{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,3,5]],"date-time":"2026-03-05T08:05:43Z","timestamp":1772697943362,"version":"3.50.1"},"reference-count":41,"publisher":"Springer Science and Business Media LLC","issue":"3","license":[{"start":{"date-parts":[[2025,10,20]],"date-time":"2025-10-20T00:00:00Z","timestamp":1760918400000},"content-version":"tdm","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by-nc-nd\/4.0"},{"start":{"date-parts":[[2025,10,20]],"date-time":"2025-10-20T00:00:00Z","timestamp":1760918400000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by-nc-nd\/4.0"}],"funder":[{"DOI":"10.13039\/501100001691","name":"Japan Society for the Promotion of Science","doi-asserted-by":"publisher","award":["JP 24K02251"],"award-info":[{"award-number":["JP 24K02251"]}],"id":[{"id":"10.13039\/501100001691","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100001691","name":"Japan Society for the Promotion of Science","doi-asserted-by":"publisher","award":["JP22K06981"],"award-info":[{"award-number":["JP22K06981"]}],"id":[{"id":"10.13039\/501100001691","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100011746","name":"NOVARTIS Foundation (Japan) for the Promotion of Science","doi-asserted-by":"publisher","award":["JP 24K10000"],"award-info":[{"award-number":["JP 24K10000"]}],"id":[{"id":"10.13039\/501100011746","id-type":"DOI","asserted-by":"publisher"}]}],"content-domain":{"domain":["link.springer.com"],"crossmark-restriction":false},"short-container-title":["Virchows Arch"],"published-print":{"date-parts":[[2026,3]]},"DOI":"10.1007\/s00428-025-04302-3","type":"journal-article","created":{"date-parts":[[2025,10,20]],"date-time":"2025-10-20T09:00:31Z","timestamp":1760950831000},"page":"569-583","update-policy":"https:\/\/doi.org\/10.1007\/springer_crossmark_policy","source":"Crossref","is-referenced-by-count":1,"title":["Spatial transcriptomic analysis of foveolar-type gastric adenoma with raspberry-like appearance"],"prefix":"10.1007","volume":"488","author":[{"ORCID":"https:\/\/orcid.org\/0000-0002-7466-7356","authenticated-orcid":false,"given":"Sohei","family":"Kitazawa","sequence":"first","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Eiji","family":"Takeshita","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Hideomi","family":"Tomida","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Teruki","family":"Miyake","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Yoshio","family":"Ikeda","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Seiya","family":"Bekku","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Ryuma","family":"Haraguchi","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Yuki","family":"Takaoka","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Teruyuki","family":"Ohno","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Mashio","family":"Taniwaki","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Yoichi","family":"Hiasa","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Riko","family":"Kitazawa","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]}],"member":"297","published-online":{"date-parts":[[2025,10,20]]},"reference":[{"key":"4302_CR1","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1007\/s00535-023-02050-2","volume":"59","author":"S Kotani","year":"2024","unstructured":"Kotani S, Shibagaki K, Hirahara N, Hasegawa N, Tanabe R, Ebisutani Y, Nonomura S, Kishimoto K, Kodama Y, Takahashi Y, Kataoka M, Oka A, Fukuba N, Mishima Y, Oshima N, Kawashima K, Ishimura N, Araki A, Kadota K, Itawaki A, Nagasaki M, Miyaoka Y, Onuma H, Ishihara S (2024) Clinicopathologic differences of gastric neoplasms between Helicobacter pylori-infected and -naive patients: a multicenter retrospective analysis. J Gastroenterol 59:1\u201310. https:\/\/doi.org\/10.1007\/s00535-023-02050-2","journal-title":"J Gastroenterol"},{"key":"4302_CR2","doi-asserted-by":"publisher","DOI":"10.1111\/den.14980","author":"K Shibagaki","year":"2025","unstructured":"Shibagaki K, Kushima R, Sekine S, Mishiro T, Kotani S, Miyaoka Y, Ishimura N, Araki A, Ohnuma H, Niino D, Ishihara S (2025) Spectrum of gastric neoplasms in Helicobacter pylori-naive patients. Dig Endosc. https:\/\/doi.org\/10.1111\/den.14980","journal-title":"Dig Endosc"},{"key":"4302_CR3","doi-asserted-by":"publisher","first-page":"423","DOI":"10.1111\/pin.13456","volume":"74","author":"K Shibagaki","year":"2024","unstructured":"Shibagaki K, Kushima R, Mishiro T, Araki A, Niino D, Ishimura N, Ishihara S (2024) Gastric dysplastic lesions in Helicobacter pylori-naive stomach: foveolar-type adenoma and intestinal-type dysplasia. Pathol Int 74:423\u2013437. https:\/\/doi.org\/10.1111\/pin.13456","journal-title":"Pathol Int"},{"key":"4302_CR4","doi-asserted-by":"publisher","first-page":"783","DOI":"10.1007\/s00428-021-03237-9","volume":"480","author":"K Shibagaki","year":"2022","unstructured":"Shibagaki K, Itawaki A, Miyaoka Y, Kishimoto K, Takahashi Y, Kotani S, Mishiro T, Oshima N, Kawashima K, Ishimura N, Onuma H, Nagasaki M, Nagase M, Araki A, Kadota K, Kushima R, Ishihara S (2022) Intestinal-type gastric dysplasia in Helicobacter pylori-naive patients. Virchows Arch 480:783\u2013792. https:\/\/doi.org\/10.1007\/s00428-021-03237-9","journal-title":"Virchows Arch"},{"key":"4302_CR5","doi-asserted-by":"publisher","first-page":"971","DOI":"10.1038\/bjc.1998.611","volume":"78","author":"MS Wu","year":"1998","unstructured":"Wu MS, Shun CT, Lee WC, Chen CJ, Wang HP, Lee WJ, Sheu JC, Lin JT (1998) Overexpression of p53 in different subtypes of intestinal metaplasia and gastric cancer. Br J Cancer 78:971\u2013973. https:\/\/doi.org\/10.1038\/bjc.1998.611","journal-title":"Br J Cancer"},{"key":"4302_CR6","first-page":"551","volume":"1","author":"M Rugge","year":"1992","unstructured":"Rugge M, Shiao YH, Correa P, Baffa R, DiMario F (1992) Immunohistochemical evidence of p53 overexpression in gastric epithelial dysplasia. Cancer Epidemiol Biomarkers Prev 1:551\u2013554","journal-title":"Cancer Epidemiol Biomarkers Prev"},{"key":"4302_CR7","doi-asserted-by":"publisher","DOI":"10.3390\/ijerph192214829","author":"S Betschart","year":"2022","unstructured":"Betschart S, Sandmeier A, Skedsmo G, Hascher T, Okan O, Dadaczynski K (2022) The importance of school leaders\u2019 attitudes and health literacy to the implementation of a health-promoting schools approach. Int J Environ Res Public Health. https:\/\/doi.org\/10.3390\/ijerph192214829","journal-title":"Int J Environ Res Public Health"},{"key":"4302_CR8","doi-asserted-by":"publisher","first-page":"1307","DOI":"10.1007\/s10120-021-01208-2","volume":"24","author":"K Imamura","year":"2021","unstructured":"Imamura K, Yao K, Nimura S, Tanabe H, Kanemitsu T, Miyaoka M, Ono Y, Ueki T, Iwashita A (2021) Characteristic endoscopic findings of gastric adenocarcinoma of fundic-gland mucosa type. Gastric Cancer 24:1307\u20131319. https:\/\/doi.org\/10.1007\/s10120-021-01208-2","journal-title":"Gastric Cancer"},{"key":"4302_CR9","doi-asserted-by":"publisher","first-page":"1250","DOI":"10.1016\/j.ajpath.2022.06.005","volume":"192","author":"T Mishiro","year":"2022","unstructured":"Mishiro T, Shibagaki K, Fukuyama C, Kataoka M, Notsu T, Yamashita N, Oka A, Nagase M, Araki A, Kawashima K, Ishimura N, Maruyama R, Kinoshita Y, Ishihara S (2022) KLF4 mutation shapes pathologic characteristics of foveolar-type gastric adenoma in Helicobacter pylori-naive patients. Am J Pathol 192:1250\u20131258. https:\/\/doi.org\/10.1016\/j.ajpath.2022.06.005","journal-title":"Am J Pathol"},{"key":"4302_CR10","doi-asserted-by":"publisher","first-page":"E784","DOI":"10.1055\/a-0854-3818","volume":"7","author":"K Shibagaki","year":"2019","unstructured":"Shibagaki K, Fukuyama C, Mikami H, Izumi D, Yamashita N, Mishiro T, Oshima N, Ishimura N, Sato S, Ishihara S, Nagase M, Araki A, Ishikawa N, Maruyama R, Kushima R, Kinoshita Y (2019) Gastric foveolar-type adenomas endoscopically showing a raspberry-like appearance in the Helicobacter pylori -uninfected stomach. Endosc Int Open 7:E784\u2013E791. https:\/\/doi.org\/10.1055\/a-0854-3818","journal-title":"Endosc Int Open"},{"key":"4302_CR11","unstructured":"Kushima R LG, Rugge M. Gastric dysplasia. In: WHO Classification of Tumours Editorial Board. Digestive System Tumours. 5th ed. Lyon: International Agency for Research on Cancer; 2019. p. 71\u201375. (2019) Gastric dysplasia WHO Classification of Tumours Editorial Board. Digestive system tumours. International Agency for Research on Cancer, Lyon, pp 71\u201375"},{"key":"4302_CR12","doi-asserted-by":"publisher","first-page":"611","DOI":"10.1016\/S0002-9440(10)64216-2","volume":"161","author":"JH Lee","year":"2002","unstructured":"Lee JH, Abraham SC, Kim HS, Nam JH, Choi C, Lee MC, Park CS, Juhng SW, Rashid A, Hamilton SR, Wu TT (2002) Inverse relationship between APC gene mutation in gastric adenomas and development of adenocarcinoma. Am J Pathol 161:611\u2013618. https:\/\/doi.org\/10.1016\/S0002-9440(10)64216-2","journal-title":"Am J Pathol"},{"key":"4302_CR13","doi-asserted-by":"publisher","first-page":"786","DOI":"10.1097\/01.MP.0000080349.37658.5E","volume":"16","author":"SC Abraham","year":"2003","unstructured":"Abraham SC, Park SJ, Lee JH, Mugartegui L, Wu TT (2003) Genetic alterations in gastric adenomas of intestinal and foveolar phenotypes. Mod Pathol 16:786\u2013795. https:\/\/doi.org\/10.1097\/01.MP.0000080349.37658.5E","journal-title":"Mod Pathol"},{"key":"4302_CR14","doi-asserted-by":"publisher","first-page":"4604","DOI":"10.1128\/MCB.16.9.4604","volume":"16","author":"JA Forsythe","year":"1996","unstructured":"Forsythe JA, Jiang BH, Iyer NV, Agani F, Leung SW, Koos RD, Semenza GL (1996) Activation of vascular endothelial growth factor gene transcription by hypoxia-inducible factor 1. Mol Cell Biol 16:4604\u20134613. https:\/\/doi.org\/10.1128\/MCB.16.9.4604","journal-title":"Mol Cell Biol"},{"key":"4302_CR15","doi-asserted-by":"publisher","first-page":"8104","DOI":"10.1073\/pnas.94.15.8104","volume":"94","author":"PH Maxwell","year":"1997","unstructured":"Maxwell PH, Dachs GU, Gleadle JM, Nicholls LG, Harris AL, Stratford IJ, Hankinson O, Pugh CW, Ratcliffe PJ (1997) Hypoxia-inducible factor-1 modulates gene expression in solid tumors and influences both angiogenesis and tumor growth. Proc Natl Acad Sci U S A 94:8104\u20138109. https:\/\/doi.org\/10.1073\/pnas.94.15.8104","journal-title":"Proc Natl Acad Sci U S A"},{"key":"4302_CR16","doi-asserted-by":"publisher","first-page":"138","DOI":"10.1002\/ijc.11680","volume":"109","author":"H Ohata","year":"2004","unstructured":"Ohata H, Kitauchi S, Yoshimura N, Mugitani K, Iwane M, Nakamura H, Yoshikawa A, Yanaoka K, Arii K, Tamai H, Shimizu Y, Takeshita T, Mohara O, Ichinose M (2004) Progression of chronic atrophic gastritis associated with Helicobacter pylori infection increases risk of gastric cancer. Int J Cancer 109:138\u2013143. https:\/\/doi.org\/10.1002\/ijc.11680","journal-title":"Int J Cancer"},{"key":"4302_CR17","doi-asserted-by":"publisher","DOI":"10.1136\/bmjopen-2017-015855","volume":"7","author":"S Hiroi","year":"2017","unstructured":"Hiroi S, Sugano K, Tanaka S, Kawakami K (2017) Impact of health insurance coverage for Helicobacter pylori gastritis on the trends in eradication therapy in Japan: retrospective observational study and simulation study based on real-world data. BMJ Open 7:e015855. https:\/\/doi.org\/10.1136\/bmjopen-2017-015855","journal-title":"BMJ Open"},{"key":"4302_CR18","doi-asserted-by":"publisher","first-page":"551","DOI":"10.1111\/den.12471","volume":"27","author":"Y Yamamoto","year":"2015","unstructured":"Yamamoto Y, Fujisaki J, Omae M, Hirasawa T, Igarashi M (2015) Helicobacter pylori-negative gastric cancer: characteristics and endoscopic findings. Dig Endosc 27:551\u2013561. https:\/\/doi.org\/10.1111\/den.12471","journal-title":"Dig Endosc"},{"key":"4302_CR19","doi-asserted-by":"publisher","first-page":"2775","DOI":"10.2169\/internalmedicine.1265-22","volume":"62","author":"U Sakamoto","year":"2023","unstructured":"Sakamoto U, Adachi K, Kishi K, Okimoto E, Ishimura N, Miura H, Ishihara S (2023) Prevalence of raspberry-type gastric foveolar-type tumor in individuals undergoing medical checkups. Intern Med 62:2775\u20132781. https:\/\/doi.org\/10.2169\/internalmedicine.1265-22","journal-title":"Intern Med"},{"key":"4302_CR20","doi-asserted-by":"publisher","first-page":"521","DOI":"10.1097\/PAS.0000000000002012","volume":"47","author":"T Naka","year":"2023","unstructured":"Naka T, Hashimoto T, Yoshida T, Yatabe Y, Sekine S (2023) KLF4 c.A1322C mutation is a consistent and specific genetic feature of raspberry-like foveolar-type adenoma of the stomach. Am J Surg Pathol 47:521\u2013523. https:\/\/doi.org\/10.1097\/PAS.0000000000002012","journal-title":"Am J Surg Pathol"},{"key":"4302_CR21","doi-asserted-by":"publisher","first-page":"2619","DOI":"10.1242\/dev.129.11.2619","volume":"129","author":"JP Katz","year":"2002","unstructured":"Katz JP, Perreault N, Goldstein BG, Lee CS, Labosky PA, Yang VW, Kaestner KH (2002) The zinc-finger transcription factor Klf4 is required for terminal differentiation of goblet cells in the colon. Development 129:2619\u20132628. https:\/\/doi.org\/10.1242\/dev.129.11.2619","journal-title":"Development"},{"key":"4302_CR22","doi-asserted-by":"publisher","first-page":"424","DOI":"10.15283\/ijsc20048","volume":"13","author":"T Katano","year":"2020","unstructured":"Katano T, Bialkowska AB, Yang VW (2020) KLF4 regulates goblet cell differentiation in BMI1(+) reserve intestinal stem cell lineage during homeostasis. Int J Stem Cells 13:424\u2013431. https:\/\/doi.org\/10.15283\/ijsc20048","journal-title":"Int J Stem Cells"},{"key":"4302_CR23","doi-asserted-by":"publisher","first-page":"G490","DOI":"10.1152\/ajpgi.90393.2008","volume":"296","author":"H Zheng","year":"2009","unstructured":"Zheng H, Pritchard DM, Yang X, Bennett E, Liu G, Liu C, Ai W (2009) Klf4 gene expression is inhibited by the notch signaling pathway that controls goblet cell differentiation in mouse gastrointestinal tract. Am J Physiol Gastrointest Liver Physiol 296:G490-498. https:\/\/doi.org\/10.1152\/ajpgi.90393.2008","journal-title":"Am J Physiol Gastrointest Liver Physiol"},{"key":"4302_CR24","doi-asserted-by":"publisher","DOI":"10.1371\/journal.pone.0032492","volume":"7","author":"T Yu","year":"2012","unstructured":"Yu T, Chen X, Zhang W, Li J, Xu R, Wang TC, Ai W, Liu C (2012) Kruppel-like factor 4 regulates intestinal epithelial cell morphology and polarity. PLoS ONE 7:e32492. https:\/\/doi.org\/10.1371\/journal.pone.0032492","journal-title":"PLoS ONE"},{"key":"4302_CR25","doi-asserted-by":"publisher","first-page":"310","DOI":"10.1016\/j.ydbio.2010.11.001","volume":"349","author":"AM Ghaleb","year":"2011","unstructured":"Ghaleb AM, McConnell BB, Kaestner KH, Yang VW (2011) Altered intestinal epithelial homeostasis in mice with intestine-specific deletion of the Kruppel-like factor 4 gene. Dev Biol 349:310\u2013320. https:\/\/doi.org\/10.1016\/j.ydbio.2010.11.001","journal-title":"Dev Biol"},{"key":"4302_CR26","doi-asserted-by":"publisher","first-page":"3712","DOI":"10.1016\/j.yexcr.2008.10.004","volume":"314","author":"M Flandez","year":"2008","unstructured":"Flandez M, Guilmeau S, Blache P, Augenlicht LH (2008) KLF4 regulation in intestinal epithelial cell maturation. Exp Cell Res 314:3712\u20133723. https:\/\/doi.org\/10.1016\/j.yexcr.2008.10.004","journal-title":"Exp Cell Res"},{"key":"4302_CR27","doi-asserted-by":"publisher","first-page":"466","DOI":"10.1006\/bbrc.2002.6475","volume":"291","author":"JR Gum Jr.","year":"2002","unstructured":"Gum JR Jr., Crawley SC, Hicks JW, Szymkowski DE, Kim YS (2002) MUC17, a novel membrane-tethered mucin. Biochem Biophys Res Commun 291:466\u2013475. https:\/\/doi.org\/10.1006\/bbrc.2002.6475","journal-title":"Biochem Biophys Res Commun"},{"key":"4302_CR28","doi-asserted-by":"publisher","first-page":"G1144","DOI":"10.1152\/ajpgi.00138.2010","volume":"300","author":"S Resta-Lenert","year":"2011","unstructured":"Resta-Lenert S, Das S, Batra SK, Ho SB (2011) Muc17 protects intestinal epithelial cells from enteroinvasive E. coli infection by promoting epithelial barrier integrity. Am J Physiol Gastrointest Liver Physiol 300:G1144-1155. https:\/\/doi.org\/10.1152\/ajpgi.00138.2010","journal-title":"Am J Physiol Gastrointest Liver Physiol"},{"key":"4302_CR29","doi-asserted-by":"publisher","first-page":"941","DOI":"10.1007\/s10120-019-00932-0","volume":"22","author":"S Lin","year":"2019","unstructured":"Lin S, Zhang Y, Hu Y, Yang B, Cui J, Huang J, Wang JM, Xing R, Lu Y (2019) Epigenetic downregulation of MUC17 by H. pylori infection facilitates NF-kappaB-mediated expression of CEACAM1-3S in human gastric cancer. Gastric Cancer 22:941\u2013954. https:\/\/doi.org\/10.1007\/s10120-019-00932-0","journal-title":"Gastric Cancer"},{"key":"4302_CR30","doi-asserted-by":"publisher","DOI":"10.1016\/j.tranon.2024.102207","volume":"51","author":"B Yang","year":"2025","unstructured":"Yang B, Xie X, Jin X, Huang X, He Y, Yin K, Ji C, Liu L, Feng Z (2025) Identification and validation of serum MUC17 as a non-invasive early warning biomarker for screening of gastric intraepithelial neoplasia. Transl Oncol 51:102207. https:\/\/doi.org\/10.1016\/j.tranon.2024.102207","journal-title":"Transl Oncol"},{"key":"4302_CR31","doi-asserted-by":"publisher","DOI":"10.3390\/ijms25031858","author":"G Leoncini","year":"2024","unstructured":"Leoncini G, Cari L, Ronchetti S, Donato F, Caruso L, Calafa C, Villanacci V (2024) Mucin expression profiles in ulcerative colitis: new insights on the histological mucosal healing. Int J Mol Sci. https:\/\/doi.org\/10.3390\/ijms25031858","journal-title":"Int J Mol Sci"},{"key":"4302_CR32","doi-asserted-by":"publisher","DOI":"10.3390\/jcm10091935","author":"E Bankole","year":"2021","unstructured":"Bankole E, Read E, Curtis MA, Neves JF, Garnett JA (2021) The relationship between mucins and ulcerative colitis: a systematic review. J Clin Med. https:\/\/doi.org\/10.3390\/jcm10091935","journal-title":"J Clin Med"},{"key":"4302_CR33","doi-asserted-by":"publisher","unstructured":"Layunta E, Javerfelt S, van de Koolwijk FC, Sivertsson M, Dolan B, Arike L, Thulin SI, Vallance BA, Pelaseyed T (2024) MUC17 is an essential small intestinal glycocalyx component that is disrupted in Crohn\u2019s disease. JCI Insight 10. https:\/\/doi.org\/10.1172\/jci.insight.181481","DOI":"10.1172\/jci.insight.181481"},{"key":"4302_CR34","doi-asserted-by":"publisher","first-page":"702","DOI":"10.1136\/jcp.2010.078717","volume":"63","author":"S Senapati","year":"2010","unstructured":"Senapati S, Ho SB, Sharma P, Das S, Chakraborty S, Kaur S, Niehans G, Batra SK (2010) Expression of intestinal MUC17 membrane-bound mucin in inflammatory and neoplastic diseases of the colon. J Clin Pathol 63:702\u2013707. https:\/\/doi.org\/10.1136\/jcp.2010.078717","journal-title":"J Clin Pathol"},{"key":"4302_CR35","doi-asserted-by":"publisher","first-page":"2054","DOI":"10.3389\/fimmu.2020.02054","volume":"11","author":"JA Grondin","year":"2020","unstructured":"Grondin JA, Kwon YH, Far PM, Haq S, Khan WI (2020) Mucins in intestinal mucosal defense and inflammation: learning from clinical and experimental studies. Front Immunol 11:2054. https:\/\/doi.org\/10.3389\/fimmu.2020.02054","journal-title":"Front Immunol"},{"key":"4302_CR36","doi-asserted-by":"publisher","first-page":"8","DOI":"10.1111\/imr.12182","volume":"260","author":"T Pelaseyed","year":"2014","unstructured":"Pelaseyed T, Bergstrom JH, Gustafsson JK, Ermund A, Birchenough GM, Schutte A, van der Post S, Svensson F, Rodriguez-Pineiro AM, Nystrom EE, Wising C, Johansson ME, Hansson GC (2014) The mucus and mucins of the goblet cells and enterocytes provide the first defense line of the gastrointestinal tract and interact with the immune system. Immunol Rev 260:8\u201320. https:\/\/doi.org\/10.1111\/imr.12182","journal-title":"Immunol Rev"},{"key":"4302_CR37","doi-asserted-by":"publisher","first-page":"996","DOI":"10.1016\/j.biocel.2010.03.001","volume":"42","author":"Y Luu","year":"2010","unstructured":"Luu Y, Junker W, Rachagani S, Das S, Batra SK, Heinrikson RL, Shekels LL, Ho SB (2010) Human intestinal MUC17 mucin augments intestinal cell restitution and enhances healing of experimental colitis. Int J Biochem Cell Biol 42:996\u20131006. https:\/\/doi.org\/10.1016\/j.biocel.2010.03.001","journal-title":"Int J Biochem Cell Biol"},{"key":"4302_CR38","doi-asserted-by":"publisher","first-page":"141","DOI":"10.1186\/s12964-023-01162-9","volume":"21","author":"X Liang","year":"2023","unstructured":"Liang X, Du W, Huang L, Xiang L, Pan W, Yang F, Zheng F, Xie Y, Geng L, Gong S, Xu W (2023) Helicobacter pylori promotes gastric intestinal metaplasia through activation of IRF3-mediated kynurenine pathway. Cell Commun Signal 21:141. https:\/\/doi.org\/10.1186\/s12964-023-01162-9","journal-title":"Cell Commun Signal"},{"key":"4302_CR39","doi-asserted-by":"publisher","first-page":"294","DOI":"10.1046\/j.1523-5378.2001.00042.x","volume":"6","author":"M Asaka","year":"2001","unstructured":"Asaka M, Sugiyama T, Nobuta A, Kato M, Takeda H, Graham DY (2001) Atrophic gastritis and intestinal metaplasia in Japan: results of a large multicenter study. Helicobacter 6:294\u2013299. https:\/\/doi.org\/10.1046\/j.1523-5378.2001.00042.x","journal-title":"Helicobacter"},{"key":"4302_CR40","doi-asserted-by":"publisher","first-page":"112","DOI":"10.1002\/(sici)1097-0215(19970220)74:1<112::aid-ijc19>3.0.co;2-h","volume":"74","author":"CA Reis","year":"1997","unstructured":"Reis CA, David L, Nielsen PA, Clausen H, Mirgorodskaya K, Roepstorff P, Sobrinho-Simoes M (1997) Immunohistochemical study of MUC5AC expression in human gastric carcinomas using a novel monoclonal antibody. Int J Cancer 74:112\u2013121. https:\/\/doi.org\/10.1002\/(sici)1097-0215(19970220)74:1%3c112::aid-ijc19%3e3.0.co;2-h","journal-title":"Int J Cancer"},{"key":"4302_CR41","doi-asserted-by":"publisher","first-page":"1399","DOI":"10.3389\/fonc.2019.01399","volume":"9","author":"D Aguilar-Cazares","year":"2019","unstructured":"Aguilar-Cazares D, Chavez-Dominguez R, Carlos-Reyes A, Lopez-Camarillo C, Hernadez de la Cruz ON, Lopez-Gonzalez JS (2019) Contribution of angiogenesis to inflammation and cancer. Front Oncol 9:1399. https:\/\/doi.org\/10.3389\/fonc.2019.01399","journal-title":"Front Oncol"}],"container-title":["Virchows Archiv"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/link.springer.com\/content\/pdf\/10.1007\/s00428-025-04302-3.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/link.springer.com\/article\/10.1007\/s00428-025-04302-3","content-type":"text\/html","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/link.springer.com\/content\/pdf\/10.1007\/s00428-025-04302-3.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,3,5]],"date-time":"2026-03-05T07:10:45Z","timestamp":1772694645000},"score":1,"resource":{"primary":{"URL":"https:\/\/link.springer.com\/10.1007\/s00428-025-04302-3"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2025,10,20]]},"references-count":41,"journal-issue":{"issue":"3","published-print":{"date-parts":[[2026,3]]}},"alternative-id":["4302"],"URL":"https:\/\/doi.org\/10.1007\/s00428-025-04302-3","relation":{},"ISSN":["0945-6317","1432-2307"],"issn-type":[{"value":"0945-6317","type":"print"},{"value":"1432-2307","type":"electronic"}],"subject":[],"published":{"date-parts":[[2025,10,20]]},"assertion":[{"value":"12 April 2025","order":1,"name":"received","label":"Received","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"23 September 2025","order":2,"name":"revised","label":"Revised","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"4 October 2025","order":3,"name":"accepted","label":"Accepted","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"20 October 2025","order":4,"name":"first_online","label":"First Online","group":{"name":"ArticleHistory","label":"Article History"}},{"order":1,"name":"Ethics","group":{"name":"EthicsHeading","label":"Declarations"}},{"value":"All procedures performed in studies involving human participants were in accordance with the ethical standards of the institutional and\/or national research committee and with the 1964 Helsinki declaration and its later amendments or comparable ethical standards. Written informed consent was obtained from all individual participants included in the study. The study protocol was approved by the Ethics Committee of the Faculty of Medicine, Ehime University (Approval No. 2311001).","order":2,"name":"Ethics","group":{"name":"EthicsHeading","label":"Ethical approval"}},{"value":"The authors declare no competing interests.","order":3,"name":"Ethics","group":{"name":"EthicsHeading","label":"Conflict of interest"}},{"value":"The funders had no role in the design of the study, in the collection, analyses, or interpretation of data; in the writing of the manuscript; or in the decision to publish the results.","order":4,"name":"Ethics","group":{"name":"EthicsHeading","label":"Disclosure"}}]}}