{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,6,15]],"date-time":"2026-06-15T14:39:58Z","timestamp":1781534398622,"version":"3.54.5"},"reference-count":205,"publisher":"MDPI AG","issue":"11","license":[{"start":{"date-parts":[[2021,5,27]],"date-time":"2021-05-27T00:00:00Z","timestamp":1622073600000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"DOI":"10.13039\/100014440","name":"Ministerio de Ciencia, Innovaci\u00f3n y Universidades","doi-asserted-by":"publisher","award":["PID2019 110570GB-I00"],"award-info":[{"award-number":["PID2019 110570GB-I00"]}],"id":[{"id":"10.13039\/100014440","id-type":"DOI","asserted-by":"publisher"}]}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":["IJMS"],"abstract":"<jats:p>Herpes simplex virus type 1 (HSV-1) is a neurotropic alphaherpesvirus that can infect the peripheral and central nervous systems, and it has been implicated in demyelinating and neurodegenerative processes. Transposable elements (TEs) are DNA sequences that can move from one genomic location to another. TEs have been linked to several diseases affecting the central nervous system (CNS), including multiple sclerosis (MS), a demyelinating disease of unknown etiology influenced by genetic and environmental factors. Exogenous viral transactivators may activate certain retrotransposons or class I TEs. In this context, several herpesviruses have been linked to MS, and one of them, HSV-1, might act as a risk factor by mediating processes such as molecular mimicry, remyelination, and activity of endogenous retroviruses (ERVs). Several herpesviruses have been involved in the regulation of human ERVs (HERVs), and HSV-1 in particular can modulate HERVs in cells involved in MS pathogenesis. This review exposes current knowledge about the relationship between HSV-1 and human ERVs, focusing on their contribution as a risk factor for MS.<\/jats:p>","DOI":"10.3390\/ijms22115738","type":"journal-article","created":{"date-parts":[[2021,5,27]],"date-time":"2021-05-27T23:16:46Z","timestamp":1622157406000},"page":"5738","update-policy":"https:\/\/doi.org\/10.3390\/mdpi_crossmark_policy","source":"Crossref","is-referenced-by-count":28,"title":["HSV-1 and Endogenous Retroviruses as Risk Factors in Demyelination"],"prefix":"10.3390","volume":"22","author":[{"ORCID":"https:\/\/orcid.org\/0000-0002-4415-6483","authenticated-orcid":false,"given":"Raquel","family":"Bello-Morales","sequence":"first","affiliation":[{"name":"Departamento de Biolog\u00eda Molecular, Universidad Aut\u00f3noma de Madrid, Cantoblanco, 28049 Madrid, Spain"},{"name":"Centro de Biolog\u00eda Molecular Severo Ochoa, CSIC-UAM, Cantoblanco, 28049 Madrid, Spain"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-0600-4152","authenticated-orcid":false,"given":"Sabina","family":"Andreu","sequence":"additional","affiliation":[{"name":"Departamento de Biolog\u00eda Molecular, Universidad Aut\u00f3noma de Madrid, Cantoblanco, 28049 Madrid, Spain"},{"name":"Centro de Biolog\u00eda Molecular Severo Ochoa, CSIC-UAM, Cantoblanco, 28049 Madrid, Spain"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"In\u00e9s","family":"Ripa","sequence":"additional","affiliation":[{"name":"Departamento de Biolog\u00eda Molecular, Universidad Aut\u00f3noma de Madrid, Cantoblanco, 28049 Madrid, Spain"},{"name":"Centro de Biolog\u00eda Molecular Severo Ochoa, CSIC-UAM, Cantoblanco, 28049 Madrid, Spain"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-2817-5956","authenticated-orcid":false,"given":"Jos\u00e9 Antonio","family":"L\u00f3pez-Guerrero","sequence":"additional","affiliation":[{"name":"Departamento de Biolog\u00eda Molecular, Universidad Aut\u00f3noma de Madrid, Cantoblanco, 28049 Madrid, Spain"},{"name":"Centro de Biolog\u00eda Molecular Severo Ochoa, CSIC-UAM, Cantoblanco, 28049 Madrid, Spain"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"1968","published-online":{"date-parts":[[2021,5,27]]},"reference":[{"key":"ref_1","unstructured":"Howley, D.M.K.a.P.M. (2007). Herpes simplex viruses. Fields Virology, Lippincott Williams & Wilkins."},{"key":"ref_2","doi-asserted-by":"crossref","unstructured":"Arvin, A., Campadelli-Fiume, G., Mocarski, E., Moore, P.S., Roizman, B., Whitley, R., and Yamanishi, K. (2007). Persistence in the population: Epidemiology, transmission. Human Herpesviruses: Biology, Therapy, and Immunoprophylaxis, Cambridge University Press.","DOI":"10.1017\/CBO9780511545313"},{"key":"ref_3","doi-asserted-by":"crossref","first-page":"481","DOI":"10.1186\/1743-422X-8-481","article-title":"Herpes simplex virus infects most cell types in vitro: Clues to its success","volume":"8","author":"Karasneh","year":"2011","journal-title":"Virol. J."},{"key":"ref_4","doi-asserted-by":"crossref","first-page":"512","DOI":"10.1007\/s13365-011-0058-x","article-title":"Checkpoints in productive and latent infections with herpes simplex virus 1: Conceptualization of the issues","volume":"17","author":"Roizman","year":"2011","journal-title":"J. Neurovirology"},{"key":"ref_5","doi-asserted-by":"crossref","unstructured":"Bello-Morales, R., Andreu, S., and Lopez-Guerrero, J.A. (2020). The Role of Herpes Simplex Virus Type 1 Infection in Demyelination of the Central Nervous System. Int. J. Mol. Sci., 21.","DOI":"10.3390\/ijms21145026"},{"key":"ref_6","doi-asserted-by":"crossref","first-page":"43","DOI":"10.1038\/s41572-018-0041-4","article-title":"Multiple sclerosis","volume":"4","author":"Filippi","year":"2018","journal-title":"Nat. Rev. Dis. Primers"},{"key":"ref_7","doi-asserted-by":"crossref","first-page":"147","DOI":"10.1038\/nrn3900","article-title":"Exploring the origins of grey matter damage in multiple sclerosis","volume":"16","author":"Calabrese","year":"2015","journal-title":"Nat. Rev. Neurosci."},{"key":"ref_8","doi-asserted-by":"crossref","first-page":"540","DOI":"10.1191\/1352458503ms965oa","article-title":"Blood-brain barrier disruption in multiple sclerosis","volume":"9","author":"Minagar","year":"2003","journal-title":"Mult. Scler."},{"key":"ref_9","doi-asserted-by":"crossref","first-page":"452","DOI":"10.1016\/j.bbadis.2015.10.010","article-title":"Molecular alterations of the blood-brain barrier under inflammatory conditions: The role of endothelial to mesenchymal transition","volume":"1862","author":"Kamermans","year":"2016","journal-title":"Biochim. Et Biophys. Acta"},{"key":"ref_10","doi-asserted-by":"crossref","first-page":"11718","DOI":"10.3390\/ijms130911718","article-title":"Environmental risk factors for multiple sclerosis: A review with a focus on molecular mechanisms","volume":"13","author":"Lucas","year":"2012","journal-title":"Int. J. Mol. Sci."},{"key":"ref_11","doi-asserted-by":"crossref","first-page":"309","DOI":"10.1007\/978-981-15-3449-2_12","article-title":"Epigenetics in Multiple Sclerosis","volume":"1253","author":"Chan","year":"2020","journal-title":"Adv. Exp. Med. Biol."},{"key":"ref_12","unstructured":"Zagon, I.S., and McLaughlin, P.J. (2017). The Genetics of Multiple Sclerosis. Multiple Sclerosis: Perspectives in Treatment and Pathogenesis, Codon Publications."},{"key":"ref_13","doi-asserted-by":"crossref","unstructured":"Donati, D. (2020). Viral infections and multiple sclerosis. Drug Discov. Today. Dis. Models.","DOI":"10.1016\/j.ddmod.2020.02.003"},{"key":"ref_14","doi-asserted-by":"crossref","unstructured":"Tarlinton, R.E., Martynova, E., Rizvanov, A.A., Khaiboullina, S., and Verma, S. (2020). Role of Viruses in the Pathogenesis of Multiple Sclerosis. Viruses, 12.","DOI":"10.3390\/v12060643"},{"key":"ref_15","doi-asserted-by":"crossref","first-page":"355","DOI":"10.1038\/s41579-019-0189-2","article-title":"Origins and evolutionary consequences of ancient endogenous retroviruses","volume":"17","author":"Johnson","year":"2019","journal-title":"Nat. Rev. Microbiol."},{"key":"ref_16","doi-asserted-by":"crossref","first-page":"2039","DOI":"10.3389\/fimmu.2018.02039","article-title":"Human Endogenous Retroviruses Are Ancient Acquired Elements Still Shaping Innate Immune Responses","volume":"9","author":"Grandi","year":"2018","journal-title":"Front. Immunol."},{"key":"ref_17","doi-asserted-by":"crossref","first-page":"20120507","DOI":"10.1098\/rstb.2012.0507","article-title":"Paleovirology of \u2032syncytins\u2032, retroviral env genes exapted for a role in placentation","volume":"368","author":"Lavialle","year":"2013","journal-title":"Philos. Trans. R. Soc. Lond. Ser. B. Biol. Sci."},{"key":"ref_18","doi-asserted-by":"crossref","first-page":"1083","DOI":"10.1126\/science.aad5497","article-title":"Regulatory evolution of innate immunity through co-option of endogenous retroviruses","volume":"351","author":"Chuong","year":"2016","journal-title":"Science"},{"key":"ref_19","doi-asserted-by":"crossref","first-page":"415","DOI":"10.1038\/ni.2872","article-title":"The enemy within: Endogenous retroelements and autoimmune disease","volume":"15","author":"Volkman","year":"2014","journal-title":"Nat. Immunol."},{"key":"ref_20","doi-asserted-by":"crossref","first-page":"405","DOI":"10.1038\/nature13804","article-title":"Primate-specific endogenous retrovirus-driven transcription defines naive-like stem cells","volume":"516","author":"Wang","year":"2014","journal-title":"Nature"},{"key":"ref_21","doi-asserted-by":"crossref","first-page":"17842","DOI":"10.1073\/pnas.2002427117","article-title":"Regulation of stem cell function and neuronal differentiation by HERV-K via mTOR pathway","volume":"117","author":"Wang","year":"2020","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_22","doi-asserted-by":"crossref","first-page":"49","DOI":"10.1016\/j.coviro.2017.07.001","article-title":"How to tame an endogenous retrovirus: HERVH and the evolution of human pluripotency","volume":"25","author":"Romer","year":"2017","journal-title":"Curr. Opin. Virol."},{"key":"ref_23","doi-asserted-by":"crossref","first-page":"379","DOI":"10.1016\/j.molmed.2018.02.007","article-title":"Human Endogenous Retroviruses in Neurological Diseases","volume":"24","author":"Kury","year":"2018","journal-title":"Trends Mol. Med."},{"key":"ref_24","doi-asserted-by":"crossref","first-page":"265","DOI":"10.3389\/fmicb.2018.00265","article-title":"Human Endogenous Retroviruses and Their Putative Role in the Development of Autoimmune Disorders Such as Multiple Sclerosis","volume":"9","author":"Groger","year":"2018","journal-title":"Front. Microbiol."},{"key":"ref_25","doi-asserted-by":"crossref","first-page":"560","DOI":"10.1177\/014107680409701202","article-title":"Human endogenous retroviruses in health and disease: A symbiotic perspective","volume":"97","author":"Ryan","year":"2004","journal-title":"J. R. Soc. Med."},{"key":"ref_26","doi-asserted-by":"crossref","first-page":"6","DOI":"10.1038\/s41389-018-0114-y","article-title":"Transactivation of human endogenous retroviruses by tumor viruses and their functions in virus-associated malignancies","volume":"8","author":"Chen","year":"2019","journal-title":"Oncogenesis"},{"key":"ref_27","doi-asserted-by":"crossref","first-page":"2590","DOI":"10.1007\/s12035-018-1255-x","article-title":"Do Human Endogenous Retroviruses Contribute to Multiple Sclerosis, and if So, How?","volume":"56","author":"Morris","year":"2019","journal-title":"Mol. Neurobiol."},{"key":"ref_28","doi-asserted-by":"crossref","unstructured":"Morandi, E., Tanasescu, R., Tarlinton, R.E., Constantinescu, C.S., Zhang, W., Tench, C., and Gran, B. (2017). The association between human endogenous retroviruses and multiple sclerosis: A systematic review and meta-analysis. PLoS ONE, 12.","DOI":"10.1371\/journal.pone.0172415"},{"key":"ref_29","doi-asserted-by":"crossref","first-page":"179","DOI":"10.1002\/rmv.465","article-title":"Association of human endogenous retroviruses with multiple sclerosis and possible interactions with herpes viruses","volume":"15","author":"Christensen","year":"2005","journal-title":"Rev. Med. Virol."},{"key":"ref_30","doi-asserted-by":"crossref","first-page":"18","DOI":"10.1111\/ane.12836","article-title":"Human endogenous retroviruses in the aetiology of MS","volume":"136","author":"Christensen","year":"2017","journal-title":"Acta Neurol. Scand."},{"key":"ref_31","doi-asserted-by":"crossref","unstructured":"Lezhnyova, V.R., Martynova, E.V., Khaiboullin, T.I., Urbanowicz, R.A., Khaiboullina, S.F., and Rizvanov, A.A. (2020). The Relationship of the Mechanisms of the Pathogenesis of Multiple Sclerosis and the Expression of Endogenous Retroviruses. Biology, 9.","DOI":"10.3390\/biology9120464"},{"key":"ref_32","doi-asserted-by":"crossref","first-page":"647","DOI":"10.3389\/fimmu.2015.00647","article-title":"Multiple Sclerosis between Genetics and Infections: Human Endogenous Retroviruses in Monocytes and Macrophages","volume":"6","author":"Morandi","year":"2015","journal-title":"Front. Immunol."},{"key":"ref_33","doi-asserted-by":"crossref","first-page":"162","DOI":"10.1016\/j.bbadis.2010.07.016","article-title":"Human endogenous retroviruses and multiple sclerosis: Innocent bystanders or disease determinants?","volume":"1812","author":"Antony","year":"2011","journal-title":"Biochim. Et Biophys. Acta"},{"key":"ref_34","doi-asserted-by":"crossref","first-page":"239","DOI":"10.1016\/j.jns.2016.11.060","article-title":"Association between human herpesvirus & human endogenous retrovirus and MS onset & progression","volume":"372","author":"Tao","year":"2017","journal-title":"J. Neurol. Sci."},{"key":"ref_35","doi-asserted-by":"crossref","first-page":"147","DOI":"10.1016\/j.jneuroim.2007.04.003","article-title":"Activation of endogenous retrovirus reverse transcriptase in multiple sclerosis patient lymphocytes by inactivated HSV-1, HHV-6 and VZV","volume":"187","author":"Brudek","year":"2007","journal-title":"J. Neuroimmunol."},{"key":"ref_36","doi-asserted-by":"crossref","first-page":"65","DOI":"10.1080\/13550280600614973","article-title":"Regulation of human endogenous retrovirus W protein expression by herpes simplex virus type 1: Implications for multiple sclerosis","volume":"12","author":"Ruprecht","year":"2006","journal-title":"J. Neurovirology"},{"key":"ref_37","doi-asserted-by":"crossref","first-page":"28","DOI":"10.1016\/j.jcv.2015.04.018","article-title":"CMV induces HERV-K and HERV-W expression in kidney transplant recipients","volume":"68","author":"Bergallo","year":"2015","journal-title":"J. Clin. Virol."},{"key":"ref_38","doi-asserted-by":"crossref","first-page":"47","DOI":"10.1016\/j.jcv.2009.05.019","article-title":"HHV-6A infection induces expression of HERV-K18-encoded superantigen","volume":"46","author":"Tai","year":"2009","journal-title":"J. Clin. Virol."},{"key":"ref_39","doi-asserted-by":"crossref","first-page":"579","DOI":"10.1016\/S1074-7613(01)00210-2","article-title":"Epstein-Barr virus transactivates the human endogenous retrovirus HERV-K18 that encodes a superantigen","volume":"15","author":"Sutkowski","year":"2001","journal-title":"Immunity"},{"key":"ref_40","doi-asserted-by":"crossref","unstructured":"Mameli, G., Poddighe, L., Mei, A., Uleri, E., Sotgiu, S., Serra, C., Manetti, R., and Dolei, A. (2012). Expression and activation by Epstein Barr virus of human endogenous retroviruses-W in blood cells and astrocytes: Inference for multiple sclerosis. PLoS ONE, 7.","DOI":"10.1371\/journal.pone.0044991"},{"key":"ref_41","doi-asserted-by":"crossref","first-page":"344","DOI":"10.1073\/pnas.36.6.344","article-title":"The origin and behavior of mutable loci in maize","volume":"36","author":"McClintock","year":"1950","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_42","doi-asserted-by":"crossref","first-page":"199","DOI":"10.1186\/s13059-018-1577-z","article-title":"Ten things you should know about transposable elements","volume":"19","author":"Bourque","year":"2018","journal-title":"Genome Biol."},{"key":"ref_43","first-page":"32","article-title":"Transposons, or Jumping Genes: Not Junk DNA?","volume":"1","author":"Pray","year":"2008","journal-title":"Nat. Educ."},{"key":"ref_44","doi-asserted-by":"crossref","first-page":"409","DOI":"10.3390\/genes3030409","article-title":"Transposable elements: From DNA parasites to architects of metazoan evolution","volume":"3","author":"Piskurek","year":"2012","journal-title":"Genes"},{"key":"ref_45","doi-asserted-by":"crossref","unstructured":"Drongitis, D., Aniello, F., Fucci, L., and Donizetti, A. (2019). Roles of Transposable Elements in the Different Layers of Gene Expression Regulation. Int. J. Mol. Sci., 20.","DOI":"10.3390\/ijms20225755"},{"key":"ref_46","doi-asserted-by":"crossref","first-page":"691","DOI":"10.1038\/nrg2640","article-title":"The impact of retrotransposons on human genome evolution","volume":"10","author":"Cordaux","year":"2009","journal-title":"Nat. Rev. Genet."},{"key":"ref_47","doi-asserted-by":"crossref","first-page":"17600","DOI":"10.1073\/pnas.0607612103","article-title":"Transposable elements and the evolution of eukaryotic genomes","volume":"103","author":"Wessler","year":"2006","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_48","unstructured":"van Regenmortel, M.H.V., and Mahy, B.W.J. (2008). Retrotransposons of Vertebrates. Encyclopedia of Virology, Academic Press. [3rd ed.]."},{"key":"ref_49","doi-asserted-by":"crossref","first-page":"MDNA3-0061-2014","DOI":"10.1128\/microbiolspec.MDNA3-0061-2014","article-title":"The Influence of LINE-1 and SINE Retrotransposons on Mammalian Genomes","volume":"3","author":"Richardson","year":"2015","journal-title":"Microbiol. Spectr."},{"key":"ref_50","doi-asserted-by":"crossref","first-page":"115","DOI":"10.2174\/138920210790886871","article-title":"DNA transposons: Nature and applications in genomics","volume":"11","year":"2010","journal-title":"Curr. Genom."},{"key":"ref_51","doi-asserted-by":"crossref","first-page":"14572","DOI":"10.1073\/pnas.0404838101","article-title":"Retroelements and the human genome: New perspectives on an old relation","volume":"101","author":"Bannert","year":"2004","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_52","doi-asserted-by":"crossref","first-page":"587","DOI":"10.1146\/annurev.genet.42.110807.091549","article-title":"The take and give between retrotransposable elements and their hosts","volume":"42","author":"Beauregard","year":"2008","journal-title":"Annu. Rev. Genet."},{"key":"ref_53","doi-asserted-by":"crossref","first-page":"610","DOI":"10.1016\/j.tig.2020.05.004","article-title":"Transposable Elements: A Common Feature of Neurodevelopmental and Neurodegenerative Disorders","volume":"36","author":"Jonsson","year":"2020","journal-title":"Trends Genet. Tig"},{"key":"ref_54","doi-asserted-by":"crossref","first-page":"51","DOI":"10.1016\/j.gde.2015.12.001","article-title":"Hide and seek: How chromatin-based pathways silence retroelements in the mammalian germline","volume":"37","author":"Molaro","year":"2016","journal-title":"Curr. Opin. Genet. Dev."},{"key":"ref_55","doi-asserted-by":"crossref","first-page":"417","DOI":"10.1038\/s41576-019-0106-6","article-title":"Regulation of transposable elements by DNA modifications","volume":"20","author":"Deniz","year":"2019","journal-title":"Nat. Rev. Genet."},{"key":"ref_56","doi-asserted-by":"crossref","first-page":"183","DOI":"10.1016\/j.tig.2007.02.006","article-title":"Which transposable elements are active in the human genome?","volume":"23","author":"Mills","year":"2007","journal-title":"Trends Genet Tig"},{"key":"ref_57","doi-asserted-by":"crossref","first-page":"715","DOI":"10.1080\/1040841X.2018.1501345","article-title":"Human endogenous retrovirus-K (HML-2): A comprehensive review","volume":"44","author":"Henderson","year":"2018","journal-title":"Crit. Rev. Microbiol."},{"key":"ref_58","doi-asserted-by":"crossref","first-page":"497","DOI":"10.1080\/24750263.2019.1695967","article-title":"Transposable elements in vertebrates: Species evolution and environmental adaptation","volume":"86","author":"Carducci","year":"2019","journal-title":"Eur. Zool. J."},{"key":"ref_59","doi-asserted-by":"crossref","unstructured":"Serrato-Capuchina, A., and Matute, D.R. (2018). The Role of Transposable Elements in Speciation. Genes, 9.","DOI":"10.3390\/genes9050254"},{"key":"ref_60","doi-asserted-by":"crossref","first-page":"1098","DOI":"10.1101\/gad.327312.119","article-title":"Host-transposon interactions: Conflict, cooperation, and cooption","volume":"33","author":"Cosby","year":"2019","journal-title":"Genes Dev."},{"key":"ref_61","doi-asserted-by":"crossref","first-page":"10863","DOI":"10.1073\/pnas.1102343108","article-title":"Selfish genetic elements, genetic conflict, and evolutionary innovation","volume":"108","author":"Werren","year":"2011","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_62","doi-asserted-by":"crossref","unstructured":"Agren, J.A., and Clark, A.G. (2018). Selfish genetic elements. PLoS Genet., 14.","DOI":"10.1371\/journal.pgen.1007700"},{"key":"ref_63","doi-asserted-by":"crossref","first-page":"34","DOI":"10.1016\/j.gde.2018.02.011","article-title":"Exaptation of transposable element coding sequences","volume":"49","author":"Bureau","year":"2018","journal-title":"Curr. Opin. Genet. Dev."},{"key":"ref_64","doi-asserted-by":"crossref","first-page":"627","DOI":"10.1016\/j.tree.2012.07.007","article-title":"Paleovirology and virally derived immunity","volume":"27","author":"Aswad","year":"2012","journal-title":"Trends Ecol. Evol."},{"key":"ref_65","doi-asserted-by":"crossref","first-page":"4","DOI":"10.1017\/S0094837300004310","article-title":"Exaptation\u2014A Missing Term in the Science of Form","volume":"8","author":"Gould","year":"1982","journal-title":"Paleobiology"},{"key":"ref_66","doi-asserted-by":"crossref","first-page":"E487","DOI":"10.1073\/pnas.1417000112","article-title":"Retroviral envelope gene captures and syncytin exaptation for placentation in marsupials","volume":"112","author":"Cornelis","year":"2015","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_67","doi-asserted-by":"crossref","first-page":"817","DOI":"10.1016\/j.tig.2017.07.011","article-title":"Transposable Element Domestication As an Adaptation to Evolutionary Conflicts","volume":"33","author":"Jangam","year":"2017","journal-title":"Trends Genet. Tig"},{"key":"ref_68","doi-asserted-by":"crossref","first-page":"760","DOI":"10.1038\/s41576-019-0165-8","article-title":"Transposable elements in human genetic disease","volume":"20","author":"Payer","year":"2019","journal-title":"Nat. Rev. Genet."},{"key":"ref_69","doi-asserted-by":"crossref","first-page":"65","DOI":"10.1016\/j.conb.2020.01.012","article-title":"Awakening the dark side: Retrotransposon activation in neurodegenerative disorders","volume":"61","author":"Zuniga","year":"2020","journal-title":"Curr. Opin. Neurobiol."},{"key":"ref_70","doi-asserted-by":"crossref","first-page":"894","DOI":"10.3389\/fneur.2019.00894","article-title":"Transposable Elements, Inflammation, and Neurological Disease","volume":"10","author":"Saleh","year":"2019","journal-title":"Front. Neurol."},{"key":"ref_71","doi-asserted-by":"crossref","unstructured":"Dolei, A., Ibba, G., Piu, C., and Serra, C. (2019). Expression of HERV Genes as Possible Biomarker and Target in Neurodegenerative Diseases. Int. J. Mol. Sci., 20.","DOI":"10.3390\/ijms20153706"},{"key":"ref_72","doi-asserted-by":"crossref","first-page":"307ra153","DOI":"10.1126\/scitranslmed.aac8201","article-title":"Human endogenous retrovirus-K contributes to motor neuron disease","volume":"7","author":"Li","year":"2015","journal-title":"Sci. Transl. Med."},{"key":"ref_73","doi-asserted-by":"crossref","first-page":"19359","DOI":"10.1073\/pnas.1921719117","article-title":"DNA methylation enables transposable element-driven genome expansion","volume":"117","author":"Zhou","year":"2020","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_74","doi-asserted-by":"crossref","first-page":"1150","DOI":"10.3389\/fgene.2019.01150","article-title":"DNA Methylation Cancer Biomarkers: Translation to the Clinic","volume":"10","author":"Locke","year":"2019","journal-title":"Front. Genet."},{"key":"ref_75","doi-asserted-by":"crossref","first-page":"245","DOI":"10.2217\/epi.10.2","article-title":"Hypomethylation of repeated DNA sequences in cancer","volume":"2","author":"Ross","year":"2010","journal-title":"Epigenomics"},{"key":"ref_76","doi-asserted-by":"crossref","first-page":"239","DOI":"10.2217\/epi.09.33","article-title":"DNA hypomethylation in cancer cells","volume":"1","author":"Ehrlich","year":"2009","journal-title":"Epigenomics"},{"key":"ref_77","doi-asserted-by":"crossref","first-page":"6883","DOI":"10.1093\/nar\/11.19.6883","article-title":"The 5-methylcytosine content of DNA from human tumors","volume":"11","author":"Slagel","year":"1983","journal-title":"Nucleic Acids Res."},{"key":"ref_78","doi-asserted-by":"crossref","unstructured":"Anwar, S.L., Wulaningsih, W., and Lehmann, U. (2017). Transposable Elements in Human Cancer: Causes and Consequences of Deregulation. Int. J. Mol. Sci., 18.","DOI":"10.3390\/ijms18050974"},{"key":"ref_79","doi-asserted-by":"crossref","first-page":"717","DOI":"10.1038\/ng.2680","article-title":"Hypomethylation marks enhancers within transposable elements","volume":"45","author":"Mukamel","year":"2013","journal-title":"Nat. Genet."},{"key":"ref_80","doi-asserted-by":"crossref","first-page":"836","DOI":"10.1038\/ng.2649","article-title":"DNA hypomethylation within specific transposable element families associates with tissue-specific enhancer landscape","volume":"45","author":"Xie","year":"2013","journal-title":"Nat. Genet."},{"key":"ref_81","doi-asserted-by":"crossref","first-page":"1395","DOI":"10.1126\/science.aah3378","article-title":"Early life experience drives structural variation of neural genomes in mice","volume":"359","author":"Bedrosian","year":"2018","journal-title":"Science"},{"key":"ref_82","doi-asserted-by":"crossref","first-page":"4892","DOI":"10.1073\/pnas.78.8.4892","article-title":"Identification and cloning of endogenous retroviral sequences present in human DNA","volume":"78","author":"Martin","year":"1981","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_83","doi-asserted-by":"crossref","first-page":"67","DOI":"10.1186\/1742-4690-3-67","article-title":"The discovery of endogenous retroviruses","volume":"3","author":"Weiss","year":"2006","journal-title":"Retrovirology"},{"key":"ref_84","doi-asserted-by":"crossref","first-page":"3715","DOI":"10.1128\/JVI.74.8.3715-3730.2000","article-title":"Identification and characterization of novel human endogenous retrovirus families by phylogenetic screening of the human genome mapping project database","volume":"74","author":"Tristem","year":"2000","journal-title":"J. Virol."},{"key":"ref_85","doi-asserted-by":"crossref","first-page":"164","DOI":"10.1006\/viro.2002.1615","article-title":"Characterization of a novel human endogenous retrovirus, HERV-H\/F, expressed in human leukemia cell lines","volume":"303","author":"Patzke","year":"2002","journal-title":"Virology"},{"key":"ref_86","doi-asserted-by":"crossref","first-page":"861","DOI":"10.1016\/S0960-9822(99)80390-X","article-title":"Many human endogenous retrovirus K (HERV-K) proviruses are unique to humans","volume":"9","author":"Barbulescu","year":"1999","journal-title":"Curr. Biol. Cb"},{"key":"ref_87","doi-asserted-by":"crossref","first-page":"90","DOI":"10.1186\/1742-4690-8-90","article-title":"Identification, characterization, and comparative genomic distribution of the HERV-K (HML-2) group of human endogenous retroviruses","volume":"8","author":"Subramanian","year":"2011","journal-title":"Retrovirology"},{"key":"ref_88","doi-asserted-by":"crossref","first-page":"8201642","DOI":"10.1155\/2016\/8201642","article-title":"Human Endogenous Retroviruses-K (HML-2) Expression Is Correlated with Prognosis and Progress of Hepatocellular Carcinoma","volume":"2016","author":"Ma","year":"2016","journal-title":"Biomed. Res. Int."},{"key":"ref_89","first-page":"70","article-title":"Human endogenous retrovirus np9 gene is over expressed in chronic lymphocytic leukemia patients","volume":"3","author":"Fischer","year":"2014","journal-title":"Leuk. Res. Rep."},{"key":"ref_90","doi-asserted-by":"crossref","first-page":"849","DOI":"10.1016\/j.yexcr.2008.12.023","article-title":"The activation of human endogenous retrovirus K (HERV-K) is implicated in melanoma cell malignant transformation","volume":"315","author":"Serafino","year":"2009","journal-title":"Exp. Cell Res."},{"key":"ref_91","doi-asserted-by":"crossref","first-page":"42","DOI":"10.1177\/1352458517737370","article-title":"The aliens inside us: HERV-W endogenous retroviruses and multiple sclerosis","volume":"24","author":"Dolei","year":"2018","journal-title":"Mult. Scler."},{"key":"ref_92","doi-asserted-by":"crossref","first-page":"7","DOI":"10.1186\/s12977-015-0232-y","article-title":"Classification and characterization of human endogenous retroviruses; mosaic forms are common","volume":"13","author":"Vargiu","year":"2016","journal-title":"Retrovirology"},{"key":"ref_93","doi-asserted-by":"crossref","first-page":"7187","DOI":"10.1128\/JVI.00544-15","article-title":"Human Endogenous Retrovirus Type K (HERV-K) Particles Package and Transmit HERV-K-Related Sequences","volume":"89","author":"Kaplan","year":"2015","journal-title":"J. Virol."},{"key":"ref_94","doi-asserted-by":"crossref","first-page":"321","DOI":"10.1006\/viro.2001.1045","article-title":"Multiple sclerosis retrovirus particles and recombinant envelope trigger an abnormal immune response in vitro, by inducing polyclonal Vbeta16 T-lymphocyte activation","volume":"287","author":"Perron","year":"2001","journal-title":"Virology"},{"key":"ref_95","doi-asserted-by":"crossref","first-page":"59","DOI":"10.1186\/s12977-018-0442-1","article-title":"Nomenclature for endogenous retrovirus (ERV) loci","volume":"15","author":"Gifford","year":"2018","journal-title":"Retrovirology"},{"key":"ref_96","doi-asserted-by":"crossref","unstructured":"Groger, V., Emmer, A., Staege, M.S., and Cynis, H. (2021). Endogenous Retroviruses in Nervous System Disorders. Pharmaceuticals, 14.","DOI":"10.3390\/ph14010070"},{"key":"ref_97","doi-asserted-by":"crossref","first-page":"45","DOI":"10.1186\/s40478-019-0698-2","article-title":"Quantitative analysis of human endogenous retrovirus-K transcripts in postmortem premotor cortex fails to confirm elevated expression of HERV-K RNA in amyotrophic lateral sclerosis","volume":"7","author":"Garson","year":"2019","journal-title":"Acta Neuropathol. Commun."},{"key":"ref_98","doi-asserted-by":"crossref","first-page":"101","DOI":"10.1186\/s40478-019-0753-z","article-title":"Technical considerations in detection of HERV-K in amyotrophic lateral sclerosis: Selection of controls and the perils of qPCR","volume":"7","author":"Li","year":"2019","journal-title":"Acta Neuropathol. Commun."},{"key":"ref_99","doi-asserted-by":"crossref","first-page":"102","DOI":"10.1186\/s40478-019-0756-9","article-title":"Response to the Letter from Garcia-Montojo and colleagues concerning our paper entitled, Quantitative analysis of human endogenous retrovirus-K transcripts in postmortem premotor cortex fails to confirm elevated expression of HERV-K RNA in amyotrophic lateral sclerosis","volume":"7","author":"Garson","year":"2019","journal-title":"Acta Neuropathol. Commun."},{"key":"ref_100","doi-asserted-by":"crossref","first-page":"1033","DOI":"10.1016\/S0140-6736(05)60075-X","article-title":"Expression of sequence variants of endogenous retrovirus RGH in particle form in multiple sclerosis","volume":"352","author":"Christensen","year":"1998","journal-title":"Lancet"},{"key":"ref_101","doi-asserted-by":"crossref","first-page":"6","DOI":"10.1191\/1352458503ms867oa","article-title":"Antibodies against a human endogenous retrovirus and the preponderance of env splice variants in multiple sclerosis patients","volume":"9","author":"Christensen","year":"2003","journal-title":"Mult. Scler."},{"key":"ref_102","doi-asserted-by":"crossref","first-page":"292","DOI":"10.59566\/IJBS.2007.3292","article-title":"Multiple Sclerosis and HERV-W\/MSRV: A Multicentric Study","volume":"3","author":"Arru","year":"2007","journal-title":"Int. J. Biomed. Sci."},{"key":"ref_103","first-page":"637","article-title":"In vitro modulation of the multiple sclerosis (MS)-associated retrovirus by cytokines: Implications for MS pathogenesis","volume":"9","author":"Serra","year":"2003","journal-title":"J. Neurovirology"},{"key":"ref_104","doi-asserted-by":"crossref","first-page":"108","DOI":"10.1016\/j.jneuroim.2009.08.015","article-title":"Effects of interferon-beta therapy on innate and adaptive immune responses to the human endogenous retroviruses HERV-H and HERV-W, cytokine production, and the lectin complement activation pathway in multiple sclerosis","volume":"215","author":"Petersen","year":"2009","journal-title":"J. Neuroimmunol."},{"key":"ref_105","doi-asserted-by":"crossref","first-page":"73","DOI":"10.1080\/13550280701801107","article-title":"Inhibition of multiple-sclerosis-associated retrovirus as biomarker of interferon therapy","volume":"14","author":"Mameli","year":"2008","journal-title":"J. Neurovirology"},{"key":"ref_106","doi-asserted-by":"crossref","first-page":"7636","DOI":"10.4049\/jimmunol.176.12.7636","article-title":"The envelope protein of a human endogenous retrovirus-W family activates innate immunity through CD14\/TLR4 and promotes Th1-like responses","volume":"176","author":"Rolland","year":"2006","journal-title":"J. Immunol."},{"key":"ref_107","doi-asserted-by":"crossref","first-page":"195","DOI":"10.1016\/j.jneuroim.2004.10.019","article-title":"Correlation between disease severity and in vitro cytokine production mediated by MSRV (multiple sclerosis associated retroviral element) envelope protein in patients with multiple sclerosis","volume":"160","author":"Rolland","year":"2005","journal-title":"J. Neuroimmunol."},{"key":"ref_108","doi-asserted-by":"crossref","unstructured":"Kriesel, J.D., Bhetariya, P.J., Chan, B.K., Wilson, T., and Fischer, K.F. (2017). Enrichment of Retroviral Sequences in Brain Tissue from Patients with Severe Demyelinating Diseases. J. Emerg. Dis. Virol., 3.","DOI":"10.16966\/2473-1846.132"},{"key":"ref_109","doi-asserted-by":"crossref","first-page":"685","DOI":"10.1002\/eji.201948423","article-title":"Syncytin-1\/HERV-W envelope is an early activation marker of leukocytes and is upregulated in multiple sclerosis patients","volume":"50","author":"Casanova","year":"2020","journal-title":"Eur. J. Immunol."},{"key":"ref_110","doi-asserted-by":"crossref","first-page":"11","DOI":"10.1016\/j.msard.2016.04.006","article-title":"Human endogenous retrovirus W in brain lesions: Rationale for targeted therapy in multiple sclerosis","volume":"8","author":"Nijland","year":"2016","journal-title":"Mult. Scler. Relat. Disord."},{"key":"ref_111","doi-asserted-by":"crossref","unstructured":"Grandi, N., and Tramontano, E. (2017). Type W Human Endogenous Retrovirus (HERV-W) Integrations and Their Mobilization by L1 Machinery: Contribution to the Human Transcriptome and Impact on the Host Physiopathology. Viruses, 9.","DOI":"10.3390\/v9070162"},{"key":"ref_112","doi-asserted-by":"crossref","first-page":"1175","DOI":"10.1128\/JVI.73.2.1175-1185.1999","article-title":"Molecular characterization and placental expression of HERV-W, a new human endogenous retrovirus family","volume":"73","author":"Blond","year":"1999","journal-title":"J. Virol."},{"key":"ref_113","doi-asserted-by":"crossref","first-page":"264","DOI":"10.1099\/vir.0.81890-0","article-title":"Brains and peripheral blood mononuclear cells of multiple sclerosis (MS) patients hyperexpress MS-associated retrovirus\/HERV-W endogenous retrovirus, but not Human herpesvirus 6","volume":"88","author":"Mameli","year":"2007","journal-title":"J. Gen. Virol."},{"key":"ref_114","doi-asserted-by":"crossref","first-page":"1645","DOI":"10.3389\/fphar.2019.01645","article-title":"Differential Expression of HERV-W in Peripheral Blood in Multiple Sclerosis and Healthy Patients in Two Different Ethnic Groups","volume":"10","author":"Tarlinton","year":"2019","journal-title":"Front. Pharmacol."},{"key":"ref_115","doi-asserted-by":"crossref","first-page":"721","DOI":"10.1002\/ana.23970","article-title":"Human endogenous retrovirus type W envelope protein inhibits oligodendroglial precursor cell differentiation","volume":"74","author":"Kremer","year":"2013","journal-title":"Ann. Neurol."},{"key":"ref_116","doi-asserted-by":"crossref","first-page":"29","DOI":"10.1016\/j.jneuroim.2015.12.006","article-title":"MSRV envelope protein is a potent, endogenous and pathogenic agonist of human toll-like receptor 4: Relevance of GNbAC1 in multiple sclerosis treatment","volume":"291","author":"Madeira","year":"2016","journal-title":"J. Neuroimmunol."},{"key":"ref_117","doi-asserted-by":"crossref","first-page":"15216","DOI":"10.1073\/pnas.1901283116","article-title":"pHERV-W envelope protein fuels microglial cell-dependent damage of myelinated axons in multiple sclerosis","volume":"116","author":"Kremer","year":"2019","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_118","doi-asserted-by":"crossref","first-page":"104","DOI":"10.1186\/1742-4690-6-104","article-title":"B cells and monocytes from patients with active multiple sclerosis exhibit increased surface expression of both HERV-H Env and HERV-W Env, accompanied by increased seroreactivity","volume":"6","author":"Brudek","year":"2009","journal-title":"Retrovirology"},{"key":"ref_119","doi-asserted-by":"crossref","unstructured":"Lopez-Guerrero, J.A., Ripa, I., Andreu, S., and Bello-Morales, R. (2020). The Role of Extracellular Vesicles in Demyelination of the Central Nervous System. Int. J. Mol. Sci., 21.","DOI":"10.3390\/ijms21239111"},{"key":"ref_120","doi-asserted-by":"crossref","first-page":"545","DOI":"10.1093\/intimm\/dxv025","article-title":"Inflammatory response of endothelial cells to a human endogenous retrovirus associated with multiple sclerosis is mediated by TLR4","volume":"27","author":"Duperray","year":"2015","journal-title":"Int. Immunol."},{"key":"ref_121","doi-asserted-by":"crossref","first-page":"1287","DOI":"10.1007\/s40264-020-00988-3","article-title":"Temelimab, an IgG4 Anti-Human Endogenous Retrovirus Monoclonal Antibody: An Early Development Safety Review","volume":"43","author":"Kornmann","year":"2020","journal-title":"Drug Saf."},{"key":"ref_122","doi-asserted-by":"crossref","first-page":"785","DOI":"10.1038\/35001608","article-title":"Syncytin is a captive retroviral envelope protein involved in human placental morphogenesis","volume":"403","author":"Mi","year":"2000","journal-title":"Nature"},{"key":"ref_123","doi-asserted-by":"crossref","first-page":"E10991","DOI":"10.1073\/pnas.1714590114","article-title":"An endogenous retroviral envelope syncytin and its cognate receptor identified in the viviparous placental Mabuya lizard","volume":"114","author":"Cornelis","year":"2017","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_124","doi-asserted-by":"crossref","first-page":"1796","DOI":"10.2174\/13816128113199990541","article-title":"Decreased expression and altered methylation of syncytin-1 gene in human placentas associated with preeclampsia","volume":"20","author":"Zhuang","year":"2014","journal-title":"Curr. Pharm. Des."},{"key":"ref_125","doi-asserted-by":"crossref","first-page":"1088","DOI":"10.1038\/nn1319","article-title":"Human endogenous retrovirus glycoprotein-mediated induction of redox reactants causes oligodendrocyte death and demyelination","volume":"7","author":"Antony","year":"2004","journal-title":"Nat. Neurosci."},{"key":"ref_126","doi-asserted-by":"crossref","first-page":"422","DOI":"10.3389\/fpsyt.2018.00422","article-title":"Human Endogenous Retroviral Envelope Protein Syncytin-1 and Inflammatory Abnormalities in Neuropsychological Diseases","volume":"9","author":"Wang","year":"2018","journal-title":"Front. Psychiatry"},{"key":"ref_127","doi-asserted-by":"crossref","first-page":"1906","DOI":"10.1007\/s00018-006-6201-9","article-title":"Syncytin is involved in breast cancer-endothelial cell fusions","volume":"63","author":"Bjerregaard","year":"2006","journal-title":"Cell. Mol. Life Sci."},{"key":"ref_128","doi-asserted-by":"crossref","first-page":"31","DOI":"10.2147\/OTT.S191041","article-title":"Upregulation of syncytin-1 promotes invasion and metastasis by activating epithelial-mesenchymal transition-related pathway in endometrial carcinoma","volume":"12","author":"Liu","year":"2019","journal-title":"Oncotargets Ther."},{"key":"ref_129","doi-asserted-by":"crossref","first-page":"23","DOI":"10.1007\/s00109-006-0104-y","article-title":"Proliferation and cell-cell fusion of endometrial carcinoma are induced by the human endogenous retroviral Syncytin-1 and regulated by TGF-beta","volume":"85","author":"Strick","year":"2007","journal-title":"J. Mol. Med."},{"key":"ref_130","doi-asserted-by":"crossref","first-page":"40983","DOI":"10.1038\/srep40983","article-title":"Up-regulation of syncytin-1 contributes to TNF-alpha-enhanced fusion between OSCC and HUVECs partly via Wnt\/beta-catenin-dependent pathway","volume":"7","author":"Yan","year":"2017","journal-title":"Sci. Rep."},{"key":"ref_131","doi-asserted-by":"crossref","first-page":"13013","DOI":"10.1073\/pnas.2132646100","article-title":"Genomewide screening for fusogenic human endogenous retrovirus envelopes identifies syncytin 2, a gene conserved on primate evolution","volume":"100","author":"Blaise","year":"2003","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_132","doi-asserted-by":"crossref","first-page":"815","DOI":"10.1095\/biolreprod.108.069765","article-title":"Functional characterization of the human placental fusogenic membrane protein syncytin 2","volume":"79","author":"Chen","year":"2008","journal-title":"Biol. Reprod."},{"key":"ref_133","doi-asserted-by":"crossref","first-page":"37","DOI":"10.1186\/1742-4690-6-37","article-title":"Analysis of transcribed human endogenous retrovirus W env loci clarifies the origin of multiple sclerosis-associated retrovirus env sequences","volume":"6","author":"Laufer","year":"2009","journal-title":"Retrovirology"},{"key":"ref_134","doi-asserted-by":"crossref","first-page":"1251","DOI":"10.1089\/aid.2006.0274","article-title":"Comparative expression of human endogenous retrovirus-W genes in multiple sclerosis","volume":"23","author":"Antony","year":"2007","journal-title":"Aids Res. Hum. Retrovir."},{"key":"ref_135","doi-asserted-by":"crossref","first-page":"4","DOI":"10.1080\/13550280802448451","article-title":"The multiple sclerosis-associated retrovirus and its HERV-W endogenous family: A biological interface between virology, genetics, and immunology in human physiology and disease","volume":"15","author":"Dolei","year":"2009","journal-title":"J. Neurovirology"},{"key":"ref_136","doi-asserted-by":"crossref","first-page":"229","DOI":"10.1034\/j.1600-0404.2000.101004229.x","article-title":"Molecular characterization of HERV-H variants associated with multiple sclerosis","volume":"101","author":"Christensen","year":"2000","journal-title":"Acta Neurol. Scand."},{"key":"ref_137","doi-asserted-by":"crossref","first-page":"175","DOI":"10.1016\/j.jneuroim.2006.09.014","article-title":"Gene-environment interactions in multiple sclerosis: Innate and adaptive immune responses to human endogenous retrovirus and herpesvirus antigens and the lectin complement activation pathway","volume":"183","author":"Christensen","year":"2007","journal-title":"J. Neuroimmunol."},{"key":"ref_138","doi-asserted-by":"crossref","first-page":"595","DOI":"10.1177\/1352458507086425","article-title":"Herpesviruses and human endogenous retroviral sequences in the cerebrospinal fluid of multiple sclerosis patients","volume":"14","author":"Bartolome","year":"2008","journal-title":"Mult. Scler."},{"key":"ref_139","doi-asserted-by":"crossref","first-page":"2383","DOI":"10.1099\/0022-1317-80-9-2383","article-title":"HERV-F, a new group of human endogenous retrovirus sequences","volume":"80","author":"Kjellman","year":"1999","journal-title":"J. Gen. Virol."},{"key":"ref_140","doi-asserted-by":"crossref","first-page":"3713","DOI":"10.1128\/JVI.06723-11","article-title":"Expression of HERV-Fc1, a human endogenous retrovirus, is increased in patients with active multiple sclerosis","volume":"86","author":"Laska","year":"2012","journal-title":"J. Virol."},{"key":"ref_141","doi-asserted-by":"crossref","unstructured":"Nexo, B.A., Christensen, T., Frederiksen, J., Moller-Larsen, A., Oturai, A.B., Villesen, P., Hansen, B., Nissen, K.K., Laska, M.J., and Petersen, T.S. (2011). The etiology of multiple sclerosis: Genetic evidence for the involvement of the human endogenous retrovirus HERV-Fc1. PLoS ONE, 6.","DOI":"10.1371\/journal.pone.0016652"},{"key":"ref_142","doi-asserted-by":"crossref","first-page":"55","DOI":"10.1007\/s12026-015-8671-z","article-title":"Are human endogenous retroviruses triggers of autoimmune diseases? Unveiling associations of three diseases and viral loci","volume":"64","author":"Nexo","year":"2016","journal-title":"Immunol. Res."},{"key":"ref_143","doi-asserted-by":"crossref","unstructured":"De la Hera, B., Varade, J., Garcia-Montojo, M., Alcina, A., Fedetz, M., Alloza, I., Astobiza, I., Leyva, L., Fernandez, O., and Izquierdo, G. (2014). Human endogenous retrovirus HERV-Fc1 association with multiple sclerosis susceptibility: A meta-analysis. PLoS ONE, 9.","DOI":"10.1371\/journal.pone.0090182"},{"key":"ref_144","doi-asserted-by":"crossref","first-page":"288","DOI":"10.1002\/ana.21117","article-title":"Environmental risk factors for multiple sclerosis. Part I: The role of infection","volume":"61","author":"Ascherio","year":"2007","journal-title":"Ann. Neurol."},{"key":"ref_145","doi-asserted-by":"crossref","first-page":"46","DOI":"10.1111\/ane.12431","article-title":"Multiple sclerosis and environmental factors: The role of vitamin D, parasites, and Epstein-Barr virus infection","volume":"132","author":"Correale","year":"2015","journal-title":"Acta Neurol. Scand."},{"key":"ref_146","doi-asserted-by":"crossref","first-page":"127","DOI":"10.1016\/j.coviro.2014.09.016","article-title":"Evidence linking HHV-6 with multiple sclerosis: An update","volume":"9","author":"Leibovitch","year":"2014","journal-title":"Curr. Opin. Virol."},{"key":"ref_147","doi-asserted-by":"crossref","first-page":"839","DOI":"10.1007\/s13365-017-0569-1","article-title":"Prevalence of a history of prior varicella\/herpes zoster infection in multiple sclerosis","volume":"23","author":"Manouchehrinia","year":"2017","journal-title":"J. Neurovirology"},{"key":"ref_148","doi-asserted-by":"crossref","first-page":"44","DOI":"10.1016\/j.clineuro.2013.12.020","article-title":"The participation of varicella zoster virus in relapses of multiple sclerosis","volume":"119","author":"Sotelo","year":"2014","journal-title":"Clin. Neurol. Neurosurg."},{"key":"ref_149","doi-asserted-by":"crossref","first-page":"27","DOI":"10.1177\/1352458513490545","article-title":"Oligoclonal bands in multiple sclerosis reactive against two herpesviruses and association with magnetic resonance imaging findings","volume":"20","author":"Virtanen","year":"2014","journal-title":"Mult. Scler."},{"key":"ref_150","doi-asserted-by":"crossref","first-page":"569","DOI":"10.1002\/ana.410090610","article-title":"Viral antibody activity of oligoclonal and polyclonal immunoglobulins synthesized within the central nervous system in multiple sclerosis","volume":"9","author":"Rostrom","year":"1981","journal-title":"Ann. Neurol."},{"key":"ref_151","doi-asserted-by":"crossref","first-page":"93","DOI":"10.1016\/j.jneuroim.2011.06.012","article-title":"Intrathecal human herpesvirus 6 antibodies in multiple sclerosis and other demyelinating diseases presenting as oligoclonal bands in cerebrospinal fluid","volume":"237","author":"Virtanen","year":"2011","journal-title":"J. Neuroimmunol."},{"key":"ref_152","doi-asserted-by":"crossref","first-page":"253","DOI":"10.1016\/0022-510X(79)90119-9","article-title":"Latent herpes simplex virus trigeminal ganglionic infection in mice and demyelination in the central nervous system","volume":"43","author":"Kristensson","year":"1979","journal-title":"J. Neurol. Sci."},{"key":"ref_153","doi-asserted-by":"crossref","first-page":"52","DOI":"10.1002\/ana.410220113","article-title":"Multifocal CNS demyelination following peripheral inoculation with herpes simplex virus type 1","volume":"22","author":"Kastrukoff","year":"1987","journal-title":"Ann. Neurol."},{"key":"ref_154","doi-asserted-by":"crossref","first-page":"432","DOI":"10.1097\/00005072-199207000-00005","article-title":"Herpes simplex virus type I (HSV I)-induced multifocal central nervous system (CNS) demyelination in mice","volume":"51","author":"Kastrukoff","year":"1992","journal-title":"J. Neuropathol. Exp. Neurol."},{"key":"ref_155","doi-asserted-by":"crossref","first-page":"4","DOI":"10.1186\/2042-4280-3-4","article-title":"The effect of mouse strain on herpes simplex virus type 1 (HSV-1) infection of the central nervous system (CNS)","volume":"3","author":"Kastrukoff","year":"2012","journal-title":"Herpesviridae"},{"key":"ref_156","doi-asserted-by":"crossref","first-page":"205","DOI":"10.1016\/0022-510X(82)90006-5","article-title":"Virus-induced demyelination in herpes simplex virus-infected mice","volume":"53","author":"Kristensson","year":"1982","journal-title":"J. Neurol. Sci."},{"key":"ref_157","doi-asserted-by":"crossref","unstructured":"Lee, D.H., Zandian, M., Kuo, J., Mott, K.R., Chen, S., Arditi, M., and Ghiasi, H. (2017). Suppression of IL-12p70 formation by IL-2 or following macrophage depletion causes T-cell autoreactivity leading to CNS demyelination in HSV-1-infected mice. PLoS Pathog., 13.","DOI":"10.1371\/journal.ppat.1006401"},{"key":"ref_158","doi-asserted-by":"crossref","first-page":"68","DOI":"10.1006\/exnr.2002.8035","article-title":"Demyelination associated with HSV-1-induced facial paralysis","volume":"178","author":"Wakisaka","year":"2002","journal-title":"Exp. Neurol."},{"key":"ref_159","doi-asserted-by":"crossref","unstructured":"Boukhvalova, M.S., Mortensen, E., Mbaye, A., Lopez, D., Kastrukoff, L., and Blanco, J.C.G. (2019). Herpes Simplex Virus 1 Induces Brain Inflammation and Multifocal Demyelination in the Cotton Rat Sigmodon hispidus. J. Virol., 94.","DOI":"10.1128\/JVI.01161-19"},{"key":"ref_160","doi-asserted-by":"crossref","first-page":"85","DOI":"10.1016\/0014-4886(64)90008-1","article-title":"Virus Isolated from the Brain of a Patient with Multiple Sclerosis","volume":"9","author":"Gudnadottir","year":"1964","journal-title":"Exp. Neurol."},{"key":"ref_161","doi-asserted-by":"crossref","first-page":"283","DOI":"10.1002\/ana.410260218","article-title":"Isolation of herpes simplex virus type 1 during first attack of multiple sclerosis","volume":"26","author":"Bergstrom","year":"1989","journal-title":"Ann. Neurol."},{"key":"ref_162","doi-asserted-by":"crossref","first-page":"249","DOI":"10.3109\/13550289609146888","article-title":"Detection of herpesviridae in postmortem multiple sclerosis brain tissue and controls by polymerase chain reaction","volume":"2","author":"Sanders","year":"1996","journal-title":"J. Neurovirology"},{"key":"ref_163","doi-asserted-by":"crossref","first-page":"125","DOI":"10.1001\/archneur.1996.00550020029012","article-title":"Herpes simplex virus in postmortem multiple sclerosis brain tissue","volume":"53","author":"Sanders","year":"1996","journal-title":"Arch. Neurol."},{"key":"ref_164","doi-asserted-by":"crossref","first-page":"437","DOI":"10.1007\/s13365-012-0110-5","article-title":"Active intrathecal herpes simplex virus type 1 (HSV-1) and human herpesvirus-6 (HHV-6) infection at onset of multiple sclerosis","volume":"18","author":"Ferro","year":"2012","journal-title":"J. Neurovirology"},{"key":"ref_165","doi-asserted-by":"crossref","first-page":"6","DOI":"10.1186\/s40893-017-0023-5","article-title":"Coincident onset of multiple sclerosis and Herpes simplex virus 1 encephalitis: A case report","volume":"2","author":"Buscarinu","year":"2017","journal-title":"Mult. Scler. Demyelinating Disord."},{"key":"ref_166","doi-asserted-by":"crossref","first-page":"31","DOI":"10.1212\/WNL.58.1.31","article-title":"A randomized, double-blind, placebo-controlled MRI study of anti-herpes virus therapy in MS","volume":"58","author":"Bech","year":"2002","journal-title":"Neurology"},{"key":"ref_167","first-page":"S109","article-title":"Molecular evidences for a role of HSV-1 in multiple sclerosis clinical acute attack","volume":"6","author":"Ferrante","year":"2000","journal-title":"J. Neurovirology"},{"key":"ref_168","first-page":"199","article-title":"Detection of human herpesviruses and polyomaviruses DNA in a group of patients with relapsing-remitting multiple sclerosis","volume":"28","author":"Pietropaolo","year":"2005","journal-title":"New Microbiol."},{"key":"ref_169","doi-asserted-by":"crossref","first-page":"229","DOI":"10.1159\/000496402","article-title":"Herpesviridae Seropositivity in Patients with Multiple Sclerosis: First Polish Study","volume":"80","author":"Czarnowska","year":"2018","journal-title":"Eur. Neurol."},{"key":"ref_170","doi-asserted-by":"crossref","first-page":"450","DOI":"10.1007\/s004150050121","article-title":"Human herpes virus 6 and human herpes virus 8 DNA sequences in brains of multiple sclerosis patients, normal adults and children","volume":"244","author":"Merelli","year":"1997","journal-title":"J. Neurol."},{"key":"ref_171","doi-asserted-by":"crossref","first-page":"253","DOI":"10.3109\/13550289509114021","article-title":"Human herpesvirus 6 polymerase chain reaction findings in human immunodeficiency virus associated neurological disease and multiple sclerosis","volume":"1","author":"Liedtke","year":"1995","journal-title":"J. Neurovirology"},{"key":"ref_172","doi-asserted-by":"crossref","first-page":"7440","DOI":"10.1073\/pnas.92.16.7440","article-title":"Plaque-associated expression of human herpesvirus 6 in multiple sclerosis","volume":"92","author":"Challoner","year":"1995","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_173","doi-asserted-by":"crossref","first-page":"2850","DOI":"10.1002\/jcp.26000","article-title":"Relationship of human herpes virus 6 and multiple sclerosis: A systematic review and meta-analysis","volume":"233","author":"Pormohammad","year":"2018","journal-title":"J. Cell. Physiol."},{"key":"ref_174","first-page":"4","article-title":"Human herpesvirus 6 and multiple sclerosis: Potential mechanisms for virus-induced disease","volume":"12","author":"Fotheringham","year":"2005","journal-title":"Herpes"},{"key":"ref_175","doi-asserted-by":"crossref","first-page":"373","DOI":"10.4103\/1673-5374.245462","article-title":"The role of Epstein-Barr virus in multiple sclerosis: From molecular pathophysiology to in vivo imaging","volume":"14","author":"Guan","year":"2019","journal-title":"Neural Regen. Res."},{"key":"ref_176","doi-asserted-by":"crossref","first-page":"239","DOI":"10.1007\/s00415-017-8656-z","article-title":"Low intrathecal antibody production despite high seroprevalence of Epstein-Barr virus in multiple sclerosis: A review of the literature","volume":"265","author":"Ruprecht","year":"2018","journal-title":"J. Neurol."},{"key":"ref_177","doi-asserted-by":"crossref","first-page":"940","DOI":"10.1038\/ni835","article-title":"A functional and structural basis for TCR cross-reactivity in multiple sclerosis","volume":"3","author":"Lang","year":"2002","journal-title":"Nat. Immunol."},{"key":"ref_178","doi-asserted-by":"crossref","first-page":"271","DOI":"10.1007\/s11481-010-9201-3","article-title":"Epstein-barr virus infection and multiple sclerosis: A review","volume":"5","author":"Ascherio","year":"2010","journal-title":"J. Neuroimmune Pharmacol."},{"key":"ref_179","doi-asserted-by":"crossref","unstructured":"Angelini, D.F., Serafini, B., Piras, E., Severa, M., Coccia, E.M., Rosicarelli, B., Ruggieri, S., Gasperini, C., Buttari, F., and Centonze, D. (2013). Increased CD8+ T cell response to Epstein-Barr virus lytic antigens in the active phase of multiple sclerosis. PLoS Pathog., 9.","DOI":"10.1371\/journal.ppat.1003220"},{"key":"ref_180","doi-asserted-by":"crossref","first-page":"e126","DOI":"10.1038\/cti.2016.87","article-title":"Defective T-cell control of Epstein-Barr virus infection in multiple sclerosis","volume":"6","author":"Pender","year":"2017","journal-title":"Clin. Transl. Immunol."},{"key":"ref_181","doi-asserted-by":"crossref","first-page":"1763","DOI":"10.1084\/jem.20072397","article-title":"EBNA1-specific T cells from patients with multiple sclerosis cross react with myelin antigens and co-produce IFN-gamma and IL-2","volume":"205","author":"Lunemann","year":"2008","journal-title":"J. Exp. Med."},{"key":"ref_182","doi-asserted-by":"crossref","first-page":"540","DOI":"10.1007\/s13337-018-0481-1","article-title":"Human herpesvirus 8 DNA detection and variant analysis in patients with multiple sclerosis","volume":"29","author":"Marashi","year":"2018","journal-title":"Virusdisease"},{"key":"ref_183","doi-asserted-by":"crossref","first-page":"731","DOI":"10.1089\/088922200308738","article-title":"Chromosomal distribution and coding capacity of the human endogenous retrovirus HERV-W family","volume":"16","author":"Voisset","year":"2000","journal-title":"Aids Res. Hum. Retrovir."},{"key":"ref_184","doi-asserted-by":"crossref","first-page":"465","DOI":"10.1016\/B978-0-444-53488-0.00022-5","article-title":"Human endogenous retroviruses and the nervous system","volume":"123","author":"Douville","year":"2014","journal-title":"Handb. Clin. Neurol."},{"key":"ref_185","doi-asserted-by":"crossref","first-page":"65","DOI":"10.1016\/j.jns.2009.04.034","article-title":"Endogenous retroviral genes, Herpesviruses and gender in Multiple Sclerosis","volume":"286","author":"Perron","year":"2009","journal-title":"J. Neurol. Sci."},{"key":"ref_186","doi-asserted-by":"crossref","first-page":"7852","DOI":"10.1128\/JVI.78.14.7852-7860.2004","article-title":"Epstein-Barr virus latent membrane protein LMP-2A is sufficient for transactivation of the human endogenous retrovirus HERV-K18 superantigen","volume":"78","author":"Sutkowski","year":"2004","journal-title":"J. Virol."},{"key":"ref_187","doi-asserted-by":"crossref","first-page":"2056","DOI":"10.4049\/jimmunol.177.4.2056","article-title":"Cutting edge: Epstein-Barr virus transactivates the HERV-K18 superantigen by docking to the human complement receptor 2 (CD21) on primary B cells","volume":"177","author":"Hsiao","year":"2006","journal-title":"J. Immunol"},{"key":"ref_188","doi-asserted-by":"crossref","first-page":"238","DOI":"10.1016\/S1471-4906(02)02207-X","article-title":"Human viral superantigens: To be or not to be transactivated?","volume":"23","author":"Lafon","year":"2002","journal-title":"Trends Immunol."},{"key":"ref_189","doi-asserted-by":"crossref","first-page":"551","DOI":"10.1016\/S0923-2516(89)80141-4","article-title":"Leptomeningeal cell line from multiple sclerosis with reverse transcriptase activity and viral particles","volume":"140","author":"Perron","year":"1989","journal-title":"Res. Virol."},{"key":"ref_190","doi-asserted-by":"crossref","first-page":"862","DOI":"10.1016\/0140-6736(91)92579-Q","article-title":"Isolation of retrovirus from patients with multiple sclerosis","volume":"337","author":"Perron","year":"1991","journal-title":"Lancet"},{"key":"ref_191","doi-asserted-by":"crossref","first-page":"337","DOI":"10.1016\/S0923-2516(06)80122-6","article-title":"In vitro transmission and antigenicity of a retrovirus isolated from a multiple sclerosis patient","volume":"143","author":"Perron","year":"1992","journal-title":"Res. Virol."},{"key":"ref_192","doi-asserted-by":"crossref","first-page":"65","DOI":"10.1099\/0022-1317-74-1-65","article-title":"Herpes simplex virus ICP0 and ICP4 immediate early proteins strongly enhance expression of a retrovirus harboured by a leptomeningeal cell line from a patient with multiple sclerosis","volume":"74","author":"Perron","year":"1993","journal-title":"J. Gen. Virol."},{"key":"ref_193","doi-asserted-by":"crossref","first-page":"7583","DOI":"10.1073\/pnas.94.14.7583","article-title":"Molecular identification of a novel retrovirus repeatedly isolated from patients with multiple sclerosis. The Collaborative Research Group on Multiple Sclerosis","volume":"94","author":"Perron","year":"1997","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_194","doi-asserted-by":"crossref","first-page":"863","DOI":"10.1016\/0140-6736(91)92580-U","article-title":"Just another dubious virus in cells from a patient with multiple sclerosis?","volume":"337","author":"Haahr","year":"1991","journal-title":"Lancet"},{"key":"ref_195","doi-asserted-by":"crossref","first-page":"33","DOI":"10.1016\/S0140-6736(98)24001-3","article-title":"Detection of virion-associated MSRV-RNA in serum of patients with multiple sclerosis","volume":"351","author":"Garson","year":"1998","journal-title":"Lancet"},{"key":"ref_196","doi-asserted-by":"crossref","first-page":"471","DOI":"10.1212\/WNL.58.3.471","article-title":"Multiple sclerosis-associated retrovirus (MSRV) in Sardinian MS patients","volume":"58","author":"Dolei","year":"2002","journal-title":"Neurology"},{"key":"ref_197","doi-asserted-by":"crossref","first-page":"112","DOI":"10.1080\/13550280390173355","article-title":"Multiple sclerosis-associated virus-related pol sequences found both in multiple sclerosis and healthy donors are more frequently expressed in multiple sclerosis patients","volume":"9","author":"Nowak","year":"2003","journal-title":"J. Neurovirology"},{"key":"ref_198","doi-asserted-by":"crossref","first-page":"1721","DOI":"10.1177\/1352458512441381","article-title":"Human endogenous retrovirus type W envelope expression in blood and brain cells provides new insights into multiple sclerosis disease","volume":"18","author":"Perron","year":"2012","journal-title":"Mult. Scler."},{"key":"ref_199","doi-asserted-by":"crossref","first-page":"1248","DOI":"10.1177\/1352458510376956","article-title":"Multiple sclerosis-associated retrovirus and progressive disability of multiple sclerosis","volume":"16","author":"Sotgiu","year":"2010","journal-title":"Mult. Scler."},{"key":"ref_200","doi-asserted-by":"crossref","first-page":"75","DOI":"10.1016\/S1016-8478(23)13710-1","article-title":"Activation of the human endogenous retrovirus W long terminal repeat by herpes simplex virus type 1 immediate early protein 1","volume":"15","author":"Lee","year":"2003","journal-title":"Mol. Cells"},{"key":"ref_201","doi-asserted-by":"crossref","first-page":"257","DOI":"10.1038\/6766","article-title":"An almost-intact human endogenous retrovirus K on human chromosome 7","volume":"21","author":"Mayer","year":"1999","journal-title":"Nat. Genet."},{"key":"ref_202","doi-asserted-by":"crossref","first-page":"1175","DOI":"10.1177\/1352458508094641","article-title":"Human endogenous retrovirus-K18 Env as a risk factor in multiple sclerosis","volume":"14","author":"Tai","year":"2008","journal-title":"Mult. Scler."},{"key":"ref_203","doi-asserted-by":"crossref","first-page":"93","DOI":"10.1016\/S0168-1702(02)00058-8","article-title":"Transactivation of the human endogenous retrovirus K long terminal repeat by herpes simplex virus type 1 immediate early protein 0","volume":"86","author":"Kwun","year":"2002","journal-title":"Virus Res."},{"key":"ref_204","doi-asserted-by":"crossref","first-page":"415","DOI":"10.1089\/088922204323048168","article-title":"Simultaneous presence of endogenous retrovirus and herpes virus antigens has profound effect on cell-mediated immune responses: Implications for multiple sclerosis","volume":"20","author":"Brudek","year":"2004","journal-title":"Aids Res. Hum. Retrovir."},{"key":"ref_205","doi-asserted-by":"crossref","first-page":"295","DOI":"10.1111\/j.1365-3083.2007.02067.x","article-title":"Synergistic immune responses induced by endogenous retrovirus and herpesvirus antigens result in increased production of inflammatory cytokines in multiple sclerosis patients","volume":"67","author":"Brudek","year":"2008","journal-title":"Scand. J. Immunol."}],"container-title":["International Journal of Molecular Sciences"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.mdpi.com\/1422-0067\/22\/11\/5738\/pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2025,10,11]],"date-time":"2025-10-11T06:09:21Z","timestamp":1760162961000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.mdpi.com\/1422-0067\/22\/11\/5738"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2021,5,27]]},"references-count":205,"journal-issue":{"issue":"11","published-online":{"date-parts":[[2021,6]]}},"alternative-id":["ijms22115738"],"URL":"https:\/\/doi.org\/10.3390\/ijms22115738","relation":{},"ISSN":["1422-0067"],"issn-type":[{"value":"1422-0067","type":"electronic"}],"subject":[],"published":{"date-parts":[[2021,5,27]]}}}