{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,9,26]],"date-time":"2026-09-26T03:40:18Z","timestamp":1790394018775,"version":"4.1.0"},"reference-count":249,"publisher":"MDPI AG","issue":"11","license":[{"start":{"date-parts":[[2019,11,19]],"date-time":"2019-11-19T00:00:00Z","timestamp":1574121600000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"DOI":"10.13039\/501100001700","name":"Ministry of Education, Culture, Sports, Science and Technology","doi-asserted-by":"publisher","award":["15H02657"],"award-info":[{"award-number":["15H02657"]}],"id":[{"id":"10.13039\/501100001700","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100001700","name":"Ministry of Education, Culture, Sports, Science and Technology","doi-asserted-by":"publisher","award":["16H06279"],"award-info":[{"award-number":["16H06279"]}],"id":[{"id":"10.13039\/501100001700","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100001700","name":"Ministry of Education, Culture, Sports, Science and Technology","doi-asserted-by":"publisher","award":["16H05191"],"award-info":[{"award-number":["16H05191"]}],"id":[{"id":"10.13039\/501100001700","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100001700","name":"Ministry of Education, Culture, Sports, Science and Technology","doi-asserted-by":"publisher","award":["18KK0266"],"award-info":[{"award-number":["18KK0266"]}],"id":[{"id":"10.13039\/501100001700","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/100000002","name":"National Institutes of Health","doi-asserted-by":"publisher","award":["DK62813"],"award-info":[{"award-number":["DK62813"]}],"id":[{"id":"10.13039\/100000002","id-type":"DOI","asserted-by":"publisher"}]}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":["Toxins"],"abstract":"<jats:p>Helicobacter pylori colonizes the gastric epithelial cells of at least half of the world\u2019s population, and it is the strongest risk factor for developing gastric complications like chronic gastritis, ulcer diseases, and gastric cancer. To successfully colonize and establish a persistent infection, the bacteria must overcome harsh gastric conditions. H. pylori has a well-developed mechanism by which it can survive in a very acidic niche. Despite bacterial factors, gastric environmental factors and host genetic constituents together play a co-operative role for gastric pathogenicity. The virulence factors include bacterial colonization factors BabA, SabA, OipA, and HopQ, and the virulence factors necessary for gastric pathogenicity include the effector proteins like CagA, VacA, HtrA, and the outer membrane vesicles. Bacterial factors are considered more important. Here, we summarize the recent information to better understand several bacterial virulence factors and their role in the pathogenic mechanism.<\/jats:p>","DOI":"10.3390\/toxins11110677","type":"journal-article","created":{"date-parts":[[2019,11,19]],"date-time":"2019-11-19T11:30:17Z","timestamp":1574163017000},"page":"677","update-policy":"https:\/\/doi.org\/10.3390\/mdpi_crossmark_policy","source":"Crossref","is-referenced-by-count":264,"title":["Helicobacter pylori Virulence Factors Exploiting Gastric Colonization and its Pathogenicity"],"prefix":"10.3390","volume":"11","author":[{"ORCID":"https:\/\/orcid.org\/0000-0003-1846-1377","authenticated-orcid":false,"given":"Shamshul","family":"Ansari","sequence":"first","affiliation":[{"name":"Department of Microbiology, Chitwan Medical College and Teaching Hospital, Bharatpur 44200, Chitwan, Nepal"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-1222-5819","authenticated-orcid":false,"given":"Yoshio","family":"Yamaoka","sequence":"additional","affiliation":[{"name":"Department of Environmental and Preventive Medicine, Oita University Faculty of Medicine, Idaigaoka, Hasama-machi, Yufu, Oita 879-5593, Japan"},{"name":"Global Oita Medical Advanced Research Center for Health, Idaigaoka, Hasama-machi, Yufu, Oita 879-5593, Japan"},{"name":"Department of Medicine, Gastroenterology and Hepatology Section, Baylor College of Medicine, 2002 Holcombe Blvd., Houston, TX 77030, USA"},{"name":"Borneo Medical and Health Research Centre, Universiti Malaysia Sabah, Kota Kinabaru, Sabah 88400, Malaysia"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"1968","published-online":{"date-parts":[[2019,11,19]]},"reference":[{"key":"ref_1","doi-asserted-by":"crossref","first-page":"162","DOI":"10.1126\/science.aad2545","article-title":"The 5300-year-old Helicobacter pylori genome of the Iceman","volume":"351","author":"Maixner","year":"2016","journal-title":"Science"},{"key":"ref_2","doi-asserted-by":"crossref","first-page":"196","DOI":"10.2183\/pjab.93.013","article-title":"Structure and function of Helicobacter pylori CagA, the first-identified bacterial protein involved in human cancer","volume":"93","author":"Hatakeyama","year":"2017","journal-title":"Proc. Jpn. Acad. Ser. B Phys. Biol. Sci."},{"key":"ref_3","doi-asserted-by":"crossref","first-page":"420","DOI":"10.1053\/j.gastro.2017.04.022","article-title":"Global Prevalence of Helicobacter pylori Infection: Systematic Review and Meta-Analysis","volume":"153","author":"Hooi","year":"2017","journal-title":"Gastroenterology"},{"key":"ref_4","doi-asserted-by":"crossref","first-page":"868","DOI":"10.1111\/apt.14561","article-title":"Systematic review with meta-analysis: The worldwide prevalence of Helicobacter pylori infection","volume":"47","author":"Zamani","year":"2018","journal-title":"Aliment. Pharmacol. Ther."},{"key":"ref_5","doi-asserted-by":"crossref","first-page":"38","DOI":"10.1080\/09674845.2016.1150666","article-title":"Intrafamilial transmission of Helicobacter pylori: Genotyping of faecal samples","volume":"73","author":"Mamishi","year":"2016","journal-title":"Br. J. Biomed. Sci."},{"key":"ref_6","doi-asserted-by":"crossref","first-page":"82","DOI":"10.4269\/ajtmh.15-0297","article-title":"Serologic Evidence for Fecal-Oral Transmission of Helicobacter pylori","volume":"94","author":"Bui","year":"2016","journal-title":"Am. J. Trop. Med. Hyg."},{"key":"ref_7","doi-asserted-by":"crossref","first-page":"e12386","DOI":"10.1111\/hel.12386","article-title":"Survival of Helicobacter pylori in gastric acidic territory","volume":"22","author":"Ansari","year":"2017","journal-title":"Helicobacter"},{"key":"ref_8","doi-asserted-by":"crossref","first-page":"199","DOI":"10.1007\/s10096-016-2810-x","article-title":"Population attributable burden of Helicobacter pylori-related gastric cancer, coronary heart disease, and ischemic stroke in China","volume":"36","author":"Jiang","year":"2017","journal-title":"Eur. J. Clin. Microbiol. Infect. Dis."},{"key":"ref_9","first-page":"59","article-title":"The prevalence of Helicobacter pylori in peptic ulcer disease","volume":"9","author":"Kuipers","year":"1995","journal-title":"Aliment. Pharmacol. Ther."},{"key":"ref_10","doi-asserted-by":"crossref","first-page":"1244","DOI":"10.1053\/gast.1996.v110.pm8613015","article-title":"Relationship between Helicobacter pylori eradication and reduced duodenal and gastric ulcer recurrence: A review","volume":"110","author":"Hopkins","year":"1996","journal-title":"Gastroenterology"},{"key":"ref_11","doi-asserted-by":"crossref","first-page":"64","DOI":"10.1053\/j.gastro.2015.09.004","article-title":"Pathobiology of Helicobacter pylori-induced gastric cancer","volume":"150","author":"Amieva","year":"2016","journal-title":"Gastroenterology"},{"key":"ref_12","doi-asserted-by":"crossref","unstructured":"Schoep, T.D., Fulurija, A., Good, F., Lu, W., Himbeck, R.P., Schwan, C., Choi, S.S., Berg, D.E., Mittl, P.R., and Benghezal, M. (2010). Surface properties of Helicobacter pylori urease complex are essential for persistence. PLoS ONE, 5.","DOI":"10.1371\/journal.pone.0015042"},{"key":"ref_13","doi-asserted-by":"crossref","first-page":"778","DOI":"10.1136\/jcp.42.7.778-b","article-title":"Ultrastructural localisation of urease in outer membrane and periplasm of Campylobacter pylori","volume":"42","author":"Bode","year":"1989","journal-title":"J. Clin. Pathol."},{"key":"ref_14","doi-asserted-by":"crossref","first-page":"339","DOI":"10.1172\/JCI9351","article-title":"Local pH elevation mediated by the intra-bacterial urease of Helicobacter pylori co-cultured with gastric cells","volume":"106","author":"Athmann","year":"2000","journal-title":"J. Clin. Investig."},{"key":"ref_15","doi-asserted-by":"crossref","first-page":"729","DOI":"10.1128\/JB.187.2.729-738.2005","article-title":"The periplasmic alpha-carbonic anhydrase activity of Helicobacter pylori is essential for acid acclimation","volume":"187","author":"Marcus","year":"2005","journal-title":"J. Bacteriol."},{"key":"ref_16","doi-asserted-by":"crossref","first-page":"482","DOI":"10.1126\/science.287.5452.482","article-title":"A H+-gated urea channel: The link between Helicobacter pylori urease and gastric colonization","volume":"287","author":"Weeks","year":"2000","journal-title":"Science"},{"key":"ref_17","doi-asserted-by":"crossref","unstructured":"Debowski, A.W., Walton, S.M., Chua, E.-G., Tay, A.C.-Y., Liao, T., Lamichhane, B., Himbeck, R., Stubbs, K.A., Marshall, B.J., and Fulurija, A. (2017). Helicobacter pylori gene silencing in vivo demonstrates urease is essential for chronic infection. PLoS Pathog., 13.","DOI":"10.1371\/journal.ppat.1006464"},{"key":"ref_18","doi-asserted-by":"crossref","first-page":"3267","DOI":"10.1099\/mic.0.28049-0","article-title":"Production of ammonium by Helicobacter pylori mediates occludin processing and disruption of tight junctions in Caco-2 cells","volume":"151","author":"Lytton","year":"2005","journal-title":"Microbiology"},{"key":"ref_19","doi-asserted-by":"crossref","first-page":"236","DOI":"10.1053\/j.gastro.2008.10.011","article-title":"Helicobacter pylori dysregulation of gastric epithelial tight junctions by urease-mediated myosin II activation","volume":"136","author":"Wroblewski","year":"2009","journal-title":"Gastroenterology"},{"key":"ref_20","doi-asserted-by":"crossref","first-page":"4378","DOI":"10.1128\/IAI.68.8.4378-4383.2000","article-title":"Helicobacter pylori urease suppresses bactericidal activity of peroxynitrite via carbon dioxide production","volume":"68","author":"Kuwahara","year":"2000","journal-title":"Infect. Immun."},{"key":"ref_21","doi-asserted-by":"crossref","first-page":"1883","DOI":"10.3389\/fmicb.2017.01883","article-title":"A New Role for Helicobacter pylori Urease: Contributions to Angiogenesis","volume":"8","author":"Uberti","year":"2017","journal-title":"Front. Microbiol."},{"key":"ref_22","doi-asserted-by":"crossref","unstructured":"Valenzuela-Valderrama, M., Cerda-Opazo, P., Backert, S., Gonz\u00e1lez, M.F., Carrasco-V\u00e9liz, N., Jorquera-Cordero, C., Wehinger, S., Canales, J., Bravo, D., and Quest, A.F.G. (2019). The Helicobacter pylori Urease Virulence Factor Is Required for the Induction of Hypoxia-Induced Factor-1\u03b1 in Gastric Cells. Cancers, 11.","DOI":"10.3390\/cancers11060799"},{"key":"ref_23","doi-asserted-by":"crossref","first-page":"e12573","DOI":"10.1111\/hel.12573","article-title":"Helicobacter pylori urease induces pro-inflammatory effects and differentiation of human endothelial cells: Cellular and molecular mechanism","volume":"24","author":"Uberti","year":"2019","journal-title":"Helicobacter"},{"key":"ref_24","doi-asserted-by":"crossref","first-page":"1918","DOI":"10.4049\/jimmunol.165.4.1918","article-title":"Helicobacter pylori urease binds to class II MHC on gastric epithelial cells and induces their apoptosis","volume":"165","author":"Fan","year":"2000","journal-title":"J. Immunol."},{"key":"ref_25","doi-asserted-by":"crossref","first-page":"88","DOI":"10.1111\/mmi.13218","article-title":"Helicobacter pylori strains vary cell shape and flagellum number to maintain robust motility in viscous environments","volume":"99","author":"Hardcastle","year":"2016","journal-title":"Mol. Microbiol."},{"key":"ref_26","doi-asserted-by":"crossref","first-page":"822","DOI":"10.1016\/j.cell.2010.03.046","article-title":"Relaxation of peptidoglycan cross-linking promotes Helicobacter pylori\u2019s helical shape and stomach colonization","volume":"141","author":"Sycuro","year":"2010","journal-title":"Cell"},{"key":"ref_27","doi-asserted-by":"crossref","first-page":"596","DOI":"10.1016\/j.mib.2007.09.009","article-title":"Bacterial morphology: Why have different shapes?","volume":"10","author":"Young","year":"2007","journal-title":"Curr. Opin. Microbiol."},{"key":"ref_28","doi-asserted-by":"crossref","first-page":"1601","DOI":"10.1128\/iai.61.5.1601-1610.1993","article-title":"Pathogenicity of Helicobacter pylori: A perspective","volume":"61","author":"Lee","year":"1993","journal-title":"Infect. Immun."},{"key":"ref_29","doi-asserted-by":"crossref","unstructured":"Sycuro, L.K., Wyckoff, T.J., Biboy, J., Born, P., Pincus, Z., Vollmer, W., and Salama, N.R. (2012). Multiple peptidoglycan modification networks modulate Helicobacter pylori\u2019s cell shape, motility, and colonization potential. PLoS Pathog., 8.","DOI":"10.1371\/journal.ppat.1002603"},{"key":"ref_30","doi-asserted-by":"crossref","first-page":"168","DOI":"10.1016\/S0966-842X(00)88913-1","article-title":"The complex flagella of gastric Helicobacter species","volume":"3","author":"Suerbaum","year":"1995","journal-title":"Trends Microbiol."},{"key":"ref_31","doi-asserted-by":"crossref","first-page":"3529","DOI":"10.1128\/IAI.71.6.3529-3539.2003","article-title":"pH-regulated gene expression of the gastric pathogen Helicobacter pylori","volume":"71","author":"Merrell","year":"2003","journal-title":"Infect. Immun."},{"key":"ref_32","doi-asserted-by":"crossref","first-page":"411","DOI":"10.1111\/j.1523-5378.2012.00974.x","article-title":"Higher Motility Enhances Bacterial Density and Inflammatory Response in Dyspeptic Patients Infected with Helicobacter pylori","volume":"17","author":"Kao","year":"2012","journal-title":"Helicobacter"},{"key":"ref_33","doi-asserted-by":"crossref","unstructured":"Forstneri\u010d, V., Ivi\u010dak-Kocjan, K., Plaper, T., Jerala, R., and Ben\u010dina, M. (2017). The role of the Cterminal D0 domain of flagellin in activation of Toll like receptor 5. PLoS Pathog., 13.","DOI":"10.1371\/journal.ppat.1006574"},{"key":"ref_34","doi-asserted-by":"crossref","first-page":"4158","DOI":"10.3748\/wjg.v23.i23.4158","article-title":"Helicobacter pylori BabA in adaptation for gastric colonization","volume":"23","author":"Ansari","year":"2017","journal-title":"World J. Gastroenterol."},{"key":"ref_35","doi-asserted-by":"crossref","first-page":"25256","DOI":"10.1074\/jbc.M111.233601","article-title":"BabA-mediated adherence is a potentiator of the Helicobacter pylori type IV secretion system activity","volume":"286","author":"Ishijima","year":"2011","journal-title":"J. Biol. Chem."},{"key":"ref_36","doi-asserted-by":"crossref","first-page":"14944","DOI":"10.1073\/pnas.1100959108","article-title":"Carcinogenic bacterial pathogen Helicobacter pylori triggers DNA double-strand breaks and a DNA damage response in its host cells","volume":"108","author":"Toller","year":"2011","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_37","doi-asserted-by":"crossref","first-page":"1519","DOI":"10.1128\/IAI.72.3.1519-1529.2004","article-title":"Helicobacter pylori and complex gangliosides","volume":"72","author":"Roche","year":"2004","journal-title":"Infect. Immun."},{"key":"ref_38","doi-asserted-by":"crossref","first-page":"414","DOI":"10.1053\/gast.2002.34781","article-title":"Importance of Helicobacter pylori oipA in clinical presentation, gastric inflammation, and mucosal interleukin 8 production","volume":"23","author":"Yamaoka","year":"2002","journal-title":"Gastroenterology"},{"key":"ref_39","doi-asserted-by":"crossref","first-page":"e8036","DOI":"10.1038\/s41598-017-08176-7","article-title":"Binding of the Helicobacter pylori OipA causes apoptosis of host cells via modulation of Bax\/Bcl-2 levels","volume":"7","author":"Teymournejad","year":"2017","journal-title":"Sci. Rep."},{"key":"ref_40","doi-asserted-by":"crossref","first-page":"e1553487","DOI":"10.1080\/2162402X.2018.1553487","article-title":"The Helicobacter pylori HopQ outer membrane protein inhibits immune cell activities","volume":"8","author":"Gur","year":"2019","journal-title":"OncoImmunology"},{"key":"ref_41","doi-asserted-by":"crossref","first-page":"1190","DOI":"10.1111\/j.1742-4658.2011.08035.x","article-title":"Role of the cag pathogenicity island encoded type IV secretion system in Helicobacter pylori pathogenesis","volume":"278","author":"Tegtmeyer","year":"2011","journal-title":"FEBS J."},{"key":"ref_42","doi-asserted-by":"crossref","first-page":"239","DOI":"10.1007\/s00535-009-0014-1","article-title":"Helicobacter pylori and gastric carcinogenesis","volume":"44","author":"Hatakeyama","year":"2009","journal-title":"J. Gastroenterol."},{"key":"ref_43","doi-asserted-by":"crossref","first-page":"1337","DOI":"10.1046\/j.1365-2958.2001.02714.x","article-title":"Systematic mutagenesis of the Helicobacter pylori cag pathogenicity island: Essential genes for cagA translocation in host cells and induction of interleukin-8","volume":"42","author":"Fischer","year":"2001","journal-title":"Mol. Microbiol."},{"key":"ref_44","doi-asserted-by":"crossref","first-page":"207","DOI":"10.1016\/j.mib.2006.02.008","article-title":"Type IV secretion systems and their effectors in bacterial pathogenesis","volume":"9","author":"Backert","year":"2006","journal-title":"Curr. Opin. Microbiol."},{"key":"ref_45","doi-asserted-by":"crossref","unstructured":"Barrozo, R.M., Cooke, C.L., Hansen, L.M., Lam, A.M., Gaddy, J.A., Johnson, E.M., Cariaga, T.A., Suarez, G., Peek, R.M., and Cover, T.L. (2013). Functional plasticity in the type IV secretion system of Helicobacter pylori. PLoS Pathog., 9.","DOI":"10.1371\/journal.ppat.1003189"},{"key":"ref_46","doi-asserted-by":"crossref","first-page":"554","DOI":"10.1111\/cmi.12055","article-title":"A novel NOD1- and CagA-independent pathway of interleukin-8 induction mediated by the Helicobacter pylori type IV secretion system","volume":"15","author":"Gorrell","year":"2013","journal-title":"Cell. Microbiol."},{"key":"ref_47","doi-asserted-by":"crossref","first-page":"1164","DOI":"10.1053\/j.gastro.2016.08.014","article-title":"CagY is an Immune-Sensitive Regulator of the Helicobacter pylori Type IV Secretion System","volume":"151","author":"Barrozo","year":"2016","journal-title":"Gastroenterology"},{"key":"ref_48","doi-asserted-by":"crossref","first-page":"22","DOI":"10.1007\/s00284-016-1016-x","article-title":"The Helicobacter pylori Cag Pathogenicity Island Protein Cag1 is Associated with the Function of T4SS","volume":"73","author":"Wang","year":"2016","journal-title":"Curr. Microbiol."},{"key":"ref_49","doi-asserted-by":"crossref","first-page":"862","DOI":"10.1038\/nature06187","article-title":"Helicobacter exploits integrin for type IV secretion and kinase activation","volume":"449","author":"Kwok","year":"2007","journal-title":"Nature"},{"key":"ref_50","doi-asserted-by":"crossref","first-page":"186","DOI":"10.1556\/1886.2016.00020","article-title":"A C-terminal coiled-coil region of cagL is responsible for Helicobacter pylori-induced IL-8 expression","volume":"6","author":"Wiedemann","year":"2016","journal-title":"Eur. J. Microbiol. Immunol."},{"key":"ref_51","doi-asserted-by":"crossref","first-page":"163","DOI":"10.1111\/j.1523-5378.2010.00759.x","article-title":"The versatility of Helicobacter pylori CagA effector protein functions: The master key hypothesis","volume":"15","author":"Backert","year":"2010","journal-title":"Helicobacter"},{"key":"ref_52","doi-asserted-by":"crossref","first-page":"683","DOI":"10.1126\/science.1067147","article-title":"SHP-2 tyrosine phosphatase as an intracellular target of Helicobacter pylori CagA protein","volume":"295","author":"Higashi","year":"2002","journal-title":"Science"},{"key":"ref_53","doi-asserted-by":"crossref","first-page":"306","DOI":"10.1016\/j.chom.2014.02.008","article-title":"Helicobacter pylori CagA and gastric cancer: A paradigm for hit- and-run carcinogenesis","volume":"15","author":"Hatakeyama","year":"2014","journal-title":"Cell Host Microbe"},{"key":"ref_54","doi-asserted-by":"crossref","first-page":"523","DOI":"10.1007\/s12192-016-0680-x","article-title":"The Helicobacter pylori cytotoxin CagA is essential for suppressing host heat shock protein expression","volume":"21","author":"Lang","year":"2016","journal-title":"Cell Stress Chaperones"},{"key":"ref_55","doi-asserted-by":"crossref","first-page":"231","DOI":"10.1016\/j.micpath.2009.08.002","article-title":"Inhibition of heat shock protein expression by Helicobacter pylori","volume":"47","author":"Axsen","year":"2009","journal-title":"Microb. Pathog."},{"key":"ref_56","doi-asserted-by":"crossref","first-page":"11","DOI":"10.1046\/j.1462-5822.2002.00163.x","article-title":"Cell vacuolization induced by Helicobacter pylori VacA cytotoxin does not depend on late endosomal SNAREs","volume":"4","author":"Moschioni","year":"2002","journal-title":"Cell. Microbiol."},{"key":"ref_57","doi-asserted-by":"crossref","first-page":"2535","DOI":"10.1128\/IAI.01370-10","article-title":"Helicobacter pylori VacA induces programmed necrosis in gastric epithelial cells","volume":"79","author":"Radin","year":"2011","journal-title":"Infect. Immun."},{"key":"ref_58","doi-asserted-by":"crossref","unstructured":"Akazawa, Y., Isomoto, H., Matsushima, K., Kanda, T., Minami, H., Yamaghchi, N., Taura, N., Shiozawa, K., Ohnita, K., and Takeshima, F. (2013). Endoplasmic reticulum stress contributes to Helicobacter pylori VacA-induced apoptosis. PLoS ONE, 8.","DOI":"10.1371\/journal.pone.0082322"},{"key":"ref_59","doi-asserted-by":"crossref","first-page":"523","DOI":"10.1139\/cjm-2013-0021","article-title":"Overexpression of Helicobacter pylori VacA N-terminal fragment induces pro-inflammatory cytokine expression and apoptosis in human monocytic cell line through activation of NF-kappa B","volume":"59","author":"Luo","year":"2013","journal-title":"Can. J. Microbiol."},{"key":"ref_60","doi-asserted-by":"crossref","first-page":"2168","DOI":"10.3760\/cma.j.issn.0366-6999.20122481","article-title":"Effects of broth culture filtrate protein of VacA + Helicobacter pylori on the proliferation and apoptosis of gastric epithelial cells","volume":"126","author":"Zhao","year":"2013","journal-title":"Chin. Med. J."},{"key":"ref_61","doi-asserted-by":"crossref","first-page":"3207","DOI":"10.1038\/s41419-017-0011-x","article-title":"Helicobacter pylori VacA induces autophagic cell death in gastric epithelial cells via the endoplasmic reticulum stress pathway","volume":"8","author":"Zhu","year":"2017","journal-title":"Cell Death Dis."},{"key":"ref_62","doi-asserted-by":"crossref","first-page":"4","DOI":"10.1186\/s12964-017-0162-5","article-title":"Bacterial serine protease HtrA as a promising new target for antimicrobial therapy?","volume":"15","author":"Wessler","year":"2017","journal-title":"Cell Commun. Signal."},{"key":"ref_63","doi-asserted-by":"crossref","first-page":"e12845","DOI":"10.1111\/cmi.12845","article-title":"Extracellular HtrA serine proteases: An emerging new strategy in bacterial pathogenesis","volume":"20","author":"Backert","year":"2018","journal-title":"Cell. Microbiol."},{"key":"ref_64","doi-asserted-by":"crossref","first-page":"10115","DOI":"10.1074\/jbc.C111.333419","article-title":"Distinct roles of secreted HtrA proteases from Gram-negative pathogens in cleaving the junctional protein and tumor suppressor E-cadherin","volume":"287","author":"Hoy","year":"2012","journal-title":"J. Biol. Chem."},{"key":"ref_65","doi-asserted-by":"crossref","first-page":"798","DOI":"10.1038\/embor.2010.114","article-title":"Helicobacter pylori HtrA is a new secreted virulence factor that cleaves E-Cadherin to disrupt intercellular adhesion","volume":"11","author":"Hoy","year":"2010","journal-title":"EMBO J."},{"key":"ref_66","doi-asserted-by":"crossref","first-page":"10244","DOI":"10.1002\/anie.201504035","article-title":"Fragmented-Based De-Novo Design Reveals a Small-Molecule Inhibitor of Helicobacter pylori HtrA","volume":"54","author":"Perna","year":"2015","journal-title":"Angew. Chem. Int. Ed."},{"key":"ref_67","doi-asserted-by":"crossref","first-page":"23264","DOI":"10.1038\/srep23264","article-title":"Identification of E-cadherin signature motifs functioning as cleavage sites for Helicobacter pylori HtrA","volume":"6","author":"Schmidt","year":"2016","journal-title":"Sci. Rep."},{"key":"ref_68","doi-asserted-by":"crossref","first-page":"925","DOI":"10.1111\/mmi.13276","article-title":"Characterization of worldwide Helicobacter pylori strains reveals genetic conservation and essentiality of serine protease HtrA","volume":"99","author":"Tegtmeyer","year":"2016","journal-title":"Mol. Microbiol."},{"key":"ref_69","doi-asserted-by":"crossref","first-page":"199","DOI":"10.3748\/wjg.v12.i2.199","article-title":"E-cadherin in gastric cancer","volume":"12","author":"Chan","year":"2006","journal-title":"World J. Gastroenterol."},{"key":"ref_70","doi-asserted-by":"crossref","first-page":"552","DOI":"10.1016\/j.chom.2017.09.005","article-title":"Helicobacter pylori Employs a Unique Basolateral Type IV Secretion Mechanism for CagA Delivery","volume":"22","author":"Tegtmeyer","year":"2017","journal-title":"Cell Host Microbe"},{"key":"ref_71","doi-asserted-by":"crossref","first-page":"60","DOI":"10.15698\/mic2018.01.611","article-title":"A novel basolateral type IV secretion model for the CagA oncoprotein of Helicobacter pylori","volume":"5","author":"Wessler","year":"2018","journal-title":"Microb. Cell"},{"key":"ref_72","doi-asserted-by":"crossref","first-page":"1837","DOI":"10.3389\/fmicb.2018.01837","article-title":"Helicobacter pylori outer membrane vesicles protect the pathogen from reactive oxygen species of the respiratory burst","volume":"9","author":"Lekmeechai","year":"2018","journal-title":"Front. Microbiol."},{"key":"ref_73","doi-asserted-by":"crossref","first-page":"607","DOI":"10.1016\/j.resmic.2012.10.020","article-title":"Offense and defense: Microbial membrane vesicles play both ways","volume":"163","author":"Macdonald","year":"2012","journal-title":"Res. Microbiol."},{"key":"ref_74","doi-asserted-by":"crossref","first-page":"5012","DOI":"10.1016\/j.vaccine.2015.05.014","article-title":"Altered host immune responses to membrane vesicles from Salmonella and gram-negative pathogens","volume":"33","author":"Laughlin","year":"2015","journal-title":"Vaccine"},{"key":"ref_75","doi-asserted-by":"crossref","first-page":"2162","DOI":"10.1128\/IAI.00190-16","article-title":"Crude preparations of Helicobacter pylori outer membrane vesicles induce upregulation of Heme Oxygenase-1 via activating Akt-Nrf2 and mTOR-IkappaB kinase-NF-kappaB pathways in dendritic cells","volume":"84","author":"Ko","year":"2016","journal-title":"Infect. Immun."},{"key":"ref_76","doi-asserted-by":"crossref","first-page":"443","DOI":"10.1046\/j.1365-2958.2003.03305.x","article-title":"A novel apoptosis-inducing protein from Helicobacter pylori","volume":"l47","author":"Shibayama","year":"2003","journal-title":"Mol. Microbiol."},{"key":"ref_77","doi-asserted-by":"crossref","first-page":"562","DOI":"10.1016\/j.bbrc.2007.02.021","article-title":"\u03b3-Glutamyltranspeptidase of Helicobacter pylori induces mitochondria-mediated apoptosis in AGS cells","volume":"355","author":"Kim","year":"2007","journal-title":"Biochem. Biophys. Res. Commun."},{"key":"ref_78","doi-asserted-by":"crossref","first-page":"1131","DOI":"10.1093\/infdis\/jit286","article-title":"Helicobacter pylori-induced loss of survivin and gastric cell viability is attributable to secreted bacterial gamma-glutamyl transpeptidase activity","volume":"208","author":"Valenzuela","year":"2013","journal-title":"J. Infect. Dis."},{"key":"ref_79","doi-asserted-by":"crossref","unstructured":"Ling, S.S.M., Khoo, L.H.B., Hwang, L.A., Yeoh, K.G., and Ho, B. (2015). Instrumental role of Helicobacter pylori \u03b3-glutamyl transpeptidase in VacA-dependent vacuolation in gastric epithelial cells. PLoS ONE, 10.","DOI":"10.1371\/journal.pone.0131460"},{"key":"ref_80","doi-asserted-by":"crossref","first-page":"317","DOI":"10.1016\/j.ijmm.2005.06.003","article-title":"Adherence properties of Helicobacter pylori: Impact on pathogenesis and adaptation to the host","volume":"295","author":"Odenbreit","year":"2005","journal-title":"Int. J. Med. Microbiol."},{"key":"ref_81","doi-asserted-by":"crossref","first-page":"80","DOI":"10.1016\/S0966-842X(02)00038-0","article-title":"The basis of persistent bacterial infections","volume":"11","author":"Rhen","year":"2003","journal-title":"Trends Microbiol."},{"key":"ref_82","doi-asserted-by":"crossref","first-page":"293","DOI":"10.1016\/S0076-6879(06)17020-2","article-title":"Helicobacter pylori adhesion to carbohydrates","volume":"417","author":"Aspholm","year":"2006","journal-title":"Methods Enzymol."},{"key":"ref_83","doi-asserted-by":"crossref","first-page":"373","DOI":"10.1126\/science.279.5349.373","article-title":"Helicobacter pylori adhesin binding fucosylated histo-blood group antigens revealed by retagging","volume":"279","author":"Ilver","year":"1998","journal-title":"Science"},{"key":"ref_84","doi-asserted-by":"crossref","first-page":"629","DOI":"10.1006\/jmbi.2001.5311","article-title":"Concerted evolution between duplicated genetic elements in Helicobacter pylori","volume":"316","author":"Pride","year":"2002","journal-title":"J. Mol. Biol."},{"key":"ref_85","doi-asserted-by":"crossref","first-page":"1892","DOI":"10.1126\/science.8018146","article-title":"Attachment of Helicobacter pylori to human gastric epithelium mediated by blood group antigens","volume":"262","author":"Falk","year":"1993","journal-title":"Science"},{"key":"ref_86","doi-asserted-by":"crossref","first-page":"3429","DOI":"10.1128\/IAI.72.6.3429-3435.2004","article-title":"Heterogeneity among Helicobacter pylori strains in expression of the outer membrane protein BabA","volume":"72","author":"Hennig","year":"2004","journal-title":"Infect. Immun."},{"key":"ref_87","doi-asserted-by":"crossref","first-page":"519","DOI":"10.1126\/science.1098801","article-title":"Functional adaptation of BabA, the H. pylori ABO blood group antigen binding adhesin","volume":"305","author":"Dailide","year":"2004","journal-title":"Science"},{"key":"ref_88","doi-asserted-by":"crossref","first-page":"1160","DOI":"10.1128\/IAI.69.2.1160-1171.2001","article-title":"Allelic Variation within Helicobacter pylori babA and babB","volume":"69","author":"Pride","year":"2001","journal-title":"Infect. Immun."},{"key":"ref_89","doi-asserted-by":"crossref","first-page":"2231","DOI":"10.1099\/mic.0.2007\/016055-0","article-title":"Expression of the Helicobacter pylori adhesin SabA is controlled via phase variation and the ArsRS signal transduction system","volume":"154","author":"Goodwin","year":"2008","journal-title":"Microbiology"},{"key":"ref_90","doi-asserted-by":"crossref","first-page":"2106","DOI":"10.1073\/pnas.0308573100","article-title":"Modification of Helicobacter pylori outer membrane protein expression during experimental infection of rhesus macaques","volume":"101","author":"Solnick","year":"2004","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_91","doi-asserted-by":"crossref","first-page":"12778","DOI":"10.1073\/pnas.96.22.12778","article-title":"Clinical relevance of the Helicobacter pylori gene for blood-group antigen-binding adhesin","volume":"96","author":"Gerhard","year":"1999","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_92","doi-asserted-by":"crossref","first-page":"151","DOI":"10.1016\/j.femsim.2004.10.010","article-title":"Correlation of the Helicobacter pylori adherence factor BabA with duodenal ulcer disease in four European countries","volume":"44","author":"Olfat","year":"2005","journal-title":"FEMS Immunol. Med. Microbiol."},{"key":"ref_93","doi-asserted-by":"crossref","first-page":"1699","DOI":"10.1080\/21505594.2018.1532243","article-title":"BabA-mediated adherence of pediatric ulcerogenic H. pylori strains to gastric mucins at neutral and acidic pH","volume":"9","author":"Rocha","year":"2018","journal-title":"Virulence"},{"key":"ref_94","doi-asserted-by":"crossref","first-page":"738","DOI":"10.1080\/21505594.2018.1440171","article-title":"Helicobacter pylori SabA binding gangliosides of human stomach","volume":"9","author":"Benktander","year":"2018","journal-title":"Virulence"},{"key":"ref_95","doi-asserted-by":"crossref","first-page":"36","DOI":"10.1111\/j.1572-0241.2006.00358.x","article-title":"Interaction between host gastric Sialy-Lewis-X and H. pylori SabA enhances H. pylori density in patients lacking gastric Lewis B antigen","volume":"101","author":"Sheu","year":"2006","journal-title":"Am. J. Gastroenterol."},{"key":"ref_96","doi-asserted-by":"crossref","first-page":"1050","DOI":"10.1111\/j.1365-2958.2012.08073.x","article-title":"Regulation of Helicobacter pylori adherence by gene conversion","volume":"84","author":"Talarico","year":"2012","journal-title":"Mol. Microbiol."},{"key":"ref_97","doi-asserted-by":"crossref","first-page":"775","DOI":"10.1136\/gut.2005.083014","article-title":"Helicobacter pylori outer membrane proteins and gastroduodenal disease","volume":"55","author":"Yamaoka","year":"2006","journal-title":"Gut"},{"key":"ref_98","doi-asserted-by":"crossref","first-page":"2228","DOI":"10.1111\/j.1440-1746.2007.04831.x","article-title":"Clinical relevance of Helicobacter pylori sabA genotype in Japanese clinical isolates","volume":"22","author":"Yanai","year":"2007","journal-title":"J. Gastroenterol. Hepatol."},{"key":"ref_99","doi-asserted-by":"crossref","first-page":"100","DOI":"10.1016\/j.micpath.2017.06.023","article-title":"Relationship between histopathological status of the Helicobacter pylori infected patients and proteases of H. pylori in isolates carrying diverse virulence genotypes","volume":"110","author":"Gharibi","year":"2017","journal-title":"Microb. Pathog."},{"key":"ref_100","doi-asserted-by":"crossref","first-page":"573","DOI":"10.1126\/science.1069076","article-title":"Helicobacter pylori SabA adhesin in persistent infection and chronic inflammation","volume":"297","author":"Mahdavi","year":"2002","journal-title":"Science"},{"key":"ref_101","doi-asserted-by":"crossref","first-page":"419","DOI":"10.1007\/s004280050269","article-title":"Helicobacter pylori infection produces reversible glycosylation changes to gastric mucins","volume":"433","author":"Ota","year":"1998","journal-title":"Virchows Arch."},{"key":"ref_102","doi-asserted-by":"crossref","first-page":"36442","DOI":"10.1038\/srep36442","article-title":"Combination of OipA, BabA, and SabA as candidate biomarkers for predicting Helicobacter pylori-related gastric cancer","volume":"6","author":"Su","year":"2016","journal-title":"Sci. Rep."},{"key":"ref_103","doi-asserted-by":"crossref","first-page":"7533","DOI":"10.1073\/pnas.130079797","article-title":"A M(r) 34,000 pro-inflammatory outer membrane protein (oipA) of Helicobacter pylori","volume":"97","author":"Yamaoka","year":"2000","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_104","doi-asserted-by":"crossref","first-page":"ftx113","DOI":"10.1093\/femspd\/ftx113","article-title":"Outer inflammatory protein a (OipA) of Helicobacter pylori is regulated by host cell contact and mediates CagA translocation and interleukin-8 response only in the presence of a functional cag pathogenicity island type IV secretion system","volume":"75","author":"Horridge","year":"2017","journal-title":"Pathog. Dis."},{"key":"ref_105","doi-asserted-by":"crossref","first-page":"1992","DOI":"10.1053\/gast.2002.37074","article-title":"Helicobacter pylori infection in mice: Role of outer membrane proteins in colonization and inflammation","volume":"123","author":"Yamaoka","year":"2002","journal-title":"Gastroenterology"},{"key":"ref_106","doi-asserted-by":"crossref","unstructured":"Braga, L.L.B.C., Batista, M.H.R., de Azevedo, O.G.R., da Silva Costa, K.C., Gomes, A.D., Rocha, G.A., and Queiroz, D.M.M. (2019). oipA \u201con\u201d status of Helicobacter pylori is associated with gastric cancer in NorthEastern Brazil. BMC Cancer, 19.","DOI":"10.1186\/s12885-018-5249-x"},{"key":"ref_107","doi-asserted-by":"crossref","first-page":"93","DOI":"10.1007\/s00203-018-1580-5","article-title":"Status (on\/off) of oipA gene: Their associations with gastritis and gastric cancer and geographic origins","volume":"201","author":"Sallas","year":"2019","journal-title":"Arch. Microbiol."},{"key":"ref_108","doi-asserted-by":"crossref","unstructured":"Liu, J., He, C., Chen, M., Wang, Z., Xing, C., and Yuan, Y. (2013). Association of presence\/absence and on\/off patterns of Helicobacter pylori oipA gene with peptic ulcer disease and gastric cancer risks: A meta-analysis. BMC Infect. Dis., 13.","DOI":"10.1186\/1471-2334-13-555"},{"key":"ref_109","doi-asserted-by":"crossref","first-page":"379","DOI":"10.1158\/0008-5472.CAN-07-0824","article-title":"Regulation of gastric carcinogenesis by Helicobacter pylori virulence factors","volume":"68","author":"Franco","year":"2008","journal-title":"Cancer Res."},{"key":"ref_110","doi-asserted-by":"crossref","first-page":"629","DOI":"10.1038\/nrgastro.2010.154","article-title":"Mechanisms of disease: Helicobacter pylori virulence factors","volume":"7","author":"Yamaoka","year":"2010","journal-title":"Nat. Rev. Gastroenterol. Hepatol."},{"key":"ref_111","doi-asserted-by":"crossref","first-page":"136","DOI":"10.1111\/hel.12107","article-title":"In vitro suppression of dendritic cells by Helicobacter pylori OipA","volume":"19","author":"Teymournejad","year":"2014","journal-title":"Helicobacter"},{"key":"ref_112","doi-asserted-by":"crossref","first-page":"1573","DOI":"10.1111\/j.1462-5822.2008.01156.x","article-title":"Role of type IV secretion in Helicobacter pylori pathogenesis","volume":"10","author":"Backert","year":"2008","journal-title":"Cell. Microbiol."},{"key":"ref_113","doi-asserted-by":"crossref","first-page":"4504","DOI":"10.1128\/JCM.40.12.4504-4511.2002","article-title":"Two different families of hopQ alleles in Helicobacter pylori","volume":"40","author":"Cao","year":"2002","journal-title":"J. Clin. Microbiol."},{"key":"ref_114","doi-asserted-by":"crossref","first-page":"16188","DOI":"10.1038\/nmicrobiol.2016.188","article-title":"Helicobacter pylori exploits human CEACAMs via HopQ fot adherence and translocation of CagA","volume":"2","author":"Koniger","year":"2016","journal-title":"Nat. Microbiol."},{"key":"ref_115","doi-asserted-by":"crossref","first-page":"527","DOI":"10.1016\/j.ijmm.2018.05.003","article-title":"HopQ impacts the integrin alpha5beta1-independent NF-kappaB activation by Helicobacter pylori in CEACAM expressing cells","volume":"308","author":"Feige","year":"2018","journal-title":"Int. J. Med. Microbiol."},{"key":"ref_116","doi-asserted-by":"crossref","first-page":"e98664","DOI":"10.15252\/embj.201798664","article-title":"The Helicobacter pylori adhesin protein HopQ exploits the dimer interface of human CEACAMs to facilitate translocation of the oncoprotein CagA","volume":"37","author":"Bonsor","year":"2018","journal-title":"EMBO J."},{"key":"ref_117","doi-asserted-by":"crossref","first-page":"37","DOI":"10.1016\/j.femsle.2005.07.023","article-title":"Analysis of hopQ alleles in East Asian and Western strains of Helicobacter pylori","volume":"251","author":"Cao","year":"2005","journal-title":"FEMS Microbiol. Lett."},{"key":"ref_118","doi-asserted-by":"crossref","first-page":"601","DOI":"10.3892\/br.2016.634","article-title":"Helicobacter pylori hopQ alleles (type I and II) in gastric cancer","volume":"4","author":"Leylabadlo","year":"2016","journal-title":"Biomed. Rep."},{"key":"ref_119","doi-asserted-by":"crossref","first-page":"57","DOI":"10.1016\/j.meegid.2015.10.027","article-title":"Helicobacter pylori outer membrane protein Q allele distribution is associated with distinct pathologies in Pakistan","volume":"37","author":"Yakoob","year":"2016","journal-title":"Infect. Genet. Evol."},{"key":"ref_120","first-page":"871601","article-title":"High Prevalence of Helicobacter pylori hopQ II Genotype Isolated from Iranian Patients with Gastroduodenal Disorders","volume":"2014","author":"Abadi","year":"2014","journal-title":"J. Pathog."},{"key":"ref_121","doi-asserted-by":"crossref","first-page":"16189","DOI":"10.1038\/nmicrobiol.2016.189","article-title":"Helicobacter pylori adhesin HopQ engages in a virulence-enhancing interaction with human CEACAMs","volume":"2","author":"Javaheri","year":"2016","journal-title":"Nat. Microbiol."},{"key":"ref_122","doi-asserted-by":"crossref","first-page":"977","DOI":"10.1016\/j.ijmm.2018.08.002","article-title":"Thiol-oxidoreductase HP0231 of Helicobacter pylori impacts HopQ-dependent CagA translocation","volume":"308","author":"Grzeszczuk","year":"2018","journal-title":"Int. J. Med. Microbiol."},{"key":"ref_123","doi-asserted-by":"crossref","first-page":"3468","DOI":"10.1017\/S0950268817002023","article-title":"Gastric lymphoma: Association with Helicobacter pylori outer membrane protein Q (HopQ) and cytotoxic-pathogenicity activity island (CPAI) genes","volume":"145","author":"Yakoob","year":"2017","journal-title":"Epidemiol. Infect."},{"key":"ref_124","doi-asserted-by":"crossref","first-page":"211","DOI":"10.1016\/j.tim.2011.01.010","article-title":"Integrin-mediated type IV secretion by Helicobacter: What makes it tick?","volume":"19","author":"Schuelein","year":"2011","journal-title":"Trends Microbiol."},{"key":"ref_125","doi-asserted-by":"crossref","first-page":"955","DOI":"10.2217\/fmb.15.32","article-title":"Composition, structure and function of the Helicobacter pylori cag pathogenicity island encoded type IV secretion system","volume":"10","author":"Backert","year":"2015","journal-title":"Future Microbiol."},{"key":"ref_126","first-page":"e02001","article-title":"Molecular and structural analysis of the Helicobacter pylori cag type IV secretion system core complex","volume":"7","author":"Pyburn","year":"2016","journal-title":"mBio"},{"key":"ref_127","doi-asserted-by":"crossref","first-page":"651","DOI":"10.1007\/s10096-010-0909-z","article-title":"Clinical relevance of cagPAI intactness in Helicobacter pylori isolates from Vietnam","volume":"29","author":"Nguyen","year":"2010","journal-title":"Eur. J. Clin. Microbiol. Inf. Dis."},{"key":"ref_128","doi-asserted-by":"crossref","unstructured":"Sanchez-Zauco, N.A., Torres, J., Perez-Figueroa, G.E., Alvarez-Arellano, L., Camorlinga-Ponce, M., G\u00f3mez, A., Giono-Cerezo, S., and Maldonado-Bernal, C. (2013). Impact of cagPAI and T4SS on the Inflammatory Response of Human Neutrophils to Helicobacter pylori Infection. PLoS ONE, 8.","DOI":"10.1371\/journal.pone.0064623"},{"key":"ref_129","doi-asserted-by":"crossref","first-page":"317","DOI":"10.1038\/nrgastro.2016.68","article-title":"Helicobacter pylori: Pathogenic enablers-toxic relationships in the stomach","volume":"13","author":"Wroblewski","year":"2016","journal-title":"Nat. Rev. Gastroenterol. Hepatol."},{"key":"ref_130","doi-asserted-by":"crossref","first-page":"252","DOI":"10.1016\/j.patbio.2015.09.004","article-title":"Association of CagPAI integrity with severeness of Helicobacter pylori infection in patients with gastritis","volume":"63","author":"Ahmadzadeh","year":"2015","journal-title":"Pathol. Biol."},{"key":"ref_131","doi-asserted-by":"crossref","first-page":"5302","DOI":"10.1128\/JCM.42.11.5302-5308.2004","article-title":"The cag pathogenicity island of Helicobacter pylori is disrupted in the majority of patient isolates from different human populations","volume":"42","author":"Kauser","year":"2004","journal-title":"J. Clin. Microbiol."},{"key":"ref_132","doi-asserted-by":"crossref","first-page":"449","DOI":"10.1128\/CMR.00054-05","article-title":"Pathogenesis of Helicobacter pylori infection","volume":"19","author":"Kusters","year":"2006","journal-title":"Clin. Microbiol. Rev."},{"key":"ref_133","doi-asserted-by":"crossref","unstructured":"Olbermann, P., Josenhans, C., Moodley, Y., Uhr, M., Stamer, C., Vauterin, M., Suerbaum, S., Achtman, M., and Linz, B. (2010). A global overview of the genetic and functional diversity in the Helicobacter pylori cag pathogenicity island. PLoS Genet., 6.","DOI":"10.1371\/journal.pgen.1001069"},{"key":"ref_134","doi-asserted-by":"crossref","first-page":"167","DOI":"10.1016\/j.meegid.2018.02.009","article-title":"Status of Helicobacter pylori cag pathogenicity island (cagPAI) integrity and significance of its individual genes","volume":"59","author":"Markovska","year":"2018","journal-title":"Infect. Genet. Evol."},{"key":"ref_135","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1016\/j.micpath.2017.12.076","article-title":"Preliminary study and bioinformatics analysis on the potential role of CagQ in type IV secretion system of H. pylori","volume":"116","author":"Yao","year":"2018","journal-title":"Microb. Pathog."},{"key":"ref_136","doi-asserted-by":"crossref","first-page":"70","DOI":"10.3389\/fcimb.2012.00070","article-title":"An RGD helper sequence in CagL of Helicobacter pylori assists in interactions with integrins and injection of CagA","volume":"2","author":"Conradi","year":"2012","journal-title":"Front. Cell. Infect. Microbiol."},{"key":"ref_137","doi-asserted-by":"crossref","first-page":"239","DOI":"10.1053\/j.gastro.2010.03.036","article-title":"Helicobacter pylori CagL activates ADAM17 to induce repression of the gastric H, K-ATPase alpha subunit","volume":"139","author":"Saha","year":"2010","journal-title":"Gastroenterology"},{"key":"ref_138","doi-asserted-by":"crossref","first-page":"23515","DOI":"10.1074\/jbc.M109.096214","article-title":"A small fibronectin-mimicking protein from bacteria induces cell spreading and focal adhesion formation","volume":"285","author":"Tegtmeyer","year":"2010","journal-title":"J. Biol. Chem."},{"key":"ref_139","doi-asserted-by":"crossref","first-page":"5","DOI":"10.1186\/s13099-019-0286-9","article-title":"CagL polymorphisms D58\/K59 are predominant in Helicobacter pylori strains isolated from Mexican patients with chronic gastritis","volume":"11","author":"Mendoza","year":"2019","journal-title":"Gut Pathog."},{"key":"ref_140","doi-asserted-by":"crossref","first-page":"1619","DOI":"10.1002\/cam4.1941","article-title":"Genetic diversity and amino acid sequence polymorphism in Helicobacter pylori CagL hypervariable motif and its association with virulence markers and gastroduodenal diseases","volume":"8","author":"Yadegar","year":"2019","journal-title":"Cancer Med."},{"key":"ref_141","doi-asserted-by":"crossref","unstructured":"Tafreshi, M., Zwickel, N., Gorrell, R.J., and Kwok, T. (2015). Preservation of Helicobacter pylori CagA translocation and host cell pro-inflammatory responses in the face of CagL hyper variability at amino acid residues 58\/59. PLoS ONE, 10.","DOI":"10.1371\/journal.pone.0133531"},{"key":"ref_142","doi-asserted-by":"crossref","first-page":"1927","DOI":"10.1093\/infdis\/jiw060","article-title":"Helicobacter pylori CagL hypervariable motif: A global analysis of geographical diversity and association with gastric cancer","volume":"213","author":"Gorrell","year":"2016","journal-title":"J. Infect. Dis."},{"key":"ref_143","doi-asserted-by":"crossref","first-page":"587","DOI":"10.1084\/jem.191.4.587","article-title":"Tyrosine phosphorylated bacterial proteins: Trojan horses for the host cell","volume":"191","author":"Covacci","year":"2000","journal-title":"J. Exp. Med."},{"key":"ref_144","doi-asserted-by":"crossref","unstructured":"Shaffer, C.L., Gaddy, J.A., Loh, J.T., Johnson, E.M., Hill, S., Hennig, E.E., McClain, M.S., McDonald, W.H., and Cover, T.L. (2011). Helicobacter pylori exploits a unique repertoire of type IV secretion system components for pilus assembly at the bacteria-host cell interface. PLoS Pathog., 7.","DOI":"10.1371\/journal.ppat.1002237"},{"key":"ref_145","doi-asserted-by":"crossref","first-page":"1166","DOI":"10.1038\/ni1131","article-title":"Nod1 responds to peptidoglycan delivered by the Helicobacter pylori cag pathogenicity island","volume":"5","author":"Viala","year":"2004","journal-title":"Nat. Immunol."},{"key":"ref_146","doi-asserted-by":"crossref","first-page":"6262","DOI":"10.1038\/onc.2016.158","article-title":"Pathogenic Helicobacter pylori strains translocate DNA and activate TLR9 via the cancer-associated cag type IV secretion system","volume":"35","author":"Varga","year":"2016","journal-title":"Oncogene"},{"key":"ref_147","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1371\/journal.ppat.1006514","article-title":"Helicobacter pylori modulates host cell responses by CagT4SS-dependent translocation of an intermediate metabolite of LPS inner core heptose biosynthesis","volume":"13","author":"Stein","year":"2017","journal-title":"PLoS Pathog."},{"key":"ref_148","doi-asserted-by":"crossref","first-page":"122","DOI":"10.1038\/s41586-018-0433-3","article-title":"Alpha-kinase 1 is a cytosolic innate immune receptor for bacterial ADP heptose","volume":"561","author":"Zhou","year":"2018","journal-title":"Nature"},{"key":"ref_149","doi-asserted-by":"crossref","first-page":"23227","DOI":"10.1038\/srep23227","article-title":"The CagA toxin of Helicobacter pylori: Abundant production but relatively low amount translocated","volume":"6","author":"Haas","year":"2016","journal-title":"Sci. Rep."},{"key":"ref_150","doi-asserted-by":"crossref","first-page":"682","DOI":"10.1016\/j.cell.2010.11.012","article-title":"Ubiquitin: Same molecule, different degradation pathways","volume":"143","author":"Clague","year":"2010","journal-title":"Cell"},{"key":"ref_151","doi-asserted-by":"crossref","first-page":"38","DOI":"10.1038\/s41598-018-37095-4","article-title":"VacA promotes CagA accumulation in gastric epithelial cells during Helicobacter pylori infection","volume":"9","author":"Abdullah","year":"2019","journal-title":"Sci. Rep."},{"key":"ref_152","doi-asserted-by":"crossref","first-page":"44","DOI":"10.1016\/j.micpath.2016.01.005","article-title":"CagA-mediated pathogenesis of Helicobacter pylori","volume":"93","author":"Tohidpour","year":"2016","journal-title":"Microb. Pathog."},{"key":"ref_153","doi-asserted-by":"crossref","first-page":"1402","DOI":"10.3748\/wjg.v20.i6.1402","article-title":"Structural and functional aspects of the Helicobacter pylori secretome","volume":"20","author":"Zanotti","year":"2014","journal-title":"World J. Gastroenterol."},{"key":"ref_154","doi-asserted-by":"crossref","first-page":"745","DOI":"10.1016\/S1097-2765(02)00681-0","article-title":"Grb2 is a key mediator of Helicobacter pylori CagA protein activities","volume":"10","author":"Mimuro","year":"2002","journal-title":"Mol. Cell"},{"key":"ref_155","doi-asserted-by":"crossref","first-page":"2157","DOI":"10.1084\/jem.20100602","article-title":"Conversion of Helicobacter pylori CagA from senescence inducer to oncogenic driver through polarity-dependent regulation of p21","volume":"207","author":"Saito","year":"2010","journal-title":"J. Exp. Med."},{"key":"ref_156","doi-asserted-by":"crossref","first-page":"280","DOI":"10.1186\/s13046-018-0962-5","article-title":"Helicobacter pylori CagA promotes epithelial mesenchymal transition in gastric carcinogenesis via triggering oncogenic YAP pathway","volume":"37","author":"Li","year":"2018","journal-title":"J. Exp. Clin. Cancer Res."},{"key":"ref_157","doi-asserted-by":"crossref","first-page":"14428","DOI":"10.1073\/pnas.222375399","article-title":"Biological activity of the Helicobacter pylori virulence factor CagA is determined by variation in the tyrosine phosphorylation sites","volume":"99","author":"Higashi","year":"2002","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_158","doi-asserted-by":"crossref","first-page":"e6620","DOI":"10.1097\/MD.0000000000006620","article-title":"Association of CagA EPIYA-D or EPIYA-C phosphorylation sites with peptic ulcer and gastric cancer risks: A meta-analysis","volume":"96","author":"Li","year":"2017","journal-title":"Medicine"},{"key":"ref_159","doi-asserted-by":"crossref","unstructured":"Zhang, X.S., Tegtmeyer, N., Traube, L., Jindal, S., Perez-Perez, G., Sticht, H., Backert, S., and Blaser, M.J. (2015). A specific A\/T polymorphism in Western tyrosine phosphorylation B-motifs regulates Helicobacter pylori CagA epithelial cell interactions. PLoS Pathog., 11.","DOI":"10.1371\/journal.ppat.1004621"},{"key":"ref_160","doi-asserted-by":"crossref","first-page":"2169","DOI":"10.1158\/1541-7786.MCR-19-0181","article-title":"CDX1 expression induced by CagA-expressing Helicobacter pylori promotes gastric tumorigenesis","volume":"17","author":"Choi","year":"2019","journal-title":"Mol. Cancer Res."},{"key":"ref_161","doi-asserted-by":"crossref","unstructured":"Grainger, S., Hryniuk, A., and Lohnes, D. (2013). Cdx1 and Cdx2 exhibit transcriptional specificity in the intestine. PLoS ONE, 8.","DOI":"10.1371\/journal.pone.0054757"},{"key":"ref_162","doi-asserted-by":"crossref","first-page":"19652","DOI":"10.1073\/pnas.1903798116","article-title":"Targeted mobilization of Lrig1+ gastric epithelial stem cell populations by a carcinogenic Helicobacter pylori type IV secretion system","volume":"116","author":"Wroblewski","year":"2019","journal-title":"Proc. Nat. Acad. Sci. USA"},{"key":"ref_163","doi-asserted-by":"crossref","first-page":"764","DOI":"10.1016\/j.chom.2012.10.014","article-title":"Reactive oxygen species-induced autophagic degradation of Helicobacter pylori CagA is specifically suppressed in cancer stem-like cells","volume":"12","author":"Tsugawa","year":"2012","journal-title":"Cell Host Microbe"},{"key":"ref_164","doi-asserted-by":"crossref","first-page":"370","DOI":"10.4161\/auto.5.3.7663","article-title":"Efect of Helicobacter pylori\u2019s vacuolating cytotoxin on the autophagy pathway in gastric epithelial cells","volume":"5","author":"Terebiznik","year":"2009","journal-title":"Autophagy"},{"key":"ref_165","doi-asserted-by":"crossref","unstructured":"Ricci, V. (2016). Relationship between VacA toxin and host cell autophagy in Helicobacter pylori infection of the human stomach: A few answers, many questions. Toxins, 8.","DOI":"10.3390\/toxins8070203"},{"key":"ref_166","doi-asserted-by":"crossref","first-page":"17771","DOI":"10.1074\/jbc.270.30.17771","article-title":"Mosaicism in vacuolating cytotoxin alleles of Helicobacter pylori association of specific VacA types with cytotoxin production and peptic ulceration","volume":"270","author":"Atherton","year":"1995","journal-title":"J. Biol. Chem."},{"key":"ref_167","doi-asserted-by":"crossref","first-page":"926","DOI":"10.1053\/j.gastro.2007.06.056","article-title":"A new Helicobacter pylori vacuolating cytotoxin determinant, the intermediate region, is associated with gastric cancer","volume":"133","author":"Rhead","year":"2007","journal-title":"Gastroenterology"},{"key":"ref_168","doi-asserted-by":"crossref","first-page":"101","DOI":"10.4161\/gmic.23797","article-title":"Polymorphism in the Helicobacter pylori CagA and VacA toxins and disease","volume":"4","author":"Bridge","year":"2003","journal-title":"Gut Microbes"},{"key":"ref_169","doi-asserted-by":"crossref","first-page":"690","DOI":"10.1128\/JCM.01815-09","article-title":"Diversity of VacA intermediate region among Helicobacter pylori strains from several regions of the world","volume":"48","author":"Chung","year":"2010","journal-title":"J. Clin. Microbiol."},{"key":"ref_170","doi-asserted-by":"crossref","first-page":"6499","DOI":"10.1128\/JB.183.22.6499-6508.2001","article-title":"A 12 amino acid segment, present in type s2 but not type s1 Helicobacter pylori VacA proteins, abolishes cytotoxin activity and alters membrane channel formation","volume":"183","author":"McClain","year":"2001","journal-title":"J. Bacteriol."},{"key":"ref_171","doi-asserted-by":"crossref","unstructured":"Trang, T.T.H., Binh, T.T., and Yamaoka, Y. (2016). Relationship between vacA Types and Development of Gastroduodenal Diseases. Toxins, 8.","DOI":"10.3390\/toxins8060182"},{"key":"ref_172","doi-asserted-by":"crossref","first-page":"211","DOI":"10.1007\/s002849900447","article-title":"Vacuolating cytotoxin (vacA) alleles of Helicobacter pylori comprise two geographically widespread types, m1 and m2, and have evolved through limited recombination","volume":"39","author":"Atherton","year":"1999","journal-title":"Curr. Microbiol."},{"key":"ref_173","doi-asserted-by":"crossref","first-page":"e3088","DOI":"10.1128\/IAI.66.7.3088-3094.1998","article-title":"Heterogeneity in levels of vacuolating cytotoxin gene (vacA) transcription among Helicobacter pylori strains","volume":"66","author":"Forsyth","year":"1998","journal-title":"Infect. Immun."},{"key":"ref_174","doi-asserted-by":"crossref","first-page":"1431","DOI":"10.1097\/MEG.0b013e328364b53e","article-title":"Helicobacter pylori CagA and VacA genotypes and gastric phenotype: A meta-analysis","volume":"25","author":"Matos","year":"2013","journal-title":"Eur. J. Gastroenterol. Hepatol."},{"key":"ref_175","doi-asserted-by":"crossref","first-page":"409","DOI":"10.1002\/2211-5463.12046","article-title":"A systematic review on the association between the Helicobacter pylori vacA i genotype and gastric disease","volume":"6","author":"Liu","year":"2016","journal-title":"FEBS Open Bio"},{"key":"ref_176","doi-asserted-by":"crossref","first-page":"1239","DOI":"10.1111\/j.1742-1241.2005.00778.x","article-title":"Frequencies of serum antibodies to Helicobacter pylori CagA and VacA in a Turkish population with various gastroduodenal diseases","volume":"60","author":"Sezikli","year":"2006","journal-title":"Int. J. Clin. Pract."},{"key":"ref_177","doi-asserted-by":"crossref","first-page":"220","DOI":"10.1016\/j.micpath.2016.08.030","article-title":"Serum VacA antibody is associated with risks of peptic ulcer and gastric cancer: A meta-analysis","volume":"99","author":"Li","year":"2016","journal-title":"Microb. Pathog."},{"key":"ref_178","doi-asserted-by":"crossref","first-page":"175","DOI":"10.1053\/j.gastro.2018.09.054","article-title":"Serologic response to Helicobacter pylori proteins associated with risk of colorectal cancer among diverse populations in the United States","volume":"156","author":"Butt","year":"2019","journal-title":"Gastroenterology"},{"key":"ref_179","doi-asserted-by":"crossref","first-page":"3754","DOI":"10.1128\/IAI.68.6.3754-3757.2000","article-title":"Cell specificity of Helicobacter pylori cytotoxin is determined by a short region in the polymorphic midregion","volume":"68","author":"Ji","year":"2000","journal-title":"Infect. Immun."},{"key":"ref_180","doi-asserted-by":"crossref","first-page":"11887","DOI":"10.1021\/bi010065u","article-title":"Two distinctive cell binding patterns by vacuolating toxin fused with glutathione S transferase: One high affinity m1 specific binding and the other lower affinity binding for variant m forms","volume":"40","author":"Wang","year":"2001","journal-title":"Biochemistry"},{"key":"ref_181","doi-asserted-by":"crossref","first-page":"6599","DOI":"10.1128\/IAI.01085-06","article-title":"Helicobacter pylori VacA Toxin Promotes Bacterial Intracellular Survival in Gastric Epithelial Cells","volume":"74","author":"Terebiznik","year":"2006","journal-title":"Infect. Immun."},{"key":"ref_182","doi-asserted-by":"crossref","first-page":"1411","DOI":"10.1038\/s41564-019-0441-6","article-title":"VacA generates a protective intracellular reservoir for Helicobacter pylori that is eliminated by activation of the lysosomal calcium channel TRPML1","volume":"4","author":"Capurro","year":"2019","journal-title":"Nat. Microbiol."},{"key":"ref_183","doi-asserted-by":"crossref","first-page":"443","DOI":"10.1016\/S1097-2765(02)00658-5","article-title":"The HtrA family of proteases: Implications for protein composition and cell fate","volume":"10","author":"Clausen","year":"2002","journal-title":"Mol. Cell"},{"key":"ref_184","doi-asserted-by":"crossref","first-page":"152","DOI":"10.1038\/nrm3065","article-title":"HTRA proteases: Regulated proteolysis in protein quality control","volume":"12","author":"Clausen","year":"2011","journal-title":"Nat. Rev. Mol. Cell Biol."},{"key":"ref_185","doi-asserted-by":"crossref","first-page":"30010","DOI":"10.1074\/jbc.M804650200","article-title":"Evolution of peptidase diversity","volume":"283","author":"Page","year":"2008","journal-title":"J. Biol. Chem."},{"key":"ref_186","doi-asserted-by":"crossref","first-page":"149","DOI":"10.1146\/annurev-micro-090110-102925","article-title":"Protein quality control in the bacterial periplasm","volume":"65","author":"Merdanovic","year":"2011","journal-title":"Annu. Rev. Microbiol."},{"key":"ref_187","doi-asserted-by":"crossref","first-page":"815","DOI":"10.1101\/gad.11.7.815","article-title":"Protein quality control: Triage by chaperone and proteases","volume":"11","author":"Gottesman","year":"1997","journal-title":"Genes Dev."},{"key":"ref_188","doi-asserted-by":"crossref","first-page":"704","DOI":"10.1016\/j.resmic.2009.08.017","article-title":"Proteases in bacterial pathogenesis","volume":"160","author":"Ingmer","year":"2009","journal-title":"Res. Microbiol."},{"key":"ref_189","doi-asserted-by":"crossref","first-page":"161","DOI":"10.1007\/978-94-007-5940-4_7","article-title":"Bacterial proteases and virulence","volume":"66","author":"Frees","year":"2013","journal-title":"Subcell. Biochem."},{"key":"ref_190","first-page":"2174","article-title":"The extracellular bacterial HtrA proteins as potential therapeutic targets and vaccine candidates","volume":"24","author":"Figaj","year":"2017","journal-title":"Curr. Med. Chem."},{"key":"ref_191","doi-asserted-by":"crossref","first-page":"961","DOI":"10.3389\/fmicb.2019.00961","article-title":"Properties of the HtrA Protease from Bacterium Helicobacter pylori Whose Activity Is Indispensable for Growth Under Stress Conditions","volume":"10","author":"Zarzecka","year":"2019","journal-title":"Front. Microbiol."},{"key":"ref_192","doi-asserted-by":"crossref","first-page":"402","DOI":"10.1002\/jobm.201200074","article-title":"The stability and activity of recombinant Helicobacter pylori HtrA under stress conditions","volume":"53","author":"Hoy","year":"2013","journal-title":"J. Basic Microbiol."},{"key":"ref_193","doi-asserted-by":"crossref","first-page":"40","DOI":"10.1186\/s13099-017-0189-6","article-title":"Overexpression of serine protease HtrA enhances disruption of adherens junctions, paracellular transmigration and type IV secretion of CagA by Helicobacter pylori","volume":"9","author":"Harrer","year":"2017","journal-title":"Gut Pathog."},{"key":"ref_194","doi-asserted-by":"crossref","first-page":"676","DOI":"10.1124\/pr.112.005983","article-title":"Classification, functions, and clinical relevance of extracellular vesicles","volume":"64","author":"Boing","year":"2012","journal-title":"Pharmacol. Rev."},{"key":"ref_195","doi-asserted-by":"crossref","first-page":"97","DOI":"10.1016\/j.semcdb.2015.02.006","article-title":"Gram-negative and Gram-positive bacterial extracellular vesicles","volume":"40","author":"Kim","year":"2015","journal-title":"Semin. Cell Dev. Biol."},{"key":"ref_196","doi-asserted-by":"crossref","first-page":"1395","DOI":"10.1111\/j.1365-2958.2009.06731.x","article-title":"Biogenesis of bacterial outer membrane vesicles","volume":"72","author":"Deatherage","year":"2009","journal-title":"Mol. Microbiol."},{"key":"ref_197","doi-asserted-by":"crossref","first-page":"1539","DOI":"10.1111\/j.1365-2958.2010.07307.x","article-title":"Biochemical and functional characterization of Helicobacter pylori vesicles","volume":"77","author":"Olofsson","year":"2010","journal-title":"Mol. Microbiol."},{"key":"ref_198","doi-asserted-by":"crossref","first-page":"395","DOI":"10.1007\/s00281-010-0231-y","article-title":"Bacterial outer membrane vesicles in disease and preventive medicine","volume":"33","author":"Unal","year":"2011","journal-title":"Semin. Immunopathol."},{"key":"ref_199","doi-asserted-by":"crossref","first-page":"220","DOI":"10.1002\/(SICI)1096-9896(199906)188:2<220::AID-PATH307>3.0.CO;2-C","article-title":"Release of Helicobacter pylori vacuolating cytotoxin by both a specific secretion pathway and budding of outer membrane vesicles. Uptake of released toxin and vesicles by gastric epithelium","volume":"188","author":"Fiocca","year":"1999","journal-title":"J. Pathol."},{"key":"ref_200","doi-asserted-by":"crossref","first-page":"259","DOI":"10.1111\/j.1574-6968.2000.tb08905.x","article-title":"A role for the bacterial outer membrane in the pathogenesis of Helicobacter pylori infection","volume":"182","author":"Keenan","year":"2000","journal-title":"FEMS Microbiol. Lett."},{"key":"ref_201","doi-asserted-by":"crossref","first-page":"785","DOI":"10.1002\/prca.200800192","article-title":"Proteomic and functional characterization of the outer membrane vesicles from the gastric pathogen Helicobacter pylori","volume":"3","author":"Mullaney","year":"2009","journal-title":"Proteom. Clin. Appl."},{"key":"ref_202","doi-asserted-by":"crossref","first-page":"e00979-14","DOI":"10.1128\/mBio.00979-14","article-title":"Uptake of Helicobacter pylori Vesicles Is Facilitated by Clathrin Dependent and Clathrin-Independent Endocytic Pathways","volume":"5","author":"Olofsson","year":"2014","journal-title":"mBio"},{"key":"ref_203","doi-asserted-by":"crossref","first-page":"9","DOI":"10.1016\/j.micinf.2011.08.007","article-title":"Composition and function of Helicobacter pylori outer membrane vesicles","volume":"14","author":"Parker","year":"2012","journal-title":"Microbes Infect."},{"key":"ref_204","doi-asserted-by":"crossref","first-page":"81","DOI":"10.1128\/MMBR.00031-09","article-title":"Virulence and immunomodulatory roles of bacterial outer membrane vesicles","volume":"74","author":"Ellis","year":"2010","journal-title":"Microbiol. Mol. Biol. Rev."},{"key":"ref_205","doi-asserted-by":"crossref","first-page":"1372","DOI":"10.1128\/IAI.01443-13","article-title":"Helicobacter pylori membrane vesicles stimulate innate pro- and anti-inflammatory responses and induce apoptosis in Jurkat T cells","volume":"82","author":"Winter","year":"2014","journal-title":"Infect. Immun."},{"key":"ref_206","doi-asserted-by":"crossref","first-page":"e330","DOI":"10.1038\/emm.2017.47","article-title":"Helicobacter pylori-derived extracellular vesicles increased in the gastric juices of gastric adenocarcinoma patients and induced inflammation mainly via specific targeting of gastrin epithelial cells","volume":"49","author":"Choi","year":"2017","journal-title":"Exp. Mol. Med."},{"key":"ref_207","doi-asserted-by":"crossref","first-page":"2400","DOI":"10.1093\/carcin\/bgn218","article-title":"Outer membrane vesicles enhance the carcinogenic potential of Helicobacter pylori","volume":"29","author":"Chitcholtan","year":"2008","journal-title":"Carcinogenesis"},{"key":"ref_208","doi-asserted-by":"crossref","first-page":"333","DOI":"10.1016\/S0009-2797(97)00170-1","article-title":"gamma-Glutamyl transpeptidase, a glutathionase: Its expression and function in carcinogenesis","volume":"111","author":"Hanigan","year":"1998","journal-title":"Chem. Biol. Interact."},{"key":"ref_209","doi-asserted-by":"crossref","first-page":"1359","DOI":"10.1046\/j.1365-2958.1999.01271.x","article-title":"Essential role of Helicobacter pylori g-glutamyltranspeptidase for the colonization of the gastric mucosa of mice","volume":"31","author":"Chevalier","year":"1999","journal-title":"Mol. Microbiol."},{"key":"ref_210","doi-asserted-by":"crossref","first-page":"4168","DOI":"10.1128\/IAI.69.6.4168-4173.2001","article-title":"g-glutamyltransferase is a Helicobacter pylori virulence factor but is not essential for colonization","volume":"69","author":"Mcgovern","year":"2001","journal-title":"Infect. Immun."},{"key":"ref_211","doi-asserted-by":"crossref","first-page":"13636","DOI":"10.1038\/s41598-017-14028-1","article-title":"Helicobacter pylori \u03b3-glutamyl transferase contributes to colonization and diferential recruitment of T cells during persistence","volume":"7","author":"Anderl","year":"2017","journal-title":"Sci. Rep."},{"key":"ref_212","doi-asserted-by":"crossref","unstructured":"Bravo, J., D\u00edaz, P., Corval\u00e1n, A.H., and Quest, A.F.G. (2019). A Novel Role for Helicobacter pylori Gamma-Glutamyltranspeptidase in Regulating Autophagy and Bacterial Internalization in Human Gastric Cells. Cancers, 11.","DOI":"10.3390\/cancers11060801"},{"key":"ref_213","doi-asserted-by":"crossref","first-page":"372","DOI":"10.1007\/s12275-010-9293-8","article-title":"Helicobacter pylori \u03b3-glutamyltranspeptidase induces cell cycle arrest at the G1-S phase transition","volume":"48","author":"Kim","year":"2010","journal-title":"J. Microbiol."},{"key":"ref_214","doi-asserted-by":"crossref","first-page":"564","DOI":"10.1053\/j.gastro.2010.03.050","article-title":"Helicobacter pylori \u03b3-glutamyl transpeptidase is a pathogenic factor in the development of peptic ulcer disease","volume":"139","author":"Gong","year":"2010","journal-title":"Gastroenterology"},{"key":"ref_215","doi-asserted-by":"crossref","first-page":"665","DOI":"10.1111\/1348-0421.12089","article-title":"Role of \u03b3-glutamyltranspeptidase in the pathogenesis of Helicobacter pylori infection","volume":"57","author":"Rimbara","year":"2013","journal-title":"Microbiol. Immunol."},{"key":"ref_216","doi-asserted-by":"crossref","first-page":"e13006","DOI":"10.1111\/cmi.13006","article-title":"The lipoprotein HP1454 of Helicobacter pylori regulates T-cell response by shaping T-cell receptor signalling","volume":"21","author":"Capitani","year":"2019","journal-title":"Cell. Microbiol."},{"key":"ref_217","doi-asserted-by":"crossref","first-page":"6073","DOI":"10.1038\/s41598-018-24406-y","article-title":"Distribution and clinical associations of integrating conjugative elements and cag pathogenicity islands of Helicobacter pylori in Indonesia","volume":"8","author":"Waskito","year":"2018","journal-title":"Sci. Rep."},{"key":"ref_218","doi-asserted-by":"crossref","first-page":"1391","DOI":"10.1053\/j.gastro.2017.12.008","article-title":"Helicobacter pylori depletes cholesterol in gastric glands to prevent interferon gamma signaling and escape the inflammatory response","volume":"154","author":"Morey","year":"2018","journal-title":"Gastroenterology"},{"key":"ref_219","doi-asserted-by":"crossref","first-page":"94","DOI":"10.1089\/fpd.2018.2480","article-title":"Genotypes and antimicrobial susceptibility profiles of hemolytic Escherichia coli from diarrheic piglets","volume":"16","author":"Jiang","year":"2019","journal-title":"Foodborne Pathog. Dis."},{"key":"ref_220","doi-asserted-by":"crossref","first-page":"2","DOI":"10.1016\/j.cmi.2016.11.012","article-title":"The role of whole genome sequencing in antimicrobial susceptibility testing of bacteria: Report from the EUCAST Subcommittee","volume":"23","author":"Ellington","year":"2017","journal-title":"Clin. Microbiol. Infect."},{"key":"ref_221","doi-asserted-by":"crossref","first-page":"49","DOI":"10.1016\/S2213-2600(15)00466-X","article-title":"Rapid, comprehensive, and affordable mycobacterial diagnosis with whole-genome sequencing: A prospective study","volume":"4","author":"Pankhurst","year":"2016","journal-title":"Lancet Respir. Med."},{"key":"ref_222","doi-asserted-by":"crossref","unstructured":"Witney, A.A., Cosgrove, C.A., Arnold, A., Hinds, J., Stoker, N.G., and Butcher, P.D. (2016). Clinical use of whole genome sequencing for Mycobacterium tuberculosis. BMC Med., 14.","DOI":"10.1186\/s12916-016-0598-2"},{"key":"ref_223","doi-asserted-by":"crossref","first-page":"e00252-19","DOI":"10.1128\/mSphere.00252-19","article-title":"Whole-genome sequencing-based characterization of 100 Listeria monocytogenes isolates collected from food processing environments over a four-year period","volume":"4","author":"Hurley","year":"2019","journal-title":"mSphere"},{"key":"ref_224","doi-asserted-by":"crossref","first-page":"1464","DOI":"10.3389\/fmicb.2019.01464","article-title":"Whole Genome Sequencing-Based Comparison of Food Isolates of Cronobacter sakazakii","volume":"10","author":"Aly","year":"2019","journal-title":"Front. Microbiol."},{"key":"ref_225","doi-asserted-by":"crossref","first-page":"1193","DOI":"10.1016\/S1473-3099(15)00062-6","article-title":"Whole-genome sequencing for prediction of Mycobacterium tuberculosis drug susceptibility and resistance: A retrospective cohort study","volume":"15","author":"Walker","year":"2015","journal-title":"Lancet Infect. Dis."},{"key":"ref_226","doi-asserted-by":"crossref","first-page":"31","DOI":"10.1038\/nrg2626","article-title":"Sequencing technologies-the next generation","volume":"11","author":"Metzker","year":"2010","journal-title":"Nat. Rev. Genet."},{"key":"ref_227","doi-asserted-by":"crossref","first-page":"387","DOI":"10.1146\/annurev.genom.9.081307.164359","article-title":"Next-generation DNA sequencing methods","volume":"9","author":"Mardis","year":"2008","journal-title":"Annu. Rev. Genom. Hum. Genet."},{"key":"ref_228","doi-asserted-by":"crossref","unstructured":"Edwards, M.S., McLaughlin, R.W., Li, J., Wan, X.-L., Liu, Y., Xie, H.-X., Hao, Y.-J., and Zheng, J.-S. (2019). Putative virulence factors of Plesiomonas shigelloides. Antonie Van Leeuwenhoek, 1\u201312.","DOI":"10.1007\/s10482-019-01303-6"},{"key":"ref_229","doi-asserted-by":"crossref","unstructured":"Lauener, F.N., Imkamp, F., Lehours, P., Buissonni\u00e8re, A., Benejat, L., Zbinden, R., Keller, P.M., and Wagner, K. (2019). Genetic Determinants and Prediction of Antibiotic Resistance Phenotypes in Helicobacter pylori. J. Clin. Med., 8.","DOI":"10.3390\/jcm8010053"},{"key":"ref_230","doi-asserted-by":"crossref","unstructured":"Tuan, V.P., Narith, D., Tshibangu-Kabamba, E., Dung, H.D.Q., Viet, P.T., Sokomoth, S., Binh, T.T., Sokhem, S., Tri, T.D., and Ngov, S. (2019). A Next-Generation Sequencing-Based Approach to Identify Genetic Determinants of Antibiotic Resistance in Cambodian Helicobacter pylori Clinical Isolates. J. Clin. Med., 8.","DOI":"10.3390\/jcm8060858"},{"key":"ref_231","doi-asserted-by":"crossref","first-page":"10","DOI":"10.1186\/s12941-018-0259-8","article-title":"Application of next-generation sequencing to characterize novel mutations in clarithromycin-susceptible Helicobacter pylori strains with A2143G of 23S rRNA gene","volume":"17","author":"Chen","year":"2018","journal-title":"Ann. Clin. Microbiol. Antimicrob."},{"key":"ref_232","doi-asserted-by":"crossref","unstructured":"Imkamp, F., Lauener, F.N., Pohl, D., Lehours, P., Vale, F.F., Jehanne, Q., Zbinden, R., Keller, P.M., and Wagner, K. (2019). Rapid Characterization of Virulence Determinants in Helicobacter pylori Isolated from Non-Atrophic Gastritis Patients by Next-Generation Sequencing. J. Clin. Med., 8.","DOI":"10.3390\/jcm8071030"},{"key":"ref_233","doi-asserted-by":"crossref","first-page":"699","DOI":"10.1016\/S1473-3099(06)70627-2","article-title":"Helicobacter pylori and antimicrobial resistance: Molecular mechanisms and clinical implications","volume":"6","author":"Gerrits","year":"2006","journal-title":"Lancet Infect. Dis."},{"key":"ref_234","doi-asserted-by":"crossref","first-page":"21","DOI":"10.1186\/s13099-017-0165-1","article-title":"Genetic variants of Helicobacter pylori type IV secretion system components CagL and CagI and their association with clinical outcomes","volume":"9","author":"Ogawa","year":"2017","journal-title":"Gut Pathog."},{"key":"ref_235","doi-asserted-by":"crossref","first-page":"661","DOI":"10.1038\/nature05911","article-title":"Genome-wide association study of 14,000 cases of seven common diseases and 3,000 shared controls","volume":"447","author":"Burton","year":"2007","journal-title":"Nature"},{"key":"ref_236","doi-asserted-by":"crossref","unstructured":"Berthenet, E., Yahara, K., Thorell, K., Pascoe, B., Meric, G., Mikhail, J.M., Engstrand, L., Enroth, H., Burette, A., and Megraud, F. (2018). A GWAS on Helicobacter pylori strains points to genetic variants associated with gastric cancer risk. BMC Biol., 16.","DOI":"10.1186\/s12915-018-0550-3"},{"key":"ref_237","doi-asserted-by":"crossref","first-page":"574","DOI":"10.1038\/ismej.2010.149","article-title":"Structure of the human gastric bacterial community in relation to Helicobacter pylori status","volume":"5","author":"Goldfarb","year":"2011","journal-title":"ISME J."},{"key":"ref_238","doi-asserted-by":"crossref","first-page":"364","DOI":"10.1111\/hel.12293","article-title":"Analysis of gastric microbiota by pyrosequencing: Minor role of bacteria other than Helicobacter pylori in the gastric carcinogenesis","volume":"21","author":"Jo","year":"2016","journal-title":"Helicobacter"},{"key":"ref_239","doi-asserted-by":"crossref","first-page":"216","DOI":"10.1136\/gutjnl-2016-312904","article-title":"The active bacterial assemblages of the upper GI tract in individuals with and without Helicobacter infection","volume":"67","author":"Schulz","year":"2018","journal-title":"Gut"},{"key":"ref_240","doi-asserted-by":"crossref","first-page":"80","DOI":"10.1126\/science.aaa4972","article-title":"Cancer and the microbiota","volume":"348","author":"Garrett","year":"2015","journal-title":"Science"},{"key":"ref_241","doi-asserted-by":"crossref","first-page":"341","DOI":"10.1038\/nri.2016.42","article-title":"Gut microbiota, metabolites and host immunity","volume":"16","author":"Rooks","year":"2016","journal-title":"Nat. Rev. Immunol."},{"key":"ref_242","doi-asserted-by":"crossref","first-page":"407","DOI":"10.1111\/hel.12145","article-title":"Differences in gastric mucosal microbiota profiling in patients with chronic gastritis, intestinal metaplasia, and gastric cancer using pyrosequencing methods","volume":"19","author":"Eun","year":"2014","journal-title":"Helicobacter"},{"key":"ref_243","doi-asserted-by":"crossref","first-page":"1024","DOI":"10.1136\/gutjnl-2017-314281","article-title":"Mucosal microbiome dysbiosis in gastric carcinogenesis","volume":"67","author":"Coker","year":"2018","journal-title":"Gut"},{"key":"ref_244","doi-asserted-by":"crossref","first-page":"226","DOI":"10.1136\/gutjnl-2017-314205","article-title":"Gastric microbial community profiling reveals a dysbiotic cancer-associated microbiota","volume":"67","author":"Ferreira","year":"2018","journal-title":"Gut"},{"key":"ref_245","doi-asserted-by":"crossref","first-page":"339","DOI":"10.1053\/gast.2000.9367","article-title":"Omeprazole, Helicobacter pylori status, and alterations in the intragastric milieu facilitating bacterial N-nitrosation","volume":"119","author":"Mowat","year":"2000","journal-title":"Gastroenterology"},{"key":"ref_246","doi-asserted-by":"crossref","first-page":"3","DOI":"10.1111\/j.1365-2036.2006.02707.x","article-title":"Review article: Proton pump inhibitors and bacterial overgrowth","volume":"23","author":"Williams","year":"2005","journal-title":"Aliment. Pharmacol. Ther."},{"key":"ref_247","doi-asserted-by":"crossref","first-page":"261","DOI":"10.1097\/MEG.0000000000000542","article-title":"Bacterial overgrowth and diversification of microbiota in gastric cancer","volume":"28","author":"Wang","year":"2016","journal-title":"Eur. J. Gastroenterol. Hepatol."},{"key":"ref_248","doi-asserted-by":"crossref","first-page":"18594","DOI":"10.1038\/srep18594","article-title":"Different gastric microbiota compositions in two human populations with high and low gastric cancer risk in Colombia","volume":"6","author":"Yang","year":"2016","journal-title":"Sci. Rep."},{"key":"ref_249","doi-asserted-by":"crossref","first-page":"202","DOI":"10.3389\/fcimb.2018.00202","article-title":"Association between gut microbiota and Helicobacter pylori-related gastric lesions in a high-risk population of gastric cancer","volume":"8","author":"Gao","year":"2018","journal-title":"Front. Cell. Infect. Microbiol."}],"container-title":["Toxins"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.mdpi.com\/2072-6651\/11\/11\/677\/pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2025,10,11]],"date-time":"2025-10-11T13:35:45Z","timestamp":1760189745000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.mdpi.com\/2072-6651\/11\/11\/677"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2019,11,19]]},"references-count":249,"journal-issue":{"issue":"11","published-online":{"date-parts":[[2019,11]]}},"alternative-id":["toxins11110677"],"URL":"https:\/\/doi.org\/10.3390\/toxins11110677","relation":{},"ISSN":["2072-6651"],"issn-type":[{"value":"2072-6651","type":"electronic"}],"subject":[],"published":{"date-parts":[[2019,11,19]]}}}