{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,10,6]],"date-time":"2026-10-06T18:51:12Z","timestamp":1791312672917,"version":"4.3.1"},"reference-count":57,"publisher":"National Academy of Sciences","issue":"41","license":[{"start":{"date-parts":[[2026,10,6]],"date-time":"2026-10-06T00:00:00Z","timestamp":1791244800000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by-nc-nd\/4.0\/"}],"funder":[{"DOI":"10.13039\/501100001665","name":"Agence Nationale de la Recherche","doi-asserted-by":"publisher","award":["ANR-20-CE12-0004"],"award-info":[{"award-number":["ANR-20-CE12-0004"]}],"id":[{"id":"10.13039\/501100001665","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100001665","name":"Agence Nationale de la Recherche","doi-asserted-by":"publisher","award":["ANR-17-CE13-0031"],"award-info":[{"award-number":["ANR-17-CE13-0031"]}],"id":[{"id":"10.13039\/501100001665","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100001665","name":"Agence Nationale de la Recherche","doi-asserted-by":"publisher","award":["ANR-22-CE44-0044"],"award-info":[{"award-number":["ANR-22-CE44-0044"]}],"id":[{"id":"10.13039\/501100001665","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100001665","name":"Agence Nationale de la Recherche","doi-asserted-by":"publisher","award":["ANR-10-LABX-62-IBEID"],"award-info":[{"award-number":["ANR-10-LABX-62-IBEID"]}],"id":[{"id":"10.13039\/501100001665","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100002915","name":"Fondation pour la Recherche M\u00e9dicale","doi-asserted-by":"publisher","award":["FDT202001010890"],"award-info":[{"award-number":["FDT202001010890"]}],"id":[{"id":"10.13039\/501100002915","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/100018694","name":"EC | Horizon Europe | Excellent Science | HORIZON EUROPE Marie Sklodowska-Curie Actions","doi-asserted-by":"publisher","award":["101208987"],"award-info":[{"award-number":["101208987"]}],"id":[{"id":"10.13039\/100018694","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/100000057","name":"HHS | NIH | National Institute of General Medical Sciences","doi-asserted-by":"publisher","award":["R35GM128674"],"award-info":[{"award-number":["R35GM128674"]}],"id":[{"id":"10.13039\/100000057","id-type":"DOI","asserted-by":"publisher"}]}],"content-domain":{"domain":["www.pnas.org"],"crossmark-restriction":true},"short-container-title":["Proc. Natl. Acad. Sci. U.S.A."],"published-print":{"date-parts":[[2026,10,13]]},"abstract":"<jats:p>Natural transformation is a widespread mechanism driving genetic exchanges in bacteria. It proceeds from the capture of extracellular DNA released by other cells. Internalized in linear single strands, this exogenous DNA is ultimately integrated into the genome by homologous recombination. It is unknown how the RecA-directed D-loop intermediate of this dedicated recombination pathway is processed. We report that resolution of the transformation D-loop depends on two endonucleases of opposing phylogenetic distribution in bacteria. One is YraN, which has coevolved and interacts with the ComM helicase, known to extend DNA recombination at the transformation D-loop. The other is CoiA, which is restricted to the Bacillota. CoiA is shown to be a resolvase of the transformation D-loop, extended by the RadA helicase in these species. We demonstrate that both YraN and CoiA act synergistically with their cognate helicases. These findings reveal that bacteria have evolved two helicase\/nuclease pairs for the maturation and recombination extension of the transformation D-loop. The wide distribution of this dual system indicates that recombination of exogenous DNA by natural transformation is an ancestral and pervasive feature of bacteria.<\/jats:p>","DOI":"10.1073\/pnas.2610795123","type":"journal-article","created":{"date-parts":[[2026,10,6]],"date-time":"2026-10-06T18:04:38Z","timestamp":1791309878000},"update-policy":"https:\/\/doi.org\/10.1073\/pnas.cm10313","source":"Crossref","is-referenced-by-count":0,"title":["Two D-loop resolution systems enable natural genetic transformation in bacteria"],"prefix":"10.1073","volume":"123","author":[{"ORCID":"https:\/\/orcid.org\/0009-0006-7852-4367","authenticated-orcid":false,"given":"L\u00e9o","family":"Hardy","sequence":"first","affiliation":[{"id":[{"id":"https:\/\/ror.org\/04zmssz18","id-type":"ROR","asserted-by":"publisher"}],"name":"Centre International de Recherche end Infectiologie, Inserm, U1111, Universit\u00e9 Claude Bernard Lyon 1, CNRS, UMR5308, \u00c9cole Normale Sup\u00e9rieure de Lyon, Univ Lyon","place":["Villeurbanne, France"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0009-0008-8868-5273","authenticated-orcid":false,"given":"Violette","family":"Morales","sequence":"additional","affiliation":[{"id":[{"id":"https:\/\/ror.org\/04rrj3a80","id-type":"ROR","asserted-by":"publisher"}],"name":"Laboratoire de Microbiologie et G\u00e9n\u00e9tique Mol\u00e9culaires, UMR5100, Centre de Biologie Int\u00e9grative, Centre Nationale de la Recherche Scientifique","place":["Toulouse, France"]},{"id":[{"id":"https:\/\/ror.org\/01ahyrz84","id-type":"ROR","asserted-by":"publisher"}],"name":"Universit\u00e9 de Toulouse","place":["Toulouse, France"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-2278-5112","authenticated-orcid":false,"given":"Clothilde J.","family":"Rousseau","sequence":"additional","affiliation":[{"id":[{"id":"https:\/\/ror.org\/04zmssz18","id-type":"ROR","asserted-by":"publisher"}],"name":"Centre International de Recherche end Infectiologie, Inserm, U1111, Universit\u00e9 Claude Bernard Lyon 1, CNRS, UMR5308, \u00c9cole Normale Sup\u00e9rieure de Lyon, Univ Lyon","place":["Villeurbanne, France"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-9555-5081","authenticated-orcid":false,"given":"Mathieu","family":"Berg\u00e9","sequence":"additional","affiliation":[{"id":[{"id":"https:\/\/ror.org\/04rrj3a80","id-type":"ROR","asserted-by":"publisher"}],"name":"Laboratoire de Microbiologie et G\u00e9n\u00e9tique Mol\u00e9culaires, UMR5100, Centre de Biologie Int\u00e9grative, Centre Nationale de la Recherche Scientifique","place":["Toulouse, France"]},{"id":[{"id":"https:\/\/ror.org\/01ahyrz84","id-type":"ROR","asserted-by":"publisher"}],"name":"Universit\u00e9 de Toulouse","place":["Toulouse, France"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0009-0008-2746-4901","authenticated-orcid":false,"given":"Fanny","family":"Mazzamurro","sequence":"additional","affiliation":[{"id":[{"id":"https:\/\/ror.org\/05f82e368","id-type":"ROR","asserted-by":"publisher"}],"name":"Institut Pasteur, CNRS UMR3525, Microbial Evolutionary Genomics, Universit\u00e9 Paris Cit\u00e9","place":["Paris, France"]},{"id":[{"id":"https:\/\/ror.org\/02en5vm52","id-type":"ROR","asserted-by":"publisher"}],"name":"Coll\u00e8ge Doctoral\u2013Sorbonne Universit\u00e9","place":["Paris, France"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0009-0009-3174-4065","authenticated-orcid":false,"given":"Julie","family":"Plantade","sequence":"additional","affiliation":[{"id":[{"id":"https:\/\/ror.org\/04zmssz18","id-type":"ROR","asserted-by":"publisher"}],"name":"Centre International de Recherche end Infectiologie, Inserm, U1111, Universit\u00e9 Claude Bernard Lyon 1, CNRS, UMR5308, \u00c9cole Normale Sup\u00e9rieure de Lyon, Univ Lyon","place":["Villeurbanne, France"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-7614-1377","authenticated-orcid":false,"given":"Triana N.","family":"Dalia","sequence":"additional","affiliation":[{"name":"Department of Biology, Indiana University","place":["Bloomington"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-2203-1230","authenticated-orcid":false,"given":"Ankur B.","family":"Dalia","sequence":"additional","affiliation":[{"name":"Department of Biology, Indiana University","place":["Bloomington"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-7704-822X","authenticated-orcid":false,"given":"Eduardo P. C.","family":"Rocha","sequence":"additional","affiliation":[{"id":[{"id":"https:\/\/ror.org\/05f82e368","id-type":"ROR","asserted-by":"publisher"}],"name":"Institut Pasteur, CNRS UMR3525, Microbial Evolutionary Genomics, Universit\u00e9 Paris Cit\u00e9","place":["Paris, France"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-9039-8021","authenticated-orcid":false,"given":"Xavier","family":"Charpentier","sequence":"additional","affiliation":[{"id":[{"id":"https:\/\/ror.org\/04zmssz18","id-type":"ROR","asserted-by":"publisher"}],"name":"Centre International de Recherche end Infectiologie, Inserm, U1111, Universit\u00e9 Claude Bernard Lyon 1, CNRS, UMR5308, \u00c9cole Normale Sup\u00e9rieure de Lyon, Univ Lyon","place":["Villeurbanne, France"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-0365-4347","authenticated-orcid":false,"given":"Patrice","family":"Polard","sequence":"additional","affiliation":[{"id":[{"id":"https:\/\/ror.org\/04rrj3a80","id-type":"ROR","asserted-by":"publisher"}],"name":"Laboratoire de Microbiologie et G\u00e9n\u00e9tique Mol\u00e9culaires, UMR5100, Centre de Biologie Int\u00e9grative, Centre Nationale de la Recherche Scientifique","place":["Toulouse, France"]},{"id":[{"id":"https:\/\/ror.org\/01ahyrz84","id-type":"ROR","asserted-by":"publisher"}],"name":"Universit\u00e9 de Toulouse","place":["Toulouse, France"]}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"341","published-online":{"date-parts":[[2026,10,6]]},"reference":[{"key":"e_1_3_4_1_2","doi-asserted-by":"publisher","DOI":"10.1038\/nrmicro3199"},{"key":"e_1_3_4_2_2","doi-asserted-by":"publisher","DOI":"10.1146\/annurev-genet-112618-043641"},{"key":"e_1_3_4_3_2","doi-asserted-by":"publisher","DOI":"10.1038\/s41564-018-0174-y"},{"key":"e_1_3_4_4_2","doi-asserted-by":"publisher","DOI":"10.1371\/journal.ppat.1003473"},{"key":"e_1_3_4_5_2","doi-asserted-by":"publisher","DOI":"10.1093\/femsre\/fuu001"},{"key":"e_1_3_4_6_2","doi-asserted-by":"publisher","DOI":"10.1111\/mmi.14718"},{"key":"e_1_3_4_7_2","doi-asserted-by":"publisher","DOI":"10.1128\/jb.00156-23"},{"key":"e_1_3_4_8_2","doi-asserted-by":"publisher","DOI":"10.1371\/journal.ppat.1004043"},{"key":"e_1_3_4_9_2","doi-asserted-by":"publisher","DOI":"10.1046\/j.1365-2958.1999.01170.x"},{"key":"e_1_3_4_10_2","doi-asserted-by":"publisher","DOI":"10.1371\/journal.pgen.1004066"},{"key":"e_1_3_4_11_2","doi-asserted-by":"publisher","DOI":"10.1111\/j.1365-2958.2004.04430.x"},{"key":"e_1_3_4_12_2","doi-asserted-by":"publisher","DOI":"10.1073\/pnas.2511180122"},{"key":"e_1_3_4_13_2","doi-asserted-by":"publisher","DOI":"10.1093\/nar\/gkaf932"},{"key":"e_1_3_4_14_2","doi-asserted-by":"publisher","DOI":"10.1038\/s41467-022-29494-z"},{"key":"e_1_3_4_15_2","doi-asserted-by":"publisher","DOI":"10.1111\/mmi.13732"},{"key":"e_1_3_4_16_2","doi-asserted-by":"publisher","DOI":"10.1073\/pnas.1315647110"},{"key":"e_1_3_4_17_2","doi-asserted-by":"publisher","DOI":"10.1128\/JB.00671-12"},{"key":"e_1_3_4_18_2","doi-asserted-by":"publisher","DOI":"10.1046\/j.1365-2958.2003.03702.x"},{"key":"e_1_3_4_19_2","doi-asserted-by":"publisher","DOI":"10.1016\/j.cell.2007.07.038"},{"key":"e_1_3_4_20_2","doi-asserted-by":"publisher","DOI":"10.1098\/rstb.2015.0528"},{"key":"e_1_3_4_21_2","doi-asserted-by":"publisher","DOI":"10.1371\/journal.pbio.1002394"},{"key":"e_1_3_4_22_2","doi-asserted-by":"publisher","DOI":"10.1038\/35084593"},{"key":"e_1_3_4_23_2","doi-asserted-by":"publisher","DOI":"10.1038\/349029a0"},{"key":"e_1_3_4_24_2","doi-asserted-by":"publisher","DOI":"10.1371\/journal.pbio.3002814"},{"key":"e_1_3_4_25_2","doi-asserted-by":"publisher","DOI":"10.1093\/molbev\/msx309"},{"key":"e_1_3_4_26_2","doi-asserted-by":"publisher","DOI":"10.1073\/pnas.2409843121"},{"key":"e_1_3_4_27_2","doi-asserted-by":"publisher","DOI":"10.1038\/ncomms15638"},{"key":"e_1_3_4_28_2","doi-asserted-by":"publisher","DOI":"10.1016\/j.cell.2019.11.021"},{"key":"e_1_3_4_29_2","doi-asserted-by":"publisher","DOI":"10.1073\/pnas.2421764122"},{"key":"e_1_3_4_30_2","doi-asserted-by":"publisher","DOI":"10.1093\/nar\/gky343"},{"key":"e_1_3_4_31_2","doi-asserted-by":"publisher","DOI":"10.1038\/s44318-024-00264-5"},{"key":"e_1_3_4_32_2","doi-asserted-by":"publisher","DOI":"10.1111\/j.1365-2958.2006.05558.x"},{"key":"e_1_3_4_33_2","doi-asserted-by":"publisher","DOI":"10.1128\/JB.00103-06"},{"key":"e_1_3_4_34_2","doi-asserted-by":"publisher","DOI":"10.1186\/1471-2105-6-172"},{"key":"e_1_3_4_35_2","doi-asserted-by":"publisher","DOI":"10.1128\/JB.180.10.2701-2710.1998"},{"key":"e_1_3_4_36_2","doi-asserted-by":"publisher","DOI":"10.1186\/gb-2001-2-12-research0054"},{"key":"e_1_3_4_37_2","doi-asserted-by":"publisher","DOI":"10.1128\/JB.00548-20"},{"key":"e_1_3_4_38_2","doi-asserted-by":"publisher","DOI":"10.1038\/s41467-020-15384-9"},{"key":"e_1_3_4_39_2","doi-asserted-by":"publisher","DOI":"10.1371\/journal.pone.0218941"},{"key":"e_1_3_4_40_2","doi-asserted-by":"publisher","DOI":"10.1093\/nar\/gkq958"},{"key":"e_1_3_4_41_2","doi-asserted-by":"publisher","DOI":"10.1093\/nar\/gks382"},{"key":"e_1_3_4_42_2","doi-asserted-by":"publisher","DOI":"10.1038\/nrm.2016.177"},{"key":"e_1_3_4_43_2","doi-asserted-by":"publisher","DOI":"10.1101\/cshperspect.a023192"},{"key":"e_1_3_4_44_2","doi-asserted-by":"publisher","DOI":"10.1073\/pnas.2213867120"},{"key":"e_1_3_4_45_2","doi-asserted-by":"publisher","DOI":"10.1126\/science.aaw6718"},{"key":"e_1_3_4_46_2","doi-asserted-by":"publisher","DOI":"10.1371\/journal.pcbi.1006891"},{"key":"e_1_3_4_47_2","doi-asserted-by":"publisher","DOI":"10.1073\/pnas.95.10.5752"},{"key":"e_1_3_4_48_2","doi-asserted-by":"publisher","DOI":"10.1016\/0022-2836(88)90055-1"},{"key":"e_1_3_4_49_2","doi-asserted-by":"publisher","DOI":"10.1017\/S0022172400031879"},{"key":"e_1_3_4_50_2","doi-asserted-by":"publisher","DOI":"10.1126\/science.adq0534"},{"key":"e_1_3_4_51_2","article-title":"A genomic view of experimental intraspecies and interspecies transformation of a rifampicin-resistance allele into Neisseria meningitidis","volume":"4","author":"Alfsnes K.","year":"2018","unstructured":"K. Alfsnes , A genomic view of experimental intraspecies and interspecies transformation of a rifampicin-resistance allele into Neisseria meningitidis. Microb. Genom. 4, e000222 (2018).","journal-title":"Microb. Genom."},{"key":"e_1_3_4_52_2","doi-asserted-by":"publisher","DOI":"10.1038\/ncomms11995"},{"key":"e_1_3_4_53_2","doi-asserted-by":"publisher","DOI":"10.1371\/journal.ppat.1002745"},{"key":"e_1_3_4_54_2","doi-asserted-by":"publisher","DOI":"10.1111\/1462-2920.15342"},{"key":"e_1_3_4_55_2","doi-asserted-by":"publisher","DOI":"10.1093\/nar\/gkv610"},{"key":"e_1_3_4_56_2","doi-asserted-by":"publisher","DOI":"10.1038\/s41587-020-0501-8"},{"key":"e_1_3_4_57_2","unstructured":"L. Hardy . Data for \u201cCharacterize recombination events.\u201d European Nucleotide Archive. https:\/\/www.ebi.ac.uk\/ena\/browser\/view\/PRJEB124975. Deposited 1 September 2026."}],"container-title":["Proceedings of the National Academy of Sciences"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/pnas.org\/doi\/pdf\/10.1073\/pnas.2610795123","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,10,6]],"date-time":"2026-10-06T18:05:19Z","timestamp":1791309919000},"score":1,"resource":{"primary":{"URL":"https:\/\/pnas.org\/doi\/10.1073\/pnas.2610795123"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,10,6]]},"references-count":57,"journal-issue":{"issue":"41","published-print":{"date-parts":[[2026,10,13]]}},"alternative-id":["10.1073\/pnas.2610795123"],"URL":"https:\/\/doi.org\/10.1073\/pnas.2610795123","relation":{},"ISSN":["0027-8424","1091-6490"],"issn-type":[{"value":"0027-8424","type":"print"},{"value":"1091-6490","type":"electronic"}],"subject":[],"published":{"date-parts":[[2026,10,6]]},"assertion":[{"value":"2026-03-27","order":0,"name":"received","label":"Received","group":{"name":"publication_history","label":"Publication History"}},{"value":"2026-07-28","order":2,"name":"accepted","label":"Accepted","group":{"name":"publication_history","label":"Publication History"}},{"value":"2026-10-06","order":3,"name":"published","label":"Published","group":{"name":"publication_history","label":"Publication History"}}],"article-number":"e2610795123"}}