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This form usually harbors recombinations in the non-coding control region (NCCR), a key region that contains sequences regulating viral replication and containing binding sites for cellular transcription factors. This form also contains mutations in the capsid protein, especially VP1. Due to the diversity of the JCPyV, a natural polymorphism also exists between the different genotypes. In this review, we aimed to summarize the main features of the archetype and prototype strains in order to facilitate the interpretation of sequence data that are increasingly generated by new sequencing technologies. This will also help to distinguish mutations associated with the natural polymorphism from those specific to the prototype form.<\/jats:p>","DOI":"10.3390\/v18030378","type":"journal-article","created":{"date-parts":[[2026,3,18]],"date-time":"2026-03-18T09:10:04Z","timestamp":1773825004000},"page":"378","update-policy":"https:\/\/doi.org\/10.3390\/mdpi_crossmark_policy","source":"Crossref","is-referenced-by-count":0,"title":["Genetic Diversity of the Polyomavirus JC and Implications for the Pathogenesis of Progressive Multifocal Leukoencephalopathy"],"prefix":"10.3390","volume":"18","author":[{"given":"Michael P.","family":"Wilczek","sequence":"first","affiliation":[{"name":"Life Science, Health, and Engineering Department, The Roux Institute, Northeastern University, Portland, ME 04101, USA"},{"name":"Observational Health Data Sciences and Informatics Center, The Roux Institute, Northeastern University, Portland, ME 04101, USA"},{"name":"Department of Chemistry and Chemical Biology, College of Science, Northeastern University, Boston, MA 02115, USA"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-7908-1576","authenticated-orcid":false,"given":"Sebastien","family":"Lhomme","sequence":"additional","affiliation":[{"name":"Department of Virology, Toulouse University Hospital, 31300 Toulouse, France"},{"name":"Toulouse Institute for Infectious and Inflammatory Diseases, INSERM UMR1291-CNRS UMR5051-Toulouse III University, 31300 Toulouse, France"},{"name":"NEO-I3D Research Group, Toulouse University Hospital, 31300 Toulouse, France"}],"role":[{"role":"author","vocabulary":"crossref"}]}],"member":"1968","published-online":{"date-parts":[[2026,3,18]]},"reference":[{"key":"ref_1","doi-asserted-by":"crossref","first-page":"2429","DOI":"10.1007\/s00705-022-05516-5","article-title":"Recent changes to virus taxonomy ratified by the International Committee on Taxonomy of Viruses (2022)","volume":"167","author":"Walker","year":"2022","journal-title":"Arch. Virol."},{"key":"ref_2","doi-asserted-by":"crossref","first-page":"104150","DOI":"10.1016\/j.meegid.2019.104150","article-title":"Evolution and molecular epidemiology of polyomaviruses","volume":"79","author":"Torres","year":"2020","journal-title":"Infect. Genet. Evol."},{"key":"ref_3","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_4","doi-asserted-by":"crossref","first-page":"10","DOI":"10.1128\/JVI.00561-18","article-title":"JC Polyomavirus: Let\u2019s Please Respect Privacy","volume":"92","author":"Imperiale","year":"2018","journal-title":"J. Virol."},{"key":"ref_5","doi-asserted-by":"crossref","unstructured":"Kean, J.M., Rao, S., Wang, M., and Garcea, R.L. (2009). Seroepidemiology of human polyomaviruses. PLoS Pathog., 5.","DOI":"10.1371\/journal.ppat.1000363"},{"key":"ref_6","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_7","first-page":"112","article-title":"Age-Dependent Seroprevalence of JCV Antibody in Children","volume":"47","author":"Hennes","year":"2016","journal-title":"Neuropediatrics"},{"key":"ref_8","first-page":"93","article-title":"Progressive multifocal leuko-encephalopathy; a hitherto unrecognized complication of chronic lymphatic leukaemia and Hodgkin\u2019s disease","volume":"81","author":"Astrom","year":"1958","journal-title":"Brain"},{"key":"ref_9","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_10","doi-asserted-by":"crossref","first-page":"349","DOI":"10.1093\/brain\/awac237","article-title":"Progressive multifocal leukoencephalopathy: Epidemiology and spectrum of predisposing conditions","volume":"146","author":"Joly","year":"2023","journal-title":"Brain"},{"key":"ref_11","doi-asserted-by":"crossref","first-page":"1691","DOI":"10.2147\/NDT.S114636","article-title":"The use of natalizumab for multiple sclerosis","volume":"13","author":"Brandstadter","year":"2017","journal-title":"Neuropsychiatr. Dis. Treat."},{"key":"ref_12","first-page":"117","article-title":"Stratification and monitoring of natalizumab-associated progressive multifocal leukoencephalopathy risk: Recommendations from an expert group","volume":"87","author":"McGuigan","year":"2016","journal-title":"J. Neurol. Neurosurg. Psychiatry"},{"key":"ref_13","doi-asserted-by":"crossref","first-page":"576","DOI":"10.1002\/ana.20431","article-title":"JC virus granule cell neuronopathy: A novel clinical syndrome distinct from progressive multifocal leukoencephalopathy","volume":"57","author":"Koralnik","year":"2005","journal-title":"Ann. Neurol."},{"key":"ref_14","doi-asserted-by":"crossref","first-page":"775","DOI":"10.1212\/01.WNL.0000081306.86961.33","article-title":"Productive infection of cerebellar granule cell neurons by JC virus in an HIV+ individual","volume":"61","author":"Corey","year":"2003","journal-title":"Neurology"},{"key":"ref_15","doi-asserted-by":"crossref","first-page":"65","DOI":"10.1007\/s00415-014-7530-5","article-title":"Heterogeneous imaging characteristics of JC virus granule cell neuronopathy (GCN): A case series and review of the literature","volume":"262","author":"Wijburg","year":"2015","journal-title":"J. Neurol."},{"key":"ref_16","doi-asserted-by":"crossref","first-page":"742","DOI":"10.1002\/ana.21619","article-title":"Fulminant JC virus encephalopathy with productive infection of cortical pyramidal neurons","volume":"65","author":"Wuthrich","year":"2009","journal-title":"Ann. Neurol."},{"key":"ref_17","doi-asserted-by":"crossref","first-page":"878","DOI":"10.1002\/ana.25608","article-title":"Fatal encephalopathy with wild-type JC virus and ruxolitinib therapy","volume":"86","author":"Reoma","year":"2019","journal-title":"Ann. Neurol."},{"key":"ref_18","doi-asserted-by":"crossref","first-page":"783","DOI":"10.1007\/s13365-017-0558-4","article-title":"Fatal ruxolitinib-related JC virus meningitis","volume":"23","author":"Ballesta","year":"2017","journal-title":"J. Neurovirol."},{"key":"ref_19","doi-asserted-by":"crossref","first-page":"964","DOI":"10.1191\/0961203305lu2229cr","article-title":"JC virus meningitis in a patient with systemic lupus erythematosus","volume":"14","author":"Viallard","year":"2005","journal-title":"Lupus"},{"key":"ref_20","doi-asserted-by":"crossref","first-page":"785","DOI":"10.1001\/jamaophthalmol.2025.2235","article-title":"JC Virus-Related Retinopathy","volume":"143","author":"Varenne","year":"2025","journal-title":"JAMA Ophthalmol."},{"key":"ref_21","doi-asserted-by":"crossref","first-page":"458","DOI":"10.1128\/jvi.51.2.458-469.1984","article-title":"Human polyomavirus JC virus genome","volume":"51","author":"Frisque","year":"1984","journal-title":"J. Virol."},{"key":"ref_22","doi-asserted-by":"crossref","unstructured":"Saribas, S., and Safak, M. (2020). A Comprehensive Proteomics Analysis of the JC Virus (JCV) Large and Small Tumor Antigen Interacting Proteins: Large T Primarily Targets the Host Protein Complexes with V-ATPase and Ubiquitin Ligase Activities While Small t Mostly Associates with Those Having Phosphatase and Chromatin-Remodeling Functions. Viruses, 12.","DOI":"10.3390\/v12101192"},{"key":"ref_23","first-page":"4287","article-title":"Detection of JC virus DNA sequences and expression of the viral regulatory protein T-antigen in tumors of the central nervous system","volume":"61","author":"Gordon","year":"2001","journal-title":"Cancer Res."},{"key":"ref_24","doi-asserted-by":"crossref","first-page":"5496","DOI":"10.1128\/jvi.68.9.5496-5508.1994","article-title":"The kinetics of simian virus 40-induced progression of quiescent cells into S phase depend on four independent functions of large T antigen","volume":"68","author":"Dickmanns","year":"1994","journal-title":"J. Virol."},{"key":"ref_25","doi-asserted-by":"crossref","first-page":"10-1128","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_26","doi-asserted-by":"crossref","unstructured":"Del Valle, L., and Pina-Oviedo, S. (2019). Human Polyomavirus JCPyV and Its Role in Progressive Multifocal Leukoencephalopathy and Oncogenesis. Front. Oncol., 9.","DOI":"10.3389\/fonc.2019.00711"},{"key":"ref_27","doi-asserted-by":"crossref","first-page":"475","DOI":"10.1093\/mutage\/gev004","article-title":"Human JC virus small tumour antigen inhibits nucleotide excision repair and sensitises cells to DNA-damaging agents","volume":"30","author":"Huang","year":"2015","journal-title":"Mutagenesis"},{"key":"ref_28","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_29","doi-asserted-by":"crossref","first-page":"3893","DOI":"10.1128\/JVI.80.8.3893-3903.2006","article-title":"Phosphorylation mutants of JC virus agnoprotein are unable to sustain the viral infection cycle","volume":"80","author":"Sariyer","year":"2006","journal-title":"J. Virol."},{"key":"ref_30","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_31","doi-asserted-by":"crossref","first-page":"302","DOI":"10.1080\/13550280152537148","article-title":"Distribution and function of JCV agnoprotein","volume":"7","author":"Okada","year":"2001","journal-title":"J. Neurovirol."},{"key":"ref_32","doi-asserted-by":"crossref","first-page":"66","DOI":"10.1016\/j.virol.2006.01.025","article-title":"Human polyomavirus JCV late leader peptide region contains important regulatory elements","volume":"349","author":"Akan","year":"2006","journal-title":"Virology"},{"key":"ref_33","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_34","doi-asserted-by":"crossref","first-page":"10846","DOI":"10.1128\/JVI.00542-09","article-title":"Regulation of gene expression in primate polyomaviruses","volume":"83","author":"White","year":"2009","journal-title":"J. Virol."},{"key":"ref_35","doi-asserted-by":"crossref","first-page":"20180301","DOI":"10.1098\/rstb.2018.0301","article-title":"Intra-patient viral evolution in polyomavirus-related diseases","volume":"374","author":"McIlroy","year":"2019","journal-title":"Philos. Trans. R. Soc. Lond. B Biol. Sci."},{"key":"ref_36","doi-asserted-by":"crossref","unstructured":"Wilczek, M.P., Pike, A.M.C., Craig, S.E., Maginnis, M.S., and King, B.L. (2022). Rearrangement in the Hypervariable Region of JC Polyomavirus Genomes Isolated from Patient Samples and Impact on Transcription Factor-Binding Sites and Disease Outcomes. Int. J. Mol. Sci., 23.","DOI":"10.3390\/ijms23105699"},{"key":"ref_37","doi-asserted-by":"crossref","first-page":"237","DOI":"10.1093\/infdis\/jir256","article-title":"Sequencing and analysis of JC virus DNA from natalizumab-treated PML patients","volume":"204","author":"Reid","year":"2011","journal-title":"J. Infect. Dis."},{"key":"ref_38","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_39","doi-asserted-by":"crossref","first-page":"105652","DOI":"10.1016\/j.jcv.2024.105652","article-title":"Differentiation of highly pathogenic strains of human JC polyomavirus in neurological patients by next generation sequencing","volume":"171","author":"Auvinen","year":"2024","journal-title":"J. Clin. Virol."},{"key":"ref_40","doi-asserted-by":"crossref","first-page":"197807","DOI":"10.1155\/2013\/197807","article-title":"Polyomavirus JC in the context of immunosuppression: A series of adaptive, DNA replication-driven recombination events in the development of progressive multifocal leukoencephalopathy","volume":"2013","author":"Johnson","year":"2013","journal-title":"Clin. Dev. Immunol."},{"key":"ref_41","doi-asserted-by":"crossref","unstructured":"Van Loy, T., Thys, K., Tritsmans, L., and Stuyver, L.J. (2013). Quasispecies analysis of JC virus DNA present in urine of healthy subjects. PLoS ONE, 8.","DOI":"10.1371\/journal.pone.0070950"},{"key":"ref_42","doi-asserted-by":"crossref","first-page":"51","DOI":"10.1080\/13550280590900382","article-title":"A case of a progressive multifocal leukoencephalopathy patient with four different JC virus transcriptional control region rearrangements in cerebrospinal fluid, blood, serum, and urine","volume":"11","author":"Delbue","year":"2005","journal-title":"J. Neurovirol."},{"key":"ref_43","doi-asserted-by":"crossref","first-page":"1340","DOI":"10.1128\/JVI.02565-14","article-title":"JC virus quasispecies analysis reveals a complex viral population underlying progressive multifocal leukoencephalopathy and supports viral dissemination via the hematogenous route","volume":"89","author":"Thys","year":"2015","journal-title":"J. Virol."},{"key":"ref_44","doi-asserted-by":"crossref","first-page":"1151","DOI":"10.1093\/infdis\/jiab639","article-title":"JC Polyomavirus Whole Genome Sequencing at the Single-Molecule Level Reveals Emerging Neurotropic Populations in Progressive Multifocal Leukoencephalopathy","volume":"226","author":"Cokelaer","year":"2022","journal-title":"J. Infect. Dis."},{"key":"ref_45","first-page":"889","article-title":"Single-Molecule Sequencing Revealing the Presence of Distinct JC Polyomavirus Populations in Patients With Progressive Multifocal Leukoencephalopathy","volume":"215","author":"Seppala","year":"2017","journal-title":"J. Infect. Dis."},{"key":"ref_46","doi-asserted-by":"crossref","first-page":"639","DOI":"10.1007\/s00705-012-1532-3","article-title":"Sequential changes in the non-coding control region sequences of JC polyomaviruses from the cerebrospinal fluid of patients with progressive multifocal leukoencephalopathy","volume":"158","author":"Nakamichi","year":"2013","journal-title":"Arch. Virol."},{"key":"ref_47","doi-asserted-by":"crossref","first-page":"274","DOI":"10.1007\/s13365-013-0167-9","article-title":"HIV-associated progressive multifocal leukoencephalopathy: Longitudinal study of JC virus non-coding control region rearrangements and host immunity","volume":"19","author":"Iannetta","year":"2013","journal-title":"J. Neurovirol."},{"key":"ref_48","doi-asserted-by":"crossref","first-page":"1880","DOI":"10.1007\/s00415-018-8931-7","article-title":"Detection of JC virus archetype in cerebrospinal fluid in a MS patient with dimethylfumarate treatment without lymphopenia or signs of PML","volume":"265","author":"Motte","year":"2018","journal-title":"J. Neurol."},{"key":"ref_49","doi-asserted-by":"crossref","first-page":"1617","DOI":"10.1212\/01.WNL.0000096147.47128.7D","article-title":"Comparison of PCR-amplified JC virus control region sequences from multiple brain regions in PML","volume":"61","author":"Yasuda","year":"2003","journal-title":"Neurology"},{"key":"ref_50","doi-asserted-by":"crossref","first-page":"42866","DOI":"10.1038\/s41598-025-26996-w","article-title":"Molecular characterization of JC virus in progressive multifocal leukoencephalopathy cases from India","volume":"15","author":"Palani","year":"2025","journal-title":"Sci. Rep."},{"key":"ref_51","doi-asserted-by":"crossref","first-page":"1499","DOI":"10.1099\/0022-1317-74-8-1499","article-title":"Human polyomavirus JC promoter\/enhancer rearrangement patterns from progressive multifocal leukoencephalopathy brain are unique derivatives of a single archetypal structure","volume":"74","author":"Ault","year":"1993","journal-title":"J. Gen. Virol."},{"key":"ref_52","doi-asserted-by":"crossref","first-page":"5062","DOI":"10.1073\/pnas.90.11.5062","article-title":"Origin of JC polyomavirus variants associated with progressive multifocal leukoencephalopathy","volume":"90","author":"Iida","year":"1993","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_53","doi-asserted-by":"crossref","first-page":"72","DOI":"10.1016\/0042-6822(92)90167-N","article-title":"Evidence of human polyomavirus BK and JC infection in normal brain tissue","volume":"191","author":"Elsner","year":"1992","journal-title":"Virology"},{"key":"ref_54","doi-asserted-by":"crossref","first-page":"139","DOI":"10.1097\/00042560-199606010-00006","article-title":"JCV-DNA and BKV-DNA in the CNS tissue and CSF of AIDS patients and normal subjects. Study of 41 cases and review of the literature","volume":"12","author":"Vago","year":"1996","journal-title":"J. Acquir. Immune Defic. Syndr. Hum. Retrovirol."},{"key":"ref_55","doi-asserted-by":"crossref","first-page":"5726","DOI":"10.1128\/jvi.66.10.5726-5734.1992","article-title":"JC virus DNA is present in many human brain samples from patients without progressive multifocal leukoencephalopathy","volume":"66","author":"White","year":"1992","journal-title":"J. Virol."},{"key":"ref_56","doi-asserted-by":"crossref","unstructured":"Lauver, M.D., and Lukacher, A.E. (2020). JCPyV VP1 Mutations in Progressive MultifocalLeukoencephalopathy: Altering Tropismor Mediating Immune Evasion?. Viruses, 12.","DOI":"10.3390\/v12101156"},{"key":"ref_57","doi-asserted-by":"crossref","first-page":"2173","DOI":"10.1007\/s007050070047","article-title":"Lack of disease-specific amino acid changes in the viral proteins of JC virus isolates from the brain with progressive multifocal leukoencephalopathy","volume":"145","author":"Kato","year":"2000","journal-title":"Arch. Virol."},{"key":"ref_58","doi-asserted-by":"crossref","unstructured":"Sunyaev, S.R., Lugovskoy, A., Simon, K., and Gorelik, L. (2009). Adaptive mutations in the JC virus protein capsid are associated with progressive multifocal leukoencephalopathy (PML). PLoS Genet., 5.","DOI":"10.1371\/journal.pgen.1000368"},{"key":"ref_59","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_60","doi-asserted-by":"crossref","first-page":"e00247-13","DOI":"10.1128\/mBio.00247-13","article-title":"Progressive multifocal leukoencephalopathy-associated mutations in the JC polyomavirus capsid disrupt lactoseries tetrasaccharide c binding","volume":"4","author":"Maginnis","year":"2013","journal-title":"mBio"},{"key":"ref_61","doi-asserted-by":"crossref","first-page":"175","DOI":"10.1099\/vir.0.037440-0","article-title":"JC virus granule cell neuronopathy is associated with VP1 C terminus mutants","volume":"93","author":"Dang","year":"2012","journal-title":"J. Gen. Virol."},{"key":"ref_62","doi-asserted-by":"crossref","first-page":"339","DOI":"10.1080\/13550280152537201","article-title":"Predicted amino acid sequences for 100 JCV strains","volume":"7","author":"Cubitt","year":"2001","journal-title":"J. Neurovirol."},{"key":"ref_63","doi-asserted-by":"crossref","first-page":"10-1128","DOI":"10.1128\/JVI.01335-16","article-title":"Deep-Sequence Identification and Role in Virus Replication of a JC Virus Quasispecies in Patients with Progressive Multifocal Leukoencephalopathy","volume":"91","author":"Takahashi","year":"2017","journal-title":"J. Virol."},{"key":"ref_64","doi-asserted-by":"crossref","unstructured":"Dang, X., Wuthrich, C., Gordon, J., Sawa, H., and Koralnik, I.J. (2012). JC virus encephalopathy is associated with a novel agnoprotein-deletion JCV variant. PLoS ONE, 7.","DOI":"10.1371\/journal.pone.0035793"},{"key":"ref_65","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_66","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_67","doi-asserted-by":"crossref","first-page":"269","DOI":"10.1006\/viro.1996.8409","article-title":"Propagation of the human polyomavirus, JCV, in human neuroblastoma cell lines","volume":"228","author":"Shinohara","year":"1997","journal-title":"Virology"},{"key":"ref_68","doi-asserted-by":"crossref","first-page":"97","DOI":"10.1016\/j.virusres.2009.09.005","article-title":"Modulation of JC virus transcription by C\/EBPbeta","volume":"146","author":"Romagnoli","year":"2009","journal-title":"Virus Res."},{"key":"ref_69","doi-asserted-by":"crossref","first-page":"2035","DOI":"10.1099\/vir.0.80863-0","article-title":"New sequence polymorphisms in the outer loops of the JC polyomavirus major capsid protein (VP1) possibly associated with progressive multifocal leukoencephalopathy","volume":"86","author":"Zheng","year":"2005","journal-title":"J. Gen. Virol."},{"key":"ref_70","doi-asserted-by":"crossref","first-page":"51","DOI":"10.1080\/13550280802425467","article-title":"JC virus VP1 loop-specific polymorphisms are associated with favorable prognosis for progressive multifocal leukoencephalopathy","volume":"15","author":"Delbue","year":"2009","journal-title":"J. Neurovirol."},{"key":"ref_71","doi-asserted-by":"crossref","first-page":"213","DOI":"10.1086\/317927","article-title":"JC virus genotypes in France: Molecular epidemiology and potential significance for progressive multifocal leukoencephalopathy","volume":"183","author":"Dubois","year":"2001","journal-title":"J. Infect. Dis."},{"key":"ref_72","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. 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