{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,2,10]],"date-time":"2026-02-10T08:09:17Z","timestamp":1770710957843,"version":"3.49.0"},"reference-count":97,"publisher":"MDPI AG","issue":"5","license":[{"start":{"date-parts":[[2022,4,24]],"date-time":"2022-04-24T00:00:00Z","timestamp":1650758400000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"DOI":"10.13039\/501100000264","name":"Canadian liver foundation","doi-asserted-by":"publisher","award":["001"],"award-info":[{"award-number":["001"]}],"id":[{"id":"10.13039\/501100000264","id-type":"DOI","asserted-by":"publisher"}]}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":["Viruses"],"abstract":"<jats:p>A human betaretrovirus (HBRV) has been linked with the autoimmune liver disease, primary biliary cholangitis (PBC), and various cancers, including breast cancer and lymphoma. HBRV is closely related to the mouse mammary tumor virus, and represents the only exogenous betaretrovirus characterized in humans to date. Evidence of infection in patients with PBC has been demonstrated through the identification of proviral integration sites in lymphoid tissue, the major reservoir of infection, as well as biliary epithelium, which is the site of the disease process. Accordingly, we tested the hypothesis that patients with PBC harbor a transmissible betaretrovirus by co-cultivation of PBC patients\u2019 lymph node homogenates with the HS578T breast cancer line. Because of the low level of HBRV replication, betaretrovirus producing cells were subcloned to optimize viral isolation and production. Evidence of infection was provided by electron microscopy, RT-PCR, in situ hybridization, cloning of the HBRV proviral genome and demonstration of more than 3400 integration sites. Further evidence of viral transmissibility was demonstrated by infection of biliary epithelial cells. While HBRV did not show a preference for integration proximal to specific genomic features, analyses of common insertion sites revealed evidence of integration proximal to cancer associated genes. These studies demonstrate the isolation of HBRV with features similar to mouse mammary tumor virus and confirm that patients with PBC display evidence of a transmissible viral infection.<\/jats:p>","DOI":"10.3390\/v14050886","type":"journal-article","created":{"date-parts":[[2022,4,26]],"date-time":"2022-04-26T02:14:39Z","timestamp":1650939279000},"page":"886","update-policy":"https:\/\/doi.org\/10.3390\/mdpi_crossmark_policy","source":"Crossref","is-referenced-by-count":15,"title":["Isolation of a Human Betaretrovirus from Patients with Primary Biliary Cholangitis"],"prefix":"10.3390","volume":"14","author":[{"given":"Mariam","family":"Goubran","sequence":"first","affiliation":[{"name":"Center of Excellence for Gastrointestinal Inflammation and Immunity Research, University of Alberta, Edmonton, AB T6G 2E1, Canada"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Weiwei","family":"Wang","sequence":"additional","affiliation":[{"name":"Center of Excellence for Gastrointestinal Inflammation and Immunity Research, University of Alberta, Edmonton, AB T6G 2E1, Canada"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Stanislav","family":"Indik","sequence":"additional","affiliation":[{"name":"Department of Virology, University of Veterinary Medicine, A-1210 Vienna, Austria"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Alexander","family":"Faschinger","sequence":"additional","affiliation":[{"name":"Department of Virology, University of Veterinary Medicine, A-1210 Vienna, Austria"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Shawn T.","family":"Wasilenko","sequence":"additional","affiliation":[{"name":"Center of Excellence for Gastrointestinal Inflammation and Immunity Research, University of Alberta, Edmonton, AB T6G 2E1, Canada"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Jasper","family":"Bintner","sequence":"additional","affiliation":[{"name":"Center of Excellence for Gastrointestinal Inflammation and Immunity Research, University of Alberta, Edmonton, AB T6G 2E1, Canada"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Eric J.","family":"Carpenter","sequence":"additional","affiliation":[{"name":"Department of Biological Sciences, University of Alberta, Edmonton, AB T6G 2E9, Canada"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Guangzhi","family":"Zhang","sequence":"additional","affiliation":[{"name":"Center of Excellence for Gastrointestinal Inflammation and Immunity Research, University of Alberta, Edmonton, AB T6G 2E1, Canada"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Paulo","family":"Nuin","sequence":"additional","affiliation":[{"name":"Department of Medical Genetics, University of Alberta, Edmonton, AB T6G 2E1, Canada"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Georgina","family":"Macintyre","sequence":"additional","affiliation":[{"name":"Center of Excellence for Gastrointestinal Inflammation and Immunity Research, University of Alberta, Edmonton, AB T6G 2E1, Canada"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Gane K.-S.","family":"Wong","sequence":"additional","affiliation":[{"name":"Center of Excellence for Gastrointestinal Inflammation and Immunity Research, University of Alberta, Edmonton, AB T6G 2E1, Canada"},{"name":"Department of Biological Sciences, University of Alberta, Edmonton, AB T6G 2E9, Canada"},{"name":"Li Ka Shing Institute of Virology, University of Alberta, Edmonton, AB T6G 2E1, Canada"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Andrew L.","family":"Mason","sequence":"additional","affiliation":[{"name":"Center of Excellence for Gastrointestinal Inflammation and Immunity Research, University of Alberta, Edmonton, AB T6G 2E1, Canada"},{"name":"Li Ka Shing Institute of Virology, University of Alberta, Edmonton, AB T6G 2E1, Canada"}],"role":[{"role":"author","vocabulary":"crossref"}]}],"member":"1968","published-online":{"date-parts":[[2022,4,24]]},"reference":[{"key":"ref_1","doi-asserted-by":"crossref","first-page":"157","DOI":"10.1016\/j.jviromet.2009.09.015","article-title":"Detection of human mammary tumor virus proteins in human breast cancer cells","volume":"163","author":"Melana","year":"2010","journal-title":"J. Virol. Methods"},{"key":"ref_2","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1186\/s13027-016-0113-6","article-title":"Mouse mammary tumor-like virus (MMTV) is present in human breast tissue before development of virally associated breast cancer","volume":"12","author":"Nartey","year":"2017","journal-title":"Infect. Agent. Cancer"},{"key":"ref_3","doi-asserted-by":"crossref","first-page":"20","DOI":"10.1186\/1750-9378-9-20","article-title":"Human mammary tumor virus (HMTV) sequences in human milk","volume":"9","author":"Nartey","year":"2014","journal-title":"Infect. Agent Cancer"},{"key":"ref_4","doi-asserted-by":"crossref","first-page":"1044","DOI":"10.1002\/jmv.21754","article-title":"DNA of mouse mammary tumor virus-like virus is present in human tumors influenced by hormones","volume":"82","author":"Johal","year":"2010","journal-title":"J. Med. Virol."},{"key":"ref_5","doi-asserted-by":"crossref","first-page":"149","DOI":"10.1007\/s10549-011-1421-6","article-title":"Mouse mammary tumor like virus sequences in breast milk from healthy lactating women","volume":"129","author":"Johal","year":"2011","journal-title":"Breast Cancer Res. Treat."},{"key":"ref_6","doi-asserted-by":"crossref","first-page":"3576","DOI":"10.1158\/0008-5472.CAN-09-4160","article-title":"Mouse mammary tumor virus-like sequences in human breast cancer","volume":"70","author":"Lawson","year":"2010","journal-title":"Cancer Res."},{"key":"ref_7","doi-asserted-by":"crossref","first-page":"7284","DOI":"10.1158\/1078-0432.CCR-04-0767","article-title":"Mouse mammary tumor virus-like RNA transcripts and DNA are found in affected cells of human breast cancer","volume":"10","author":"Ford","year":"2004","journal-title":"Clin. Cancer Res."},{"key":"ref_8","first-page":"1118","article-title":"Mouse mammary tumor virus-like gene sequences in breast tumors of Australian and Vietnamese women","volume":"9","author":"Ford","year":"2003","journal-title":"Clin. Cancer Res."},{"key":"ref_9","doi-asserted-by":"crossref","first-page":"141","DOI":"10.3389\/fonc.2018.00141","article-title":"Association of mouse mammary tumor virus with human breast cancer: Histology, immunohistochemistry and polymerase chain reaction analyses","volume":"8","author":"Lawson","year":"2018","journal-title":"Front. Oncol."},{"key":"ref_10","doi-asserted-by":"crossref","first-page":"1287","DOI":"10.1136\/jcp.2005.035907","article-title":"Presence of mouse mammary tumour-like virus gene sequences may be associated with morphology of specific human breast cancer","volume":"59","author":"Lawson","year":"2006","journal-title":"J. Clin. Pathol."},{"key":"ref_11","first-page":"1754","article-title":"Identification of a proviral structure in human breast cancer","volume":"61","author":"Liu","year":"2001","journal-title":"Cancer Res."},{"key":"ref_12","doi-asserted-by":"crossref","first-page":"4105","DOI":"10.1158\/0008-5472.CAN-03-3880","article-title":"A mouse mammary tumor virus-like long terminal repeat superantigen in human breast cancer","volume":"64","author":"Wang","year":"2004","journal-title":"Cancer Res."},{"key":"ref_13","doi-asserted-by":"crossref","first-page":"171","DOI":"10.1007\/s007050170201","article-title":"MMTV-like env gene sequences in human breast cancer","volume":"146","author":"Wang","year":"2001","journal-title":"Arch. Virol."},{"key":"ref_14","first-page":"1041","article-title":"Detection of MMTV-like LTR and LTR-env gene sequences in human breast cancer","volume":"18","author":"Wang","year":"2001","journal-title":"Int. J. Oncol."},{"key":"ref_15","doi-asserted-by":"crossref","first-page":"436","DOI":"10.1002\/path.1997","article-title":"MMTV-like sequences in human breast cancer: A fluorescent PCR\/laser microdissection approach","volume":"209","author":"Zammarchi","year":"2006","journal-title":"J. Pathol."},{"key":"ref_16","doi-asserted-by":"crossref","first-page":"1588","DOI":"10.1016\/j.ajpath.2011.08.003","article-title":"Mouse mammary tumor virus in human breast cancer red herring or smoking gun?","volume":"179","author":"Mason","year":"2011","journal-title":"Am. J. Pathol."},{"key":"ref_17","doi-asserted-by":"crossref","first-page":"611","DOI":"10.1038\/229611a0","article-title":"Search for a human breast cancer virus","volume":"229","author":"Moore","year":"1971","journal-title":"Nature"},{"key":"ref_18","first-page":"5173","article-title":"Detection of mammary tumor virus env gene-like sequences in human breast cancer","volume":"55","author":"Wang","year":"1995","journal-title":"Cancer Res."},{"key":"ref_19","doi-asserted-by":"crossref","first-page":"393","DOI":"10.1111\/apt.13054","article-title":"Frequent proviral integration of the human betaretrovirus in biliary epithelium of patients with autoimmune and idiopathic liver disease","volume":"41","author":"Wang","year":"2015","journal-title":"Aliment. Pharmacol. Ther."},{"key":"ref_20","doi-asserted-by":"crossref","first-page":"151","DOI":"10.1002\/hep.20024","article-title":"Cloning the human betaretrovirus proviral genome from patients with primary biliary cirrhosis","volume":"39","author":"Xu","year":"2004","journal-title":"Hepatology"},{"key":"ref_21","doi-asserted-by":"crossref","first-page":"8454","DOI":"10.1073\/pnas.1433063100","article-title":"Does a betaretrovirus infection trigger primary biliary cirrhosis?","volume":"100","author":"Xu","year":"2003","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_22","doi-asserted-by":"crossref","first-page":"548","DOI":"10.1016\/j.jhep.2008.10.026","article-title":"Mouse mammary tumour virus-like virus (MMTV-LV) is present within the liver in a wide range of hepatic disorders and unrelated to nuclear p53 expression or hepatocarcinogenesis","volume":"50","author":"Johal","year":"2009","journal-title":"J. Hepatol."},{"key":"ref_23","doi-asserted-by":"crossref","first-page":"27","DOI":"10.1016\/j.bpg.2018.06.001","article-title":"Is PBC a viral infectious disease?","volume":"34\u201335","author":"Mason","year":"2018","journal-title":"Best Pract. Res. Clin. Gastroenterol."},{"key":"ref_24","doi-asserted-by":"crossref","first-page":"127","DOI":"10.1002\/j.1460-2075.1986.tb04186.x","article-title":"A new site of integration for mouse mammary tumor virus proviral DNA common to BALB\/cf(C3H) mammary and kidney adenocarcinomas","volume":"5","author":"Garcia","year":"1986","journal-title":"EMBO J."},{"key":"ref_25","doi-asserted-by":"crossref","first-page":"1442","DOI":"10.1111\/liv.12699","article-title":"Impact of combination antiretroviral therapy in the NOD.c3c4 mouse model of autoimmune biliary disease","volume":"35","author":"Sharon","year":"2015","journal-title":"Liver Int."},{"key":"ref_26","doi-asserted-by":"crossref","first-page":"876","DOI":"10.1016\/j.jhep.2011.01.037","article-title":"Mouse mammary tumor virus in anti-mitochondrial antibody producing mouse models","volume":"55","author":"Zhang","year":"2011","journal-title":"J. Hepatol."},{"key":"ref_27","doi-asserted-by":"crossref","first-page":"15","DOI":"10.1186\/s13100-017-0099-7","article-title":"Identification of a novel HERV-K(HML10): Comprehensive characterization and comparative analysis in non-human primates provide insights about HML10 proviruses structure and diffusion","volume":"8","author":"Grandi","year":"2017","journal-title":"Mob. DNA"},{"key":"ref_28","doi-asserted-by":"crossref","first-page":"1312","DOI":"10.1016\/j.jhep.2010.12.009","article-title":"The evidence supports a viral aetiology for primary biliary cirrhosis","volume":"54","author":"Mason","year":"2011","journal-title":"J. Hepatol."},{"key":"ref_29","doi-asserted-by":"crossref","first-page":"1315","DOI":"10.1016\/j.jhep.2010.12.010","article-title":"The evidence does not support a viral etiology for primary biliary cirrhosis","volume":"54","author":"Selmi","year":"2011","journal-title":"J. Hepatol."},{"key":"ref_30","doi-asserted-by":"crossref","first-page":"18355","DOI":"10.18632\/oncotarget.4567","article-title":"Human saliva as route of inter-human infection for mouse mammary tumor virus","volume":"6","author":"Mazzanti","year":"2015","journal-title":"Oncotarget"},{"key":"ref_31","doi-asserted-by":"crossref","first-page":"15978","DOI":"10.18632\/aging.103780","article-title":"A human MMTV-like betaretrovirus linked to breast cancer has been present in humans at least since the copper age","volume":"12","author":"Lessi","year":"2020","journal-title":"Aging"},{"key":"ref_32","doi-asserted-by":"crossref","first-page":"12386","DOI":"10.1073\/pnas.192360099","article-title":"Mouse transferrin receptor 1 is the cell entry receptor for mouse mammary tumor virus","volume":"99","author":"Ross","year":"2002","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_33","doi-asserted-by":"crossref","first-page":"169","DOI":"10.1002\/rmv.427","article-title":"A human murine mammary tumour virus-like agent is an unconvincing aetiological agent for human breast cancer","volume":"14","author":"Mant","year":"2004","journal-title":"Rev. Med. Virol."},{"key":"ref_34","doi-asserted-by":"crossref","first-page":"73","DOI":"10.1186\/1742-4690-4-73","article-title":"Rapid spread of mouse mammary tumor virus in cultured human breast cells","volume":"4","author":"Indik","year":"2007","journal-title":"Retrovirology"},{"key":"ref_35","doi-asserted-by":"crossref","first-page":"6651","DOI":"10.1158\/0008-5472.CAN-04-2609","article-title":"Mouse mammary tumor virus infects human cells","volume":"65","author":"Indik","year":"2005","journal-title":"Cancer Res."},{"key":"ref_36","doi-asserted-by":"crossref","first-page":"1360","DOI":"10.1128\/JVI.02098-07","article-title":"Mouse mammary tumor virus integration site selection in human and mouse genomes","volume":"82","author":"Faschinger","year":"2008","journal-title":"J. Virol."},{"key":"ref_37","doi-asserted-by":"crossref","first-page":"1565","DOI":"10.1016\/S0140-6736(15)00154-3","article-title":"Primary biliary cirrhosis","volume":"386","author":"Carey","year":"2015","journal-title":"Lancet"},{"key":"ref_38","doi-asserted-by":"crossref","first-page":"1620","DOI":"10.1016\/S0140-6736(97)10290-2","article-title":"Detection of retroviral antibodies in primary biliary cirrhosis and other idiopathic biliary disorders","volume":"351","author":"Mason","year":"1998","journal-title":"Lancet"},{"key":"ref_39","doi-asserted-by":"crossref","first-page":"375","DOI":"10.1111\/j.1478-3231.2008.01888.x","article-title":"Duplication of MER115 on chromosome 4 in patients with primary biliary cirrhosis","volume":"29","author":"Xu","year":"2009","journal-title":"Liver Int."},{"key":"ref_40","doi-asserted-by":"crossref","first-page":"493","DOI":"10.1053\/j.gastro.2004.05.033","article-title":"Lack of immunological or molecular evidence for a role of mouse mammary tumor retrovirus in primary biliary cirrhosis","volume":"127","author":"Selmi","year":"2004","journal-title":"Gastroenterology"},{"key":"ref_41","doi-asserted-by":"crossref","first-page":"157","DOI":"10.1128\/MMBR.00033-07","article-title":"Human RNA \u201cRumor\u201d Viruses: The Search for Novel Human Retroviruses in Chronic Disease","volume":"72","author":"Voisset","year":"2008","journal-title":"Microbiol. Mol. Biol. Rev."},{"key":"ref_42","first-page":"PO-1822","article-title":"Cellular immune responses to human betaretrovirus in patients with primary biliary cholangitis [ILC abstract]","volume":"75","author":"Abofayed","year":"2021","journal-title":"J. Hepatol."},{"key":"ref_43","first-page":"8958192","article-title":"Seroprevalence of human betaretrovirus surface protein antibodies in patients with breast cancer and liver disease","volume":"2020","author":"Zhang","year":"2020","journal-title":"J. Oncol."},{"key":"ref_44","doi-asserted-by":"crossref","first-page":"31","DOI":"10.3138\/canlivj.2018-0020","article-title":"Randomized controlled trial: Combination antiretroviral therapy with Tenofovir-Emtricitabine and Lopinavir-Ritonavir in patients with primary biliary cholangitis","volume":"2","author":"Lytvyak","year":"2019","journal-title":"Can. Liver J."},{"key":"ref_45","doi-asserted-by":"crossref","first-page":"349","DOI":"10.3748\/wjg.v22.i1.349","article-title":"Combination antiretroviral studies for patients with primary biliary cirrhosis","volume":"22","author":"Lytvyak","year":"2016","journal-title":"World J. Gastroenterol."},{"key":"ref_46","doi-asserted-by":"crossref","first-page":"886","DOI":"10.1111\/j.1365-2036.2008.03799.x","article-title":"Clinical trial: Randomized controlled study of zidovudine and lamivudine for patients with primary biliary cirrhosis stabilized on ursodiol","volume":"28","author":"Mason","year":"2008","journal-title":"Aliment. Pharmacol. Ther."},{"key":"ref_47","doi-asserted-by":"crossref","first-page":"97","DOI":"10.1055\/s-2007-1007187","article-title":"Ductular expression of autoantigens in primary biliary cirrhosis","volume":"17","author":"Joplin","year":"1997","journal-title":"Semin. Liver Dis."},{"key":"ref_48","doi-asserted-by":"crossref","first-page":"1595","DOI":"10.1016\/S0140-6736(05)61042-2","article-title":"Expression of pyruvate-dehydrogenase complex PDC-E2 on biliary epithelial cells induced by lymph nodes from primary biliary cirrhosis","volume":"352","author":"Sadamoto","year":"1998","journal-title":"Lancet"},{"key":"ref_49","doi-asserted-by":"crossref","first-page":"650","DOI":"10.1099\/jgv.0.000006","article-title":"C3H strain of mouse mammary tumour virus, like GR strain, infects human mammary epithelial cells, albeit less efficiently than murine mammary epithelial cells","volume":"96","author":"Konstantoulas","year":"2015","journal-title":"J. Gen. Virol."},{"key":"ref_50","doi-asserted-by":"crossref","first-page":"43","DOI":"10.1186\/s12977-015-0168-2","article-title":"Single amino acid substitution (G42E) in the receptor binding domain of mouse mammary tumour virus envelope protein facilitates infection of non-murine cells in a transferrin receptor 1-independent manner","volume":"12","author":"Konstantoulas","year":"2015","journal-title":"Retrovirology"},{"key":"ref_51","doi-asserted-by":"crossref","first-page":"1795","DOI":"10.1093\/jnci\/58.6.1795","article-title":"Two syngeneic cell lines from human breast tissue: The aneuploid mammary epithelial (Hs578T) and the diploid myoepithelial (Hs578Bst) cell lines","volume":"58","author":"Hackett","year":"1977","journal-title":"J. Natl. Cancer Inst."},{"key":"ref_52","doi-asserted-by":"crossref","first-page":"1320","DOI":"10.18632\/oncotarget.682","article-title":"Genes affected by mouse mammary tumor virus (MMTV) proviral insertions in mouse mammary tumors are deregulated or mutated in primary human mammary tumors","volume":"3","author":"Callahan","year":"2012","journal-title":"Oncotarget"},{"key":"ref_53","doi-asserted-by":"crossref","first-page":"875","DOI":"10.1136\/gut.35.7.875","article-title":"Isolation and culture of biliary epithelial cells","volume":"35","author":"Joplin","year":"1994","journal-title":"Gut"},{"key":"ref_54","doi-asserted-by":"crossref","first-page":"587","DOI":"10.1111\/j.1365-2036.2007.03402.x","article-title":"Reverse transcriptase activity in patients with primary biliary cirrhosis and other autoimmune liver disorders","volume":"26","author":"McDermid","year":"2007","journal-title":"Aliment. Pharmacol. Ther."},{"key":"ref_55","doi-asserted-by":"crossref","unstructured":"Kearney, M.F., Spindler, J., Wiegand, A., Shao, W., Anderson, E.M., Maldarelli, F., Ruscetti, F.W., Mellors, J.W., Hughes, S.H., and Le Grice, S.F. (2012). Multiple sources of contamination in samples from patients reported to have XMRV infection. PLoS ONE, 7.","DOI":"10.1371\/journal.pone.0030889"},{"key":"ref_56","doi-asserted-by":"crossref","first-page":"47","DOI":"10.1186\/s12977-016-0277-6","article-title":"Retrovirus Integration Database (RID): A public database for retroviral insertion sites into host genomes","volume":"13","author":"Shao","year":"2016","journal-title":"Retrovirology"},{"key":"ref_57","doi-asserted-by":"crossref","unstructured":"Zhu, L.J., Gazin, C., Lawson, N.D., Pag\u00e8s, H., Lin, S.M., Lapointe, D.S., and Green, M.R. (2010). ChIPpeakAnno: A Bioconductor package to annotate ChIP-seq and ChIP-chip data. BMC Bioinform., 11.","DOI":"10.1186\/1471-2105-11-237"},{"key":"ref_58","doi-asserted-by":"crossref","first-page":"153","DOI":"10.1038\/ng950","article-title":"High-throughput retroviral tagging to identify components of specific signaling pathways in cancer","volume":"32","author":"Mikkers","year":"2002","journal-title":"Nat. Genet."},{"key":"ref_59","doi-asserted-by":"crossref","first-page":"D844","DOI":"10.1093\/nar\/gku770","article-title":"The Candidate Cancer Gene Database: A database of cancer driver genes from forward genetic screens in mice","volume":"43","author":"Abbott","year":"2015","journal-title":"Nucleic Acids Res."},{"key":"ref_60","doi-asserted-by":"crossref","first-page":"420","DOI":"10.1093\/bioinformatics\/bts715","article-title":"In silico identification of oncogenic potential of fyn-related kinase in hepatocellular carcinoma","volume":"29","author":"Chen","year":"2013","journal-title":"Bioinformatics"},{"key":"ref_61","doi-asserted-by":"crossref","first-page":"555","DOI":"10.1038\/s41568-020-0290-x","article-title":"A compendium of mutational cancer driver genes","volume":"20","author":"Mularoni","year":"2020","journal-title":"Nat. Rev. Cancer"},{"key":"ref_62","doi-asserted-by":"crossref","first-page":"696","DOI":"10.1038\/s41568-018-0060-1","article-title":"The COSMIC Cancer Gene Census: Describing genetic dysfunction across all human cancers","volume":"18","author":"Sondka","year":"2018","journal-title":"Nat. Rev. Cancer"},{"key":"ref_63","doi-asserted-by":"crossref","first-page":"446","DOI":"10.1054\/bjoc.1999.0941","article-title":"Breast cancer incidence highest in the range of one species of house mouse, Mus domesticus","volume":"82","author":"Stewart","year":"2000","journal-title":"Br. J. Cancer"},{"key":"ref_64","doi-asserted-by":"crossref","unstructured":"Hagen, B., Kraase, M., Indikova, I., and Indik, S. (2019). A high rate of polymerization during synthesis of mouse mammary tumor virus DNA alleviates hypermutation by APOBEC3 proteins. PLoS Pathog., 15.","DOI":"10.1371\/journal.ppat.1007533"},{"key":"ref_65","doi-asserted-by":"crossref","first-page":"1652","DOI":"10.1038\/s41598-019-38733-1","article-title":"Plasmid DNA contaminant in molecular reagents","volume":"9","author":"Wally","year":"2019","journal-title":"Sci. Rep."},{"key":"ref_66","doi-asserted-by":"crossref","first-page":"886","DOI":"10.1016\/j.celrep.2013.09.040","article-title":"The BET family of proteins targets moloney murine leukemia virus integration near transcription start sites","volume":"5","author":"Demeulemeester","year":"2013","journal-title":"Cell Rep."},{"key":"ref_67","doi-asserted-by":"crossref","unstructured":"Mitchell, R.S., Beitzel, B.F., Schroder, A.R., Shinn, P., Chen, H., Berry, C.C., Ecker, J.R., and Bushman, F.D. (2004). Retroviral DNA integration: ASLV, HIV, and MLV show distinct target site preferences. PLoS Biol., 2.","DOI":"10.1371\/journal.pbio.0020234"},{"key":"ref_68","doi-asserted-by":"crossref","unstructured":"Gilroy, K.L., Terry, A., Naseer, A., de Ridder, J., Allahyar, A., Wang, W., Carpenter, E., Mason, A., Wong, G.K., and Cameron, E.R. (2016). Gamma-Retrovirus Integration Marks Cell Type-Specific Cancer Genes: A Novel Profiling Tool in Cancer Genomics. PLoS ONE, 11.","DOI":"10.1371\/journal.pone.0154070"},{"key":"ref_69","doi-asserted-by":"crossref","first-page":"1767","DOI":"10.1101\/gad.1565107","article-title":"LEDGF\/p75 functions downstream from preintegration complex formation to effect gene-specific HIV-1 integration","volume":"21","author":"Shun","year":"2007","journal-title":"Genes Dev."},{"key":"ref_70","doi-asserted-by":"crossref","first-page":"2988","DOI":"10.1158\/0008-5472.CAN-10-4026","article-title":"Genome-wide DNA methylation profiling of CpG islands in breast cancer identifies novel genes associated with tumorigenicity","volume":"71","author":"Hill","year":"2011","journal-title":"Cancer Res."},{"key":"ref_71","doi-asserted-by":"crossref","first-page":"1010428317705754","DOI":"10.1177\/1010428317705754","article-title":"PTEN: A potential prognostic marker in virus-induced hepatocellular carcinoma","volume":"39","author":"Khalid","year":"2017","journal-title":"Tumour Biol."},{"key":"ref_72","doi-asserted-by":"crossref","first-page":"135","DOI":"10.1080\/0284186X.2018.1529428","article-title":"Identifying breast cancer susceptibility genes\u2014A review of the genetic background in familial breast cancer","volume":"58","author":"Wendt","year":"2019","journal-title":"Acta Oncol."},{"key":"ref_73","doi-asserted-by":"crossref","first-page":"1032","DOI":"10.18632\/aging.100855","article-title":"The PTEN tumor suppressor gene and its role in lymphoma pathogenesis","volume":"7","author":"Wang","year":"2015","journal-title":"Aging"},{"key":"ref_74","doi-asserted-by":"crossref","first-page":"1011","DOI":"10.1080\/08941939.2020.1728443","article-title":"High SPINK1 Expression Predicts Poor Prognosis and Promotes Cell Proliferation and Metastasis of Hepatocellular Carcinoma","volume":"34","author":"Huang","year":"2021","journal-title":"J. Investig. Surg."},{"key":"ref_75","doi-asserted-by":"crossref","first-page":"812","DOI":"10.1097\/CAD.0000000000001070","article-title":"Long noncoding RNA ZFPM2-AS1 promotes the proliferation, migration, and invasion of hepatocellular carcinoma cells by regulating the miR-576-3p\/HIF-1\u03b1 axis","volume":"32","author":"Song","year":"2021","journal-title":"Anticancer Drugs"},{"key":"ref_76","first-page":"11139","article-title":"LncRNA ZFPM2-AS1 aggravates the malignant development of breast cancer via upregulating JMJD6","volume":"24","author":"Zhao","year":"2020","journal-title":"Eur. Rev. Med. Pharmacol. Sci."},{"key":"ref_77","doi-asserted-by":"crossref","first-page":"625","DOI":"10.1002\/jcp.25466","article-title":"MicroRNA-30b Suppresses Epithelial-Mesenchymal Transition and Metastasis of Hepatoma Cells","volume":"232","author":"Sun","year":"2017","journal-title":"J. Cell Physiol."},{"key":"ref_78","doi-asserted-by":"crossref","first-page":"5259","DOI":"10.1158\/0008-5472.CAN-17-3058","article-title":"miRNA-30 Family Members Inhibit Breast Cancer Invasion, Osteomimicry, and Bone Destruction by Directly Targeting Multiple Bone Metastasis-Associated Genes","volume":"78","author":"Croset","year":"2018","journal-title":"Cancer Res."},{"key":"ref_79","doi-asserted-by":"crossref","first-page":"7656","DOI":"10.1038\/sj.onc.1209043","article-title":"Retroviral insertional mutagenesis: Past, present and future","volume":"24","author":"Uren","year":"2005","journal-title":"Oncogene"},{"key":"ref_80","doi-asserted-by":"crossref","unstructured":"Yang, J., Liu, Y., Wang, B., Lan, H., Liu, Y., Chen, F., Zhang, J., and Luo, J. (2017). Sumoylation in p27kip1 via RanBP2 promotes cancer cell growth in cholangiocarcinoma cell line QBC939. BMC Mol. Biol., 18.","DOI":"10.1186\/s12867-017-0100-5"},{"key":"ref_81","doi-asserted-by":"crossref","first-page":"2243","DOI":"10.1038\/s41388-021-01704-w","article-title":"The RanBP2\/RanGAP1-SUMO complex gates \u03b2-arrestin2 nuclear entry to regulate the Mdm2-p53 signaling axis","volume":"40","author":"Leverve","year":"2021","journal-title":"Oncogene"},{"key":"ref_82","doi-asserted-by":"crossref","first-page":"722","DOI":"10.1073\/pnas.1615735114","article-title":"piggyBac mediates efficient in vivo CRISPR library screening for tumorigenesis in mice","volume":"114","author":"Xu","year":"2017","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_83","doi-asserted-by":"crossref","first-page":"7575862","DOI":"10.1155\/2020\/7575862","article-title":"Coexpression Network Analysis of Genes Related to the Characteristics of Tumor Stemness in Triple-Negative Breast Cancer","volume":"2020","author":"Suo","year":"2020","journal-title":"Biomed Res. Int."},{"key":"ref_84","doi-asserted-by":"crossref","first-page":"30453","DOI":"10.18632\/oncotarget.5852","article-title":"Mitochondrial mass, a new metabolic biomarker for stem-like cancer cells: Understanding WNT\/FGF-driven anabolic signaling","volume":"6","author":"Lamb","year":"2015","journal-title":"Oncotarget"},{"key":"ref_85","doi-asserted-by":"crossref","first-page":"992","DOI":"10.1038\/sj.onc.1203276","article-title":"MMTV-induced mammary tumorigenesis: Gene discovery, progression to malignancy and cellular pathways","volume":"19","author":"Callahan","year":"2000","journal-title":"Oncogene"},{"key":"ref_86","doi-asserted-by":"crossref","first-page":"565","DOI":"10.1136\/gut.49.4.565","article-title":"Identification of novel molecules and pathogenic pathways in primary biliary cirrhosis: cDNA array analysis of intrahepatic differential gene expression","volume":"49","author":"Shackel","year":"2001","journal-title":"Gut"},{"key":"ref_87","doi-asserted-by":"crossref","first-page":"1409","DOI":"10.1002\/hep.25788","article-title":"Primary biliary cirrhosis and cancer risk: A systematic review and meta-analysis","volume":"56","author":"Liang","year":"2012","journal-title":"Hepatology"},{"key":"ref_88","doi-asserted-by":"crossref","first-page":"321","DOI":"10.1136\/gutjnl-2014-308351","article-title":"Stratification of hepatocellular carcinoma risk in primary biliary cirrhosis: A multicentre international study","volume":"65","author":"Trivedi","year":"2016","journal-title":"Gut"},{"key":"ref_89","first-page":"42","article-title":"Autoimmune liver diseases, hepatocellular carcinoma, and recurrence of autoimmunity post-liver transplantation","volume":"7","author":"Wang","year":"2021","journal-title":"Hepatoma Res."},{"key":"ref_90","doi-asserted-by":"crossref","first-page":"633","DOI":"10.1038\/314633a0","article-title":"Rearrangements of the cellular p53 gene in erythroleukaemic cells transformed by Friend virus","volume":"314","author":"Mowat","year":"1985","journal-title":"Nature"},{"key":"ref_91","doi-asserted-by":"crossref","first-page":"1141","DOI":"10.1158\/1541-7786.MCR-13-0244","article-title":"Cancer gene discovery: Exploiting insertional mutagenesis","volume":"11","author":"Ranzani","year":"2013","journal-title":"Mol. Cancer Res."},{"key":"ref_92","doi-asserted-by":"crossref","first-page":"262","DOI":"10.1016\/j.clbc.2020.01.007","article-title":"Prognostic and Predictive Impact of Beta-2 Adrenergic Receptor Expression in HER2-Positive Breast Cancer","volume":"20","author":"Caparica","year":"2020","journal-title":"Clin. Breast Cancer"},{"key":"ref_93","doi-asserted-by":"crossref","first-page":"9239","DOI":"10.1038\/s41598-020-66197-1","article-title":"DNA methylation markers panel can improve prediction of response to neoadjuvant chemotherapy in luminal B breast cancer","volume":"10","author":"Sigin","year":"2020","journal-title":"Sci. Rep."},{"key":"ref_94","doi-asserted-by":"crossref","unstructured":"Pearlman, A., Rahman, M.T., Upadhyay, K., Loke, J., and Ostrer, H. (2019). Ectopic Otoconin 90 expression in triple negative breast cancer cell lines is associated with metastasis functions. PLoS ONE, 14.","DOI":"10.1371\/journal.pone.0211737"},{"key":"ref_95","doi-asserted-by":"crossref","first-page":"5","DOI":"10.1007\/s12185-018-2518-z","article-title":"Oncogenic transcriptional program driven by TAL1 in T-cell acute lymphoblastic leukemia","volume":"109","author":"Tan","year":"2019","journal-title":"Int J. Hematol."},{"key":"ref_96","first-page":"1356","article-title":"Clinical significance of aberrant DEUP1 promoter methylation in hepatocellular carcinoma","volume":"18","author":"Yu","year":"2019","journal-title":"Oncol. Lett."},{"key":"ref_97","doi-asserted-by":"crossref","first-page":"e2014622","DOI":"10.1001\/jamanetworkopen.2020.14622","article-title":"Prediction of Overall Survival Among female patients with breast cancer using a prognostic signature based on 8 DNA repair-related genes","volume":"3","author":"Zhang","year":"2020","journal-title":"JAMA Netw. 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