{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,10,6]],"date-time":"2026-10-06T03:16:19Z","timestamp":1791256579691,"version":"4.1.0"},"reference-count":64,"publisher":"Springer Science and Business Media LLC","issue":"13","license":[{"start":{"date-parts":[[2022,6,29]],"date-time":"2022-06-29T00:00:00Z","timestamp":1656460800000},"content-version":"tdm","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by-nc-nd\/4.0\/"},{"start":{"date-parts":[[2022,6,29]],"date-time":"2022-06-29T00:00:00Z","timestamp":1656460800000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by-nc-nd\/4.0\/"}],"funder":[{"DOI":"10.13039\/501100017268","name":"Berlin Institute of Health","doi-asserted-by":"publisher","award":["Junior Clinician Scientist Programm"],"award-info":[{"award-number":["Junior Clinician Scientist Programm"]}],"id":[{"id":"10.13039\/501100017268","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100001659","name":"Deutsche Forschungsgemeinschaft","doi-asserted-by":"publisher","award":["Si 1983\u20102\/1"],"award-info":[{"award-number":["Si 1983\u20102\/1"]}],"id":[{"id":"10.13039\/501100001659","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100001659","name":"Deutsche Forschungsgemeinschaft","doi-asserted-by":"publisher","award":["1983\u20103\/1"],"award-info":[{"award-number":["1983\u20103\/1"]}],"id":[{"id":"10.13039\/501100001659","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100001659","name":"Deutsche Forschungsgemeinschaft","doi-asserted-by":"publisher","award":["1983\u20104\/1"],"award-info":[{"award-number":["1983\u20104\/1"]}],"id":[{"id":"10.13039\/501100001659","id-type":"DOI","asserted-by":"publisher"}]}],"content-domain":{"domain":["link.springer.com"],"crossmark-restriction":false},"short-container-title":["EMBO J"],"published-print":{"date-parts":[[2022,7,4]]},"DOI":"10.15252\/embj.2021109996","type":"journal-article","created":{"date-parts":[[2022,6,29]],"date-time":"2022-06-29T11:56:32Z","timestamp":1656503792000},"update-policy":"https:\/\/doi.org\/10.1007\/springer_crossmark_policy","source":"Crossref","is-referenced-by-count":37,"title":["Gastric stem cells promote inflammation and gland remodeling in response to Helicobacter pylori via Rspo3\u2010Lgr4 axis"],"prefix":"10.1038","volume":"41","author":[{"ORCID":"https:\/\/orcid.org\/0000-0003-4261-0416","authenticated-orcid":false,"given":"Jonas","family":"Wizenty","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Stefanie","family":"M\u00fcllerke","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Marina","family":"Kolesnichenko","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-7205-732X","authenticated-orcid":false,"given":"Julian","family":"Heuberger","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Manqiang","family":"Lin","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Anne\u2010Sophie","family":"Fischer","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Hans\u2010Joachim","family":"Mollenkopf","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Hilmar","family":"Berger","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-6206-0226","authenticated-orcid":false,"given":"Frank","family":"Tacke","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-4772-0761","authenticated-orcid":false,"given":"Michael","family":"Sigal","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"297","published-online":{"date-parts":[[2022,6,29]]},"reference":[{"key":"BFEMBJ2021109996_CR1","doi-asserted-by":"crossref","first-page":"387","DOI":"10.1016\/j.stem.2012.05.023","volume":"11","author":"R van Amerongen","year":"2012","unstructured":"van Amerongen R, Bowman AN, Nusse R (2012) Developmental stage and time dictate the fate of Wnt\/beta\u2010catenin\u2010responsive stem cells in the mammary gland. Cell Stem Cell 11: 387\u2013400","journal-title":"Cell Stem Cell"},{"key":"BFEMBJ2021109996_CR2","doi-asserted-by":"crossref","first-page":"306","DOI":"10.1053\/j.gastro.2007.11.009","volume":"134","author":"MR Amieva","year":"2008","unstructured":"Amieva MR, El\u2010Omar EM (2008) Host\u2010bacterial interactions in Helicobacter pylori infection. Gastroenterology 134: 306\u2013323","journal-title":"Gastroenterology"},{"key":"BFEMBJ2021109996_CR3","doi-asserted-by":"crossref","first-page":"1430","DOI":"10.1126\/science.1081919","volume":"300","author":"MR Amieva","year":"2003","unstructured":"Amieva MR, Vogelmann R, Covacci A, Tompkins LS, Nelson WJ, Falkow S (2003) Disruption of the epithelial apical\u2010junctional complex by Helicobacter pylori CagA. Science 300: 1430\u20131434","journal-title":"Science"},{"key":"BFEMBJ2021109996_CR4","doi-asserted-by":"crossref","first-page":"E3282","DOI":"10.1073\/pnas.1509091112","volume":"112","author":"ET Arena","year":"2015","unstructured":"Arena ET, Campbell\u2010Valois FX, Tinevez JY, Nigro G, Sachse M, Moya\u2010Nilges M, Nothelfer K, Marteyn B, Shorte SL, Sansonetti PJ (2015) Bioimage analysis of shigella infection reveals targeting of colonic crypts. Proc Natl Acad Sci U S A 112: E3282\u2013E3290","journal-title":"Proc Natl Acad Sci U S A"},{"key":"BFEMBJ2021109996_CR5","doi-asserted-by":"crossref","first-page":"199","DOI":"10.1053\/j.gastro.2010.06.047","volume":"140","author":"IC Arnold","year":"2011","unstructured":"Arnold IC, Lee JY, Amieva MR, Roers A, Flavell RA, Sparwasser T, M\u00fcller A (2011) Tolerance rather than immunity protects from Helicobacter pylori\u2010induced gastric preneoplasia. Gastroenterology 140: 199\u2013209","journal-title":"Gastroenterology"},{"key":"BFEMBJ2021109996_CR6","doi-asserted-by":"crossref","first-page":"155","DOI":"10.1046\/j.1462-5822.2000.00043.x","volume":"2","author":"S Backert","year":"2000","unstructured":"Backert S, Ziska E, Brinkmann V, Zimny\u2010Arndt U, Fauconnier A, Jungblut PR, Naumann M, Meyer TF (2000) Translocation of the Helicobacter pylori CagA protein in gastric epithelial cells by a type IV secretion apparatus. Cell Microbiol 2: 155\u2013164","journal-title":"Cell Microbiol"},{"key":"BFEMBJ2021109996_CR7","doi-asserted-by":"crossref","first-page":"25","DOI":"10.1016\/j.stem.2009.11.013","volume":"6","author":"N Barker","year":"2010","unstructured":"Barker N, Huch M, Kujala P, van de Wetering M, Snippert HJ, van Es JH, Sato T, Stange DE, Begthel H, van den Born M et\u00a0al (2010) Lgr5(+ve) stem cells drive self\u2010renewal in the stomach and build long\u2010lived gastric units in\u00a0vitro. Cell Stem Cell 6: 25\u201336","journal-title":"Cell Stem Cell"},{"key":"BFEMBJ2021109996_CR8","doi-asserted-by":"crossref","first-page":"1003","DOI":"10.1038\/nature06196","volume":"449","author":"N Barker","year":"2007","unstructured":"Barker N, van Es JH, Kuipers J, Kujala P, van den Born M, Cozijnsen M, Haegebarth A, Korving J, Begthel H, Peters PJ et\u00a0al (2007) Identification of stem cells in small intestine and colon by marker gene Lgr5. Nature 449: 1003\u20131007","journal-title":"Nature"},{"key":"BFEMBJ2021109996_CR9","doi-asserted-by":"crossref","first-page":"5117","DOI":"10.1038\/s41467-020-18857-z","volume":"11","author":"M Bauer","year":"2020","unstructured":"Bauer M, Nascakova Z, Mihai A\u2010I, Cheng PF, Levesque MP, Lampart S, Hurwitz R, Pfannkuch L, Dobrovolna J, Jacobs M et\u00a0al (2020) The ALPK1\/TIFA\/NF\u2010\u03baB axis links a bacterial carcinogen to R\u2010loop\u2010induced replication stress. Nat Commun 11: 5117","journal-title":"Nat Commun"},{"key":"BFEMBJ2021109996_CR10","doi-asserted-by":"crossref","first-page":"289","DOI":"10.1111\/j.2517-6161.1995.tb02031.x","volume":"57","author":"Y Benjamini","year":"1995","unstructured":"Benjamini Y, Hochberg Y (1995) Controlling the false discovery rate: A practical and powerful approach to multiple testing. J R Stat Soc B Methodol 57: 289\u2013300","journal-title":"J R Stat Soc B Methodol"},{"key":"BFEMBJ2021109996_CR11","doi-asserted-by":"crossref","first-page":"1307","DOI":"10.1016\/j.cell.2018.10.008","volume":"175","author":"M Biton","year":"2018","unstructured":"Biton M, Haber AL, Rogel N, Burgin G, Beyaz S, Schnell A, Ashenberg O, Su CW, Smillie C, Shekhar K et\u00a0al (2018) T helper cell cytokines modulate intestinal stem cell renewal and differentiation. Cell 175: 1307\u20131320.e1322","journal-title":"Cell"},{"key":"BFEMBJ2021109996_CR12","doi-asserted-by":"crossref","DOI":"10.1242\/dev.199683","volume":"148","author":"C Brischetto","year":"2021","unstructured":"Brischetto C, Krieger K, Klotz C, Krahn I, Kunz S, Kolesnichenko M, Mucka P, Heuberger J, Scheidereit C, Schmidt\u2010Ullrich R (2021) NF\u2010\u03baB determines Paneth versus goblet cell fate decision in the small intestine. Development 148: dev199683","journal-title":"Development"},{"key":"BFEMBJ2021109996_CR13","doi-asserted-by":"crossref","DOI":"10.1155\/2020\/4069354","volume":"2020","author":"P Brooks","year":"2020","unstructured":"Brooks P, Zur Bruegge T, Boyle EC, Kalies S, Villarreal SN, Liese A, Bleich A, Buettner M (2020) CD14 and ALPK1 affect expression of tight junction components and proinflammatory mediators upon bacterial stimulation in a colonic 3D organoid model. Stem Cells Int 2020: 4069354","journal-title":"Stem Cells Int"},{"key":"BFEMBJ2021109996_CR14","unstructured":"Busslinger GA, Weusten BLA, Bogte A, Begthel H, Brosens LAA, Clevers H (2021a) Gene Expression Omnibus GSE157694 (https:\/\/www.ncbi.nlm.nih.gov\/geo\/query\/acc.cgi?acc=GSE157694). [DATASET]"},{"key":"BFEMBJ2021109996_CR15","doi-asserted-by":"crossref","DOI":"10.1016\/j.celrep.2021.108819","volume":"34","author":"GA Busslinger","year":"2021","unstructured":"Busslinger GA, Weusten BLA, Bogte A, Begthel H, Brosens LAA, Clevers H (2021b) Human gastrointestinal epithelia of the esophagus, stomach, and duodenum resolved at single\u2010cell resolution. Cell Rep 34: 108819","journal-title":"Cell Rep"},{"key":"BFEMBJ2021109996_CR16","doi-asserted-by":"crossref","first-page":"411","DOI":"10.1038\/nbt.4096","volume":"36","author":"A Butler","year":"2018","unstructured":"Butler A, Hoffman P, Smibert P, Papalexi E, Satija R (2018) Integrating single\u2010cell transcriptomic data across different conditions, technologies, and species. Nat Biotechnol 36: 411\u2013420","journal-title":"Nat Biotechnol"},{"key":"BFEMBJ2021109996_CR17","doi-asserted-by":"crossref","DOI":"10.15252\/embj.201899984","volume":"38","author":"D Castillo\u2010Azofeifa","year":"2019","unstructured":"Castillo\u2010Azofeifa D, Fazio EN, Nattiv R, Good HJ, Wald T, Pest MA, de Sauvage FJ, Klein OD, Asfaha S (2019) Atoh1(+) secretory progenitors possess renewal capacity independent of Lgr5(+) cells during colonic regeneration. EMBO J 38: e99984","journal-title":"EMBO J"},{"key":"BFEMBJ2021109996_CR18","doi-asserted-by":"crossref","first-page":"6539","DOI":"10.1093\/emboj\/cdf660","volume":"21","author":"LF Chen","year":"2002","unstructured":"Chen LF, Mu Y, Greene WC (2002) Acetylation of RelA at discrete sites regulates distinct nuclear functions of NF\u2010kappaB. EMBO J 21: 6539\u20136548","journal-title":"EMBO J"},{"key":"BFEMBJ2021109996_CR19","doi-asserted-by":"crossref","first-page":"490","DOI":"10.1038\/ng1031","volume":"32","author":"GA Churchill","year":"2002","unstructured":"Churchill GA (2002) Fundamentals of experimental design for cDNA microarrays. Nat Genet 32: 490\u2013495","journal-title":"Nat Genet"},{"key":"BFEMBJ2021109996_CR20","doi-asserted-by":"crossref","first-page":"S37","DOI":"10.1097\/00000478-199501000-00005","volume":"19","author":"P Correa","year":"1995","unstructured":"Correa P (1995) Helicobacter pylori and gastric carcinogenesis. Am J Surg Pathol 19: S37\u2013S43","journal-title":"Am J Surg Pathol"},{"key":"BFEMBJ2021109996_CR21","doi-asserted-by":"crossref","first-page":"192","DOI":"10.1046\/j.1365-2249.2000.01374.x","volume":"122","author":"M Eck","year":"2000","unstructured":"Eck M, Schmausser B, Scheller K, Toksoy A, Kraus M, Menzel T, M\u00fcller\u2010Hermelink HK, Gillitzer R (2000) CXC chemokines Gro(alpha)\/IL\u20108 and IP\u201010\/MIG in Helicobacter pylori gastritis. Clin Exp Immunol 122: 192\u2013199","journal-title":"Clin Exp Immunol"},{"key":"BFEMBJ2021109996_CR22","doi-asserted-by":"crossref","first-page":"S35","DOI":"10.1016\/S0016-5085(97)80009-1","volume":"113","author":"PB Ernst","year":"1997","unstructured":"Ernst PB, Crowe SE, Reyes VE (1997) How does Helicobacter pylori cause mucosal damage? The inflammatory response. Gastroenterology 113: S35\u2013S42 discussion S50","journal-title":"Gastroenterology"},{"key":"BFEMBJ2021109996_CR23","doi-asserted-by":"crossref","first-page":"1088","DOI":"10.1016\/j.celrep.2019.12.094","volume":"30","author":"D Fast","year":"2020","unstructured":"Fast D, Petkau K, Ferguson M, Shin M, Galenza A, Kostiuk B, Pukatzki S, Foley E (2020) Vibrio cholerae\u2010symbiont interactions inhibit intestinal repair in drosophila. Cell Rep 30: 1088\u20131100.e1085","journal-title":"Cell Rep"},{"key":"BFEMBJ2021109996_CR24","volume":"7","author":"AS Fischer","year":"2019","unstructured":"Fischer AS, Sigal M (2019) The role of Wnt and R\u2010spondin in the stomach during health and disease. Biomedicine 7: 44","journal-title":"Biomedicine"},{"key":"BFEMBJ2021109996_CR25","doi-asserted-by":"crossref","DOI":"10.1371\/journal.pbio.3000231","volume":"17","author":"C Fung","year":"2019","unstructured":"Fung C, Tan S, Nakajima M, Skoog EC, Camarillo\u2010Guerrero LF, Klein JA, Lawley TD, Solnick JV, Fukami T, Amieva MR (2019) High\u2010resolution mapping reveals that microniches in the gastric glands control Helicobacter pylori colonization of the stomach. PLoS Biol 17: e3000231","journal-title":"PLoS Biol"},{"key":"BFEMBJ2021109996_CR26","doi-asserted-by":"crossref","first-page":"E3173","DOI":"10.1073\/pnas.1713510115","volume":"115","author":"G Greicius","year":"2018","unstructured":"Greicius G, Kabiri Z, Sigmundsson K, Liang C, Bunte R, Singh MK, Virshup DM (2018) PDGFRalpha(+) pericryptal stromal cells are the critical source of Wnts and RSPO3 for murine intestinal stem cells in\u00a0vivo. Proc Natl Acad Sci U S A 115: E3173\u2013E3181","journal-title":"Proc Natl Acad Sci U S A"},{"key":"BFEMBJ2021109996_CR27","doi-asserted-by":"crossref","first-page":"27","DOI":"10.1016\/j.immuni.2019.06.025","volume":"51","author":"FR Greten","year":"2019","unstructured":"Greten FR, Grivennikov SI (2019) Inflammation and cancer: Triggers, mechanisms, and consequences. Immunity 51: 27\u201341","journal-title":"Immunity"},{"key":"BFEMBJ2021109996_CR28","doi-asserted-by":"crossref","DOI":"10.26508\/lsa.202101187","volume":"5","author":"J Grinat","year":"2022","unstructured":"Grinat J, Kosel F, Goveas N, Kranz A, Alexopoulou D, Rajewsky K, Sigal M, Stewart AF, Heuberger J (2022) Epigenetic modifier balances Mapk and Wnt signalling in differentiation of goblet and Paneth cells. Life Sci Alliance 5: e202101187","journal-title":"Life Sci Alliance"},{"key":"BFEMBJ2021109996_CR29","doi-asserted-by":"crossref","first-page":"4368","DOI":"10.1038\/s41467-019-12349-5","volume":"10","author":"C Harnack","year":"2019","unstructured":"Harnack C, Berger H, Antanaviciute A, Vidal R, Sauer S, Simmons A, Meyer TF, Sigal M (2019) R\u2010spondin 3 promotes stem cell recovery and epithelial regeneration in the colon. Nat Commun 10: 4368","journal-title":"Nat Commun"},{"key":"BFEMBJ2021109996_CR30","doi-asserted-by":"crossref","first-page":"74","DOI":"10.1186\/1471-2164-15-74","volume":"15","author":"A Herbst","year":"2014","unstructured":"Herbst A, Jurinovic V, Krebs S, Thieme SE, Blum H, G\u00f6ke B, Kolligs FT (2014) Comprehensive analysis of \u03b2\u2010catenin target genes in colorectal carcinoma cell lines with deregulated Wnt\/\u03b2\u2010catenin signaling. BMC Genomics 15: 74","journal-title":"BMC Genomics"},{"key":"BFEMBJ2021109996_CR31","doi-asserted-by":"crossref","DOI":"10.1186\/2045-824X-6-21","volume":"6","author":"BP Herring","year":"2014","unstructured":"Herring BP, Hoggatt AM, Burlak C, Offermanns S (2014) Previously differentiated medial vascular smooth muscle cells contribute to neointima formation following vascular injury. Vasc Cell 6: 21","journal-title":"Vasc Cell"},{"key":"BFEMBJ2021109996_CR32","doi-asserted-by":"crossref","first-page":"1095","DOI":"10.1136\/gutjnl-2016-311606","volume":"66","author":"J Hilkens","year":"2017","unstructured":"Hilkens J, Timmer NC, Boer M, Ikink GJ, Schewe M, Sacchetti A, Koppens MAJ, Song JY, Bakker ERM (2017) RSPO3 expands intestinal stem cell and niche compartments and drives tumorigenesis. Gut 66: 1095\u20131105","journal-title":"Gut"},{"key":"BFEMBJ2021109996_CR33","doi-asserted-by":"crossref","DOI":"10.1128\/mBio.00098-11","volume":"2","author":"MR Howitt","year":"2011","unstructured":"Howitt MR, Lee JY, Lertsethtakarn P, Vogelmann R, Joubert LM, Ottemann KM, Amieva MR (2011) ChePep controls Helicobacter pylori infection of the gastric glands and chemotaxis in the Epsilonproteobacteria. MBio 2: e00098\u201011","journal-title":"MBio"},{"key":"BFEMBJ2021109996_CR34","doi-asserted-by":"crossref","first-page":"1577","DOI":"10.1038\/s41467-022-29176-w","volume":"13","author":"M Kapalczynska","year":"2022","unstructured":"Kapalczynska M, Lin M, Maertzdorf J, Heuberger J, Muellerke S, Zuo X, Vidal R, Shureiqi I, Fischer AS, Sauer S et\u00a0al (2022) BMP feed\u2010forward loop promotes terminal differentiation in gastric glands and is interrupted by H. pylori\u2010driven inflammation. Nat Commun 13: 1577","journal-title":"Nat Commun"},{"key":"BFEMBJ2021109996_CR35","doi-asserted-by":"crossref","DOI":"10.1128\/mBio.01705-16","volume":"7","author":"D Keilberg","year":"2016","unstructured":"Keilberg D, Zavros Y, Shepherd B, Salama NR, Ottemann KM (2016) Spatial and temporal shifts in bacterial biogeography and gland occupation during the development of a chronic infection. mBio 7: e01705\u201016","journal-title":"mBio"},{"key":"BFEMBJ2021109996_CR36","doi-asserted-by":"crossref","DOI":"10.15252\/embj.2019104296","volume":"40","author":"M Kolesnichenko","year":"2021","unstructured":"Kolesnichenko M, Mikuda N, H\u00f6pken UE, K\u00e4rgel E, Uyar B, Tufan AB, Milanovic M, Sun W, Krahn I, Schleich K et\u00a0al (2021) Transcriptional repression of NFKBIA triggers constitutive IKK\u2010 and proteasome\u2010independent p65\/RelA activation in senescence. EMBO J 40: e104296","journal-title":"EMBO J"},{"key":"BFEMBJ2021109996_CR37","doi-asserted-by":"crossref","first-page":"293","DOI":"10.1038\/nature10337","volume":"476","author":"W de Lau","year":"2011","unstructured":"de Lau W, Barker N, Low TY, Koo BK, Li VS, Teunissen H, Kujala P, Haegebarth A, Peters PJ, van de Wetering M et\u00a0al (2011) Lgr5 homologues associate with Wnt receptors and mediate R\u2010spondin signalling. Nature 476: 293\u2013297","journal-title":"Nature"},{"key":"BFEMBJ2021109996_CR38","doi-asserted-by":"crossref","first-page":"1067","DOI":"10.1053\/j.gastro.2021.12.252","volume":"162","author":"D Liabeuf","year":"2022","unstructured":"Liabeuf D, Oshima M, Stange DE, Sigal M (2022) Stem cells, Helicobacter pylori, and mutational landscape: Utility of preclinical models to understand carcinogenesis and to direct Management of Gastric Cancer. Gastroenterology 162: 1067\u20131087","journal-title":"Gastroenterology"},{"key":"BFEMBJ2021109996_CR39","doi-asserted-by":"crossref","first-page":"417","DOI":"10.1016\/j.cels.2015.12.004","volume":"1","author":"A Liberzon","year":"2015","unstructured":"Liberzon A, Birger C, Thorvaldsd\u00f3ttir H, Ghandi M, Mesirov JP, Tamayo P (2015) The molecular signatures database (MSigDB) hallmark gene set collection. Cell Syst 1: 417\u2013425","journal-title":"Cell Syst"},{"key":"BFEMBJ2021109996_CR40","doi-asserted-by":"crossref","first-page":"8794","DOI":"10.1074\/jbc.M112.436204","volume":"288","author":"S Liu","year":"2013","unstructured":"Liu S, Qian Y, Li L, Wei G, Guan Y, Pan H, Guan X, Zhang L, Lu X, Zhao Y et\u00a0al (2013) Lgr4 gene deficiency increases susceptibility and severity of dextran sodium sulfate\u2010induced inflammatory bowel disease in mice. J Biol Chem 288: 8794\u20138803 discussion 8804","journal-title":"J Biol Chem"},{"key":"BFEMBJ2021109996_CR41","doi-asserted-by":"crossref","DOI":"10.15252\/embj.201798658","volume":"37","author":"N Mikuda","year":"2018","unstructured":"Mikuda N, Kolesnichenko M, Beaudette P, Popp O, Uyar B, Sun W, Tufan AB, Perder B, Akalin A, Chen W et\u00a0al (2018) The I\u03baB kinase complex is a regulator of mRNA stability. EMBO J 37: e98658","journal-title":"EMBO J"},{"key":"BFEMBJ2021109996_CR42","doi-asserted-by":"crossref","first-page":"761","DOI":"10.1016\/j.stem.2013.04.006","volume":"12","author":"KB Myant","year":"2013","unstructured":"Myant KB, Cammareri P, McGhee EJ, Ridgway RA, Huels DJ, Cordero JB, Schwitalla S, Kalna G, Ogg EL, Athineos D et\u00a0al (2013) ROS production and NF\u2010\u03baB activation triggered by RAC1 facilitate WNT\u2010driven intestinal stem cell proliferation and colorectal cancer initiation. Cell Stem Cell 12: 761\u2013773","journal-title":"Cell Stem Cell"},{"key":"BFEMBJ2021109996_CR43","doi-asserted-by":"crossref","first-page":"741","DOI":"10.1002\/dvg.22040","volume":"50","author":"S Neufeld","year":"2012","unstructured":"Neufeld S, Rosin JM, Ambasta A, Hui K, Shaneman V, Crowder R, Vickerman L, Cobb J (2012) A conditional allele of Rspo3 reveals redundant function of R\u2010spondins during mouse limb development. Genesis 50: 741\u2013749","journal-title":"Genesis"},{"key":"BFEMBJ2021109996_CR44","doi-asserted-by":"crossref","first-page":"792","DOI":"10.1016\/j.chom.2014.05.003","volume":"15","author":"G Nigro","year":"2014","unstructured":"Nigro G, Rossi R, Commere PH, Jay P, Sansonetti PJ (2014) The cytosolic bacterial peptidoglycan sensor Nod2 affords stem cell protection and links microbes to gut epithelial regeneration. Cell Host Microbe 15: 792\u2013798","journal-title":"Cell Host Microbe"},{"key":"BFEMBJ2021109996_CR45","doi-asserted-by":"crossref","first-page":"297","DOI":"10.1136\/gut.40.3.297","volume":"40","author":"J Parsonnet","year":"1997","unstructured":"Parsonnet J, Friedman GD, Orentreich N, Vogelman H (1997) Risk for gastric cancer in people with CagA positive or CagA negative Helicobacter pylori infection. Gut 40: 297\u2013301","journal-title":"Gut"},{"key":"BFEMBJ2021109996_CR46","doi-asserted-by":"crossref","DOI":"10.1093\/nar\/29.9.e45","volume":"29","author":"MW Pfaffl","year":"2001","unstructured":"Pfaffl MW (2001) A new mathematical model for relative quantification in real\u2010time RT\u2010PCR. Nucleic Acids Res 29: e45","journal-title":"Nucleic Acids Res"},{"key":"BFEMBJ2021109996_CR47","doi-asserted-by":"crossref","first-page":"9087","DOI":"10.1096\/fj.201802555R","volume":"33","author":"L Pfannkuch","year":"2019","unstructured":"Pfannkuch L, Hurwitz R, Traulsen J, Sigulla J, Poeschke M, Matzner L, Kosma P, Schmid M, Meyer TF (2019) ADP heptose, a novel pathogen\u2010associated molecular pattern identified in Helicobacter pylori. FASEB J 33: 9087\u20139099","journal-title":"FASEB J"},{"key":"BFEMBJ2021109996_CR48","doi-asserted-by":"crossref","first-page":"467","DOI":"10.1038\/ncb3337","volume":"18","author":"L Planas\u2010Paz","year":"2016","unstructured":"Planas\u2010Paz L, Orsini V, Boulter L, Calabrese D, Pikiolek M, Nigsch F, Xie Y, Roma G, Donovan A, Marti P et\u00a0al (2016) The RSPO\u2010LGR4\/5\u2010ZNRF3\/RNF43 module controls liver zonation and size. Nat Cell Biol 18: 467\u2013479","journal-title":"Nat Cell Biol"},{"key":"BFEMBJ2021109996_CR49","doi-asserted-by":"crossref","first-page":"3843","DOI":"10.1242\/dev.128.19.3843","volume":"128","author":"R Schmidt\u2010Ullrich","year":"2001","unstructured":"Schmidt\u2010Ullrich R, Aebischer T, H\u00fclsken J, Birchmeier W, Klemm U, Scheidereit C (2001) Requirement of NF\u2010kappaB\/Rel for the development of hair follicles and other epidermal appendices. Development 128: 3843\u20133853","journal-title":"Development"},{"key":"BFEMBJ2021109996_CR50","doi-asserted-by":"crossref","first-page":"25","DOI":"10.1016\/j.cell.2012.12.012","volume":"152","author":"S Schwitalla","year":"2013","unstructured":"Schwitalla S, Fingerle AA, Cammareri P, Nebelsiek T, G\u00f6ktuna SI, Ziegler PK, Canli O, Heijmans J, Huels DJ, Moreaux G et\u00a0al (2013) Intestinal tumorigenesis initiated by dedifferentiation and acquisition of stem\u2010cell\u2010like properties. Cell 152: 25\u201338","journal-title":"Cell"},{"key":"BFEMBJ2021109996_CR51","doi-asserted-by":"crossref","first-page":"3864","DOI":"10.1093\/hmg\/ddn286","volume":"17","author":"S Segditsas","year":"2008","unstructured":"Segditsas S, Sieber O, Deheragoda M, East P, Rowan A, Jeffery R, Nye E, Clark S, Spencer\u2010Dene B, Stamp G et\u00a0al (2008) Putative direct and indirect Wnt targets identified through consistent gene expression changes in APC\u2010mutant intestinal adenomas from humans and mice. Hum Mol Genet 17: 3864\u20133875","journal-title":"Hum Mol Genet"},{"key":"BFEMBJ2021109996_CR52","doi-asserted-by":"publisher","DOI":"10.1101\/060012","author":"AA Sergushichev","year":"2016","unstructured":"Sergushichev AA (2016) An algorithm for fast preranked gene set enrichment analysis using cumulative statistic calculation. bioRxiv https:\/\/doi.org\/10.1101\/060012 [PREPRINT]","journal-title":"bioRxiv"},{"key":"BFEMBJ2021109996_CR53","doi-asserted-by":"crossref","first-page":"S2","DOI":"10.1136\/gut.43.2008.S2","volume":"43","author":"T Shimoyama","year":"1998","unstructured":"Shimoyama T, Crabtree JE (1998) Bacterial factors and immune pathogenesis in Helicobacter pylori infection. Gut 43: S2\u2013S5","journal-title":"Gut"},{"key":"BFEMBJ2021109996_CR54","doi-asserted-by":"crossref","first-page":"451","DOI":"10.1038\/nature23642","volume":"548","author":"M Sigal","year":"2017","unstructured":"Sigal M, Logan CY, Kapalczynska M, Mollenkopf HJ, Berger H, Wiedenmann B, Nusse R, Amieva MR, Meyer TF (2017) Stromal R\u2010spondin orchestrates gastric epithelial stem cells and gland homeostasis. Nature 548: 451\u2013455","journal-title":"Nature"},{"key":"BFEMBJ2021109996_CR55","doi-asserted-by":"crossref","first-page":"812","DOI":"10.1038\/s41556-019-0339-9","volume":"21","author":"M Sigal","year":"2019","unstructured":"Sigal M, Reines MDM, Mullerke S, Fischer C, Kapalczynska M, Berger H, Bakker ERM, Mollenkopf HJ, Rothenberg ME, Wiedenmann B et\u00a0al (2019) R\u2010spondin\u20103 induces secretory, antimicrobial Lgr5(+) cells in the stomach. Nat Cell Biol 21: 812\u2013823","journal-title":"Nat Cell Biol"},{"key":"BFEMBJ2021109996_CR56","doi-asserted-by":"crossref","first-page":"1392","DOI":"10.1053\/j.gastro.2015.02.049","volume":"148","author":"M Sigal","year":"2015","unstructured":"Sigal M, Rothenberg ME, Logan CY, Lee JY, Honaker RW, Cooper RL, Passarelli B, Camorlinga M, Bouley DM, Alvarez G et\u00a0al (2015) Helicobacter pylori activates and expands Lgr5(+) stem cells through direct colonization of the gastric glands. Gastroenterology 148: 1392\u20131404.e1321","journal-title":"Gastroenterology"},{"key":"BFEMBJ2021109996_CR57","doi-asserted-by":"crossref","first-page":"15545","DOI":"10.1073\/pnas.0506580102","volume":"102","author":"A Subramanian","year":"2005","unstructured":"Subramanian A, Tamayo P, Mootha VK, Mukherjee S, Ebert BL, Gillette MA, Paulovich A, Pomeroy SL, Golub TR, Lander ES et\u00a0al (2005) Gene set enrichment analysis: a knowledge\u2010based approach for interpreting genome\u2010wide expression profiles. Proc Natl Acad Sci U S A 102: 15545\u201315550","journal-title":"Proc Natl Acad Sci U S A"},{"key":"BFEMBJ2021109996_CR58","doi-asserted-by":"crossref","DOI":"10.1371\/journal.pone.0021800","volume":"6","author":"F Supek","year":"2011","unstructured":"Supek F, Bo\u0161njak M, \u0160kunca N, \u0160muc T (2011) REVIGO summarizes and visualizes long lists of gene ontology terms. PLoS One 6: e21800","journal-title":"PLoS One"},{"key":"BFEMBJ2021109996_CR59","doi-asserted-by":"crossref","first-page":"437","DOI":"10.1038\/s41586-020-1973-x","volume":"578","author":"SH Tan","year":"2020","unstructured":"Tan SH, Swathi Y, Tan S, Goh J, Seishima R, Murakami K, Oshima M, Tsuji T, Phuah P, Tan LT et\u00a0al (2020) AQP5 enriches for stem cells and cancer origins in the distal stomach. Nature 578: 437\u2013443","journal-title":"Nature"},{"key":"BFEMBJ2021109996_CR60","doi-asserted-by":"crossref","first-page":"255","DOI":"10.1038\/nature10408","volume":"478","author":"H Tian","year":"2011","unstructured":"Tian H, Biehs B, Warming S, Leong KG, Rangell L, Klein OD, de Sauvage FJ (2011) A reserve stem cell population in small intestine renders Lgr5\u2010positive cells dispensable. Nature 478: 255\u2013259","journal-title":"Nature"},{"key":"BFEMBJ2021109996_CR61","first-page":"1512","volume":"143","author":"P Tomann","year":"2016","unstructured":"Tomann P, Paus R, Millar SE, Scheidereit C, Schmidt\u2010Ullrich R (2016) Lhx2 is a direct NF\u2010\u03baB target gene that promotes primary hair follicle placode down\u2010growth. Development 143: 1512\u20131522","journal-title":"Development"},{"key":"BFEMBJ2021109996_CR62","doi-asserted-by":"crossref","first-page":"1030","DOI":"10.1038\/nm1480","volume":"12","author":"C Wunder","year":"2006","unstructured":"Wunder C, Churin Y, Winau F, Warnecke D, Vieth M, Lindner B, Z\u00e4hringer U, Mollenkopf HJ, Heinz E, Meyer TF (2006) Cholesterol glucosylation promotes immune evasion by Helicobacter pylori. Nat Med 12: 1030\u20131038","journal-title":"Nat Med"},{"key":"BFEMBJ2021109996_CR63","doi-asserted-by":"crossref","first-page":"238","DOI":"10.1038\/nature22313","volume":"545","author":"KS Yan","year":"2017","unstructured":"Yan KS, Janda CY, Chang J, Zheng GXY, Larkin KA, Luca VC, Chia LA, Mah AT, Han A, Terry JM et\u00a0al (2017) Non\u2010equivalence of Wnt and R\u2010spondin ligands during Lgr5(+) intestinal stem\u2010cell self\u2010renewal. Nature 545: 238\u2013242","journal-title":"Nature"},{"key":"BFEMBJ2021109996_CR64","doi-asserted-by":"crossref","first-page":"2384","DOI":"10.1016\/j.celrep.2017.08.039","volume":"20","author":"S Zimmermann","year":"2017","unstructured":"Zimmermann S, Pfannkuch L, Al\u2010Zeer MA, Bartfeld S, Koch M, Liu J, Rechner C, Soerensen M, Sokolova O, Zamyatina A et\u00a0al (2017) ALPK1\u2010 and TIFA\u2010dependent innate immune response triggered by the Helicobacter pylori type IV secretion system. Cell Rep 20: 2384\u20132395","journal-title":"Cell Rep"}],"container-title":["The EMBO Journal"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/link.springer.com\/content\/pdf\/10.15252\/embj.2021109996.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/onlinelibrary.wiley.com\/doi\/full-xml\/10.15252\/embj.2021109996","content-type":"application\/xml","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/link.springer.com\/article\/10.15252\/embj.2021109996\/fulltext.html","content-type":"text\/html","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/www.embopress.org\/doi\/pdf\/10.15252\/embj.2021109996","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"},{"URL":"https:\/\/link.springer.com\/content\/pdf\/10.15252\/embj.2021109996.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,5,20]],"date-time":"2026-05-20T20:21:07Z","timestamp":1779308467000},"score":1,"resource":{"primary":{"URL":"https:\/\/link.springer.com\/10.15252\/embj.2021109996"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2022,6,29]]},"references-count":64,"journal-issue":{"issue":"13","published-print":{"date-parts":[[2022,7,4]]}},"alternative-id":["BFEMBJ2021109996"],"URL":"https:\/\/doi.org\/10.15252\/embj.2021109996","relation":{},"ISSN":["0261-4189","1460-2075"],"issn-type":[{"value":"0261-4189","type":"print"},{"value":"1460-2075","type":"electronic"}],"subject":[],"published":{"date-parts":[[2022,6,29]]},"assertion":[{"value":"22 October 2021","order":1,"name":"received","label":"Received","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"17 May 2022","order":2,"name":"accepted","label":"Accepted","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"29 June 2022","order":3,"name":"first_online","label":"First Online","group":{"name":"ArticleHistory","label":"Article History"}}],"article-number":"EMBJ2021109996"}}