{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,9,29]],"date-time":"2026-09-29T13:30:04Z","timestamp":1790688604717,"version":"4.1.0"},"reference-count":18,"publisher":"Springer Science and Business Media LLC","issue":"1","license":[{"start":{"date-parts":[[2016,3,9]],"date-time":"2016-03-09T00:00:00Z","timestamp":1457481600000},"content-version":"tdm","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0"},{"start":{"date-parts":[[2016,3,9]],"date-time":"2016-03-09T00:00:00Z","timestamp":1457481600000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0"}],"content-domain":{"domain":["link.springer.com"],"crossmark-restriction":false},"short-container-title":["Sci Rep"],"abstract":"<jats:title>Abstract<\/jats:title><jats:p><jats:italic>Vibrio cholerae<\/jats:italic> O1 El Tor biotype strains are responsible for three multiyear epidemics of cholera in China during the seventh ongoing pandemic. The presence of the integrative conjugative element SXT is strongly correlated with resistance to nalidixic acid, tetracycline, and trimethoprim-sulfamethoxazole in these strains. Here, we sequenced the conserved genes of the SXT element, including <jats:italic>eex<\/jats:italic>, <jats:italic>setR<\/jats:italic>, and <jats:italic>int<\/jats:italic>, from 59 <jats:italic>V. cholerae<\/jats:italic> O1 El Tor strains and extracted and assembled the intact SXT sequences from the 11 genome sequenced strains. These elements had characteristics distinct from those of previously reported integrative conjugative elements (ICEs). They could be clearly divided into two types based on the clustering of conserved genes and gene structures of the elements, showing their possibly independent derivation and evolution. These two types were present before and after 2005, respectively, demonstrating the type substitution that occurred in 2005. Four to six antibiotic-resistant genes were found on the SXT elements, including genes resistant to tetracycline, trimethoprim-sulfamethoxazole, and multiple drugs. In summary, our findings demonstrated the roles of the SXT element in the emergence of multidrug resistance in epidemic O1 El Tor <jats:italic>V. cholerae<\/jats:italic> strains in China.<\/jats:p>","DOI":"10.1038\/srep22733","type":"journal-article","created":{"date-parts":[[2016,3,9]],"date-time":"2016-03-09T10:09:57Z","timestamp":1457518197000},"update-policy":"https:\/\/doi.org\/10.1007\/springer_crossmark_policy","source":"Crossref","is-referenced-by-count":50,"title":["Variations in SXT elements in epidemic Vibrio cholerae O1 El Tor strains in China"],"prefix":"10.1038","volume":"6","author":[{"given":"Ruibai","family":"Wang","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Dong","family":"Yu","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Junjie","family":"Yue","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Biao","family":"Kan","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"297","published-online":{"date-parts":[[2016,3,9]]},"reference":[{"key":"BFsrep22733_CR1","doi-asserted-by":"publisher","first-page":"361","DOI":"10.1016\/j.ijantimicag.2012.06.010","volume":"40","author":"R Wang","year":"2012","unstructured":"Wang, R. et al. Antibiotic resistance of Vibrio cholerae O1 El Tor strains from the seventh pandemic in China, 1961\u20132010. Int. J. Antimicrob. Agents. 40, 361\u2013364 (2012).","journal-title":"Int. J. Antimicrob. Agents"},{"key":"BFsrep22733_CR2","doi-asserted-by":"publisher","first-page":"4259","DOI":"10.1128\/JB.184.15.4259-4269.2002","volume":"184","author":"JW Beaber","year":"2002","unstructured":"Beaber, J. W., Hochhut, B. & Waldor, M. K. Genomic and functional analyses of SXT, an integrating antibiotic resistance gene transfer element derived from Vibrio cholerae . J. Bacteriol. 184, 4259\u20134269 (2002).","journal-title":"J. Bacteriol."},{"key":"BFsrep22733_CR3","doi-asserted-by":"publisher","first-page":"5158","DOI":"10.1128\/JB.184.18.5158-5169.2002","volume":"184","author":"D Boltner","year":"2002","unstructured":"Boltner, D., MacMahon, C., Pembroke, J. T., Strike, P. & Osborn, A. M. R391: a conjugative integrating mosaic comprised of phage, plasmid, and transposon elements. J. Bacteriol. 184, 5158\u20135169 (2002).","journal-title":"J. Bacteriol."},{"key":"BFsrep22733_CR4","doi-asserted-by":"publisher","first-page":"173","DOI":"10.1016\/j.plasmid.2006.01.001","volume":"55","author":"V Burrus","year":"2006","unstructured":"Burrus, V., Marrero, J. & Waldor, M. K. The current ICE age: biology and evolution of SXT-related integrating conjugative elements. Plasmid. 55, 173\u2013183 (2006).","journal-title":"Plasmid."},{"key":"BFsrep22733_CR5","doi-asserted-by":"publisher","first-page":"99","DOI":"10.1046\/j.1365-2958.1999.01330.x","volume":"32","author":"B Hochhut","year":"1999","unstructured":"Hochhut, B. & Waldor, M. K. Site-specific integration of the conjugal Vibrio cholerae SXT element into prfC . Mol. Microbiol. 32, 99\u2013110 (1999).","journal-title":"Mol. Microbiol."},{"key":"BFsrep22733_CR6","doi-asserted-by":"publisher","first-page":"4157","DOI":"10.1128\/jb.178.14.4157-4165.1996","volume":"178","author":"MK Waldor","year":"1996","unstructured":"Waldor, M. K., Tschape, H. & Mekalanos, J. J. A new type of conjugative transposon encodes resistance to sulfamethoxazole, trimethoprim, and streptomycin in Vibrio cholerae O139. J. Bacteriol. 178, 4157\u20134165 (1996).","journal-title":"J. Bacteriol."},{"key":"BFsrep22733_CR7","doi-asserted-by":"publisher","first-page":"510","DOI":"10.1111\/j.1462-2920.2009.02094.x","volume":"12","author":"E Bordeleau","year":"2010","unstructured":"Bordeleau, E., Brouillette, E., Robichaud, N. & Burrus, V. Beyond antibiotic resistance: integrating conjugative elements of the SXT\/R391 family that encode novel diguanylate cyclases participate to c-di-GMP signalling in Vibrio cholerae . Environ. Microbiol. 12, 510\u2013523 (2010).","journal-title":"Environ. Microbiol."},{"key":"BFsrep22733_CR8","doi-asserted-by":"publisher","first-page":"e1000786","DOI":"10.1371\/journal.pgen.1000786","volume":"5","author":"RA Wozniak","year":"2009","unstructured":"Wozniak, R. A. et al. Comparative ICE genomics: insights into the evolution of the SXT\/R391 family of ICEs. PLoS Genet. 5, e1000786 (2009).","journal-title":"PLoS Genet."},{"key":"BFsrep22733_CR9","doi-asserted-by":"publisher","first-page":"e1005072","DOI":"10.1371\/journal.pgen.1005072","volume":"11","author":"X Didelot","year":"2015","unstructured":"Didelot, X. et al. The role of china in the global spread of the current cholera pandemic. PLoS Genet. 11, e1005072 (2015).","journal-title":"PLoS Genet."},{"key":"BFsrep22733_CR10","doi-asserted-by":"publisher","first-page":"241","DOI":"10.1016\/j.femsle.2004.11.012","volume":"242","author":"AM Ahmed","year":"2005","unstructured":"Ahmed, A. M., Shinoda, S. & Shimamoto, T. A variant type of Vibrio cholerae SXT element in a multidrug-resistant strain of Vibrio fluvialis . FEMS Microbiol. Lett. 242, 241\u2013247 (2005).","journal-title":"FEMS Microbiol. Lett."},{"key":"BFsrep22733_CR11","doi-asserted-by":"publisher","first-page":"6469","DOI":"10.1128\/JB.00522-07","volume":"189","author":"J Marrero","year":"2007","unstructured":"Marrero, J. & Waldor, M. K. Determinants of entry exclusion within Eex and TraG are cytoplasmic. J. Bacteriol. 189, 6469\u20136473 (2007).","journal-title":"J. Bacteriol."},{"key":"BFsrep22733_CR12","doi-asserted-by":"publisher","first-page":"e1000439","DOI":"10.1371\/journal.pgen.1000439","volume":"5","author":"RA Wozniak","year":"2009","unstructured":"Wozniak, R. A. & Waldor, M. K. A toxin-antitoxin system promotes the maintenance of an integrative conjugative element. PLoS Genet. 5, e1000439 (2009).","journal-title":"PLoS Genet."},{"key":"BFsrep22733_CR13","doi-asserted-by":"publisher","first-page":"3054","DOI":"10.1128\/AEM.72.4.3054-3057.2006","volume":"72","author":"V Burrus","year":"2006","unstructured":"Burrus, V., Quezada-Calvillo, R., Marrero, J. & Waldor, M. K. SXT-related integrating conjugative element in New World Vibrio cholerae . Appl. Environ. Microbiol. 72, 3054\u20133057 (2006).","journal-title":"Appl. Environ. Microbiol."},{"key":"BFsrep22733_CR14","doi-asserted-by":"publisher","first-page":"80","DOI":"10.1111\/j.1574-6968.2006.00452.x","volume":"264","author":"JT Pembroke","year":"2006","unstructured":"Pembroke, J. T. & Piterina, A. V. A novel ICE in the genome of Shewanella putrefaciens W3-18-1: comparison with the SXT\/R391 ICE-like elements. FEMS Microbiol. Lett. 264, 80\u201388 (2006).","journal-title":"FEMS Microbiol. Lett."},{"key":"BFsrep22733_CR15","doi-asserted-by":"publisher","first-page":"457","DOI":"10.1111\/1574-6941.12007","volume":"83","author":"M Balado","year":"2013","unstructured":"Balado, M., Lemos, M. L. & Osorio, C. R. Integrating conjugative elements of the SXT\/R391 family from fish-isolated Vibrios encode restriction-modification systems that confer resistance to bacteriophages. FEMS Microbiol. Ecol. 83, 457\u2013467 (2013).","journal-title":"FEMS Microbiol. Ecol."},{"key":"BFsrep22733_CR16","doi-asserted-by":"publisher","first-page":"4027","DOI":"10.1128\/JB.01981-07","volume":"190","author":"MM Pearson","year":"2008","unstructured":"Pearson, M. M. et al. Complete genome sequence of uropathogenic Proteus mirabilis, a master of both adherence and motility. J. Bacteriol. 190, 4027\u20134037 (2008).","journal-title":"J. Bacteriol."},{"key":"BFsrep22733_CR17","doi-asserted-by":"publisher","first-page":"10485","DOI":"10.1073\/pnas.1509097112","volume":"112","author":"AB Dalia","year":"2015","unstructured":"Dalia, A. B., Seed, K. D., Calderwood, S. B. & Camilli, A. A globally distributed mobile genetic element inhibits natural transformation of Vibrio cholerae . Proc. Natl. Acad. Sci. USA 112, 10485\u201310490 (2015).","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"BFsrep22733_CR18","doi-asserted-by":"publisher","first-page":"2991","DOI":"10.1128\/AAC.45.11.2991-3000.2001","volume":"45","author":"B Hochhut","year":"2001","unstructured":"Hochhut, B. et al. Molecular analysis of antibiotic resistance gene clusters in Vibrio cholerae O139 and O1 SXT constins. Antimicrob. Agents Chemother. 45, 2991\u20133000 (2001).","journal-title":"Antimicrob. Agents Chemother."}],"container-title":["Scientific Reports"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.nature.com\/articles\/srep22733.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/www.nature.com\/articles\/srep22733","content-type":"text\/html","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/www.nature.com\/articles\/srep22733.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2023,1,5]],"date-time":"2023-01-05T03:54:38Z","timestamp":1672890878000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.nature.com\/articles\/srep22733"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2016,3,9]]},"references-count":18,"journal-issue":{"issue":"1","published-online":{"date-parts":[[2016,3,31]]}},"alternative-id":["BFsrep22733"],"URL":"https:\/\/doi.org\/10.1038\/srep22733","relation":{},"ISSN":["2045-2322"],"issn-type":[{"value":"2045-2322","type":"electronic"}],"subject":[],"published":{"date-parts":[[2016,3,9]]},"assertion":[{"value":"9 November 2015","order":1,"name":"received","label":"Received","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"18 February 2016","order":2,"name":"accepted","label":"Accepted","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"9 March 2016","order":3,"name":"first_online","label":"First Online","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"The authors declare no competing financial interests.","order":1,"name":"Ethics","group":{"name":"EthicsHeading","label":"Competing interests"}}],"article-number":"22733"}}