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Temporal expression of chlamydial genes during the intracellular infection is proposed to be regulated by changes in <jats:styled-content style=\"fixed-case\">DNA<\/jats:styled-content> supercoiling levels. To understand how chlamydial supercoiling levels are regulated, we purified and analyzed three putative <jats:styled-content style=\"fixed-case\"><jats:italic>C<\/jats:italic><\/jats:styled-content><jats:italic>hlamydia trachomatis<\/jats:italic> topoisomerases. As predicted by sequence homology, <jats:styled-content style=\"fixed-case\">CT<\/jats:styled-content>189\/190 are the two subunits of <jats:styled-content style=\"fixed-case\">DNA<\/jats:styled-content> gyrase, whereas <jats:styled-content style=\"fixed-case\">CT<\/jats:styled-content>643 is a topoisomerase <jats:styled-content style=\"fixed-case\">I<\/jats:styled-content>. <jats:styled-content style=\"fixed-case\">CT<\/jats:styled-content>660\/661 have been predicted to form a second <jats:styled-content style=\"fixed-case\">DNA<\/jats:styled-content> gyrase, but the reconstitute holoenzyme decatenated and relaxed <jats:styled-content style=\"fixed-case\">DNA<\/jats:styled-content>, indicating that the proteins are subunits of topoisomerase <jats:styled-content style=\"fixed-case\">IV<\/jats:styled-content>. Promoter analysis showed that each topoisomerase is transcribed from its own operon by the major chlamydial <jats:styled-content style=\"fixed-case\">RNA<\/jats:styled-content> polymerase. Surprisingly, all three topoisomerase promoters had higher activity from a more supercoiled <jats:styled-content style=\"fixed-case\">DNA<\/jats:styled-content> template. This supercoiling\u2010responsivesness is consistent with negative feedback control of topoisomerase <jats:styled-content style=\"fixed-case\">I<\/jats:styled-content> and topoisomerase <jats:styled-content style=\"fixed-case\">IV<\/jats:styled-content> expression, which is typical of other bacteria. However, activation of the chlamydial gyrase promoter by increased supercoiling is unorthodox compared with the relaxation\u2010induced transcription of gyrase in other bacteria. We present a model in which supercoiling levels during the intracellular chlamydial developmental cycle are regulated by unusual positive feedback control of the gyrase promoter and the temporal expression of three topoisomerases.<\/jats:p>","DOI":"10.1111\/mmi.13241","type":"journal-article","created":{"date-parts":[[2015,10,8]],"date-time":"2015-10-08T13:41:46Z","timestamp":1444311706000},"page":"484-496","source":"Crossref","is-referenced-by-count":14,"title":["Functional analysis of three topoisomerases that regulate <scp>DNA<\/scp> supercoiling levels in <scp><i>C<\/i><\/scp><i>hlamydia<\/i>"],"prefix":"10.1111","volume":"99","author":[{"ORCID":"https:\/\/orcid.org\/0000-0002-3872-0236","authenticated-orcid":false,"given":"Emilie","family":"Orillard","sequence":"first","affiliation":[{"name":"Department of Microbiology and Molecular Genetics University of California  Irvine CA 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