{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,10,23]],"date-time":"2026-10-23T19:22:08Z","timestamp":1792783328202,"version":"4.5.0"},"reference-count":40,"publisher":"Springer Science and Business Media LLC","license":[{"start":{"date-parts":[[2026,10,8]],"date-time":"2026-10-08T00:00:00Z","timestamp":1791417600000},"content-version":"tdm","delay-in-days":0,"URL":"https:\/\/www.springernature.com\/gp\/researchers\/text-and-data-mining"},{"start":{"date-parts":[[2026,10,8]],"date-time":"2026-10-08T00:00:00Z","timestamp":1791417600000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/www.springernature.com\/gp\/researchers\/text-and-data-mining"}],"funder":[{"name":"National Natural Science Foundation of China","award":["32402059"],"award-info":[{"award-number":["32402059"]}],"id":[{"id":"https:\/\/ror.org\/01h0zpd94","id-type":"ROR","asserted-by":"crossref"}]},{"name":"Hainan University Scientific Research Start-up Fund Project","award":["XJ2300006565"],"award-info":[{"award-number":["XJ2300006565"]}],"id":[{"id":"https:\/\/ror.org\/03q648j11","id-type":"ROR","asserted-by":"crossref"}]}],"content-domain":{"domain":["link.springer.com"],"crossmark-restriction":false},"short-container-title":["Probiotics & Antimicro. Prot."],"DOI":"10.1007\/s12602-026-11239-9","type":"journal-article","created":{"date-parts":[[2026,10,8]],"date-time":"2026-10-08T08:04:19Z","timestamp":1791446659000},"update-policy":"https:\/\/doi.org\/10.1007\/springer_crossmark_policy","source":"Crossref","is-referenced-by-count":0,"title":["Selective Protein and Lipid Cargo Sorting into Extracellular Vesicles of Lacticaseibacillus paracasei subsp. paracasei"],"prefix":"10.1007","author":[{"given":"Huixue","family":"Hu","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Ming","family":"Yang","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0009-0002-6014-4455","authenticated-orcid":false,"given":"Jiamu","family":"Kang","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"297","published-online":{"date-parts":[[2026,10,8]]},"reference":[{"key":"11239_CR1","doi-asserted-by":"publisher","first-page":"2939","DOI":"10.1007\/s00253-016-7343-9","volume":"100","author":"P Alvarez-Sieiro","year":"2016","unstructured":"Alvarez-Sieiro P, Montalb\u00e1n-L\u00f3pez M, Mu D, Kuipers OP (2016) Bacteriocins of lactic acid bacteria: extending the family. Appl Microbiol Biotechnol 100:2939\u20132951","journal-title":"Appl Microbiol Biotechnol"},{"key":"11239_CR2","doi-asserted-by":"publisher","first-page":"173","DOI":"10.1016\/j.chom.2023.01.009","volume":"31","author":"EM Brown","year":"2023","unstructured":"Brown EM, Clardy J, Xavier RJ (2023) Gut microbiome lipid metabolism and its impact on host physiology. Cell Host Microbe 31:173\u2013186","journal-title":"Cell Host Microbe"},{"key":"11239_CR3","doi-asserted-by":"publisher","first-page":"1928","DOI":"10.1111\/jipb.13894","volume":"67","author":"J Deng","year":"2025","unstructured":"Deng J, Li W, Wang Z, Zeng J, Cai Q (2025) KatB, a bacterial extracellular vesicles (EVs)-secreted catalase, detoxifies reactive oxygen species (ROS) and promotes pathogen proliferation in plants. J Integr Plant Biol 67:1928\u20131946","journal-title":"J Integr Plant Biol"},{"key":"11239_CR4","doi-asserted-by":"crossref","unstructured":"Dom\u00ednguez Rubio AP, Mart\u00ednez JH, Mart\u00ednez Casillas DC, Coluccio Leskow F, Piuri M, P\u00e9rez OE (2017) Lactobacillus casei BL23 produces microvesicles carrying proteins that have been associated with its probiotic effect. Front Microbiol 8:1783","DOI":"10.3389\/fmicb.2017.01783"},{"key":"11239_CR5","doi-asserted-by":"publisher","first-page":"425","DOI":"10.1111\/tra.12488","volume":"18","author":"A Guerrero-Mandujano","year":"2017","unstructured":"Guerrero-Mandujano A, Hern\u00e1ndez-Cortez C, Ibarra JA, Castro-Escarpulli G (2017) The outer membrane vesicles: Secretion system type zero. Traffic 18:425\u2013432","journal-title":"Traffic"},{"key":"11239_CR6","doi-asserted-by":"publisher","first-page":"723433","DOI":"10.3389\/fimmu.2021.723433","volume":"12","author":"NA Harrison","year":"2021","unstructured":"Harrison NA, Gardner CL, da Silva DR, Gonzalez CF, Lorca GL (2021) Identification of Biomarkers for Systemic Distribution of Nanovesicles From Lactobacillus johnsonii N6.2. Front Immunol 12:723433","journal-title":"Front Immunol"},{"key":"11239_CR7","doi-asserted-by":"publisher","first-page":"305","DOI":"10.1186\/s12866-020-01988-0","volume":"20","author":"JL Herndon","year":"2020","unstructured":"Herndon JL, Peters RE, Hofer RN, Simmons TB, Symes SJ, Giles DK (2020) Exogenous polyunsaturated fatty acids (PUFAs) promote changes in growth, phospholipid composition, membrane permeability and virulence phenotypes in Escherichia coli. BMC Microbiol 20:305","journal-title":"BMC Microbiol"},{"key":"11239_CR8","doi-asserted-by":"publisher","first-page":"2708","DOI":"10.1038\/s41467-025-57631-x","volume":"16","author":"M-G Hong","year":"2025","unstructured":"Hong M-G, Song E-J, Yoon HJ, Chung W-H, Seo HY, Kim D, Lee D, Seo J-G, Lee H, Kim SI, Kim GJ, Kim K-N, Lee S-N, Kim KS, Nam Y-D (2025) Clade-specific extracellular vesicles from Akkermansia muciniphila mediate competitive colonization via direct inhibition and immune stimulation. Nat Commun 16:2708","journal-title":"Nat Commun"},{"key":"11239_CR9","doi-asserted-by":"crossref","unstructured":"Hu N, Lei M, Zhao X, Wang Y, Zhang Y, Wang S (2020) Analysis of microbiota in Hainan Yucha during fermentation by 16S rRNA gene high-throughput sequencing. J Food Process Preserv 44:e14523","DOI":"10.1111\/jfpp.14523"},{"key":"11239_CR10","doi-asserted-by":"publisher","first-page":"2741","DOI":"10.1080\/10408398.2020.1858269","volume":"62","author":"Y Hu","year":"2022","unstructured":"Hu Y, Zhang L, Wen R, Chen Q, Kong B (2022) Role of lactic acid bacteria in flavor development in traditional Chinese fermented foods: A review. Crit Rev Food Sci Nutr 62:2741\u20132755","journal-title":"Crit Rev Food Sci Nutr"},{"key":"11239_CR11","doi-asserted-by":"publisher","first-page":"1700","DOI":"10.1080\/07388551.2025.2496300","volume":"45","author":"C Huang","year":"2025","unstructured":"Huang C, Cao W, Zhou S, Deng Y (2025) Biogenesis mechanisms, regulatory strategies, and applications of bacterial extracellular vesicles. Crit Rev Biotechnol 45:1700\u20131716","journal-title":"Crit Rev Biotechnol"},{"key":"11239_CR12","doi-asserted-by":"publisher","first-page":"e70077","DOI":"10.1002\/jev2.70077","volume":"14","author":"X Huang","year":"2025","unstructured":"Huang X, Hu Z, Shang W, Chen J, Hu Q, Zhou Y, Ding R, Yin J, Li M, Liu H, Dou J, Peng H, Rao Y, Liu L, Wang Y, Tan L, Yang Y, Wu J, Xiao C, Yang Y, Rao X (2025) Beta-Lactam Antibiotics Promote Extracellular Vesicle Production of Staphylococcus aureus Through ROS-Mediated Lipid Metabolic Reprogramming. J Extracell Vesicles 14:e70077","journal-title":"J Extracell Vesicles"},{"key":"11239_CR13","doi-asserted-by":"publisher","first-page":"e61","DOI":"10.1002\/jex2.61","volume":"1","author":"R Juodeikis","year":"2022","unstructured":"Juodeikis R, Jones E, Deery E, Beal DM, Stentz R, Kr\u00e4utler B, Carding SR, Warren MJ (2022) Nutrient smuggling: Commensal gut bacteria-derived extracellular vesicles scavenge vitamin B12 and related cobamides for microbe and host acquisition. J Extracell Biology 1:e61","journal-title":"J Extracell Biology"},{"key":"11239_CR14","doi-asserted-by":"publisher","first-page":"e2416652122","DOI":"10.1073\/pnas.2416652122","volume":"122","author":"Y Kawagishi","year":"2025","unstructured":"Kawagishi Y, Murase K, Grebenshchikova A, Iibushi J, Ma C, Kimeu TM, Minowa-Nozawa A, Nozawa T, Nakagawa I (2025) Bacterial extracellular vesicles target different bacterial species, impairing cell division and diminishing their pathogenicity. \u200cProceedings Natl Acad Sci United States Am 122:e2416652122","journal-title":"\u200cProceedings Natl Acad Sci United States Am"},{"key":"11239_CR15","doi-asserted-by":"publisher","first-page":"136685","DOI":"10.1016\/j.foodchem.2023.136685","volume":"427","author":"B-H Lee","year":"2023","unstructured":"Lee B-H, Chen Y-Z, Shen T-L, Pan T-M, Hsu W-H (2023) Proteomic characterization of extracellular vesicles derived from lactic acid bacteria. Food Chem 427:136685","journal-title":"Food Chem"},{"key":"11239_CR16","doi-asserted-by":"crossref","unstructured":"Lee B-H, Hu Y-F, Chu Y-T, Wu Y-S, Hsu W-H, Nan F-H (2024) Lactic acid bacteria-fermented diet containing bacterial extracellular vesicles inhibited pathogenic bacteria in striped beakfish (Oplegnathus fasciatus). Fermentation 10(1):49","DOI":"10.3390\/fermentation10010049"},{"key":"11239_CR17","doi-asserted-by":"publisher","first-page":"128104","DOI":"10.1016\/j.foodchem.2020.128104","volume":"340","author":"BH Lee","year":"2021","unstructured":"Lee BH, Wu SC, Shen TL, Hsu YY, Chen CH, Hsu WH (2021) The applications of Lactobacillus plantarum-derived extracellular vesicles as a novel natural antibacterial agent for improving quality and safety in tuna fish. Food Chem 340:128104","journal-title":"Food Chem"},{"key":"11239_CR18","doi-asserted-by":"publisher","first-page":"118955","DOI":"10.1016\/j.biopha.2025.118955","volume":"194","author":"S Lee","year":"2026","unstructured":"Lee S, Jo E, Lee E-Y, Choi D (2026) Proteomics of extracellular vesicles derived from Lactobacillus: Implications in gut immune homeostasis and therapeutic potential. Biomed Pharmacother 194:118955","journal-title":"Biomed Pharmacother"},{"key":"11239_CR19","doi-asserted-by":"crossref","unstructured":"Lemos Junior WJF, Sant\u2019Ana AS, Di Cagno R, Campanaro S, Gobbetti M (2025) Five years after the lactobacillus reclassification: genomic and functional insights into Levilactobacillus, Lacticaseibacillus, Limosilactobacillus, Lactiplantibacillus, and Ligilactobacillus in fermented foods. Trends Food Sci. Technol 163","DOI":"10.1016\/j.tifs.2025.105118"},{"key":"11239_CR20","doi-asserted-by":"publisher","first-page":"e70131","DOI":"10.1002\/jev2.70131","volume":"14","author":"J Li","year":"2025","unstructured":"Li J, Li C, Han Y, Hu Y, Yang J, Xu H, Chen X, Yang M, Zuo J, Tang Y, Lei C, Li C, Wang H (2025) Unveiling a Novel Mechanism of Enhanced Secretion, Cargo Loading, and Accelerated Dynamics of Bacterial Extracellular Vesicles Following Antibiotic Exposure. J Extracell Vesicles 14:e70131","journal-title":"J Extracell Vesicles"},{"key":"11239_CR21","doi-asserted-by":"publisher","first-page":"2427311","DOI":"10.1080\/19490976.2024.2427311","volume":"16","author":"M Li","year":"2024","unstructured":"Li M, Mao B, Tang X, Zhang Q, Zhao J, Chen W, Cui S (2024) Lactic acid bacteria derived extracellular vesicles: emerging bioactive nanoparticles in modulating host health. Gut Microbes 16:2427311","journal-title":"Gut Microbes"},{"key":"11239_CR22","doi-asserted-by":"publisher","first-page":"100925","DOI":"10.1016\/j.crfs.2024.100925","volume":"9","author":"P Li","year":"2024","unstructured":"Li P, Bai Y, Li S, Zhang Z (2024) Characterisation of kefir-derived lactic acid bacteria and their extracellular vesicles. Curr Res Food Sci 9:100925","journal-title":"Curr Res Food Sci"},{"key":"11239_CR23","doi-asserted-by":"publisher","first-page":"eadn4526","DOI":"10.1126\/sciadv.adn4526","volume":"10","author":"Y Li","year":"2024","unstructured":"Li Y, Wang Y, Lin X, Sun S, Wu A, Ge Y, Yuan M, Wang J, Deng X, Tian Y (2024) Algicidal bacteria-derived membrane vesicles as shuttles mediating cross-kingdom interactions between bacteria and algae. Sci Adv 10:eadn4526","journal-title":"Sci Adv"},{"key":"11239_CR24","doi-asserted-by":"publisher","first-page":"e12212","DOI":"10.1002\/jev2.12212","volume":"11","author":"Y Liu","year":"2022","unstructured":"Liu Y, Liu Q, Zhao L, Dickey SW, Wang H, Xu R, Chen T, Jian Y, Wang X, Lv H, Otto M, Li M (2022) Essential role of membrane vesicles for biological activity of the bacteriocin micrococcin P1. J Extracell Vesicles 11:e12212","journal-title":"J Extracell Vesicles"},{"key":"11239_CR25","doi-asserted-by":"publisher","first-page":"104038","DOI":"10.1016\/j.fm.2022.104038","volume":"106","author":"A Mencher","year":"2022","unstructured":"Mencher A, Mejias-Ortiz M, Morales P, Tronchoni J, Gonzalez R (2022) Protein content of the Oenococcus oeni extracellular vesicles-enriched fraction. Food Microbiol 106:104038","journal-title":"Food Microbiol"},{"key":"11239_CR26","doi-asserted-by":"publisher","first-page":"8","DOI":"10.1111\/1751-7915.13657","volume":"14","author":"P Morales","year":"2021","unstructured":"Morales P, Mencher A, Tronchoni J, Gonzalez R (2021) Extracellular vesicles in food biotechnology. Microb Biotechnol 14:8\u201311","journal-title":"Microb Biotechnol"},{"key":"11239_CR27","doi-asserted-by":"crossref","unstructured":"Moravec AR, Siv AW, Hobby CR, Lindsay EN, Norbash LV, Shults DJ, Symes SJK, Giles DK (2017) Exogenous polyunsaturated fatty acids impact membrane remodeling and affect virulence phenotypes among pathogenic Vibrio species. applied and environmental microbiology 83","DOI":"10.1128\/AEM.01415-17"},{"key":"11239_CR28","doi-asserted-by":"publisher","first-page":"27263","DOI":"10.1038\/s41598-025-12873-z","volume":"15","author":"S Park","year":"2025","unstructured":"Park S, Mok J, Yu H-M, An H-J, Choi G-H, Lee Y-S, Kwon K-J, Choi S-J, Kim K-H, Kim S-J, Park J (2025) Comparative and pharmacological investigation of bEVs from eight Lactobacillales strains. Sci Rep 15:27263","journal-title":"Sci Rep"},{"key":"11239_CR29","doi-asserted-by":"publisher","first-page":"261","DOI":"10.1007\/s11274-025-04481-w","volume":"41","author":"K Peng","year":"2025","unstructured":"Peng K, Liu L, Gao S (2025) Harnessing microbial nanobiotics: Lactobacillus extracellular vesicles as next-generation therapeutics across physiological systems. World J Microbiol Biotechnol 41:261","journal-title":"World J Microbiol Biotechnol"},{"key":"11239_CR30","doi-asserted-by":"publisher","first-page":"1165202","DOI":"10.3389\/fmicb.2023.1165202","volume":"14","author":"G P\u00e9rez Mart\u00ednez","year":"2023","unstructured":"P\u00e9rez Mart\u00ednez G, Giner-P\u00e9rez L, Castillo-Romero KF (2023) Bacterial extracellular vesicles and associated functional proteins in fermented dairy products with Lacticaseibacillus paracasei. Front Microbiol 14:1165202","journal-title":"Front Microbiol"},{"key":"11239_CR31","doi-asserted-by":"publisher","first-page":"145640","DOI":"10.1016\/j.foodchem.2025.145640","volume":"493","author":"F Qiao","year":"2025","unstructured":"Qiao F, Ma X, Sun T, Li B, Wang S, Xuan S, Gong P, Hung W, Zhang L, Lin K (2025) Exogenous supplementation of key membrane lipid precursors enhances freeze-drying resistance in Lacticaseibacillus paracasei K56 by modulating membrane composition. Food Chem 493:145640","journal-title":"Food Chem"},{"key":"11239_CR32","doi-asserted-by":"crossref","unstructured":"Razim A, Laimer-Digruber A, Schmid AM, Edelbacher TV, Paschal ME, Weinmayer T, Thaler M, Skalska ME, Migda\u0142 P, Morandi M, Daniel C, Srutkova D, Schwarzer M, Hrdy J, Ehling-Schulz M, G\u00f3rska S, Inic-Kanada A, Wiedermann U, Schabussova I (2025) Extracellular vesicles as indicators of environmental stress response in Lactiplantibacillus plantarum. Extracellular vesicles and circulating nucleic acids, 2026.7(2). 786\u2013821","DOI":"10.20517\/evcna.2026.26"},{"key":"11239_CR33","doi-asserted-by":"publisher","first-page":"10515","DOI":"10.1038\/ncomms10515","volume":"7","author":"S Roier","year":"2016","unstructured":"Roier S, Zingl FG, Cakar F, Durakovic S, Kohl P, Eichmann TO, Klug L, Gadermaier B, Weinzerl K, Prassl R, Lass A, Daum G, Reidl J, Feldman MF, Schild S (2016) A novel mechanism for the biogenesis of outer membrane vesicles in Gram-negative bacteria. Nat Commun 7:10515","journal-title":"Nat Commun"},{"key":"11239_CR34","doi-asserted-by":"publisher","first-page":"637","DOI":"10.1016\/j.chom.2018.10.001","volume":"24","author":"Y Teng","year":"2018","unstructured":"Teng Y, Ren Y, Sayed M, Hu X, Lei C, Kumar A, Hutchins E, Mu J, Deng Z, Luo C, Sundaram K, Sriwastva MK, Zhang L, Hsieh M, Reiman R, Haribabu B, Yan J, Jala VR, Miller DM, Van Keuren-Jensen K, Merchant ML, McClain CJ, Park JW, Egilmez NK, Zhang HG (2018) Plant-Derived Exosomal MicroRNAs Shape the Gut Microbiota. Cell Host Microbe 24:637\u2013652e638","journal-title":"Cell Host Microbe"},{"key":"11239_CR35","doi-asserted-by":"publisher","first-page":"3586","DOI":"10.1099\/mic.0.2007\/009563-0","volume":"153","author":"M Turk","year":"2007","unstructured":"Turk M, Montiel V, \u017digon D, Plemenita\u0161 A, Ramos J (2007) Plasma membrane composition of Debaryomyces hansenii adapts to changes in pH and external salinity. Microbiology 153:3586\u20133592","journal-title":"Microbiology"},{"key":"11239_CR36","doi-asserted-by":"publisher","first-page":"10301","DOI":"10.1021\/acs.jafc.1c03896","volume":"69","author":"D Wang","year":"2021","unstructured":"Wang D, Chen H, Yang H, Yao S, Wu C (2021) Incorporation of Exogenous Fatty Acids Enhances the Salt Tolerance of Food Yeast Zygosaccharomyces rouxii. J Agric Food Chem 69:10301\u201310310","journal-title":"J Agric Food Chem"},{"key":"11239_CR37","doi-asserted-by":"publisher","first-page":"D384","DOI":"10.1093\/nar\/gkac1096","volume":"51","author":"J Wang","year":"2023","unstructured":"Wang J, Chitsaz F, Derbyshire MK, Gonzales NR, Gwadz M, Lu S, Marchler GH, Song JS, Thanki N, Yamashita RA, Yang M, Zhang D, Zheng C, Lanczycki CJ, Marchler-Bauer A (2023) The conserved domain database in 2023. Nucleic Acids Res 51:D384\u2013d388","journal-title":"Nucleic Acids Res"},{"key":"11239_CR38","doi-asserted-by":"publisher","first-page":"144776","DOI":"10.1016\/j.foodchem.2025.144776","volume":"489","author":"Y Wang","year":"2025","unstructured":"Wang Y, Hu L, Gao B, Yang R, Huang Y, Li P, Shang N (2025) Lactococcus lactis-derived extracellular vesicles: A novel nanodelivery system enhance the properties of nisin Z. Food Chem 489:144776","journal-title":"Food Chem"},{"key":"11239_CR39","doi-asserted-by":"crossref","unstructured":"Xie J, Li Q, Nie S (2024) Bacterial extracellular vesicles: An emerging postbiotic. Trends Food Sci Technol 143:104275","DOI":"10.1016\/j.tifs.2023.104275"},{"key":"11239_CR40","doi-asserted-by":"publisher","first-page":"91","DOI":"10.1186\/s40168-023-01540-y","volume":"11","author":"D Zhang","year":"2023","unstructured":"Zhang D, Zhang J, Kalimuthu S, Liu J, Song Z-M, He B-b, Cai P, Zhong Z, Feng C, Neelakantan P, Li Y-X (2023) A systematically biosynthetic investigation of lactic acid bacteria reveals diverse antagonistic bacteriocins that potentially shape the human microbiome. Microbiome 11:91","journal-title":"Microbiome"}],"container-title":["Probiotics and Antimicrobial Proteins"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/link.springer.com\/content\/pdf\/10.1007\/s12602-026-11239-9.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/link.springer.com\/article\/10.1007\/s12602-026-11239-9","content-type":"text\/html","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/link.springer.com\/content\/pdf\/10.1007\/s12602-026-11239-9.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,10,8]],"date-time":"2026-10-08T08:04:21Z","timestamp":1791446661000},"score":1,"resource":{"primary":{"URL":"https:\/\/link.springer.com\/10.1007\/s12602-026-11239-9"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,10,8]]},"references-count":40,"alternative-id":["11239"],"URL":"https:\/\/doi.org\/10.1007\/s12602-026-11239-9","relation":{},"ISSN":["1867-1306","1867-1314"],"issn-type":[{"value":"1867-1306","type":"print"},{"value":"1867-1314","type":"electronic"}],"subject":[],"published":{"date-parts":[[2026,10,8]]},"assertion":[{"value":"3 June 2026","order":1,"name":"received","label":"Received","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"28 September 2026","order":2,"name":"accepted","label":"Accepted","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"8 October 2026","order":3,"name":"first_online","label":"First Online","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"The authors declare that they have no known competing financial interests.","order":1,"name":"Ethics","label":"Declaration of competing interest","group":{"name":"EthicsHeading","label":"Declarations"}}]}}