{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,8,19]],"date-time":"2026-08-19T06:27:52Z","timestamp":1787120872329,"version":"build-2736575974"},"reference-count":79,"publisher":"Frontiers Media SA","license":[{"start":{"date-parts":[[2026,8,19]],"date-time":"2026-08-19T00:00:00Z","timestamp":1787097600000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"content-domain":{"domain":["frontiersin.org"],"crossmark-restriction":true},"short-container-title":["Front. Microbiol."],"abstract":"<jats:sec>\n                    <jats:title>Background<\/jats:title>\n                    <jats:p>\n                      <jats:italic>Helicobacter pylori<\/jats:italic>\n                      lipopolysaccharide (LPS) contributes to bacterial fitness, immune evasion, host colonization, and virulence. The O-antigen region contains Lewis antigens and other glycans that facilitate adaptation to the gastric environment. HP1581 is predicted to encode a WecA-like UDP-\n                      <jats:italic>N<\/jats:italic>\n                      -acetylglucosamine-1-phosphate transferase that catalyzes the first step of O-antigen biosynthesis. However, its role in LPS assembly and\n                      <jats:italic>H. pylori<\/jats:italic>\n                      pathogenesis remains unclear.\n                    <\/jats:p>\n                  <\/jats:sec>\n                  <jats:sec>\n                    <jats:title>Methods<\/jats:title>\n                    <jats:p>\n                      An\n                      <jats:italic>HP1581<\/jats:italic>\n                      (\n                      <jats:italic>wecA<\/jats:italic>\n                      ) knockout mutant and a complemented strain were constructed in\n                      <jats:italic>H. pylori<\/jats:italic>\n                      strain 26695 to investigate the role of HP1581 in LPS biosynthesis, bacterial physiology, outer membrane vesicle (OMV) composition, host cell interactions, and virulence. Biofilm formation and swimming motility were examined using O-antigen-deficient mutants of the motile\n                      <jats:italic>H. pylori<\/jats:italic>\n                      G27 strain, including\n                      <jats:italic>HP1581<\/jats:italic>\n                      (\n                      <jats:italic>wecA<\/jats:italic>\n                      ),\n                      <jats:italic>HP1206<\/jats:italic>\n                      (\n                      <jats:italic>wzk<\/jats:italic>\n                      ), and\n                      <jats:italic>HP1039<\/jats:italic>\n                      (\n                      <jats:italic>waaL<\/jats:italic>\n                      ) mutants.\n                    <\/jats:p>\n                  <\/jats:sec>\n                  <jats:sec>\n                    <jats:title>Results<\/jats:title>\n                    <jats:p>\n                      Disruption of\n                      <jats:italic>HP1581<\/jats:italic>\n                      abolished O-antigen biosynthesis and eliminated Lewis X and Lewis Y antigen expression, resulting in a truncated LPS structure lacking the O-antigen region. The mutant displayed increased sensitivity to sodium dodecyl sulfate and novobiocin, elevated surface hydrophobicity and autoaggregation, and reduced survival during late-stage growth. Loss of\n                      <jats:italic>HP1581<\/jats:italic>\n                      significantly impaired bacterial adhesion and internalization in AGS cells and attenuated the induction of the elongated (hummingbird) phenotype. In addition, OMVs from the mutant contained substantially lower levels of CagA and VacA. In the\n                      <jats:italic>Galleria mellonella<\/jats:italic>\n                      infection model, the mutant exhibited reduced persistence and attenuated virulence. Furthermore, O-antigen-deficient mutants of the motile\n                      <jats:italic>H. pylori<\/jats:italic>\n                      G27 strain, including\n                      <jats:italic>HP1581<\/jats:italic>\n                      (\n                      <jats:italic>wecA<\/jats:italic>\n                      ),\n                      <jats:italic>HP1206<\/jats:italic>\n                      (\n                      <jats:italic>wzk<\/jats:italic>\n                      ), and\n                      <jats:italic>HP1039<\/jats:italic>\n                      (\n                      <jats:italic>waaL<\/jats:italic>\n                      ) mutants, exhibited enhanced biofilm formation despite reduced swimming motility. Biofilm stability was found to depend primarily on extracellular proteins rather than extracellular DNA.\n                    <\/jats:p>\n                  <\/jats:sec>\n                  <jats:sec>\n                    <jats:title>Conclusion<\/jats:title>\n                    <jats:p>\n                      HP1581 (WecA) is essential for O-antigen biosynthesis and plays critical roles in maintaining outer membrane integrity, regulating bacterial surface properties, promoting host cell interactions, modulating virulence-associated factors, and influencing biofilm development. These findings identify HP1581 (WecA)-dependent O-antigen biosynthesis as a key determinant of\n                      <jats:italic>H. pylori<\/jats:italic>\n                      virulence and persistence and highlight HP1581 as a potential therapeutic target.\n                    <\/jats:p>\n                  <\/jats:sec>","DOI":"10.3389\/fmicb.2026.1909795","type":"journal-article","created":{"date-parts":[[2026,8,19]],"date-time":"2026-08-19T05:47:37Z","timestamp":1787118457000},"update-policy":"https:\/\/doi.org\/10.3389\/crossmark-policy","source":"Crossref","is-referenced-by-count":0,"title":["HP1581 (WecA)-dependent O-antigen biosynthesis governs adhesion, virulence, and biofilm dynamics in Helicobacter pylori"],"prefix":"10.3389","volume":"17","author":[{"given":"Tram Thi Hong","family":"Nguyen","sequence":"first","affiliation":[{"name":"Institute of Molecular Medicine, National Tsing Hua University","place":["Hsinchu, Taiwan"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Mamoona","family":"Hassan","sequence":"additional","affiliation":[{"name":"Institute of Molecular Medicine, National Tsing Hua University","place":["Hsinchu, Taiwan"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Kai-Wen","family":"Teng","sequence":"additional","affiliation":[{"name":"Institute of Molecular Medicine, National Tsing Hua University","place":["Hsinchu, Taiwan"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Steffen","family":"Backert","sequence":"additional","affiliation":[{"name":"Division of Microbiology, Department of Biology, Friedrich-Alexander-Universit\u00e4t Erlangen-N\u00fcrnberg","place":["Erlangen, Germany"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Song-Wei","family":"Wang","sequence":"additional","affiliation":[{"name":"Division of Gastroenterology, Department of Internal Medicine, Kaohsiung Medical University Hospital","place":["Kaohsiung, Taiwan"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Chung-Jung","family":"Liu","sequence":"additional","affiliation":[{"name":"Division of Gastroenterology, Department of Internal Medicine, Kaohsiung Medical University Hospital","place":["Kaohsiung, Taiwan"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Deng-Chyang","family":"Wu","sequence":"additional","affiliation":[{"name":"Division of Gastroenterology, Department of Internal Medicine, Kaohsiung Medical University Hospital","place":["Kaohsiung, Taiwan"]},{"name":"School of Medicine, College of Medicine, Kaohsiung Medical University","place":["Kaohsiung, Taiwan"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Chao-Hung","family":"Kuo","sequence":"additional","affiliation":[{"name":"Division of Gastroenterology, Department of Internal Medicine, Kaohsiung Medical University Hospital","place":["Kaohsiung, Taiwan"]},{"name":"School of Medicine, College of Medicine, Kaohsiung Medical University","place":["Kaohsiung, Taiwan"]},{"name":"Division of Gastroenterology, Department of Internal Medicine, Kaohsiung Show Chwan Memorial Hospital","place":["Kaohsiung, Taiwan"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Mou-Chieh","family":"Kao","sequence":"additional","affiliation":[{"name":"Institute of Molecular Medicine, National Tsing Hua University","place":["Hsinchu, Taiwan"]},{"name":"Department of Life Science, National Tsing Hua University","place":["Hsinchu, Taiwan"]},{"name":"School of Medicine, National Tsing Hua University","place":["Hsinchu, Taiwan"]}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"1965","published-online":{"date-parts":[[2026,8,19]]},"reference":[{"key":"B1","doi-asserted-by":"publisher","first-page":"7141","DOI":"10.1128\/JB.00676-08","article-title":"Purification and characterization of the bacterial UDP-GlcNAc: undecaprenyl-phosphate GlcNAc-1-phosphate transferase WecA","volume":"190","author":"Al-Dabbagh","year":"2008","journal-title":"J. Bacteriol."},{"key":"B2","doi-asserted-by":"publisher","first-page":"777","DOI":"10.1093\/glycob\/cwg106","article-title":"Occurrence of a nontypable Helicobacter pylori strain lacking Lewis blood group O antigens and DD-heptoglycan: evidence for the role of the core \u03b11,6-glucan chain in colonization","volume":"13","author":"Altman","year":"2003","journal-title":"Glycobiology"},{"key":"B3","doi-asserted-by":"publisher","first-page":"2031","DOI":"10.1128\/IAI.64.6.2031-2040.1996","article-title":"Potential role of molecular mimicry between Helicobacter pylori lipopolysaccharide and host Lewis blood group antigens in autoimmunity","volume":"64","author":"Appelmelk","year":"1996","journal-title":"Infect. Immun"},{"key":"B4","doi-asserted-by":"publisher","first-page":"1130","DOI":"10.1111\/j.1365-2958.2010.07393.x","article-title":"Helicobacter pylori HP0518 affects flagellin glycosylation to alter bacterial motility","volume":"78","author":"Asakura","year":"2010","journal-title":"Mol. Microbiol."},{"key":"B5","doi-asserted-by":"publisher","first-page":"163","DOI":"10.1007\/978-1-4939-6536-6_14","article-title":"\u201cUsing Galleria mellonella as an infection model for Campylobacter jejuni pathogenesis,\u201d","author":"Askoura","year":"2017","journal-title":"Campylobacter jejuni: Methods and Protocols"},{"key":"B6","doi-asserted-by":"publisher","first-page":"447","DOI":"10.1128\/JB.01416-08","article-title":"The complete genome sequence of Helicobacter pylori strain G27","volume":"191","author":"Baltrus","year":"2009","journal-title":"J. Bacteriol."},{"key":"B7","doi-asserted-by":"publisher","first-page":"517","DOI":"10.1016\/j.tim.2014.05.002","article-title":"Biofilms, flagella, and mechanosensing of surfaces by bacteria","volume":"22","author":"Belas","year":"2014","journal-title":"Trends Microbiol."},{"key":"B8","doi-asserted-by":"publisher","first-page":"218","DOI":"10.1109\/CBMS58004.2023.00220","article-title":"\u201cComparing HISAT and STAR-based pipelines for RNA-Seq data analysis: a real experience,\u201d","author":"Bianchi","year":"2023","journal-title":"2023 IEEE 36th International Symposium on Computer-Based Medical Systems (CBMS)"},{"key":"B9","doi-asserted-by":"publisher","first-page":"57","DOI":"10.1007\/5584_2019_359","article-title":"Roles of adhesion to epithelial cells in gastric colonization by Helicobacter pylori","volume":"11","author":"Bonsor","year":"2019","journal-title":"Adv. Microbiol. Infect. Dis. Public Health"},{"key":"B10","doi-asserted-by":"publisher","first-page":"179","DOI":"10.1016\/S0168-6445(99)00004-2","article-title":"Physico-chemistry of initial microbial adhesive interactions\u2014its mechanisms and methods for study","volume":"23","author":"Bos","year":"1999","journal-title":"FEMS Microbiol. Rev."},{"key":"B11","doi-asserted-by":"publisher","first-page":"2208928","DOI":"10.1080\/19336950.2023.2208928","article-title":"Identification of SLC2A3 as a prognostic indicator correlated with the NF-\u03baB\/EMT axis and immune response in head and neck squamous cell carcinoma","volume":"17","author":"Chai","year":"2023","journal-title":"Channels"},{"key":"B12","doi-asserted-by":"publisher","first-page":"497","DOI":"10.1016\/j.bbrc.2011.01.060","article-title":"Effects of a HP0859 (rfaD) knockout mutation on lipopolysaccharide structure of Helicobacter pylori 26695 and the bacterial adhesion on AGS cells","volume":"405","author":"Chang","year":"2011","journal-title":"Biochem. Biophys. Res. Commun."},{"key":"B13","doi-asserted-by":"publisher","first-page":"1610","DOI":"10.1080\/21505594.2021.1938449","article-title":"Helicobacter pylori GmhB enzyme involved in ADP-heptose biosynthesis pathway is essential for lipopolysaccharide biosynthesis and bacterial virulence","volume":"12","author":"Chiu","year":"2021","journal-title":"Virulence"},{"key":"B14","doi-asserted-by":"publisher","first-page":"1150625","DOI":"10.3389\/fmicb.2023.1150625","article-title":"Lipopolysaccharide structure modulates cationic biocide susceptibility and crystalline biofilm formation in Proteus mirabilis","volume":"14","author":"Clarke","year":"2023","journal-title":"Front. Microbiol."},{"key":"B15","doi-asserted-by":"publisher","first-page":"808","DOI":"10.1016\/j.bbapap.2008.11.005","article-title":"Outer membrane permeability and antibiotic resistance","volume":"1794","author":"Delcour","year":"2009","journal-title":"Biochim. Biophys. Acta Proteins Proteomics"},{"key":"B16","doi-asserted-by":"publisher","first-page":"881","DOI":"10.3201\/eid0809.020063","article-title":"Biofilms: microbial life on surfaces","volume":"8","author":"Donlan","year":"2002","journal-title":"Emerg. Infect. Dis."},{"key":"B17","doi-asserted-by":"publisher","first-page":"e2417572121","DOI":"10.1073\/pnas.2417572121","article-title":"Proteome-wide bioinformatic annotation and functional validation of the monotopic phosphoglycosyl transferase superfamily","volume":"121","author":"Durand","year":"2024","journal-title":"Proc. Natl. Acad. Sci. U.S.A."},{"key":"B18","doi-asserted-by":"publisher","first-page":"1530","DOI":"10.1046\/j.1365-2958.2000.01823.x","article-title":"Lewis X structures in the O antigen side-chain promote adhesion of Helicobacter pylori to the gastric epithelium","volume":"35","author":"Edwards","year":"2000","journal-title":"Mol. Microbiol."},{"key":"B19","doi-asserted-by":"publisher","first-page":"563","DOI":"10.1038\/nrmicro.2016.94","article-title":"Biofilms: an emergent form of bacterial life","volume":"14","author":"Flemming","year":"2016","journal-title":"Nat. Rev. Microbiol."},{"key":"B20","doi-asserted-by":"publisher","first-page":"37","DOI":"10.1016\/j.carres.2012.03.013","article-title":"Synthesis and immunomodulatory activities of Helicobacter pylori lipophilic terminus of lipopolysaccharide including lipid A","volume":"356","author":"Fujimoto","year":"2012","journal-title":"Carbohydr. Res."},{"key":"B21","doi-asserted-by":"publisher","first-page":"e01349","DOI":"10.1128\/mBio.01349-15","article-title":"Helicobacter pylori resists the antimicrobial activity of calprotectin via lipid A modification and associated biofilm formation","volume":"6","author":"Gaddy","year":"2015","journal-title":"MBio"},{"key":"B22","doi-asserted-by":"publisher","first-page":"e0221431","DOI":"10.1371\/journal.pone.0221431","article-title":"Flagella-dependent inhibition of biofilm formation by sub-inhibitory concentration of polymyxin B in Vibrio cholerae","volume":"14","author":"Giacomucci","year":"2019","journal-title":"PLoS ONE"},{"key":"B23","doi-asserted-by":"publisher","first-page":"1369","DOI":"10.3389\/fmicb.2015.01369","volume":"6","author":"Grande","year":"2015","journal-title":"Helicobacter pyloriFront. Microbiol."},{"key":"B24","doi-asserted-by":"publisher","first-page":"141","DOI":"10.1016\/S2468-1253(21)00400-3","article-title":"Lyophilised oral faecal microbiota transplantation for ulcerative colitis (LOTUS): a randomised, double-blind, placebo-controlled trial","volume":"7","author":"Haifer","year":"2022","journal-title":"Lancet Gastroenterol. Hepatol."},{"key":"B25","doi-asserted-by":"publisher","first-page":"95","DOI":"10.1038\/nrmicro821","article-title":"Bacterial biofilms: from the natural environment to infectious diseases","volume":"2","author":"Hall-Stoodley","year":"2004","journal-title":"Nat. Rev. Microbiol."},{"key":"B26","doi-asserted-by":"publisher","first-page":"56","DOI":"10.1038\/s41522-020-00167-3","article-title":"Genetic requirements and transcriptomics of Helicobacter pylori biofilm formation on abiotic and biotic surfaces","volume":"6","author":"Hathroubi","year":"2020","journal-title":"NPJ Biofilms Microbiomes"},{"key":"B27","doi-asserted-by":"publisher","first-page":"e00001","DOI":"10.1128\/MMBR.00001-18","article-title":"Helicobacter pylori biofilm formation and its potential role in pathogenesis","volume":"82","author":"Hathroubi","year":"","journal-title":"Microbiol. Mol. Biol. Rev."},{"key":"B28","doi-asserted-by":"publisher","first-page":"10","DOI":"10.1128\/mBio.01973-18","article-title":"Helicobacter pylori biofilm involves a multigene stress-biased response, including a structural role for flagella","volume":"9","author":"Hathroubi","year":"","journal-title":"MBio"},{"key":"B29","doi-asserted-by":"publisher","first-page":"102","DOI":"10.1016\/j.dci.2015.12.006","article-title":"Insect immunology and hematopoiesis","volume":"58","author":"Hillyer","year":"2016","journal-title":"Dev. Comp. Immunol."},{"key":"B30","doi-asserted-by":"publisher","first-page":"2534","DOI":"10.1128\/IAI.71.5.2534-2541.2003","article-title":"Development of an interleukin-12-deficient mouse model that is permissive for colonization by a motile KE26695 strain of Helicobacter pylori","volume":"71","author":"Hoffman","year":"2003","journal-title":"Infect. Immun."},{"key":"B31","doi-asserted-by":"publisher","first-page":"e1000819","DOI":"10.1371\/journal.ppat.1000819","article-title":"Helicobacter pylori lipopolysaccharide is synthesized via a novel pathway with an evolutionary connection to protein N-glycosylation","volume":"6","author":"Hug","year":"2010","journal-title":"PLoS Pathog."},{"key":"B32","doi-asserted-by":"publisher","first-page":"88","DOI":"10.3389\/fcimb.2017.00088","article-title":"The bioinformatics analysis of comparative genomics of Mycobacterium tuberculosis complex (MTBC) provides insight into dissimilarities between intraspecific groups differing in host association, virulence, and epitope diversity","volume":"7","author":"Jia","year":"2017","journal-title":"Front. Cell. Infect. Microbiol."},{"key":"B33","doi-asserted-by":"publisher","first-page":"383","DOI":"10.1080\/21505594.2017.1397871","article-title":"From moths to caterpillars: ideal conditions for Galleria mellonella rearing for in vivo microbiological studies","volume":"9","author":"Jorj\u00e3o","year":"2018","journal-title":"Virulence"},{"key":"B34","doi-asserted-by":"publisher","first-page":"1048","DOI":"10.1111\/1574-6976.12070","article-title":"Progress in understanding the assembly process of bacterial O-antigen","volume":"38","author":"Kalynych","year":"2014","journal-title":"FEMS Microbiol. Rev."},{"key":"B35","doi-asserted-by":"publisher","first-page":"14","DOI":"10.1016\/j.bj.2015.06.002","article-title":"Helicobacter pylori infection: an overview of bacterial virulence factors and pathogenesis","volume":"39","author":"Kao","year":"2016","journal-title":"Biomed. J."},{"key":"B36","doi-asserted-by":"publisher","first-page":"37","DOI":"10.22203\/eCM.v008a05","article-title":"Concise review of mechanisms of bacterial adhesion to biomaterials and of techniques used in estimating bacteria\u2013material interactions","volume":"8","author":"Katsikogianni","year":"2004","journal-title":"Eur. Cell. Mater."},{"key":"B37","doi-asserted-by":"publisher","first-page":"112","DOI":"10.3389\/fcimb.2014.00112","article-title":"How microorganisms use hydrophobicity and what does this mean for human needs?","volume":"4","author":"Krasowska","year":"2014","journal-title":"Front. Cell. Infect. Microbiol."},{"key":"B38","doi-asserted-by":"publisher","first-page":"1062","DOI":"10.3390\/pathogens9121062","article-title":"Biofilm formation as a complex result of virulence and adaptive responses of Helicobacter pylori","volume":"9","author":"Krzy\u017cek","year":"2020","journal-title":"Pathogens"},{"key":"B39","doi-asserted-by":"publisher","first-page":"2120747","DOI":"10.1080\/19490976.2022.2120747","article-title":"Biomaterials for Helicobacter pylori therapy: therapeutic potential and future perspectives","volume":"14","author":"Lai","year":"2022","journal-title":"Gut Microbes"},{"key":"B40","doi-asserted-by":"publisher","first-page":"445","DOI":"10.1111\/hel.12301","article-title":"Lipopolysaccharide structure and biosynthesis in Helicobacter pylori","volume":"21","author":"Li","year":"2016","journal-title":"Helicobacter"},{"key":"B41","doi-asserted-by":"publisher","first-page":"145","DOI":"10.3390\/biomedicines10010145","article-title":"The effects of HP0044 and HP1275 knockout mutations on the structure and function of lipopolysaccharide in Helicobacter pylori strain 26695","volume":"10","author":"Liu","year":"","journal-title":"Biomedicines"},{"key":"B42","doi-asserted-by":"publisher","first-page":"2730","DOI":"10.3390\/ijms23052730","article-title":"C2H2 zinc finger proteins response to abiotic stress in plants","volume":"23","author":"Liu","year":"","journal-title":"Int. J. Mol. Sci."},{"key":"B43","doi-asserted-by":"publisher","first-page":"820","DOI":"10.1093\/glycob\/cwx064","article-title":"Bacterial phosphoglycosyl transferases: initiators of glycan biosynthesis at the membrane interface","volume":"27","author":"Lukose","year":"2017","journal-title":"Glycobiology"},{"key":"B44","doi-asserted-by":"publisher","first-page":"19","DOI":"10.1038\/s41572-023-00431-8","article-title":"Helicobacter pylori infection","volume":"9","author":"Malfertheiner","year":"2023","journal-title":"Nat. Rev. Dis. Primers"},{"key":"B45","doi-asserted-by":"publisher","first-page":"656439","DOI":"10.3389\/fmolb.2021.656439","article-title":"Adhesion of Helicobacter species to the human gastric mucosa: a deep look into glycans role","volume":"8","author":"Matos","year":"2021","journal-title":"Front. Mol. Biosci."},{"key":"B46","doi-asserted-by":"publisher","first-page":"101","DOI":"10.3390\/toxins9030101","article-title":"Helicobacter pylori outer membrane protein-related pathogenesis","volume":"9","author":"Matsuo","year":"2017","journal-title":"Toxins"},{"key":"B47","doi-asserted-by":"publisher","first-page":"109","DOI":"10.3389\/fimmu.2013.00109","article-title":"Structural modifications of bacterial lipopolysaccharide that facilitate gram-negative bacteria evasion of host innate immunity","volume":"4","author":"Matsuura","year":"2013","journal-title":"Front. Immunol."},{"key":"B48","doi-asserted-by":"publisher","first-page":"958","DOI":"10.1099\/0022-1317-51-11-958","article-title":"The Helicobacter pylori flbA flagellar biosynthesis and regulatory gene is required for motility and virulence and modulates urease of H. pylori and Proteus mirabilis","volume":"51","author":"McGEE","year":"2002","journal-title":"J. Med. Microbiol."},{"key":"B49","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1016\/j.adengl.2020.12.014","article-title":"Clinical and histopathologic characteristics of the main causes of vascular occusion\u2014part i: thrombi","volume":"112","author":"Merino","year":"2021","journal-title":"Actas Dermo Sifiliogr\u00e1ficas"},{"key":"B50","doi-asserted-by":"publisher","first-page":"39","DOI":"10.1046\/j.1365-2036.1996.22164004.x","article-title":"The role of lipopolysaccharide in Helicobacter pylori pathogenesis","volume":"10","author":"Moran","year":"1996","journal-title":"Aliment. Pharmacol. Ther."},{"key":"B51","doi-asserted-by":"publisher","first-page":"e51241","DOI":"10.1371\/journal.pone.0051241","article-title":"Enhanced biofilm formation by Escherichia coli LPS mutants defective in Hep biosynthesis","volume":"7","author":"Nakao","year":"2012","journal-title":"PLoS ONE"},{"key":"B52","doi-asserted-by":"publisher","first-page":"6145","DOI":"10.1128\/IAI.00261-06","article-title":"Porphyromonas gingivalis galE is involved in lipopolysaccharide O-antigen synthesis and biofilm formation","volume":"74","author":"Nakao","year":"2006","journal-title":"Infect. Immun."},{"key":"B53","doi-asserted-by":"publisher","first-page":"2548620","DOI":"10.1080\/21505594.2025.2548620","article-title":"Roles of the lipopolysaccharide biosynthesis-related gene HP0858 in the fitness of Helicobacter pylori and its virulence in Galleria mellonella","volume":"16","author":"Nguyen","year":"2025","journal-title":"Virulence"},{"key":"B54","doi-asserted-by":"publisher","first-page":"237","DOI":"10.1093\/dnares\/dsm024","article-title":"A genome-wide approach to identify the genes involved in biofilm formation in E. coli","volume":"14","author":"Niba","year":"2007","journal-title":"DNA Res."},{"key":"B55","doi-asserted-by":"publisher","first-page":"593","DOI":"10.1128\/MMBR.67.4.593-656.2003","article-title":"Molecular basis of bacterial outer membrane permeability revisited","volume":"67","author":"Nikaido","year":"2003","journal-title":"Microbiol. Mol. Biol. Rev."},{"key":"B56","doi-asserted-by":"publisher","first-page":"2863","DOI":"10.1073\/pnas.0511119103","article-title":"An enzymatic ruler modulates Lewis antigen glycosylation of Helicobacter pylori LPS during persistent infection","volume":"103","author":"Nilsson","year":"2006","journal-title":"Proc. Natl. Acad. Sci. U.S.A."},{"key":"B57","doi-asserted-by":"publisher","first-page":"51","DOI":"10.1007\/978-1-62703-005-2_8","article-title":"\u201cGenetic manipulation of a naturally competent bacterium, Helicobacter pylori,\u201d","author":"Noto","year":"2012","journal-title":"Helicobacter Species: Methods and Protocols"},{"key":"B58","doi-asserted-by":"publisher","first-page":"1477","DOI":"10.1099\/jmm.0.46660-0","article-title":"Mutation of luxS affects motility and infectivity of Helicobacter pylori in gastric mucosa of a Mongolian gerbil model","volume":"55","author":"Osaki","year":"2006","journal-title":"J. Med. Microbiol."},{"key":"B59","doi-asserted-by":"publisher","first-page":"295","DOI":"10.1046\/j.1365-2958.1998.01062.x","article-title":"Flagellar and twitching motility are necessary for Pseudomonas aeruginosa biofilm development","volume":"30","author":"O'Toole","year":"1998","journal-title":"Mol. Microbiol."},{"key":"B60","doi-asserted-by":"publisher","first-page":"e2018289118","DOI":"10.1073\/pnas.2018289118","article-title":"Glycoconjugate pathway connections revealed by sequence similarity network analysis of the monotopic phosphoglycosyl transferases","volume":"118","author":"O'Toole","year":"2021","journal-title":"Proc. Natl. Acad. Sci. U.S.A."},{"key":"B61","doi-asserted-by":"publisher","first-page":"29R","DOI":"10.1093\/glycob\/cwi065","article-title":"Modeling bacterial UDP-HexNAc: polyprenol-P HexNAc-1-P transferases","volume":"15","author":"Price","year":"2005","journal-title":"Glycobiology"},{"key":"B62","doi-asserted-by":"publisher","first-page":"94","DOI":"10.1016\/S0966-842X(02)00034-3","article-title":"Bacterial coaggregation: an integral process in the development of multi-species biofilms","volume":"11","author":"Rickard","year":"2003","journal-title":"Trends Microbiol."},{"key":"B63","doi-asserted-by":"publisher","first-page":"501","DOI":"10.1161\/CIRCRESAHA.119.315215","article-title":"Longitudinal RNA-Seq analysis of the repeatability of gene expression and splicing in human platelets identifies a platelet SELP splice QTL","volume":"126","author":"Rondina","year":"2020","journal-title":"Circ. Res."},{"key":"B64","doi-asserted-by":"publisher","first-page":"315","DOI":"10.1093\/glycob\/7.2.315","volume":"7","author":"Rush","year":"1997","journal-title":"E. coli. Glycobiology"},{"key":"B65","doi-asserted-by":"publisher","first-page":"661","DOI":"10.1099\/jmm.0.026658-0","article-title":"Galleria mellonella as an infection model for Campylobacter jejuni virulence","volume":"60","author":"Senior","year":"2011","journal-title":"J. Med. Microbiol."},{"key":"B66","doi-asserted-by":"publisher","first-page":"125","DOI":"10.12688\/f1000research.27504.1","article-title":"Isolation and primary culture of Galleria mellonella hemocytes for infection studies","volume":"2138","author":"Senior","year":"2020","journal-title":"Methods Mol. Biol."},{"key":"B67","doi-asserted-by":"publisher","first-page":"8587","DOI":"10.1038\/s41598-024-59234-w","article-title":"Probing the expression and adhesion of glycans involved in Helicobacter pylori infection","volume":"14","author":"Sijmons","year":"2024","journal-title":"Sci. Rep."},{"key":"B68","doi-asserted-by":"publisher","first-page":"a000414","DOI":"10.1101\/cshperspect.a000414","article-title":"The bacterial cell envelope","volume":"2","author":"Silhavy","year":"2010","journal-title":"Cold Spring Harb. Perspect. Biol."},{"key":"B69","doi-asserted-by":"publisher","first-page":"539","DOI":"10.1038\/41483","article-title":"The complete genome sequence of the gastric pathogen Helicobacter pylori","volume":"388","author":"Tomb","year":"1997","journal-title":"Nature"},{"key":"B70","doi-asserted-by":"publisher","first-page":"3","DOI":"10.3390\/jof5010003","article-title":"Immune response of Galleria mellonella against human fungal pathogens","volume":"5","author":"Trevijano-Contador","year":"2018","journal-title":"J. Fungi"},{"key":"B71","doi-asserted-by":"publisher","first-page":"140","DOI":"10.3934\/microbiol.2018.1.140","article-title":"Bacterial autoaggregation","volume":"4","author":"Trunk","year":"2018","journal-title":"APMIS Microbiol"},{"key":"B72","doi-asserted-by":"publisher","first-page":"451","DOI":"10.1111\/j.1348-0421.1995.tb02228.x","article-title":"The release of outer membrane vesicles from the strains of enterotoxigenic Escherichia coli","volume":"39","author":"Wai","year":"1995","journal-title":"Microbiol. Immunol."},{"key":"B73","doi-asserted-by":"publisher","first-page":"e0166835","DOI":"10.1371\/journal.pone.0166835","article-title":"Comparative genomics revealed multiple Helicobacter pylori genes associated with biofilm formation in vitro","volume":"11","author":"Wong","year":"2016","journal-title":"PLoS ONE"},{"key":"B74","doi-asserted-by":"publisher","first-page":"361","DOI":"10.1007\/s00253-005-0263-8","article-title":"Motility influences biofilm architecture in Escherichia coli","volume":"72","author":"Wood","year":"2006","journal-title":"Appl. Microbiol. Biotechnol."},{"key":"B75","doi-asserted-by":"publisher","first-page":"914791","DOI":"10.1155\/2015\/914791","article-title":"Biofilm formation by Helicobacter pylori and its involvement for antibiotic resistance","volume":"2015","author":"Yonezawa","year":"2015","journal-title":"Biomed Res. Int."},{"key":"B76","doi-asserted-by":"publisher","first-page":"794","DOI":"10.1016\/j.bbrc.2016.06.137","article-title":"Functional characterization of Helicobacter pylori 26695 sedoheptulose 7-phosphate isomerase encoded by hp0857 and its association with lipopolysaccharide biosynthesis and adhesion","volume":"477","author":"Yu","year":"2016","journal-title":"Biochem. Biophys. Res. Commun."},{"key":"B77","doi-asserted-by":"publisher","first-page":"102701","DOI":"10.1016\/j.ebiom.2020.102701","article-title":"Microtubule associated protein 9 inhibits liver tumorigenesis by suppressing ERCC3","volume":"53","author":"Zhang","year":"2020","journal-title":"EBioMedicine"},{"key":"B78","doi-asserted-by":"publisher","first-page":"4075","DOI":"10.1007\/s00253-014-5552-7","article-title":"O-antigen polymerase adopts a distributive mechanism for lipopolysaccharide biosynthesis","volume":"98","author":"Zhao","year":"2014","journal-title":"Appl. Microbiol. Biotechnol."},{"key":"B79","doi-asserted-by":"publisher","first-page":"100148","DOI":"10.1016\/j.imj.2024.100148","article-title":"Leveraging computer-aided design and artificial intelligence to develop a next-generation multi-epitope tuberculosis vaccine candidate","volume":"3","author":"Zhuang","year":"2024","journal-title":"Infect. Med."}],"container-title":["Frontiers in Microbiology"],"original-title":[],"link":[{"URL":"https:\/\/www.frontiersin.org\/articles\/10.3389\/fmicb.2026.1909795\/full","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,8,19]],"date-time":"2026-08-19T05:47:38Z","timestamp":1787118458000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.frontiersin.org\/articles\/10.3389\/fmicb.2026.1909795\/full"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,8,19]]},"references-count":79,"alternative-id":["10.3389\/fmicb.2026.1909795"],"URL":"https:\/\/doi.org\/10.3389\/fmicb.2026.1909795","relation":{},"ISSN":["1664-302X"],"issn-type":[{"value":"1664-302X","type":"electronic"}],"subject":[],"published":{"date-parts":[[2026,8,19]]},"article-number":"1909795"}}