{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,5,20]],"date-time":"2026-05-20T15:13:25Z","timestamp":1779290005817,"version":"3.51.4"},"reference-count":150,"publisher":"MDPI AG","issue":"5","license":[{"start":{"date-parts":[[2026,5,19]],"date-time":"2026-05-19T00:00:00Z","timestamp":1779148800000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":["Viruses"],"abstract":"<jats:p>BK and JC Polyomaviruses (BKPyV and JCPyV) are ubiquitous human pathogens capable of establishing lifelong, asymptomatic persistence in the majority of the global population. While decades of research have focused on their lytic replication cycles and the development of severe diseases, such as polyomavirus-associated nephropathy (PVAN) caused by BKPyV and progressive multifocal leukoencephalopathy (PML) caused by JCPyV, their primary evolutionary strategy is one of persistence rather than pathogenesis. This review shifts the perspective from a replication-centric framework towards an evolutionary persistence model, detailing the multi-layered host and viral determinants that maintain the homeostatic balance. At the cellular level, viral genomes are restricted by chromatinization into minichromosomes and host S-phase licensing. These constraints are reinforced by innate immune sensing and adaptive T-cell and antibody responses that curtail systemic dissemination while permitting periodic, low-level urinary shedding, which is essential for horizontal transmission. In addition to these host barriers, the viruses utilize intrinsic regulatory mechanisms to prevent excessive replication and immune detection, including the stable archetype non-coding control region (NCCR), viral microRNAs that downregulate early gene expression, and the small t antigen (STAg). Finally, we address unresolved questions regarding the full spectrum of cellular reservoirs, the molecular triggers of reactivation, and the ecological factors shaping their transmission routes. Understanding these maintenance mechanisms is crucial for refining clinical interventions and managing the rare, devastating transitions from silent persistence to lytic disease.<\/jats:p>","DOI":"10.3390\/v18050571","type":"journal-article","created":{"date-parts":[[2026,5,20]],"date-time":"2026-05-20T14:12:54Z","timestamp":1779286374000},"page":"571","update-policy":"https:\/\/doi.org\/10.3390\/mdpi_crossmark_policy","source":"Crossref","is-referenced-by-count":0,"title":["Living on the Edge: The Goldilocks Zone of Polyomavirus Replication and Persistence"],"prefix":"10.3390","volume":"18","author":[{"ORCID":"https:\/\/orcid.org\/0000-0002-8928-4884","authenticated-orcid":false,"given":"Wenqing","family":"Yuan","sequence":"first","affiliation":[{"name":"Department of Molecular Biology, Cell Biology, and Biochemistry, Brown University, Providence, RI 02912, USA"},{"name":"Graduate Program in Pathobiology, Brown University, Providence, RI 02912, USA"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-2855-5668","authenticated-orcid":false,"given":"Sheila A.","family":"Haley","sequence":"additional","affiliation":[{"name":"Department of Molecular Biology, Cell Biology, and Biochemistry, Brown University, Providence, RI 02912, USA"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-1218-6458","authenticated-orcid":false,"given":"Michael J.","family":"Imperiale","sequence":"additional","affiliation":[{"name":"Department of Microbiology and Immunology, University of Michigan, Ann Arbor, MI 48109, USA"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-3763-9073","authenticated-orcid":false,"given":"Walter J.","family":"Atwood","sequence":"additional","affiliation":[{"name":"Department of Molecular Biology, Cell Biology, and Biochemistry, Brown University, Providence, RI 02912, USA"}],"role":[{"role":"author","vocabulary":"crossref"}]}],"member":"1968","published-online":{"date-parts":[[2026,5,19]]},"reference":[{"key":"ref_1","doi-asserted-by":"crossref","first-page":"178","DOI":"10.1080\/1040841X.2016.1189873","article-title":"45 years after the discovery of human polyomaviruses BK and JC: Time to speed up the understanding of associated diseases and treatment approaches","volume":"43","author":"Barth","year":"2017","journal-title":"Crit. Rev. Microbiol."},{"key":"ref_2","doi-asserted-by":"crossref","unstructured":"Atkinson, A.L., and Atwood, W.J. (2020). Fifty Years of JC Polyomavirus: A Brief Overview and Remaining Questions. Viruses, 12.","DOI":"10.3390\/v12090969"},{"key":"ref_3","doi-asserted-by":"crossref","first-page":"503","DOI":"10.1128\/CMR.00074-16","article-title":"BK Polyomavirus: Clinical Aspects, Immune Regulation, and Emerging Therapies","volume":"30","author":"Ambalathingal","year":"2017","journal-title":"Clin. Microbiol. Rev."},{"key":"ref_4","doi-asserted-by":"crossref","first-page":"349","DOI":"10.1146\/annurev-virology-101416-041439","article-title":"Progressive Multifocal Leukoencephalopathy: Endemic Viruses and Lethal Brain Disease","volume":"4","author":"Haley","year":"2017","journal-title":"Annu. Rev. Virol."},{"key":"ref_5","doi-asserted-by":"crossref","first-page":"1253","DOI":"10.1016\/S0140-6736(71)91776-4","article-title":"New human papovavirus (B.K.) isolated from urine after renal transplantation","volume":"1","author":"Gardner","year":"1971","journal-title":"Lancet"},{"key":"ref_6","doi-asserted-by":"crossref","first-page":"1257","DOI":"10.1016\/S0140-6736(71)91777-6","article-title":"Cultivation of papova-like virus from human brain with progressive multifocal leucoencephalopathy","volume":"1","author":"Padgett","year":"1971","journal-title":"Lancet"},{"key":"ref_7","doi-asserted-by":"crossref","first-page":"1361","DOI":"10.1128\/JVI.80.3.1361-1366.2006","article-title":"Identification of gangliosides GD1b and GT1b as receptors for BK virus","volume":"80","author":"Low","year":"2006","journal-title":"J. Virol."},{"key":"ref_8","doi-asserted-by":"crossref","first-page":"309","DOI":"10.1016\/j.chom.2010.09.004","article-title":"Structure-function analysis of the human JC polyomavirus establishes the LSTc pentasaccharide as a functional receptor motif","volume":"8","author":"Neu","year":"2010","journal-title":"Cell Host Microbe"},{"key":"ref_9","doi-asserted-by":"crossref","first-page":"13490","DOI":"10.1128\/JVI.02252-13","article-title":"5-HT2 receptors facilitate JC polyomavirus entry","volume":"87","author":"Assetta","year":"2013","journal-title":"J. Virol."},{"key":"ref_10","doi-asserted-by":"crossref","first-page":"1380","DOI":"10.1126\/science.1103492","article-title":"The human polyomavirus, JCV, uses serotonin receptors to infect cells","volume":"306","author":"Elphick","year":"2004","journal-title":"Science"},{"key":"ref_11","doi-asserted-by":"crossref","unstructured":"Mehmood, K., Wilczek, M.P., DuShane, J.K., Parent, M.T., Mayberry, C.L., Wallace, J.N., Levasseur, F.L., Fong, T.M., Hess, S.T., and Maginnis, M.S. (2022). Dynamics and Patterning of 5-Hydroxytryptamine 2 Subtype Receptors in JC Polyomavirus Entry. Viruses, 14.","DOI":"10.3390\/v14122597"},{"key":"ref_12","doi-asserted-by":"crossref","first-page":"e02139-20","DOI":"10.1128\/JVI.02139-20","article-title":"GRK2 mediates beta-arrestin interactions with 5-HT(2) receptors for JC polyomavirus endocytosis","volume":"95","author":"Mayberry","year":"2021","journal-title":"J. Virol."},{"key":"ref_13","doi-asserted-by":"crossref","first-page":"841","DOI":"10.1084\/jem.20072097","article-title":"Polyomavirus BK with rearranged noncoding control region emerge in vivo in renal transplant patients and increase viral replication and cytopathology","volume":"205","author":"Gosert","year":"2008","journal-title":"J. Exp. Med."},{"key":"ref_14","doi-asserted-by":"crossref","first-page":"10448","DOI":"10.1128\/JVI.00614-10","article-title":"Rearranged JC virus noncoding control regions found in progressive multifocal leukoencephalopathy patient samples increase virus early gene expression and replication rate","volume":"84","author":"Gosert","year":"2010","journal-title":"J. Virol."},{"key":"ref_15","doi-asserted-by":"crossref","first-page":"261","DOI":"10.1007\/s00705-003-0214-6","article-title":"Unambiguous identification of JC polyomavirus strains transmitted from parents to children","volume":"149","author":"Zheng","year":"2004","journal-title":"Arch. Virol."},{"key":"ref_16","doi-asserted-by":"crossref","first-page":"61","DOI":"10.1093\/aje\/kwv155","article-title":"Seroepidemiology of Human Polyomaviruses in a US Population","volume":"183","author":"Gossai","year":"2016","journal-title":"Am. J. Epidemiol."},{"key":"ref_17","doi-asserted-by":"crossref","first-page":"837","DOI":"10.1086\/597126","article-title":"Prevalence of polyomavirus BK and JC infection and replication in 400 healthy blood donors","volume":"199","author":"Egli","year":"2009","journal-title":"J. Infect. Dis."},{"key":"ref_18","doi-asserted-by":"crossref","first-page":"1044","DOI":"10.1099\/vir.0.028571-0","article-title":"Serological evidence of vertical transmission of JC and BK polyomaviruses in humans","volume":"92","author":"Boldorini","year":"2011","journal-title":"J. Gen. Virol."},{"key":"ref_19","doi-asserted-by":"crossref","first-page":"640","DOI":"10.1111\/j.1469-0691.2004.00882.x","article-title":"Prevalence and patterns of polyomavirus urinary excretion in immunocompetent adults and children","volume":"10","author":"Polo","year":"2004","journal-title":"Clin. Microbiol. Infect."},{"key":"ref_20","doi-asserted-by":"crossref","first-page":"26.2. 1","DOI":"10.1002\/0471143030.cb2602s42","article-title":"BK virus (BKV): Infection, propagation, quantitation, purification, labeling, and analysis of cell entry","volume":"42","author":"Moriyama","year":"2009","journal-title":"Curr. Protoc. Cell Biol."},{"key":"ref_21","doi-asserted-by":"crossref","first-page":"32","DOI":"10.1186\/s12985-025-02643-8","article-title":"Clinical and molecular analysis of JCPyV and BKPyV infection and associated risk of urothelial carcinoma development in the upper tract","volume":"22","author":"Chao","year":"2025","journal-title":"Virol. J."},{"key":"ref_22","doi-asserted-by":"crossref","first-page":"204","DOI":"10.1016\/j.micpath.2016.06.015","article-title":"BK and JC virus infections in healthy patients compared to kidney transplant recipients in Tunisia","volume":"97","author":"Boukoum","year":"2016","journal-title":"Microb. Pathog."},{"key":"ref_23","doi-asserted-by":"crossref","first-page":"160","DOI":"10.1007\/0-387-32957-9_11","article-title":"Polyomavirus-associated nephropathy in renal transplantation: Critical issues of screening and management","volume":"577","author":"Hirsch","year":"2006","journal-title":"Adv. Exp. Med. Biol."},{"key":"ref_24","doi-asserted-by":"crossref","unstructured":"Mojumder, A., Nguyen, K.W., and Sullivan, C.S. (2026). Divergent Fates of Kidney-Resident Polyomaviruses: Stable Shedding Versus Near-Silent Persistence. Viruses, 18.","DOI":"10.64898\/2026.02.12.705499"},{"key":"ref_25","doi-asserted-by":"crossref","unstructured":"Blois, S., Goetz, B.M., Mojumder, A., and Sullivan, C.S. (2025). Shedding dynamics of a DNA virus population during acute and long-term persistent infection. PLoS Pathog., 21.","DOI":"10.1101\/2025.03.31.646279"},{"key":"ref_26","doi-asserted-by":"crossref","unstructured":"Moens, U., Prezioso, C., and Pietropaolo, V. (2020). Genetic Diversity of the Noncoding Control Region of the Novel Human Polyomaviruses. Viruses, 12.","DOI":"10.3390\/v12121406"},{"key":"ref_27","doi-asserted-by":"crossref","first-page":"1571","DOI":"10.1099\/0022-1317-76-7-1571","article-title":"Subpopulations of non-coding control region variants within a cell culture-passaged stock of BK virus: Sequence comparisons and biological characteristics","volume":"76","author":"Johnsen","year":"1995","journal-title":"J. Gen. Virol."},{"key":"ref_28","doi-asserted-by":"crossref","first-page":"261","DOI":"10.1007\/BF01701816","article-title":"Noncoding control region of naturally occurring BK virus variants: Sequence comparison and functional analysis","volume":"10","author":"Moens","year":"1995","journal-title":"Virus Genes"},{"key":"ref_29","first-page":"150","article-title":"Noncoding Control Region Pattern of BK polyomavirus in Kidney Transplant Patients With Nephropathy","volume":"15","author":"Emami","year":"2017","journal-title":"Exp. Clin. Transplant."},{"key":"ref_30","doi-asserted-by":"crossref","unstructured":"Haghi Navand, A., Jalilian, S., Ahmadi Angali, K., Karimi Babaahmadi, M., Talaiezadeh, A., and Makvandi, M. (2024). A new evaluation of the rearranged non-coding control region of JC virus in patients with colorectal cancer. BMC Cancer, 24.","DOI":"10.1186\/s12885-024-12684-x"},{"key":"ref_31","doi-asserted-by":"crossref","unstructured":"Yang, J.F., and You, J. (2020). Regulation of Polyomavirus Transcription by Viral and Cellular Factors. Viruses, 12.","DOI":"10.3390\/v12101072"},{"key":"ref_32","doi-asserted-by":"crossref","first-page":"6205","DOI":"10.1093\/emboj\/cdg612","article-title":"Large T antigen on the simian virus 40 origin of replication: A 3D snapshot prior to DNA replication","volume":"22","author":"Valle","year":"2003","journal-title":"EMBO J."},{"key":"ref_33","doi-asserted-by":"crossref","first-page":"1544","DOI":"10.1074\/jbc.M109.064311","article-title":"Large T antigen promotes JC virus replication in G2-arrested cells by inducing ATM- and ATR-mediated G2 checkpoint signaling","volume":"285","author":"Orba","year":"2010","journal-title":"J. Biol. Chem."},{"key":"ref_34","doi-asserted-by":"crossref","unstructured":"Bollag, B., Hofstetter, C.A., Reviriego-Mendoza, M.M., and Frisque, R.J. (2010). JC virus small T antigen binds phosphatase PP2A and Rb family proteins and is required for efficient viral DNA replication activity. PLoS ONE, 5.","DOI":"10.1371\/journal.pone.0010606"},{"key":"ref_35","doi-asserted-by":"crossref","first-page":"e0007723","DOI":"10.1128\/jvi.00077-23","article-title":"Regulation of Virus Replication by BK Polyomavirus Small T Antigen","volume":"97","author":"Zou","year":"2023","journal-title":"J. Virol."},{"key":"ref_36","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1016\/j.virol.2013.10.031","article-title":"Human polyoma JC virus minor capsid proteins, VP2 and VP3, enhance large T antigen binding to the origin of viral DNA replication: Evidence for their involvement in regulation of the viral DNA replication","volume":"449","author":"Saribas","year":"2014","journal-title":"Virology"},{"key":"ref_37","doi-asserted-by":"crossref","first-page":"12079","DOI":"10.1128\/JVI.01456-06","article-title":"NFAT4 is required for JC virus infection of glial cells","volume":"80","author":"Manley","year":"2006","journal-title":"J. Virol."},{"key":"ref_38","doi-asserted-by":"crossref","first-page":"1722","DOI":"10.1128\/JVI.01918-09","article-title":"Transcriptional regulation of BK virus by nuclear factor of activated T cells","volume":"84","author":"Jordan","year":"2010","journal-title":"J. Virol."},{"key":"ref_39","doi-asserted-by":"crossref","first-page":"3396","DOI":"10.1128\/JVI.03625-14","article-title":"Sp1 sites in the noncoding control region of BK polyomavirus are key regulators of bidirectional viral early and late gene expression","volume":"89","author":"Bethge","year":"2015","journal-title":"J. Virol."},{"key":"ref_40","doi-asserted-by":"crossref","first-page":"9918","DOI":"10.1128\/JVI.72.12.9918-9923.1998","article-title":"Detection of JC virus DNA in human tonsil tissue: Evidence for site of initial viral infection","volume":"72","author":"Monaco","year":"1998","journal-title":"J. Virol."},{"key":"ref_41","doi-asserted-by":"crossref","first-page":"143","DOI":"10.1002\/jmv.1890080208","article-title":"The persistence of papovavirus BK DNA sequences in normal human renal tissue","volume":"8","author":"Heritage","year":"1981","journal-title":"J. Med. Virol."},{"key":"ref_42","doi-asserted-by":"crossref","first-page":"59","DOI":"10.1016\/j.jcv.2015.08.003","article-title":"Mechanisms of BK virus infection of renal cells and therapeutic implications","volume":"71","author":"Mbianda","year":"2015","journal-title":"J. Clin. Virol."},{"key":"ref_43","doi-asserted-by":"crossref","first-page":"37","DOI":"10.1038\/s41582-020-00427-y","article-title":"Progressive multifocal leukoencephalopathy and the spectrum of JC virus-related disease","volume":"17","author":"Cortese","year":"2021","journal-title":"Nat. Rev. Neurol."},{"key":"ref_44","doi-asserted-by":"crossref","first-page":"471","DOI":"10.1128\/CMR.05031-11","article-title":"Molecular biology, epidemiology, and pathogenesis of progressive multifocal leukoencephalopathy, the JC virus-induced demyelinating disease of the human brain","volume":"25","author":"Ferenczy","year":"2012","journal-title":"Clin. Microbiol. Rev."},{"key":"ref_45","doi-asserted-by":"crossref","first-page":"266","DOI":"10.1016\/j.virol.2008.09.027","article-title":"The role of polyomaviruses in human disease","volume":"384","author":"Jiang","year":"2009","journal-title":"Virology"},{"key":"ref_46","doi-asserted-by":"crossref","first-page":"29","DOI":"10.1016\/j.coviro.2014.03.012","article-title":"Polyomavirus miRNAs: The beginning","volume":"7","author":"Imperiale","year":"2014","journal-title":"Curr. Opin. Virol."},{"key":"ref_47","doi-asserted-by":"crossref","first-page":"839719","DOI":"10.1155\/2013\/839719","article-title":"New insights on human polyomavirus JC and pathogenesis of progressive multifocal leukoencephalopathy","volume":"2013","author":"Bellizzi","year":"2013","journal-title":"Clin. Dev. Immunol."},{"key":"ref_48","doi-asserted-by":"crossref","first-page":"235","DOI":"10.1016\/j.virol.2011.10.026","article-title":"Efficient propagation of archetype BK and JC polyomaviruses","volume":"422","author":"Broekema","year":"2012","journal-title":"Virology"},{"key":"ref_49","doi-asserted-by":"crossref","first-page":"261","DOI":"10.1016\/j.semcancer.2009.02.013","article-title":"JC virus: An oncogenic virus in animals and humans?","volume":"19","author":"Maginnis","year":"2009","journal-title":"Semin. Cancer Biol."},{"key":"ref_50","doi-asserted-by":"crossref","first-page":"e1815","DOI":"10.1212\/WNL.0000000000005529","article-title":"Progressive multifocal leukoencephalopathy after fingolimod treatment","volume":"90","author":"Berger","year":"2018","journal-title":"Neurology"},{"key":"ref_51","doi-asserted-by":"crossref","first-page":"e13528","DOI":"10.1111\/ctr.13528","article-title":"BK polyomavirus in solid organ transplantation-Guidelines from the American Society of Transplantation Infectious Diseases Community of Practice","volume":"33","author":"Hirsch","year":"2019","journal-title":"Clin. Transplant."},{"key":"ref_52","doi-asserted-by":"crossref","first-page":"485","DOI":"10.1097\/00007890-200208270-00009","article-title":"Quantitation of viral DNA in renal allograft tissue from patients with BK virus nephropathy","volume":"74","author":"Randhawa","year":"2002","journal-title":"Transplantation"},{"key":"ref_53","doi-asserted-by":"crossref","first-page":"115","DOI":"10.1002\/jmv.10450","article-title":"Population-based study of antibody to the human polyomaviruses BKV and JCV and the simian polyomavirus SV40","volume":"71","author":"Knowles","year":"2003","journal-title":"J. Med. Virol."},{"key":"ref_54","doi-asserted-by":"crossref","first-page":"825","DOI":"10.1681\/ASN.2010080877","article-title":"Inhibitory interactions between BK and JC virus among kidney transplant recipients","volume":"22","author":"Cheng","year":"2011","journal-title":"J. Am. Soc. Nephrol."},{"key":"ref_55","doi-asserted-by":"crossref","unstructured":"Alcendor, D.J. (2019). BK Polyomavirus Virus Glomerular Tropism: Implications for Virus Reactivation from Latency and Amplification during Immunosuppression. J. Clin. Med., 8.","DOI":"10.3390\/jcm8091477"},{"key":"ref_56","doi-asserted-by":"crossref","first-page":"9200","DOI":"10.1128\/JVI.00609-10","article-title":"JC virus latency in the brain and extraneural organs of patients with and without progressive multifocal leukoencephalopathy","volume":"84","author":"Tan","year":"2010","journal-title":"J. Virol."},{"key":"ref_57","doi-asserted-by":"crossref","first-page":"7004","DOI":"10.1128\/jvi.70.10.7004-7012.1996","article-title":"JC virus infection of hematopoietic progenitor cells, primary B lymphocytes, and tonsillar stromal cells: Implications for viral latency","volume":"70","author":"Monaco","year":"1996","journal-title":"J. Virol."},{"key":"ref_58","doi-asserted-by":"crossref","first-page":"127","DOI":"10.3109\/13550280009013156","article-title":"JC virus binds to primary human glial cells, tonsillar stromal cells, and B-lymphocytes, but not to T lymphocytes","volume":"6","author":"Wei","year":"2000","journal-title":"J. Neurovirol."},{"key":"ref_59","doi-asserted-by":"crossref","first-page":"419","DOI":"10.1016\/S0002-9440(10)63132-X","article-title":"Differential distribution of the JC virus receptor-type sialic acid in normal human tissues","volume":"164","author":"Eash","year":"2004","journal-title":"Am. J. Pathol."},{"key":"ref_60","doi-asserted-by":"crossref","first-page":"1228","DOI":"10.1053\/gast.2000.19269","article-title":"JC virus DNA sequences are frequently present in the human upper and lower gastrointestinal tract","volume":"119","author":"Ricciardiello","year":"2000","journal-title":"Gastroenterology"},{"key":"ref_61","doi-asserted-by":"crossref","first-page":"7484","DOI":"10.1073\/pnas.96.13.7484","article-title":"JC virus DNA is present in the mucosa of the human colon and in colorectal cancers","volume":"96","author":"Laghi","year":"1999","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_62","doi-asserted-by":"crossref","first-page":"379","DOI":"10.1002\/ana.21443","article-title":"Detection of JC virus DNA fragments but not proteins in normal brain tissue","volume":"64","author":"Gentilella","year":"2008","journal-title":"Ann. Neurol."},{"key":"ref_63","doi-asserted-by":"crossref","first-page":"59","DOI":"10.3109\/13550289809113482","article-title":"Progressive multifocal leukoencephalopathy in patients with HIV infection","volume":"4","author":"Berger","year":"1998","journal-title":"J. Neurovirol."},{"key":"ref_64","doi-asserted-by":"crossref","first-page":"1430","DOI":"10.1212\/WNL.0b013e31828c2fa1","article-title":"PML diagnostic criteria: Consensus statement from the AAN Neuroinfectious Disease Section","volume":"80","author":"Berger","year":"2013","journal-title":"Neurology"},{"key":"ref_65","doi-asserted-by":"crossref","unstructured":"An, P., Saenz Robles, M.T., Duray, A.M., Cantalupo, P.G., and Pipas, J.M. (2019). Human polyomavirus BKV infection of endothelial cells results in interferon pathway induction and persistence. PLoS Pathog., 15.","DOI":"10.1371\/journal.ppat.1007505"},{"key":"ref_66","doi-asserted-by":"crossref","first-page":"275","DOI":"10.1146\/annurev-virology-092818-015716","article-title":"Hitchhiking of Viral Genomes on Cellular Chromosomes","volume":"6","author":"Coursey","year":"2019","journal-title":"Annu. Rev. Virol."},{"key":"ref_67","doi-asserted-by":"crossref","unstructured":"Majumder, K., and Morales, A.J. (2021). Utilization of Host Cell Chromosome Conformation by Viral Pathogens: Knowing When to Hold and When to Fold. Front. Immunol., 12.","DOI":"10.3389\/fimmu.2021.633762"},{"key":"ref_68","doi-asserted-by":"crossref","first-page":"1126","DOI":"10.1073\/pnas.75.3.1126","article-title":"Minichromosome from BK virus as a template for transcription in vitro","volume":"75","author":"Meneguzzi","year":"1978","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_69","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1016\/j.virol.2015.04.009","article-title":"Global profiling of histone modifications in the polyomavirus BK virion minichromosome","volume":"483","author":"Fang","year":"2015","journal-title":"Virology"},{"key":"ref_70","doi-asserted-by":"crossref","unstructured":"Needham, J.M., Perritt, S.E., and Thompson, S.R. (2024). Single-cell analysis reveals host S phase drives large T antigen expression during BK polyomavirus infection. PLoS Pathog., 20.","DOI":"10.1371\/journal.ppat.1012663"},{"key":"ref_71","doi-asserted-by":"crossref","first-page":"5708","DOI":"10.1128\/JVI.00300-09","article-title":"Restriction of human polyomavirus BK virus DNA replication in murine cells and extracts","volume":"83","author":"Mahon","year":"2009","journal-title":"J. Virol."},{"key":"ref_72","doi-asserted-by":"crossref","first-page":"265","DOI":"10.1159\/000496278","article-title":"The Role of Renal Interstitial Cells in Proximal Tubular Regeneration","volume":"141","author":"Castrop","year":"2019","journal-title":"Nephron"},{"key":"ref_73","doi-asserted-by":"crossref","first-page":"3752","DOI":"10.1093\/ndt\/gfn470","article-title":"BK virus large T and VP-1 expression in infected human renal allografts","volume":"23","author":"Seemayer","year":"2008","journal-title":"Nephrol. Dial. Transplant."},{"key":"ref_74","doi-asserted-by":"crossref","unstructured":"Lorentzen, E.M., Henriksen, S., and Rinaldo, C.H. (2023). Modelling BK Polyomavirus dissemination and cytopathology using polarized human renal tubule epithelial cells. PLoS Pathog., 19.","DOI":"10.1371\/journal.ppat.1011622"},{"key":"ref_75","doi-asserted-by":"crossref","first-page":"e00903","DOI":"10.1128\/mBio.00903-16","article-title":"JC Polyomavirus Infection of Primary Human Renal Epithelial Cells Is Controlled by a Type I IFN-Induced Response","volume":"7","author":"Assetta","year":"2016","journal-title":"mBio"},{"key":"ref_76","doi-asserted-by":"crossref","first-page":"73","DOI":"10.1016\/j.virol.2009.10.047","article-title":"Global effects of BKV infection on gene expression in human primary kidney epithelial cells","volume":"397","author":"Abend","year":"2010","journal-title":"Virology"},{"key":"ref_77","doi-asserted-by":"crossref","first-page":"712","DOI":"10.1086\/520518","article-title":"Interferon- alpha and - beta restrict polyomavirus JC replication in primary human fetal glial cells: Implications for progressive multifocal leukoencephalopathy therapy","volume":"196","author":"Co","year":"2007","journal-title":"J. Infect. Dis."},{"key":"ref_78","doi-asserted-by":"crossref","first-page":"457","DOI":"10.1016\/j.virol.2005.10.012","article-title":"JC virus induces altered patterns of cellular gene expression: Interferon-inducible genes as major transcriptional targets","volume":"345","author":"Verma","year":"2006","journal-title":"Virology"},{"key":"ref_79","doi-asserted-by":"crossref","first-page":"e0062824","DOI":"10.1128\/spectrum.00628-24","article-title":"JCPyV infection of primary choroid plexus epithelial cells reduces expression of critical junctional proteins and increases expression of barrier disrupting inflammatory cytokines","volume":"12","author":"Haley","year":"2024","journal-title":"Microbiol. Spectr."},{"key":"ref_80","doi-asserted-by":"crossref","first-page":"100","DOI":"10.1038\/ki.2011.311","article-title":"Activation of innate immune defense mechanisms contributes to polyomavirus BK-associated nephropathy","volume":"81","author":"Ribeiro","year":"2012","journal-title":"Kidney Int."},{"key":"ref_81","doi-asserted-by":"crossref","first-page":"101257","DOI":"10.1016\/j.isci.2020.101257","article-title":"BK Polyomavirus Evades Innate Immune Sensing by Disrupting the Mitochondrial Network and Promotes Mitophagy","volume":"23","author":"Manzetti","year":"2020","journal-title":"iScience"},{"key":"ref_82","doi-asserted-by":"crossref","first-page":"e00620-21","DOI":"10.1128\/mBio.00620-21","article-title":"The Small t Antigen of JC Virus Antagonizes RIG-I-Mediated Innate Immunity by Inhibiting TRIM25\u2019s RNA Binding Ability","volume":"12","author":"Chiang","year":"2021","journal-title":"mBio"},{"key":"ref_83","doi-asserted-by":"crossref","first-page":"13009","DOI":"10.1128\/JVI.01786-13","article-title":"The T antigen locus of Merkel cell polyomavirus downregulates human Toll-like receptor 9 expression","volume":"87","author":"Shahzad","year":"2013","journal-title":"J. Virol."},{"key":"ref_84","doi-asserted-by":"crossref","first-page":"100682","DOI":"10.1016\/j.bj.2023.100682","article-title":"Interferon-alpha and MxA inhibit BK polyomavirus replication by interaction with polyomavirus large T antigen","volume":"47","author":"Wu","year":"2024","journal-title":"Biomed. J."},{"key":"ref_85","doi-asserted-by":"crossref","unstructured":"Hou, Y., Hu, C., Shi, Y., Zhou, X., Zhu, T., and Wu, N. (2025). ISG20 Restricts BK Polyomavirus Infection and Engages in Reciprocal Regulation with Viral Large T Antigen. Microorganisms, 13.","DOI":"10.3390\/microorganisms13112540"},{"key":"ref_86","doi-asserted-by":"crossref","first-page":"628","DOI":"10.1016\/j.chom.2018.04.005","article-title":"Characterization of BK Polyomaviruses from Kidney Transplant Recipients Suggests a Role for APOBEC3 in Driving In-Host Virus Evolution","volume":"23","author":"Peretti","year":"2018","journal-title":"Cell Host Microbe"},{"key":"ref_87","doi-asserted-by":"crossref","first-page":"eaea6124","DOI":"10.1126\/sciadv.aea6124","article-title":"Virus-induced APOBEC3 transmutagenesis in bladder cancer initiation","volume":"11","author":"Hatton","year":"2025","journal-title":"Sci. Adv."},{"key":"ref_88","doi-asserted-by":"crossref","unstructured":"Poulain, F., Lejeune, N., Willemart, K., and Gillet, N.A. (2020). Footprint of the host restriction factors APOBEC3 on the genome of human viruses. PLoS Pathog., 16.","DOI":"10.1371\/journal.ppat.1008718"},{"key":"ref_89","doi-asserted-by":"crossref","first-page":"5850","DOI":"10.1128\/JVI.01979-15","article-title":"Emerging Role of PML Nuclear Bodies in Innate Immune Signaling","volume":"90","author":"Scherer","year":"2016","journal-title":"J. Virol."},{"key":"ref_90","doi-asserted-by":"crossref","first-page":"109866","DOI":"10.1016\/j.virol.2023.109866","article-title":"Human neurotropic polyomavirus, JC virus, late coding region encodes a novel nuclear protein, ORF4, which targets the promyelocytic leukemia nuclear bodies (PML-NBs) and modulates their reorganization","volume":"587","author":"Saribas","year":"2023","journal-title":"Virology"},{"key":"ref_91","doi-asserted-by":"crossref","first-page":"279","DOI":"10.1016\/j.virol.2009.05.017","article-title":"Modulation of PML protein expression regulates JCV infection","volume":"390","author":"Gasparovic","year":"2009","journal-title":"Virology"},{"key":"ref_92","doi-asserted-by":"crossref","unstructured":"Shishido-Hara, Y., and Yaoi, T. (2025). JC Virus Agnogene Regulates Histone-Modifying Enzymes via PML-NBs: Transcriptomics in VLP-Expressing Cells. Viruses, 17.","DOI":"10.3390\/v17101399"},{"key":"ref_93","doi-asserted-by":"crossref","first-page":"442","DOI":"10.1097\/NEN.0000000000000066","article-title":"JC virus inclusions in progressive multifocal leukoencephalopathy: Scaffolding promyelocytic leukemia nuclear bodies grow with cell cycle transition through an S-to-G2-like state in enlarging oligodendrocyte nuclei","volume":"73","author":"Yazawa","year":"2014","journal-title":"J. Neuropathol. Exp. Neurol."},{"key":"ref_94","doi-asserted-by":"crossref","first-page":"58","DOI":"10.1016\/j.yexcr.2004.03.045","article-title":"PML-nuclear bodies accumulate DNA in response to polyomavirus BK and simian virus 40 replication","volume":"298","author":"Visted","year":"2004","journal-title":"Exp. Cell Res."},{"key":"ref_95","doi-asserted-by":"crossref","first-page":"2591","DOI":"10.1111\/ajt.14282","article-title":"BK Polyomavirus-Specific 9mer CD8 T Cell Responses Correlate with Clearance of BK Viremia in Kidney Transplant Recipients: First Report From the Swiss Transplant Cohort Study","volume":"17","author":"Leboeuf","year":"2017","journal-title":"Am. J. Transplant."},{"key":"ref_96","doi-asserted-by":"crossref","first-page":"1131","DOI":"10.1111\/j.1600-6143.2007.01754.x","article-title":"Polyomavirus BK-specific cellular immune response to VP1 and large T-antigen in kidney transplant recipients","volume":"7","author":"Binggeli","year":"2007","journal-title":"Am. J. Transplant."},{"key":"ref_97","doi-asserted-by":"crossref","first-page":"499","DOI":"10.4049\/jimmunol.168.1.499","article-title":"Association of prolonged survival in HLA-A2+ progressive multifocal leukoencephalopathy patients with a CTL response specific for a commonly recognized JC virus epitope","volume":"168","author":"Koralnik","year":"2002","journal-title":"J. Immunol."},{"key":"ref_98","doi-asserted-by":"crossref","unstructured":"Perkins, M.R., Ryschkewitsch, C., Liebner, J.C., Monaco, M.C., Himelfarb, D., Ireland, S., Roque, A., Edward, H.L., Jensen, P.N., and Remington, G. (2012). Changes in JC virus-specific T cell responses during natalizumab treatment and in natalizumab-associated progressive multifocal leukoencephalopathy. PLoS Pathog., 8.","DOI":"10.1371\/journal.ppat.1003014"},{"key":"ref_99","doi-asserted-by":"crossref","unstructured":"Pastrana, D.V., Brennan, D.C., Cuburu, N., Storch, G.A., Viscidi, R.P., Randhawa, P.S., and Buck, C.B. (2012). Neutralization serotyping of BK polyomavirus infection in kidney transplant recipients. PLoS Pathog., 8.","DOI":"10.1371\/journal.ppat.1002650"},{"key":"ref_100","doi-asserted-by":"crossref","first-page":"161","DOI":"10.1111\/ajt.13880","article-title":"Pretransplantation Donor-Recipient Pair Seroreactivity Against BK Polyomavirus Predicts Viremia and Nephropathy After Kidney Transplantation","volume":"17","author":"Wunderink","year":"2017","journal-title":"Am. J. Transplant."},{"key":"ref_101","doi-asserted-by":"crossref","first-page":"137","DOI":"10.1007\/s13365-013-0222-6","article-title":"Immune surveillance and response to JC virus infection and PML","volume":"20","author":"Beltrami","year":"2014","journal-title":"J. Neurovirol."},{"key":"ref_102","doi-asserted-by":"crossref","first-page":"306ra151","DOI":"10.1126\/scitranslmed.aab1720","article-title":"JC polyomavirus mutants escape antibody-mediated neutralization","volume":"7","author":"Ray","year":"2015","journal-title":"Sci. Transl. Med."},{"key":"ref_103","doi-asserted-by":"crossref","first-page":"193","DOI":"10.1128\/JCM.01645-06","article-title":"Age-related urinary excretion of BK polyomavirus by nonimmunocompromised individuals","volume":"45","author":"Zhong","year":"2007","journal-title":"J. Clin. Microbiol."},{"key":"ref_104","doi-asserted-by":"crossref","first-page":"e9","DOI":"10.1086\/504947","article-title":"JC virus detection in bodily fluids: Clues to transmission","volume":"43","author":"Berger","year":"2006","journal-title":"Clin. Infect. Dis."},{"key":"ref_105","doi-asserted-by":"crossref","first-page":"280","DOI":"10.1080\/13550280152537102","article-title":"A classification scheme for human polyomavirus JCV variants based on the nucleotide sequence of the noncoding regulatory region","volume":"7","author":"Jensen","year":"2001","journal-title":"J. Neurovirol."},{"key":"ref_106","doi-asserted-by":"crossref","first-page":"250","DOI":"10.1002\/rmv.1747","article-title":"Serological cross-reactivity between human polyomaviruses","volume":"23","author":"Moens","year":"2013","journal-title":"Rev. Med. Virol."},{"key":"ref_107","doi-asserted-by":"crossref","first-page":"682","DOI":"10.1038\/nature03576","article-title":"SV40-encoded microRNAs regulate viral gene expression and reduce susceptibility to cytotoxic T cells","volume":"435","author":"Sullivan","year":"2005","journal-title":"Nature"},{"key":"ref_108","doi-asserted-by":"crossref","first-page":"9823","DOI":"10.1128\/JVI.01144-08","article-title":"Evolutionarily conserved function of a viral microRNA","volume":"82","author":"Seo","year":"2008","journal-title":"J. Virol."},{"key":"ref_109","doi-asserted-by":"crossref","unstructured":"Takahashi, K., Sato, Y., Sekizuka, T., Kuroda, M., Suzuki, T., Hasegawa, H., and Katano, H. (2020). High expression of JC polyomavirus-encoded microRNAs in progressive multifocal leukoencephalopathy tissues and its repressive role in virus replication. PLoS Pathog., 16.","DOI":"10.1371\/journal.ppat.1008523"},{"key":"ref_110","doi-asserted-by":"crossref","unstructured":"Mishra, R., Kumar, A., Ingle, H., and Kumar, H. (2019). The Interplay Between Viral-Derived miRNAs and Host Immunity During Infection. Front. Immunol., 10.","DOI":"10.3389\/fimmu.2019.03079"},{"key":"ref_111","doi-asserted-by":"crossref","first-page":"93","DOI":"10.1016\/j.chom.2011.01.008","article-title":"An identical miRNA of the human JC and BK polyoma viruses targets the stress-induced ligand ULBP3 to escape immune elimination","volume":"9","author":"Bauman","year":"2011","journal-title":"Cell Host Microbe"},{"key":"ref_112","doi-asserted-by":"crossref","first-page":"12683","DOI":"10.1128\/JVI.01933-14","article-title":"Naturally arising strains of polyomaviruses with severely attenuated microRNA expression","volume":"88","author":"Chen","year":"2014","journal-title":"J. Virol."},{"key":"ref_113","doi-asserted-by":"crossref","first-page":"268","DOI":"10.1186\/1743-422X-10-268","article-title":"The miRNA world of polyomaviruses","volume":"10","author":"Lagatie","year":"2013","journal-title":"Virol. J."},{"key":"ref_114","doi-asserted-by":"crossref","first-page":"70","DOI":"10.1016\/j.jcv.2018.02.007","article-title":"BK polyomavirus microRNA expression and sequence variation in polyomavirus-associated nephropathy","volume":"102","author":"Virtanen","year":"2018","journal-title":"J. Clin. Virol."},{"key":"ref_115","doi-asserted-by":"crossref","unstructured":"Cho, U.S., Morrone, S., Sablina, A.A., Arroyo, J.D., Hahn, W.C., and Xu, W. (2007). Structural basis of PP2A inhibition by small t antigen. PLoS Biol., 5.","DOI":"10.1371\/journal.pbio.0050202"},{"key":"ref_116","doi-asserted-by":"crossref","first-page":"464","DOI":"10.1016\/j.virol.2008.02.020","article-title":"Dephosphorylation of JC virus agnoprotein by protein phosphatase 2A: Inhibition by small t antigen","volume":"375","author":"Sariyer","year":"2008","journal-title":"Virology"},{"key":"ref_117","doi-asserted-by":"crossref","unstructured":"Zhou, X., Zhu, C., and Li, H. (2023). BK polyomavirus: Latency, reactivation, diseases and tumorigenesis. Front. Cell Infect. Microbiol., 13.","DOI":"10.3389\/fcimb.2023.1263983"},{"key":"ref_118","doi-asserted-by":"crossref","first-page":"617","DOI":"10.1182\/bloodadvances.2019000802","article-title":"Presentation of BK polyomavirus-associated hemorrhagic cystitis after allogeneic hematopoietic cell transplantation","volume":"4","author":"Imlay","year":"2020","journal-title":"Blood Adv."},{"key":"ref_119","doi-asserted-by":"crossref","first-page":"464","DOI":"10.4103\/ijn.ijn_150_22","article-title":"A Rare Cause of Encephalopathy Post Renal Transplant: BK Polyoma Virus Encephalitis","volume":"33","author":"Jeemon","year":"2023","journal-title":"Indian. J. Nephrol."},{"key":"ref_120","doi-asserted-by":"crossref","unstructured":"Wang, Y., Fang, Y., Yan, Z., Xia, R., Zeng, W., Deng, W., Xu, J., Feng, X., Peng, J., and Miao, Y. (2023). Fatal BK polyomavirus-associated pneumonia: Report of two cases with literature review. BMC Infect. Dis., 23.","DOI":"10.1186\/s12879-023-08577-2"},{"key":"ref_121","doi-asserted-by":"crossref","first-page":"560","DOI":"10.1002\/ana.24371","article-title":"Persistence and pathogenesis of the neurotropic polyomavirus JC","volume":"77","author":"Wollebo","year":"2015","journal-title":"Ann. Neurol."},{"key":"ref_122","doi-asserted-by":"crossref","first-page":"467","DOI":"10.1016\/S1474-4422(18)30040-1","article-title":"Pathogenesis of progressive multifocal leukoencephalopathy and risks associated with treatments for multiple sclerosis: A decade of lessons learned","volume":"17","author":"Major","year":"2018","journal-title":"Lancet Neurol."},{"key":"ref_123","doi-asserted-by":"crossref","first-page":"6221","DOI":"10.1128\/JVI.00396-13","article-title":"Differentiation of human fetal multipotential neural progenitor cells to astrocytes reveals susceptibility factors for JC virus","volume":"87","author":"Ferenczy","year":"2013","journal-title":"J. Virol."},{"key":"ref_124","doi-asserted-by":"crossref","unstructured":"O\u2019Hara, B.A., Gee, G.V., Atwood, W.J., and Haley, S.A. (2018). Susceptibility of Primary Human Choroid Plexus Epithelial Cells and Meningeal Cells to Infection by JC Virus. J. Virol., 92.","DOI":"10.1128\/JVI.00105-18"},{"key":"ref_125","doi-asserted-by":"crossref","first-page":"e0061625","DOI":"10.1128\/jvi.00616-25","article-title":"Neurovascular pericytes are susceptible to infection by JC polyomavirus","volume":"99","author":"Garabian","year":"2025","journal-title":"J. Virol."},{"key":"ref_126","doi-asserted-by":"crossref","first-page":"3394","DOI":"10.1128\/JVI.77.6.3394-3401.2003","article-title":"Glial cell-specific regulation of the JC virus early promoter by histone deacetylase inhibitors","volume":"77","author":"Kim","year":"2003","journal-title":"J. Virol."},{"key":"ref_127","doi-asserted-by":"crossref","first-page":"264","DOI":"10.1186\/1743-422X-10-264","article-title":"Epigenetic regulation of polyomavirus JC","volume":"10","author":"Wollebo","year":"2013","journal-title":"Virol. J."},{"key":"ref_128","doi-asserted-by":"crossref","first-page":"736","DOI":"10.1016\/0042-6822(92)90040-V","article-title":"Persistence of archetypal JC virus DNA in normal renal tissue derived from tumor-bearing patients","volume":"186","author":"Tominaga","year":"1992","journal-title":"Virology"},{"key":"ref_129","doi-asserted-by":"crossref","first-page":"103","DOI":"10.1093\/infdis\/jir198","article-title":"Progressive multifocal leukoencephalopathy (PML) development is associated with mutations in JC virus capsid protein VP1 that change its receptor specificity","volume":"204","author":"Gorelik","year":"2011","journal-title":"J. Infect. Dis."},{"key":"ref_130","doi-asserted-by":"crossref","first-page":"5177","DOI":"10.1128\/JVI.03221-13","article-title":"Lymphocyte gene expression and JC virus noncoding control region sequences are linked with the risk of progressive multifocal leukoencephalopathy","volume":"88","author":"Marshall","year":"2014","journal-title":"J. Virol."},{"key":"ref_131","doi-asserted-by":"crossref","first-page":"23","DOI":"10.1186\/s12985-020-1295-5","article-title":"JCPyV NCCR analysis in PML patients with different risk factors: Exploring common rearrangements as essential changes for neuropathogenesis","volume":"17","author":"Ciardi","year":"2020","journal-title":"Virol. J."},{"key":"ref_132","doi-asserted-by":"crossref","first-page":"659","DOI":"10.1099\/0022-1317-78-3-659","article-title":"JC virus regulatory region rearrangements and genotypes in progressive multifocal leukoencephalopathy: Two independent aspects of virus variation","volume":"78","author":"Agostini","year":"1997","journal-title":"J. Gen. Virol."},{"key":"ref_133","doi-asserted-by":"crossref","first-page":"e0152725","DOI":"10.1128\/spectrum.01527-25","article-title":"Therapeutic potential of flavonols in the treatment of JCPyV","volume":"13","author":"MacLure","year":"2025","journal-title":"Microbiol. Spectr."},{"key":"ref_134","doi-asserted-by":"crossref","unstructured":"Biffi, R., Benoit, S.W., Sariyer, I.K., and Safak, M. (2024). JC virus small tumor antigen promotes S phase entry and cell cycle progression. Tumour Virus Res., 18.","DOI":"10.1016\/j.tvr.2024.200298"},{"key":"ref_135","doi-asserted-by":"crossref","first-page":"404","DOI":"10.1007\/s11481-010-9203-1","article-title":"Molecular regulation of JC virus tropism: Insights into potential therapeutic targets for progressive multifocal leukoencephalopathy","volume":"5","author":"Marshall","year":"2010","journal-title":"J. Neuroimmune Pharmacol."},{"key":"ref_136","doi-asserted-by":"crossref","first-page":"1196","DOI":"10.1097\/00002030-200106150-00026","article-title":"BK virus encephalitis in a patient with AIDS and lymphoma","volume":"15","author":"Lesprit","year":"2001","journal-title":"AIDS"},{"key":"ref_137","doi-asserted-by":"crossref","first-page":"8200","DOI":"10.1073\/pnas.1301907110","article-title":"miRNA regulation of BK polyomavirus replication during early infection","volume":"110","author":"Broekema","year":"2013","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_138","doi-asserted-by":"crossref","first-page":"7","DOI":"10.1186\/s12985-014-0220-1","article-title":"Agnoprotein of polyomavirus BK interacts with proliferating cell nuclear antigen and inhibits DNA replication","volume":"12","author":"Gerits","year":"2015","journal-title":"Virol. J."},{"key":"ref_139","doi-asserted-by":"crossref","first-page":"250","DOI":"10.1086\/514032","article-title":"Analysis of the systemic and intrathecal humoral immune response in progressive multifocal leukoencephalopathy","volume":"176","author":"Weber","year":"1997","journal-title":"J. Infect. Dis."},{"key":"ref_140","doi-asserted-by":"crossref","first-page":"1057","DOI":"10.1111\/j.1348-0421.1982.tb00254.x","article-title":"Prevalence rate and age of acquisition of antibodies against JC virus and BK virus in human sera","volume":"26","author":"Taguchi","year":"1982","journal-title":"Microbiol. Immunol."},{"key":"ref_141","doi-asserted-by":"crossref","first-page":"1737","DOI":"10.1128\/CVI.05175-11","article-title":"Age-specific seroprevalence of Merkel cell polyomavirus, BK virus, and JC virus","volume":"18","author":"Viscidi","year":"2011","journal-title":"Clin. Vaccine Immunol."},{"key":"ref_142","doi-asserted-by":"crossref","first-page":"553","DOI":"10.1016\/j.jcv.2013.08.035","article-title":"JC and BK polyomavirus-specific immunoglobulin G responses in patients thymectomized in early childhood","volume":"58","author":"Prelog","year":"2013","journal-title":"J. Clin. Virol."},{"key":"ref_143","doi-asserted-by":"crossref","first-page":"1560","DOI":"10.1002\/jmv.23841","article-title":"Seroprevalence rates of BKV, JCV, and MCPyV polyomaviruses in the general Czech Republic population","volume":"86","author":"Sroller","year":"2014","journal-title":"J. Med. Virol."},{"key":"ref_144","doi-asserted-by":"crossref","first-page":"498","DOI":"10.1080\/13550280390218887","article-title":"Analysis of the excreted JC virus strains and their potential oral transmission","volume":"9","author":"Major","year":"2003","journal-title":"J. Neurovirol."},{"key":"ref_145","doi-asserted-by":"crossref","first-page":"692","DOI":"10.1099\/vir.0.036798-0","article-title":"Individuals infected with JC polyomavirus do not present detectable JC virus DNA in oropharyngeal fluids","volume":"93","author":"Matos","year":"2012","journal-title":"J. Gen. Virol."},{"key":"ref_146","doi-asserted-by":"crossref","first-page":"559","DOI":"10.1128\/JVI.02777-13","article-title":"Replication of oral BK virus in human salivary gland cells","volume":"88","author":"Madden","year":"2014","journal-title":"J. Virol."},{"key":"ref_147","doi-asserted-by":"crossref","first-page":"1459","DOI":"10.1002\/jmv.23319","article-title":"Dynamics of urinary polyomavirus shedding in healthy adult women","volume":"84","author":"Kling","year":"2012","journal-title":"J. Med. Virol."},{"key":"ref_148","doi-asserted-by":"crossref","first-page":"e02374-17","DOI":"10.1128\/AEM.02374-17","article-title":"Fate of the Urinary Tract Virus BK Human Polyomavirus in Source-Separated Urine","volume":"84","author":"Goetsch","year":"2018","journal-title":"Appl. Environ. Microbiol."},{"key":"ref_149","doi-asserted-by":"crossref","first-page":"63","DOI":"10.1016\/j.virol.2012.12.015","article-title":"Human polyomaviruses in disease and cancer","volume":"437","author":"Dalianis","year":"2013","journal-title":"Virology"},{"key":"ref_150","doi-asserted-by":"crossref","first-page":"56","DOI":"10.1016\/j.jcv.2018.02.015","article-title":"Reactivation of BK polyomavirus during pregnancy, vertical transmission, and clinical significance: A meta-analysis","volume":"102","author":"Cheungpasitporn","year":"2018","journal-title":"J. Clin. 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