{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,9,17]],"date-time":"2026-09-17T06:03:35Z","timestamp":1789625015086,"version":"build-2803163510"},"reference-count":89,"publisher":"Frontiers Media SA","license":[{"start":{"date-parts":[[2026,9,17]],"date-time":"2026-09-17T00:00:00Z","timestamp":1789603200000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"content-domain":{"domain":["frontiersin.org"],"crossmark-restriction":true},"short-container-title":["Front. Microbiol."],"abstract":"<jats:p>\n                    Horizontal gene transfer (HGT) mediated by conjugative plasmids is a major driver of antimicrobial resistance (AMR) dissemination in aquatic environments. In\n                    <jats:italic>Aeromonas salmonicida<\/jats:italic>\n                    , plasmid-mediated HGT is largely driven by the pRAS plasmid family. Here we present genomic and experimental characterization of pRAS2 from\n                    <jats:italic>A. salmonicida<\/jats:italic>\n                    strain 1682\/92 and demonstrate its role in helper-dependent mobilization of the co-resident pRAS3. Whole genome sequencing (WGS) divided the pRAS2 backbone into three modules. Module A encodes replication, maintenance and inheritance stability functions, including transposase, resolvase, partitioning and antitoxin-associated genes. Module B encodes a canonical type IV secretion system (T4SS) conjugation module containing\n                    <jats:italic>virB2<\/jats:italic>\n                    -\n                    <jats:italic>virB6<\/jats:italic>\n                    ,\n                    <jats:italic>virB8<\/jats:italic>\n                    -\n                    <jats:italic>virB11<\/jats:italic>\n                    and\n                    <jats:italic>virD4<\/jats:italic>\n                    with\n                    <jats:italic>virB2<\/jats:italic>\n                    and\n                    <jats:italic>virB5<\/jats:italic>\n                    forming the transfer pilus. Module C encodes the resistance determinants\n                    <jats:italic>sul2<\/jats:italic>\n                    ,\n                    <jats:italic>aph(6)-Id<\/jats:italic>\n                    ,\n                    <jats:italic>neo<\/jats:italic>\n                    and Tet 31. Modules A and B were organized as in plasmids of\n                    <jats:italic>Vibrio alginolyticus<\/jats:italic>\n                    ,\n                    <jats:italic>Edwardsiella piscicida<\/jats:italic>\n                    and\n                    <jats:italic>Piscirickettsia salmonis<\/jats:italic>\n                    , indicating that pRAS2 shares a conserved backbone with plasmids of other aquatic bacteria, including fish pathogens isolated on different continents. Module C was the most variable region:\n                    <jats:italic>sul2<\/jats:italic>\n                    and\n                    <jats:italic>aph(6)-Id<\/jats:italic>\n                    were shared with pEIB202 of\n                    <jats:italic>E. piscicida<\/jats:italic>\n                    , whereas the tetracycline resistance determinant remained the same, being Tet 31 in pRAS2 as well as in pEIB202, p3PS10 and p3PS11, but absent from pVb1978 (\n                    <jats:italic>Vibrio alginolyticus<\/jats:italic>\n                    ), pVIM-20710 (\n                    <jats:italic>Enterobacter hormaechei<\/jats:italic>\n                    ) and p4PS9. The pRAS3 plasmids of strains 1682\/92 and 2131\/90, used as donors in conjugation experiments with an\n                    <jats:italic>Escherichia coli<\/jats:italic>\n                    DH5\u03b1 recipient, carried the same mobilization-associated (MOB) genes and the tetracycline resistance determinant Tet C. Transfer of pRAS3 occurred only from strain 1682\/92, which carried both pRAS2 and pRAS3; no transconjugant colonies were recovered from strain 2131\/90, which carried pRAS3 alone. WGS showed an identical organization of the MOB,\n                    <jats:italic>tetA<\/jats:italic>\n                    (C) and\n                    <jats:italic>tetR<\/jats:italic>\n                    (C) genes in the 1682\/92 donor and its transconjugant, providing experimental evidence that pRAS2 functions as a conjugative helper plasmid mediating mobilization of pRAS3. Conjugation trials performed with these strains between 1993 and 1995 further show that pRAS2 is self-transmissible, transferring autonomously at a frequency approximately three orders of magnitude higher than that of its co-mobilization of pRAS3. This demonstrates plasmid-plasmid cooperation within the pRAS plasmid family as a mechanism supporting the spread of AMR genes in aquatic bacterial communities.\n                  <\/jats:p>","DOI":"10.3389\/fmicb.2026.1872994","type":"journal-article","created":{"date-parts":[[2026,9,17]],"date-time":"2026-09-17T05:31:05Z","timestamp":1789623065000},"update-policy":"https:\/\/doi.org\/10.3389\/crossmark-policy","source":"Crossref","is-referenced-by-count":0,"title":["The R plasmid pRAS2 in the fish pathogen Aeromonas salmonicida is a global mediator of antibiotic resistance in aquaculture and can be used by pRAS3 for mobilization"],"prefix":"10.3389","volume":"17","author":[{"given":"Surya Prasad","family":"Dahal","sequence":"first","affiliation":[{"name":"Faculty of Biosciences and Aquaculture, Nord University","place":["Bod\u00f8, Norway"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Hetron Mweemba","family":"Munang\u2019andu","sequence":"additional","affiliation":[{"name":"Faculty of Biosciences and Aquaculture, Nord University","place":["Bod\u00f8, Norway"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Asta","family":"Solberg","sequence":"additional","affiliation":[{"name":"Department of Paraclinical Sciences, Faculty of Veterinary Medicine, Norwegian University of Life Sciences","place":["\u00c5s, Norway"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Henning","family":"S\u00f8rum","sequence":"additional","affiliation":[{"name":"Department of Paraclinical Sciences, Faculty of Veterinary Medicine, Norwegian University of Life Sciences","place":["\u00c5s, Norway"]}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"1965","published-online":{"date-parts":[[2026,9,17]]},"reference":[{"key":"B1","doi-asserted-by":"publisher","first-page":"D517","DOI":"10.1093\/nar\/gkz935","article-title":"CARD 2020: Antibiotic resistome surveillance with the comprehensive antibiotic resistance database.","volume":"48","author":"Alcock","year":"2020","journal-title":"Nucleic Acids Res."},{"key":"B2","volume-title":"FastQC: A Quality Control Tool for High Throughput Sequence Data. 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