{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,10,28]],"date-time":"2026-10-28T07:43:19Z","timestamp":1793173399784,"version":"4.5.0"},"reference-count":174,"publisher":"MDPI AG","issue":"8","license":[{"start":{"date-parts":[[2026,8,3]],"date-time":"2026-08-03T00:00:00Z","timestamp":1785715200000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":["Biomolecules"],"abstract":"<jats:p>Bacterial extracellular vesicles (BEVs) are nano-sized lipid bilayer particles secreted by bacteria, capable of carrying various proteins, lipids, nucleic acids, and pathogen-associated molecular patterns (PAMPs). The biosynthetic pathway of BEVs determines their load components, physicochemical properties, and different biological activities. Increasing evidence indicates that BEVs play an important role in mediating host immune responses and the dynamic regulation of bacterial biofilms. Additionally, BEVs may serve as a molecular bridge between the two. BEVs derived from pathogens can trigger pro-inflammatory cascades, assist bacteria in immune evasion, and further accelerate the maturation of biofilms, forming a vicious cycle of persistent infection and inflammatory damage. In contrast, BEVs derived from probiotics can maintain host immune homeostasis and exert direct anti-biofilm and synergistic antibacterial effects, thereby breaking the pathological cycle. However, significant methodological research bottlenecks have greatly hindered the comparability and clinical translation of BEVs research. This article systematically summarizes the classification of BEVs and their biosynthetic mechanisms, compares the differential effects of BEVs from pathogenic bacteria and probiotic bacteria on immunity, clarifies the dual regulatory role of BEVs throughout the life cycle of biofilms, and highlights the bridging function of BEVs in the immune\u2013biofilm interaction. Additionally, this article also discusses the current development of BEVs in clinical translation applications, such as vaccine development, antibiotic delivery, and mucosal inflammation intervention, and outlines the key industrial and clinical challenges faced in the future development of BEVs-based therapeutic approaches.<\/jats:p>","DOI":"10.3390\/biom16081132","type":"journal-article","created":{"date-parts":[[2026,8,3]],"date-time":"2026-08-03T11:50:21Z","timestamp":1785757821000},"page":"1132","update-policy":"https:\/\/doi.org\/10.3390\/mdpi_crossmark_policy","source":"Crossref","is-referenced-by-count":1,"title":["Bacterial Extracellular Vesicles at the Crossroads of Immune Regulation and Biofilm Dynamics: Biogenesis, Comparative Analysis, and Translational Challenges"],"prefix":"10.3390","volume":"16","author":[{"given":"Qingyu","family":"Zhang","sequence":"first","affiliation":[{"name":"Faculty of Applied Science, Universiti Teknologi MARA Shah Alam, Shah Alam 40450, Selangor, Malaysia"},{"name":"Department of Food Research, Zhejiang Business College, Hangzhou 310059, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Beilei","family":"Zhang","sequence":"additional","affiliation":[{"name":"Sanmen Food and Drug Inspection and Testing Center, Taizhou 317100, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-4942-8457","authenticated-orcid":false,"given":"Mohd Shafiq","family":"Aazmi","sequence":"additional","affiliation":[{"name":"Faculty of Applied Science, Universiti Teknologi MARA Shah Alam, Shah Alam 40450, Selangor, Malaysia"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Lin","family":"Chen","sequence":"additional","affiliation":[{"name":"Department of Food Research, Zhejiang Business College, Hangzhou 310059, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-1208-6599","authenticated-orcid":false,"given":"Mohd Fakharul Zaman Raja","family":"Yahya","sequence":"additional","affiliation":[{"name":"Faculty of Applied Science, Universiti Teknologi MARA Shah Alam, Shah Alam 40450, Selangor, Malaysia"},{"name":"Integrative Pharmacogenomics Institute (iPROMISE), Universiti Teknologi MARA Selangor Branch, Puncak Alam Campus, Bandar Puncak Alam 42300, Selangor, Malaysia"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"1968","published-online":{"date-parts":[[2026,8,3]]},"reference":[{"key":"ref_1","doi-asserted-by":"crossref","first-page":"415","DOI":"10.1038\/s41579-023-00875-5","article-title":"Composition and functions of bacterial membrane vesicles","volume":"21","author":"Toyofuku","year":"2023","journal-title":"Nat. Rev. Microbiol."},{"key":"ref_2","doi-asserted-by":"crossref","first-page":"1948","DOI":"10.1128\/IAI.06014-11","article-title":"Membrane vesicle release in bacteria, eukaryotes, and archaea: A conserved yet underappreciated aspect of microbial life","volume":"80","author":"Deatherage","year":"2012","journal-title":"Infect. Immun."},{"key":"ref_3","doi-asserted-by":"crossref","first-page":"97","DOI":"10.1016\/j.semcdb.2015.02.006","article-title":"Gram-negative and Gram-positive bacterial extracellular vesicles","volume":"40","author":"Kim","year":"2015","journal-title":"Semin. Cell Dev. Biol."},{"key":"ref_4","doi-asserted-by":"crossref","first-page":"e12406","DOI":"10.1002\/jev2.12406","article-title":"Differential temporal release and lipoprotein loading in B. thetaiotaomicron bacterial extracellular vesicles","volume":"13","author":"Juodeikis","year":"2024","journal-title":"J. Extracell. Vesicles"},{"key":"ref_5","doi-asserted-by":"crossref","first-page":"e0168622","DOI":"10.1128\/aem.01686-22","article-title":"Proteomic Profiling Reveals Distinct Bacterial Extracellular Vesicle Subpopulations with Possibly Unique Functionality","volume":"89","author":"McMillan","year":"2023","journal-title":"Appl. Environ. Microbiol."},{"key":"ref_6","doi-asserted-by":"crossref","first-page":"e1800209","DOI":"10.1002\/pmic.201800209","article-title":"Helicobacter pylori Growth Stage Determines the Size, Protein Composition, and Preferential Cargo Packaging of Outer Membrane Vesicles","volume":"19","author":"Zavan","year":"2019","journal-title":"Proteomics"},{"key":"ref_7","doi-asserted-by":"crossref","unstructured":"Wei, S., Jiao, D., and Xing, W. (2022). A rapid method for isolation of bacterial extracellular vesicles from culture media using epsilon-poly-L-lysine that enables immunological function research. Front. Immunol., 13.","DOI":"10.3389\/fimmu.2022.930510"},{"key":"ref_8","doi-asserted-by":"crossref","unstructured":"Wang, Y., Hoffmann, J.P., Baker, S.M., Bentrup, K.H.Z., Wimley, W.C., Fuselier, J.A., Bitoun, J.P., and Morici, L.A. (2021). Inhibition of Streptococcus mutans biofilms with bacterial-derived outer membrane vesicles. BMC Microbiol., 21.","DOI":"10.1186\/s12866-021-02296-x"},{"key":"ref_9","doi-asserted-by":"crossref","first-page":"3786","DOI":"10.1039\/D4TB01899K","article-title":"Bacterial outer membrane vesicles in tumor prevention and treatment: Advancements in research and application","volume":"13","author":"Liu","year":"2025","journal-title":"J. Mater. Chem. B"},{"key":"ref_10","doi-asserted-by":"crossref","first-page":"e70069","DOI":"10.1002\/jev2.70069","article-title":"Nanobody Engineered and Photosensitiser Loaded Bacterial Outer Membrane Vesicles Potentiate Antitumour Immunity and Immunotherapy","volume":"14","author":"Xia","year":"2025","journal-title":"J. Extracell. Vesicles"},{"key":"ref_11","doi-asserted-by":"crossref","first-page":"375","DOI":"10.1016\/j.jare.2024.03.003","article-title":"Gut microbiome-derived bacterial extracellular vesicles in patients with solid tumours","volume":"68","author":"Mishra","year":"2025","journal-title":"J. Adv. Res."},{"key":"ref_12","doi-asserted-by":"crossref","unstructured":"Zhang, R., Li, G., Wu, Y., Wang, X., and Luan, Q. (2024). Pathogenic mechanisms and potential applications of extracellular vesicles from periodontal pathogens in periodontitis. Front. Immunol., 15.","DOI":"10.3389\/fimmu.2024.1513983"},{"key":"ref_13","doi-asserted-by":"crossref","first-page":"13","DOI":"10.1038\/s41579-018-0112-2","article-title":"Types and origins of bacterial membrane vesicles","volume":"17","author":"Toyofuku","year":"2019","journal-title":"Nat. Rev. Microbiol."},{"key":"ref_14","doi-asserted-by":"crossref","first-page":"25","DOI":"10.1186\/s40104-020-00532-4","article-title":"Lactobacillus reuteri-derived extracellular vesicles maintain intestinal immune homeostasis against lipopolysaccharide-induced inflammatory responses in broilers","volume":"12","author":"Hu","year":"2021","journal-title":"J. Anim. Sci. Biotechnol."},{"key":"ref_15","doi-asserted-by":"crossref","first-page":"6600","DOI":"10.7150\/thno.132287","article-title":"Bacterial extracellular vesicles as multifaceted regulators of human diseases and emerging therapeutic platforms","volume":"16","author":"Wang","year":"2026","journal-title":"Theranostics"},{"key":"ref_16","doi-asserted-by":"crossref","unstructured":"Bril\u2019kov, M.S., Stenbakk, V., Jakubec, M., Vasskog, T., Kristoffersen, T., Cavanagh, J.P., Ericson, J.U., Isaksson, J., and Flaten, G.E. (2023). Bacterial extracellular vesicles: Towards realistic models for bacterial membranes in molecular interaction studies by surface plasmon resonance. Front Mol. Biosci., 10.","DOI":"10.3389\/fmolb.2023.1277963"},{"key":"ref_17","doi-asserted-by":"crossref","first-page":"e20230017","DOI":"10.1002\/INMD.20230017","article-title":"Bacterial extracellular vesicles: A position paper by the microbial vesicles task force of the Chinese society for extracellular vesicles","volume":"1","author":"Wen","year":"2023","journal-title":"Interdiscip. Med."},{"key":"ref_18","doi-asserted-by":"crossref","first-page":"312","DOI":"10.15698\/mic2020.12.737","article-title":"Extracellular vesicles: An emerging platform in gram-positive bacteria","volume":"7","author":"Bose","year":"2020","journal-title":"Microb. Cell"},{"key":"ref_19","doi-asserted-by":"crossref","first-page":"e70301","DOI":"10.1002\/jev2.70301","article-title":"Host Factor Induced Bacterial Extracellular Vesicles Promote Horizontal Gene Transfer in Vibrio cholerae","volume":"15","author":"Fleischhacker","year":"2026","journal-title":"J. Extracell. Vesicles"},{"key":"ref_20","doi-asserted-by":"crossref","first-page":"547","DOI":"10.3724\/SP.J.1123.2024.10017","article-title":"Lipidomics analysis of glycine-induced bacterial outer membrane vesicles","volume":"43","author":"Song","year":"2025","journal-title":"Chin. J. Chromatogr."},{"key":"ref_21","doi-asserted-by":"crossref","unstructured":"Aktar, S., Okamoto, Y., Ueno, S., Tahara, Y.O., Imaizumi, M., Shintani, M., Miyata, M., Futamata, H., Nojiri, H., and Tashiro, Y. (2021). Incorporation of Plasmid DNA Into Bacterial Membrane Vesicles by Peptidoglycan Defects in Escherichia coli. Front. Microbiol., 12.","DOI":"10.3389\/fmicb.2021.747606"},{"key":"ref_22","first-page":"55146","article-title":"Legionella pneumophila Outer Membrane Vesicles: Isolation and Analysis of Their Pro-inflammatory Potential on Macrophages","volume":"120","author":"Jung","year":"2017","journal-title":"J. Vis. Exp."},{"key":"ref_23","first-page":"169","article-title":"Bacterial extracellular vesicles as bioactive nanocarriers for drug delivery: Advances and perspectives","volume":"14","author":"Liu","year":"2021","journal-title":"Bioact. Mater."},{"key":"ref_24","doi-asserted-by":"crossref","unstructured":"Chudzik, A., Migda\u0142, P., and Pa\u015bciak, M. (2022). Different Cutibacterium acnes Phylotypes Release Distinct Extracellular Vesicles. Int. J. Mol. Sci., 23.","DOI":"10.3390\/ijms23105797"},{"key":"ref_25","doi-asserted-by":"crossref","unstructured":"Villageliu, D.N., and Samuelson, D.R. (2022). The Role of Bacterial Membrane Vesicles in Human Health and Disease. Front. Microbiol., 13.","DOI":"10.3389\/fmicb.2022.828704"},{"key":"ref_26","doi-asserted-by":"crossref","unstructured":"Verbunt, J., Mennens, L., Jocken, J., Blaak, E.E., Savelkoul, P., and Stassen, F.R.M. (2026). From food to vesicle: Nutritional influences on gut microbial inflammatory signaling. Front. Nutr., 13.","DOI":"10.3389\/fnut.2026.1756462"},{"key":"ref_27","doi-asserted-by":"crossref","first-page":"e0018722","DOI":"10.1128\/msphere.00187-22","article-title":"Contribution of Membrane Vesicle to Reprogramming of Bacterial Membrane Fluidity in Pseudomonas aeruginosa","volume":"7","author":"Mozaheb","year":"2022","journal-title":"mSphere"},{"key":"ref_28","doi-asserted-by":"crossref","first-page":"e70350","DOI":"10.1111\/1751-7915.70350","article-title":"Bacterial Extracellular Vesicles in Biomedical Research and Clinical Translation","volume":"19","year":"2026","journal-title":"Microb. Biotechnol."},{"key":"ref_29","unstructured":"(2024). Microbial Vesicles Task Force of the Chinese Society for Extracellular Vesicles Affiliated with the Chinese Research Hospital Association; Society of Extracellular Vesicles Affiliated with Guangdong Precision Medicine Application Association. Chinese expert consensus for nomenclature, isolation and detection of bacterial extracellular vesicles. Natl. Med. J. China, 104, 2910\u20132927."},{"key":"ref_30","doi-asserted-by":"crossref","unstructured":"Barbieri, G., Maurizi, L., Zini, M., Fratini, F., Pietrantoni, A., Bellini, I., Cavallero, S., D\u2019Intino, E., Rinaldi, F., and Chiani, P. (2026). Biophysical Features of Outer Membrane Vesicles (OMVs) from Pathogenic Escherichia coli: Methodological Implications for Reproducible OMV Characterization. Antibiotics, 15.","DOI":"10.3390\/antibiotics15020117"},{"key":"ref_31","doi-asserted-by":"crossref","first-page":"36405","DOI":"10.31083\/FBL36405","article-title":"Extracellular Vesicles: Recent Advances and Perspectives","volume":"30","author":"Ragni","year":"2025","journal-title":"Front Biosci. (Landmark Ed.)"},{"key":"ref_32","doi-asserted-by":"crossref","unstructured":"Bhanu, P., Godwin, A.K., Umar, S., and Mahoney, D.E. (2025). Bacterial Extracellular Vesicles in Oncology: Molecular Mechanisms and Future Clinical Applications. Cancers, 17.","DOI":"10.3390\/cancers17111774"},{"key":"ref_33","doi-asserted-by":"crossref","first-page":"123","DOI":"10.1007\/978-1-0716-1302-3_13","article-title":"Isolation of Outer Membrane Vesicles from Helicobacter pylori","volume":"2283","author":"Windle","year":"2021","journal-title":"Methods Mol. Biol."},{"key":"ref_34","doi-asserted-by":"crossref","first-page":"1723","DOI":"10.1093\/ibd\/izaf107","article-title":"Therapeutic Potential and Translational Challenges for Bacterial Extracellular Vesicles in Inflammatory Bowel Disease","volume":"31","author":"Pirolli","year":"2025","journal-title":"Inflamm. Bowel Dis."},{"key":"ref_35","doi-asserted-by":"crossref","first-page":"114822","DOI":"10.1016\/j.jconrel.2026.114822","article-title":"Optimizing genetic engineering approaches for protein loading into bacterial extracellular vesicles for vaginal drug delivery","volume":"393","author":"Steinman","year":"2026","journal-title":"J. Control. Release"},{"key":"ref_36","first-page":"265","article-title":"Engineering strategies and biomedical applications of bacterial extracellular vesicles","volume":"6","author":"Liang","year":"2025","journal-title":"Biomater. Transl."},{"key":"ref_37","doi-asserted-by":"crossref","first-page":"e2506011","DOI":"10.4014\/jmb.2506.06011","article-title":"Standardizing Bacterial Extracellular Vesicle Purification: A Call for Consensus","volume":"35","author":"Choi","year":"2025","journal-title":"J. Microbiol. Biotechnol."},{"key":"ref_38","doi-asserted-by":"crossref","first-page":"1324731","DOI":"10.1080\/20013078.2017.1324731","article-title":"Isolation of membrane vesicles from prokaryotes: A technical and biological comparison reveals heterogeneity","volume":"6","author":"Chang","year":"2017","journal-title":"J. Extracell. Vesicles"},{"key":"ref_39","doi-asserted-by":"crossref","first-page":"e0127321","DOI":"10.1128\/Spectrum.01273-21","article-title":"Considerations for the Analysis of Bacterial Membrane Vesicles: Methods of Vesicle Production and Quantification Can Influence Biological and Experimental Outcomes","volume":"9","author":"Bitto","year":"2021","journal-title":"Microbiol. Spectr."},{"key":"ref_40","doi-asserted-by":"crossref","unstructured":"Ste\u0107, A., Jo\u0144ca, J., Waleron, K., Waleron, M., P\u0142oska, A., Kalinowski, L., Wielgomas, B., and Dziomba, S. (2022). Quality Control of Bacterial Extracellular Vesicles with Total Protein Content Assay, Nanoparticles Tracking Analysis, and Capillary Electrophoresis. Int. J. Mol. Sci., 23.","DOI":"10.3390\/ijms23084347"},{"key":"ref_41","doi-asserted-by":"crossref","first-page":"219","DOI":"10.1007\/978-1-0716-4055-5_14","article-title":"Dose Determination and Administration of Bacterial Extracellular Vesicles for In Vivo Preclinical Studies","volume":"2843","author":"Lathia","year":"2024","journal-title":"Methods Mol. Biol."},{"key":"ref_42","doi-asserted-by":"crossref","unstructured":"Petersen, A.P., and Zierden, H.C. (2024). Molecular Approaches to Measure Bacterial Extracellular Vesicles in Tissue Samples. Bacterial Extracellular Vesicles: Methods and Protocols, Springer.","DOI":"10.1007\/978-1-0716-4055-5_16"},{"key":"ref_43","doi-asserted-by":"crossref","first-page":"18752","DOI":"10.1038\/s41598-023-45628-9","article-title":"Molecular imaging of bacterial outer membrane vesicles based on bacterial surface display","volume":"13","author":"Hajdrik","year":"2023","journal-title":"Sci. Rep."},{"key":"ref_44","doi-asserted-by":"crossref","first-page":"239","DOI":"10.1007\/978-1-0716-4055-5_15","article-title":"Assessing In Vivo Bacterial Extracellular Vesicle (BEV) Biodistribution Using Fluorescent Lipophilic Membrane Stains","volume":"2843","author":"Jones","year":"2024","journal-title":"Methods Mol. Biol."},{"key":"ref_45","doi-asserted-by":"crossref","first-page":"1900111","DOI":"10.1002\/adtp.201900111","article-title":"Chemical Modulation of Bioengineered Exosomes for Tissue-Specific Biodistribution","volume":"2","author":"Hwang","year":"2019","journal-title":"Adv. Ther."},{"key":"ref_46","doi-asserted-by":"crossref","first-page":"456","DOI":"10.1002\/smll.201401803","article-title":"In vivo kinetic biodistribution of nano-sized outer membrane vesicles derived from bacteria","volume":"11","author":"Jang","year":"2015","journal-title":"Small"},{"key":"ref_47","doi-asserted-by":"crossref","first-page":"6104","DOI":"10.1021\/acs.molpharmaceut.3c00460","article-title":"Elucidation of the Tissue Distribution and Host Immunostimulatory Activity of Exogenously Administered Probiotic-Derived Extracellular Vesicles for Immunoadjuvant","volume":"20","author":"Morishita","year":"2023","journal-title":"Mol. Pharm."},{"key":"ref_48","doi-asserted-by":"crossref","unstructured":"Turner, L., Bitto, N.J., Steer, D.L., Lo, C., D\u2019Costa, K., Ramm, G., Shambrook, M., Hill, A.F., Ferrero, R.L., and Kaparakis-Liaskos, M. (2018). Helicobacter pylori Outer Membrane Vesicle Size Determines Their Mechanisms of Host Cell Entry and Protein Content. Front. Immunol., 9.","DOI":"10.3389\/fimmu.2018.01466"},{"key":"ref_49","doi-asserted-by":"crossref","first-page":"103994","DOI":"10.1016\/j.jprot.2020.103994","article-title":"Characterization and proteomic analysis of outer membrane vesicles from a commensal microbe, Enterobacter cloacae","volume":"231","author":"Bhar","year":"2021","journal-title":"J. Proteom."},{"key":"ref_50","doi-asserted-by":"crossref","unstructured":"Jan, A.T. (2017). Outer Membrane Vesicles (OMVs) of Gram-negative Bacteria: A Perspective Update. Front. Microbiol., 8.","DOI":"10.3389\/fmicb.2017.01053"},{"key":"ref_51","doi-asserted-by":"crossref","first-page":"375","DOI":"10.1038\/nri3837","article-title":"Immune modulation by bacterial outer membrane vesicles","volume":"15","author":"Ferrero","year":"2015","journal-title":"Nat. Rev. Immunol."},{"key":"ref_52","first-page":"1462","article-title":"The extracellular vesicles from gram-positive bacteria: A review","volume":"38","author":"Ge","year":"2022","journal-title":"Chin. J. Biotechnol."},{"key":"ref_53","doi-asserted-by":"crossref","first-page":"21381","DOI":"10.1038\/s41598-025-99160-z","article-title":"Extracellular vesicles derived from Lactobacillus gasseri GFC-1220 alleviate inflammation via the TLR4\/NF-\u03baB signaling pathway in LPS-stimulated RAW264.7 macrophages","volume":"15","author":"Koh","year":"2025","journal-title":"Sci. Rep."},{"key":"ref_54","doi-asserted-by":"crossref","first-page":"620","DOI":"10.1038\/nrmicro3480","article-title":"Through the wall: Extracellular vesicles in Gram-positive bacteria, mycobacteria and fungi","volume":"13","author":"Brown","year":"2015","journal-title":"Nat. Rev. Microbiol."},{"key":"ref_55","doi-asserted-by":"crossref","unstructured":"Yang, J., Hwang, I., Lee, E., Shin, S.J., Lee, E.J., Rhee, J.H., and Yu, J.W. (2020). Bacterial Outer Membrane Vesicle-Mediated Cytosolic Delivery of Flagellin Triggers Host NLRC4 Canonical Inflammasome Signaling. Front. Immunol., 11.","DOI":"10.3389\/fimmu.2020.581165"},{"key":"ref_56","doi-asserted-by":"crossref","first-page":"e01188-17","DOI":"10.1128\/mBio.01188-17","article-title":"Inflammasome Activation by Bacterial Outer Membrane Vesicles Requires Guanylate Binding Proteins","volume":"8","author":"Finethy","year":"2017","journal-title":"mBio"},{"key":"ref_57","doi-asserted-by":"crossref","unstructured":"Bielaszewska, M., R\u00fcter, C., Bauwens, A., Greune, L., Jarosch, K.A., Steil, D., Zhang, W., He, X., Lloubes, R., and Fruth, A. (2017). Host cell interactions of outer membrane vesicle-associated virulence factors of enterohemorrhagic Escherichia coli O157: Intracellular delivery, trafficking and mechanisms of cell injury. PLoS Pathog., 13.","DOI":"10.1371\/journal.ppat.1006159"},{"key":"ref_58","doi-asserted-by":"crossref","first-page":"177","DOI":"10.1007\/978-1-0716-1339-9_8","article-title":"Virulence Factor Cargo and Host Cell Interactions of Shiga Toxin-Producing Escherichia coli Outer Membrane Vesicles","volume":"2291","author":"Bielaszewska","year":"2021","journal-title":"Methods Mol. Biol."},{"key":"ref_59","doi-asserted-by":"crossref","first-page":"240","DOI":"10.1021\/acsomega.1c04549","article-title":"Outer Membrane Vesicles Secreted by Helicobacter pylori Transmitting Gastric Pathogenic Virulence Factors","volume":"7","author":"Wei","year":"2021","journal-title":"ACS Omega"},{"key":"ref_60","doi-asserted-by":"crossref","first-page":"e98089","DOI":"10.15252\/embj.201798089","article-title":"LPS targets host guanylate-binding proteins to the bacterial outer membrane for non-canonical inflammasome activation","volume":"37","author":"Santos","year":"2018","journal-title":"EMBO J."},{"key":"ref_61","doi-asserted-by":"crossref","unstructured":"Amatya, S.B., Salmi, S., Kainulainen, V., Karihtala, P., and Reunanen, J. (2021). Bacterial Extracellular Vesicles in Gastrointestinal Tract Cancer: An Unexplored Territory. Cancers, 13.","DOI":"10.3390\/cancers13215450"},{"key":"ref_62","doi-asserted-by":"crossref","first-page":"6059","DOI":"10.1038\/s41598-025-90466-6","article-title":"Mastitis-related Staphylococcus aureus-derived extracellular vesicles induce a pro-inflammatory response in bovine monocyte-derived macrophages","volume":"15","author":"Silvestrelli","year":"2025","journal-title":"Sci. Rep."},{"key":"ref_63","doi-asserted-by":"crossref","unstructured":"Alipoor, S.D., and Elieh-Ali-Komi, D. (2024). Significance of extracellular vesicles in orchestration of immune responses in Mycobacterium tuberculosis infection. Front. Cell. Infect. Microbiol., 14.","DOI":"10.3389\/fcimb.2024.1398077"},{"key":"ref_64","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1111\/omi.12272","article-title":"Characterization and immunostimulatory activity of extracellular vesicles from Filifactor alocis","volume":"35","author":"Kim","year":"2020","journal-title":"Mol. Oral Microbiol."},{"key":"ref_65","doi-asserted-by":"crossref","unstructured":"Wang, H., Sun, J., Ma, G., You, F., Heng, B.C., Bai, Y., and Deng, X. (2025). The role and mechanism of bacterial outer membrane vesicles in the development of periodontitis. Front. Microbiol., 16.","DOI":"10.3389\/fmicb.2025.1654137"},{"key":"ref_66","doi-asserted-by":"crossref","first-page":"106994","DOI":"10.1016\/j.micpath.2024.106994","article-title":"Streptococcus mutans outer membrane vesicles affect inflammasome activation and the glycolysis of macrophages","volume":"196","author":"Song","year":"2024","journal-title":"Microb. Pathog."},{"key":"ref_67","doi-asserted-by":"crossref","unstructured":"Molina-Tijeras, J.A., G\u00e1lvez, J., and Rodr\u00edguez-Cabezas, M.E. (2019). The Immunomodulatory Properties of Extracellular Vesicles Derived from Probiotics: A Novel Approach for the Management of Gastrointestinal Diseases. Nutrients, 11.","DOI":"10.3390\/nu11051038"},{"key":"ref_68","doi-asserted-by":"crossref","first-page":"216","DOI":"10.1186\/s10020-024-00959-9","article-title":"Probiotic bacteria-released extracellular vesicles enhance macrophage phagocytosis in polymicrobial sepsis by activating the FPR1\/2 pathway","volume":"30","author":"Zhu","year":"2024","journal-title":"Mol. Med."},{"key":"ref_69","doi-asserted-by":"crossref","unstructured":"Hu, R., Lin, H., Li, J., Zhao, Y., Wang, M., Sun, X., Min, Y., Gao, Y., and Yang, M. (2020). Probiotic Escherichia coli Nissle 1917-derived outer membrane vesicles enhance immunomodulation and antimicrobial activity in RAW264.7 macrophages. BMC Microbiol., 20.","DOI":"10.1186\/s12866-020-01953-x"},{"key":"ref_70","unstructured":"Wang, M. (2021). Construction of Extracellular Vesicle Surface Display System of Intestinal Lactobacillus and Its Effect on the Immune Function of Porcine Intestinal Epithelial Cells. [Master\u2019s Thesis, Northwest A&F University]. Available online: https:\/\/link.cnki.net\/doi\/10.27409\/d.cnki.gxbnu.2021.000904."},{"key":"ref_71","doi-asserted-by":"crossref","unstructured":"\u00d1ahui Palomino, R.A., Vanpouille, C., Costantini, P.E., and Margolis, L. (2021). Microbiota-host communications: Bacterial extracellular vesicles as a common language. PLoS Pathog., 17.","DOI":"10.1371\/journal.ppat.1009508"},{"key":"ref_72","doi-asserted-by":"crossref","unstructured":"Liu, Y., Defourny, K., Smid, E.J., and Abee, T. (2018). Gram-Positive Bacterial Extracellular Vesicles and Their Impact on Health and Disease. Front. Microbiol., 9.","DOI":"10.3389\/fmicb.2018.01502"},{"key":"ref_73","doi-asserted-by":"crossref","unstructured":"Alvarez, C.S., Gim\u00e9nez, R., Ca\u00f1as, M.A., Vera, R., D\u00edaz-Garrido, N., Badia, J., and Baldom\u00e0, L. (2019). Extracellular vesicles and soluble factors secreted by Escherichia coli Nissle 1917 and ECOR63 protect against enteropathogenic E. coli-induced intestinal epithelial barrier dysfunction. BMC Microbiol., 19.","DOI":"10.1186\/s12866-019-1534-3"},{"key":"ref_74","doi-asserted-by":"crossref","unstructured":"F\u00e1brega, M.J., Aguilera, L., Gim\u00e9nez, R., Varela, E., Alexandra Ca\u00f1as, M., Antol\u00edn, M., Bad\u00eda, J., and Baldom\u00e0, L. (2016). Activation of Immune and Defense Responses in the Intestinal Mucosa by Outer Membrane Vesicles of Commensal and Probiotic Escherichia coli Strains. Front. Microbiol., 7.","DOI":"10.3389\/fmicb.2016.00705"},{"key":"ref_75","first-page":"401","article-title":"Oral probiotic extracellular vesicle therapy mitigates Influenza A Virus infection via blunting IL-17 signaling","volume":"45","author":"Zhou","year":"2024","journal-title":"Bioact. Mater."},{"key":"ref_76","doi-asserted-by":"crossref","first-page":"612","DOI":"10.1186\/s12951-025-03702-6","article-title":"Lactobacillus johnsonii-derived extracellular vesicles restore mucosal immunity via taurine-linked Th17\/Treg and IgA\/IgG regulation in colitis","volume":"23","author":"Zhao","year":"2025","journal-title":"J. Nanobiotechnology"},{"key":"ref_77","doi-asserted-by":"crossref","unstructured":"Li, M., Lee, K., Hsu, M., Nau, G., Mylonakis, E., and Ramratnam, B. (2017). Lactobacillus-derived extracellular vesicles enhance host immune responses against vancomycin-resistant enterococci. BMC Microbiol., 17.","DOI":"10.1186\/s12866-017-0977-7"},{"key":"ref_78","doi-asserted-by":"crossref","first-page":"2566255","DOI":"10.1080\/21505594.2025.2566255","article-title":"Deciphering the immunomodulatory cross-talk: Bacterial extracellular vesicles in gut homeostasis","volume":"16","author":"Yi","year":"2025","journal-title":"Virulence"},{"key":"ref_79","doi-asserted-by":"crossref","first-page":"23","DOI":"10.12938\/bmfh.18-015","article-title":"IgA-enhancing effects of membrane vesicles derived from Lactobacillus sakei subsp. sakei NBRC15893","volume":"38","author":"Miyoshi","year":"2019","journal-title":"Biosci. Microbiota Food Health"},{"key":"ref_80","doi-asserted-by":"crossref","first-page":"509","DOI":"10.1016\/j.chom.2012.08.004","article-title":"Outer membrane vesicles of a human commensal mediate immune regulation and disease protection","volume":"12","author":"Shen","year":"2012","journal-title":"Cell Host Microbe"},{"key":"ref_81","doi-asserted-by":"crossref","first-page":"1116","DOI":"10.1126\/science.aad9948","article-title":"Gene-microbiota interactions contribute to the pathogenesis of inflammatory bowel disease","volume":"352","author":"Chu","year":"2016","journal-title":"Science"},{"key":"ref_82","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1080\/19490976.2017.1344810","article-title":"Outer membrane vesicles blebbing contributes to B. vulgatus mpk-mediated immune response silencing","volume":"9","author":"Maerz","year":"2018","journal-title":"Gut Microbes"},{"key":"ref_83","doi-asserted-by":"crossref","unstructured":"F\u00e1brega, M.J., Rodr\u00edguez-Nogales, A., Garrido-Mesa, J., Algieri, F., Bad\u00eda, J., Gim\u00e9nez, R., G\u00e1lvez, J., and Baldom\u00e0, L. (2017). Intestinal Anti-inflammatory Effects of Outer Membrane Vesicles from Escherichia coli Nissle 1917 in DSS-Experimental Colitis in Mice. Front. Microbiol., 8.","DOI":"10.3389\/fmicb.2017.01274"},{"key":"ref_84","doi-asserted-by":"crossref","first-page":"423","DOI":"10.1038\/s12276-019-0359-3","article-title":"Lactobacillus paracasei-derived extracellular vesicles attenuate the intestinal inflammatory response by augmenting the endoplasmic reticulum stress pathway","volume":"52","author":"Choi","year":"2020","journal-title":"Exp. Mol. Med."},{"key":"ref_85","doi-asserted-by":"crossref","unstructured":"Lee, K.S., Kim, Y., Lee, J.H., Shon, S., Kim, A., Pham, A.V.Q., Kim, C., Kim, D.H., Kim, Y.K., and Cho, E.G. (2023). Human Probiotic Lactobacillus paracasei-Derived Extracellular Vesicles Improve Tumor Necrosis Factor-\u03b1-Induced Inflammatory Phenotypes in Human Skin. Cells, 12.","DOI":"10.3390\/cells12242789"},{"key":"ref_86","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1016\/j.micpath.2017.06.016","article-title":"The inhibitory impacts of Lactobacillus rhamnosus GG-derived extracellular vesicles on the growth of hepatic cancer cells","volume":"110","author":"Behzadi","year":"2017","journal-title":"Microb. Pathog."},{"key":"ref_87","doi-asserted-by":"crossref","first-page":"88","DOI":"10.24191\/scl.v19i1.6353","article-title":"A Mini Review on Biofilm-Mediated Water Pollution: Mechanistic Interactions, Preventive Approaches, and Removal Strategies","volume":"19","author":"Syaida","year":"2025","journal-title":"Sci. Lett."},{"key":"ref_88","doi-asserted-by":"crossref","first-page":"108426","DOI":"10.1016\/j.micpath.2026.108426","article-title":"Exploring the antibacterial and antibiofilm potential of graphene nanosheets synthesized from candlenut shell waste","volume":"214","author":"Azmi","year":"2026","journal-title":"Microb. Pathog."},{"key":"ref_89","doi-asserted-by":"crossref","first-page":"108613","DOI":"10.1016\/j.micpath.2026.108613","article-title":"Structural, proteomic, and metabolomic insights into the antibacterial efficacy of nalidixic acid against Corynebacterium pseudotuberculosis","volume":"217","author":"Syaida","year":"2026","journal-title":"Microb. Pathog."},{"key":"ref_90","doi-asserted-by":"crossref","first-page":"247","DOI":"10.1038\/s41579-019-0158-9","article-title":"Bacteria and archaea on Earth and their abundance in biofilms","volume":"17","author":"Flemming","year":"2019","journal-title":"Nat. Rev. Microbiol."},{"key":"ref_91","first-page":"6631","article-title":"Formation of bacterial biofilm and anti-biofilm activity of antimicrobial peptides","volume":"33","author":"Zhang","year":"2021","journal-title":"Chin. J. Anim. Nutr."},{"key":"ref_92","doi-asserted-by":"crossref","first-page":"39","DOI":"10.1016\/j.micpath.2015.02.007","article-title":"Drug resistance of bacterial dental biofilm and the potential use of natural compounds as alternative for prevention and treatment","volume":"80","author":"Kouidhi","year":"2015","journal-title":"Microb. Pathog."},{"key":"ref_93","doi-asserted-by":"crossref","unstructured":"Yahya, M.F.Z.R., Jalil, M.T.M., Jamil, N.M., Nor, N.H.M., Alhajj, N., Siburian, R., and Majid, N.A. (2025). Biofilms and multidrug resistance: An emerging crisis and the need for multidisciplinary interventions. Front. Bioeng. Biotechnol., 13.","DOI":"10.3389\/fbioe.2025.1625356"},{"key":"ref_94","doi-asserted-by":"crossref","first-page":"753","DOI":"10.1007\/s12275-017-7274-x","article-title":"Antibiofilm agents: A new perspective for antimicrobial strategy","volume":"55","author":"Li","year":"2017","journal-title":"J. Microbiol."},{"key":"ref_95","doi-asserted-by":"crossref","first-page":"8649","DOI":"10.1007\/s00253-020-10873-9","article-title":"Antibiotic resistance related to biofilm formation in Streptococcus suis","volume":"104","author":"Yi","year":"2020","journal-title":"Appl. Microbiol. Biotechnol."},{"key":"ref_96","doi-asserted-by":"crossref","first-page":"98","DOI":"10.1016\/j.colsurfb.2016.11.031","article-title":"In-situ, time-lapse study of extracellular polymeric substance discharge in Streptococcus mutans biofilm","volume":"150","author":"Liu","year":"2017","journal-title":"Colloids Surf. B Biointerfaces"},{"key":"ref_97","first-page":"e006","article-title":"Antibiofilm potential of lactic acid bacteria: Mechanism of cell-free supernatant, metabolite insights, and prospects for edible coating applications in food quality control","volume":"5","author":"Yahya","year":"2025","journal-title":"Food Mater. Res."},{"key":"ref_98","doi-asserted-by":"crossref","first-page":"97","DOI":"10.55230\/mabjournal.v52i2.2614","article-title":"FTIR spectroscopic study of inhibition of chloroxylenol-based disinfectant against Salmonella enterica serovar Thyphimurium biofilm","volume":"52","author":"Johari","year":"2023","journal-title":"Malays. Appl. Biol."},{"key":"ref_99","doi-asserted-by":"crossref","first-page":"459","DOI":"10.1016\/j.colsurfb.2016.06.039","article-title":"Investigation of extracellular polymeric substances (EPS) properties of P. aeruginosa and B. subtilis and their role in bacterial adhesion","volume":"146","author":"Harimawan","year":"2016","journal-title":"Colloids Surf. B Biointerfaces"},{"key":"ref_100","doi-asserted-by":"crossref","first-page":"274","DOI":"10.1016\/j.biortech.2015.06.110","article-title":"Extracellular polymeric substances govern the development of biofilm and mass transfer of polycyclic aromatic hydrocarbons for improved biodegradation","volume":"193","author":"Zhang","year":"2015","journal-title":"Bioresour. Technol."},{"key":"ref_101","doi-asserted-by":"crossref","first-page":"493","DOI":"10.4155\/fmc.15.6","article-title":"Biofilm formation mechanisms and targets for developing antibiofilm agents","volume":"7","author":"Rabin","year":"2015","journal-title":"Future Med. Chem."},{"key":"ref_102","doi-asserted-by":"crossref","first-page":"11353","DOI":"10.1073\/pnas.1503058112","article-title":"Pel is a cationic exopolysaccharide that cross-links extracellular DNA in the Pseudomonas aeruginosa biofilm matrix","volume":"112","author":"Jennings","year":"2015","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_103","doi-asserted-by":"crossref","first-page":"126829","DOI":"10.1016\/j.micres.2021.126829","article-title":"Biofilm patterns in gram-positive and gram-negative bacteria","volume":"251","author":"Ruhal","year":"2021","journal-title":"Microbiol. Res."},{"key":"ref_104","doi-asserted-by":"crossref","first-page":"fnv117","DOI":"10.1093\/femsle\/fnv117","article-title":"The relationship between biofilm and outer membrane vesicles: A novel therapy overview","volume":"362","author":"Wang","year":"2015","journal-title":"FEMS Microbiol. Lett."},{"key":"ref_105","doi-asserted-by":"crossref","first-page":"14656","DOI":"10.1038\/s41598-023-41663-8","article-title":"Photoacoustic based evaluation of viscoelastic properties of Gram-negative and Gram-positive bacterial colonies","volume":"13","author":"Hosseindokht","year":"2023","journal-title":"Sci. Rep."},{"key":"ref_106","doi-asserted-by":"crossref","unstructured":"Wells, M., Schneider, R., Bhattarai, B., Currie, H., Chavez, B., Christopher, G., Rumbaugh, K., and Gordon, V. (2023). Perspective: The viscoelastic properties of biofilm infections and mechanical interactions with phagocytic immune cells. Front. Cell. Infect. Microbiol., 13.","DOI":"10.3389\/fcimb.2023.1102199"},{"key":"ref_107","doi-asserted-by":"crossref","unstructured":"Balmuri, S.R., Noaman, S., Usman, H., and Niepa, T.H.R. (2024). Altering the interfacial rheology of Pseudomonas aeruginosa and Staphylococcus aureus with N-acetyl cysteine and cysteamine. Front. Cell. Infect. Microbiol., 13.","DOI":"10.3389\/fcimb.2023.1338477"},{"key":"ref_108","doi-asserted-by":"crossref","unstructured":"Zarnowski, R., Sanchez, H., Covelli, A.S., Dominguez, E., Jaromin, A., Bernhardt, J., Mitchell, K.F., Heiss, C., Azadi, P., and Mitchell, A. (2018). Candida albicans biofilm-induced vesicles confer drug resistance through matrix biogenesis. PLoS Biol., 16.","DOI":"10.1371\/journal.pbio.2006872"},{"key":"ref_109","doi-asserted-by":"crossref","first-page":"4193","DOI":"10.1007\/s00253-019-09812-0","article-title":"Quantification of outer membrane vesicles: A potential tool to compare response in Pseudomonas putida KT2440 to stress caused by alkanols","volume":"103","author":"Eberlein","year":"2019","journal-title":"Appl. Microbiol. Biotechnol."},{"key":"ref_110","doi-asserted-by":"crossref","first-page":"1517","DOI":"10.2217\/fmb.15.63","article-title":"Biogenesis and function of Porphyromonas gingivalis outer membrane vesicles","volume":"10","author":"Xie","year":"2015","journal-title":"Future Microbiol."},{"key":"ref_111","doi-asserted-by":"crossref","first-page":"13","DOI":"10.1016\/j.actbio.2024.02.029","article-title":"Bacterial extracellular vesicles: Modulation of biofilm and virulence properties","volume":"178","author":"Jeong","year":"2024","journal-title":"Acta Biomater."},{"key":"ref_112","doi-asserted-by":"crossref","unstructured":"Cecil, J.D., Sirisaengtaksin, N., O\u2019Brien-Simpson, N.M., and Krachler, A.M. (2019). Outer Membrane Vesicle-Host Cell Interactions. Microbiol. Spectr., 7, 10.1128\/microbiolspec.PSIB-0001-2018.","DOI":"10.1128\/microbiolspec.PSIB-0001-2018"},{"key":"ref_113","doi-asserted-by":"crossref","first-page":"100183","DOI":"10.1016\/j.bioflm.2024.100183","article-title":"Dual roles of the conditional extracellular vesicles derived from Pseudomonas aeruginosa biofilms: Promoting and inhibiting bacterial biofilm growth","volume":"7","author":"Saad","year":"2024","journal-title":"Biofilm"},{"key":"ref_114","doi-asserted-by":"crossref","unstructured":"Ma, G., Ding, Y., Wu, Q., Zhang, J., Liu, M., Wang, Z., Wang, Z., Wu, S., Yang, X., and Li, Y. (2022). Yersinia enterocolitica-Derived Outer Membrane Vesicles Inhibit Initial Stage of Biofilm Formation. Microorganisms, 10.","DOI":"10.3390\/microorganisms10122357"},{"key":"ref_115","doi-asserted-by":"crossref","unstructured":"Seike, S., Kobayashi, H., Ueda, M., Takahashi, E., Okamoto, K., and Yamanaka, H. (2021). Outer Membrane Vesicles Released From Aeromonas Strains Are Involved in the Biofilm Formation. Front. Microbiol., 11.","DOI":"10.3389\/fmicb.2020.613650"},{"key":"ref_116","doi-asserted-by":"crossref","unstructured":"Vitse, J., and Devreese, B. (2020). The Contribution of Membrane Vesicles to Bacterial Pathogenicity in Cystic Fibrosis Infections and Healthcare Associated Pneumonia. Front. Microbiol., 11.","DOI":"10.3389\/fmicb.2020.00630"},{"key":"ref_117","doi-asserted-by":"crossref","first-page":"1199","DOI":"10.1016\/j.ijmm.2014.08.013","article-title":"Pseudomonas quinolone signalling system: A component of quorum sensing cascade is a crucial player in the acute urinary tract infection caused by Pseudomonas aeruginosa","volume":"304","author":"Bala","year":"2014","journal-title":"Int. J. Med. Microbiol. IJMM"},{"key":"ref_118","doi-asserted-by":"crossref","first-page":"e12507","DOI":"10.1002\/jev2.12507","article-title":"The activity of the quorum sensing regulator HapR is modulated by the bacterial extracellular vesicle (BEV)-associated protein ObfA of Vibrio cholerae","volume":"13","author":"Ebenberger","year":"2024","journal-title":"J. Extracell. Vesicles"},{"key":"ref_119","doi-asserted-by":"crossref","first-page":"1009","DOI":"10.1111\/1462-2920.15278","article-title":"Membrane vesicles from a Dietzia bacterium containing multiple cargoes and their roles in iron delivery","volume":"23","author":"Wang","year":"2021","journal-title":"Environ. Microbiol."},{"key":"ref_120","doi-asserted-by":"crossref","first-page":"32481","DOI":"10.1074\/jbc.C114.617142","article-title":"Peptidyl arginine deiminase from Porphyromonas gingivalis abolishes anaphylatoxin C5a activity","volume":"289","author":"Bielecka","year":"2014","journal-title":"J. Biol. Chem."},{"key":"ref_121","doi-asserted-by":"crossref","first-page":"5670","DOI":"10.1039\/D5NA00248F","article-title":"Biofilm formation on collagen substrates modulates Streptococcus mutans bacterial extracellular nanovesicle production and cargo","volume":"7","author":"Saavedra","year":"2025","journal-title":"Nanoscale Adv."},{"key":"ref_122","doi-asserted-by":"crossref","first-page":"823","DOI":"10.1080\/1040841X.2024.2427656","article-title":"Extracellular vesicle production by oral bacteria related to dental caries and periodontal disease: Role in microbe-host and interspecies interactions","volume":"51","author":"Saavedra","year":"2025","journal-title":"Crit. Rev. Microbiol."},{"key":"ref_123","doi-asserted-by":"crossref","unstructured":"Kudryakova, I., Afoshin, A., Tarlachkov, S., Leontyevskaya, E., Suzina, N., and Leontyevskaya Vasilyeva, N. (2023). Lysobacter gummosus 10.1.1, a Producer of Antimicrobial Agents. Microorganisms, 11.","DOI":"10.3390\/microorganisms11122853"},{"key":"ref_124","doi-asserted-by":"crossref","first-page":"645","DOI":"10.1007\/s00248-020-01614-6","article-title":"Relationship Between Membrane Vesicles, Extracellular ATP and Biofilm Formation in Antarctic Gram-Negative Bacteria","volume":"81","author":"Baeza","year":"2021","journal-title":"Microb. Ecol."},{"key":"ref_125","doi-asserted-by":"crossref","first-page":"5478","DOI":"10.1128\/JB.180.20.5478-5483.1998","article-title":"Gram-negative bacteria produce membrane vesicles which are capable of killing other bacteria","volume":"180","author":"Li","year":"1998","journal-title":"J. Bacteriol."},{"key":"ref_126","doi-asserted-by":"crossref","first-page":"101175","DOI":"10.1016\/j.crfs.2025.101175","article-title":"Extracellular vesicles from Lactiplantibacillus plantarum inhibit the biofilm formation of Listeria monocytogenes and reduce bacterial contamination on romaine lettuce","volume":"11","author":"Park","year":"2025","journal-title":"Curr. Res. Food Sci."},{"key":"ref_127","doi-asserted-by":"crossref","first-page":"2742","DOI":"10.1099\/mic.0.060343-0","article-title":"Predatory activity of Myxococcus xanthus outer-membrane vesicles and properties of their hydrolase cargo","volume":"158","author":"Evans","year":"2012","journal-title":"Microbiology"},{"key":"ref_128","doi-asserted-by":"crossref","unstructured":"Berleman, J.E., Allen, S., Danielewicz, M.A., Remis, J.P., Gorur, A., Cunha, J., Hadi, M.Z., Zusman, D.R., Northen, T.R., and Witkowska, H.E. (2014). The lethal cargo of Myxococcus xanthus outer membrane vesicles. Front. Microbiol., 5.","DOI":"10.3389\/fmicb.2014.00474"},{"key":"ref_129","doi-asserted-by":"crossref","first-page":"1476","DOI":"10.1128\/AAC.42.6.1476","article-title":"Delivery of the non-membrane-permeative antibiotic gentamicin into mammalian cells by using Shigella flexneri membrane vesicles","volume":"42","author":"Kadurugamuwa","year":"1998","journal-title":"Antimicrob. Agents Chemother."},{"key":"ref_130","doi-asserted-by":"crossref","first-page":"2767","DOI":"10.1128\/jb.178.10.2767-2774.1996","article-title":"Bacteriolytic effect of membrane vesicles from Pseudomonas aeruginosa on other bacteria including pathogens: Conceptually new antibiotics","volume":"178","author":"Kadurugamuwa","year":"1996","journal-title":"J. Bacteriol."},{"key":"ref_131","doi-asserted-by":"crossref","first-page":"e70194","DOI":"10.1002\/jev2.70194","article-title":"High-Yield Outer Membrane Vesicles Derived From Probiotics as a Nanoplatform for Precise Treatment and Prophylaxis of Pseudomonas aeruginosa Infection","volume":"14","author":"Yang","year":"2025","journal-title":"J. Extracell. Vesicles"},{"key":"ref_132","doi-asserted-by":"crossref","first-page":"100216","DOI":"10.1016\/j.bioflm.2024.100216","article-title":"Bacterial extracellular vesicle: A non-negligible component in biofilm life cycle and challenges in biofilm treatments","volume":"8","author":"Chen","year":"2024","journal-title":"Biofilm"},{"key":"ref_133","doi-asserted-by":"crossref","first-page":"2704","DOI":"10.1039\/C7MB00365J","article-title":"Staphylococcus aureus extracellular vesicles (EVs): Surface-binding antagonists of biofilm formation","volume":"13","author":"Im","year":"2017","journal-title":"Mol. BioSyst."},{"key":"ref_134","doi-asserted-by":"crossref","first-page":"2242","DOI":"10.1093\/ofid\/ofw172.1790","article-title":"Gram-Negative Bacterial Outer Membrane Vesicles Inhibit Growth of Multidrug-Resistant Organisms and Induce Wound-Healing Cytokines","volume":"3","author":"Baker","year":"2016","journal-title":"Open Forum Infect. Dis."},{"key":"ref_135","doi-asserted-by":"crossref","unstructured":"Federica, D., Cosimato, I., Salzano, F., Mensitieri, F., Andretta, V., Santoro, E., Boccia, G., Folliero, V., and Franci, G. (2025). Adaptations of Bacterial Extracellular Vesicles in Response to Antibiotic Pressure. Int. J. Mol. Sci., 26.","DOI":"10.3390\/ijms26115025"},{"key":"ref_136","doi-asserted-by":"crossref","first-page":"77","DOI":"10.1016\/j.biochi.2025.07.024","article-title":"Mechanisms of outer membrane vesicles in bacterial drug resistance: Insights and implications","volume":"238","author":"Zhang","year":"2025","journal-title":"Biochimie"},{"key":"ref_137","doi-asserted-by":"crossref","unstructured":"Kosgodage, U.S., Matewele, P., Awamaria, B., Kraev, I., Warde, P., Mastroianni, G., Nunn, A.V., Guy, G.W., Bell, J.D., and Inal, J.M. (2019). Cannabidiol Is a Novel Modulator of Bacterial Membrane Vesicles. Front. Cell. Infect. Microbiol., 9.","DOI":"10.3389\/fcimb.2019.00324"},{"key":"ref_138","doi-asserted-by":"crossref","first-page":"20","DOI":"10.1186\/s13099-023-00543-2","article-title":"Emerging role of bacterial outer membrane vesicle in gastrointestinal tract","volume":"15","author":"Tian","year":"2023","journal-title":"Gut Pathog."},{"key":"ref_139","doi-asserted-by":"crossref","unstructured":"Olivo-Mart\u00ednez, Y., Mart\u00ednez-Ruiz, S., Cordero, C., Badia, J., and Baldoma, L. (2024). Extracellular Vesicles of the Probiotic Escherichia coli Nissle 1917 Reduce PepT1 Levels in IL-1\u03b2-Treated Caco-2 Cells via Upregulation of miR-193a-3p. Nutrients, 16.","DOI":"10.3390\/nu16162719"},{"key":"ref_140","doi-asserted-by":"crossref","unstructured":"Gong, S., Zeng, R., Liu, L., Wang, R., Xue, M., Dong, H., Wu, Z., and Zhang, Y. (2024). Extracellular vesicles from a novel Lactiplantibacillus plantarum strain suppress inflammation and promote M2 macrophage polarization. Front. Immunol., 15.","DOI":"10.3389\/fimmu.2024.1459213"},{"key":"ref_141","doi-asserted-by":"crossref","unstructured":"Zhang, Q., He, K., Aazmi, M.S., and Raja Yahya, M.F.Z. (2026). Clostridium butyricum extracellular vesicles remodel the transcriptional network of pyroptosis-related genes in LPS-stimulated macrophages. Front. Immunol., 16.","DOI":"10.3389\/fimmu.2025.1686347"},{"key":"ref_142","doi-asserted-by":"crossref","first-page":"3005","DOI":"10.1016\/j.tibtech.2025.05.022","article-title":"Application of bacterial extracellular vesicles in gastrointestinal diseases","volume":"43","author":"Wang","year":"2025","journal-title":"Trends Biotechnol."},{"key":"ref_143","doi-asserted-by":"crossref","unstructured":"Macia, L., Nanan, R., Hosseini-Beheshti, E., and Grau, G.E. (2019). Host- and Microbiota-Derived Extracellular Vesicles, Immune Function, and Disease Development. Int. J. Mol. Sci., 21.","DOI":"10.3390\/ijms21010107"},{"key":"ref_144","doi-asserted-by":"crossref","first-page":"101","DOI":"10.1007\/978-3-030-67171-6_6","article-title":"Pathogenesis Mediated by Bacterial Membrane Vesicles","volume":"97","author":"Gilmore","year":"2021","journal-title":"Subcell. Biochem."},{"key":"ref_145","doi-asserted-by":"crossref","first-page":"419","DOI":"10.1042\/BST20240150","article-title":"Recent advances in understanding the role of extracellular vesicles from probiotics in intestinal immunity signaling","volume":"53","author":"Kurata","year":"2025","journal-title":"Biochem. Soc. Trans."},{"key":"ref_146","doi-asserted-by":"crossref","first-page":"621","DOI":"10.1111\/jfd.13159","article-title":"Identification and characterization of outer membrane vesicles from the fish pathogen Vibrio ordalii","volume":"43","author":"Irgang","year":"2020","journal-title":"J. Fish. Dis."},{"key":"ref_147","doi-asserted-by":"crossref","first-page":"e2300269","DOI":"10.1002\/pmic.202300269","article-title":"The effect of altered pH growth conditions on the production, composition, and proteomes of Helicobacter pylori outer membrane vesicles","volume":"24","author":"Johnston","year":"2024","journal-title":"Proteomics"},{"key":"ref_148","doi-asserted-by":"crossref","first-page":"e70101","DOI":"10.1111\/1758-2229.70101","article-title":"Functional Cargo in Membrane Vesicles From a Citrus Pathogen","volume":"17","author":"Araujo","year":"2025","journal-title":"Environ. Microbiol. Rep."},{"key":"ref_149","doi-asserted-by":"crossref","unstructured":"Marquez-Paradas, E., Torrecillas-Lopez, M., Barrera-Chamorro, L., Del Rio-Vazquez, J.L., Gonzalez-de la Rosa, T., and Montserrat-de la Paz, S. (2025). Microbiota-derived extracellular vesicles: Current knowledge, gaps, and challenges in precision nutrition. Front. Immunol., 16, Erratum in Front. Immunol. 2025\u00a016, 1599029.","DOI":"10.3389\/fimmu.2025.1514726"},{"key":"ref_150","doi-asserted-by":"crossref","first-page":"3424","DOI":"10.1002\/pmic.201000212","article-title":"Proteomic analysis of outer membrane vesicles derived from Pseudomonas aeruginosa","volume":"11","author":"Choi","year":"2011","journal-title":"Proteomics"},{"key":"ref_151","doi-asserted-by":"crossref","unstructured":"Atshan, S.S., Shamsudin, M.N., Sekawi, Z., Thian Lung, L.T., Barantalab, F., Liew, Y.K., Alreshidi, M.A., Abduljaleel, S.A., and Hamat, R.A. (2015). Comparative proteomic analysis of extracellular proteins expressed by various clonal types of Staphylococcus aureus and during planktonic growth and biofilm development. Front Microbiol., 6.","DOI":"10.3389\/fmicb.2015.00524"},{"key":"ref_152","doi-asserted-by":"crossref","first-page":"100","DOI":"10.1016\/j.molp.2025.11.003","article-title":"Cross-kingdom RNA trafficking from bacteria to fungi enables plant protection against fungal pathogens","volume":"19","author":"Wu","year":"2026","journal-title":"Mol. Plant"},{"key":"ref_153","doi-asserted-by":"crossref","first-page":"90","DOI":"10.1186\/s40168-025-02078-x","article-title":"Faecalibacterium prausnitzii-derived outer membrane vesicles reprogram gut microbiota metabolism to alleviate Porcine Epidemic Diarrhea Virus infection","volume":"13","author":"Xing","year":"2025","journal-title":"Microbiome"},{"key":"ref_154","doi-asserted-by":"crossref","first-page":"791","DOI":"10.20517\/evcna.2025.32","article-title":"Special delivery-extracellular vesicles released by commensal gut bacteria deliver bioactive protein to distal organs","volume":"6","author":"Jones","year":"2025","journal-title":"Extracell. Vesicles Circ. Nucl. Acids"},{"key":"ref_155","doi-asserted-by":"crossref","first-page":"1021","DOI":"10.1042\/BST20180114","article-title":"Fantastic voyage: The journey of intestinal microbiota-derived microvesicles through the body","volume":"46","author":"Stentz","year":"2018","journal-title":"Biochem Soc. Trans."},{"key":"ref_156","doi-asserted-by":"crossref","unstructured":"Kameli, N. (2025). Vesicle-Mediated Transfer of CTX-M \u03b2-Lactamase Genes and Proteins Confers Ampicillin Resistance in Escherichia coli. Int. J. Mol. Sci., 26.","DOI":"10.3390\/ijms262110601"},{"key":"ref_157","doi-asserted-by":"crossref","unstructured":"Kroniger, T., Mehanny, M., Schl\u00fcter, R., Trautwein-Schult, A., K\u00f6llner, B., and Becher, D. (2022). Effect of Iron Limitation, Elevated Temperature, and Florfenicol on the Proteome and Vesiculation of the Fish Pathogen Aeromonas salmonicida. Microorganisms, 10.","DOI":"10.3390\/microorganisms10091735"},{"key":"ref_158","doi-asserted-by":"crossref","unstructured":"Ca\u00f1as, M.A., F\u00e1brega, M.J., Gim\u00e9nez, R., Badia, J., and Baldom\u00e0, L. (2018). Outer Membrane Vesicles From Probiotic and Commensal Escherichia coli Activate NOD1-Mediated Immune Responses in Intestinal Epithelial Cells. Front. Microbiol., 9.","DOI":"10.3389\/fmicb.2018.00498"},{"key":"ref_159","doi-asserted-by":"crossref","unstructured":"Mao, H., Gong, T., Sun, Y., Yang, S., Qiao, X., and Yang, D. (2023). Bacterial growth stage determines the yields, protein composition, and periodontal pathogenicity of Porphyromonas gingivalis outer membrane vesicles. Front. Cell. Infect. Microbiol., 13.","DOI":"10.3389\/fcimb.2023.1193198"},{"key":"ref_160","doi-asserted-by":"crossref","first-page":"4029","DOI":"10.7150\/ijbs.94156","article-title":"Helicobacter pylori Outer Membrane Vesicles: Biogenesis, Composition, and Biological Functions","volume":"20","author":"Li","year":"2024","journal-title":"Int. J. Biol. Sci."},{"key":"ref_161","doi-asserted-by":"crossref","first-page":"e0181922","DOI":"10.1128\/spectrum.01819-22","article-title":"Outer Membrane Vesicles of Actinobacillus pleuropneumoniae Exert Immunomodulatory Effects on Porcine Alveolar Macrophages","volume":"10","author":"Zhu","year":"2022","journal-title":"Microbiol. Spectr."},{"key":"ref_162","doi-asserted-by":"crossref","first-page":"19259","DOI":"10.1021\/acs.jafc.4c04223","article-title":"Immunomodulatory Effects of Probiotic-Derived Extracellular Vesicles: Opportunities and Challenges","volume":"72","author":"Chen","year":"2024","journal-title":"J. Agric. Food Chem."},{"key":"ref_163","doi-asserted-by":"crossref","unstructured":"Alvarez, C.S., Badia, J., Bosch, M., Gim\u00e9nez, R., and Baldom\u00e0, L. (2016). Outer Membrane Vesicles and Soluble Factors Released by Probiotic Escherichia coli Nissle 1917 and Commensal ECOR63 Enhance Barrier Function by Regulating Expression of Tight Junction Proteins in Intestinal Epithelial Cells. Front. Microbiol., 7.","DOI":"10.3389\/fmicb.2016.01981"},{"key":"ref_164","doi-asserted-by":"crossref","first-page":"e0043922","DOI":"10.1128\/iai.00439-22","article-title":"Role of Bacterial Extracellular Vesicles in Manipulating Infection","volume":"91","author":"Zhao","year":"2023","journal-title":"Infect. Immun."},{"key":"ref_165","doi-asserted-by":"crossref","first-page":"127788","DOI":"10.1016\/j.micres.2024.127788","article-title":"Effects of bacterial extracellular vesicles derived from oral and gastrointestinal pathogens on systemic diseases","volume":"285","author":"Zhang","year":"2024","journal-title":"Microbiol. Res."},{"key":"ref_166","doi-asserted-by":"crossref","unstructured":"Behrouzi, A., Vaziri, F., Riazi Rad, F., Amanzadeh, A., Fateh, A., Moshiri, A., Khatami, S., and Siadat, S.D. (2018). Comparative study of pathogenic and non-pathogenic Escherichia coli outer membrane vesicles and prediction of host-interactions with TLR signaling pathways. BMC Res. Notes, 11.","DOI":"10.1186\/s13104-018-3648-3"},{"key":"ref_167","doi-asserted-by":"crossref","first-page":"e0170721","DOI":"10.1128\/mBio.01707-21","article-title":"Bacterial Outer Membrane Vesicles as a Versatile Tool in Vaccine Research and the Fight against Antimicrobial Resistance","volume":"12","author":"Zhu","year":"2021","journal-title":"mBio"},{"key":"ref_168","doi-asserted-by":"crossref","first-page":"1080","DOI":"10.1021\/acs.molpharmaceut.0c01011","article-title":"Characterizing Different Probiotic-Derived Extracellular Vesicles as a Novel Adjuvant for Immunotherapy","volume":"18","author":"Morishita","year":"2021","journal-title":"Mol. Pharm."},{"key":"ref_169","first-page":"1301","article-title":"Antitumor effect of Escherichia coli-derived outer membrane vesicles on neuroblastoma in vitro and in vivo","volume":"54","author":"Jin","year":"2022","journal-title":"Acta Biochim. Biophys. Sin."},{"key":"ref_170","doi-asserted-by":"crossref","first-page":"e12514","DOI":"10.1002\/jev2.12514","article-title":"Enhancing protective immunity against bacterial infection via coating nano-Rehmannia glutinosa polysaccharide with outer membrane vesicles","volume":"13","author":"Huang","year":"2024","journal-title":"J. Extracell. Vesicles"},{"key":"ref_171","doi-asserted-by":"crossref","first-page":"13867","DOI":"10.1039\/D5TB01708D","article-title":"Bacterial outer membrane vesicles in immune modulation: From mechanisms to applications","volume":"13","author":"Wang","year":"2025","journal-title":"J. Mater. Chem. B"},{"key":"ref_172","doi-asserted-by":"crossref","first-page":"174","DOI":"10.1007\/s11274-024-03963-7","article-title":"Bacterial extracellular vesicles: Biotechnological perspective for enhanced productivity","volume":"40","author":"Geiger","year":"2024","journal-title":"World J. Microbiol. Biotechnol."},{"key":"ref_173","doi-asserted-by":"crossref","first-page":"1","DOI":"10.20517\/evcna.2024.76","article-title":"Consistency in bacterial extracellular vesicle production: Key to their application in human health","volume":"6","author":"Dai","year":"2025","journal-title":"Extracell. Vesicles Circ. Nucleic Acids"},{"key":"ref_174","doi-asserted-by":"crossref","first-page":"626","DOI":"10.1038\/s41467-017-00729-8","article-title":"Bacterial outer membrane vesicles suppress tumor by interferon-\u03b3-mediated antitumor response","volume":"8","author":"Kim","year":"2017","journal-title":"Nat. Commun."}],"container-title":["Biomolecules"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.mdpi.com\/2218-273X\/16\/8\/1132\/pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,8,6]],"date-time":"2026-08-06T04:19:16Z","timestamp":1785989956000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.mdpi.com\/2218-273X\/16\/8\/1132"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,8,3]]},"references-count":174,"journal-issue":{"issue":"8","published-online":{"date-parts":[[2026,8]]}},"alternative-id":["biom16081132"],"URL":"https:\/\/doi.org\/10.3390\/biom16081132","relation":{},"ISSN":["2218-273X"],"issn-type":[{"value":"2218-273X","type":"electronic"}],"subject":[],"published":{"date-parts":[[2026,8,3]]}}}