{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,9,21]],"date-time":"2026-09-21T19:01:38Z","timestamp":1790017298009,"version":"4.0.1"},"update-to":[{"DOI":"10.1371\/journal.pgen.1012311","type":"new_version","label":"New version","source":"publisher","updated":{"date-parts":[[2026,9,21]],"date-time":"2026-09-21T00:00:00Z","timestamp":1789948800000}}],"reference-count":84,"publisher":"Public Library of Science (PLoS)","issue":"9","license":[{"start":{"date-parts":[[2026,9,15]],"date-time":"2026-09-15T00:00:00Z","timestamp":1789430400000},"content-version":"vor","delay-in-days":0,"URL":"http:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"DOI":"10.13039\/501100001809","name":"National Natural Science Foundation of China","doi-asserted-by":"publisher","award":["32370115 and 32570118"],"award-info":[{"award-number":["32370115 and 32570118"]}],"id":[{"id":"10.13039\/501100001809","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100007129","name":"Natural Science Foundation of Shandong Province","doi-asserted-by":"publisher","award":["ZR2026QA16"],"award-info":[{"award-number":["ZR2026QA16"]}],"id":[{"id":"10.13039\/501100007129","id-type":"DOI","asserted-by":"publisher"}]},{"name":"National Key Research and Development Project of China","award":["2023YFC3403500 and 2024YFD2401701"],"award-info":[{"award-number":["2023YFC3403500 and 2024YFD2401701"]}]},{"name":"Southern Marine Science and Engineering Guangdong Laboratory","award":["SML2024SP022 and SML2024SP002"],"award-info":[{"award-number":["SML2024SP022 and SML2024SP002"]}]}],"content-domain":{"domain":["www.plosgenetics.org"],"crossmark-restriction":false},"short-container-title":["PLoS Genet"],"abstract":"<jats:p>\n                    Lipoprotein and OmpW are two major components of the outer membrane (OM) in Gram-negative bacteria and play essential roles in various physiological processes, e.g., protein secretion, folding, and localization. They typically assist in maintaining client proteins in a folded state or form pore channels during secretion. However, how OM proteins facilitate the secretion of proteins that lack classical signal peptides remains elusive. Here, we demonstrate that they contribute to the secretion of unconventional B-type dye-decolorizing peroxidase (DypB\n                    <jats:sub>2985<\/jats:sub>\n                    ) in\n                    <jats:italic>Pseudomonas putida<\/jats:italic>\n                    . The lipoprotein, Lpp\n                    <jats:sub>1528<\/jats:sub>\n                    , which contains a Sec signal peptide, is translocated to the periplasm through the Sec pathway and anchored in the inner leaflet of the OM. Lpp\n                    <jats:sub>1528<\/jats:sub>\n                    recognizes the C-terminal hydrophobic region of DypB\n                    <jats:sub>2985<\/jats:sub>\n                    in the cytoplasm and facilitates its coupling to the Sec machinery for inner membrane translocation, despite DypB\n                    <jats:sub>2985<\/jats:sub>\n                    lacking a canonical N-terminal Sec signal peptide. Following translocation, the two proteins appear to dissociate in the periplasm. Subsequently, another OM protein, OmpW\n                    <jats:sub>4836<\/jats:sub>\n                    , recognizes the N-terminal hydrophobic region of periplasmic DypB\n                    <jats:sub>2985<\/jats:sub>\n                    and mediates its incorporation into outer membrane vesicles (OMVs) for extracellular delivery. Our study reveals the crosstalk between the Sec pathway and OMVs in the secretion of a non-canonical peroxidase, in which OM proteins, acting as the molecular tethers, mediate the stepwise secretion process. It expands our understanding of non-classical protein secretion mechanisms and bacterial survival strategies. Moreover, the identified OMV sorting mechanism offers potential for the further functionalization of OMVs as versatile biotechnological platforms.\n                  <\/jats:p>","DOI":"10.1371\/journal.pgen.1012311","type":"journal-article","created":{"date-parts":[[2026,9,15]],"date-time":"2026-09-15T18:18:23Z","timestamp":1789496303000},"page":"e1012311","update-policy":"https:\/\/doi.org\/10.1371\/journal.pgen.corrections_policy","source":"Crossref","is-referenced-by-count":0,"title":["Outer membrane proteins mediate unconventional secretion of Pseudomonas peroxidase through crosstalk between the Sec pathway and outer membrane vesicles"],"prefix":"10.1371","volume":"22","author":[{"given":"Congying","family":"Liang","sequence":"first","affiliation":[{"name":"Institute of Marine Science and Technology, Shandong University","place":["Qingdao, China"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Wenping","family":"Zhu","sequence":"additional","affiliation":[{"name":"Institute of Marine Science and Technology, Shandong University","place":["Qingdao, China"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-9663-6382","authenticated-orcid":true,"given":"Lu","family":"Lin","sequence":"additional","affiliation":[{"name":"Institute of Marine Science and Technology, Shandong University","place":["Qingdao, China"]},{"name":"Southern Marine Science and Engineering Guangdong Laboratory (Zhuhai)","place":["Zhuhai, China"]},{"name":"Shandong Key Laboratory of Intelligent Marine Engineering Geology, Environment and Equipment","place":["Qingdao, China"]}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"340","published-online":{"date-parts":[[2026,9,15]]},"reference":[{"issue":"7","key":"pgen.1012311.ref001","doi-asserted-by":"crossref","first-page":"734","DOI":"10.1038\/ncb0707-734","article-title":"Threonine phosphorylation times bacterial secretion","volume":"9","author":"HD Kulasekara","year":"2007","journal-title":"Nat Cell Biol"},{"key":"pgen.1012311.ref002","doi-asserted-by":"crossref","first-page":"107790","DOI":"10.1016\/j.micpath.2025.107790","article-title":"Bacterial protein secretion systems: Mechanisms, functions, and roles in virulence","volume":"206","author":"H Jamali","year":"2025","journal-title":"Microb Pathog"},{"issue":"9","key":"pgen.1012311.ref003","doi-asserted-by":"crossref","first-page":"1735","DOI":"10.1016\/j.bbamem.2007.07.015","article-title":"Sec- and Tat-mediated protein secretion across the bacterial cytoplasmic membrane--distinct translocases and 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