{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,9,19]],"date-time":"2026-09-19T07:57:04Z","timestamp":1789804624310,"version":"4.0.1"},"update-to":[{"DOI":"10.1371\/journal.ppat.1013514","type":"new_version","label":"New version","source":"publisher","updated":{"date-parts":[[2025,10,9]],"date-time":"2025-10-09T00:00:00Z","timestamp":1759968000000}}],"reference-count":50,"publisher":"Public Library of Science (PLoS)","issue":"9","license":[{"start":{"date-parts":[[2025,9,30]],"date-time":"2025-09-30T00:00:00Z","timestamp":1759190400000},"content-version":"vor","delay-in-days":0,"URL":"http:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"DOI":"10.13039\/100009619","name":"Japan Agency for Medical Research and Development","doi-asserted-by":"publisher","award":["23fk0108614h"],"award-info":[{"award-number":["23fk0108614h"]}],"id":[{"id":"10.13039\/100009619","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100020959","name":"JST-Mirai Program","doi-asserted-by":"publisher","award":["JPMJSP2111"],"award-info":[{"award-number":["JPMJSP2111"]}],"id":[{"id":"10.13039\/501100020959","id-type":"DOI","asserted-by":"publisher"}]}],"content-domain":{"domain":["www.plospathogens.org"],"crossmark-restriction":false},"short-container-title":["PLoS Pathog"],"abstract":"<jats:p>\n                    <jats:italic>Borrelia miyamotoi<\/jats:italic>\n                    is a hard tick-borne spirochete genetically related to relapsing fever\n                    <jats:italic>Borrelia<\/jats:italic>\n                    and the etiological agent of an emerging infectious disease in humans. Like relapsing fever\n                    <jats:italic>Borrelia<\/jats:italic>\n                    ,\n                    <jats:italic>B. miyamotoi<\/jats:italic>\n                    carries clusters of gene cassettes encoding variable major proteins (Vmps) on multiple linear plasmids and shows antigenic variation in mammalian hosts by switching the expression\n                    <jats:italic>vmp<\/jats:italic>\n                    gene cassette. However, it remains unknown how the switch occurs in\n                    <jats:italic>B. miyamotoi<\/jats:italic>\n                    . Here we determined the whole genome sequences of Japanese\n                    <jats:italic>B. miyamotoi<\/jats:italic>\n                    strains to identify the repertoire and arrangement of\n                    <jats:italic>vmp<\/jats:italic>\n                    gene cassettes on five linear plasmids, and based on this information, analyzed\n                    <jats:italic>B. miyamotoi<\/jats:italic>\n                    clones reisolated from experimentally infected mice. Our analyses revealed that the switch occurred by replacing the expression cassette and its downstream silent cassettes with the long segment from archival plasmid. As the result of this long segment conversion, the first cassette became the expression cassette. Notably, this phenomenon was not due to single gene conversion but the replacement of a long (up to 16\u2009kb or more) plasmid segment. We also show that while bacterial elimination depended on the presence of specific antibodies, the segment conversion was detected at five days post-infection, earlier than antibody production in mice, and even in severe combined immunodeficient mice. These results provide novel insights into the mechanisms that\n                    <jats:italic>Borrelia<\/jats:italic>\n                    evolved to survive and persist in mammalian hosts.\n                  <\/jats:p>","DOI":"10.1371\/journal.ppat.1013514","type":"journal-article","created":{"date-parts":[[2025,9,30]],"date-time":"2025-09-30T17:45:55Z","timestamp":1759254355000},"page":"e1013514","update-policy":"https:\/\/doi.org\/10.1371\/journal.ppat.corrections_policy","source":"Crossref","is-referenced-by-count":3,"title":["Antigenic variation is caused by long plasmid segment conversion in a hard tick-borne relapsing fever Borrelia miyamotoi"],"prefix":"10.1371","volume":"21","author":[{"given":"Tomohi","family":"Takeuchi","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Yasuhiro","family":"Gotoh","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Tetsuya","family":"Hayashi","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Hiroki","family":"Kawabata","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-5919-3738","authenticated-orcid":true,"given":"Ai","family":"Takano","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"340","published-online":{"date-parts":[[2025,9,30]]},"reference":[{"key":"ppat.1013514.ref001","doi-asserted-by":"crossref","first-page":"11","DOI":"10.1016\/j.ttbdis.2012.06.007","article-title":"Exploring gaps in our knowledge on Lyme borreliosis spirochaetes--updates on complex heterogeneity, ecology, and pathogenicity","volume":"4","author":"J Franke","year":"2013","journal-title":"Ticks Tick Borne Dis"},{"issue":"1","key":"ppat.1013514.ref002","doi-asserted-by":"crossref","first-page":"134","DOI":"10.1111\/j.1462-2920.2009.02054.x","article-title":"Isolation and characterization of a novel Borrelia group of tick-borne borreliae from imported reptiles and their associated ticks","volume":"12","author":"A Takano","year":"2010","journal-title":"Environ Microbiol"},{"issue":"1","key":"ppat.1013514.ref003","doi-asserted-by":"crossref","first-page":"105","DOI":"10.1186\/s12862-021-01838-1","article-title":"The evolution of hard tick-borne relapsing fever borreliae is correlated with vector species rather than geographical distance","volume":"21","author":"R Nakao","year":"2021","journal-title":"BMC Ecol Evol"},{"issue":"4","key":"ppat.1013514.ref004","doi-asserted-by":"crossref","first-page":"804","DOI":"10.1099\/00207713-45-4-804","article-title":"Genetic and phenotypic analysis of Borrelia miyamotoi sp. nov., isolated from the ixodid tick Ixodes persulcatus, the vector for Lyme disease in Japan","volume":"45","author":"M Fukunaga","year":"1995","journal-title":"Int J Syst Bacteriol"},{"issue":"3","key":"ppat.1013514.ref005","doi-asserted-by":"crossref","first-page":"240","DOI":"10.1056\/NEJMoa1209039","article-title":"Meningoencephalitis from Borrelia miyamotoi in an immunocompromised patient","volume":"368","author":"JL Gugliotta","year":"2013","journal-title":"N Engl J Med"},{"issue":"9892","key":"ppat.1013514.ref006","doi-asserted-by":"crossref","first-page":"658","DOI":"10.1016\/S0140-6736(13)61644-X","article-title":"A case of meningoencephalitis by the relapsing fever spirochaete Borrelia miyamotoi in Europe","volume":"382","author":"JWR Hovius","year":"2013","journal-title":"Lancet"},{"issue":"10","key":"ppat.1013514.ref007","doi-asserted-by":"crossref","first-page":"1816","DOI":"10.3201\/eid1710.101474","article-title":"Humans infected with relapsing fever spirochete Borrelia miyamotoi, Russia","volume":"17","author":"AE Platonov","year":"2011","journal-title":"Emerg Infect Dis"},{"issue":"8","key":"ppat.1013514.ref008","doi-asserted-by":"crossref","first-page":"1391","DOI":"10.3201\/eid2008.131761","article-title":"Human infections with Borrelia miyamotoi, Japan","volume":"20","author":"K Sato","year":"2014","journal-title":"Emerg Infect Dis"},{"issue":"5","key":"ppat.1013514.ref009","doi-asserted-by":"crossref","first-page":"1297","DOI":"10.1084\/jem.156.5.1297","article-title":"Antigenic variation of Borrelia hermsii","volume":"156","author":"HG Stoenner","year":"1982","journal-title":"J Exp Med"},{"issue":"1","key":"ppat.1013514.ref010","doi-asserted-by":"crossref","first-page":"29","DOI":"10.1093\/infdis\/117.1.29","article-title":"Experimental relapsing fever initiated by Borrelia hermsi. 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