{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,7,3]],"date-time":"2026-07-03T04:21:42Z","timestamp":1783052502375,"version":"3.54.6"},"reference-count":61,"publisher":"Springer Science and Business Media LLC","issue":"1","license":[{"start":{"date-parts":[[2026,5,11]],"date-time":"2026-05-11T00:00:00Z","timestamp":1778457600000},"content-version":"tdm","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by-nc-nd\/4.0"},{"start":{"date-parts":[[2026,7,3]],"date-time":"2026-07-03T00:00:00Z","timestamp":1783036800000},"content-version":"vor","delay-in-days":53,"URL":"https:\/\/creativecommons.org\/licenses\/by-nc-nd\/4.0"}],"funder":[{"name":"Basic Science Research Program through the NRF of Korea funded by the Ministry of Education","award":["RS-2022-NR075350"],"award-info":[{"award-number":["RS-2022-NR075350"]}]},{"name":"Bio & Medical Technology Development Program of the National Research Foundation (NRF) funded by the Korean government","award":["RS-2023-00223501"],"award-info":[{"award-number":["RS-2023-00223501"]}]},{"name":"a National Research Foundation (NRF) grant funded by the Korean government","award":["NRF-2021R1C1C1010631"],"award-info":[{"award-number":["NRF-2021R1C1C1010631"]}]}],"content-domain":{"domain":["link.springer.com"],"crossmark-restriction":false},"short-container-title":["BMC Microbiol"],"DOI":"10.1186\/s12866-026-05130-4","type":"journal-article","created":{"date-parts":[[2026,5,11]],"date-time":"2026-05-11T02:13:38Z","timestamp":1778465618000},"update-policy":"https:\/\/doi.org\/10.1007\/springer_crossmark_policy","source":"Crossref","is-referenced-by-count":0,"title":["Genomic characterization of clinical Helicobacter pylori strains from Korea: insights into phylogeny, pan-genome structure, and virulence factors"],"prefix":"10.1186","volume":"26","author":[{"given":"Hi Eun","family":"Jung","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Chang Seol","family":"Bang","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Ji Yong","family":"Ahn","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Hwoon-Yong","family":"Jung","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Sun Mi","family":"Lee","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"So Won","family":"Kim","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Dong Woo","family":"Chae","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Eun Jeong","family":"Gong","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"297","published-online":{"date-parts":[[2026,5,11]]},"reference":[{"key":"5130_CR1","doi-asserted-by":"publisher","first-page":"1311","DOI":"10.1016\/S0140-6736(84)91816-6","volume":"1","author":"BJ Marshall","year":"1984","unstructured":"Marshall BJ, Warren JR. Unidentified curved bacilli in the stomach of patients with gastritis and peptic ulceration. Lancet. 1984;1:1311\u20135.","journal-title":"Lancet"},{"key":"5130_CR2","doi-asserted-by":"publisher","first-page":"133","DOI":"10.1016\/j.ijmm.2004.06.014","volume":"294","author":"S Suerbaum","year":"2004","unstructured":"Suerbaum S, Achtman M. Helicobacter pylori: recombination, population structure and human migrations. Int J Med Microbiol. 2004;294:133\u20139.","journal-title":"Int J Med Microbiol"},{"key":"5130_CR3","doi-asserted-by":"publisher","first-page":"1175","DOI":"10.1056\/NEJMra020542","volume":"347","author":"S Suerbaum","year":"2002","unstructured":"Suerbaum S, Michetti P. Helicobacter pylori infection. N Engl J Med. 2002;347:1175\u201386.","journal-title":"N Engl J Med"},{"key":"5130_CR4","doi-asserted-by":"publisher","first-page":"261","DOI":"10.1053\/j.gastro.2024.12.033","volume":"169","author":"AC Ford","year":"2025","unstructured":"Ford AC, Yuan Y, Park JY, Forman D, Moayyedi P. Eradication therapy to prevent gastric cancer in Helicobacterpylori-positive individuals: Systematic review and meta-analysis of randomized controlled trials and observational studies. Gastroenterology. 2025;169:261\u201376.","journal-title":"Gastroenterology"},{"key":"5130_CR5","doi-asserted-by":"publisher","first-page":"605","DOI":"10.1053\/j.gastro.2023.12.022","volume":"166","author":"YC Chen","year":"2024","unstructured":"Chen YC, Malfertheiner P, Yu HT, Kuo CL, Chang YY, Meng FT, et al. Global prevalence of Helicobacter pylori infection and incidence of gastric cancer between 1980 and 2022. Gastroenterology. 2024;166:605\u201319.","journal-title":"Gastroenterology"},{"key":"5130_CR6","first-page":"229","volume":"74","author":"F Bray","year":"2024","unstructured":"Bray F, Laversanne M, Sung H, Ferlay J, Siegel RL, Soerjomataram I, et al. Global cancer statistics 2022: GLOBOCAN estimates of incidence and mortality worldwide for 36 cancers in 185 countries. CA Cancer J Clin. 2024;74:229\u201363.","journal-title":"CA Cancer J Clin"},{"key":"5130_CR7","doi-asserted-by":"publisher","first-page":"160","DOI":"10.5230\/jgc.2022.22.e21","volume":"22","author":"SH Park","year":"2022","unstructured":"Park SH, Kang MJ, Yun EH, Jung KW. Epidemiology of gastric cancer in Korea: Trends in incidence and survival based on Korea central cancer registry data (1999\u20132019). J Gastric Cancer. 2022;22:160\u20138.","journal-title":"J Gastric Cancer"},{"key":"5130_CR8","doi-asserted-by":"publisher","first-page":"629","DOI":"10.1038\/nrgastro.2010.154","volume":"7","author":"Y Yamaoka","year":"2010","unstructured":"Yamaoka Y. Mechanisms of disease: Helicobacter pylori virulence factors. Nat reviews Gastroenterol Hepatol. 2010;7:629\u201341.","journal-title":"Nat reviews Gastroenterol Hepatol"},{"key":"5130_CR9","doi-asserted-by":"publisher","first-page":"14648","DOI":"10.1073\/pnas.93.25.14648","volume":"93","author":"S Censini","year":"1996","unstructured":"Censini S, Lange C, Xiang Z, Crabtree JE, Ghiara P, Borodovsky M, et al. cag, a pathogenicity island of Helicobacter pylori, encodes type I-specific and disease-associated virulence factors. Proc Natl Acad Sci USA. 1996;93:14648\u201353.","journal-title":"Proc Natl Acad Sci USA"},{"key":"5130_CR10","doi-asserted-by":"publisher","first-page":"50","DOI":"10.1038\/s41423-019-0339-5","volume":"17","author":"A Takahashi-Kanemitsu","year":"2020","unstructured":"Takahashi-Kanemitsu A, Knight CT, Hatakeyama M. Molecular anatomy and pathogenic actions of Helicobacter pylori CagA that underpin gastric carcinogenesis. Cell Mol Immunol. 2020;17:50\u201363.","journal-title":"Cell Mol Immunol"},{"key":"5130_CR11","doi-asserted-by":"publisher","first-page":"104135","DOI":"10.1016\/j.meegid.2019.104135","volume":"78","author":"A Hanafiah","year":"2020","unstructured":"Hanafiah A, Lopes BS. Genetic diversity and virulence characteristics of Helicobacter pylori isolates in different human ethnic groups. Infect Genet evolution: J Mol Epidemiol evolutionary Genet Infect Dis. 2020;78:104135.","journal-title":"Infect Genet evolution: J Mol Epidemiol evolutionary Genet Infect Dis"},{"key":"5130_CR12","first-page":"683","volume-title":"SHP-2 tyrosine phosphatase as an intracellular target of Helicobacter pylori CagA protein","author":"H Higashi","year":"2002","unstructured":"Higashi H, Tsutsumi R, Muto S, Sugiyama T, Azuma T, Asaka M, et al. SHP-2 tyrosine phosphatase as an intracellular target of Helicobacter pylori CagA protein. Volume 295. New York, NY: Science; 2002. pp. 683\u20136."},{"key":"5130_CR13","doi-asserted-by":"publisher","first-page":"959","DOI":"10.1128\/JCM.02330-08","volume":"47","author":"KR Jones","year":"2009","unstructured":"Jones KR, Joo YM, Jang S, Yoo YJ, Lee HS, Chung IS, et al. Polymorphism in the CagA EPIYA motif impacts development of gastric cancer. J Clin Microbiol. 2009;47:959\u201368.","journal-title":"J Clin Microbiol"},{"key":"5130_CR14","doi-asserted-by":"publisher","first-page":"1062","DOI":"10.1099\/jmm.0.2008\/001818-0","volume":"57","author":"RH Argent","year":"2008","unstructured":"Argent RH, Hale JL, El-Omar EM, Atherton JC. Differences in Helicobacter pylori CagA tyrosine phosphorylation motif patterns between western and East Asian strains, and influences on interleukin-8 secretion. J Med Microbiol. 2008;57:1062\u20137.","journal-title":"J Med Microbiol"},{"key":"5130_CR15","doi-asserted-by":"publisher","first-page":"820","DOI":"10.1086\/381782","volume":"189","author":"T Azuma","year":"2004","unstructured":"Azuma T, Yamazaki S, Yamakawa A, Ohtani M, Muramatsu A, Suto H, et al. Association between diversity in the Src homology 2 domain\u2013containing tyrosine phosphatase binding site of Helicobacter pylori CagA protein and gastric atrophy and cancer. J Infect Dis. 2004;189:820\u20137.","journal-title":"J Infect Dis"},{"key":"5130_CR16","doi-asserted-by":"publisher","first-page":"14428","DOI":"10.1073\/pnas.222375399","volume":"99","author":"H Higashi","year":"2002","unstructured":"Higashi H, Tsutsumi R, Fujita A, Yamazaki S, Asaka M, Azuma T, et al. Biological activity of the Helicobacter pylori virulence factor CagA is determined by variation in the tyrosine phosphorylation sites. Proc Natl Acad Sci USA. 2002;99:14428\u201333.","journal-title":"Proc Natl Acad Sci USA"},{"key":"5130_CR17","doi-asserted-by":"crossref","unstructured":"McClain MS, Beckett AC, Cover TL. Helicobacter pylori Vacuolating Toxin and Gastric Cancer. Toxins 2017;9.","DOI":"10.3390\/toxins9100316"},{"key":"5130_CR18","doi-asserted-by":"publisher","first-page":"92","DOI":"10.3389\/fcimb.2012.00092","volume":"2","author":"SL Palframan","year":"2012","unstructured":"Palframan SL, Kwok T, Gabriel K. Vacuolating cytotoxin A (VacA), a key toxin for Helicobacter pylori pathogenesis. Front Cell Infect Microbiol. 2012;2:92.","journal-title":"Front Cell Infect Microbiol"},{"key":"5130_CR19","doi-asserted-by":"publisher","first-page":"926","DOI":"10.1053\/j.gastro.2007.06.056","volume":"133","author":"JL Rhead","year":"2007","unstructured":"Rhead JL, Letley DP, Mohammadi M, Hussein N, Mohagheghi MA, Eshagh Hosseini M, et al. A new Helicobacter pylori vacuolating cytotoxin determinant, the intermediate region, is associated with gastric cancer. Gastroenterology. 2007;133:926\u201336.","journal-title":"Gastroenterology"},{"key":"5130_CR20","first-page":"1582","volume-title":"Traces of human migrations in Helicobacter pylori populations","author":"D Falush","year":"2003","unstructured":"Falush D, Wirth T, Linz B, Pritchard JK, Stephens M, Kidd M, et al. Traces of human migrations in Helicobacter pylori populations. Volume 299. New York, NY: Science; 2003. pp. 1582\u20135."},{"key":"5130_CR21","doi-asserted-by":"publisher","first-page":"e1006546","DOI":"10.1371\/journal.pgen.1006546","volume":"13","author":"K Thorell","year":"2017","unstructured":"Thorell K, Yahara K, Berthenet E, Lawson DJ, Mikhail J, Kato I, et al. Rapid evolution of distinct Helicobacter pylori subpopulations in the Americas. PLoS Genet. 2017;13:e1006546.","journal-title":"PLoS Genet"},{"key":"5130_CR22","doi-asserted-by":"publisher","first-page":"915","DOI":"10.1038\/nature05562","volume":"445","author":"B Linz","year":"2007","unstructured":"Linz B, Balloux F, Moodley Y, Manica A, Liu H, Roumagnac P, et al. An African origin for the intimate association between humans and Helicobacter pylori. Nature. 2007;445:915\u20138.","journal-title":"Nature"},{"key":"5130_CR23","doi-asserted-by":"crossref","unstructured":"Vital JS, Tanoeiro L, Lopes-Oliveira R, Vale FF. Biomarker Characterization and Prediction of Virulence and Antibiotic Resistance from Helicobacter pylori Next Generation Sequencing Data. Biomolecules 2022;12.","DOI":"10.3390\/biom12050691"},{"key":"5130_CR24","doi-asserted-by":"publisher","first-page":"8184","DOI":"10.1038\/s41467-023-43562-y","volume":"14","author":"K Thorell","year":"2023","unstructured":"Thorell K, Mu\u00f1oz-Ram\u00edrez ZY, Wang D, Sandoval-Motta S, Boscolo Agostini R, Ghirotto S, et al. The Helicobacter pylori Genome Project: insights into H. pylori population structure from analysis of a worldwide collection of complete genomes. Nat Commun. 2023;14:8184.","journal-title":"Nat Commun"},{"key":"5130_CR25","doi-asserted-by":"publisher","first-page":"104","DOI":"10.1186\/1471-2180-11-104","volume":"11","author":"M Kawai","year":"2011","unstructured":"Kawai M, Furuta Y, Yahara K, Tsuru T, Oshima K, Handa N, et al. Evolution in an oncogenic bacterial species with extreme genome plasticity: Helicobacter pylori East Asian genomes. BMC Microbiol. 2011;11:104.","journal-title":"BMC Microbiol"},{"key":"5130_CR26","doi-asserted-by":"publisher","first-page":"1","DOI":"10.3904\/kjim.2004.19.1.1","volume":"19","author":"JW Kim","year":"2004","unstructured":"Kim JW, Kim JG, Chae SL, Cha YJ, Park SM. High prevalence of multiple strain colonization of Helicobacter pylori in Korean patients: DNA diversity among clinical isolates from the gastric corpus, antrum and duodenum. Korean J Intern Med. 2004;19:1\u20139.","journal-title":"Korean J Intern Med"},{"key":"5130_CR27","doi-asserted-by":"publisher","first-page":"e0137078","DOI":"10.1371\/journal.pone.0137078","volume":"10","author":"A Kim","year":"2015","unstructured":"Kim A, Servetas SL, Kang J, Kim J, Jang S, Cha HJ, et al. Helicobacter pylori bab Paralog Distribution and Association with cagA, vacA, and homA\/B Genotypes in American and South Korean Clinical Isolates. PLoS ONE. 2015;10:e0137078.","journal-title":"PLoS ONE"},{"key":"5130_CR28","doi-asserted-by":"publisher","first-page":"e12733","DOI":"10.1111\/hel.12733","volume":"25","author":"EJ Gong","year":"2020","unstructured":"Gong EJ, Ahn JY, Jung DK, Lee SM, Pih GY, Kim GH, et al. Isolation of Helicobacter pylori using leftover tissue in the rapid urease test kit. Helicobacter. 2020;25:e12733.","journal-title":"Helicobacter"},{"key":"5130_CR29","unstructured":"EUCAST. Clinical breakpoints-breakpoints and guidance. Available online: https:\/\/www.eucast.org\/clinical_breakpoints."},{"key":"5130_CR30","first-page":"2114","volume":"30","author":"AM Bolger","year":"2014","unstructured":"Bolger AM, Lohse M, Usadel B. Trimmomatic: a flexible trimmer for Illumina sequence data. Bioinf (Oxford England). 2014;30:2114\u201320.","journal-title":"Bioinf (Oxford England)"},{"key":"5130_CR31","unstructured":"Li H. Aligning sequence reads, clone sequences and assembly contigs with BWA-MEM. arXiv preprint arXiv:13033997 2013."},{"key":"5130_CR32","doi-asserted-by":"publisher","first-page":"e1005595","DOI":"10.1371\/journal.pcbi.1005595","volume":"13","author":"RR Wick","year":"2017","unstructured":"Wick RR, Judd LM, Gorrie CL, Holt KE. Unicycler: Resolving bacterial genome assemblies from short and long sequencing reads. PLoS Comput Biol. 2017;13:e1005595.","journal-title":"PLoS Comput Biol"},{"key":"5130_CR33","first-page":"2032","volume":"31","author":"A Tarasov","year":"2015","unstructured":"Tarasov A, Vilella AJ, Cuppen E, Nijman IJ, Prins P. Sambamba: fast processing of NGS alignment formats. Bioinf (Oxford England). 2015;31:2032\u20134.","journal-title":"Bioinf (Oxford England)"},{"key":"5130_CR34","first-page":"2078","volume":"25","author":"H Li","year":"2009","unstructured":"Li H, Handsaker B, Wysoker A, Fennell T, Ruan J, Homer N, et al. The Sequence Alignment\/Map format and SAMtools. Bioinf (Oxford England). 2009;25:2078\u20139.","journal-title":"Bioinf (Oxford England)"},{"key":"5130_CR35","doi-asserted-by":"publisher","first-page":"e102","DOI":"10.1002\/cpbi.102","volume":"70","author":"A Prjibelski","year":"2020","unstructured":"Prjibelski A, Antipov D, Meleshko D, Lapidus A, Korobeynikov A. Using SPAdes De Novo Assembler. Curr Protoc Bioinf. 2020;70:e102.","journal-title":"Curr Protoc Bioinf"},{"key":"5130_CR36","first-page":"3094","volume":"34","author":"H Li","year":"2018","unstructured":"Li H. Minimap2: pairwise alignment for nucleotide sequences. Bioinf (Oxford England). 2018;34:3094\u2013100.","journal-title":"Bioinf (Oxford England)"},{"key":"5130_CR37","first-page":"2068","volume":"30","author":"T Seemann","year":"2014","unstructured":"Seemann T. Prokka: rapid prokaryotic genome annotation. Bioinf (Oxford England). 2014;30:2068\u20139.","journal-title":"Bioinf (Oxford England)"},{"key":"5130_CR38","doi-asserted-by":"publisher","first-page":"W484","DOI":"10.1093\/nar\/gkad326","volume":"51","author":"JR Grant","year":"2023","unstructured":"Grant JR, Enns E, Marinier E, Mandal A, Herman EK, Chen CY, et al. Proksee: in-depth characterization and visualization of bacterial genomes. Nucleic Acids Res. 2023;51:W484\u201392.","journal-title":"Nucleic Acids Res"},{"key":"5130_CR39","doi-asserted-by":"publisher","first-page":"1792","DOI":"10.1093\/nar\/gkh340","volume":"32","author":"RC Edgar","year":"2004","unstructured":"Edgar RC. MUSCLE: multiple sequence alignment with high accuracy and high throughput. Nucleic Acids Res. 2004;32:1792\u20137.","journal-title":"Nucleic Acids Res"},{"key":"5130_CR40","doi-asserted-by":"publisher","first-page":"3022","DOI":"10.1093\/molbev\/msab120","volume":"38","author":"K Tamura","year":"2021","unstructured":"Tamura K, Stecher G, Kumar S. MEGA11: Molecular Evolutionary Genetics Analysis Version 11. Mol Biol Evol. 2021;38:3022\u20137.","journal-title":"Mol Biol Evol"},{"key":"5130_CR41","first-page":"3691","volume":"31","author":"AJ Page","year":"2015","unstructured":"Page AJ, Cummins CA, Hunt M, Wong VK, Reuter S, Holden MT, et al. Roary: rapid large-scale prokaryote pan genome analysis. Bioinf (Oxford England). 2015;31:3691\u20133.","journal-title":"Bioinf (Oxford England)"},{"key":"5130_CR42","doi-asserted-by":"publisher","first-page":"D325","DOI":"10.1093\/nar\/gki008","volume":"33","author":"L Chen","year":"2005","unstructured":"Chen L, Yang J, Yu J, Yao Z, Sun L, Shen Y, et al. VFDB: a reference database for bacterial virulence factors. Nucleic Acids Res. 2005;33:D325\u20138.","journal-title":"Nucleic Acids Res"},{"key":"5130_CR43","doi-asserted-by":"publisher","first-page":"17771","DOI":"10.1074\/jbc.270.30.17771","volume":"270","author":"JC Atherton","year":"1995","unstructured":"Atherton JC, Cao P, Peek RM Jr., Tummuru MK, Blaser MJ, Cover TL. Mosaicism in vacuolating cytotoxin alleles of Helicobacter pylori. Association of specific vacA types with cytotoxin production and peptic ulceration. J Biol Chem. 1995;270:17771\u20137.","journal-title":"J Biol Chem"},{"key":"5130_CR44","doi-asserted-by":"publisher","first-page":"1710","DOI":"10.1128\/jcm.35.7.1710-1714.1997","volume":"35","author":"Y Ito","year":"1997","unstructured":"Ito Y, Azuma T, Ito S, Miyaji H, Hirai M, Yamazaki Y, et al. Analysis and typing of the vacA gene from cagA-positive strains of Helicobacter pylori isolated in Japan. J Clin Microbiol. 1997;35:1710\u20134.","journal-title":"J Clin Microbiol"},{"key":"5130_CR45","doi-asserted-by":"publisher","first-page":"2274","DOI":"10.1128\/JCM.37.7.2274-2279.1999","volume":"37","author":"Y Yamaoka","year":"1999","unstructured":"Yamaoka Y, Kodama T, Gutierrez O, Kim JG, Kashima K, Graham DY. Relationship between Helicobacter pylori iceA, cagA, and vacA status and clinical outcome: studies in four different countries. J Clin Microbiol. 1999;37:2274\u20139.","journal-title":"J Clin Microbiol"},{"key":"5130_CR46","doi-asserted-by":"publisher","first-page":"2979","DOI":"10.1128\/JCM.37.9.2979-2982.1999","volume":"37","author":"JC Atherton","year":"1999","unstructured":"Atherton JC, Cover TL, Twells RJ, Morales MR, Hawkey CJ, Blaser MJ. Simple and accurate PCR-based system for typing vacuolating cytotoxin alleles of Helicobacter pylori. J Clin Microbiol. 1999;37:2979\u201382.","journal-title":"J Clin Microbiol"},{"key":"5130_CR47","doi-asserted-by":"publisher","first-page":"539","DOI":"10.1038\/41483","volume":"388","author":"JF Tomb","year":"1997","unstructured":"Tomb JF, White O, Kerlavage AR, Clayton RA, Sutton GG, Fleischmann RD, et al. The complete genome sequence of the gastric pathogen Helicobacter pylori. Nature. 1997;388:539\u201347.","journal-title":"Nature"},{"key":"5130_CR48","doi-asserted-by":"publisher","first-page":"12619","DOI":"10.1073\/pnas.95.21.12619","volume":"95","author":"S Suerbaum","year":"1998","unstructured":"Suerbaum S, Smith JM, Bapumia K, Morelli G, Smith NH, Kunstmann E, et al. Free recombination within Helicobacter pylori. Proc Natl Acad Sci USA. 1998;95:12619\u201324.","journal-title":"Proc Natl Acad Sci USA"},{"key":"5130_CR49","doi-asserted-by":"publisher","first-page":"960","DOI":"10.1007\/s10620-005-9030-z","volume":"52","author":"KD Choi","year":"2007","unstructured":"Choi KD, Kim N, Lee DH, Kim JM, Kim JS, Jung HC, et al. Analysis of the 3\u2019 variable region of the cagA gene of Helicobacter pylori isolated in Koreans. Dig Dis Sci. 2007;52:960\u20136.","journal-title":"Dig Dis Sci"},{"key":"5130_CR50","doi-asserted-by":"publisher","first-page":"232","DOI":"10.1046\/j.1523-5378.2002.00094.x","volume":"7","author":"JS Ko","year":"2002","unstructured":"Ko JS, Seo JK. cag pathogenicity island of Helicobacter pylori in Korean children. Helicobacter. 2002;7:232\u20136.","journal-title":"Helicobacter"},{"key":"5130_CR51","doi-asserted-by":"publisher","first-page":"1093","DOI":"10.1007\/s42770-020-00292-3","volume":"51","author":"W Boonyanugomol","year":"2020","unstructured":"Boonyanugomol W, Kongkasame W, Palittapongarnpim P, Baik SC, Jung MH, Shin MK, et al. Genetic variation in the cag pathogenicity island of Helicobacter pylori strains detected from gastroduodenal patients in Thailand. Braz J Microbiol. 2020;51:1093\u2013101.","journal-title":"Braz J Microbiol"},{"key":"5130_CR52","doi-asserted-by":"publisher","first-page":"e17881","DOI":"10.1016\/j.heliyon.2023.e17881","volume":"9","author":"M Yu","year":"2023","unstructured":"Yu M, Xu M, Shen Y, Liu Y, Xu C, Feng T, et al. Hp0521 inhibited the virulence of H. pylori 26,695 strain via regulating CagA expression. Heliyon. 2023;9:e17881.","journal-title":"Heliyon"},{"key":"5130_CR53","doi-asserted-by":"publisher","first-page":"154","DOI":"10.1007\/s00284-024-03653-5","volume":"81","author":"RSP Rao","year":"2024","unstructured":"Rao RSP, Ghate SD, Pinto L, Suravajhala P, Patil P, Shetty P, et al. Extent of Virulence and Antibiotic Resistance Genes in Helicobacter pylori and Campylobacteria. Curr Microbiol. 2024;81:154.","journal-title":"Curr Microbiol"},{"key":"5130_CR54","doi-asserted-by":"publisher","first-page":"1164","DOI":"10.1053\/j.gastro.2016.08.014","volume":"151","author":"RM Barrozo","year":"2016","unstructured":"Barrozo RM, Hansen LM, Lam AM, Skoog EC, Martin ME, Cai LP, et al. CagY Is an Immune-Sensitive Regulator of the Helicobacter pylori Type IV Secretion System. Gastroenterology. 2016;151:1164\u2013e753.","journal-title":"Gastroenterology"},{"key":"5130_CR55","doi-asserted-by":"publisher","first-page":"1349","DOI":"10.1084\/jem.20030381","volume":"198","author":"RA Aras","year":"2003","unstructured":"Aras RA, Fischer W, Perez-Perez GI, Crosatti M, Ando T, Haas R, et al. Plasticity of repetitive DNA sequences within a bacterial (Type IV) secretion system component. J Exp Med. 2003;198:1349\u201360.","journal-title":"J Exp Med"},{"key":"5130_CR56","doi-asserted-by":"publisher","first-page":"e1003189","DOI":"10.1371\/journal.ppat.1003189","volume":"9","author":"RM Barrozo","year":"2013","unstructured":"Barrozo RM, Cooke CL, Hansen LM, Lam AM, Gaddy JA, Johnson EM, et al. Functional plasticity in the type IV secretion system of Helicobacter pylori. PLoS Pathog. 2013;9:e1003189.","journal-title":"PLoS Pathog"},{"key":"5130_CR57","doi-asserted-by":"publisher","first-page":"441","DOI":"10.1038\/nrmicro1658","volume":"5","author":"S Suerbaum","year":"2007","unstructured":"Suerbaum S, Josenhans C. Helicobacter pylori evolution and phenotypic diversification in a changing host. Nat Rev Microbiol. 2007;5:441\u201352.","journal-title":"Nat Rev Microbiol"},{"key":"5130_CR58","doi-asserted-by":"publisher","first-page":"e0159419","DOI":"10.1371\/journal.pone.0159419","volume":"11","author":"I Uchiyama","year":"2016","unstructured":"Uchiyama I, Albritton J, Fukuyo M, Kojima KK, Yahara K, Kobayashi I. A Novel Approach to Helicobacter pylori Pan-Genome Analysis for Identification of Genomic Islands. PLoS ONE. 2016;11:e0159419.","journal-title":"PLoS ONE"},{"key":"5130_CR59","doi-asserted-by":"publisher","first-page":"249","DOI":"10.1128\/JB.188.1.249-254.2006","volume":"188","author":"C Kraft","year":"2006","unstructured":"Kraft C, Stack A, Josenhans C, Niehus E, Dietrich G, Correa P, et al. Genomic changes during chronic Helicobacter pylori infection. J Bacteriol. 2006;188:249\u201354.","journal-title":"J Bacteriol"},{"key":"5130_CR60","doi-asserted-by":"publisher","first-page":"e1001036","DOI":"10.1371\/journal.pgen.1001036","volume":"6","author":"G Morelli","year":"2010","unstructured":"Morelli G, Didelot X, Kusecek B, Schwarz S, Bahlawane C, Falush D, et al. Microevolution of Helicobacter pylori during prolonged infection of single hosts and within families. PLoS Genet. 2010;6:e1001036.","journal-title":"PLoS Genet"},{"key":"5130_CR61","doi-asserted-by":"publisher","first-page":"3984","DOI":"10.3748\/wjg.15.3984","volume":"15","author":"QJ Dong","year":"2009","unstructured":"Dong QJ, Wang Q, Xin YN, Li N, Xuan SY. Comparative genomics of Helicobacter pylori. World J Gastroenterol. 2009;15:3984\u201391.","journal-title":"World J Gastroenterol"}],"container-title":["BMC Microbiology"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/link.springer.com\/article\/10.1186\/s12866-026-05130-4","content-type":"text\/html","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/link.springer.com\/content\/pdf\/10.1186\/s12866-026-05130-4.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/link.springer.com\/content\/pdf\/10.1186\/s12866-026-05130-4.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,7,3]],"date-time":"2026-07-03T04:11:07Z","timestamp":1783051867000},"score":1,"resource":{"primary":{"URL":"https:\/\/link.springer.com\/10.1186\/s12866-026-05130-4"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,5,11]]},"references-count":61,"journal-issue":{"issue":"1","published-online":{"date-parts":[[2026,12]]}},"alternative-id":["5130"],"URL":"https:\/\/doi.org\/10.1186\/s12866-026-05130-4","relation":{},"ISSN":["1471-2180"],"issn-type":[{"value":"1471-2180","type":"electronic"}],"subject":[],"published":{"date-parts":[[2026,5,11]]},"assertion":[{"value":"5 December 2025","order":1,"name":"received","label":"Received","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"4 May 2026","order":2,"name":"accepted","label":"Accepted","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"11 May 2026","order":3,"name":"first_online","label":"First Online","group":{"name":"ArticleHistory","label":"Article History"}},{"order":1,"name":"Ethics","group":{"name":"EthicsHeading","label":"Declarations"}},{"value":"The study protocol was approved by the Institutional Review Board (IRB) of Hallym University Chuncheon Sacred Hospital (No. 2025-08-009). The IRB waived the requirement for written informed consent, as this study involved only previously collected, anonymized\n                      H. pylori\n                      bacterial strains and de-identified patient information, in accordance with national research ethics guidelines. All study protocols and methods were performed in accordance with the relevant guidelines and regulations, including the principles of the Declaration of Helsinki.\u201d","order":2,"name":"Ethics","group":{"name":"EthicsHeading","label":"Ethics approval and consent to participate"}},{"value":"Not applicable.","order":3,"name":"Ethics","group":{"name":"EthicsHeading","label":"Consent for publication"}},{"value":"The authors declare no competing interests.","order":4,"name":"Ethics","group":{"name":"EthicsHeading","label":"Competing interests"}}],"article-number":"588"}}