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Throughout the lifecycle of Vibrio cholerae, beneficial and detrimental activities of the T6SS have been described, but its role in biofilm-associated interactions remains poorly understood. Here, we show that biofilm formation is accompanied by significant repression of T6SS genes. Expression analyses across multiple V. cholerae isolates revealed consistent downregulation of the major T6SS gene cluster in biofilm-derived cells relative to planktonic cultures. Consistent with this biofilm-dependent repression, a loss of the T6SS did not affect V. cholerae biofilm formation. Notably, biofilm-derived V. cholerae wild-type isolates and their isogenic T6SS-deletion mutants displayed comparable resistance to attacks by T6SS-reactive Pseudomonas aeruginosa. These findings suggest that T6SS repression during biofilm growth provides a fitness advantage for V. cholerae by limiting susceptibility to T6SS-mediated counterattacks from neighboring heterologous T6SS+ predatory species. Our results, therefore, identify conditional T6SS silencing as a potential adaptive strategy that promotes survival within multispecies microbial communities, particularly in environments containing bacteria with highly active or more potent T6SS machineries.<\/jats:p>","DOI":"10.3390\/microorganisms14092024","type":"journal-article","created":{"date-parts":[[2026,9,11]],"date-time":"2026-09-11T10:51:37Z","timestamp":1789123897000},"page":"2024","update-policy":"https:\/\/doi.org\/10.3390\/mdpi_crossmark_policy","source":"Crossref","is-referenced-by-count":0,"title":["Repression of the Type VI Secretion System (T6SS) During Biofilm Formation Provides a Fitness Advantage for Vibrio cholerae"],"prefix":"10.3390","volume":"14","author":[{"ORCID":"https:\/\/orcid.org\/0009-0005-5843-3831","authenticated-orcid":false,"given":"Stephan P.","family":"Ebenberger","sequence":"first","affiliation":[{"name":"Institute of Molecular Biosciences, University of Graz, Humboldtstrasse 50, 8010 Graz, Austria"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-9314-5193","authenticated-orcid":false,"given":"Joao P.","family":"Pombo","sequence":"additional","affiliation":[{"name":"Institute of Molecular Biosciences, University of Graz, Humboldtstrasse 50, 8010 Graz, Austria"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0009-0005-6416-2548","authenticated-orcid":false,"given":"Alexander","family":"Rechberger","sequence":"additional","affiliation":[{"name":"Institute of Molecular Biosciences, University of Graz, Humboldtstrasse 50, 8010 Graz, Austria"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Dominik","family":"Fleischhacker","sequence":"additional","affiliation":[{"name":"Institute of Molecular Biosciences, University of Graz, Humboldtstrasse 50, 8010 Graz, Austria"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-7842-0177","authenticated-orcid":false,"given":"Stefan","family":"Schild","sequence":"additional","affiliation":[{"name":"Institute of Molecular Biosciences, University of Graz, Humboldtstrasse 50, 8010 Graz, Austria"},{"name":"BioTechMed Graz, Austria"},{"name":"Field of Excellence Biohealth, University of Graz, 8010 Graz, Austria"},{"name":"Austrian Agency for Health and Food Safety (AGES), Institute for Medical Microbiology and Hygiene, Beethovenstra\u00dfe 6, 8010 Graz, Austria"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"1968","published-online":{"date-parts":[[2026,9,11]]},"reference":[{"key":"ref_1","doi-asserted-by":"crossref","first-page":"267","DOI":"10.1016\/j.tim.2016.12.003","article-title":"Rules of Engagement: The Type VI Secretion System in Vibrio cholerae","volume":"25","author":"Joshi","year":"2017","journal-title":"Trends Microbiol."},{"key":"ref_2","doi-asserted-by":"crossref","unstructured":"Montero, D.A., Vidal, R.M., Velasco, J., George, S., Lucero, Y., G\u00f3mez, L.A., Carre\u00f1o, L.J., Garc\u00eda-Betancourt, R., and O\u2019Ryan, M. (2023). Vibrio cholerae, classification, pathogenesis, immune response, and trends in vaccine development. Front. Med., 10.","DOI":"10.3389\/fmed.2023.1155751"},{"key":"ref_3","doi-asserted-by":"crossref","unstructured":"Boyer, F., Fichant, G., Berthod, J., Vandenbrouck, Y., and Attree, I. (2009). Dissecting the bacterial type VI secretion system by a genome wide in silico analysis: What can be learned from available microbial genomic resources?. BMC Genom., 10.","DOI":"10.1186\/1471-2164-10-104"},{"key":"ref_4","doi-asserted-by":"crossref","first-page":"309","DOI":"10.1007\/s10529-023-03354-2","article-title":"Bacterial type VI secretion system (T6SS): An evolved molecular weapon with diverse functionality","volume":"45","author":"Singh","year":"2023","journal-title":"Biotechnol. Lett."},{"key":"ref_5","doi-asserted-by":"crossref","first-page":"182","DOI":"10.1038\/nature10846","article-title":"Type VI secretion requires a dynamic contractile phage tail-like structure","volume":"483","author":"Basler","year":"2012","journal-title":"Nature"},{"key":"ref_6","doi-asserted-by":"crossref","first-page":"15508","DOI":"10.1073\/pnas.0706532104","article-title":"Type VI secretion system translocates a phage tail spike-like protein into target cells where it cross-links actin","volume":"104","author":"Pukatzki","year":"2007","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_7","doi-asserted-by":"crossref","unstructured":"Felisberto-Rodrigues, C., Durand, E., Aschtgen, M.-S., Blangy, S., Ortiz-Lombardia, M., Douzi, B., Cambillau, C., and Cascales, E. (2011). Towards a structural comprehension of bacterial type VI secretion systems: Characterization of the TssJ-TssM complex of an Escherichia coli pathovar. PLoS Pathog., 7.","DOI":"10.1371\/journal.ppat.1002386"},{"key":"ref_8","doi-asserted-by":"crossref","first-page":"1600262","DOI":"10.1002\/bies.201600262","article-title":"TssA: The cap protein of the Type VI secretion system tail","volume":"39","author":"Zoued","year":"2017","journal-title":"BioEssays"},{"key":"ref_9","doi-asserted-by":"crossref","first-page":"1613","DOI":"10.15252\/embj.201694024","article-title":"TssA forms a gp6-like ring attached to the type VI secretion sheath","volume":"35","author":"Planamente","year":"2016","journal-title":"EMBO J."},{"key":"ref_10","doi-asserted-by":"crossref","unstructured":"Brunet, Y.R., Zoued, A., Boyer, F., Douzi, B., and Cascales, E. (2015). The Type VI Secretion TssEFGK-VgrG Phage-Like Baseplate Is Recruited to the TssJLM Membrane Complex via Multiple Contacts and Serves As Assembly Platform for Tail Tube\/Sheath Polymerization. PLoS Genet., 11.","DOI":"10.1371\/journal.pgen.1005545"},{"key":"ref_11","doi-asserted-by":"crossref","first-page":"20","DOI":"10.1016\/j.celrep.2014.05.034","article-title":"Structure of the VipA\/B type VI secretion complex suggests a contraction-state-specific recycling mechanism","volume":"8","author":"Kube","year":"2014","journal-title":"Cell Rep."},{"key":"ref_12","doi-asserted-by":"crossref","unstructured":"Br\u00f6ms, J.E., Ishikawa, T., Wai, S.N., and Sj\u00f6stedt, A. (2013). A functional VipA-VipB interaction is required for the type VI secretion system activity of Vibrio cholerae O1 strain A1552. BMC Microbiol., 13.","DOI":"10.1186\/1471-2180-13-96"},{"key":"ref_13","doi-asserted-by":"crossref","first-page":"1203","DOI":"10.1099\/mic.0.000842","article-title":"PAAR proteins act as the \u2018sorting hat\u2019 of the type VI secretion system","volume":"165","author":"Wood","year":"2019","journal-title":"Microbiology"},{"key":"ref_14","doi-asserted-by":"crossref","unstructured":"Cianfanelli, F.R., Alcoforado Diniz, J., Guo, M., Cesare, V.d., Trost, M., and Coulthurst, S.J. (2016). VgrG and PAAR Proteins Define Distinct Versions of a Functional Type VI Secretion System. PLoS Pathog., 12.","DOI":"10.1371\/journal.ppat.1005735"},{"key":"ref_15","doi-asserted-by":"crossref","first-page":"350","DOI":"10.1038\/nature12453","article-title":"PAAR-repeat proteins sharpen and diversify the type VI secretion system spike","volume":"500","author":"Shneider","year":"2013","journal-title":"Nature"},{"key":"ref_16","doi-asserted-by":"crossref","first-page":"1528","DOI":"10.1073\/pnas.0510322103","article-title":"Identification of a conserved bacterial protein secretion system in Vibrio cholerae using the Dictyostelium host model system","volume":"103","author":"Pukatzki","year":"2006","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_17","doi-asserted-by":"crossref","first-page":"315","DOI":"10.1038\/emboj.2008.269","article-title":"Remodelling of VipA\/VipB tubules by ClpV-mediated threading is crucial for type VI protein secretion","volume":"28","author":"Pietrosiuk","year":"2009","journal-title":"EMBO J."},{"key":"ref_18","doi-asserted-by":"crossref","first-page":"4112","DOI":"10.1111\/1462-2920.14976","article-title":"The Vibrio cholerae type VI secretion system: Toxins, regulators and consequences","volume":"22","author":"Crisan","year":"2020","journal-title":"Environ. Microbiol."},{"key":"ref_19","doi-asserted-by":"crossref","unstructured":"Zheng, J., Ho, B., and Mekalanos, J.J. (2011). Genetic analysis of anti-amoebae and anti-bacterial activities of the type VI secretion system in Vibrio cholerae. PLoS ONE, 6.","DOI":"10.1371\/journal.pone.0023876"},{"key":"ref_20","doi-asserted-by":"crossref","first-page":"e00075","DOI":"10.1128\/mBio.00075-15","article-title":"Secretome analysis of Vibrio cholerae type VI secretion system reveals a new effector-immunity pair","volume":"6","author":"Altindis","year":"2015","journal-title":"mBio"},{"key":"ref_21","doi-asserted-by":"crossref","unstructured":"Miyata, S.T., Unterweger, D., Rudko, S.P., and Pukatzki, S. (2013). Dual expression profile of type VI secretion system immunity genes protects pandemic Vibrio cholerae. PLoS Pathog., 9.","DOI":"10.1371\/journal.ppat.1003752"},{"key":"ref_22","doi-asserted-by":"crossref","first-page":"597","DOI":"10.1038\/nrmicro1461","article-title":"Social evolution theory for microorganisms","volume":"4","author":"West","year":"2006","journal-title":"Nat. Rev. Microbiol."},{"key":"ref_23","doi-asserted-by":"crossref","first-page":"e48212","DOI":"10.7554\/eLife.48212","article-title":"Neighbor predation linked to natural competence fosters the transfer of large genomic regions in Vibrio cholerae","volume":"8","author":"Matthey","year":"2019","journal-title":"eLife"},{"key":"ref_24","doi-asserted-by":"crossref","first-page":"163","DOI":"10.1186\/s13059-019-1765-5","article-title":"Analysis of Vibrio cholerae genomes identifies new type VI secretion system gene clusters","volume":"20","author":"Crisan","year":"2019","journal-title":"Genome Biol."},{"key":"ref_25","doi-asserted-by":"crossref","first-page":"R545","DOI":"10.1016\/j.cub.2019.03.068","article-title":"Drivers of Spatial Structure in Social Microbial Communities","volume":"29","author":"Yanni","year":"2019","journal-title":"Curr. Biol."},{"key":"ref_26","doi-asserted-by":"crossref","first-page":"210","DOI":"10.1126\/science.aap8775","article-title":"Antagonism toward the intestinal microbiota and its effect on Vibrio cholerae virulence","volume":"359","author":"Zhao","year":"2018","journal-title":"Science"},{"key":"ref_27","doi-asserted-by":"crossref","first-page":"14371","DOI":"10.1038\/ncomms14371","article-title":"Killing by Type VI secretion drives genetic phase separation and correlates with increased cooperation","volume":"8","author":"McNally","year":"2017","journal-title":"Nat. Commun."},{"key":"ref_28","doi-asserted-by":"crossref","first-page":"63","DOI":"10.1126\/science.1260064","article-title":"The type VI secretion system of Vibrio cholerae fosters horizontal gene transfer","volume":"347","author":"Borgeaud","year":"2015","journal-title":"Science"},{"key":"ref_29","doi-asserted-by":"crossref","first-page":"2941","DOI":"10.1128\/IAI.01266-10","article-title":"Vibrio cholerae requires the type VI secretion system virulence factor VasX to kill Dictyostelium discoideum","volume":"79","author":"Miyata","year":"2011","journal-title":"Infect. Immun."},{"key":"ref_30","doi-asserted-by":"crossref","first-page":"652","DOI":"10.1016\/j.chom.2013.11.001","article-title":"Tn-Seq analysis of Vibrio cholerae intestinal colonization reveals a role for T6SS-mediated antibacterial activity in the host","volume":"14","author":"Fu","year":"2013","journal-title":"Cell Host Microbe"},{"key":"ref_31","doi-asserted-by":"crossref","first-page":"7099","DOI":"10.1073\/pnas.1802165115","article-title":"Commensal pathogen competition impacts host viability","volume":"115","author":"Fast","year":"2018","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_32","doi-asserted-by":"crossref","first-page":"E3779","DOI":"10.1073\/pnas.1720133115","article-title":"The Vibrio cholerae type VI secretion system can modulate host intestinal mechanics to displace gut bacterial symbionts","volume":"115","author":"Logan","year":"2018","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_33","doi-asserted-by":"crossref","first-page":"2623","DOI":"10.1073\/pnas.1222783110","article-title":"Identification of T6SS-dependent effector and immunity proteins by Tn-seq in Vibrio cholerae","volume":"110","author":"Dong","year":"2013","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_34","doi-asserted-by":"crossref","first-page":"21128","DOI":"10.1073\/pnas.1014998107","article-title":"Quorum sensing and a global regulator TsrA control expression of type VI secretion and virulence in Vibrio cholerae","volume":"107","author":"Zheng","year":"2010","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_35","doi-asserted-by":"crossref","first-page":"4365","DOI":"10.1073\/pnas.0915156107","article-title":"In vivo actin cross-linking induced by Vibrio cholerae type VI secretion system is associated with intestinal inflammation","volume":"107","author":"Ma","year":"2010","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_36","doi-asserted-by":"crossref","first-page":"5395","DOI":"10.1038\/s41467-020-19017-z","article-title":"The evolution of tit-for-tat in bacteria via the type VI secretion system","volume":"11","author":"Smith","year":"2020","journal-title":"Nat. Commun."},{"key":"ref_37","doi-asserted-by":"crossref","first-page":"884","DOI":"10.1016\/j.cell.2013.01.042","article-title":"Tit-for-tat: Type VI secretion system counterattack during bacterial cell-cell interactions","volume":"152","author":"Basler","year":"2013","journal-title":"Cell"},{"key":"ref_38","doi-asserted-by":"crossref","unstructured":"Pombo, J.P., Ebenberger, S.P., M\u00fcller, A.M., Wolinski, H., and Schild, S. (2022). Impact of Gene Repression on Biofilm Formation of Vibrio cholerae. Front. Microbiol., 13.","DOI":"10.3389\/fmicb.2022.912297"},{"key":"ref_39","doi-asserted-by":"crossref","unstructured":"Soria-Bustos, J., Ares, M.A., G\u00f3mez-Aldapa, C.A., Gonz\u00e1lez-Y-Merchand, J.A., Gir\u00f3n, J.A., and La Cruz, M.A.d. (2020). Two Type VI Secretion Systems of Enterobacter cloacae Are Required for Bacterial Competition, Cell Adherence, and Intestinal Colonization. Front. Microbiol., 11.","DOI":"10.3389\/fmicb.2020.560488"},{"key":"ref_40","doi-asserted-by":"crossref","first-page":"12","DOI":"10.1006\/plas.1999.1433","article-title":"Genetic system for reversible integration of DNA constructs and lacZ gene fusions into the Escherichia coli chromosome","volume":"43","author":"Platt","year":"2000","journal-title":"Plasmid"},{"key":"ref_41","doi-asserted-by":"crossref","first-page":"784","DOI":"10.1038\/nbt1183-784","article-title":"A Broad Host Range Mobilization System for In Vivo Genetic Engineering: Transposon Mutagenesis in Gram Negative Bacteria","volume":"1","author":"Simon","year":"1983","journal-title":"Nat. Biotechnol."},{"key":"ref_42","doi-asserted-by":"crossref","first-page":"1288","DOI":"10.1128\/jb.171.3.1288-1293.1989","article-title":"Identification of toxS, a regulatory gene whose product enhances toxR-mediated activation of the cholera toxin promoter","volume":"171","author":"Miller","year":"1989","journal-title":"J. Bacteriol."},{"key":"ref_43","doi-asserted-by":"crossref","first-page":"1655","DOI":"10.1099\/00221287-148-6-1655","article-title":"Evolutionary and functional analyses of variants of the toxin-coregulated pilus protein TcpA from toxigenic Vibrio cholerae non-O1\/non-O139 serogroup isolates","volume":"148","author":"Boyd","year":"2002","journal-title":"Microbiology"},{"key":"ref_44","doi-asserted-by":"crossref","first-page":"1029","DOI":"10.1093\/infdis\/169.5.1029","article-title":"Spread of Vibrio cholerae O139 Bengal in India","volume":"169","author":"Nair","year":"1994","journal-title":"J. Infect. Dis."},{"key":"ref_45","doi-asserted-by":"crossref","first-page":"224","DOI":"10.1128\/iai.42.1.224-230.1983","article-title":"Molecular epidemiological studies of United States Gulf Coast Vibrio cholerae strains: Integration site of mutator vibriophage VcA-3","volume":"42","author":"Goldberg","year":"1983","journal-title":"Infect. Immun."},{"key":"ref_46","doi-asserted-by":"crossref","first-page":"225","DOI":"10.1016\/j.chom.2019.12.002","article-title":"Outer membrane vesiculation facilitates surface exchange and in vivo adaptation of Vibrio cholerae","volume":"27","author":"Zingl","year":"2020","journal-title":"Cell Host Microbe"},{"key":"ref_47","first-page":"572","article-title":"Genetic recombination in Pseudomonas aeruginosa","volume":"13","author":"Holloway","year":"1955","journal-title":"J. Gen. Microbiol."},{"key":"ref_48","doi-asserted-by":"crossref","first-page":"e0111523","DOI":"10.1128\/spectrum.01115-23","article-title":"Characterization of the Inflammatory Response Evoked by Bacterial Membrane Vesicles in Intestinal Cells Reveals an RIPK2-Dependent Activation by Enterotoxigenic Escherichia coli Vesicles","volume":"11","author":"Thapa","year":"2023","journal-title":"Microbiol. Spectr."},{"key":"ref_49","doi-asserted-by":"crossref","first-page":"4310","DOI":"10.1128\/iai.59.12.4310-4317.1991","article-title":"Construction of an eae deletion mutant of enteropathogenic Escherichia coli by using a positive-selection suicide vector","volume":"59","author":"Donnenberg","year":"1991","journal-title":"Infect. Immun."},{"key":"ref_50","doi-asserted-by":"crossref","first-page":"7027","DOI":"10.1128\/JB.00948-09","article-title":"A novel regulatory protein involved in motility of Vibrio cholerae","volume":"191","author":"Moisi","year":"2009","journal-title":"J. Bacteriol."},{"key":"ref_51","doi-asserted-by":"crossref","first-page":"61","DOI":"10.1016\/0378-1119(89)90359-4","article-title":"Engineering hybrid genes without the use of restriction enzymes: Gene splicing by overlap extension","volume":"77","author":"Horton","year":"1989","journal-title":"Gene"},{"key":"ref_52","doi-asserted-by":"crossref","first-page":"1015","DOI":"10.1111\/j.1365-2958.2011.07867.x","article-title":"Extracellular nucleases and extracellular DNA play important roles in Vibrio cholerae biofilm formation","volume":"82","author":"Seper","year":"2011","journal-title":"Mol. Microbiol."},{"key":"ref_53","doi-asserted-by":"crossref","first-page":"61","DOI":"10.1016\/S0091-679X(08)61676-3","article-title":"Analysis of membrane protein topology using alkaline phosphatase and beta-galactosidase gene fusions","volume":"34","author":"Manoil","year":"1991","journal-title":"Methods Cell Biol."},{"key":"ref_54","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_55","doi-asserted-by":"crossref","first-page":"19520","DOI":"10.1073\/pnas.1012931107","article-title":"The Vibrio cholerae type VI secretion system displays antimicrobial properties","volume":"107","author":"MacIntyre","year":"2010","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_56","doi-asserted-by":"crossref","first-page":"8069","DOI":"10.1038\/s41598-018-26342-3","article-title":"Extraction and quantification of biofilm bacteria: Method optimized for urinary catheters","volume":"8","author":"Mandakhalikar","year":"2018","journal-title":"Sci. Rep."},{"key":"ref_57","doi-asserted-by":"crossref","first-page":"3560","DOI":"10.1128\/IAI.00048-10","article-title":"Growth in a biofilm induces a hyperinfectious phenotype in Vibrio cholerae","volume":"78","author":"Tamayo","year":"2010","journal-title":"Infect. Immun."},{"key":"ref_58","doi-asserted-by":"crossref","first-page":"3549","DOI":"10.1038\/ncomms4549","article-title":"The Vibrio cholerae type VI secretion system employs diverse effector modules for intraspecific competition","volume":"5","author":"Unterweger","year":"2014","journal-title":"Nat. Commun."},{"key":"ref_59","doi-asserted-by":"crossref","first-page":"e0058423","DOI":"10.1128\/msphere.00584-23","article-title":"The type VI secretion system of the emerging pathogen Stenotrophomonas maltophilia complex has antibacterial properties","volume":"8","author":"Crisan","year":"2023","journal-title":"mSphere"},{"key":"ref_60","doi-asserted-by":"crossref","first-page":"7997","DOI":"10.1073\/pnas.1808469115","article-title":"Exopolysaccharide protects Vibrio cholerae from exogenous attacks by the type 6 secretion system","volume":"115","author":"Toska","year":"2018","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_61","doi-asserted-by":"crossref","first-page":"1052","DOI":"10.1038\/s41396-023-01401-4","article-title":"Collective protection against the type VI secretion system in bacteria","volume":"17","author":"Granato","year":"2023","journal-title":"ISME J."},{"key":"ref_62","doi-asserted-by":"crossref","unstructured":"Li, X., Gu, N., Huang, T.Y., Zhong, F., and Peng, G. (2022). Pseudomonas aeruginosa: A typical biofilm forming pathogen and an emerging but underestimated pathogen in food processing. Front. Microbiol., 13.","DOI":"10.3389\/fmicb.2022.1114199"},{"key":"ref_63","doi-asserted-by":"crossref","unstructured":"Misra, T., Tare, M., and Jha, P.N. (2022). Insights Into the Dynamics and Composition of Biofilm Formed by Environmental Isolate of Enterobacter cloacae. Front. Microbiol., 13.","DOI":"10.3389\/fmicb.2022.877060"},{"key":"ref_64","doi-asserted-by":"crossref","first-page":"2294","DOI":"10.1128\/JCM.39.6.2294-2297.2001","article-title":"Biofilm formation by gram-negative bacteria on central venous catheter connectors: Effect of conditioning films in a laboratory model","volume":"39","author":"Murga","year":"2001","journal-title":"J. Clin. Microbiol."}],"container-title":["Microorganisms"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.mdpi.com\/2076-2607\/14\/9\/2024\/pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,9,11]],"date-time":"2026-09-11T11:08:34Z","timestamp":1789124914000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.mdpi.com\/2076-2607\/14\/9\/2024"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,9,11]]},"references-count":64,"journal-issue":{"issue":"9","published-online":{"date-parts":[[2026,9]]}},"alternative-id":["microorganisms14092024"],"URL":"https:\/\/doi.org\/10.3390\/microorganisms14092024","relation":{},"ISSN":["2076-2607"],"issn-type":[{"value":"2076-2607","type":"electronic"}],"subject":[],"published":{"date-parts":[[2026,9,11]]}}}