{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,10,6]],"date-time":"2026-10-06T06:06:33Z","timestamp":1791266793662,"version":"4.1.0"},"reference-count":101,"publisher":"MDPI AG","issue":"3","license":[{"start":{"date-parts":[[2016,7,5]],"date-time":"2016-07-05T00:00:00Z","timestamp":1467676800000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":["Microorganisms"],"abstract":"<jats:p>Listeria monocytogenes is a foodborne pathogen able to persist in food industry and is responsible for a severe illness called listeriosis. The ability of L. monocytogenes to persist in environments is due to its capacity to form biofilms that are a sessile community of microorganisms embedded in a self-produced matrix of extracellular polymeric substances (EPS\u2019s). In this review, we summarized recent efforts performed in order to better characterize the polymeric substances that compose the extracellular matrix (ECM) of L. monocytogenes biofilms. EPS extraction and analysis led to the identification of polysaccharides, proteins, extracellular DNA, and other molecules within the listerial ECM. All this knowledge will be useful for increasing food protection, suggesting effective strategies for the minimization of persistence of L. monocytogenes in food industry environments.<\/jats:p>","DOI":"10.3390\/microorganisms4030022","type":"journal-article","created":{"date-parts":[[2016,7,5]],"date-time":"2016-07-05T10:06:19Z","timestamp":1467713179000},"page":"22","update-policy":"https:\/\/doi.org\/10.3390\/mdpi_crossmark_policy","source":"Crossref","is-referenced-by-count":104,"title":["A Look inside the Listeria monocytogenes Biofilms Extracellular Matrix"],"prefix":"10.3390","volume":"4","author":[{"given":"Angelo","family":"Colagiorgi","sequence":"first","affiliation":[{"name":"Department of Food Science, University of Parma, Via del Taglio 10, Parma 43126, Italy"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Pierluigi","family":"Di Ciccio","sequence":"additional","affiliation":[{"name":"Department of Food Science, University of Parma, Via del Taglio 10, Parma 43126, Italy"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-2253-3647","authenticated-orcid":false,"given":"Emanuela","family":"Zanardi","sequence":"additional","affiliation":[{"name":"Department of Food Science, University of Parma, Via del Taglio 10, Parma 43126, Italy"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-7351-0381","authenticated-orcid":false,"given":"Sergio","family":"Ghidini","sequence":"additional","affiliation":[{"name":"Department of Food Science, University of Parma, Via del Taglio 10, Parma 43126, Italy"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Adriana","family":"Ianieri","sequence":"additional","affiliation":[{"name":"Department of Food Science, University of Parma, Via del Taglio 10, Parma 43126, Italy"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"1968","published-online":{"date-parts":[[2016,7,5]]},"reference":[{"key":"ref_1","doi-asserted-by":"crossref","first-page":"319","DOI":"10.1089\/fpd.2006.3.319","article-title":"Listeria monocytogenes in multiple habitats and host populations: Review of available data for mathematical modeling","volume":"3","author":"Ivanek","year":"2006","journal-title":"Foodborne Pathog. Dis."},{"key":"ref_2","unstructured":"Doyle, M.P., Buechat, L.R., and Montville, T.J. (1997). Food Microbiology\u2014Fundamentals and Frontiers, American Society for Microbiology (ASM) Press."},{"key":"ref_3","doi-asserted-by":"crossref","first-page":"6721","DOI":"10.1128\/JB.186.20.6721-6727.2004","article-title":"Flagellin from Listeria monocytogenes is glycosylated with beta-O-linked N-acetylglucosamine","volume":"186","author":"Schirm","year":"2004","journal-title":"J. Bacteriol."},{"key":"ref_4","doi-asserted-by":"crossref","first-page":"203","DOI":"10.1056\/NEJM198301273080407","article-title":"Epidemic listeriosis\u2014Evidence for transmission by food","volume":"308","author":"Schlech","year":"1983","journal-title":"N. Engl. J. Med."},{"key":"ref_5","doi-asserted-by":"crossref","first-page":"7","DOI":"10.3201\/eid1701.P11101","article-title":"Foodborne illness acquired in the united states-major pathogens","volume":"17","author":"Scallan","year":"2011","journal-title":"Emerg. Infect. Dis."},{"key":"ref_6","unstructured":"ECDC (2014). Food- and Waterborne Diseases and Zoonoses, ECDC."},{"key":"ref_7","doi-asserted-by":"crossref","unstructured":"EFSA (2015). The european union summary report on trends and sources of zoonoses, zoonotic agents and food-borne outbreaks in 2013. EFSA J., 13, 165.","DOI":"10.2903\/j.efsa.2015.3991"},{"key":"ref_8","unstructured":"CDC (2014). Incidence and trends of infection with pathogens transmitted commonly through food-foodborne diseases active surveillance network, 10 US sites, 2006\u20132013. Morb. Mortal. Wkly. Rep., 63, 328\u2013332."},{"key":"ref_9","doi-asserted-by":"crossref","first-page":"1073","DOI":"10.1016\/S1473-3099(14)70870-9","article-title":"The global burden of listeriosis: A systematic review and meta-analysis","volume":"14","author":"Devleesschauwer","year":"2014","journal-title":"Lancet Infect. Dis."},{"key":"ref_10","doi-asserted-by":"crossref","unstructured":"EFSA (2015). The european union summary report on trends and sources of zoonoses, zoonotic agents and food-borne outbreaks in 2014. EFSA J., 13, 191.","DOI":"10.2903\/j.efsa.2015.4329"},{"key":"ref_11","unstructured":"CDC (2014). Foodborne Diseases Active Surveillance Network (Foodnet): Foodnet Surveillance Report for 2012 (Final Report)."},{"key":"ref_12","doi-asserted-by":"crossref","first-page":"S396","DOI":"10.1093\/cid\/cis268","article-title":"Invasive listeriosis in the foodborne diseases active surveillance network (foodnet), 2004\u20132009: Further targeted prevention needed for higher-risk groups","volume":"54","author":"Silk","year":"2012","journal-title":"Clin. Infect. Dis. Off. Publ. Infect. Dis. Soc. Am."},{"key":"ref_13","unstructured":"Painter, J., and Slutsker, L. (2007). Listeria, Listeriosis and Food Safety, CRC press. [3rd ed.]."},{"key":"ref_14","doi-asserted-by":"crossref","first-page":"1503","DOI":"10.1017\/S0950268810000294","article-title":"Pregnancy-associated listeriosis","volume":"138","author":"Jackson","year":"2010","journal-title":"Epidemiol. Infect."},{"key":"ref_15","doi-asserted-by":"crossref","first-page":"693","DOI":"10.1093\/infdis\/170.3.693","article-title":"A point-source foodborne listeriosis outbreak: Documented incubation period and possible mild illness","volume":"170","author":"Riedo","year":"1994","journal-title":"J. Infect. Dis."},{"key":"ref_16","doi-asserted-by":"crossref","first-page":"100","DOI":"10.1056\/NEJM199701093360204","article-title":"An outbreak of gastroenteritis and fever due to Listeria monocytogenes in milk","volume":"336","author":"Dalton","year":"1997","journal-title":"N. Engl. J. Med."},{"key":"ref_17","doi-asserted-by":"crossref","first-page":"410","DOI":"10.1542\/pir.29.11.410","article-title":"Listeriosis","volume":"29","author":"Jacobson","year":"2008","journal-title":"Pediatr. Rev. Am. Acad. Pediatr."},{"key":"ref_18","doi-asserted-by":"crossref","unstructured":"Simjee, S. (2007). Foodborne Diseases, Humana Press.","DOI":"10.1007\/978-1-59745-501-5"},{"key":"ref_19","doi-asserted-by":"crossref","first-page":"476","DOI":"10.1128\/mr.55.3.476-511.1991","article-title":"Listeria monocytogenes, a food-borne pathogen","volume":"55","author":"Farber","year":"1991","journal-title":"Microbiol. Rev."},{"key":"ref_20","doi-asserted-by":"crossref","first-page":"584","DOI":"10.1128\/CMR.14.3.584-640.2001","article-title":"Listeria pathogenesis and molecular virulence determinants","volume":"14","author":"Kuhn","year":"2001","journal-title":"Clin. Microbiol. Rev."},{"key":"ref_21","doi-asserted-by":"crossref","first-page":"827","DOI":"10.4315\/0362-028X-59.8.827","article-title":"Growth of Listeria monocytogenes as a biofilm on various food-processing surfaces","volume":"59","author":"Blackman","year":"1996","journal-title":"J. Food Prot."},{"key":"ref_22","doi-asserted-by":"crossref","first-page":"1552","DOI":"10.1111\/j.1365-2672.2007.03688.x","article-title":"Influence of temperature on biofilm formation by Listeria monocytogenes on various food-contact surfaces: Relationship with motility and cell surface hydrophobicity","volume":"104","author":"Piccolomini","year":"2008","journal-title":"J. Appl. Microbiol."},{"key":"ref_23","doi-asserted-by":"crossref","first-page":"1969","DOI":"10.1128\/aem.57.7.1969-1973.1991","article-title":"Characterization of physicochemical forces involved in adhesion of Listeria monocytogenes to surfaces","volume":"57","author":"Mafu","year":"1991","journal-title":"Appl. Environ. Microbiol."},{"key":"ref_24","doi-asserted-by":"crossref","first-page":"167","DOI":"10.1128\/CMR.15.2.167-193.2002","article-title":"Biofilms: Survival mechanisms of clinically relevant microorganisms","volume":"15","author":"Donlan","year":"2002","journal-title":"Clin. Microbiol. Rev."},{"key":"ref_25","doi-asserted-by":"crossref","first-page":"623","DOI":"10.1038\/nrmicro2415","article-title":"The biofilm matrix","volume":"8","author":"Flemming","year":"2010","journal-title":"Nat. Rev. Microbiol."},{"key":"ref_26","first-page":"222","article-title":"The biofilm matrix\u2014An immobilized but dynamic microbial environment","volume":"9","author":"Sutherland","year":"2001","journal-title":"TIM"},{"key":"ref_27","doi-asserted-by":"crossref","first-page":"1995","DOI":"10.1111\/j.1462-2920.2012.02753.x","article-title":"Synthesis of multiple Pseudomonas aeruginosa biofilm matrix exopolysaccharides is post-transcriptionally regulated","volume":"14","author":"Ma","year":"2012","journal-title":"Environ. Microbiol."},{"key":"ref_28","first-page":"231","article-title":"Bacterial biofilms, other structures seen as mainstream concepts","volume":"2","author":"Schaudinn","year":"2007","journal-title":"Microbe"},{"key":"ref_29","first-page":"207","article-title":"Staphylococcal biofilms","volume":"322","author":"Otto","year":"2008","journal-title":"Curr. Top. Microbiol. Immunol."},{"key":"ref_30","doi-asserted-by":"crossref","first-page":"236","DOI":"10.1126\/science.1222981","article-title":"Molecular architecture and assembly principles of Vibrio cholerae biofilms","volume":"337","author":"Berk","year":"2012","journal-title":"Science"},{"key":"ref_31","doi-asserted-by":"crossref","first-page":"1204","DOI":"10.4315\/0362-028X-63.9.1204","article-title":"Persistent Listeria monocytogenes strains show enhanced adherence to food contact surface after short contact times","volume":"63","author":"Lunden","year":"2000","journal-title":"J. Food Prot."},{"key":"ref_32","doi-asserted-by":"crossref","first-page":"25","DOI":"10.1111\/j.1574-6968.2002.tb11155.x","article-title":"Comparison of protein patterns of Listeria monocytogenes grown in biofilm or in planktonic mode by proteomic analysis","volume":"210","author":"Namane","year":"2002","journal-title":"FEMS Microbiol. Lett."},{"key":"ref_33","doi-asserted-by":"crossref","first-page":"95","DOI":"10.1038\/nrmicro821","article-title":"Bacterial biofilms: From the natural environment to infectious diseases","volume":"2","author":"Costerton","year":"2004","journal-title":"Nat. Rev. Microbiol."},{"key":"ref_34","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1016\/j.ijfoodmicro.2011.01.005","article-title":"Review\u2014Persistence of Listeria monocytogenes in food industry equipment and premises","volume":"145","author":"Carpentier","year":"2011","journal-title":"Int. J. Food Microbiol."},{"key":"ref_35","doi-asserted-by":"crossref","first-page":"7336","DOI":"10.1128\/AEM.69.12.7336-7342.2003","article-title":"Variation in biofilm formation among strains of Listeria monocytogenes","volume":"69","author":"Borucki","year":"2003","journal-title":"Appl. Environ. Microbiol."},{"key":"ref_36","doi-asserted-by":"crossref","first-page":"2950","DOI":"10.1128\/AEM.68.6.2950-2958.2002","article-title":"Microtiter plate assay for assessment of Listeria monocytogenes biofilm formation","volume":"68","author":"Djordjevic","year":"2002","journal-title":"Appl. Environ. Microbiol."},{"key":"ref_37","doi-asserted-by":"crossref","first-page":"282","DOI":"10.1016\/j.ijfoodmicro.2007.09.006","article-title":"Effect of temperature and growth media on the attachment of Listeria monocytogenes to stainless steel","volume":"120","author":"Mai","year":"2007","journal-title":"Int. J. Food Microbiol."},{"key":"ref_38","doi-asserted-by":"crossref","first-page":"92","DOI":"10.4315\/0362-028X-68.1.92","article-title":"Formation of biofilms by Listeria monocytogenes under various growth conditions","volume":"68","author":"Moltz","year":"2005","journal-title":"J. Food Prot."},{"key":"ref_39","doi-asserted-by":"crossref","first-page":"103","DOI":"10.1016\/S0168-1605(00)00406-2","article-title":"Comparative evaluation of adhesion and biofilm formation of different Listeria monocytogenes strains","volume":"62","author":"Chae","year":"2000","journal-title":"Int. J. Food Microbiol."},{"key":"ref_40","doi-asserted-by":"crossref","first-page":"1569","DOI":"10.4315\/0362-028X-70.7.1569","article-title":"Variable adhesion of Listeria monocytogenes isolates from food-processing facilities and clinical cases to inert surfaces","volume":"70","author":"Tresse","year":"2007","journal-title":"J. Food Prot."},{"key":"ref_41","doi-asserted-by":"crossref","first-page":"380","DOI":"10.1016\/j.fm.2006.06.006","article-title":"Assessing biofilm formation by Listeria monocytogenes strains","volume":"24","author":"Harvey","year":"2007","journal-title":"Food Microbiol."},{"key":"ref_42","doi-asserted-by":"crossref","first-page":"262","DOI":"10.1111\/j.1365-2672.2004.02296.x","article-title":"Impact of cleaning and disinfection agents on biofilm structure and on microbial transfer to a solid model food","volume":"97","author":"Midelet","year":"2004","journal-title":"J. Appl. Microbiol."},{"key":"ref_43","doi-asserted-by":"crossref","first-page":"2313","DOI":"10.1128\/AEM.72.4.2313-2321.2006","article-title":"Construction and analysis of fractional multifactorial designs to study attachment strength and transfer of Listeria monocytogenes from pure or mixed biofilms after contact with a solid model food","volume":"72","author":"Midelet","year":"2006","journal-title":"Appl. Environ. Microbiol."},{"key":"ref_44","doi-asserted-by":"crossref","first-page":"1028","DOI":"10.1139\/W10-093","article-title":"Characterization of Pseudomonas aeruginosa fatty acid profiles in biofilms and batch planktonic cultures","volume":"56","author":"Chao","year":"2010","journal-title":"Can. J. Microbiol."},{"key":"ref_45","doi-asserted-by":"crossref","first-page":"155","DOI":"10.1111\/j.1574-6968.2003.tb11512.x","article-title":"Different cellular fatty acid pattern behaviours of two strains of Listeria monocytogenes Scott A and CNL 895807 under different temperature and salinity conditions","volume":"218","author":"Chihib","year":"2003","journal-title":"FEMS Microbiol. Lett."},{"key":"ref_46","doi-asserted-by":"crossref","first-page":"9669","DOI":"10.1021\/jm300908t","article-title":"Role of the CAI-1 fatty acid tail in the Vibrio cholerae quorum sensing response","volume":"55","author":"Perez","year":"2012","journal-title":"J. Med. Chem."},{"key":"ref_47","doi-asserted-by":"crossref","first-page":"835","DOI":"10.1111\/j.1462-2920.2010.02378.x","article-title":"Molecular biology of surface colonization by Listeria monocytogenes: An additional facet of an opportunistic Gram-positive foodborne pathogen","volume":"13","author":"Renier","year":"2011","journal-title":"Environ. Microbiol."},{"key":"ref_48","doi-asserted-by":"crossref","first-page":"728","DOI":"10.1128\/AEM.68.2.728-737.2002","article-title":"Listeria monocytogenes LO28: Surface physicochemical properties and ability to form biofilms at different temperatures and growth phases","volume":"68","author":"Chavant","year":"2002","journal-title":"Appl. Environ. Microbiol."},{"key":"ref_49","doi-asserted-by":"crossref","first-page":"203","DOI":"10.1016\/S0378-1097(03)00752-3","article-title":"A three-tiered approach to differentiate Listeria monocytogenes biofilm-forming abilities","volume":"228","author":"Marsh","year":"2003","journal-title":"FEMS Microbiol. Lett."},{"key":"ref_50","doi-asserted-by":"crossref","first-page":"4491","DOI":"10.1128\/AEM.00255-08","article-title":"Listeria monocytogenes EGD-e biofilms: No mushrooms but a network of knitted chains","volume":"74","author":"Rieu","year":"2008","journal-title":"Appl. Environ. Microbiol."},{"key":"ref_51","doi-asserted-by":"crossref","first-page":"1804","DOI":"10.1128\/AEM.03173-14","article-title":"Exploring the diversity of Listeria monocytogenes biofilm architecture by high-throughput confocal laser scanning microscopy and the predominance of the honeycomb-like morphotype","volume":"81","author":"Guilbaud","year":"2015","journal-title":"Appl. Environ. Microbiol."},{"key":"ref_52","doi-asserted-by":"crossref","first-page":"64","DOI":"10.1016\/j.mimet.2010.04.006","article-title":"The biofilm architecture of sixty opportunistic pathogens deciphered using a high throughput CLSM method","volume":"82","author":"Bridier","year":"2010","journal-title":"J. Microbiol. Methods"},{"key":"ref_53","doi-asserted-by":"crossref","first-page":"5629","DOI":"10.1128\/JB.186.17.5629-5639.2004","article-title":"The enterococcus faecalis fsr two-component system controls biofilm development through production of gelatinase","volume":"186","author":"Hancock","year":"2004","journal-title":"J. Bacteriol."},{"key":"ref_54","doi-asserted-by":"crossref","first-page":"1143","DOI":"10.1007\/s11274-009-0271-4","article-title":"Development of a biofilm model for Listeria monocytogenes EGD-e","volume":"26","author":"Zameer","year":"2010","journal-title":"World J. Microbiol. Biotechnol."},{"key":"ref_55","doi-asserted-by":"crossref","unstructured":"Chen, L.H., K\u00f6seoglu, V.K., Guvener, Z.T., Myers-Morales, T., Reed, J.M., D\u2019Orazio, S.E., Miller, K.W., and Gomelsky, M. (2014). Cyclic di-GMP-dependent signaling pathways in the pathogenic firmicute Listeria monocytogenes. PLoS Pathog., 10.","DOI":"10.1371\/journal.ppat.1004301"},{"key":"ref_56","doi-asserted-by":"crossref","first-page":"909","DOI":"10.1111\/mmi.12140","article-title":"Cycles of light and dark co-ordinate reversible colony differentiation in Listeria monocytogenes","volume":"87","author":"Tiensuu","year":"2013","journal-title":"Mol. Microbiol."},{"key":"ref_57","doi-asserted-by":"crossref","first-page":"197","DOI":"10.1139\/w04-129","article-title":"Proteomic and microscopic analysis of biofilms formed by Listeria monocytogenes 568","volume":"51","author":"Hefford","year":"2005","journal-title":"Can. J. Microbiol."},{"key":"ref_58","first-page":"E2209","article-title":"Antibody to a conserved antigenic target is protective against diverse prokaryotic and eukaryotic pathogens","volume":"110","author":"Skurnik","year":"2013","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_59","doi-asserted-by":"crossref","first-page":"1120","DOI":"10.1111\/jam.12127","article-title":"Quantification of the extracellular matrix of the Listeria monocytogenes biofilms of different phylogenic lineages with optimization of culture conditions","volume":"114","author":"Combrouse","year":"2013","journal-title":"J. Appl. Microbiol."},{"key":"ref_60","doi-asserted-by":"crossref","unstructured":"Brauge, T., Sadovskaya, I., Faille, C., Benezech, T., Maes, E., Guerardel, Y., and Midelet-Bourdin, G. (2016). Teichoic acid is the major polysaccharide present in the Listeria monocytogenes biofilm matrix. FEMS Microbiol. Lett., 363.","DOI":"10.1093\/femsle\/fnv229"},{"key":"ref_61","doi-asserted-by":"crossref","first-page":"728","DOI":"10.1111\/mmi.12966","article-title":"Listeria monocytogenes exopolysaccharide: Origin, structure, biosynthetic machinery and c-di-GMP-dependent regulation","volume":"96","author":"Koseoglu","year":"2015","journal-title":"Mol. Microbiol."},{"key":"ref_62","doi-asserted-by":"crossref","first-page":"1428","DOI":"10.3389\/fmicb.2015.01428","article-title":"Dnase-sensitive and -resistant modes of biofilm formation by Listeria monocytogenes","volume":"6","author":"Zetzmann","year":"2015","journal-title":"Front. Microbiol."},{"key":"ref_63","doi-asserted-by":"crossref","first-page":"302","DOI":"10.1016\/S0076-6879(01)36597-7","article-title":"Isolation and biochemical characterization of extracellular polymeric substances from Pseudomonas aeruginosa","volume":"336","author":"Wingender","year":"2001","journal-title":"Method Enzymol."},{"key":"ref_64","doi-asserted-by":"crossref","first-page":"1749","DOI":"10.1016\/0043-1354(95)00323-1","article-title":"Extraction of extracellular polymers from activated sludge using a cation exchange resin","volume":"30","author":"Frolund","year":"1996","journal-title":"Water Res."},{"key":"ref_65","doi-asserted-by":"crossref","first-page":"915","DOI":"10.1128\/jb.116.2.915-924.1973","article-title":"Production and characterization of the slime polysaccharide of Pseudomonas aeruginosa","volume":"116","author":"Evans","year":"1973","journal-title":"J. Bacteriol."},{"key":"ref_66","doi-asserted-by":"crossref","first-page":"3007","DOI":"10.1128\/IAI.73.5.3007-3017.2005","article-title":"Extracellular carbohydrate-containing polymers of a model biofilm-producing strain, Staphylococcus epidermidis rp62a","volume":"73","author":"Sadovskaya","year":"2005","journal-title":"Infect. Immun."},{"key":"ref_67","doi-asserted-by":"crossref","unstructured":"Alonso, A.N., Perry, K.J., Regeimbal, J.M., Regan, P.M., and Higgins, D.E. (2014). Identification of Listeria monocytogenes determinants required for biofilm formation. PLoS ONE, 9.","DOI":"10.1371\/journal.pone.0113696"},{"key":"ref_68","doi-asserted-by":"crossref","first-page":"45","DOI":"10.1016\/j.micpath.2008.01.007","article-title":"Protease treatment affects both invasion ability and biofilm formation in Listeria monocytogenes","volume":"45","author":"Longhi","year":"2008","journal-title":"Microb. Pathog."},{"key":"ref_69","doi-asserted-by":"crossref","first-page":"26","DOI":"10.1016\/j.ijfoodmicro.2014.06.025","article-title":"Dnase i and proteinase k impair Listeria monocytogenes biofilm formation and induce dispersal of pre-existing biofilms","volume":"187","author":"Nguyen","year":"2014","journal-title":"Int. J. Food Microbiol."},{"key":"ref_70","doi-asserted-by":"crossref","first-page":"183","DOI":"10.1177\/039463200902200121","article-title":"Expression of internalin a and biofilm formation among Listeria monocytogenes clinical isolates","volume":"22","author":"Franciosa","year":"2009","journal-title":"Int. J. Immunopathol. Pharmacol."},{"key":"ref_71","doi-asserted-by":"crossref","first-page":"5451","DOI":"10.1128\/AEM.02419-07","article-title":"Listeria monocytogenes biofilm-associated protein (BapL) may contribute to surface attachment of L. monocytogenes but is absent from many field isolates","volume":"74","author":"Jordan","year":"2008","journal-title":"Appl. Environ. Microbiol."},{"key":"ref_72","doi-asserted-by":"crossref","first-page":"4418","DOI":"10.1128\/JB.01967-06","article-title":"Flagellar motility is critical for Listeria monocytogenes biofilm formation","volume":"189","author":"Lemon","year":"2007","journal-title":"J. Bacteriol."},{"key":"ref_73","doi-asserted-by":"crossref","first-page":"6075","DOI":"10.1128\/AEM.01592-13","article-title":"Comparison of Listeria monocytogenes exoproteomes from biofilm and planktonic state: Lmo2504, a protein associated with biofilms","volume":"79","author":"Lourenco","year":"2013","journal-title":"Appl. Environ. Microbiol."},{"key":"ref_74","doi-asserted-by":"crossref","first-page":"3625","DOI":"10.1128\/AEM.02361-09","article-title":"The role of extra-cellular DNA during biofilm formation of Listeria monocytogenes","volume":"76","author":"Harmsen","year":"2010","journal-title":"Appl. Environ. Microbiol."},{"key":"ref_75","doi-asserted-by":"crossref","first-page":"341","DOI":"10.3109\/1040841X.2013.841639","article-title":"The role of extracellular DNA in the establishment, maintenance and perpetuation of bacterial biofilms","volume":"41","author":"Okshevsky","year":"2015","journal-title":"Crit. Rev. Microbiol."},{"key":"ref_76","doi-asserted-by":"crossref","first-page":"1467","DOI":"10.1016\/j.carres.2004.03.017","article-title":"Structural elucidation of the extracellular and cell-wall teichoic acids of Staphylococcus epidermidis RP62A, a reference biofilm-positive strain","volume":"339","author":"Sadovskaya","year":"2004","journal-title":"Carbohydr. Res."},{"key":"ref_77","first-page":"63","article-title":"Synthase-dependent exopolysaccharide secretion in gram-negative bacteria","volume":"21","author":"Whitney","year":"2013","journal-title":"TIM"},{"key":"ref_78","doi-asserted-by":"crossref","first-page":"876","DOI":"10.1111\/j.1365-2958.2007.05817.x","article-title":"The second messenger bis-(3\u2032-5\u2032)-cyclic-GMP and its PilZ domain-containing receptor Alg44 are required for alginate biosynthesis in Pseudomonas aeruginosa","volume":"65","author":"Merighi","year":"2007","journal-title":"Mol. Microbiol."},{"key":"ref_79","doi-asserted-by":"crossref","first-page":"1474","DOI":"10.1111\/j.1365-2958.2007.05879.x","article-title":"A cyclic-di-GMP receptor required for bacterial exopolysaccharide production","volume":"65","author":"Lee","year":"2007","journal-title":"Mol. Microbiol."},{"key":"ref_80","doi-asserted-by":"crossref","first-page":"279","DOI":"10.1038\/325279a0","article-title":"Regulation of cellulose synthesis in Acetobacter xylinum by cyclic diguanylic acid","volume":"325","author":"Ross","year":"1987","journal-title":"Nature"},{"key":"ref_81","doi-asserted-by":"crossref","first-page":"30310","DOI":"10.1074\/jbc.C600179200","article-title":"The pilz domain is a receptor for the second messenger c-di-GMP: The PilZ domain protein ycgr controls motility in enterobacteria","volume":"281","author":"Ryjenkov","year":"2006","journal-title":"J. Biol. Chem."},{"key":"ref_82","doi-asserted-by":"crossref","first-page":"181","DOI":"10.1038\/nature11744","article-title":"Crystallographic snapshot of cellulose synthesis and membrane translocation","volume":"493","author":"Morgan","year":"2013","journal-title":"Nature"},{"key":"ref_83","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1128\/MMBR.00043-12","article-title":"Cyclic di-GMP: The first 25 years of a universal bacterial second messenger","volume":"77","author":"Romling","year":"2013","journal-title":"Microbiol. Mol. Biol. Rev."},{"key":"ref_84","doi-asserted-by":"crossref","first-page":"9113","DOI":"10.1039\/c2ob26724a","article-title":"The bacterial second messenger c-di-GMP: Probing interactions with protein and RNA binding partners using cyclic dinucleotide analogs","volume":"10","author":"Shanahan","year":"2012","journal-title":"Org. Biomol. Chem."},{"key":"ref_85","doi-asserted-by":"crossref","first-page":"1286","DOI":"10.4315\/0362-028X-61.10.1286","article-title":"Influence of environmental stress on the kinetics and strength of attachment of Listeria monocytogenes Scott A to Buna-N rubber and stainless steel","volume":"61","author":"Smoot","year":"1998","journal-title":"J. Food Prot."},{"key":"ref_86","doi-asserted-by":"crossref","first-page":"5076","DOI":"10.1021\/pr1003642","article-title":"Comprehensive appraisal of the extracellular proteins from a monoderm bacterium: Theoretical and empirical exoproteomes of Listeria monocytogenes egd-e by secretomics","volume":"9","author":"Desvaux","year":"2010","journal-title":"J. Proteome Res."},{"key":"ref_87","doi-asserted-by":"crossref","first-page":"377","DOI":"10.1128\/MMBR.00039-06","article-title":"Listeria monocytogenes surface proteins: From genome predictions to function","volume":"71","author":"Bierne","year":"2007","journal-title":"Microbiol. Mol. Biol. Rev."},{"key":"ref_88","first-page":"21","article-title":"Towards the identification of the common features of bacterial biofilm development","volume":"9","author":"Lasa","year":"2006","journal-title":"Int. Microbiol."},{"key":"ref_89","doi-asserted-by":"crossref","first-page":"2888","DOI":"10.1128\/JB.183.9.2888-2896.2001","article-title":"Bap, a Staphylococcus aureus surface protein involved in biofilm formation","volume":"183","author":"Cucarella","year":"2001","journal-title":"J. Bacteriol."},{"key":"ref_90","doi-asserted-by":"crossref","first-page":"4538","DOI":"10.1128\/AEM.67.10.4538-4545.2001","article-title":"The enterococcal surface protein, Esp, is involved in Enterococcus faecalis biofilm formation","volume":"67","author":"Valle","year":"2001","journal-title":"Appl. Environ. Microbiol."},{"key":"ref_91","doi-asserted-by":"crossref","first-page":"1322","DOI":"10.1111\/j.1365-2958.2005.04907.x","article-title":"Bapa, a large secreted protein required for biofilm formation and host colonization of Salmonella enterica serovar enteritidis","volume":"58","author":"Latasa","year":"2005","journal-title":"Mol. Microbiol."},{"key":"ref_92","doi-asserted-by":"crossref","first-page":"905","DOI":"10.1046\/j.1365-2958.2003.03615.x","article-title":"Transition from reversible to irreversible attachment during biofilm formation by Pseudomonas fluorescens WCS365 requires an ABC transporter and a large secreted protein","volume":"49","author":"Hinsa","year":"2003","journal-title":"Mol. Microbiol."},{"key":"ref_93","doi-asserted-by":"crossref","first-page":"6030","DOI":"10.1128\/JB.00155-08","article-title":"Loss of flagellum-based motility by Listeria monocytogenes results in formation of hyperbiofilms","volume":"190","author":"Todhanakasem","year":"2008","journal-title":"J. Bacteriol."},{"key":"ref_94","first-page":"2171","article-title":"Temperature-dependent expression of flagella of Listeria monocytogenes studied by electron microscopy, SDS-PAGE and western blotting","volume":"134","author":"Peel","year":"1988","journal-title":"J. Gen. Microbiol."},{"key":"ref_95","doi-asserted-by":"crossref","unstructured":"Hall-Stoodley, L., Nistico, L., Sambanthamoorthy, K., Dice, B., Nguyen, D., Mershon, W.J., Johnson, C., Ze Hu, F., Stoodley, P., and Ehrlich, G.D. (2008). Characterization of biofilm matrix, degradation by dnase treatment and evidence of capsule downregulation in Streptococcus pneumoniae clinical isolates. BMC Microbiol., 8.","DOI":"10.1186\/1471-2180-8-173"},{"key":"ref_96","doi-asserted-by":"crossref","first-page":"470","DOI":"10.1128\/AEM.02073-07","article-title":"Differential roles of poly-N-acetylglucosamine surface polysaccharide and extracellular DNA in Staphylococcus aureus and Staphylococcus epidermidis biofilms","volume":"74","author":"Izano","year":"2008","journal-title":"Appl. Environ. Microbiol."},{"key":"ref_97","doi-asserted-by":"crossref","unstructured":"Mann, E.E., Rice, K.C., Boles, B.R., Endres, J.L., Ranjit, D., Chandramohan, L., Tsang, L.H., Smeltzer, M.S., Horswill, A.R., and Bayles, K.W. (2009). Modulation of eDNA release and degradation affects Staphylococcus aureus biofilm maturation. PLoS ONE, 4.","DOI":"10.1371\/journal.pone.0005822"},{"key":"ref_98","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_99","doi-asserted-by":"crossref","first-page":"2355","DOI":"10.1128\/JB.01493-14","article-title":"Streptococcus mutans extracellular DNA is upregulated during growth in biofilms, actively released via membrane vesicles, and influenced by components of the protein secretion machinery","volume":"196","author":"Liao","year":"2014","journal-title":"J. Bacteriol."},{"key":"ref_100","doi-asserted-by":"crossref","first-page":"259","DOI":"10.1016\/j.ijfoodmicro.2013.05.025","article-title":"Diversity assessment of Listeria monocytogenes biofilm formation: Impact of growth condition, serotype and strain origin","volume":"165","author":"Kadam","year":"2013","journal-title":"Int. J. Food Microbiol."},{"key":"ref_101","doi-asserted-by":"crossref","first-page":"7945","DOI":"10.1128\/JB.00858-07","article-title":"The EPS matrix: The \u201chouse of biofilm cells\u201d","volume":"189","author":"Flemming","year":"2007","journal-title":"J. Bacteriol."}],"container-title":["Microorganisms"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.mdpi.com\/2076-2607\/4\/3\/22\/pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2025,10,11]],"date-time":"2025-10-11T19:25:26Z","timestamp":1760210726000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.mdpi.com\/2076-2607\/4\/3\/22"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2016,7,5]]},"references-count":101,"journal-issue":{"issue":"3","published-online":{"date-parts":[[2016,9]]}},"alternative-id":["microorganisms4030022"],"URL":"https:\/\/doi.org\/10.3390\/microorganisms4030022","relation":{},"ISSN":["2076-2607"],"issn-type":[{"value":"2076-2607","type":"electronic"}],"subject":[],"published":{"date-parts":[[2016,7,5]]}}}