{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,11,9]],"date-time":"2026-11-09T23:30:03Z","timestamp":1794267003181,"version":"4.9.0"},"reference-count":56,"publisher":"Springer Science and Business Media LLC","issue":"1","license":[{"start":{"date-parts":[[2026,5,23]],"date-time":"2026-05-23T00:00:00Z","timestamp":1779494400000},"content-version":"tdm","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by-nc-nd\/4.0"},{"start":{"date-parts":[[2026,7,9]],"date-time":"2026-07-09T00:00:00Z","timestamp":1783555200000},"content-version":"vor","delay-in-days":47,"URL":"https:\/\/creativecommons.org\/licenses\/by-nc-nd\/4.0"}],"funder":[{"DOI":"10.13039\/501100001809","name":"National Natural Science Foundation of China","doi-asserted-by":"crossref","award":["82372310 and 82172315"],"award-info":[{"award-number":["82372310 and 82172315"]}],"id":[{"id":"10.13039\/501100001809","id-type":"DOI","asserted-by":"crossref"}]}],"content-domain":{"domain":["link.springer.com"],"crossmark-restriction":false},"short-container-title":["BMC Microbiol"],"DOI":"10.1186\/s12866-026-05146-w","type":"journal-article","created":{"date-parts":[[2026,5,23]],"date-time":"2026-05-23T01:43:53Z","timestamp":1779500633000},"update-policy":"https:\/\/doi.org\/10.1007\/springer_crossmark_policy","source":"Crossref","is-referenced-by-count":1,"title":["Lineage-specific type 1 fimbriae in hypervirulent ST23 Klebsiella pneumoniae target \u03b13\u03b21 integrin to promote intestinal epithelial invasion"],"prefix":"10.1186","volume":"26","author":[{"given":"Qian","family":"Zeng","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Ying","family":"Zhang","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Shujing","family":"Li","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Xinyue","family":"Fan","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Su","family":"Wang","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Jiayuan","family":"Liu","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Cui","family":"Tai","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Meng","family":"Li","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Yuanyuan","family":"Wei","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Yunkun","family":"Huang","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Yi","family":"Li","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Lefu","family":"Lan","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Xiaofei","family":"Jiang","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"297","published-online":{"date-parts":[[2026,5,23]]},"reference":[{"issue":"11","key":"5146_CR1","doi-asserted-by":"publisher","first-page":"881","DOI":"10.1016\/s1473-3099(12)70205-0","volume":"12","author":"LK Siu","year":"2012","unstructured":"Siu LK, Yeh KM, Lin JC, Fung CP, Chang FY. Klebsiella pneumoniae liver abscess: a new invasive syndrome. Lancet Infect Dis. 2012;12(11):881\u20137. https:\/\/doi.org\/10.1016\/s1473-3099(12)70205-0.","journal-title":"Lancet Infect Dis"},{"key":"5146_CR2","doi-asserted-by":"publisher","unstructured":"Russo TA, Marr CM. Hypervirulent Klebsiella pneumoniae. Clin Microbiol Rev. 2019;32(3). https:\/\/doi.org\/10.1128\/cmr.00001-19.","DOI":"10.1128\/cmr.00001-19"},{"issue":"1","key":"5146_CR3","doi-asserted-by":"publisher","first-page":"2703","DOI":"10.1038\/s41467-018-05114-7","volume":"9","author":"MMC Lam","year":"2018","unstructured":"Lam MMC, Wyres KL, Duch\u00eane S, Wick RR, Judd LM, Gan YH, et al. Population genomics of hypervirulent Klebsiella pneumoniae clonal-group 23 reveals early emergence and rapid global dissemination. Nat Commun. 2018;9(1):2703. https:\/\/doi.org\/10.1038\/s41467-018-05114-7.","journal-title":"Nat Commun"},{"key":"5146_CR4","doi-asserted-by":"publisher","first-page":"5215","DOI":"10.2147\/jir.S334581","volume":"14","author":"Y Zheng","year":"2021","unstructured":"Zheng Y, Ding Y, Xu M, Chen H, Zhang H, Liu Y, et al. Gut Microbiota Contributes to Host Defense Against Klebsiella pneumoniae-Induced Liver Abscess. J Inflamm Res. 2021;14:5215\u201325. https:\/\/doi.org\/10.2147\/jir.S334581.","journal-title":"J Inflamm Res"},{"issue":"7","key":"5146_CR5","doi-asserted-by":"publisher","first-page":"952","DOI":"10.1007\/s00284-018-1471-7","volume":"75","author":"N Chen","year":"2018","unstructured":"Chen N, Ling ZX, Jin TT, Li M, Zhao S, Zheng LS, et al. Altered Profiles of Gut Microbiota in Klebsiella pneumoniae-Induced Pyogenic Liver Abscess. Curr Microbiol. 2018;75(7):952\u20139. https:\/\/doi.org\/10.1007\/s00284-018-1471-7.","journal-title":"Curr Microbiol"},{"issue":"3","key":"5146_CR6","doi-asserted-by":"publisher","first-page":"629","DOI":"10.1128\/mmbr.00078-15","volume":"80","author":"MK Paczosa","year":"2016","unstructured":"Paczosa MK, Mecsas J. Klebsiella pneumoniae: Going on the Offense with a Strong Defense. Microbiol Mol Biol Rev. 2016;80(3):629\u201361. https:\/\/doi.org\/10.1128\/mmbr.00078-15.","journal-title":"Microbiol Mol Biol Rev"},{"issue":"1","key":"5146_CR7","doi-asserted-by":"publisher","first-page":"2114739","DOI":"10.1080\/19490976.2022.2114739","volume":"14","author":"J Yang","year":"2022","unstructured":"Yang J, Li Y, Tang N, Li J, Zhou J, Lu S, et al. The human gut serves as a reservoir of hypervirulent Klebsiella pneumoniae. Gut Microbes. 2022;14(1):2114739. https:\/\/doi.org\/10.1080\/19490976.2022.2114739.","journal-title":"Gut Microbes"},{"issue":"9","key":"5146_CR8","doi-asserted-by":"publisher","first-page":"373","DOI":"10.1007\/s00203-024-04092-3","volume":"206","author":"MI Isidro-Coxca","year":"2024","unstructured":"Isidro-Coxca MI, Ortiz-Jim\u00e9nez S, Puente JL. Type 1 fimbria and P pili: regulatory mechanisms of the prototypical members of the chaperone-usher fimbrial family. Arch Microbiol. 2024;206(9):373. https:\/\/doi.org\/10.1007\/s00203-024-04092-3.","journal-title":"Arch Microbiol"},{"issue":"9","key":"5146_CR9","doi-asserted-by":"publisher","first-page":"4055","DOI":"10.1128\/iai.00494-08","volume":"76","author":"C Struve","year":"2008","unstructured":"Struve C, Bojer M, Krogfelt KA. Characterization of Klebsiella pneumoniae type 1 fimbriae by detection of phase variation during colonization and infection and impact on virulence. Infect Immun. 2008;76(9):4055\u201365. https:\/\/doi.org\/10.1128\/iai.00494-08.","journal-title":"Infect Immun"},{"issue":"5393","key":"5146_CR10","doi-asserted-by":"publisher","first-page":"1494","DOI":"10.1126\/science.282.5393.1494","volume":"282","author":"MA Mulvey","year":"1998","unstructured":"Mulvey MA, Lopez-Boado YS, Wilson CL, Roth R, Parks WC, Heuser J, et al. Induction and evasion of host defenses by type 1-piliated uropathogenic Escherichia coli. Science. 1998;282(5393):1494\u20137. https:\/\/doi.org\/10.1126\/science.282.5393.1494.","journal-title":"Science"},{"issue":"12","key":"5146_CR11","doi-asserted-by":"publisher","first-page":"2803","DOI":"10.1093\/emboj\/19.12.2803","volume":"19","author":"JJ Martinez","year":"2000","unstructured":"Martinez JJ, Mulvey MA, Schilling JD, Pinkner JS, Hultgren SJ. Type 1 pilus-mediated bacterial invasion of bladder epithelial cells. Embo j. 2000;19(12):2803\u201312. https:\/\/doi.org\/10.1093\/emboj\/19.12.2803.","journal-title":"Embo j"},{"issue":"7659","key":"5146_CR12","doi-asserted-by":"publisher","first-page":"528","DOI":"10.1038\/nature22972","volume":"546","author":"CN Spaulding","year":"2017","unstructured":"Spaulding CN, Klein RD, Ruer S, Kau AL, Schreiber HL, Cusumano ZT, et al. Selective depletion of uropathogenic E. coli from the gut by a FimH antagonist. Nature. 2017;546(7659):528\u201332. https:\/\/doi.org\/10.1038\/nature22972.","journal-title":"Nature"},{"issue":"15","key":"5146_CR13","doi-asserted-by":"publisher","first-page":"8922","DOI":"10.1073\/pnas.95.15.8922","volume":"95","author":"EV Sokurenko","year":"1998","unstructured":"Sokurenko EV, Chesnokova V, Dykhuizen DE, Ofek I, Wu XR, Krogfelt KA, et al. Pathogenic adaptation of Escherichia coli by natural variation of the FimH adhesin. Proc Natl Acad Sci U S A. 1998;95(15):8922\u20136. https:\/\/doi.org\/10.1073\/pnas.95.15.8922.","journal-title":"Proc Natl Acad Sci U S A"},{"issue":"52","key":"5146_CR14","doi-asserted-by":"publisher","first-page":"22439","DOI":"10.1073\/pnas.0902179106","volume":"106","author":"SL Chen","year":"2009","unstructured":"Chen SL, Hung CS, Pinkner JS, Walker JN, Cusumano CK, Li Z, et al. Positive selection identifies an in vivo role for FimH during urinary tract infection in addition to mannose binding. Proc Natl Acad Sci U S A. 2009;106(52):22439\u201344. https:\/\/doi.org\/10.1073\/pnas.0902179106.","journal-title":"Proc Natl Acad Sci U S A"},{"issue":"9","key":"5146_CR15","doi-asserted-by":"publisher","first-page":"e0017222","DOI":"10.1128\/jb.00172-22","volume":"204","author":"S Venkitapathi","year":"2022","unstructured":"Venkitapathi S, Wijesundara YH, Cornelius SA, Herbert FC, Gassensmith JJ, Zimmern PE, et al. Conserved FimK Truncation Coincides with Increased Expression of Type 3 Fimbriae and Cultured Bladder Epithelial Cell Association in Klebsiella quasipneumoniae. J Bacteriol. 2022;204(9):e0017222. https:\/\/doi.org\/10.1128\/jb.00172-22.","journal-title":"J Bacteriol"},{"issue":"11","key":"5146_CR16","doi-asserted-by":"publisher","first-page":"5016","DOI":"10.1128\/iai.00585-09","volume":"77","author":"C Struve","year":"2009","unstructured":"Struve C, Bojer M, Krogfelt KA. Identification of a conserved chromosomal region encoding Klebsiella pneumoniae type 1 and type 3 fimbriae and assessment of the role of fimbriae in pathogenicity. Infect Immun. 2009;77(11):5016\u201324. https:\/\/doi.org\/10.1128\/iai.00585-09.","journal-title":"Infect Immun"},{"issue":"14","key":"5146_CR17","doi-asserted-by":"publisher","first-page":"4492","DOI":"10.1128\/jb.00315-09","volume":"191","author":"KM Wu","year":"2009","unstructured":"Wu KM, Li LH, Yan JJ, Tsao N, Liao TL, Tsai HC, et al. Genome sequencing and comparative analysis of Klebsiella pneumoniae NTUH-K2044, a strain causing liver abscess and meningitis. J Bacteriol. 2009;191(14):4492\u2013501. https:\/\/doi.org\/10.1128\/jb.00315-09.","journal-title":"J Bacteriol"},{"issue":"7","key":"5146_CR18","doi-asserted-by":"publisher","first-page":"1841","DOI":"10.1128\/jb.00043-12","volume":"194","author":"P Liu","year":"2012","unstructured":"Liu P, Li P, Jiang X, Bi D, Xie Y, Tai C, et al. Complete genome sequence of Klebsiella pneumoniae subsp. pneumoniae HS11286, a multidrug-resistant strain isolated from human sputum. J Bacteriol. 2012;194(7):1841\u20132. https:\/\/doi.org\/10.1128\/jb.00043-12.","journal-title":"J Bacteriol"},{"issue":"12","key":"5146_CR19","doi-asserted-by":"publisher","first-page":"6640","DOI":"10.1073\/pnas.120163297","volume":"97","author":"KA Datsenko","year":"2000","unstructured":"Datsenko KA, Wanner BL. One-step inactivation of chromosomal genes in Escherichia coli K-12 using PCR products. Proc Natl Acad Sci U S A. 2000;97(12):6640\u20135. https:\/\/doi.org\/10.1073\/pnas.120163297.","journal-title":"Proc Natl Acad Sci U S A"},{"issue":"1","key":"5146_CR20","doi-asserted-by":"publisher","first-page":"13","DOI":"10.1186\/s12941-021-00418-x","volume":"20","author":"S Shadkam","year":"2021","unstructured":"Shadkam S, Goli HR, Mirzaei B, Gholami M, Ahanjan M. Correlation between antimicrobial resistance and biofilm formation capability among Klebsiella pneumoniae strains isolated from hospitalized patients in Iran. Ann Clin Microbiol Antimicrob. 2021;20(1):13. https:\/\/doi.org\/10.1186\/s12941-021-00418-x.","journal-title":"Ann Clin Microbiol Antimicrob"},{"issue":"2","key":"5146_CR21","doi-asserted-by":"publisher","first-page":"e1000303","DOI":"10.1371\/journal.ppat.1000303","volume":"5","author":"CM M\u00fcller","year":"2009","unstructured":"M\u00fcller CM, Aberg A, Strasevi\u00e7iene J, Emody L, Uhlin BE, Balsalobre C. Type 1 fimbriae, a colonization factor of uropathogenic Escherichia coli, are controlled by the metabolic sensor CRP-cAMP. PLoS Pathog. 2009;5(2):e1000303. https:\/\/doi.org\/10.1371\/journal.ppat.1000303.","journal-title":"PLoS Pathog"},{"issue":"7","key":"5146_CR22","doi-asserted-by":"publisher","first-page":"e100","DOI":"10.1371\/journal.ppat.0030100","volume":"3","author":"DS Eto","year":"2007","unstructured":"Eto DS, Jones TA, Sundsbak JL, Mulvey MA. Integrin-mediated host cell invasion by type 1-piliated uropathogenic Escherichia coli. PLoS Pathog. 2007;3(7):e100. https:\/\/doi.org\/10.1371\/journal.ppat.0030100.","journal-title":"PLoS Pathog"},{"issue":"10","key":"5146_CR23","doi-asserted-by":"publisher","first-page":"1887","DOI":"10.1161\/atvbaha.123.319119","volume":"43","author":"Y Yu","year":"2023","unstructured":"Yu Y, Leng Y, Song X, Mu J, Ma L, Yin L, et al. Extracellular Matrix Stiffness Regulates Microvascular Stability by Controlling Endothelial Paracrine Signaling to Determine Pericyte Fate. Arterioscler Thromb Vasc Biol. 2023;43(10):1887\u201399. https:\/\/doi.org\/10.1161\/atvbaha.123.319119.","journal-title":"Arterioscler Thromb Vasc Biol"},{"issue":"8016","key":"5146_CR24","doi-asserted-by":"publisher","first-page":"493","DOI":"10.1038\/s41586-024-07487-w","volume":"630","author":"J Abramson","year":"2024","unstructured":"Abramson J, Adler J, Dunger J, Evans R, Green T, Pritzel A, et al. Accurate structure prediction of biomolecular interactions with AlphaFold 3. Nature. 2024;630(8016):493\u2013500. https:\/\/doi.org\/10.1038\/s41586-024-07487-w.","journal-title":"Nature"},{"issue":"W1","key":"5146_CR25","doi-asserted-by":"publisher","first-page":"W365","DOI":"10.1093\/nar\/gkx407","volume":"45","author":"Y Yan","year":"2017","unstructured":"Yan Y, Zhang D, Zhou P, Li B, Huang SY. HDOCK: a web server for protein-protein and protein-DNA\/RNA docking based on a hybrid strategy. Nucleic Acids Res. 2017;45(W1):W365\u201373. https:\/\/doi.org\/10.1093\/nar\/gkx407.","journal-title":"Nucleic Acids Res"},{"issue":"1","key":"5146_CR26","doi-asserted-by":"publisher","first-page":"268","DOI":"10.1002\/pro.3781","volume":"29","author":"BHM Mooers","year":"2020","unstructured":"Mooers BHM. Shortcuts for faster image creation in PyMOL. Protein Sci. 2020;29(1):268\u201376. https:\/\/doi.org\/10.1002\/pro.3781.","journal-title":"Protein Sci"},{"key":"5146_CR27","doi-asserted-by":"publisher","first-page":"117394","DOI":"10.1016\/j.jep.2023.117394","volume":"321","author":"G Devi","year":"2024","unstructured":"Devi G, Gorki V, Walter NS, Sivangula S, Sobhia ME, Jachak S, et al. Exploring the efficacy of ethnomedicinal plants of Himalayan region against the malaria parasite. J Ethnopharmacol. 2024;321:117394. https:\/\/doi.org\/10.1016\/j.jep.2023.117394.","journal-title":"J Ethnopharmacol"},{"issue":"1","key":"5146_CR28","doi-asserted-by":"publisher","first-page":"168","DOI":"10.1186\/s12943-024-02086-9","volume":"23","author":"J Zhou","year":"2024","unstructured":"Zhou J, Song Q, Li H, Han Y, Pu Y, Li L, et al. Targeting circ-0034880-enriched tumor extracellular vesicles to impede SPP1(high)CD206(+) pro-tumor macrophages mediated pre-metastatic niche formation in colorectal cancer liver metastasis. Mol Cancer. 2024;23(1):168. https:\/\/doi.org\/10.1186\/s12943-024-02086-9.","journal-title":"Mol Cancer"},{"issue":"W1","key":"5146_CR29","doi-asserted-by":"publisher","first-page":"W420","DOI":"10.1093\/nar\/gkac373","volume":"50","author":"Y Xie","year":"2022","unstructured":"Xie Y, Li H, Luo X, Li H, Gao Q, Zhang L, et al. IBS 2.0: an upgraded illustrator for the visualization of biological sequences. Nucleic Acids Res. 2022;50(W1):W420\u20136. https:\/\/doi.org\/10.1093\/nar\/gkac373.","journal-title":"Nucleic Acids Res"},{"issue":"3","key":"5146_CR30","doi-asserted-by":"publisher","first-page":"467","DOI":"10.1007\/s10096-016-2820-8","volume":"36","author":"T Feenstra","year":"2017","unstructured":"Feenstra T, Th\u00f8gersen MS, Wieser E, Peschel A, Ball MJ, Brandes R, et al. Adhesion of Escherichia coli under flow conditions reveals potential novel effects of FimH mutations. Eur J Clin Microbiol Infect Dis. 2017;36(3):467\u201378. https:\/\/doi.org\/10.1007\/s10096-016-2820-8.","journal-title":"Eur J Clin Microbiol Infect Dis"},{"issue":"8","key":"5146_CR31","doi-asserted-by":"publisher","first-page":"991","DOI":"10.2217\/fmb.12.74","volume":"7","author":"CN Murphy","year":"2012","unstructured":"Murphy CN, Clegg S. Klebsiella pneumoniae and type 3 fimbriae: nosocomial infection, regulation and biofilm formation. Future Microbiol. 2012;7(8):991\u20131002. https:\/\/doi.org\/10.2217\/fmb.12.74.","journal-title":"Future Microbiol"},{"issue":"3","key":"5146_CR32","doi-asserted-by":"publisher","first-page":"542","DOI":"10.1016\/j.jmb.2013.10.029","volume":"426","author":"MJ Walczak","year":"2014","unstructured":"Walczak MJ, Puorger C, Glockshuber R, Wider G. Intramolecular donor strand complementation in the E. coli type 1 pilus subunit FimA explains the existence of FimA monomers as off-pathway products of pilus assembly that inhibit host cell apoptosis. J Mol Biol. 2014;426(3):542\u20139. https:\/\/doi.org\/10.1016\/j.jmb.2013.10.029.","journal-title":"J Mol Biol"},{"issue":"3","key":"5146_CR33","doi-asserted-by":"publisher","first-page":"520","DOI":"10.1016\/j.jmb.2011.07.044","volume":"412","author":"C Puorger","year":"2011","unstructured":"Puorger C, Vetsch M, Wider G, Glockshuber R. Structure, folding and stability of FimA, the main structural subunit of type 1 pili from uropathogenic Escherichia coli strains. J Mol Biol. 2011;412(3):520\u201335. https:\/\/doi.org\/10.1016\/j.jmb.2011.07.044.","journal-title":"J Mol Biol"},{"issue":"6","key":"5146_CR34","doi-asserted-by":"publisher","first-page":"673","DOI":"10.1016\/s0092-8674(02)00971-6","volume":"110","author":"RO Hynes","year":"2002","unstructured":"Hynes RO. Integrins: bidirectional, allosteric signaling machines. Cell. 2002;110(6):673\u201387. https:\/\/doi.org\/10.1016\/s0092-8674(02)00971-6.","journal-title":"Cell"},{"key":"5146_CR35","doi-asserted-by":"publisher","first-page":"1367898","DOI":"10.3389\/fmicb.2024.1367898","volume":"15","author":"GV De Gaetano","year":"2024","unstructured":"De Gaetano GV, Lentini G, Coppolino F, Fam\u00e0 A, Pietrocola G, Beninati C. Engagement of \u03b1(3)\u03b2(1) and \u03b1(2)\u03b2(1) integrins by hypervirulent Streptococcus agalactiae in invasion of polarized enterocytes. Front Microbiol. 2024;15:1367898. https:\/\/doi.org\/10.3389\/fmicb.2024.1367898.","journal-title":"Front Microbiol"},{"issue":"1","key":"5146_CR36","doi-asserted-by":"publisher","first-page":"6040","DOI":"10.1038\/s41467-023-41721-9","volume":"14","author":"F Homma","year":"2023","unstructured":"Homma F, Huang J, van der Hoorn RAL. AlphaFold-Multimer predicts cross-kingdom interactions at the plant-pathogen interface. Nat Commun. 2023;14(1):6040. https:\/\/doi.org\/10.1038\/s41467-023-41721-9.","journal-title":"Nat Commun"},{"key":"5146_CR37","doi-asserted-by":"publisher","unstructured":"Day CJ, Lo AW, Hartley-Tassell LE, Argente MP, Poole J, King NP, et al. Discovery of Bacterial Fimbria-Glycan Interactions Using Whole-Cell Recombinant Escherichia coli Expression. mBio. 2021;12(1). https:\/\/doi.org\/10.1128\/mBio.03664-20.","DOI":"10.1128\/mBio.03664-20"},{"issue":"6","key":"5146_CR38","doi-asserted-by":"publisher","first-page":"e0240022","DOI":"10.1128\/spectrum.02400-22","volume":"10","author":"S Wang","year":"2022","unstructured":"Wang S, Ding Q, Zhang Y, Zhang A, Wang Q, Wang R, et al. Evolution of Virulence, Fitness, and Carbapenem Resistance Transmission in ST23 Hypervirulent Klebsiella pneumoniae with the Capsular Polysaccharide Synthesis Gene wcaJ Inserted via Insertion Sequence Elements. Microbiol Spectr. 2022;10(6):e0240022. https:\/\/doi.org\/10.1128\/spectrum.02400-22.","journal-title":"Microbiol Spectr"},{"issue":"3","key":"5146_CR39","doi-asserted-by":"publisher","first-page":"470","DOI":"10.1093\/infdis\/jiy531","volume":"219","author":"R Liu","year":"2019","unstructured":"Liu R, Wu C, Li L, Chi F, Zhang T, Xu Y, et al. CD48 and \u03b17 Nicotinic Acetylcholine Receptor Synergistically Regulate FimH-Mediated Escherichia coli K1 Penetration and Neutrophil Transmigration Across Human Brain Microvascular Endothelial Cells. J Infect Dis. 2019;219(3):470\u20139. https:\/\/doi.org\/10.1093\/infdis\/jiy531.","journal-title":"J Infect Dis"},{"key":"5146_CR40","doi-asserted-by":"publisher","unstructured":"Kallas P, Haugen HJ, Gadegaard N, Stormonth-Darling J, Hulander M, Andersson M, et al. Adhesion of Escherichia Coli to Nanostructured Surfaces and the Role of Type 1 Fimbriae. Nanomaterials (Basel). 2020;10(11). https:\/\/doi.org\/10.3390\/nano10112247.","DOI":"10.3390\/nano10112247"},{"issue":"1","key":"5146_CR41","doi-asserted-by":"publisher","first-page":"269","DOI":"10.1007\/s00441-009-0834-6","volume":"339","author":"M Barczyk","year":"2010","unstructured":"Barczyk M, Carracedo S, Gullberg D, Integrins. Cell Tissue Res. 2010;339(1):269\u201380. https:\/\/doi.org\/10.1007\/s00441-009-0834-6.","journal-title":"Cell Tissue Res"},{"key":"5146_CR42","doi-asserted-by":"publisher","unstructured":"Zhou Y, Zhou Z, Zheng L, Gong Z, Li Y, Jin Y, et al. Urinary Tract Infections Caused by Uropathogenic Escherichia coli: Mechanisms of Infection and Treatment Options. Int J Mol Sci. 2023;24(13). https:\/\/doi.org\/10.3390\/ijms241310537.","DOI":"10.3390\/ijms241310537"},{"issue":"5","key":"5146_CR43","doi-asserted-by":"publisher","first-page":"489","DOI":"10.1016\/j.ceb.2005.08.013","volume":"17","author":"S Sousa","year":"2005","unstructured":"Sousa S, Lecuit M, Cossart P. Microbial strategies to target, cross or disrupt epithelia. Curr Opin Cell Biol. 2005;17(5):489\u201398. https:\/\/doi.org\/10.1016\/j.ceb.2005.08.013.","journal-title":"Curr Opin Cell Biol"},{"issue":"11","key":"5146_CR44","doi-asserted-by":"publisher","first-page":"799","DOI":"10.1038\/nri2653","volume":"9","author":"JR Turner","year":"2009","unstructured":"Turner JR. Intestinal mucosal barrier function in health and disease. Nat Rev Immunol. 2009;9(11):799\u2013809. https:\/\/doi.org\/10.1038\/nri2653.","journal-title":"Nat Rev Immunol"},{"issue":"5","key":"5146_CR45","doi-asserted-by":"publisher","first-page":"861","DOI":"10.1016\/0092-8674(90)90099-z","volume":"60","author":"RR Isberg","year":"1990","unstructured":"Isberg RR, Leong JM. Multiple beta 1 chain integrins are receptors for invasin, a protein that promotes bacterial penetration into mammalian cells. Cell. 1990;60(5):861\u201371. https:\/\/doi.org\/10.1016\/0092-8674(90)90099-z.","journal-title":"Cell"},{"issue":"4","key":"5146_CR46","doi-asserted-by":"publisher","first-page":"715","DOI":"10.1016\/j.cell.2006.02.012","volume":"124","author":"J Pizarro-Cerd\u00e1","year":"2006","unstructured":"Pizarro-Cerd\u00e1 J, Cossart P. Bacterial adhesion and entry into host cells. Cell. 2006;124(4):715\u201327. https:\/\/doi.org\/10.1016\/j.cell.2006.02.012.","journal-title":"Cell"},{"issue":"7","key":"5146_CR47","doi-asserted-by":"publisher","first-page":"4404","DOI":"10.1128\/iai.73.7.4404-4409.2005","volume":"73","author":"T Tenenbaum","year":"2005","unstructured":"Tenenbaum T, Bloier C, Adam R, Reinscheid DJ, Schroten H. Adherence to and invasion of human brain microvascular endothelial cells are promoted by fibrinogen-binding protein FbsA of Streptococcus agalactiae. Infect Immun. 2005;73(7):4404\u20139. https:\/\/doi.org\/10.1128\/iai.73.7.4404-4409.2005.","journal-title":"Infect Immun"},{"key":"5146_CR48","doi-asserted-by":"publisher","first-page":"462","DOI":"10.1038\/ncomms1474","volume":"2","author":"A Banerjee","year":"2011","unstructured":"Banerjee A, Kim BJ, Carmona EM, Cutting AS, Gurney MA, Carlos C, et al. Bacterial Pili exploit integrin machinery to promote immune activation and efficient blood-brain barrier penetration. Nat Commun. 2011;2:462. https:\/\/doi.org\/10.1038\/ncomms1474.","journal-title":"Nat Commun"},{"issue":"2","key":"5146_CR49","doi-asserted-by":"publisher","first-page":"417","DOI":"10.1128\/iai.29.2.417-424.1980","volume":"29","author":"Z Bar-Shavit","year":"1980","unstructured":"Bar-Shavit Z, Goldman R, Ofek I, Sharon N, Mirelman D. Mannose-binding activity of Escherichia coli: a determinant of attachment and ingestion of the bacteria by macrophages. Infect Immun. 1980;29(2):417\u201324. https:\/\/doi.org\/10.1128\/iai.29.2.417-424.1980.","journal-title":"Infect Immun"},{"issue":"18","key":"5146_CR50","doi-asserted-by":"publisher","first-page":"9827","DOI":"10.1073\/pnas.93.18.9827","volume":"93","author":"I Connell","year":"1996","unstructured":"Connell I, Agace W, Klemm P, Schembri M, M\u0103rild S, Svanborg C. Type 1 fimbrial expression enhances Escherichia coli virulence for the urinary tract. Proc Natl Acad Sci U S A. 1996;93(18):9827\u201332. https:\/\/doi.org\/10.1073\/pnas.93.18.9827.","journal-title":"Proc Natl Acad Sci U S A"},{"key":"5146_CR51","doi-asserted-by":"publisher","first-page":"8754","DOI":"10.1038\/ncomms9754","volume":"6","author":"M Yue","year":"2015","unstructured":"Yue M, Han X, De Masi L, Zhu C, Ma X, Zhang J, et al. Allelic variation contributes to bacterial host specificity. Nat Commun. 2015;6:8754. https:\/\/doi.org\/10.1038\/ncomms9754.","journal-title":"Nat Commun"},{"issue":"3","key":"5146_CR52","doi-asserted-by":"publisher","first-page":"407","DOI":"10.1016\/s0092-8674(02)00628-1","volume":"108","author":"SM Akula","year":"2002","unstructured":"Akula SM, Pramod NP, Wang FZ, Chandran B. Integrin alpha3beta1 (CD 49c\/29) is a cellular receptor for Kaposi\u2019s sarcoma-associated herpesvirus (KSHV\/HHV-8) entry into the target cells. Cell. 2002;108(3):407\u201319. https:\/\/doi.org\/10.1016\/s0092-8674(02)00628-1.","journal-title":"Cell"},{"key":"5146_CR53","doi-asserted-by":"publisher","first-page":"179","DOI":"10.1186\/1471-2180-10-179","volume":"10","author":"C Schroll","year":"2010","unstructured":"Schroll C, Barken KB, Krogfelt KA, Struve C. Role of type 1 and type 3 fimbriae in Klebsiella pneumoniae biofilm formation. BMC Microbiol. 2010;10:179. https:\/\/doi.org\/10.1186\/1471-2180-10-179.","journal-title":"BMC Microbiol"},{"issue":"4","key":"5146_CR54","doi-asserted-by":"publisher","first-page":"1485","DOI":"10.1128\/aac.01275-10","volume":"55","author":"YK Tsai","year":"2011","unstructured":"Tsai YK, Fung CP, Lin JC, Chen JH, Chang FY, Chen TL, et al. Klebsiella pneumoniae outer membrane porins OmpK35 and OmpK36 play roles in both antimicrobial resistance and virulence. Antimicrob Agents Chemother. 2011;55(4):1485\u201393. https:\/\/doi.org\/10.1128\/aac.01275-10.","journal-title":"Antimicrob Agents Chemother"},{"key":"5146_CR55","doi-asserted-by":"publisher","first-page":"128262","DOI":"10.1016\/j.micres.2025.128262","volume":"299","author":"W Zeng","year":"2025","unstructured":"Zeng W, Sun Y, Zhang J, Yao Z, Cao J, Hu P, et al. Outer membrane protein A mediates Klebsiella pneumoniae penetration of the blood-brain barrier and induces bacterial meningitis. Microbiol Res. 2025;299:128262. https:\/\/doi.org\/10.1016\/j.micres.2025.128262.","journal-title":"Microbiol Res"},{"issue":"4","key":"5146_CR56","doi-asserted-by":"publisher","first-page":"375","DOI":"10.1093\/intimm\/dxm003","volume":"19","author":"R Graveline","year":"2007","unstructured":"Graveline R, Segura M, Radzioch D, Gottschalk M. TLR2-dependent recognition of Streptococcus suis is modulated by the presence of capsular polysaccharide which modifies macrophage responsiveness. Int Immunol. 2007;19(4):375\u201389. https:\/\/doi.org\/10.1093\/intimm\/dxm003.","journal-title":"Int Immunol"}],"container-title":["BMC Microbiology"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/link.springer.com\/article\/10.1186\/s12866-026-05146-w","content-type":"text\/html","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/link.springer.com\/content\/pdf\/10.1186\/s12866-026-05146-w.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/link.springer.com\/content\/pdf\/10.1186\/s12866-026-05146-w.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,7,9]],"date-time":"2026-07-09T04:13:07Z","timestamp":1783570387000},"score":1,"resource":{"primary":{"URL":"https:\/\/link.springer.com\/10.1186\/s12866-026-05146-w"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,5,23]]},"references-count":56,"journal-issue":{"issue":"1","published-online":{"date-parts":[[2026,12]]}},"alternative-id":["5146"],"URL":"https:\/\/doi.org\/10.1186\/s12866-026-05146-w","relation":{},"ISSN":["1471-2180"],"issn-type":[{"value":"1471-2180","type":"electronic"}],"subject":[],"published":{"date-parts":[[2026,5,23]]},"assertion":[{"value":"14 November 2025","order":1,"name":"received","label":"Received","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"6 May 2026","order":2,"name":"accepted","label":"Accepted","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"23 May 2026","order":3,"name":"first_online","label":"First Online","group":{"name":"ArticleHistory","label":"Article History"}},{"order":1,"name":"Ethics","group":{"name":"EthicsHeading","label":"Declarations"}},{"value":"This study was reviewed and approved by the Ethics Committee of Huashan Hospital, affiliated with Fudan University (Approval No. 2021\u2009\u2212\u2009484). As the research exclusively utilized anonymous bacterial strains\u2014classified as anonymous recorded materials in accordance with Item 34 of the Measures for Ethical Review of Biomedical Research Involving Humans (National Health and Family Planning Commission of China, 2016)\u2014the requirement for written informed consent was waived by the ethics review board in compliance with both national legislation and institutional policies. The work involved cell-based experiments, bacterial strain construction, protein-binding predictions, and in vitro affinity measurements. No human participants or vertebrate animals were directly involved in these experiments. All methods were performed in accordance with the relevant guidelines and regulations.","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":"609"}}