{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,5,21]],"date-time":"2026-05-21T18:18:19Z","timestamp":1779387499199,"version":"3.53.1"},"update-to":[{"DOI":"10.1371\/journal.ppat.1013584","type":"new_version","label":"New version","source":"publisher","updated":{"date-parts":[[2026,5,21]],"date-time":"2026-05-21T00:00:00Z","timestamp":1779321600000}}],"reference-count":96,"publisher":"Public Library of Science (PLoS)","issue":"5","license":[{"start":{"date-parts":[[2026,5,18]],"date-time":"2026-05-18T00:00:00Z","timestamp":1779062400000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/publicdomain\/zero\/1.0\/"}],"funder":[{"DOI":"10.13039\/100007917","name":"Agricultural Research Service","doi-asserted-by":"publisher","award":["This research was supported in part by the USDA-ARS CRIS project 5030-32000-225-00D."],"award-info":[{"award-number":["This research was supported in part by the USDA-ARS CRIS project 5030-32000-225-00D."]}],"id":[{"id":"10.13039\/100007917","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/100012651","name":"College of Veterinary Medicine, University of Florida","doi-asserted-by":"publisher","id":[{"id":"10.13039\/100012651","id-type":"DOI","asserted-by":"publisher"}]}],"content-domain":{"domain":["www.plospathogens.org"],"crossmark-restriction":false},"short-container-title":["PLoS Pathog"],"abstract":"<jats:p>\n                    Shiga toxin-producing\n                    <jats:italic>Escherichia coli<\/jats:italic>\n                    O157:H7 (O157), a foodborne human pathogen, persists at the rectoanal junction (RAJ) of the bovine intestinal tract, in asymptomatic cattle reservoirs. Identifying mechanisms used by O157 for initial adherence before persistence at the RAJ could help develop effective O157 control modalities. We recently established the role of carbon starvation-inducible lipoprotein (Slp) in initial adherence of O157 to Caco-2 cells, with the human polymeric immunoglobulin receptor (pIgR) protein as the Slp-receptor. Here, we evaluated the role of Slp in O157 adherence to the bovine RAJ using the RAJ squamous epithelial (RSE) cell- and RAJ-\n                    <jats:italic>in vitro<\/jats:italic>\n                    Organ Culture (IVOC)- adherence assays. The wild-type O157 strain EDL932 (EDL932-WT), it\u2019s isogenic\n                    <jats:italic>slp<\/jats:italic>\n                    deletion mutant (EDL932 \u0394\n                    <jats:italic>slp<\/jats:italic>\n                    ), and the\n                    <jats:italic>slp<\/jats:italic>\n                    complemented mutant (EDL932 \u0394\n                    <jats:italic>slp<\/jats:italic>\n                    -p:\n                    <jats:italic>slp<\/jats:italic>\n                    ), were tested with no bacteria controls. Adherence was verified by culture and immunofluorescence (IF) staining of O157. Tissue integrity was determined using nuclear\/cell staining dyes and histopathological examination. All test strains adhered in a diffuse-moderate pattern on RSE cells. However, differential adherence was observed on the RAJ-IVOC with the strains preferentially adhering to the columnar cells. Additionally, EDL932-WT and EDL932 \u0394\n                    <jats:italic>slp<\/jats:italic>\n                    -p:\n                    <jats:italic>slp<\/jats:italic>\n                    strains adhered in slightly greater numbers than the EDL932 \u0394\n                    <jats:italic>slp<\/jats:italic>\n                    strain to the RAJ-IVOC, causing disruptions primarily in the columnar region of otherwise intact RAJ-IVOC tissues. Interestingly, pIgR was also predominantly detected by IF microscopy and RNAscope\n                    <jats:italic>in situ<\/jats:italic>\n                    hybridization at the columnar region of the RAJ-IVOC tissue.\n                    <jats:italic>In silico<\/jats:italic>\n                    modeling demonstrated the possibility of a bovine pIgR- bacterial Slp interaction. Hence, our observations support the role for Slp in the initial adherence of O157 to the columnar cells at the bovine RAJ, unlike the squamous cells where the loss of\n                    <jats:italic>slp<\/jats:italic>\n                    did not impact attachment. In addition, a possible mucosal immune-interference resulting from the bovine pIgR-Slp interaction may contribute towards long-term O157 colonization of cattle.\n                  <\/jats:p>","DOI":"10.1371\/journal.ppat.1013584","type":"journal-article","created":{"date-parts":[[2026,5,18]],"date-time":"2026-05-18T17:42:21Z","timestamp":1779126141000},"page":"e1013584","update-policy":"https:\/\/doi.org\/10.1371\/journal.ppat.corrections_policy","source":"Crossref","is-referenced-by-count":0,"title":["The carbon starvation-inducible lipoprotein (Slp) influences differential adherence of Escherichia coli O157:H7 at the bovine rectoanal junction"],"prefix":"10.1371","volume":"22","author":[{"ORCID":"https:\/\/orcid.org\/0000-0001-6750-7867","authenticated-orcid":true,"given":"Indira T.","family":"Kudva","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Erika N.","family":"Biernbaum","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Eric D.","family":"Cassmann","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Mitchell V.","family":"Palmer","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Lekshmi K.","family":"Edison","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Jessy","family":"Castellanos-Gell","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Subhashinie","family":"Kariyawasam","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"340","published-online":{"date-parts":[[2026,5,18]]},"reference":[{"issue":"2","key":"ppat.1013584.ref001","doi-asserted-by":"crossref","first-page":"35","DOI":"10.14252\/foodsafetyfscj.2016029","article-title":"Shiga Toxin (Verotoxin)-producing Escherichia coli and Foodborne Disease: A Review","volume":"5","author":"J Terajima","year":"2017","journal-title":"Food Saf (Tokyo)"},{"key":"ppat.1013584.ref002","doi-asserted-by":"crossref","first-page":"273","DOI":"10.3389\/fcimb.2020.00273","article-title":"Recent Updates on Outbreaks of Shiga Toxin-Producing Escherichia coli and Its Potential Reservoirs","volume":"10","author":"J-S Kim","year":"2020","journal-title":"Front Cell Infect Microbiol"},{"issue":"6","key":"ppat.1013584.ref003","doi-asserted-by":"crossref","first-page":"877","DOI":"10.3390\/microorganisms8060877","article-title":"An Overview of the Elusive Passenger in the Gastrointestinal Tract of Cattle: The Shiga Toxin Producing Escherichia coli","volume":"8","author":"P Sapountzis","year":"2020","journal-title":"Microorganisms"},{"key":"ppat.1013584.ref004","volume-title":"Escherichia coli O157:H7 and other Shiga toxin-producing E. coli strains","author":"JB Kaper","year":"1998"},{"issue":"5","key":"ppat.1013584.ref005","doi-asserted-by":"crossref","DOI":"10.1128\/microbiolspec.EHEC-0009-2013","article-title":"Enterohemorrhagic Escherichia coli Pathogenesis and the Host Response","volume":"2","author":"D Karpman","year":"2014","journal-title":"Microbiol Spectr"},{"issue":"5","key":"ppat.1013584.ref006","doi-asserted-by":"crossref","DOI":"10.1128\/microbiolspec.EHEC-0002-2013","article-title":"Risk Factors for Shiga Toxin-Producing Escherichia coli-Associated Human Diseases","volume":"2","author":"M Rivas","year":"2014","journal-title":"Microbiol Spectr"},{"issue":"15","key":"ppat.1013584.ref007","doi-asserted-by":"crossref","first-page":"1402","DOI":"10.1056\/NEJMra2108739","article-title":"Shiga Toxin-Producing Escherichia coli and the Hemolytic-Uremic Syndrome","volume":"389","author":"SB Freedman","year":"2023","journal-title":"N Engl J Med"},{"issue":"10","key":"ppat.1013584.ref008","doi-asserted-by":"crossref","first-page":"2009","DOI":"10.3390\/microorganisms12102009","article-title":"Host-Pathogen Interactions during Shiga Toxin-Producing Escherichia coli Adherence and Colonization in the Bovine Gut: A Comprehensive Review","volume":"12","author":"LK Edison","year":"2024","journal-title":"Microorganisms"},{"issue":"3","key":"ppat.1013584.ref009","doi-asserted-by":"crossref","first-page":"1505","DOI":"10.1128\/IAI.71.3.1505-1512.2003","article-title":"Lymphoid follicle-dense mucosa at the terminal rectum is the principal site of colonization of enterohemorrhagic Escherichia coli O157:H7 in the bovine host","volume":"71","author":"SW Naylor","year":"2003","journal-title":"Infect Immun"},{"issue":"8","key":"ppat.1013584.ref010","first-page":"248","article-title":"Colonisation of Escherichia coli O157:H7 on squamous epithelial cells at the rectal-anal junction","volume":"155","author":"JF Pohlenz","year":"2004","journal-title":"Vet Rec"},{"issue":"4","key":"ppat.1013584.ref011","doi-asserted-by":"crossref","first-page":"1380","DOI":"10.1128\/AEM.02242-06","article-title":"Escherichia coli O157:H7 colonization at the rectoanal junction of long-duration culture-positive cattle","volume":"73","author":"JY Lim","year":"2007","journal-title":"Appl Environ Microbiol"},{"issue":"15","key":"ppat.1013584.ref012","doi-asserted-by":"crossref","first-page":"5278","DOI":"10.1128\/AEM.00400-10","article-title":"Distribution of Shiga-toxigenic Escherichia coli O157 in the gastrointestinal tract of naturally O157-shedding cattle at necropsy","volume":"76","author":"JE Keen","year":"2010","journal-title":"Appl Environ Microbiol"},{"issue":"3","key":"ppat.1013584.ref013","doi-asserted-by":"crossref","first-page":"365","DOI":"10.1007\/s00441-005-1080-1","article-title":"Phenotypic and functional characterisation of follicle-associated epithelium of rectal lymphoid tissue","volume":"321","author":"A Mahajan","year":"2005","journal-title":"Cell Tissue Res"},{"issue":"5","key":"ppat.1013584.ref014","doi-asserted-by":"crossref","first-page":"1283","DOI":"10.1111\/j.1365-2672.2011.05139.x","article-title":"Bovine recto-anal junction squamous epithelial (RSE) cell adhesion assay for studying Escherichia coli O157 adherence","volume":"111","author":"IT Kudva","year":"2011","journal-title":"J Appl Microbiol"},{"key":"ppat.1013584.ref015","doi-asserted-by":"crossref","first-page":"103","DOI":"10.1186\/1471-2180-12-103","article-title":"Proteins other than the locus of enterocyte effacement-encoded proteins contribute to Escherichia coli O157:H7 adherence to bovine rectoanal junction stratified squamous epithelial cells","volume":"12","author":"IT Kudva","year":"2012","journal-title":"BMC Microbiol"},{"issue":"4","key":"ppat.1013584.ref016","doi-asserted-by":"crossref","first-page":"375","DOI":"10.1089\/fpd.2012.1382","article-title":"Adherence of non-O157 Shiga toxin-producing Escherichia coli to bovine recto-anal junction squamous epithelial cells appears to be mediated by mechanisms distinct from those used by O157","volume":"10","author":"IT Kudva","year":"2013","journal-title":"Foodborne Pathog Dis"},{"issue":"11","key":"ppat.1013584.ref017","doi-asserted-by":"crossref","first-page":"1829","DOI":"10.1002\/pmic.201400432","article-title":"The Escherichia coli O157:H7 cattle immunoproteome includes outer membrane protein A (OmpA), a modulator of adherence to bovine rectoanal junction squamous epithelial (RSE) cells","volume":"15","author":"IT Kudva","year":"2015","journal-title":"Proteomics"},{"issue":"1","key":"ppat.1013584.ref018","doi-asserted-by":"crossref","DOI":"10.1128\/AEM.02594-16","article-title":"Curli Temper Adherence of Escherichia coli O157:H7 to Squamous Epithelial Cells from the Bovine Recto-Anal Junction in a Strain-Dependent Manner","volume":"83","author":"IT Kudva","year":"2016","journal-title":"Appl Environ Microbiol"},{"key":"ppat.1013584.ref019","first-page":"2368154","article-title":"Supershed Escherichia coli O157:H7 Has Potential for Increased Persistence on the Rectoanal Junction Squamous Epithelial Cells and Antibiotic Resistance","volume":"2020","author":"RA Mir","year":"2020","journal-title":"Int J Microbiol"},{"issue":"6","key":"ppat.1013584.ref020","doi-asserted-by":"crossref","first-page":"1059","DOI":"10.1111\/j.1365-2958.1994.tb00383.x","article-title":"Characterization of the carbon starvation-inducible and stationary phase-inducible gene slp encoding an outer membrane lipoprotein in Escherichia coli","volume":"11","author":"DM Alexander","year":"1994","journal-title":"Mol Microbiol"},{"issue":"1","key":"ppat.1013584.ref021","doi-asserted-by":"crossref","first-page":"51","DOI":"10.1111\/j.1574-6968.2000.tb09401.x","article-title":"Purification and analysis of expression of the stationary phase-inducible slp lipoprotein in Escherichia coli: role of the Mar system","volume":"193","author":"GP Price","year":"2000","journal-title":"FEMS Microbiol Lett"},{"issue":"41","key":"ppat.1013584.ref022","doi-asserted-by":"crossref","first-page":"34409","DOI":"10.1074\/jbc.M506479200","article-title":"Protein complexes of the Escherichia coli cell envelope","volume":"280","author":"F Stenberg","year":"2005","journal-title":"J Biol Chem"},{"issue":"2","key":"ppat.1013584.ref023","first-page":"301","article-title":"Acid stress response in enteropathogenic gammaproteobacteria: an aptitude for survival","volume":"88","author":"B Zhao","year":"2010","journal-title":"Biochem Cell Biol"},{"issue":"4","key":"ppat.1013584.ref024","doi-asserted-by":"crossref","first-page":"1004","DOI":"10.1128\/AEM.07033-11","article-title":"Evolutionary silence of the acid chaperone protein HdeB in enterohemorrhagic Escherichia coli O157:H7","volume":"78","author":"MQ Carter","year":"2012","journal-title":"Appl Environ Microbiol"},{"issue":"6","key":"ppat.1013584.ref025","doi-asserted-by":"crossref","DOI":"10.1371\/journal.pone.0216791","article-title":"The Escherichia coli O157:H7 carbon starvation-inducible lipoprotein Slp contributes to initial adherence in vitro via the human polymeric immunoglobulin receptor","volume":"14","author":"C Fedorchuk","year":"2019","journal-title":"PLoS One"},{"issue":"1","key":"ppat.1013584.ref026","doi-asserted-by":"crossref","first-page":"253","DOI":"10.1046\/j.1365-2958.2003.03432.x","article-title":"Global gene expression in Escherichia coli biofilms","volume":"48","author":"MA Schembri","year":"2003","journal-title":"Mol Microbiol"},{"key":"ppat.1013584.ref027","article-title":"Carbon starvation-induced lipoprotein Slp directs the synthesis of catalase and expression of OxyR regulator to protect against hydrogen peroxide stress in Escherichia coli","author":"X Li","year":"2018","journal-title":"bioRxiv"},{"issue":"23","key":"ppat.1013584.ref028","doi-asserted-by":"crossref","first-page":"12745","DOI":"10.1128\/JVI.02495-13","article-title":"The protein interaction network of bacteriophage lambda with its host, Escherichia coli","volume":"87","author":"S Blasche","year":"2013","journal-title":"J Virol"},{"issue":"1","key":"ppat.1013584.ref029","doi-asserted-by":"crossref","first-page":"376","DOI":"10.4049\/jimmunol.175.1.376","article-title":"Regulation of the polymeric Ig receptor by signaling through TLRs 3 and 4: linking innate and adaptive immune responses","volume":"175","author":"TA Schneeman","year":"2005","journal-title":"J Immunol"},{"key":"ppat.1013584.ref030","doi-asserted-by":"crossref","first-page":"64","DOI":"10.1111\/j.0105-2896.2005.00290.x","article-title":"Mucosal immunoglobulins","volume":"206","author":"JM Woof","year":"2005","journal-title":"Immunol Rev"},{"issue":"2","key":"ppat.1013584.ref031","doi-asserted-by":"crossref","first-page":"148","DOI":"10.1016\/j.bbrc.2009.01.176","article-title":"Human Fcalpha\/muR and pIgR distribute differently in intestinal tissues","volume":"381","author":"R Wang","year":"2009","journal-title":"Biochem Biophys Res Commun"},{"issue":"5","key":"ppat.1013584.ref032","doi-asserted-by":"crossref","first-page":"237","DOI":"10.3390\/v10050237","article-title":"The Role of the Polymeric Immunoglobulin Receptor and Secretory Immunoglobulins during Mucosal Infection and Immunity","volume":"10","author":"H Turula","year":"2018","journal-title":"Viruses"},{"issue":"1","key":"ppat.1013584.ref033","doi-asserted-by":"crossref","first-page":"68","DOI":"10.1038\/s41419-023-05589-3","article-title":"Polymeric immunoglobulin receptor deficiency exacerbates autoimmune hepatitis by inducing intestinal dysbiosis and barrier dysfunction","volume":"14","author":"H Lin","year":"2023","journal-title":"Cell Death Dis"},{"issue":"2","key":"ppat.1013584.ref034","doi-asserted-by":"crossref","first-page":"397","DOI":"10.1074\/mcp.M113.035600","article-title":"Analysis of the human tissue-specific expression by genome-wide integration of transcriptomics and antibody-based proteomics","volume":"13","author":"L Fagerberg","year":"2014","journal-title":"Mol Cell Proteomics"},{"issue":"1","key":"ppat.1013584.ref035","doi-asserted-by":"crossref","DOI":"10.1016\/S0960-9822(00)00041-5","article-title":"Polymeric Ig receptor: defender of the fort or Trojan horse?","volume":"11","author":"CS Kaetzel","year":"2001","journal-title":"Current Biology"},{"issue":"46","key":"ppat.1013584.ref036","doi-asserted-by":"crossref","first-page":"35615","DOI":"10.1074\/jbc.M110.172999","article-title":"Polymeric immunoglobulin receptor-mediated invasion of Streptococcus pneumoniae into host cells requires a coordinate signaling of SRC family of protein-tyrosine kinases, ERK, and c-Jun N-terminal kinase","volume":"285","author":"V Agarwal","year":"2010","journal-title":"J Biol Chem"},{"issue":"7","key":"ppat.1013584.ref037","doi-asserted-by":"crossref","first-page":"2380","DOI":"10.1128\/AEM.02262-06","article-title":"Expression profiles of bovine genes in the rectoanal junction mucosa during colonization with Escherichia coli O157:H7","volume":"73","author":"J Li","year":"2007","journal-title":"Appl Environ Microbiol"},{"issue":"10","key":"ppat.1013584.ref038","doi-asserted-by":"crossref","first-page":"2562","DOI":"10.3390\/microorganisms11102562","article-title":"Comparative Transcriptome Analysis of Shiga Toxin-Producing Escherichia coli O157:H7 on Bovine Rectoanal Junction Cells and Human Colonic Epithelial Cells during Initial Adherence","volume":"11","author":"LK Edison","year":"2023","journal-title":"Microorganisms"},{"issue":"6","key":"ppat.1013584.ref039","doi-asserted-by":"crossref","first-page":"598","DOI":"10.1038\/mi.2011.37","article-title":"Regulation of the polymeric immunoglobulin receptor and IgA transport: new advances in environmental factors that stimulate pIgR expression and its role in mucosal immunity","volume":"4","author":"F-E Johansen","year":"2011","journal-title":"Mucosal Immunol"},{"key":"ppat.1013584.ref040","volume-title":"Animal Nutrition Handbook","author":"LI Chiba","year":"2014"},{"key":"ppat.1013584.ref041","article-title":"Recto-Anal Junction (RAJ) and Fecal Microbiomes of Cattle Experimentally Challenged With Escherichia coli O157:H7","volume":"11","author":"RA Mir","year":"2020","journal-title":"Front Microbiol"},{"issue":"5","key":"ppat.1013584.ref042","doi-asserted-by":"crossref","first-page":"1289","DOI":"10.3390\/microorganisms11051289","article-title":"Bovine Rectoanal Junction In Vitro Organ Culture Model System to Study Shiga Toxin-Producing Escherichia coli Adherence","volume":"11","author":"IT Kudva","year":"2023","journal-title":"Microorganisms"},{"issue":"12","key":"ppat.1013584.ref043","doi-asserted-by":"crossref","first-page":"1322","DOI":"10.1128\/CVI.00205-06","article-title":"Bovine immune response to shiga-toxigenic Escherichia coli O157:H7","volume":"13","author":"MA Hoffman","year":"2006","journal-title":"Clin Vaccine Immunol"},{"issue":"1","key":"ppat.1013584.ref044","doi-asserted-by":"crossref","first-page":"1049","DOI":"10.1186\/s12864-016-3374-y","article-title":"Potential immunosuppressive effects of Escherichia coli O157:H7 experimental infection on the bovine host","volume":"17","author":"E Kieckens","year":"2016","journal-title":"BMC Genomics"},{"issue":"6","key":"ppat.1013584.ref045","doi-asserted-by":"crossref","first-page":"807","DOI":"10.1093\/bioinformatics\/btt038","article-title":"SwarmDock: a server for flexible protein-protein docking","volume":"29","author":"M Torchala","year":"2013","journal-title":"Bioinformatics"},{"issue":"12","key":"ppat.1013584.ref046","doi-asserted-by":"crossref","first-page":"2143","DOI":"10.1002\/prot.24369","article-title":"A Markov-chain model description of binding funnels to enhance the ranking of docked solutions","volume":"81","author":"M Torchala","year":"2013","journal-title":"Proteins"},{"key":"ppat.1013584.ref047","doi-asserted-by":"crossref","first-page":"181","DOI":"10.1007\/978-1-4939-0366-5_13","article-title":"Predicting the structure of protein-protein complexes using the SwarmDock Web Server","volume":"1137","author":"M Torchala","year":"2014","journal-title":"Methods Mol Biol"},{"issue":"10","key":"ppat.1013584.ref048","doi-asserted-by":"crossref","first-page":"5416","DOI":"10.1128\/IAI.70.10.5416-5427.2002","article-title":"Identification and characterization of lpfABCC\u2019DE, a fimbrial operon of enterohemorrhagic Escherichia coli O157:H7","volume":"70","author":"AG Torres","year":"2002","journal-title":"Infect Immun"},{"issue":"15","key":"ppat.1013584.ref049","doi-asserted-by":"crossref","first-page":"8951","DOI":"10.1073\/pnas.1537100100","article-title":"Bacteria-host communication: the language of hormones","volume":"100","author":"V Sperandio","year":"2003","journal-title":"Proc Natl Acad Sci U S A"},{"issue":"1","key":"ppat.1013584.ref050","doi-asserted-by":"crossref","first-page":"18","DOI":"10.1128\/IAI.73.1.18-29.2005","article-title":"Adherence of diarrheagenic Escherichia coli strains to epithelial cells","volume":"73","author":"AG Torres","year":"2005","journal-title":"Infect Immun"},{"issue":"4","key":"ppat.1013584.ref051","doi-asserted-by":"crossref","first-page":"2233","DOI":"10.1128\/IAI.74.4.2233-2244.2006","article-title":"Cloning, expression, and characterization of fimbrial operon F9 from enterohemorrhagic Escherichia coli O157:H7","volume":"74","author":"AS Low","year":"2006","journal-title":"Infect Immun"},{"issue":"7","key":"ppat.1013584.ref052","doi-asserted-by":"crossref","first-page":"1741","DOI":"10.1016\/j.micinf.2006.02.012","article-title":"Long polar fimbriae and tissue tropism in Escherichia coli O157:H7","volume":"8","author":"R Fitzhenry","year":"2006","journal-title":"Microbes Infect"},{"issue":"5","key":"ppat.1013584.ref053","doi-asserted-by":"crossref","first-page":"57","DOI":"10.1051\/vetres\/2010029","article-title":"Initial adherence of EPEC, EHEC and VTEC to host cells","volume":"41","author":"M Bardiau","year":"2010","journal-title":"Vet Res"},{"issue":"3","key":"ppat.1013584.ref054","doi-asserted-by":"crossref","first-page":"903","DOI":"10.1128\/IAI.05907-11","article-title":"Molecular mechanisms that mediate colonization of Shiga toxin-producing Escherichia coli strains","volume":"80","author":"MJ Farfan","year":"2012","journal-title":"Infect Immun"},{"issue":"3","key":"ppat.1013584.ref055","doi-asserted-by":"crossref","DOI":"10.1128\/microbiolspec.EHEC-0003-2013","article-title":"Enterohemorrhagic Escherichia coli Adhesins","volume":"2","author":"BD McWilliams","year":"2014","journal-title":"Microbiol Spectr"},{"issue":"11","key":"ppat.1013584.ref056","doi-asserted-by":"crossref","DOI":"10.1128\/spectrum.01270-24","article-title":"Investigating the adherence factors of Escherichia coli at the bovine recto-anal junction","volume":"12","author":"EM Nawrocki","year":"2024","journal-title":"Microbiol Spectr"},{"issue":"12","key":"ppat.1013584.ref057","doi-asserted-by":"crossref","first-page":"4302","DOI":"10.1128\/iai.59.12.4302-4309.1991","article-title":"The eae gene of enteropathogenic Escherichia coli encodes a 94-kilodalton membrane protein, the expression of which is influenced by the EAF plasmid","volume":"59","author":"AE Jerse","year":"1991","journal-title":"Infect Immun"},{"issue":"3","key":"ppat.1013584.ref058","doi-asserted-by":"crossref","first-page":"662","DOI":"10.1128\/JCM.36.3.662-668.1998","article-title":"Detection of intimins a, b, g, and d, four intimin derivatives expressed by attaching and effacing microbial pathogens","volume":"36","author":"J Adu-Bobie","year":"1998","journal-title":"J Clin Microbiol"},{"issue":"3","key":"ppat.1013584.ref059","doi-asserted-by":"crossref","first-page":"1400","DOI":"10.1128\/IAI.68.3.1400-1407.2000","article-title":"Iha: a novel Escherichia coli O157:H7 adherence-conferring molecule encoded on a recently acquired chromosomal island of conserved structure","volume":"68","author":"PI Tarr","year":"2000","journal-title":"Infect Immun"},{"issue":"6","key":"ppat.1013584.ref060","doi-asserted-by":"crossref","first-page":"207","DOI":"10.1016\/j.micpath.2006.07.006","article-title":"Expression of the Escherichia coli IrgA homolog adhesin is regulated by the ferric uptake regulation protein","volume":"41","author":"RA Rashid","year":"2006","journal-title":"Microb Pathog"},{"issue":"11","key":"ppat.1013584.ref061","doi-asserted-by":"crossref","first-page":"3519","DOI":"10.1172\/JCI30727","article-title":"Intestinal adherence associated with type IV pili of enterohemorrhagic Escherichia coli O157:H7","volume":"117","author":"J Xicohtencatl-Cortes","year":"2007","journal-title":"J Clin Invest"},{"key":"ppat.1013584.ref062","article-title":"Cell-surface localized nucleolin is a eucaryotic receptor for the adhesin intimin-g of enterohemorrhagic Escherichia coli","author":"JF Sinclair","year":"2000","journal-title":"J Biol Chem"},{"key":"ppat.1013584.ref063","doi-asserted-by":"crossref","first-page":"237","DOI":"10.1128\/9781555817619.ch17","article-title":"Tissue Tropism in Intestinal Colonization","volume-title":"Colonization of Mucosal Surfaces","author":"EL Hartland","year":"2014"},{"issue":"2","key":"ppat.1013584.ref064","doi-asserted-by":"crossref","first-page":"180","DOI":"10.1136\/gut.50.2.180","article-title":"Intimin type influences the site of human intestinal mucosal colonisation by enterohaemorrhagic Escherichia coli O157:H7","volume":"50","author":"RJ 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Escherichia coli O157:H7 Super-Shedders","volume":"11","author":"O Wang","year":"2016","journal-title":"PLoS One"},{"issue":"1","key":"ppat.1013584.ref080","doi-asserted-by":"crossref","first-page":"21535","DOI":"10.1038\/s41598-020-78752-x","article-title":"Early immune innate hallmarks and microbiome changes across the gut during Escherichia coli O157: H7 infection in cattle","volume":"10","author":"M Larz\u00e1bal","year":"2020","journal-title":"Sci Rep"},{"issue":"12","key":"ppat.1013584.ref081","doi-asserted-by":"crossref","first-page":"1945","DOI":"10.1007\/s00726-017-2410-9","article-title":"L-Glutamine and L-arginine protect against enterotoxigenic Escherichia coli infection via intestinal innate immunity in mice","volume":"49","author":"G Liu","year":"2017","journal-title":"Amino Acids"},{"issue":"4","key":"ppat.1013584.ref082","doi-asserted-by":"crossref","first-page":"346","DOI":"10.1089\/fpd.2011.1021","article-title":"In vitro adherence patterns of Shigella serogroups to bovine recto-anal junction squamous epithelial (RSE) cells are similar to those of Escherichia coli O157","volume":"9","author":"IT Kudva","year":"2012","journal-title":"Foodborne Pathog Dis"},{"issue":"4","key":"ppat.1013584.ref083","doi-asserted-by":"crossref","first-page":"1363","DOI":"10.1128\/aem.61.4.1363-1370.1995","article-title":"Effect of diet on the shedding of Escherichia coli O157:H7 in a sheep model","volume":"61","author":"IT Kudva","year":"1995","journal-title":"Appl Environ Microbiol"},{"issue":"10","key":"ppat.1013584.ref084","doi-asserted-by":"crossref","first-page":"3878","DOI":"10.1128\/aem.63.10.3878-3886.1997","article-title":"Evaluation of dietary influences on Escherichia coli O157:H7 shedding by sheep","volume":"63","author":"IT Kudva","year":"1997","journal-title":"Appl Environ Microbiol"},{"issue":"4","key":"ppat.1013584.ref085","doi-asserted-by":"crossref","first-page":"892","DOI":"10.1128\/jcm.35.4.892-899.1997","article-title":"Characterization of Escherichia coli O157:H7 and other Shiga toxin-producing E. coli serotypes isolated from sheep","volume":"35","author":"IT Kudva","year":"1997","journal-title":"J Clin Microbiol"},{"key":"ppat.1013584.ref086","doi-asserted-by":"crossref","first-page":"140105","DOI":"10.1155\/2012\/140105","article-title":"Polymorphic Amplified Typing Sequences and Pulsed-Field Gel Electrophoresis Yield Comparable Results in the Strain Typing of a Diverse Set of Bovine Escherichia coli O157:H7 Isolates","volume":"2012","author":"IT Kudva","year":"2012","journal-title":"Int J Microbiol"},{"key":"ppat.1013584.ref087","doi-asserted-by":"crossref","first-page":"123","DOI":"10.2174\/1874285801307010123","article-title":"Polymorphic Amplified Typing Sequences (PATS) Strain Typing System Accurately Discriminates a Set of Temporally and Spatially Disparate Escherichia coli O157 Isolates Associated with Human Infection","volume":"7","author":"IT Kudva","year":"2013","journal-title":"Open Microbiol 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a novel strain typing system for Escherichia coli O157:H7","volume":"42","author":"IT Kudva","year":"2004","journal-title":"J Clin Microbiol"},{"key":"ppat.1013584.ref091","doi-asserted-by":"crossref","first-page":"266","DOI":"10.1186\/1746-6148-9-266","article-title":"Bison and bovine rectoanal junctions exhibit similar cellular architecture and Escherichia coli O157 adherence patterns","volume":"9","author":"IT Kudva","year":"2013","journal-title":"BMC Vet Res"},{"issue":"1","key":"ppat.1013584.ref092","doi-asserted-by":"crossref","first-page":"22","DOI":"10.1016\/j.jmoldx.2011.08.002","article-title":"RNAscope: a novel in situ RNA analysis platform for formalin-fixed, paraffin-embedded tissues","volume":"14","author":"F Wang","year":"2012","journal-title":"J Mol Diagn"},{"issue":"7873","key":"ppat.1013584.ref093","doi-asserted-by":"crossref","first-page":"583","DOI":"10.1038\/s41586-021-03819-2","article-title":"Highly accurate protein structure prediction with 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