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The conjugation operon present on the Bacillus subtilis conjugative plasmid pLS20 is over 32 kb long. This operon starts with a 456-nt leader region, which is followed by the first two genes of the operon, encoding a two-component processive antitermination system. Here, we demonstrate that the long leader region encodes a transcriptional attenuator that we named cATTpLS20. In vivo and in vitro analyses showed that the attenuator is composed of three segments: a stem\u2013loop structure with a long imperfect stem that is preceded by a sequence that may form a weak stem\u2013loop and followed by a sequence that may form an intrinsic terminator. Sequences of the upstream stem\u2013loop and downstream terminator are complementary, allowing further extension of the long stem, thereby generating an antiterminator conformation. Based on similarity with a zipper, we coined this the zipper-type attenuator. Similar zipper-type attenuators are present upstream of the first genes of conjugation operons of all pLS20 family plasmids, as well as on many other conjugative plasmids in Gram-positive bacteria, suggesting that a common attenuation mechanism regulates expression of many conjugative operons.<\/jats:p>","DOI":"10.1093\/nar\/gkag726","type":"journal-article","created":{"date-parts":[[2026,7,9]],"date-time":"2026-07-09T11:59:12Z","timestamp":1783598352000},"source":"Crossref","is-referenced-by-count":0,"title":["Expression of conjugation genes is controlled by a novel zipper-type transcriptional attenuation mechanism"],"prefix":"10.1093","volume":"54","author":[{"given":"Daniel","family":"Gonz\u00e1lez-\u00c1lvarez","sequence":"first","affiliation":[{"name":"Microbes in health and welfare, Centro de Biolog\u00eda Molecular Severo Ochoa (CSIC-UAM), C. Nicol\u00e1s Cabrera 1, Universidad Aut\u00f3noma de Madrid , Cantoblanco, Madrid 28049 ,","place":["Spain"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-5679-9083","authenticated-orcid":false,"given":"Andr\u00e9s","family":"Miguel-Arribas","sequence":"additional","affiliation":[{"name":"Microbes in health and welfare, Centro de Biolog\u00eda Molecular Severo Ochoa (CSIC-UAM), C. 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Nicol\u00e1s Cabrera 1, Universidad Aut\u00f3noma de Madrid , Cantoblanco, Madrid 28049 ,","place":["Spain"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"David","family":"Abia","sequence":"additional","affiliation":[{"name":"Bioinformatics Facility, Centro de Biolog\u00eda Molecular Severo Ochoa, (CSIC-UAM), C. Nicol\u00e1s Cabrera 1, Universidad Aut\u00f3noma de Madrid , Canto Blanco, Madrid 28049 ,","place":["Spain"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-7024-2334","authenticated-orcid":false,"given":"Anne","family":"de\u00a0Jong","sequence":"additional","affiliation":[{"name":"Department of Molecular Genetics, Groningen Biomolecular Sciences and Biotechnology Institute, University of Groningen , Groningen 9747AG ,","place":["The Netherlands"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Ling Juan","family":"Wu","sequence":"additional","affiliation":[{"name":"Centre for Bacterial Cell Biology, Biosciences Institute, Newcastle University , Richardson Road, Newcastle Upon Tyne NE2 4AX ,","place":["United Kingdom"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-2481-1062","authenticated-orcid":false,"given":"Paul","family":"Babitzke","sequence":"additional","affiliation":[{"name":"Department of Biochemistry and Molecular Biology, Center for RNA Molecular Biology, Pennsylvania State University, University Park , PA 16802 ,","place":["United States"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-1842-0049","authenticated-orcid":false,"given":"Wilfried J J","family":"Meijer","sequence":"additional","affiliation":[{"name":"Microbes in health and welfare, Centro de Biolog\u00eda Molecular Severo Ochoa (CSIC-UAM), C. 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