{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,10,1]],"date-time":"2026-10-01T04:59:55Z","timestamp":1790830795032,"version":"4.1.0"},"reference-count":73,"publisher":"Frontiers Media SA","license":[{"start":{"date-parts":[[2026,10,1]],"date-time":"2026-10-01T00:00:00Z","timestamp":1790812800000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"DOI":"10.13039\/501100001659","name":"Deutsche Forschungsgemeinschaft","doi-asserted-by":"publisher","id":[{"id":"10.13039\/501100001659","id-type":"DOI","asserted-by":"publisher"}]}],"content-domain":{"domain":["frontiersin.org"],"crossmark-restriction":true},"short-container-title":["Front. Cell. Infect. Microbiol."],"abstract":"<jats:sec>\n                    <jats:title>Introduction<\/jats:title>\n                    <jats:p>Bacterial membrane vesicles (BMVs) are key mediators of host-pathogen interaction and intercellular communication. However, their characterization has been conducted almost exclusively under planktonic growth conditions, even though biofilm formation is the predominant mode of bacterial persistence in chronic and device-associated infections. How the shift from planktonic to biofilm growth reshapes BMV composition and function, and how this is affecting the host, remains poorly understood.<\/jats:p>\n                  <\/jats:sec>\n                  <jats:sec>\n                    <jats:title>Methods<\/jats:title>\n                    <jats:p>\n                      Here, we systematically compared BMVs from the ESKAPE pathogens\n                      <jats:italic>Staphylococcus aureus<\/jats:italic>\n                      and\n                      <jats:italic>Pseudomonas aeruginosa<\/jats:italic>\n                      , representing Gram-positive and Gram-negative bacteria, across both growth conditions. Particle sizes and number-weighted size distributions were obtained by dynamic light scattering and nanoparticle tracking analysis, respectively.\n                    <\/jats:p>\n                  <\/jats:sec>\n                  <jats:sec>\n                    <jats:title>Results<\/jats:title>\n                    <jats:p>\n                      Cryogenic electron micrographs showed intact, membrane-enclosed vesicles for all four isolations. Biofilm-derived BMVs exhibited a more negative surface charge compared to their planktonic counterparts. Quantitative proteomics identified growth mode as a strong determinant of cargo: planktonic BMVs were enriched in ribosomal proteins, whereas biofilm-derived BMVs carried central metabolic enzymes. Notably, exopolysaccharide biosynthesis proteins of the Psl, Pel, and Ica systems were more abundant in planktonic than in biofilm BMVs.\n                      <jats:italic>P. aeruginosa<\/jats:italic>\n                      BMVs isolated from biofilms carried heme and iron acquisition systems, proteins of the oxidative stress defense, and the secreted proteases LasB and AprA, while planktonic\n                      <jats:italic>S. aureus<\/jats:italic>\n                      BMVs were enriched in secreted virulence factors, highlighting species-level differences. Functionally, all preparations reduced metabolic activity in epithelial cells with only limited lactate dehydrogenase release, identifying metabolic suppression rather than membrane rupture as the dominant effect. In the case of\n                      <jats:italic>P. aeruginosa<\/jats:italic>\n                      , biofilm-derived BMVs suppressed metabolic activity significantly more than planktonic ones. In human macrophages, planktonic BMVs elicited the strongest IL-1\u03b2 and TNF-\u03b1 responses, whereas biofilm-derived vesicles induced earlier secretion of IL-6 and, for\n                      <jats:italic>P. aeruginosa<\/jats:italic>\n                      , a distinctly reduced release of the anti-inflammatory cytokine IL-10. These effects persisted under polymyxin B, which reduced but did not abolish the overall cytokine responses.\n                    <\/jats:p>\n                  <\/jats:sec>\n                  <jats:sec>\n                    <jats:title>Discussion<\/jats:title>\n                    <jats:p>This indicates that vesicle cargo does not simply mirror the biofilm matrix, pointing to a contribution of vesicle-associated components beyond lipopolysaccharide. Together, these data establish a comparative framework linking bacterial growth mode to BMV composition and host cell response, advancing the understanding of vesicle-mediated processes in chronic infection.<\/jats:p>\n                  <\/jats:sec>","DOI":"10.3389\/fcimb.2026.1881620","type":"journal-article","created":{"date-parts":[[2026,10,1]],"date-time":"2026-10-01T04:42:32Z","timestamp":1790829752000},"update-policy":"https:\/\/doi.org\/10.3389\/crossmark-policy","source":"Crossref","is-referenced-by-count":0,"title":["Proteomic profiling reveals growth condition-driven functional differences in bacterial membrane vesicles from planktonic and biofilm cultures"],"prefix":"10.3389","volume":"16","author":[{"given":"Nathalie","family":"Jung","sequence":"first","affiliation":[{"name":"Institute of Pharmaceutical Technology, Goethe University Frankfurt","place":["Frankfurt am Main, Germany"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Lennart","family":"Christe","sequence":"additional","affiliation":[{"name":"Institute of Pharmaceutical Technology, Goethe University Frankfurt","place":["Frankfurt am Main, Germany"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Stefanie","family":"Gier","sequence":"additional","affiliation":[{"name":"Institute of Pharmaceutical Technology, Goethe University Frankfurt","place":["Frankfurt am Main, Germany"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Jana Romy","family":"Friedrich","sequence":"additional","affiliation":[{"name":"Institute of Pharmaceutical Technology, Goethe University Frankfurt","place":["Frankfurt am Main, Germany"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Fadi","family":"Haddad","sequence":"additional","affiliation":[{"name":"Institute of Pharmaceutical Technology, Goethe University Frankfurt","place":["Frankfurt am Main, Germany"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Pauline","family":"Roth","sequence":"additional","affiliation":[{"name":"Buchmann Institute for Molecular Life Sciences and Institute for Biophysics, Goethe University Frankfurt","place":["Frankfurt am Main, Germany"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Jo\u00e3o","family":"Mello-Viera","sequence":"additional","affiliation":[{"name":"Molecular Signaling Group, Buchmann Institute for Molecular Life Sciences, Goethe University Frankfurt","place":["Frankfurt am Main, Germany"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Thorsten","family":"Mosler","sequence":"additional","affiliation":[{"name":"Molecular Signaling Group, Institute of Biochemistry II, Medical Faculty, Goethe University Frankfurt","place":["Frankfurt am Main, Germany"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Stefan","family":"Schiller","sequence":"additional","affiliation":[{"name":"Institute of Pharmaceutical Technology, Goethe University Frankfurt","place":["Frankfurt am Main, Germany"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Achilleas S.","family":"Frangakis","sequence":"additional","affiliation":[{"name":"Buchmann Institute for Molecular Life Sciences and Institute for Biophysics, Goethe University Frankfurt","place":["Frankfurt am Main, Germany"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Ivan","family":"\u0110iki\u0107","sequence":"additional","affiliation":[{"name":"Molecular Signaling Group, Buchmann Institute for Molecular Life Sciences, Goethe University Frankfurt","place":["Frankfurt am Main, Germany"]},{"name":"Molecular Signaling Group, Institute of Biochemistry II, Medical Faculty, Goethe University Frankfurt","place":["Frankfurt am Main, Germany"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Maike","family":"Windbergs","sequence":"additional","affiliation":[{"name":"Institute of Pharmaceutical Technology, Goethe University Frankfurt","place":["Frankfurt am Main, Germany"]}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"1965","published-online":{"date-parts":[[2026,10,1]]},"reference":[{"key":"B1","doi-asserted-by":"publisher","first-page":"590","DOI":"10.1089\/mdr.2017.0147","article-title":"Vancomycin-resistant enterococci: a review of antimicrobial resistance mechanisms and perspectives of human and animal health","volume":"24","author":"Ahmed","year":"2018","journal-title":"Microb. Drug Resist."},{"key":"B2","doi-asserted-by":"publisher","DOI":"10.1016\/J.SJBS.2023.103604","article-title":"Antimicrobial resistance in methicillin-resistant Staphylococcus aureus","volume":"30","author":"Ali Alghamdi","year":"2023","journal-title":"Saudi J. Biol. Sci."},{"key":"B3","doi-asserted-by":"publisher","DOI":"10.3390\/IJMS25052957","article-title":"Extraribosomal functions of bacterial ribosomal proteins\u2014an updat","volume":"25","author":"Aseev","year":"2024","journal-title":"Int. J. Mol. Sci."},{"key":"B4","doi-asserted-by":"publisher","first-page":"3407","DOI":"10.1021\/acsinfecdis.5c00664","article-title":"Mechanistic insights into the role of extracellular vesicles at the bacterial\u2013host interface","volume":"11","author":"Battista","year":"2025","journal-title":"ACS Infect. Dis."},{"key":"B5","doi-asserted-by":"publisher","first-page":"2400","DOI":"10.1016\/J.MICINF.2006.05.001","article-title":"Purification of outer membrane vesicles from Pseudomonas aeruginosa and their activation of an IL-8 response","volume":"8","author":"Bauman","year":"2006","journal-title":"Microbes Infect."},{"key":"B6","doi-asserted-by":"publisher","DOI":"10.1186\/1471-2180-9-26","article-title":"Pseudomonas aeruginosa vesicles associate with and are internalized by human lung epithelial cells","volume":"9","author":"Bauman","year":"2009","journal-title":"BMC Microbiol."},{"key":"B7","doi-asserted-by":"publisher","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":"2021","journal-title":"Microb. Cell."},{"key":"B8","doi-asserted-by":"publisher","DOI":"10.1128\/msphere.00676-21","article-title":"Temperature influences the composition and cytotoxicity of extracellular vesicles in Staphylococcus aureus","volume":"6","author":"Briaud","year":"2021","journal-title":"mSphere"},{"key":"B9","doi-asserted-by":"publisher","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":"B10","doi-asserted-by":"publisher","DOI":"10.3390\/biology13020109","article-title":"Bacterial biofilm in chronic wounds and possible therapeutic approaches","volume":"13","author":"Cavallo","year":"2024","journal-title":"Biol. (Basel)."},{"key":"B11","doi-asserted-by":"publisher","DOI":"10.1128\/microbiolspec.psib-0001-2018","article-title":"Outer membrane vesicle-host cell interactions","volume":"7","author":"Cecil","year":"2019","journal-title":"Microbiol. Spectr."},{"key":"B12","doi-asserted-by":"publisher","first-page":"1606","DOI":"10.1038\/s41467-021-21748-6","article-title":"Biofilm formation in the lung contributes to virulence and drug tolerance of Mycobacterium tuberculosis","volume":"12","author":"Chakraborty","year":"2021","journal-title":"Nat. Commun."},{"key":"B13","doi-asserted-by":"publisher","DOI":"10.3389\/FMICB.2020.619038","article-title":"The ribosome as a switchboard for bacterial stress response","volume":"11","author":"Cheng-Guang","year":"2021","journal-title":"Front. Microbiol."},{"key":"B14","doi-asserted-by":"publisher","first-page":"698","DOI":"10.1111\/1574-6976.12057","article-title":"Phenol-soluble modulins \u2013 critical determinants of staphylococcal virulence","volume":"38","author":"Cheung","year":"2014","journal-title":"FEMS Microbiol. Rev."},{"key":"B15","doi-asserted-by":"publisher","DOI":"10.1371\/JOURNAL.PONE.0212275","article-title":"Analysis of Pseudomonas aeruginosa biofilm membrane vesicles supports multiple mechanisms of biogenesis","volume":"14","author":"Cooke","year":"2019","journal-title":"PloS One"},{"key":"B16","doi-asserted-by":"publisher","DOI":"10.3389\/fmicb.2022.1049819","article-title":"The gene regulatory network of Staphylococcus aureus ST239-SCCmecIII strain Bmb9393 and assessment of genes associated with the biofilm in diverse backgrounds","volume":"13","author":"Costa","year":"2023","journal-title":"Front. Microbiol."},{"key":"B17","doi-asserted-by":"publisher","first-page":"4207","DOI":"10.1021\/acs.jproteome.5b00312","article-title":"Proteome profiles of outer membrane vesicles and extracellular matrix of Pseudomonas aeruginosa biofilms","volume":"14","author":"Couto","year":"2015","journal-title":"J. Proteome Res."},{"key":"B18","doi-asserted-by":"publisher","first-page":"1794","DOI":"10.1021\/pr101065j","article-title":"Andromeda: a peptide search engine integrated into the MaxQuant environment","volume":"10","author":"Cox","year":"2011","journal-title":"J. Proteome Res."},{"key":"B19","doi-asserted-by":"publisher","DOI":"10.3389\/fmolb.2021.699222","article-title":"Selective nutrient transport in bacteria: multicomponent transporter systems reign supreme","volume":"8","author":"Davies","year":"2021","journal-title":"Front. Mol. Biosci."},{"key":"B20","doi-asserted-by":"publisher","first-page":"9410","DOI":"10.1038\/s41467-024-53279-1","article-title":"Mapping bacterial extracellular vesicle research: insights, best practices and knowledge gaps","volume":"15","author":"De Langhe","year":"2024","journal-title":"Nat. Commun."},{"key":"B21","doi-asserted-by":"publisher","first-page":"1797","DOI":"10.1007\/s10096-025-05148-y","article-title":"The virulence toolkit of Staphylococcus aureus: a comprehensive review of toxin diversity, molecular mechanisms, and clinical implications","volume":"44","author":"Di Bella","year":"2025","journal-title":"Eur. J. Clin. Microbiol. Infect. Dis."},{"key":"B22","doi-asserted-by":"publisher","first-page":"207","DOI":"10.1038\/nmeth1019","article-title":"Target-decoy search strategy for increased confidence in large-scale protein identifications by mass spectrometry","volume":"4","author":"Elias","year":"2007","journal-title":"Nat. Methods"},{"key":"B23","doi-asserted-by":"publisher","first-page":"3822","DOI":"10.1128\/IAI.00433-10\/","article-title":"Naturally produced outer membrane vesicles from Pseudomonas aeruginosa elicit a potent innate immune response via combined sensing of both lipopolysaccharide and protein components","volume":"78","author":"Ellis","year":"2010","journal-title":"Infect. Immun."},{"key":"B24","doi-asserted-by":"publisher","first-page":"796","DOI":"10.1007\/s11095-010-0073-2","article-title":"Critical evaluation of nanoparticle tracking analysis (NTA) by NanoSight for the measurement of nanoparticles and protein aggregates","volume":"27","author":"Filipe","year":"2010","journal-title":"Pharm. Res."},{"key":"B25","doi-asserted-by":"publisher","DOI":"10.1128\/spectrum.00804-21","article-title":"The de novo purine biosynthesis pathway is the only commonly regulated cellular pathway during biofilm formation in TSB-based medium in Staphylococcus aureus and Enterococcus faecalis","volume":"9","author":"G\u00e9linas","year":"2021","journal-title":"Microbiol. Spectr."},{"key":"B26","doi-asserted-by":"publisher","first-page":"259","DOI":"10.1099\/jmm.0.000429","article-title":"Role and mechanism of the Hsp70 molecular chaperone machines in bacterial pathogens","volume":"66","author":"Ghazaei","year":"2017","journal-title":"J. Med. Microbiol."},{"key":"B27","doi-asserted-by":"publisher","DOI":"10.1128\/MMBR.00091-21","article-title":"DNA repair in Staphylococcus aureus","volume":"85","author":"Ha","year":"2021","journal-title":"Microbiol. Mol. Biol. Rev."},{"key":"B28","doi-asserted-by":"publisher","first-page":"947","DOI":"10.1016\/J.BBRC.2013.11.012","article-title":"Biofilm vs. planktonic bacterial mode of growth: which do human macrophages prefer","volume":"441","author":"Hern\u00e1ndez-Jim\u00e9nez","year":"2013","journal-title":"Biochem. Biophys. Res. Commun."},{"key":"B29","doi-asserted-by":"publisher","first-page":"10967","DOI":"10.1021\/acsomega.4c00043","article-title":"Neutralizing Staphylococcus aureus PAMPs that trigger cytokine release from THP-1 monocytes","volume":"9","author":"Heydarian","year":"2024","journal-title":"ACS Omega"},{"key":"B30","doi-asserted-by":"publisher","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":"B31","doi-asserted-by":"publisher","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":"B32","doi-asserted-by":"publisher","DOI":"10.1128\/spectrum.05179-22","article-title":"Planktonic and biofilm-derived Pseudomonas aeruginosa outer membrane vesicles facilitate horizontal gene transfer of plasmid DNA","volume":"11","author":"Johnston","year":"2023","journal-title":"Microbiol. Spectr."},{"key":"B33","doi-asserted-by":"publisher","DOI":"10.1128\/mmbr.00032-22","article-title":"Outer membrane vesicles: biogenesis, functions, and issues","volume":"86","author":"Juodeikis","year":"2022","journal-title":"Microbiol. Mol. Biol. Rev."},{"key":"B34","doi-asserted-by":"publisher","first-page":"747","DOI":"10.1007\/S00203-019-01643-X","article-title":"Actively detached Pseudomonas aeruginosa biofilm cell susceptibility to benzalkonium chloride and associated resistance mechanism","volume":"201","author":"Khelissa","year":"2019","journal-title":"Arch. Microbiol."},{"key":"B35","doi-asserted-by":"publisher","first-page":"101","DOI":"10.1128\/mmbr.69.1.101-123.2005","article-title":"Initiation of protein synthesis in bacteria","volume":"69","author":"Laursen","year":"2005","journal-title":"Microbiol. Mol. Biol. Rev."},{"key":"B36","doi-asserted-by":"publisher","first-page":"5425","DOI":"10.1002\/PMIC.200900338","article-title":"Gram-positive bacteria produce membrane vesicles: proteomics-based characterization of Staphylococcus aureus-derived membrane vesicles","volume":"9","author":"Lee","year":"2009","journal-title":"Proteomics"},{"key":"B37","doi-asserted-by":"publisher","DOI":"10.3389\/fcimb.2022.926758","article-title":"Virulence factors of Pseudomonas aeruginosa and antivirulence strategies to combat its drug resistance","volume":"12","author":"Liao","year":"2022","journal-title":"Front. Cell. Infect. Microbiol."},{"key":"B38","doi-asserted-by":"publisher","first-page":"115","DOI":"10.1186\/s12866-019-1485-8","article-title":"Evaluation of the interaction between polymyxin B and Pseudomonas aeruginosa biofilm and planktonic cells: reactive oxygen species induction and zeta potential","volume":"19","author":"Lima","year":"2019","journal-title":"BMC Microbiol."},{"key":"B39","doi-asserted-by":"publisher","first-page":"R17","DOI":"10.1186\/cc13697","article-title":"Pseudomonas aeruginosa serotypes in nosocomial pneumonia: prevalence and clinical outcomes","volume":"18","author":"Lu","year":"2014","journal-title":"Crit. Care"},{"key":"B40","doi-asserted-by":"publisher","first-page":"212","DOI":"10.1007\/s11274-023-03651-y","article-title":"Metabolomic profiling of bacterial biofilm: trends, challenges, and an emerging antibiofilm target","volume":"39","author":"Malviya","year":"2023","journal-title":"World J. Microbiol. Biotechnol."},{"key":"B41","doi-asserted-by":"publisher","first-page":"1182","DOI":"10.1017\/S1431927603445911","article-title":"SerialEM: a program for automated tilt series acquisition on Tecnai microscopes using prediction of specimen position","volume":"9","author":"Mastronarde","year":"2003","journal-title":"Microsc. Microanal."},{"key":"B42","doi-asserted-by":"publisher","first-page":"89","DOI":"10.1034\/j.1600-065x.2000.917309.x","article-title":"Innate immune recognition: mechanisms and pathways","volume":"173","author":"Medzhitov","year":"2000","journal-title":"Immunol. Rev."},{"key":"B43","doi-asserted-by":"publisher","first-page":"732","DOI":"10.1017\/S002217240001158X","article-title":"The estimation of the bactericidal power of the blood","volume":"38","author":"Miles","year":"1938","journal-title":"J. Hyg. (Lond.)"},{"key":"B44","doi-asserted-by":"publisher","first-page":"239","DOI":"10.3934\/microbiol.2022019","article-title":"Biofilms: formation, drug resistance and alternatives to conventional approaches","volume":"8","author":"Mirghani","year":"2022","journal-title":"AIMS Microbiol."},{"key":"B45","doi-asserted-by":"publisher","first-page":"142","DOI":"10.1111\/MMI.14606","article-title":"Escherichia coli small heat shock protein IbpA is an aggregation-sensor that self-regulates its own expression at posttranscriptional levels","volume":"115","author":"Miwa","year":"2021","journal-title":"Mol. Microbiol."},{"key":"B46","doi-asserted-by":"publisher","DOI":"10.3389\/fmicb.2014.00464","article-title":"Antimicrobial targets localize to the extracellular vesicle-associated proteome of Pseudomonas aeruginosa grown in a biofilm","volume":"5","author":"Park","year":"2014","journal-title":"Front. Microbiol."},{"key":"B47","doi-asserted-by":"publisher","DOI":"10.3390\/antibiotics12010012","article-title":"A review of biofilm formation of Staphylococcus aureus and its regulation mechanism","volume":"12","author":"Peng","year":"2022","journal-title":"Antibiotics"},{"key":"B48","doi-asserted-by":"publisher","first-page":"D543","DOI":"10.1093\/nar\/gkae1011","article-title":"The PRIDE database at 20 years: 2025 update","volume":"53","author":"Perez-Riverol","year":"2025","journal-title":"Nucleic Acids Res."},{"key":"B49","doi-asserted-by":"publisher","DOI":"10.3390\/antibiotics10070825","article-title":"Prevalence and impact of biofilms on bloodstream and urinary tract infections: a systematic review and meta-analysis","volume":"10","author":"Pinto","year":"2021","journal-title":"Antibiotics"},{"key":"B50","doi-asserted-by":"publisher","DOI":"10.1128\/mbio.00623-19","article-title":"Metabolic remodeling during biofilm development of Bacillus subtilis","volume":"10","author":"Pisithkul","year":"2019","journal-title":"mBio"},{"key":"B51","doi-asserted-by":"publisher","first-page":"290","DOI":"10.1038\/nmeth.4169","article-title":"Cryosparc: algorithms for rapid unsupervised cryo-EM structure determination","volume":"14","author":"Punjani","year":"2017","journal-title":"Nat. Methods"},{"key":"B52","doi-asserted-by":"publisher","first-page":"199","DOI":"10.1038\/s41392-022-01056-1","article-title":"Pseudomonas aeruginosa: pathogenesis, virulence factors, antibiotic resistance, interaction with host, technology advances and emerging therapeutics","volume":"7","author":"Qin","year":"2022","journal-title":"Signal. Transduction Targeting Ther."},{"key":"B53","doi-asserted-by":"publisher","DOI":"10.3390\/ijms23126415","article-title":"Proteome of Staphylococcus aureus biofilm changes significantly with aging","volume":"23","author":"Rahman","year":"2022","journal-title":"Int. J. Mol. Sci."},{"key":"B54","doi-asserted-by":"publisher","first-page":"1230","DOI":"10.1111\/MMI.12255","article-title":"Paralogous chemoreceptors mediate chemotaxis towards protein amino acids and the non-protein amino acid gamma-aminobutyrate (GABA)","volume":"88","author":"Rico-Jim\u00e9nez","year":"2013","journal-title":"Mol. Microbiol."},{"key":"B55","doi-asserted-by":"publisher","DOI":"10.1016\/j.colsurfb.2023.113249","article-title":"Comparative electrokinetic properties of extracellular vesicles produced by yeast and bacteria","volume":"225","author":"Rogers","year":"2023","journal-title":"Colloids Surf. B. Biointerfaces"},{"key":"B56","doi-asserted-by":"publisher","first-page":"605","DOI":"10.1038\/nrmicro3525","article-title":"Outer-membrane vesicles from Gram-negative bacteria: Biogenesis and functions","volume":"13","author":"Schwechheimer","year":"2015","journal-title":"Nat. Rev. Microbiol."},{"key":"B57","doi-asserted-by":"publisher","first-page":"157","DOI":"10.1016\/J.BJ.2020.05.006","article-title":"Role of bacterial infections in extracellular vesicles release and impact on immune response","volume":"44","author":"Spencer","year":"2020","journal-title":"Biomed. J."},{"key":"B58","doi-asserted-by":"publisher","DOI":"10.3389\/FCIMB.2014.00171","article-title":"Protein-based biofilm matrices in Staphylococci","volume":"4","author":"Speziale","year":"2014","journal-title":"Front. Cell. Infect. Microbiol."},{"key":"B59","doi-asserted-by":"publisher","first-page":"6746","DOI":"10.1128\/JB.183.23.6746-6751.2001","article-title":"Biofilms and planktonic cells of Pseudomonas aeruginosa have similar resistance to killing by antimicrobials","volume":"183","author":"Spoering","year":"2001","journal-title":"J. Bacteriol."},{"key":"B60","doi-asserted-by":"publisher","first-page":"774","DOI":"10.1038\/s41564-019-0378-9","article-title":"Pseudomonas aeruginosa orchestrates twitching motility by sequential control of type IV pili movements","volume":"4","author":"Tal\u00e0","year":"2019","journal-title":"Nat. Microbiol."},{"key":"B61","doi-asserted-by":"publisher","first-page":"8467","DOI":"10.1038\/s41598-020-64952-y","article-title":"Extracellular vesicles produced by human and animal Staphylococcus aureus strains share a highly conserved core proteome","volume":"10","author":"Tartaglia","year":"2020","journal-title":"Sci. Rep."},{"key":"B62","doi-asserted-by":"publisher","DOI":"10.3389\/fmicb.2017.00571\/","article-title":"Interaction of bacterial membrane vesicles with specific species and their potential for delivery to target cells","volume":"8","author":"Tashiro","year":"2017","journal-title":"Front. Microbiol."},{"key":"B63","doi-asserted-by":"publisher","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":"B64","doi-asserted-by":"publisher","DOI":"10.1128\/mbio.00543-18","article-title":"A biofilm matrix-associated protease inhibitor protects Pseudomonas aeruginosa from proteolytic attack","volume":"9","author":"Tseng","year":"2018","journal-title":"mBio"},{"key":"B65","doi-asserted-by":"publisher","first-page":"2301","DOI":"10.1038\/NPROT.2016.136","article-title":"The MaxQuant computational platform for mass spectrometry-based shotgun proteomics","volume":"11","author":"Tyanova","year":"2016","journal-title":"Nat. Protoc."},{"key":"B66","doi-asserted-by":"publisher","DOI":"10.1128\/mbio.03571-22","article-title":"Contribution of extracellular membrane vesicles to the secretome of Staphylococcus aureus","volume":"14","author":"Uppu","year":"2023","journal-title":"mBio"},{"key":"B67","doi-asserted-by":"publisher","first-page":"1106","DOI":"10.1016\/j.cell.2016.04.015","article-title":"Bacterial outer membrane vesicles mediate cytosolic localization of LPS and caspase-11 activation","volume":"165","author":"Vanaja","year":"2016","journal-title":"Cell."},{"key":"B68","doi-asserted-by":"publisher","first-page":"3212","DOI":"10.1039\/d2tb02700c","article-title":"Imitating the microenvironment of native biofilms using nanofibrous scaffolds to emulate chronic wound infections","volume":"11","author":"W\u00e4chter","year":"2023","journal-title":"J. Mater. Chem. B"},{"key":"B69","doi-asserted-by":"publisher","first-page":"3174","DOI":"10.1073\/pnas.1915829117","article-title":"Orchestration of human macrophage NLRP3 inflammasome activation by Staphylococcus aureus extracellular vesicles","volume":"117","author":"Wang","year":"2020","journal-title":"Proc. Natl. Acad. Sci. U.S.A."},{"key":"B70","doi-asserted-by":"publisher","first-page":"309","DOI":"10.1111\/J.1348-0421.2009.00130.X","article-title":"Characterization of the orf1-tolQRA operon in Pseudomonas aeruginosa","volume":"53","author":"Wei","year":"2009","journal-title":"Microbiol. Immunol."},{"key":"B71","doi-asserted-by":"publisher","first-page":"1085","DOI":"10.1111\/mmi.14479","article-title":"Identification of the main glutamine and glutamate transporters in Staphylococcus aureus and their impact on c-di-AMP production","volume":"113","author":"Zeden","year":"2020","journal-title":"Mol. Microbiol."},{"key":"B72","doi-asserted-by":"publisher","first-page":"249","DOI":"10.1080\/1061186X.2020.1824235","article-title":"Iron metabolism in Pseudomonas aeruginosa biofilm and the involved iron-targeted anti-biofilm strategies","volume":"29","author":"Zhang","year":"2021","journal-title":"J. Drug Targeting"},{"key":"B73","doi-asserted-by":"publisher","DOI":"10.3389\/fcimb.2015.00040","article-title":"Extracellular matrix-associated proteins form an integral and dynamic system during Pseudomonas aeruginosa biofilm development","volume":"5","author":"Zhang","year":"2015","journal-title":"Front. Cell. Infect. Microbiol."}],"container-title":["Frontiers in Cellular and Infection Microbiology"],"original-title":[],"link":[{"URL":"https:\/\/www.frontiersin.org\/articles\/10.3389\/fcimb.2026.1881620\/full","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,10,1]],"date-time":"2026-10-01T04:42:37Z","timestamp":1790829757000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.frontiersin.org\/articles\/10.3389\/fcimb.2026.1881620\/full"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,10,1]]},"references-count":73,"alternative-id":["10.3389\/fcimb.2026.1881620"],"URL":"https:\/\/doi.org\/10.3389\/fcimb.2026.1881620","relation":{},"ISSN":["2235-2988"],"issn-type":[{"value":"2235-2988","type":"electronic"}],"subject":[],"published":{"date-parts":[[2026,10,1]]},"article-number":"1881620"}}