, Yolken R. 2008. Polymorphisms in human endogenous retrovirus K-18 and risk of type 2 diabetes in individuals with schizophrenia. Schizophr Res. 104(1–3):121–126.
  • Diem O, Schaffner M, Seifarth W, Leib-Mosch C. 2012. Influence of antipsychotic drugs on human endogenous retrovirus (HERV) transcription in brain cells. PLoS One. 7(1):e30054.
  • Domansky AN, Kopantzev EP, Snezhkov EV, Lebedev YB, Leib-Mosch C, Sverdlov ED. 2000. Solitary HERV-K LTRs possess bi-directional promoter activity and contain a negative regulatory element in the U5 region. FEBS Lett. 472(2–3):191–195.
  • Douville R, Liu J, Rothstein J, Nath A. 2011. Identification of active loci of a human endogenous retrovirus in neurons of patients with amyotrophic lateral sclerosis. Ann Neurol. 69(1):141–151.
  • Dube D, Contreras-Galindo R, He S, King SR, Gonzalez-Hernandez MJ, Gitlin SD, Kaplan MH, Markovitz DM. 2014. Genomic flexibility of human endogenous retrovirus type K. J Virol. 88(17):9673–9682.
  • Dunn CA, Romanish MT, Gutierrez LE, van de Lagemaat LN, Mager DL. 2006. Transcription of two human genes from a bidirectional endogenous retrovirus promoter. Gene. 366(2):335–342.
  • Ehlhardt S, Seifert M, Schneider J, Ojak A, Zang KD, Mehraein Y. 2006. Human endogenous retrovirus HERV-K(HML-2) Rec expression and transcriptional activities in normal and rheumatoid arthritis synovia. J Rheumatol. 33(1):16–23.
  • Esqueda D, Xu F, Moore Y, Yang Z, Huang G, Lennon PA, Hu PC, Dong J. 2013. Lack of correlation between HERV-K expression and HIV-1 viral load in plasma specimens. Ann Clin Lab Sci. 43(2):122–125.
  • Faulkner GJ, Carninci P. 2009. Altruistic functions for selfish DNA. Cell Cycle. 8(18):2895–2900.
  • Feuchter A, Mager D. 1990. Functional heterogeneity of a large family of human LTR-like promoters and enhancers. Nucleic Acids Res. 18(5):1261–1270.
  • Flockerzi A, Ruggieri A, Frank O, Sauter M, Maldener E, Kopper B, Wullich B, Seifarth W, Muller-Lantzsch N, Leib-Mosch C et al. 2008. Expression patterns of transcribed human endogenous retrovirus HERV-K(HML-2) loci in human tissues and the need for a HERV Transcriptome Project. BMC Genomics. 9:354.
  • Fuchs NV, Kraft M, Tondera C, Hanschmann KM, Lower J, Lower R. 2011. Expression of the human endogenous retrovirus (HERV) group HML-2/HERV-K does not depend on canonical promoter elements but is regulated by transcription factors Sp1 and Sp3. J Virol. 85(7):3436–3448.
  • Fuchs NV, Loewer S, Daley GQ, Izsvak Z, Lower J, Lower R. 2013. Human endogenous retrovirus K (HML-2) RNA and protein expression is a marker for human embryonic and induced pluripotent stem cells. Retrovirology. 10:115.
  • Garrison KE, Jones RB, Meiklejohn DA, Anwar N, Ndhlovu LC, Chapman JM, Erickson AL, Agrawal A, Spotts G, Hecht FM et al. 2007. T cell responses to human endogenous retroviruses in HIV-1 infection. PLoS Pathog. 3(11):e165.
  • Gautam P, Yu T, Loh YH. 2017. Regulation of ERVs in pluripotent stem cells and reprogramming. Curr Opin Genet Dev. 46:194–201.
  • George M, Schwecke T, Beimforde N, Hohn O, Chudak C, Zimmermann A, Kurth R, Naumann D, Bannert N. 2011. Identification of the protease cleavage sites in a reconstituted Gag polyprotein of an HERV-K(HML-2) element. Retrovirology. 8:30.
  • Gogvadze E, Stukacheva E, Buzdin A, Sverdlov E. 2009. Human-specific modulation of transcriptional activity provided by endogenous retroviral insertions. J Virol. 83(12):6098–6105.
  • Gonzalez-Cao M, Iduma P, Karachaliou N, Santarpia M, Blanco J, Rosell R. 2016. Human endogenous retroviruses and cancer. Cancer Biol Med. 13(4):483–488.
  • Gonzalez-Hernandez MJ, Swanson MD, Contreras-Galindo R, Cookinham S, King SR, Noel RJ, Kaplan MH, Markovitz DM. 2012. Expression of human endogenous retrovirus type K (HML-2) is activated by the TAT protein of HIV-1. J Virol. 86(15):7790–7805.
  • Grow EJ, Flynn RA, Chavez SL, Bayless NL, Wossidlo M, Wesche DJ, Martin L, Ware CB, Blish CA, Chang HY et al. 2015. Intrinsic retroviral reactivation in human preimplantation embryos and pluripotent cells. Nature. 522(7555):221–225.
  • Guasch G, Popovici C, Mugneret F, Chaffanet M, Pontarotti P, Birnbaum D, Pebusque MJ. 2003. Endogenous retroviral sequence is fused to FGFR1 kinase in the 8p12 stem-cell myeloproliferative disorder with t(8;19)(p12;q13.3). Blood. 101(1):286–288.
  • Hanke K, Chudak C, Kurth R, Bannert N. 2013. The Rec protein of HERV-K(HML-2) upregulates androgen receptor activity by binding to the human small glutamine-rich tetratricopeptide repeat protein (hSGT). Int J Cancer. 132(3):556–567.
  • Hanke K, Hohn O, Liedgens L, Fiddeke K, Wamara J, Kurth R, Bannert N. 2013. Staufen-1 interacts with the human endogenous retrovirus family HERV-K(HML-2) rec and gag proteins and increases virion production. J Virol. 87(20):11019–11030.
  • Hanke K, Kramer P, Seeher S, Beimforde N, Kurth R, Bannert N. 2009. Reconstitution of the ancestral glycoprotein of human endogenous retrovirus k and modulation of its functional activity by truncation of the cytoplasmic domain. J Virol. 83(24):12790–12800.
  • Harris JM, Haynes RH, McIntosh EM. 1997. A consensus sequence for a functional human endogenous retrovirus K (HERV-K) dUTPase. Biochem Cell Biol. 75(2):143–151.
  • Henzy JE, Coffin JM. 2013. Betaretroviral envelope subunits are noncovalently associated and restricted to the mammalian class. J Virol. 87(4):1937–1946.
  • Herbst H, Sauter M, Mueller-Lantzsch N. 1996. Expression of human endogenous retrovirus K elements in germ cell and trophoblastic tumors. Am J Pathol. 149(5):1727–1735.
  • Hohn O, Hanke K, Bannert N. 2013. HERV-K(HML-2), the best preserved family of HERVs: endogenization, expression, and implications in health and disease. Front Oncol. 3:246.
  • Holt MP, Shevach EM, Punkosdy GA. 2013. Endogenous mouse mammary tumor viruses (mtv): new roles for an old virus in cancer, infection, and immunity. Front Oncol. 3:287.
  • Hughes JF, Coffin JM. 2001. Evidence for genomic rearrangements mediated by human endogenous retroviruses during primate evolution. Nat Genet. 29(4):487–489.
  • Hughes JF, Coffin JM. 2004. Human endogenous retrovirus K solo-LTR formation and insertional polymorphisms: implications for human and viral evolution. Proc Natl Acad Sci U S A. 101(6):1668–1672.
  • Huh JW, Kim DS, Kang DW, Ha HS, Ahn K, Noh YN, Min DS, Chang KT, Kim HS. 2008. Transcriptional regulation of GSDML gene by antisense-oriented HERV-H LTR element. Arch Virol. 153(6):1201–1205.
  • Ikeda H, Odaka T. 1984. A cell membrane "gp70" associated with Fv-4 gene: immunological characterization, and tissue and strain distribution. Virology. 133(1):65–76.
  • Ishida T, Obata Y, Ohara N, Matsushita H, Sato S, Uenaka A, Saika T, Miyamura T, Chayama K, Nakamura Y et al. 2008. Identification of the HERV-K gag antigen in prostate cancer by SEREX using autologous patient serum and its immunogenicity. Cancer Immun. 8:15.
  • Jones RB, Garrison KE, Mujib S, Mihajlovic V, Aidarus N, Hunter DV, Martin E, John VM, Zhan W, Faruk NF et al. 2012. HERV-K-specific T cells eliminate diverse HIV-1/2 and SIV primary isolates. J Clin Invest. 122(12):4473–4489.
  • Kammerer U, Germeyer A, Stengel S, Kapp M, Denner J. 2011. Human endogenous retrovirus K (HERV-K) is expressed in villous and extravillous cytotrophoblast cells of the human placenta. J Reprod Immunol. 91(1–2):1–8. English.
  • Kapitonov VV, Jurka J. 1999. The long terminal repeat of an endogenous retrovirus induces alternative splicing and encodes an additional carboxy-terminal sequence in the human leptin receptor. J Mol Evol. 48(2):248–251.
  • Karamitros T, Paraskevis D, Hatzakis A, Psichogiou M, Elefsiniotis I, Hurst T, Geretti AM, Beloukas A, Frater J, Klenerman P et al. 2016. A contaminant-free assessment of Endogenous Retroviral RNA in human plasma. Sci Rep. 6:33598.
  • Katoh I, Mirova A, Kurata S, Murakami Y, Horikawa K, Nakakuki N, Sakai T, Hashimoto K, Maruyama A, Yonaga T et al. 2011. Activation of the long terminal repeat of human endogenous retrovirus K by melanoma-specific transcription factor MITF-M. Neoplasia. 13(11):1081–1092.
  • Kempf L, Nicodemus KK, Kolachana B, Vakkalanka R, Verchinski BA, Egan MF, Straub RE, Mattay VA, Callicott JH, Weinberger DR et al. 2008. Functional polymorphisms in PRODH are associated with risk and protection for schizophrenia and fronto-striatal structure and function. PLoS Genet. 4(11):e1000252.
  • Khodosevich K, Lebedev Y, Sverdlov ED. 2004. Large-scale determination of the methylation status of retrotransposons in different tissues using a methylation tags approach. Nucleic Acids Res. 32(3):e31.
  • Kidd JM, Cooper GM, Donahue WF, Hayden HS, Sampas N, Graves T, Hansen N, Teague B, Alkan C, Antonacci F et al. 2008. Mapping and sequencing of structural variation from eight human genomes. Nature. 453(7191):56–64.
  • Kitamura Y, Ayukawa T, Ishikawa T, Kanda T, Yoshiike K. 1996. Human endogenous retrovirus K10 encodes a functional integrase. J Virol. 70(5):3302–3306.
  • Knossl M, Lower R, Lower J. 1999. Expression of the human endogenous retrovirus HTDV/HERV-K is enhanced by cellular transcription factor YY1. J Virol. 73(2):1254–1261.
  • Koppe B, Menendez-Arias L, Oroszlan S. 1994. Expression and purification of the mouse mammary tumor virus gag-pro transframe protein p30 and characterization of its dUTPase activity. J Virol. 68(4):2313–2319.
  • Kovalskaya E, Buzdin A, Gogvadze E, Vinogradova T, Sverdlov E. 2006. Functional human endogenous retroviral LTR transcription start sites are located between the R and U5 regions. Virology. 346(2):373–378.
  • Kramer P, Lausch V, Volkwein A, Hanke K, Hohn O, Bannert N. 2016. The human endogenous retrovirus K(HML-2) has a broad envelope-mediated cellular tropism and is prone to inhibition at a post-entry, pre-integration step. Virology. 487:121–128.
  • Krishnamurthy J, Rabinovich BA, Mi T, Switzer KC, Olivares S, Maiti SN, Plummer JB, Singh H, Kumaresan PR, Huls HM et al. 2015. Genetic engineering of T cells to target HERV-K, an ancient retrovirus on melanoma. Clin Cancer Res. 21(14):3241–3251.
  • Krzysztalowska-Wawrzyniak M, Ostanek M, Clark J, Binczak-Kuleta A, Ostanek L, Kaczmarczyk M, Loniewska B, Wyrwicz LS, Brzosko M, Ciechanowicz A. 2011. The distribution of human endogenous retrovirus K-113 in health and autoimmune diseases in Poland. Rheumatology (Oxford). 50(7):1310–1314.
  • Kurth R, Bannert N. 2010. Beneficial and detrimental effects of human endogenous retroviruses. Int J Cancer. 126(2):306–314.
  • Laderoute MP, Giulivi A, Larocque L, Bellfoy D, Hou Y, Wu HX, Fowke K, Wu J, Diaz-Mitoma F. 2007. The replicative activity of human endogenous retrovirus K102 (HERV-K102) with HIV viremia. AIDS. 21(18):2417–2424.
  • Laderoute MP, Larocque LJ, Giulivi A, Diaz-Mitoma F. 2015. Further evidence that human endogenous retrovirus K102 is a replication competent foamy virus that may antagonize HIV-1 replication. Open AIDS J. 9:112–122.
  • Langner JS, Fuchs NV, Hoffmann J, Wittmann A, Brutschy B, Lower R, Suess B. 2012. Biochemical analysis of the complex between the tetrameric export adapter protein Rec of HERV-K/HML-2 and the responsive RNA element RcRE pck30. J Virol. 86(17):9079–9087.
  • Lee E, Iskow R, Yang L, Gokcumen O, Haseley P, Luquette LJ, 3rd, Lohr JG, Harris CC, Ding L, Wilson RK et al. 2012. Landscape of somatic retrotransposition in human cancers. Science. 337(6097):967–971.
  • Lee YN, Bieniasz PD. 2007. Reconstitution of an infectious human endogenous retrovirus. PLoS Pathog. 3(1):e10.
  • Lemaitre C, Harper F, Pierron G, Heidmann T, Dewannieux M. 2014. The HERV-K human endogenous retrovirus envelope protein antagonizes Tetherin antiviral activity. J Virol. 88(23):13626–13637.
  • Lemaitre C, Tsang J, Bireau C, Heidmann T, Dewannieux M. 2017. A human endogenous retrovirus-derived gene that can contribute to oncogenesis by activating the ERK pathway and inducing migration and invasion. PLoS Pathog. 13(6):e1006451.
  • Li M, Radvanyi LG, Yin B, Li J, Chivukula R, Lin K, Lu Y, Shen J, Chang DZ, Li D et al. 2017. Down-regulation of human endogenous retrovirus type K (HERV-K) viral env RNA in pancreatic cancer cells decreases cell proliferation and tumor growth. Clin Cancer Res. 23:5892–5911.
  • Li W, Lee MH, Henderson L, Tyagi R, Bachani M, Steiner J, Campanac E, Hoffman DA, von Geldern G, Johnson K et al. 2015. Human endogenous retrovirus-K contributes to motor neuron disease. Sci Transl Med. 7(307):307ra153.
  • Li XJ, Hanson C, Cmarik JL, Ruscetti S. 2009. Neurodegeneration induced by PVC-211 murine leukemia virus is associated with increased levels of vascular endothelial growth factor and macrophage inflammatory protein 1 alpha and is inhibited by blocking activation of microglia. J Virol. 83(10):4912–4922.
  • Liang X, Fu C, Cui W, Ober-Blobaum JL, Zahner SP, Shrikant PA, Clausen BE, Flavell RA, Mellman I, Jiang A. 2014. Beta-catenin mediates tumor-induced immunosuppression by inhibiting cross-priming of CD8(+) T cells. J Leukoc Biol. 95(1):179–190.
  • Linial ML. 1999. Foamy viruses are unconventional retroviruses. J Virol. 73(3):1747–1755.
  • Longo MC, Berninger MS, Hartley JL. 1990. Use of uracil DNA glycosylase to control carry-over contamination in polymerase chain-reactions. Gene. 93(1):125–128.
  • Lori F, Veronese FD, Devico AL, Lusso P, Reitz MS, Gallo RC. 1992. Viral-DNA carried by human-immunodeficiency-virus type-1 virions. J Virol. 66(8):5067–5074.
  • Lower R, Lower J, Frank H, Harzmann R, Kurth R. 1984. Human teratocarcinomas cultured in vitro produce unique retrovirus-like viruses. J Gen Virol. 65(Pt 5):887–898.
  • Lower R, Lower J, Tondera-Koch C, Kurth R. 1993. A general method for the identification of transcribed retrovirus sequences (R-U5 PCR) reveals the expression of the human endogenous retrovirus loci HERV-H and HERV-K in teratocarcinoma cells. Virology. 192(2):501–511.
  • Lower R, Tonjes RR, Korbmacher C, Kurth R, Lower J. 1995. Identification of a Rev-related protein by analysis of spliced transcripts of the human endogenous retroviruses HTDV/HERV-K. J Virol. 69(1):141–149.
  • Macfarlane C, Simmonds P. 2004. Allelic variation of HERV-K(HML-2) endogenous retroviral elements in human populations. J Mol Evol. 59(5):642–656.
  • Magin C, Lower R, Lower J. 1999. cORF and RcRE, the Rev/Rex and RRE/RxRE homologues of the human endogenous retrovirus family HTDV/HERV-K. J Virol. 73(11):9496–9507.
  • Magin-Lachmann C, Hahn S, Strobel H, Held U, Lower J, Lower R. 2001. Rec (formerly corf) function requires interaction with a complex, folded RNA structure within its responsive element rather than binding to a discrete specific binding site. J Virol. 75(21):10359–10371.
  • Manghera M, Douville RN. 2013. Endogenous retrovirus-K promoter: a landing strip for inflammatory transcription factors? Retrovirology. 10:16.
  • Manghera M, Ferguson J, Douville R. 2015. ERVK polyprotein processing and reverse transcriptase expression in human cell line models of neurological disease. Viruses. 7(1):320–332.
  • Manghera M, Ferguson-Parry J, Lin R, Douville RN. 2016. NF-kappaB and IRF1 induce endogenous retrovirus k expression via interferon-stimulated response elements in its 5' long terminal repeat. J Virol. 90(20):9338–9349.
  • Marchi E, Kanapin A, Magiorkinis G, Belshaw R. 2014. Unfixed endogenous retroviral insertions in the human population. J Virol. 88(17):9529–9537.
  • Marguerat S, Wang WY, Todd JA, Conrad B. 2004. Association of human endogenous retrovirus K-18 polymorphisms with type 1 diabetes. Diabetes. 53(3):852–854.
  • Martin MA, Bryan T, Rasheed S, Khan AS. 1981. Identification and cloning of endogenous retroviral sequences present in human DNA. Proc Natl Acad Sci U S A. 78(8):4892–4896.
  • Mayer J, Ehlhardt S, Seifert M, Sauter M, Muller-Lantzsch N, Mehraein Y, Zang KD, Meese E. 2004. Human endogenous retrovirus HERV-K(HML-2) proviruses with Rec protein coding capacity and transcriptional activity. Virology. 322(1):190–198.
  • Mayer J, Meese EU. 2003. Presence of dUTPase in the various human endogenous retrovirus K (HERV-K) families. J Mol Evol. 57(6):642–649.
  • Mayer J, Sauter M, Racz A, Scherer D, Mueller-Lantzsch N, Meese E. 1999. An almost-intact human endogenous retrovirus K on human chromosome 7. Nat Genet. 21(3):257–258.
  • McClintock B. 1950. The origin and behavior of mutable loci in maize. Proc Natl Acad Sci U S A. 36(6):344–355.
  • McCormick AL, Brown RH, Jr., Cudkowicz ME, Al-Chalabi A, Garson JA. 2008. Quantification of reverse transcriptase in ALS and elimination of a novel retroviral candidate. Neurology. 70(4):278–283.
  • McIntosh EM, Haynes RH. 1996. HIV and human endogenous retroviruses: an hypothesis with therapeutic implications. Acta Biochim Pol. 43(4):583–592.
  • Medstrand P, Blomberg J. 1993. Characterization of novel reverse transcriptase encoding human endogenous retroviral sequences similar to type A and type B retroviruses: differential transcription in normal human tissues. J Virol. 67(11):6778–6787.
  • Medstrand P, Mager DL. 1998. Human-specific integrations of the HERV-K endogenous retrovirus family. J Virol. 72(12):9782–9787.
  • Medstrand P, Mager DL, Yin H, Dietrich U, Blomberg J. 1997. Structure and genomic organization of a novel human endogenous retrovirus family: HERV-K (HML-6). J Gen Virol. 78(Pt 7):1731–1744.
  • Michaud HA, de Mulder M, SenGupta D, Deeks SG, Martin JN, Pilcher CD, Hecht FM, Sacha JB, Nixon DF. 2014. Trans-activation, post-transcriptional maturation, and induction of antibodies to HERV-K (HML-2) envelope transmembrane protein in HIV-1 infection. Retrovirology. 11:10.
  • Moelling K, Broecker F. 2015. The reverse transcriptase-RNase H: from viruses to antiviral defense. Ann N Y Acad Sci. 1341:126–135.
  • Monde K, Contreras-Galindo R, Kaplan MH, Markovitz DM, Ono A. 2012. Human endogenous retrovirus K Gag coassembles with HIV-1 Gag and reduces the release efficiency and infectivity of HIV-1. J Virol. 86(20):11194–11208.
  • Monde K, Terasawa H, Nakano Y, Soheilian F, Nagashima K, Maeda Y, Ono A. 2017. Molecular mechanisms by which HERV-K Gag interferes with HIV-1 Gag assembly and particle infectivity. Retrovirology. 14(1):27.
  • Morozov VA, Dao Thi VL, Denner J. 2013. The transmembrane protein of the human endogenous retrovirus–K (HERV-K) modulates cytokine release and gene expression. PLoS One. 8(8):e70399.
  • Moyes D, Griffiths DJ, Venables PJ. 2007. Insertional polymorphisms: a new lease of life for endogenous retroviruses in human disease. Trends Genet. 23(7):326–333.
  • Mueller-Lantzsch N, Sauter M, Weiskircher A, Kramer K, Best B, Buck M, Grasser F. 1993. Human endogenous retroviral element K10 (HERV-K10) encodes a full-length gag homologous 73-kDa protein and a functional protease. AIDS Res Hum Retroviruses. 9(4):343–350.
  • Muradrasoli S, Forsman A, Hu L, Blikstad V, Blomberg J. 2006. Development of real-time PCRs for detection and quantitation of human MMTV-like (HML) sequences HML expression in human tissues. J Virol Methods. 136(1–2):83–92.
  • Muster T, Waltenberger A, Grassauer A, Hirschl S, Caucig P, Romirer I, Fodinger D, Seppele H, Schanab O, Magin-Lachmann C et al. 2003. An endogenous retrovirus derived from human melanoma cells. Cancer Res. 63(24):8735–8741.
  • Ono M. 1986. Molecular cloning and long terminal repeat sequences of human endogenous retrovirus genes related to types A and B retrovirus genes. J Virol. 58(3):937–944.
  • Ono M, Yasunaga T, Miyata T, Ushikubo H. 1986. Nucleotide sequence of human endogenous retrovirus genome related to the mouse mammary tumor virus genome. J Virol. 60(2):589–598.
  • Oricchio E, Sciamanna I, Beraldi R, Tolstonog GV, Schumann GG, Spadafora C. 2007. Distinct roles for LINE-1 and HERV-K retroelements in cell proliferation, differentiation and tumor progression. Oncogene. 26(29):4226–4233.
  • Ormsby CE, Sengupta D, Tandon R, Deeks SG, Martin JN, Jones RB, Ostrowski MA, Garrison KE, Vazquez-Perez JA, Reyes-Teran G et al. 2012. Human endogenous retrovirus expression is inversely associated with chronic immune activation in HIV-1 infection. PLoS One. 7(8):e41021.
  • Otowa T, Tochigi M, Rogers M, Umekage T, Kato N, Sasaki T. 2006. Insertional polymorphism of endogenous retrovirus HERV-K115 in schizophrenia. Neurosci Lett. 408(3):226–229.
  • Pani MA, Wood JP, Bieda K, Toenjes RR, Usadel KH, Badenhoop K. 2002. The variable endogenous retroviral insertion in the human complement C4 gene: a transmission study in type I diabetes mellitus. Hum Immunol. 63(6):481–484.
  • Perot P, Mugnier N, Montgiraud C, Gimenez J, Jaillard M, Bonnaud B, Mallet F. 2012. Microarray-based sketches of the HERV transcriptome landscape. PLoS One. 7(6):e40194.
  • Ponferrada VG, Mauck BS, Wooley DP. 2003. The envelope glycoprotein of human endogenous retrovirus HERV-W induces cellular resistance to spleen necrosis virus. Arch Virol. 148(4):659–675.
  • Reis BS, Jungbluth AA, Frosina D, Holz M, Ritter E, Nakayama E, Ishida T, Obata Y, Carver B, Scher H et al. 2013. Prostate cancer progression correlates with increased humoral immune response to a human endogenous retrovirus GAG protein. Clin Cancer Res. 19(22):6112–6125.
  • Rethwilm A. 2003. The replication strategy of foamy viruses. Curr Top Microbiol Immunol. 277:1–26.
  • Reus K, Mayer J, Sauter M, Scherer D, Muller-Lantzsch N, Meese E. 2001. Genomic organization of the human endogenous retrovirus HERV-K(HML-2.HOM) (ERVK6) on chromosome 7. Genomics. 72(3):314–320.
  • Reus K, Mayer J, Sauter M, Zischler H, Muller-Lantzsch N, Meese E. 2001. HERV-K(OLD): ancestor sequences of the human endogenous retrovirus family HERV-K(HML-2). J Virol. 75(19):8917–8926.
  • Robinson D, Van Allen EM, Wu YM, Schultz N, Lonigro RJ, Mosquera JM, Montgomery B, Taplin ME, Pritchard CC, Attard G et al. 2015. Integrative clinical genomics of advanced prostate cancer. Cell. 161(5):1215–1228.
  • Robinson LR, Whelan SP. 2016. Infectious entry pathway mediated by the human endogenous retrovirus K envelope protein. J Virol. 90(7):3640–3649.
  • Romano CM, Ramalho RF, Zanotto PM. 2006. Tempo and mode of ERV-K evolution in human and chimpanzee genomes. Arch Virol. 151(11):2215–2228.
  • Ruggieri A, Maldener E, Sauter M, Mueller-Lantzsch N, Meese E, Fackler OT, Mayer J. 2009. Human endogenous retrovirus HERV-K(HML-2) encodes a stable signal peptide with biological properties distinct from Rec. Retrovirology. 6:17.
  • Schmitt K, Heyne K, Roemer K, Meese E, Mayer J. 2015. HERV-K(HML-2) rec and np9 transcripts not restricted to disease but present in many normal human tissues. Mobile DNA-Uk. 6:4.
  • Schmitt K, Reichrath J, Roesch A, Meese E, Mayer J. 2013. Transcriptional profiling of human endogenous retrovirus group HERV-K(HML-2) loci in melanoma. Genome Biol Evol. 5(2):307–328.
  • Schommer S, Sauter M, Krausslich HG, Best B, Mueller-Lantzsch N. 1996. Characterization of the human endogenous retrovirus K proteinase. J Gen Virol. 77(Pt 2):375–379.
  • Schulte AM, Lai S, Kurtz A, Czubayko F, Riegel AT, Wellstein A. 1996. Human trophoblast and choriocarcinoma expression of the growth factor pleiotrophin attributable to germ-line insertion of an endogenous retrovirus. Proc Natl Acad Sci U S A. 93(25):14759–14764.
  • Seifarth W, Frank O, Zeilfelder U, Spiess B, Greenwood AD, Hehlmann R, Leib-Mosch C. 2005. Comprehensive analysis of human endogenous retrovirus transcriptional activity in human tissues with a retrovirus-specific microarray. J Virol. 79(1):341–352.
  • SenGupta D, Tandon R, Vieira RG, Ndhlovu LC, Lown-Hecht R, Ormsby CE, Loh L, Jones RB, Garrison KE, Martin JN et al. 2011. Strong human endogenous retrovirus-specific T cell responses are associated with control of HIV-1 in chronic infection. J Virol. 85(14):6977–6985.
  • Steele AJ, Al-Chalabi A, Ferrante K, Cudkowicz ME, Brown RH, Jr., Garson JA. 2005. Detection of serum reverse transcriptase activity in patients with ALS and unaffected blood relatives. Neurology. 64(3):454–458.
  • Subramanian RP, Wildschutte JH, Russo C, Coffin JM. 2011. Identification, characterization, and comparative genomic distribution of the HERV-K (HML-2) group of human endogenous retroviruses. Retrovirology. 8:90.
  • Sugimoto J, Matsuura N, Kinjo Y, Takasu N, Oda T, Jinno Y. 2001. Transcriptionally active HERV-K genes: identification, isolation, and chromosomal mapping. Genomics. 72(2):137–144.
  • Suntsova M, Gogvadze EV, Salozhin S, Gaifullin N, Eroshkin F, Dmitriev SE, Martynova N, Kulikov K, Malakhova G, Tukhbatova G et al. 2013. Human-specific endogenous retroviral insert serves as an enhancer for the schizophrenia-linked gene PRODH. P Natl Acad Sci U S A. 110(48):19472–19477.
  • Suryawanshi A, Manicassamy S. 2015. Tumors induce immune tolerance through activation of beta-catenin/TCF4 signaling in dendritic cells: a novel therapeutic target for cancer immunotherapy. Oncoimmunology. 4(12):e1052932.
  • Tandon R, SenGupta D, Ndhlovu LC, Vieira RG, Jones RB, York VA, Vieira VA, Sharp ER, Wiznia AA, Ostrowski MA et al. 2011. Identification of human endogenous retrovirus-specific T cell responses in vertically HIV-1-infected subjects. J Virol. 85(21):11526–11531.
  • Terry SN, Manganaro L, Cuesta-Dominguez A, Brinzevich D, Simon V, Mulder LCF. 2017. Expression of HERV-K108 envelope interferes with HIV-1 production. Virology. 509:52–59.
  • Tomlins SA, Laxman B, Dhanasekaran SM, Helgeson BE, Cao X, Morris DS, Menon A, Jing X, Cao Q, Han B et al. 2007. Distinct classes of chromosomal rearrangements create oncogenic ETS gene fusions in prostate cancer. Nature. 448(7153):595–599.
  • Tonjes RR, Czauderna F, Kurth R. 1999. Genome-wide screening, cloning, chromosomal assignment, and expression of full-length human endogenous retrovirus type K. J Virol. 73(11):9187–9195.
  • Trono D. 1992. Partial reverse transcripts in virions from human immunodeficiency and murine leukemia viruses. J Virol. 66(8):4893–4900.
  • Turner G, Barbulescu M, Su M, Jensen-Seaman MI, Kidd KK, Lenz J. 2001. Insertional polymorphisms of full-length endogenous retroviruses in humans. Curr Biol. 11(19):1531–1535.
  • Tyagi R, Li W, Parades D, Bianchet MA, Nath A. 2017. Inhibition of human endogenous retrovirus-K by antiretroviral drugs. Retrovirology. 14(1):21.
  • van der Kuyl AC. 2012. HIV infection and HERV expression: a review. Retrovirology. 9:6.
  • Vertessy BG, Toth J. 2009. Keeping uracil out of DNA: physiological role, structure and catalytic mechanism of dUTPases. Acc Chem Res. 42(1):97–106.
  • Vincendeau M, Gottesdorfer I, Schreml JM, Wetie AG, Mayer J, Greenwood AD, Helfer M, Kramer S, Seifarth W, Hadian K et al. 2015. Modulation of human endogenous retrovirus (HERV) transcription during persistent and de novo HIV-1 infection. Retrovirology. 12:27.
  • Viola MV, Frazier M, White L, Brody J, Spiegelman S. 1975. RNA-instructed DNA polymerase activity in a cytoplasmic particulate fraction in brains from Guamanian patients. J Exp Med. 142(2):483–494.
  • Vogetseder W, Dumfahrt A, Mayersbach P, Schonitzer D, Dierich MP. 1993. Antibodies in human sera recognizing a recombinant outer membrane protein encoded by the envelope gene of the human endogenous retrovirus K. AIDS Res Hum Retroviruses. 9(7):687–694.
  • Wang T, Zeng J, Lowe CB, Sellers RG, Salama SR, Yang M, Burgess SM, Brachmann RK, Haussler D. 2007. Species-specific endogenous retroviruses shape the transcriptional network of the human tumor suppressor protein p53. Proc Natl Acad Sci U S A. 104(47):18613–18618.
  • Wang-Johanning F, Liu J, Rycaj K, Huang M, Tsai K, Rosen DG, Chen DT, Lu DW, Barnhart KF, Johanning GL. 2007. Expression of multiple human endogenous retrovirus surface envelope proteins in ovarian cancer. Int J Cancer. 120(1):81–90.
  • Watanabe T. 2017. Adult T-cell leukemia: molecular basis for clonal expansion and transformation of HTLV-1-infected T cells. Blood. 129(9):1071–1081.
  • Weiss RA. 1969a. The host range of Bryan strain Rous sarcoma virus synthesized in the absence of helper virus. J Gen Virol. 5:511–528.
  • Weiss RA. 1969b. Interference and neutralization studies with Bryan strain Rous sarcoma virus synthesized in the absence of helper virus. J Gen Virol. 5:529–539.
  • Weiss RA. 2006. The discovery of endogenous retroviruses. Retrovirology. 3:67.
  • Whitelaw E, Martin DIK. 2001. Retrotransposons as epigenetic mediators of phenotypic variation in mammals. Nat Genet. 27(4):361–365.
  • Wildschutte JH, Ram D, Subramanian R, Stevens VL, Coffin JM. 2014. The distribution of insertionally polymorphic endogenous retroviruses in breast cancer patients and cancer-free controls. Retrovirology. 11:62.
  • Wildschutte JH, Williams ZH, Montesion M, Subramanian RP, Kidd JM, Coffin JM. 2016. Discovery of unfixed endogenous retrovirus insertions in diverse human populations. Proc Natl Acad Sci U S A. 113(16):E2326–E2334.
  • Wilkinson D, Mager D, Leong JC. 1994. Endogenous human retroviruses. In: Levy J, editor. The retroviridae. New York: Plenum Press; p. 465–535.
  • Wodrich H, Krausslich HG. 2001. Nucleocytoplasmic RNA transport in retroviral replication. Results Probl Cell Differ. 34:197–217.
  • Wolf G, Yang P, Fuchtbauer AC, Fuichtbauer EM, Silva AM, Park C, Wu WR, Nielsen AL, Pedersen FS, Macfarlan TS. 2015. The KRAB zinc finger protein ZFP809 is required to initiate epigenetic silencing of endogenous retroviruses. Gene Dev. 29(5):538–554.
  • Yang BX, El Farran CA, Guo HC, Yu T, Fang HT, Wang HF, Schlesinger S, Seah YF, Goh GY, Neo SP et al. 2015. Systematic identification of factors for provirus silencing in embryonic stem cells. Cell. 163(1):230–245.
  • Yin H, Medstrand P, Kristofferson A, Dietrich U, Aman P, Blomberg J. 1999. Characterization of human MMTV-like (HML) elements similar to a sequence that was highly expressed in a human breast cancer: further definition of the HML-6 group. Virology. 256(1):22–35.
  • Yu SF, Baldwin DN, Gwynn SR, Yendapalli S, Linial ML. 1996. Human foamy virus replication: a pathway distinct from that of retroviruses and hepadnaviruses. Science. 271(5255):1579–1582.
  • Yu SF, Sullivan MD, Linial ML. 1999. Evidence that the human foamy virus genome is DNA. J Virol. 73(2):1565–1572.
  • Zhou F, Krishnamurthy J, Wei Y, Li M, Hunt K, Johanning GL, Cooper LJ, Wang-Johanning F. 2015. Chimeric antigen receptor T cells targeting HERV-K inhibit breast cancer and its metastasis through downregulation of Ras. Oncoimmunology. 4(11):e1047582.