{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,6,4]],"date-time":"2026-06-04T20:58:36Z","timestamp":1780606716851,"version":"3.54.1"},"reference-count":59,"publisher":"Springer Science and Business Media LLC","issue":"1","license":[{"start":{"date-parts":[[2025,7,30]],"date-time":"2025-07-30T00:00:00Z","timestamp":1753833600000},"content-version":"tdm","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0"},{"start":{"date-parts":[[2025,7,30]],"date-time":"2025-07-30T00:00:00Z","timestamp":1753833600000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0"}],"funder":[{"DOI":"10.13039\/501100009244","name":"Stockholm University","doi-asserted-by":"crossref","id":[{"id":"10.13039\/501100009244","id-type":"DOI","asserted-by":"crossref"}]}],"content-domain":{"domain":["link.springer.com"],"crossmark-restriction":false},"short-container-title":["BMC Microbiol"],"abstract":"<jats:title>Abstract<\/jats:title>\n          <jats:sec>\n            <jats:title>Background<\/jats:title>\n            <jats:p>\n              <jats:italic>Neisseria meningitidis<\/jats:italic> asymptomatically colonizes the nasopharyngeal mucosa, but occasionally, the bacteria disseminate to cause sepsis and meningitis. In the epithelial cell layer, the pathogen co-colonizes with other resident inhabitants, such as <jats:italic>Lactobacillus<\/jats:italic> species that are part of the nasopharyngeal-oral microbiota. In this study, we investigated the interaction between lactobacilli and meningococci.<\/jats:p>\n          <\/jats:sec>\n          <jats:sec>\n            <jats:title>Results<\/jats:title>\n            <jats:p>We demonstrated that <jats:italic>Lactobacillus crispatus<\/jats:italic> strain MV24 can co-aggregate with meningococci, whereas all other <jats:italic>Lactobacillus<\/jats:italic> strains tested did not co-aggregate. The binding ability of <jats:italic>L. crispatus<\/jats:italic> was not strain- or serogroup-specific but was dependent on the ability of meningococci to form microcolonies. The finding that <jats:italic>N. meningitidis<\/jats:italic> lacking pili did not co-aggregate with <jats:italic>L. crispatus<\/jats:italic>, but that hyperpiliated <jats:italic>N. meningitidis<\/jats:italic> exhibited strong co-aggregation, led us to examine the interaction between purified meningococcal pili and lactobacilli. Our results showed that <jats:italic>L. crispatus<\/jats:italic> MV24 can bind to purified meningococcal Class I and II pili, explaining the aggregative clusters observed under the microscope. Co-aggregation with <jats:italic>L. crispatus<\/jats:italic> disrupted microcolony formation, and increased the killing of meningococci by LL-37, hBD2 and cephalexin. Further, co-aggregation had the added effect of impeding motility.<\/jats:p>\n          <\/jats:sec>\n          <jats:sec>\n            <jats:title>Conclusion<\/jats:title>\n            <jats:p>\n              <jats:italic>N. meningitidis<\/jats:italic> pili bind to <jats:italic>L. crispatus<\/jats:italic>, which interferes with the meningococcal microcolonies and increases sensitivity to antimicrobial agents. Taken together, our findings suggest that <jats:italic>L. crispatus<\/jats:italic> MV24 may have a beneficial effect on the host through co-aggregating with meningococci.<\/jats:p>\n          <\/jats:sec>","DOI":"10.1186\/s12866-025-04201-2","type":"journal-article","created":{"date-parts":[[2025,7,30]],"date-time":"2025-07-30T11:34:01Z","timestamp":1753875241000},"update-policy":"https:\/\/doi.org\/10.1007\/springer_crossmark_policy","source":"Crossref","is-referenced-by-count":2,"title":["Pilus-mediated co-aggregation with Lactobacillus crispatus increases meningococcal susceptibility to antimicrobial agents by interfering with microcolony formation"],"prefix":"10.1186","volume":"25","author":[{"given":"Kenny","family":"Lidberg","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Sarah","family":"Pilheden","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Samuddi","family":"Nawarathne","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Katharina","family":"Rauscher","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Ann-Beth","family":"Jonsson","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"297","published-online":{"date-parts":[[2025,7,30]]},"reference":[{"issue":"1","key":"4201_CR1","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1007\/s10096-017-3076-7","volume":"37","author":"S Esposito","year":"2018","unstructured":"Esposito S, Principi N. Impact of nasopharyngeal microbiota on the development of respiratory tract diseases. Eur J Clin Microbiol Infect Dis. 2018;37(1):1\u20137. https:\/\/doi.org\/10.1007\/s10096-017-3076-7.","journal-title":"Eur J Clin Microbiol Infect Dis"},{"issue":"2","key":"4201_CR2","doi-asserted-by":"publisher","first-page":"e17035","DOI":"10.1371\/journal.pone.0017035","volume":"6","author":"D Bogaert","year":"2011","unstructured":"Bogaert D, Keijser B, Huse S, Rossen J, Veenhoven R, van Gils E, et al. Variability and diversity of nasopharyngeal microbiota in children: a metagenomic analysis. PLoS ONE. 2011;6(2):e17035. https:\/\/doi.org\/10.1371\/journal.pone.0017035.","journal-title":"PLoS ONE"},{"issue":"Suppl 2","key":"4201_CR3","doi-asserted-by":"publisher","first-page":"B64","DOI":"10.1016\/j.vaccine.2009.04.061","volume":"27","author":"DA Caugant","year":"2009","unstructured":"Caugant DA, Maiden MC. Meningococcal carriage and disease\u2013population biology and evolution. Vaccine. 2009;27(Suppl 2):B64\u201370. https:\/\/doi.org\/10.1016\/j.vaccine.2009.04.061.","journal-title":"Vaccine"},{"key":"4201_CR4","doi-asserted-by":"publisher","first-page":"652","DOI":"10.1186\/1471-2164-11-652","volume":"11","author":"JS Bennett","year":"2010","unstructured":"Bennett JS, Bentley SD, Vernikos GS, Quail MA, Cherevach I, White B, et al. Independent evolution of the core and accessory gene sets in the genus neisseria: insights gained from the genome of neisseria lactamica isolate 020\u2013\u200906. BMC Genomics. 2010;11:652. https:\/\/doi.org\/10.1186\/1471-2164-11-652.","journal-title":"BMC Genomics"},{"issue":"3","key":"4201_CR5","doi-asserted-by":"publisher","first-page":"149","DOI":"10.1038\/nrmicro.2016.178","volume":"15","author":"M Coureuil","year":"2017","unstructured":"Coureuil M, L\u00e9cuyer H, Bourdoulous S, Nassif X. A journey into the brain: insight into how bacterial pathogens cross blood-brain barriers. Nat Rev Microbiol. 2017;15(3):149\u201359. https:\/\/doi.org\/10.1038\/nrmicro.2016.178.","journal-title":"Nat Rev Microbiol"},{"key":"4201_CR6","doi-asserted-by":"publisher","first-page":"139","DOI":"10.1016\/B978-0-12-407697-6.00004-0","volume":"303","author":"EP Lillehoj","year":"2013","unstructured":"Lillehoj EP, Kato K, Lu W, Kim KC. Cellular and molecular biology of airway mucins. Int Rev Cell Mol Biol. 2013;303:139\u2013202. https:\/\/doi.org\/10.1016\/B978-0-12-407697-6.00004-0.","journal-title":"Int Rev Cell Mol Biol"},{"key":"4201_CR7","doi-asserted-by":"publisher","DOI":"10.1128\/mSphere.00494-19","author":"M Audry","year":"2019","unstructured":"Audry M, Robbe-Masselot C, Barnier JP, Gachet B, Saubam\u00e9a B, Schmitt A, et al. Airway mucus restricts Neisseria meningitidis away from nasopharyngeal epithelial cells and protects the mucosa from inflammation. mSphere. 2019. https:\/\/doi.org\/10.1128\/mSphere.00494-19.","journal-title":"mSphere"},{"issue":"12","key":"4201_CR8","doi-asserted-by":"publisher","first-page":"3861","DOI":"10.1128\/JB.01313-08","volume":"191","author":"A Jones","year":"2009","unstructured":"Jones A, Georg M, Maudsdotter L, Jonsson AB. Endotoxin, capsule, and bacterial attachment contribute to Neisseria meningitidis resistance to the human antimicrobial peptide LL-37. J Bacteriol. 2009;191(12):3861\u20138. https:\/\/doi.org\/10.1128\/JB.01313-08.","journal-title":"J Bacteriol"},{"key":"4201_CR9","doi-asserted-by":"publisher","first-page":"323","DOI":"10.1016\/bs.ampbs.2015.05.001","volume":"66","author":"JR Laver","year":"2015","unstructured":"Laver JR, Hughes SE, Read RC. Neisserial molecular adaptations to the nasopharyngeal niche. Adv Microb Physiol. 2015;66:323\u201355. https:\/\/doi.org\/10.1016\/bs.ampbs.2015.05.001.","journal-title":"Adv Microb Physiol"},{"issue":"5","key":"4201_CR10","doi-asserted-by":"publisher","first-page":"1442","DOI":"10.1084\/jem.152.5.1442","volume":"152","author":"MH Mulks","year":"1980","unstructured":"Mulks MH, Plaut AG, Feldman HA, Frangione B. IgA proteases of two distinct specificities are released by Neisseria meningitidis. J Exp Med. 1980;152(5):1442\u20137. https:\/\/doi.org\/10.1084\/jem.152.5.1442.","journal-title":"J Exp Med"},{"issue":"10","key":"4201_CR11","doi-asserted-by":"publisher","first-page":"5001","DOI":"10.1128\/IAI.66.10.5001-5007.1998","volume":"66","author":"P Ayala","year":"1998","unstructured":"Ayala P, Lin L, Hopper S, Fukuda M, So M. Infection of epithelial cells by pathogenic neisseriae reduces the levels of multiple lysosomal constituents. Infect Immun. 1998;66(10):5001\u20137. https:\/\/doi.org\/10.1128\/IAI.66.10.5001-5007.1998.","journal-title":"Infect Immun"},{"issue":"9","key":"4201_CR12","doi-asserted-by":"publisher","first-page":"e1005162","DOI":"10.1371\/journal.ppat.1005162","volume":"11","author":"J Gault","year":"2015","unstructured":"Gault J, Ferber M, Machata S, Imhaus AF, Malosse C, Charles-Orszag A, et al. Neisseria meningitidis type IV pili composed of sequence invariable pilins are masked by multisite glycosylation. PLoS Pathog. 2015;11(9):e1005162. https:\/\/doi.org\/10.1371\/journal.ppat.1005162.","journal-title":"PLoS Pathog"},{"issue":"1","key":"4201_CR13","doi-asserted-by":"publisher","first-page":"2414","DOI":"10.1038\/s41467-024-46677-y","volume":"15","author":"D Fernandez-Martinez","year":"2024","unstructured":"Fernandez-Martinez D, Kong Y, Goussard S, Zavala A, Gastineau P, Rey M, et al. Cryo-EM structures of type IV pili complexed with nanobodies reveal immune escape mechanisms. Nat Commun. 2024;15(1):2414. https:\/\/doi.org\/10.1038\/s41467-024-46677-y.","journal-title":"Nat Commun"},{"issue":"1","key":"4201_CR14","doi-asserted-by":"publisher","first-page":"65","DOI":"10.1111\/j.1365-2958.2004.04372.x","volume":"55","author":"S Helaine","year":"2005","unstructured":"Helaine S, Carbonnelle E, Prouvensier L, Beretti JL, Nassif X, Pelicic V. Pilx, a pilus-associated protein essential for bacterial aggregation, is a key to pilus-facilitated attachment of neisseria meningitidis to human cells. Mol Microbiol. 2005;55(1):65\u201377. https:\/\/doi.org\/10.1111\/j.1365-2958.2004.04372.x.","journal-title":"Mol Microbiol"},{"issue":"4","key":"4201_CR15","doi-asserted-by":"publisher","first-page":"613","DOI":"10.1007\/s00018-008-8477-4","volume":"66","author":"T Proft","year":"2009","unstructured":"Proft T, Baker EN. Pili in Gram-negative and Gram-positive bacteria - structure, assembly and their role in disease. Cell Mol Life Sci. 2009;66(4):613\u201335. https:\/\/doi.org\/10.1007\/s00018-008-8477-4.","journal-title":"Cell Mol Life Sci"},{"issue":"1","key":"4201_CR16","doi-asserted-by":"publisher","first-page":"54","DOI":"10.1111\/j.1365-2958.2004.04364.x","volume":"55","author":"E Carbonnelle","year":"2005","unstructured":"Carbonnelle E, Helaine S, Prouvensier L, Nassif X, Pelicic V. Type IV pilus biogenesis in neisseria meningitidis: PilW is involved in a step occurring after pilus assembly, essential for fibre stability and function. Mol Microbiol. 2005;55(1):54\u201364. https:\/\/doi.org\/10.1111\/j.1365-2958.2004.04364.x.","journal-title":"Mol Microbiol"},{"issue":"4","key":"4201_CR17","doi-asserted-by":"publisher","first-page":"827","DOI":"10.1111\/j.1365-2958.2008.06197.x","volume":"68","author":"V Pelicic","year":"2008","unstructured":"Pelicic V. Type IV pili: e pluribus unum? Mol Microbiol. 2008;68(4):827\u201337. https:\/\/doi.org\/10.1111\/j.1365-2958.2008.06197.x.","journal-title":"Mol Microbiol."},{"issue":"7","key":"4201_CR18","doi-asserted-by":"publisher","DOI":"10.1371\/journal.pone.0021749","volume":"6","author":"A Kuwae","year":"2011","unstructured":"Kuwae A, Sjolinder H, Eriksson J, Eriksson S, Chen Y, Jonsson AB. NafA negatively controls neisseria meningitidis piliation. PLoS One. 2011;6(7): e21749. https:\/\/doi.org\/10.1371\/journal.pone.0021749.","journal-title":"PLoS One"},{"issue":"2","key":"4201_CR19","doi-asserted-by":"publisher","first-page":"e1009251","DOI":"10.1371\/journal.ppat.1009251","volume":"17","author":"T Cronenberg","year":"2021","unstructured":"Cronenberg T, Hennes M, Wielert I, Maier B. Antibiotics modulate attractive interactions in bacterial colonies affecting survivability under combined treatment. PLoS Pathog. 2021;17(2):e1009251. https:\/\/doi.org\/10.1371\/journal.ppat.1009251.","journal-title":"PLoS Pathog"},{"issue":"1","key":"4201_CR20","doi-asserted-by":"publisher","first-page":"e3003022","DOI":"10.1371\/journal.pbio.3003022","volume":"23","author":"I Wielert","year":"2025","unstructured":"Wielert I, Kraus-R\u00f6mer S, Volkmann TE, Craig L, Higgins PG, Maier B. Pilin antigenic variants impact gonococcal lifestyle and antibiotic tolerance by modulating interbacterial forces. PLoS Biol. 2025;23(1):e3003022. https:\/\/doi.org\/10.1371\/journal.pbio.3003022.","journal-title":"PLoS Biol"},{"key":"4201_CR21","doi-asserted-by":"publisher","first-page":"697232","DOI":"10.3389\/fmicb.2021.697232","volume":"12","author":"GM Wassing","year":"2021","unstructured":"Wassing GM, Lidberg K, Sigurl\u00e1sd\u00f3ttir S, Frey J, Schroeder K, Ilehag N, et al. DNA blocks the lethal effect of human Beta-Defensin 2 against. Front Microbiol. 2021;12. https:\/\/doi.org\/10.3389\/fmicb.2021.697232.","journal-title":"Front Microbiol."},{"issue":"10","key":"4201_CR22","doi-asserted-by":"publisher","first-page":"2000","DOI":"10.1111\/all.13449","volume":"73","author":"M Dzidic","year":"2018","unstructured":"Dzidic M, Abrahamsson TR, Artacho A, Collado MC, Mira A, Jenmalm MC. Oral microbiota maturation during the first 7 years of life in relation to allergy development. Allergy. 2018;73(10):2000\u201311. https:\/\/doi.org\/10.1111\/all.13449.","journal-title":"Allergy"},{"issue":"6","key":"4201_CR23","doi-asserted-by":"publisher","first-page":"4076","DOI":"10.1016\/S0006-3495(03)74820-6","volume":"85","author":"P Sch\u00e4r-Zammaretti","year":"2003","unstructured":"Sch\u00e4r-Zammaretti P, Ubbink J. The cell wall of lactic acid bacteria: surface constituents and macromolecular conformations. Biophys J. 2003;85(6):4076\u201392. https:\/\/doi.org\/10.1016\/S0006-3495(03)74820-6.","journal-title":"Biophys J"},{"issue":"12","key":"4201_CR24","doi-asserted-by":"publisher","first-page":"5225","DOI":"10.1007\/s00253-013-4962-2","volume":"97","author":"U Hyn\u00f6nen","year":"2013","unstructured":"Hyn\u00f6nen U, Palva A. Lactobacillus surface layer proteins: structure, function and applications. Appl Microbiol Biotechnol. 2013;97(12):5225\u201343. https:\/\/doi.org\/10.1007\/s00253-013-4962-2.","journal-title":"Appl Microbiol Biotechnol"},{"key":"4201_CR25","doi-asserted-by":"publisher","DOI":"10.1128\/mBio.00460-13","author":"SS Witkin","year":"2013","unstructured":"Witkin SS, Mendes-Soares H, Linhares IM, Jayaram A, Ledger WJ, Forney LJ. Influence of vaginal bacteria and D- and L-lactic acid isomers on vaginal extracellular matrix metalloproteinase inducer: implications for protection against upper genital tract infections. MBio. 2013. https:\/\/doi.org\/10.1128\/mBio.00460-13.","journal-title":"MBio"},{"issue":"9","key":"4201_CR26","doi-asserted-by":"publisher","first-page":"1369","DOI":"10.1086\/597390","volume":"199","author":"DF Muench","year":"2009","unstructured":"Muench DF, Kuch DJ, Wu H, Begum AA, Veit SJ, Pelletier ME, et al. Hydrogen peroxide-producing lactobacilli inhibit gonococci in vitro but not during experimental genital tract infection. J Infect Dis. 2009;199(9):1369\u201378. https:\/\/doi.org\/10.1086\/597390.","journal-title":"J Infect Dis"},{"issue":"5","key":"4201_CR27","doi-asserted-by":"publisher","first-page":"1985","DOI":"10.1128\/IAI.66.5.1985-1989.1998","volume":"66","author":"S Boris","year":"1998","unstructured":"Boris S, Su\u00e1rez JE, V\u00e1zquez F, Barb\u00e9s C. Adherence of human vaginal lactobacilli to vaginal epithelial cells and interaction with uropathogens. Infect Immun. 1998;66(5):1985\u20139.","journal-title":"Infect Immun"},{"issue":"1","key":"4201_CR28","doi-asserted-by":"publisher","first-page":"31","DOI":"10.1080\/1040841X.2017.1306688","volume":"44","author":"S Kovachev","year":"2018","unstructured":"Kovachev S. Defence factors of vaginal lactobacilli. Crit Rev Microbiol. 2018;44(1):31\u20139. https:\/\/doi.org\/10.1080\/1040841X.2017.1306688.","journal-title":"Crit Rev Microbiol"},{"issue":"3","key":"4201_CR29","doi-asserted-by":"publisher","first-page":"87","DOI":"10.1016\/j.ijfoodmicro.2010.07.032","volume":"143","author":"W Turpin","year":"2010","unstructured":"Turpin W, Humblot C, Thomas M, Guyot JP. Lactobacilli as multifaceted probiotics with poorly disclosed molecular mechanisms. Int J Food Microbiol. 2010;143(3):87\u2013102. https:\/\/doi.org\/10.1016\/j.ijfoodmicro.2010.07.032.","journal-title":"Int J Food Microbiol"},{"issue":"2","key":"4201_CR30","doi-asserted-by":"publisher","first-page":"111","DOI":"10.1111\/j.1574-695X.1997.tb01079.x","volume":"19","author":"V Kmet","year":"1997","unstructured":"Kmet V, Lucchini F. Aggregation-promoting factor in human vaginal Lactobacillus strains. FEMS Immunol Med Microbiol. 1997;19(2):111\u20134. https:\/\/doi.org\/10.1111\/j.1574-695X.1997.tb01079.x.","journal-title":"FEMS Immunol Med Microbiol"},{"issue":"1","key":"4201_CR31","doi-asserted-by":"publisher","first-page":"11","DOI":"10.1046\/j.1365-2958.1997.4601823.x","volume":"25","author":"M Rahman","year":"1997","unstructured":"Rahman M, Kallstrom H, Normark S, Jonsson AB. Pilc of pathogenic neisseria is associated with the bacterial cell surface. Mol Microbiol. 1997;25(1):11\u201325.","journal-title":"Mol Microbiol"},{"issue":"10","key":"4201_CR32","doi-asserted-by":"publisher","first-page":"e0029621","DOI":"10.1128\/IAI.00296-21","volume":"89","author":"S Sigurl\u00e1sd\u00f3ttir","year":"2021","unstructured":"Sigurl\u00e1sd\u00f3ttir S, Lidberg K, Zuo F, Newcombe J, McFadden J, Jonsson AB. Lactate-Induced dispersal of. Infect Immun. 2021;89(10):e0029621. https:\/\/doi.org\/10.1128\/IAI.00296-21.","journal-title":"Infect Immun"},{"issue":"5","key":"4201_CR33","doi-asserted-by":"publisher","first-page":"1501","DOI":"10.1128\/IAI.01463-15","volume":"84","author":"J Engman","year":"2016","unstructured":"Engman J, Negrea A, Sigurl\u00e1sd\u00f3ttir S, Ge\u00f6rg M, Eriksson J, Eriksson OS, et al. Neisseria meningitidis polynucleotide phosphorylase affects aggregation, adhesion, and virulence. Infect Immun. 2016;84(5):1501\u201313. https:\/\/doi.org\/10.1128\/IAI.01463-15.","journal-title":"Infect Immun"},{"issue":"1","key":"4201_CR34","doi-asserted-by":"publisher","first-page":"155","DOI":"10.1128\/IAI.71.1.155-162.2003","volume":"71","author":"B Albiger","year":"2003","unstructured":"Albiger B, Johansson L, Jonsson AB. Lipooligosaccharide-deficient Neisseria meningitidis shows altered pilus-associated characteristics. Infect Immun. 2003;71(1):155\u201362.","journal-title":"Infect Immun"},{"issue":"6777","key":"4201_CR35","doi-asserted-by":"publisher","first-page":"502","DOI":"10.1038\/35006655","volume":"404","author":"J Parkhill","year":"2000","unstructured":"Parkhill J, Achtman M, James KD, Bentley SD, Churcher C, Klee SR, et al. Complete DNA sequence of a serogroup A strain of neisseria meningitidis Z2491. Nature. 2000;404(6777):502\u20136. https:\/\/doi.org\/10.1038\/35006655.","journal-title":"Nature"},{"issue":"4","key":"4201_CR36","doi-asserted-by":"publisher","first-page":"2082","DOI":"10.1128\/iai.68.4.2082-2095.2000","volume":"68","author":"SR Klee","year":"2000","unstructured":"Klee SR, Nassif X, Kusecek B, Merker P, Beretti JL, Achtman M, et al. Molecular and biological analysis of eight genetic Islands that distinguish neisseria meningitidis from the closely related pathogen neisseria gonorrhoeae. Infect Immun. 2000;68(4):2082\u201395. https:\/\/doi.org\/10.1128\/iai.68.4.2082-2095.2000.","journal-title":"Infect Immun"},{"issue":"8","key":"4201_CR37","doi-asserted-by":"publisher","first-page":"3704","DOI":"10.1128\/AAC.00624-13","volume":"57","author":"OS Eriksson","year":"2013","unstructured":"Eriksson OS, Georg M, Sjolinder H, Sillard R, Lindberg S, Langel U, et al. Identification of cell-penetrating peptides that are bactericidal to neisseria meningitidis and prevent inflammatory responses upon infection. Antimicrob Agents Chemother. 2013;57(8):3704\u201312. https:\/\/doi.org\/10.1128\/AAC.00624-13.","journal-title":"Antimicrob Agents Chemother"},{"issue":"6","key":"4201_CR38","doi-asserted-by":"publisher","first-page":"1746","DOI":"10.1128\/JCM.36.6.1746-1749.1998","volume":"36","author":"A Bart","year":"1998","unstructured":"Bart A, Schuurman IG, Achtman M, Caugant DA, Dankert J, van der Ende A. Randomly amplified polymorphic DNA genotyping of serogroup A meningococci yields results similar to those obtained by multilocus enzyme electrophoresis and reveals new genotypes. J Clin Microbiol. 1998;36(6):1746\u20139. https:\/\/doi.org\/10.1128\/JCM.36.6.1746-1749.1998.","journal-title":"J Clin Microbiol"},{"issue":"5","key":"4201_CR39","doi-asserted-by":"publisher","first-page":"1345","DOI":"10.1111\/j.1365-2958.2001.02324.x","volume":"39","author":"CR Johnson","year":"2001","unstructured":"Johnson CR, Newcombe J, Thorne S, Borde HA, Eales-Reynolds LJ, Gorringe AR, et al. Generation and characterization of a PhoP homologue mutant of neisseria meningitidis. Mol Microbiol. 2001;39(5):1345\u201355. https:\/\/doi.org\/10.1111\/j.1365-2958.2001.02324.x.","journal-title":"Mol Microbiol"},{"issue":"5459","key":"4201_CR40","doi-asserted-by":"publisher","first-page":"1809","DOI":"10.1126\/science.287.5459.1809","volume":"287","author":"H Tettelin","year":"2000","unstructured":"Tettelin H, Saunders NJ, Heidelberg J, Jeffries AC, Nelson KE, Eisen JA, et al. Complete genome sequence of Neisseria meningitidis serogroup B strain MC58. Science. 2000;287(5459):1809\u201315. https:\/\/doi.org\/10.1126\/science.287.5459.1809.","journal-title":"Science"},{"issue":"Pt 1","key":"4201_CR41","doi-asserted-by":"publisher","first-page":"77","DOI":"10.1099\/ijs.0.63083-0","volume":"55","author":"S Roos","year":"2005","unstructured":"Roos S, Engstrand L, Jonsson H. Lactobacillus gastricus sp. nov., Lactobacillus antri sp. nov., Lactobacillus kalixensis sp. nov. and Lactobacillus ultunensis sp. nov., isolated from human stomach mucosa. Int J Syst Evol Microbiol. 2005;55(Pt 1):77\u201382. https:\/\/doi.org\/10.1099\/ijs.0.63083-0.","journal-title":"Int J Syst Evol Microbiol"},{"issue":"5","key":"4201_CR42","doi-asserted-by":"publisher","first-page":"1526","DOI":"10.1128\/IAI.00163-16","volume":"84","author":"N de Klerk","year":"2016","unstructured":"de Klerk N, Maudsdotter L, Gebreegziabher H, Saroj SD, Eriksson B, Eriksson OS, et al. Lactobacilli reduce Helicobacter pylori attachment to host gastric epithelial cells by inhibiting adhesion gene expression. Infect Immun. 2016;84(5):1526\u201335. https:\/\/doi.org\/10.1128\/IAI.00163-16.","journal-title":"Infect Immun"},{"issue":"4","key":"4201_CR43","doi-asserted-by":"publisher","first-page":"2782","DOI":"10.1099\/ijsem.0.004107","volume":"70","author":"J Zheng","year":"2020","unstructured":"Zheng J, Wittouck S, Salvetti E, Franz CMAP, Harris HMB, Mattarelli P, et al. A taxonomic note on the genus. Int J Syst Evol Microbiol. 2020;70(4):2782\u2013858. https:\/\/doi.org\/10.1099\/ijsem.0.004107.","journal-title":"Int J Syst Evol Microbiol"},{"issue":"3","key":"4201_CR44","doi-asserted-by":"publisher","first-page":"596","DOI":"10.1128\/jb.96.3.596-605.1968","volume":"96","author":"DS Kellogg","year":"1968","unstructured":"Kellogg DS, Cohen IR, Norins LC, Schroeter AL, Reising G. Neisseria gonorrhoeae. II. Colonial variation and pathogenicity during 35 months in vitro. J Bacteriol. 1968;96(3):596\u2013605.","journal-title":"J Bacteriol"},{"issue":"6","key":"4201_CR45","doi-asserted-by":"publisher","first-page":"981","DOI":"10.1046\/j.1365-2672.2003.01915.x","volume":"94","author":"B Kos","year":"2003","unstructured":"Kos B, Suskovi\u0107 J, Vukovi\u0107 S, Simpraga M, Frece J, Matosi\u0107 S. Adhesion and aggregation ability of probiotic strain Lactobacillus acidophilus M92. J Appl Microbiol. 2003;94(6):981\u20137. https:\/\/doi.org\/10.1046\/j.1365-2672.2003.01915.x.","journal-title":"J Appl Microbiol"},{"issue":"12","key":"4201_CR46","doi-asserted-by":"publisher","first-page":"4154","DOI":"10.1128\/IAI.00423-12","volume":"80","author":"H Takahashi","year":"2012","unstructured":"Takahashi H, Yanagisawa T, Kim KS, Yokoyama S, Ohnishi M. Meningococcal PilV potentiates neisseria meningitidis type IV pilus-mediated internalization into human endothelial and epithelial cells. Infect Immun. 2012;80(12):4154\u201366. https:\/\/doi.org\/10.1128\/IAI.00423-12.","journal-title":"Infect Immun"},{"key":"4201_CR47","doi-asserted-by":"publisher","first-page":"73","DOI":"10.1007\/978-1-4939-0473-0_9","volume":"1149","author":"L Turnbull","year":"2014","unstructured":"Turnbull L, Whitchurch CB. Motility assay: twitching motility. Methods Mol Biol. 2014;1149:73\u201386. https:\/\/doi.org\/10.1007\/978-1-4939-0473-0_9.","journal-title":"Methods Mol Biol"},{"issue":"7","key":"4201_CR48","doi-asserted-by":"publisher","first-page":"2538","DOI":"10.1128\/IAI.06354-11","volume":"80","author":"J Eriksson","year":"2012","unstructured":"Eriksson J, Eriksson OS, Jonsson AB. Loss of meningococcal PilU delays microcolony formation and attenuates virulence in vivo. Infect Immun. 2012;80(7):2538\u201347. https:\/\/doi.org\/10.1128\/IAI.06354-11.","journal-title":"Infect Immun"},{"issue":"7","key":"4201_CR49","doi-asserted-by":"publisher","first-page":"4017","DOI":"10.1073\/pnas.96.7.4017","volume":"96","author":"C Pujol","year":"1999","unstructured":"Pujol C, Eugene E, Marceau M, Nassif X. The meningococcal PilT protein is required for induction of intimate attachment to epithelial cells following pilus-mediated adhesion. Proc Natl Acad Sci U S A. 1999;96(7):4017\u201322.","journal-title":"Proc Natl Acad Sci U S A"},{"issue":"4","key":"4201_CR50","doi-asserted-by":"publisher","first-page":"e1006251","DOI":"10.1371\/journal.ppat.1006251","volume":"13","author":"S Sigurlasdottir","year":"2017","unstructured":"Sigurlasdottir S, Engman J, Eriksson OS, Saroj SD, Zguna N, Lloris-Garcera P, et al. Host cell-derived lactate functions as an effector molecule in neisseria meningitidis microcolony dispersal. PLoS Pathog. 2017;13(4):e1006251. https:\/\/doi.org\/10.1371\/journal.ppat.1006251.","journal-title":"PLoS Pathog"},{"issue":"3","key":"4201_CR51","doi-asserted-by":"publisher","first-page":"e0007622","DOI":"10.1128\/mmbr.00076-22","volume":"86","author":"PK Singh","year":"2022","unstructured":"Singh PK, Little J, Donnenberg MS. Landmark discoveries and recent advances in type IV Pilus research. Microbiol Mol Biol Rev. 2022;86(3):e0007622. https:\/\/doi.org\/10.1128\/mmbr.00076-22.","journal-title":"Microbiol Mol Biol Rev"},{"issue":"3","key":"4201_CR52","doi-asserted-by":"publisher","first-page":"e1008372","DOI":"10.1371\/journal.ppat.1008372","volume":"16","author":"R Custodio","year":"2020","unstructured":"Custodio R, Johnson E, Liu G, Tang CM, Exley RM. Commensal neisseria cinerea impairs neisseria meningitidis microcolony development and reduces pathogen colonisation of epithelial cells. PLoS Pathog. 2020;16(3):e1008372. https:\/\/doi.org\/10.1371\/journal.ppat.1008372.","journal-title":"PLoS Pathog"},{"issue":"6800","key":"4201_CR53","doi-asserted-by":"publisher","first-page":"98","DOI":"10.1038\/35024105","volume":"407","author":"AJ Merz","year":"2000","unstructured":"Merz AJ, So M, Sheetz MP. Pilus Retraction powers bacterial twitching motility. Nature. 2000;407(6800):98\u2013102.","journal-title":"Nature"},{"issue":"4","key":"4201_CR54","doi-asserted-by":"publisher","first-page":"e100","DOI":"10.1371\/journal.pbio.0030100","volume":"3","author":"HL Howie","year":"2005","unstructured":"Howie HL, Glogauer M, So M. The N. gonorrhoeae type IV Pilus stimulates mechanosensitive pathways and cytoprotection through a pilT-dependent mechanism. PLoS Biol. 2005;3(4):e100. https:\/\/doi.org\/10.1371\/journal.pbio.0030100.","journal-title":"PLoS Biol"},{"key":"4201_CR55","doi-asserted-by":"publisher","unstructured":"Lidberg K, Pilheden S, Relloso Ortiz de Uriarte M, Jonsson AB. Internalization of. Microorganisms. 2025;13(3). https:\/\/doi.org\/10.3390\/microorganisms13030479.","DOI":"10.3390\/microorganisms13030479"},{"issue":"12","key":"4201_CR56","doi-asserted-by":"publisher","first-page":"6718","DOI":"10.4049\/jimmunol.163.12.6718","volume":"163","author":"DA O\u2019Neil","year":"1999","unstructured":"O\u2019Neil DA, Porter EM, Elewaut D, Anderson GM, Eckmann L, Ganz T, et al. Expression and regulation of the human beta-defensins hBD-1 and hBD-2 in intestinal epithelium. J Immunol. 1999;163(12):6718\u201324.","journal-title":"J Immunol"},{"issue":"1","key":"4201_CR57","doi-asserted-by":"publisher","first-page":"106","DOI":"10.1046\/j.0022-202x.2001.01401.x","volume":"117","author":"RS Ali","year":"2001","unstructured":"Ali RS, Falconer A, Ikram M, Bissett CE, Cerio R, Quinn AG. Expression of the peptide antibiotics human beta defensin-1 and human beta defensin-2 in normal human skin. J Invest Dermatol. 2001;117(1):106\u201311. https:\/\/doi.org\/10.1046\/j.0022-202x.2001.01401.x.","journal-title":"J Invest Dermatol"},{"key":"4201_CR58","doi-asserted-by":"publisher","DOI":"10.3389\/fmicb.2021.697232","volume":"12","author":"GM Wassing","year":"2021","unstructured":"Wassing GM, Lidberg K, Sigurlasdottir S, Frey J, Schroeder K, Ilehag N, et al. DNA blocks the lethal effect of human beta-defensin 2 against Neisseria meningitidis. Front Microbiol. 2021;12: 697232. https:\/\/doi.org\/10.3389\/fmicb.2021.697232.","journal-title":"Front Microbiol"},{"issue":"6","key":"4201_CR59","doi-asserted-by":"publisher","first-page":"e1010561","DOI":"10.1371\/journal.ppat.1010561","volume":"18","author":"LI Hu","year":"2022","unstructured":"Hu LI, Stohl EA, Seifert HS. The neisseria gonorrhoeae type IV Pilus promotes resistance to hydrogen peroxide- and LL-37-mediated killing by modulating the availability of intracellular, labile iron. PLoS Pathog. 2022;18(6):e1010561. https:\/\/doi.org\/10.1371\/journal.ppat.1010561.","journal-title":"PLoS Pathog"}],"container-title":["BMC Microbiology"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/link.springer.com\/content\/pdf\/10.1186\/s12866-025-04201-2.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/link.springer.com\/article\/10.1186\/s12866-025-04201-2\/fulltext.html","content-type":"text\/html","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/link.springer.com\/content\/pdf\/10.1186\/s12866-025-04201-2.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2025,9,8]],"date-time":"2025-09-08T07:36:40Z","timestamp":1757317000000},"score":1,"resource":{"primary":{"URL":"https:\/\/bmcmicrobiol.biomedcentral.com\/articles\/10.1186\/s12866-025-04201-2"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2025,7,30]]},"references-count":59,"journal-issue":{"issue":"1","published-online":{"date-parts":[[2025,12]]}},"alternative-id":["4201"],"URL":"https:\/\/doi.org\/10.1186\/s12866-025-04201-2","relation":{},"ISSN":["1471-2180"],"issn-type":[{"value":"1471-2180","type":"electronic"}],"subject":[],"published":{"date-parts":[[2025,7,30]]},"assertion":[{"value":"24 April 2025","order":1,"name":"received","label":"Received","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"9 July 2025","order":2,"name":"accepted","label":"Accepted","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"30 July 2025","order":3,"name":"first_online","label":"First Online","group":{"name":"ArticleHistory","label":"Article History"}},{"order":1,"name":"Ethics","group":{"name":"EthicsHeading","label":"Declarations"}},{"value":"Not applicable.","order":2,"name":"Ethics","group":{"name":"EthicsHeading","label":"Ethics approval and consent to participate"}},{"value":"Not applicable.","order":3,"name":"Ethics","group":{"name":"EthicsHeading","label":"Consent for publication"}},{"value":"The authors declare no competing interests.","order":4,"name":"Ethics","group":{"name":"EthicsHeading","label":"Competing interests"}}],"article-number":"467"}}