{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,7,13]],"date-time":"2026-07-13T15:17:45Z","timestamp":1783955865545,"version":"3.55.0"},"reference-count":73,"publisher":"Springer Science and Business Media LLC","issue":"1","license":[{"start":{"date-parts":[[2023,1,8]],"date-time":"2023-01-08T00:00:00Z","timestamp":1673136000000},"content-version":"tdm","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0"},{"start":{"date-parts":[[2023,1,8]],"date-time":"2023-01-08T00:00:00Z","timestamp":1673136000000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0"}],"funder":[{"DOI":"10.13039\/100006229","name":"Oak Ridge Institute for Science and Education","doi-asserted-by":"publisher","id":[{"id":"10.13039\/100006229","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/100000002","name":"National Institutes of Health","doi-asserted-by":"crossref","id":[{"id":"10.13039\/100000002","id-type":"DOI","asserted-by":"crossref"}]}],"content-domain":{"domain":["link.springer.com"],"crossmark-restriction":false},"short-container-title":["Sci Rep"],"abstract":"<jats:title>Abstract<\/jats:title><jats:p>The novel coronavirus SARS-CoV-2 continues to cause death and disease throughout the world, underscoring the necessity of understanding the virus and host immune response. From the start of the pandemic, a prominent pattern of central nervous system (CNS) pathologies, including demyelination, has emerged, suggesting an underlying mechanism of viral mimicry to CNS proteins. We hypothesized that immunodominant epitopes of SARS-CoV-2 share homology with proteins associated with multiple sclerosis (MS). Using PEPMatch, a newly developed bioinformatics package which predicts peptide similarity within specific amino acid mismatching parameters consistent with published MHC binding capacity, we discovered that nucleocapsid protein shares significant overlap with 22 MS-associated proteins, including myelin proteolipid protein (PLP). Further computational evaluation demonstrated that this overlap may have critical implications for T cell responses in MS patients and is likely unique to SARS-CoV-2 among the major human coronaviruses. Our findings substantiate the hypothesis of viral molecular mimicry in the pathogenesis of MS and warrant further experimental exploration.<\/jats:p>","DOI":"10.1038\/s41598-022-27348-8","type":"journal-article","created":{"date-parts":[[2023,1,8]],"date-time":"2023-01-08T11:02:27Z","timestamp":1673175747000},"update-policy":"https:\/\/doi.org\/10.1007\/springer_crossmark_policy","source":"Crossref","is-referenced-by-count":19,"title":["Sequence similarity between SARS-CoV-2 nucleocapsid and multiple sclerosis-associated proteins provides insight into viral neuropathogenesis following infection"],"prefix":"10.1038","volume":"13","author":[{"given":"Camille M.","family":"Lake","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Joseph J.","family":"Breen","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"297","published-online":{"date-parts":[[2023,1,8]]},"reference":[{"key":"27348_CR1","unstructured":"Coronavirus disease (COVID-19)\u2014World Health Organization. https:\/\/www.who.int\/emergencies\/diseases\/novel-coronavirus-2019."},{"key":"27348_CR2","doi-asserted-by":"publisher","first-page":"1062","DOI":"10.1007\/s12028-020-01049-4","volume":"34","author":"G Aghagoli","year":"2021","unstructured":"Aghagoli, G. et al. Neurological involvement in COVID-19 and potential mechanisms: A review. Neurocrit. Care 34, 1062\u20131071 (2021).","journal-title":"Neurocrit. Care"},{"key":"27348_CR3","doi-asserted-by":"publisher","DOI":"10.1001\/jamanetworkopen.2021.11417","volume":"4","author":"T Nasserie","year":"2021","unstructured":"Nasserie, T., Hittle, M. & Goodman, S. N. Assessment of the frequency and variety of persistent symptoms among patients with COVID-19: A Systematic review. JAMA Netw. Open 4, e2111417 (2021).","journal-title":"JAMA Netw. Open"},{"key":"27348_CR4","doi-asserted-by":"publisher","DOI":"10.1007\/s00415-021-10752-x","author":"II Ismail","year":"2021","unstructured":"Ismail, I. I. & Salama, S. Association of CNS demyelination and COVID-19 infection: an updated systematic review. J. Neurol. https:\/\/doi.org\/10.1007\/s00415-021-10752-x (2021).","journal-title":"J. Neurol."},{"key":"27348_CR5","unstructured":"Number of people with MS | Atlas of MS. https:\/\/www.atlasofms.org\/map\/global\/epidemiology\/number-of-people-with-ms."},{"key":"27348_CR6","first-page":"95","volume":"2","author":"R Liblau","year":"2000","unstructured":"Liblau, R. & Gautam, A. M. HLA, molecular mimicry and multiple sclerosis. Rev. Immunogenet. 2, 95\u2013104 (2000).","journal-title":"Rev. Immunogenet."},{"key":"27348_CR7","doi-asserted-by":"publisher","first-page":"185","DOI":"10.1056\/NEJMcibr035136","volume":"349","author":"H Wekerle","year":"2003","unstructured":"Wekerle, H. & Hohlfeld, R. Molecular mimicry in multiple sclerosis. N. Engl. J. Med. 349, 185\u2013186 (2003).","journal-title":"N. Engl. J. Med."},{"key":"27348_CR8","doi-asserted-by":"publisher","first-page":"227","DOI":"10.1111\/j.1600-065X.2011.01076.x","volume":"245","author":"EML Chastain","year":"2012","unstructured":"Chastain, E. M. L. & Miller, S. D. Molecular mimicry as an inducing trigger for CNS autoimmune demyelinating disease. Immunol. Rev. 245, 227\u2013238 (2012).","journal-title":"Immunol. Rev."},{"key":"27348_CR9","doi-asserted-by":"crossref","unstructured":"Dykema, A. G. et al. Functional characterization of CD4+ T cell receptors crossreactive for SARS-CoV-2 and endemic coronaviruses. J. Clin. Investig. 131, (2021).","DOI":"10.1172\/JCI146922"},{"key":"27348_CR10","doi-asserted-by":"publisher","first-page":"2574","DOI":"10.1056\/NEJMc2009191","volume":"382","author":"G Toscano","year":"2020","unstructured":"Toscano, G. et al. Guillain\u2013Barr\u00e9 syndrome associated with SARS-CoV-2. N. Engl. J. Med. 382, 2574\u20132576 (2020).","journal-title":"N. Engl. J. Med."},{"key":"27348_CR11","doi-asserted-by":"publisher","first-page":"2719","DOI":"10.4049\/jimmunol.169.5.2719","volume":"169","author":"JK Olson","year":"2002","unstructured":"Olson, J. K., Eagar, T. N. & Miller, S. D. Functional activation of myelin-specific T cells by virus-induced molecular mimicry. J. Immunol. 169, 2719\u20132726 (2002).","journal-title":"J. Immunol."},{"key":"27348_CR12","doi-asserted-by":"publisher","first-page":"2031","DOI":"10.1002\/eji.200939568","volume":"39","author":"G Krishnamoorthy","year":"2009","unstructured":"Krishnamoorthy, G. & Wekerle, H. EAE: An immunologist\u2019s magic eye. Eur. J. Immunol. 39, 2031\u20132035 (2009).","journal-title":"Eur. J. Immunol."},{"key":"27348_CR13","doi-asserted-by":"publisher","first-page":"6693","DOI":"10.4049\/jimmunol.166.11.6693","volume":"166","author":"SK Joshi","year":"2001","unstructured":"Joshi, S. K., Suresh, P. R. & Chauhan, V. S. Flexibility in MHC and TCR recognition: Degenerate specificity at the T cell level in the recognition of promiscuous th epitopes exhibiting no primary sequence homology. J. Immunol. 166, 6693\u20136703 (2001).","journal-title":"J. Immunol."},{"key":"27348_CR14","doi-asserted-by":"publisher","first-page":"940","DOI":"10.1038\/ni835","volume":"3","author":"HLE Lang","year":"2002","unstructured":"Lang, H. L. E. et al. A functional and structural basis for TCR cross-reactivity in multiple sclerosis. Nat. Immunol. 3, 940\u2013943 (2002).","journal-title":"Nat. Immunol."},{"key":"27348_CR15","doi-asserted-by":"publisher","first-page":"2346","DOI":"10.1073\/pnas.80.8.2346","volume":"80","author":"RS Fujinami","year":"1983","unstructured":"Fujinami, R. S., Oldstone, M. B., Wroblewska, Z., Frankel, M. E. & Koprowski, H. Molecular mimicry in virus infection: crossreaction of measles virus phosphoprotein or of herpes simplex virus protein with human intermediate filaments. PNAS 80, 2346\u20132350 (1983).","journal-title":"PNAS"},{"key":"27348_CR16","doi-asserted-by":"publisher","first-page":"533","DOI":"10.1002\/ana.410270513","volume":"27","author":"A Nath","year":"1990","unstructured":"Nath, A. & Wolinsky, J. S. Antibody response to rubella virus structural proteins in multiple sclerosis. Ann. Neurol. 27, 533\u2013536 (1990).","journal-title":"Ann. Neurol."},{"key":"27348_CR17","doi-asserted-by":"crossref","unstructured":"Marrama, D., Mahita, J., Sette, A. & Peters, B. Lack of evidence of significant homology of SARS-CoV-2 spike sequences to myocarditis-associated antigens. eBioMedicine 75, (2022).","DOI":"10.1016\/j.ebiom.2021.103807"},{"key":"27348_CR18","doi-asserted-by":"publisher","first-page":"982","DOI":"10.4049\/jimmunol.155.2.982","volume":"155","author":"S Markovic-Plese","year":"1995","unstructured":"Markovic-Plese, S. et al. T cell recognition of immunodominant and cryptic proteolipid protein epitopes in humans. J. Immunol. 155, 982\u2013992 (1995).","journal-title":"J. Immunol."},{"key":"27348_CR19","doi-asserted-by":"publisher","first-page":"270","DOI":"10.1177\/135245859600100503","volume":"1","author":"C Pelfrey","year":"1996","unstructured":"Pelfrey, C., Tranquill, L., Vogt, A. & McFarland, H. T cell response to two immunodominant proteolipid protein (PLP) peptides in multiple sclerosis patients and healthy controls. Mult. Scler. 1, 270\u2013278 (1996).","journal-title":"Mult. Scler."},{"key":"27348_CR20","doi-asserted-by":"publisher","first-page":"172","DOI":"10.1016\/S0165-5728(97)00260-9","volume":"84","author":"JL Trotter","year":"1998","unstructured":"Trotter, J. L. et al. T cell recognition of myelin proteolipid protein and myelin proteolipid protein peptides in the peripheral blood of multiple sclerosis and control subjects. J. Neuroimmunol. 84, 172\u2013178 (1998).","journal-title":"J. Neuroimmunol."},{"key":"27348_CR21","doi-asserted-by":"publisher","first-page":"897","DOI":"10.1016\/j.immuni.2009.09.025","volume":"31","author":"WA Macdonald","year":"2009","unstructured":"Macdonald, W. A. et al. T cell allorecognition via molecular mimicry. Immunity 31, 897\u2013908 (2009).","journal-title":"Immunity"},{"key":"27348_CR22","doi-asserted-by":"publisher","first-page":"742","DOI":"10.1212\/WNL.51.3.742","volume":"51","author":"BG Weinshenker","year":"1998","unstructured":"Weinshenker, B. G. et al. Major histocompatibility complex class II alleles and the course and outcome of MS: A population-based study. Neurology 51, 742\u2013747 (1998).","journal-title":"Neurology"},{"key":"27348_CR23","doi-asserted-by":"crossref","unstructured":"Zivadinov, R. et al. HLA\u2010DRB1*1501, \u2010DQB1*0301, \u2010DQB1*0302, \u2010DQB1*0602, and \u2010DQB1*0603 Alleles are Associated With More Severe Disease Outcome on Mri in Patients With Multiple Sclerosis. in International Review of Neurobiology vol. 79 521\u2013535 (Academic Press, 2007).","DOI":"10.1016\/S0074-7742(07)79023-2"},{"key":"27348_CR24","doi-asserted-by":"publisher","DOI":"10.1371\/journal.pgen.1003926","volume":"9","author":"NA Patsopoulos","year":"2013","unstructured":"Patsopoulos, N. A. et al. Fine-mapping the genetic association of the major histocompatibility complex in multiple sclerosis: HLA and non-HLA effects. PLoS Genet 9, e1003926 (2013).","journal-title":"PLoS Genet"},{"key":"27348_CR25","doi-asserted-by":"publisher","first-page":"325","DOI":"10.1007\/s00251-011-0513-0","volume":"63","author":"J Greenbaum","year":"2011","unstructured":"Greenbaum, J. et al. Functional classification of class II human leukocyte antigen (HLA) molecules reveals seven different supertypes and a surprising degree of repertoire sharing across supertypes. Immunogenetics 63, 325\u2013335 (2011).","journal-title":"Immunogenetics"},{"key":"27348_CR26","unstructured":"Selecting thresholds (cut-offs) for MHC class I and II binding predictions. IEDB Solutions Center https:\/\/help.iedb.org\/hc\/en-us\/articles\/114094151811-Selecting-thresholds-cut-offs-for-MHC-class-I-and-II-binding-predictions."},{"key":"27348_CR27","doi-asserted-by":"publisher","first-page":"125","DOI":"10.1016\/0168-1702(92)90039-C","volume":"22","author":"P Jouvenne","year":"1992","unstructured":"Jouvenne, P., Mounir, S., Stewart, J. N., Richardson, C. D. & Talbot, P. J. Sequence analysis of human coronavirus 229E mRNAs 4 and 5: Evidence for polymorphism and homology with myelin basic protein. Virus Res 22, 125\u2013141 (1992).","journal-title":"Virus Res"},{"key":"27348_CR28","doi-asserted-by":"publisher","first-page":"525","DOI":"10.1002\/ana.410310511","volume":"31","author":"RS Murray","year":"1992","unstructured":"Murray, R. S., Brown, B., Brain, D. & Cabirac, G. F. Detection of coronavirus RNA and antigen in multiple sclerosis brain. Ann. Neurol. 31, 525\u2013533 (1992).","journal-title":"Ann. Neurol."},{"key":"27348_CR29","doi-asserted-by":"publisher","DOI":"10.1007\/0-387-25518-4_43","author":"PJ Talbot","year":"2005","unstructured":"Talbot, P. J., Boucher, A., Duquette, P. & Gruslin, E. Coronaviruses and Neuroantigens: myelin proteins, myelin genes. Exp. Models Mult. Scler. https:\/\/doi.org\/10.1007\/0-387-25518-4_43 (2005).","journal-title":"Exp. Models Mult. Scler."},{"key":"27348_CR30","doi-asserted-by":"publisher","first-page":"767","DOI":"10.1016\/S1474-4422(20)30221-0","volume":"19","author":"MA Ellul","year":"2020","unstructured":"Ellul, M. A. et al. Neurological associations of COVID-19. Lancet Neurol. 19, 767\u2013783 (2020).","journal-title":"Lancet Neurol."},{"key":"27348_CR31","doi-asserted-by":"publisher","first-page":"18","DOI":"10.1016\/j.bbi.2020.03.031","volume":"87","author":"Y Wu","year":"2020","unstructured":"Wu, Y. et al. Nervous system involvement after infection with COVID-19 and other coronaviruses. Brain Behav. Immun. 87, 18\u201322 (2020).","journal-title":"Brain Behav. Immun."},{"key":"27348_CR32","doi-asserted-by":"publisher","unstructured":"Thieme, C. et al. The SARS-COV-2 T-Cell Immunity is Directed Against the Spike, Membrane, and Nucleocapsid Protein and Associated with COVID 19 Severity. https:\/\/papers.ssrn.com\/abstract=3606763 (2020). https:\/\/doi.org\/10.2139\/ssrn.3606763.","DOI":"10.2139\/ssrn.3606763"},{"key":"27348_CR33","doi-asserted-by":"publisher","first-page":"1489","DOI":"10.1016\/j.cell.2020.05.015","volume":"181","author":"A Grifoni","year":"2020","unstructured":"Grifoni, A. et al. Targets of T cell responses to SARS-CoV-2 coronavirus in humans with COVID-19 disease and unexposed individuals. Cell 181, 1489-1501.e15 (2020).","journal-title":"Cell"},{"key":"27348_CR34","doi-asserted-by":"publisher","DOI":"10.1016\/j.xcrm.2021.100204","volume":"2","author":"A Tarke","year":"2021","unstructured":"Tarke, A. et al. Comprehensive analysis of T cell immunodominance and immunoprevalence of SARS-CoV-2 epitopes in COVID-19 cases. Cell Rep. Med. 2, 100204 (2021).","journal-title":"Cell Rep. Med."},{"key":"27348_CR35","doi-asserted-by":"publisher","first-page":"2554","DOI":"10.1038\/s41423-021-00767-9","volume":"18","author":"SC Jordan","year":"2021","unstructured":"Jordan, S. C. et al. T cell immune responses to SARS-CoV-2 and variants of concern (Alpha and Delta) in infected and vaccinated individuals. Cell Mol. Immunol. 18, 2554\u20132556 (2021).","journal-title":"Cell Mol. Immunol."},{"key":"27348_CR36","doi-asserted-by":"publisher","unstructured":"Gangaev, A. et al. Profound CD8 T cell responses towards the SARS-CoV-2 ORF1ab in COVID-19 patients. https:\/\/www.researchsquare.com\/article\/rs-33197\/v1 (2020). https:\/\/doi.org\/10.21203\/rs.3.rs-33197\/v1.","DOI":"10.21203\/rs.3.rs-33197\/v1"},{"key":"27348_CR37","doi-asserted-by":"publisher","first-page":"182","DOI":"10.1016\/j.micinf.2020.05.013","volume":"22","author":"B Tilocca","year":"2020","unstructured":"Tilocca, B. et al. Immunoinformatic analysis of the SARS-CoV-2 envelope protein as a strategy to assess cross-protection against COVID-19. Microbes Infect. 22, 182\u2013187 (2020).","journal-title":"Microbes Infect."},{"key":"27348_CR38","doi-asserted-by":"publisher","first-page":"266","DOI":"10.1016\/0014-5793(86)81502-2","volume":"207","author":"SY Shaw","year":"1986","unstructured":"Shaw, S. Y., Laursen, R. A. & Lees, M. B. Analogous amino acid sequences in myelin proteolipid and viral proteins. FEBS Lett. 207, 266\u2013270 (1986).","journal-title":"FEBS Lett."},{"key":"27348_CR39","doi-asserted-by":"publisher","first-page":"5666","DOI":"10.4049\/jimmunol.150.12.5666","volume":"150","author":"S Amor","year":"1993","unstructured":"Amor, S., Baker, D., Groome, N. & Turk, J. L. Identification of a major encephalitogenic epitope of proteolipid protein (residues 56\u201370) for the induction of experimental allergic encephalomyelitis in Biozzi AB\/H and nonobese diabetic mice. J. Immunol. 150, 5666\u20135672 (1993).","journal-title":"J. Immunol."},{"key":"27348_CR40","doi-asserted-by":"publisher","first-page":"1447","DOI":"10.1093\/brain\/120.8.1447","volume":"120","author":"J Greer","year":"1997","unstructured":"Greer, J. Increased immunoreactivity to two overlapping peptides of myelin proteolipid protein in multiple sclerosis. Brain 120, 1447\u20131460 (1997).","journal-title":"Brain"},{"key":"27348_CR41","doi-asserted-by":"crossref","unstructured":"Dayhoff, M. O., Barker, W. C. & Hunt, L. T. [47] Establishing homologies in protein sequences. in Methods in Enzymology vol. 91 524\u2013545 (Academic Press, 1983).","DOI":"10.1016\/S0076-6879(83)91049-2"},{"key":"27348_CR42","doi-asserted-by":"publisher","unstructured":"Fran\u00e7a, L. C. et al. Molecular Mimicry between Zika virus and central nervous system inflammatory demyelinating disorders: the role of NS5 Zika virus epitope and PLP autoantigens. at https:\/\/doi.org\/10.21203\/rs.2.22564\/v1 (2022).","DOI":"10.21203\/rs.2.22564\/v1"},{"key":"27348_CR43","doi-asserted-by":"publisher","first-page":"83","DOI":"10.1016\/S1074-7613(02)00340-0","volume":"17","author":"X He","year":"2002","unstructured":"He, X. et al. Structural snapshot of aberrant antigen presentation linked to autoimmunity: The immunodominant epitope of MBP complexed with I-Au. Immunity 17, 83\u201394 (2002).","journal-title":"Immunity"},{"key":"27348_CR44","unstructured":"IPD-IMGT\/HLA Database. https:\/\/www.ebi.ac.uk\/ipd\/imgt\/hla\/alleles\/."},{"key":"27348_CR45","doi-asserted-by":"publisher","first-page":"247","DOI":"10.1038\/gene.2014.82","volume":"16","author":"NMF van Gerven","year":"2015","unstructured":"van Gerven, N. M. F. et al. HLA-DRB1*03:01 and HLA-DRB1*04:01 modify the presentation and outcome in autoimmune hepatitis type-1. Genes Immun 16, 247\u2013252 (2015).","journal-title":"Genes Immun"},{"key":"27348_CR46","doi-asserted-by":"publisher","DOI":"10.1016\/j.jneuroim.2020.577243","volume":"344","author":"F Hu","year":"2020","unstructured":"Hu, F. et al. Novel findings of HLA association with anti-LGI1 encephalitis: HLA-DRB1*03:01 and HLA-DQB1*02:01. J. Neuroimmunol. 344, 577243 (2020).","journal-title":"J. Neuroimmunol."},{"key":"27348_CR47","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1016\/j.jneuroim.2017.05.018","volume":"310","author":"MP Alvarenga","year":"2017","unstructured":"Alvarenga, M. P. et al. The HLA DRB1*03:01 allele is associated with NMO regardless of the NMO-IgG status in Brazilian patients from Rio de Janeiro. J. Neuroimmunol. 310, 1\u20137 (2017).","journal-title":"J. Neuroimmunol."},{"key":"27348_CR48","doi-asserted-by":"publisher","first-page":"E1703","DOI":"10.1210\/jc.2011-0645","volume":"96","author":"B Skinningsrud","year":"2011","unstructured":"Skinningsrud, B. et al. Multiple loci in the HLA complex are associated with Addison\u2019s disease. J. Clin. Endocrinol. Metab. 96, E1703\u2013E1708 (2011).","journal-title":"J. Clin. Endocrinol. Metab."},{"key":"27348_CR49","doi-asserted-by":"crossref","unstructured":"Bhattarai, P., Allen, H., Aggarwal, A., Madden, D. & Dalton, K. Unmasking of Addison\u2019s disease in COVID-19. SAGE Open Med. Case Rep. 9, 2050313X211027758 (2021).","DOI":"10.1177\/2050313X211027758"},{"key":"27348_CR50","doi-asserted-by":"publisher","unstructured":"O\u011fuz, S. H. & G\u00fcrlek, A. Emergence of Autoimmune Type 1 Diabetes and Acute Adrenal Crisis Following COVID-19. https:\/\/www.researchsquare.com\/article\/rs-1015570\/v1 (2021). https:\/\/doi.org\/10.21203\/rs.3.rs-1015570\/v1.","DOI":"10.21203\/rs.3.rs-1015570\/v1"},{"key":"27348_CR51","doi-asserted-by":"publisher","DOI":"10.1016\/j.aace.2021.11.001","author":"J S\u00e1nchez","year":"2021","unstructured":"S\u00e1nchez, J., Cohen, M., Zapater, J. L. & Eisenberg, Y. Primary adrenal insufficiency after COVID-19 infection. AACE Clin. Case Rep. https:\/\/doi.org\/10.1016\/j.aace.2021.11.001 (2021).","journal-title":"AACE Clin. Case Rep."},{"key":"27348_CR52","doi-asserted-by":"publisher","first-page":"14","DOI":"10.1111\/tan.14284","volume":"98","author":"DJ Langton","year":"2021","unstructured":"Langton, D. J. et al. The influence of HLA genotype on the severity of COVID-19 infection. HLA 98, 14\u201322 (2021).","journal-title":"HLA"},{"key":"27348_CR53","doi-asserted-by":"publisher","first-page":"84","DOI":"10.1186\/s40001-021-00563-1","volume":"26","author":"F Migliorini","year":"2021","unstructured":"Migliorini, F. et al. Association between HLA genotypes and COVID-19 susceptibility, severity and progression: a comprehensive review of the literature. Eur. J. Med. Res. 26, 84 (2021).","journal-title":"Eur. J. Med. Res."},{"key":"27348_CR54","doi-asserted-by":"crossref","unstructured":"Tavasolian, F. et al. HLA, Immune response, and susceptibility to COVID-19. Front. Immunol. 11, (2021).","DOI":"10.3389\/fimmu.2020.601886"},{"key":"27348_CR55","doi-asserted-by":"publisher","first-page":"55","DOI":"10.1007\/s12079-017-0445-z","volume":"12","author":"M Pasello","year":"2018","unstructured":"Pasello, M., Manara, M. C. & Scotlandi, K. CD99 at the crossroads of physiology and pathology. J. Cell Commun. Signal 12, 55\u201368 (2018).","journal-title":"J. Cell Commun. Signal"},{"key":"27348_CR56","doi-asserted-by":"publisher","first-page":"2100133","DOI":"10.1002\/pmic.202100133","volume":"21","author":"J Siwy","year":"2021","unstructured":"Siwy, J. et al. CD99 and polymeric immunoglobulin receptor peptides deregulation in critical COVID-19: A potential link to molecular pathophysiology?. Proteomics 21, 2100133 (2021).","journal-title":"Proteomics"},{"key":"27348_CR57","doi-asserted-by":"publisher","DOI":"10.1038\/s41586-022-04421-w","author":"J Di Domizio","year":"2022","unstructured":"Di Domizio, J. et al. The cGAS-STING pathway drives type I IFN immunopathology in COVID-19. Nature https:\/\/doi.org\/10.1038\/s41586-022-04421-w (2022).","journal-title":"Nature"},{"key":"27348_CR58","doi-asserted-by":"publisher","first-page":"1594","DOI":"10.1016\/j.immuni.2021.06.002","volume":"54","author":"M Schwabenland","year":"2021","unstructured":"Schwabenland, M. et al. Deep spatial profiling of human COVID-19 brains reveals neuroinflammation with distinct microanatomical microglia-T-cell interactions. Immunity 54, 1594-1610.e11 (2021).","journal-title":"Immunity"},{"key":"27348_CR59","doi-asserted-by":"publisher","first-page":"119","DOI":"10.1007\/s00441-003-0751-z","volume":"314","author":"B Engelhardt","year":"2003","unstructured":"Engelhardt, B. Development of the blood-brain barrier. Cell Tissue Res. 314, 119\u2013129 (2003).","journal-title":"Cell Tissue Res."},{"key":"27348_CR60","doi-asserted-by":"publisher","first-page":"327","DOI":"10.1083\/jcb.200305032","volume":"163","author":"P Wong","year":"2003","unstructured":"Wong, P. & Coulombe, P. A. Loss of keratin 6 (K6) proteins reveals a function for intermediate filaments during wound repair. J. Cell Biol. 163, 327\u2013337 (2003).","journal-title":"J. Cell Biol."},{"key":"27348_CR61","doi-asserted-by":"crossref","unstructured":"Libbey, J. E., McCoy, L. L. & Fujinami, R. S. Molecular Mimicry in Multiple Sclerosis. in International Review of Neurobiology vol. 79 127\u2013147 (Academic Press, 2007).","DOI":"10.1016\/S0074-7742(07)79006-2"},{"key":"27348_CR62","doi-asserted-by":"publisher","DOI":"10.1126\/science.abj8222","author":"K Bjornevik","year":"2022","unstructured":"Bjornevik, K. et al. Longitudinal analysis reveals high prevalence of Epstein-Barr virus associated with multiple sclerosis. Science https:\/\/doi.org\/10.1126\/science.abj8222 (2022).","journal-title":"Science"},{"key":"27348_CR63","doi-asserted-by":"publisher","DOI":"10.1038\/s41586-022-04432-7","author":"TV Lanz","year":"2022","unstructured":"Lanz, T. V. et al. Clonally expanded B cells in multiple sclerosis bind EBV EBNA1 and GlialCAM. Nature https:\/\/doi.org\/10.1038\/s41586-022-04432-7 (2022).","journal-title":"Nature"},{"key":"27348_CR64","doi-asserted-by":"publisher","first-page":"22","DOI":"10.1186\/s41983-021-00277-5","volume":"57","author":"NM Shalaby","year":"2021","unstructured":"Shalaby, N. M. & Shehata, H. S. Could SARS-CoV-2 herald a surge of multiple sclerosis?. Egypt J. Neurol. Psychiatry Neurosurg. 57, 22 (2021).","journal-title":"Egypt J. Neurol. Psychiatry Neurosurg."},{"key":"27348_CR65","doi-asserted-by":"publisher","DOI":"10.1016\/j.xcrm.2021.100288","volume":"2","author":"E Song","year":"2021","unstructured":"Song, E. et al. Divergent and self-reactive immune responses in the CNS of COVID-19 patients with neurological symptoms. Cell Rep. Med. 2, 100288 (2021).","journal-title":"Cell Rep. Med."},{"key":"27348_CR66","doi-asserted-by":"publisher","first-page":"1382","DOI":"10.1373\/clinchem.2005.051045","volume":"51","author":"Y Liang","year":"2005","unstructured":"Liang, Y. et al. Comprehensive antibody epitope mapping of the nucleocapsid protein of severe acute respiratory syndrome (SARS) coronavirus: Insight into the humoral immunity of SARS. Clin. Chem. 51, 1382\u20131396 (2005).","journal-title":"Clin. Chem."},{"key":"27348_CR67","doi-asserted-by":"publisher","first-page":"45","DOI":"10.1016\/j.bbrc.2021.01.073","volume":"543","author":"VAJ Smits","year":"2021","unstructured":"Smits, V. A. J. et al. The Nucleocapsid protein triggers the main humoral immune response in COVID-19 patients. Biochem. Biophys. Res. Commun. 543, 45\u201349 (2021).","journal-title":"Biochem. Biophys. Res. Commun."},{"key":"27348_CR68","doi-asserted-by":"publisher","first-page":"1490","DOI":"10.4049\/jimmunol.146.5.1490","volume":"146","author":"J Sun","year":"1991","unstructured":"Sun, J. et al. T and B cell responses to myelin-oligodendrocyte glycoprotein in multiple sclerosis. J. Immunol. 146, 1490\u20131495 (1991).","journal-title":"J. Immunol."},{"key":"27348_CR69","unstructured":"AlphaFold Protein Structure Database. https:\/\/alphafold.ebi.ac.uk\/."},{"key":"27348_CR70","doi-asserted-by":"publisher","first-page":"14","DOI":"10.1002\/pro.3235","volume":"27","author":"TD Goddard","year":"2018","unstructured":"Goddard, T. D. et al. UCSF ChimeraX: Meeting modern challenges in visualization and analysis. Protein Sci. 27, 14\u201325 (2018).","journal-title":"Protein Sci."},{"key":"27348_CR71","doi-asserted-by":"publisher","first-page":"70","DOI":"10.1002\/pro.3943","volume":"30","author":"EF Pettersen","year":"2021","unstructured":"Pettersen, E. F. et al. UCSF ChimeraX: Structure visualization for researchers, educators, and developers. Protein Sci. 30, 70\u201382 (2021).","journal-title":"Protein Sci."},{"key":"27348_CR72","doi-asserted-by":"publisher","first-page":"405","DOI":"10.1146\/annurev.immunol.15.1.405","volume":"15","author":"AB Rickinson","year":"1997","unstructured":"Rickinson, A. B. & Moss, D. J. Human cytotoxic T lymphocyte responses to Epstein-Barr virus infection. Annu. Rev. Immunol. 15, 405\u2013431 (1997).","journal-title":"Annu. Rev. Immunol."},{"key":"27348_CR73","doi-asserted-by":"publisher","first-page":"456","DOI":"10.1016\/j.humimm.2004.02.014","volume":"65","author":"P Moss","year":"2004","unstructured":"Moss, P. & Khan, N. CD8+ T-cell immunity to cytomegalovirus. Hum. Immunol. 65, 456\u2013464 (2004).","journal-title":"Hum. Immunol."}],"container-title":["Scientific Reports"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.nature.com\/articles\/s41598-022-27348-8.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/www.nature.com\/articles\/s41598-022-27348-8","content-type":"text\/html","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/www.nature.com\/articles\/s41598-022-27348-8.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2023,12,4]],"date-time":"2023-12-04T06:08:00Z","timestamp":1701670080000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.nature.com\/articles\/s41598-022-27348-8"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2023,1,8]]},"references-count":73,"journal-issue":{"issue":"1","published-online":{"date-parts":[[2023,12]]}},"alternative-id":["27348"],"URL":"https:\/\/doi.org\/10.1038\/s41598-022-27348-8","relation":{},"ISSN":["2045-2322"],"issn-type":[{"value":"2045-2322","type":"electronic"}],"subject":[],"published":{"date-parts":[[2023,1,8]]},"assertion":[{"value":"19 May 2022","order":1,"name":"received","label":"Received","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"30 December 2022","order":2,"name":"accepted","label":"Accepted","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"8 January 2023","order":3,"name":"first_online","label":"First Online","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"The authors declare no competing interests.","order":1,"name":"Ethics","group":{"name":"EthicsHeading","label":"Competing interests"}}],"article-number":"389"}}