{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,10,29]],"date-time":"2026-10-29T04:23:16Z","timestamp":1793247796305,"version":"4.5.0"},"reference-count":40,"publisher":"Springer Science and Business Media LLC","issue":"1","license":[{"start":{"date-parts":[[2026,1,9]],"date-time":"2026-01-09T00:00:00Z","timestamp":1767916800000},"content-version":"tdm","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0"},{"start":{"date-parts":[[2026,1,9]],"date-time":"2026-01-09T00:00:00Z","timestamp":1767916800000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0"}],"funder":[{"DOI":"10.13039\/501100001809","name":"National Natural Science Foundation of China","doi-asserted-by":"publisher","award":["82100586"],"award-info":[{"award-number":["82100586"]}],"id":[{"id":"10.13039\/501100001809","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100001809","name":"National Natural Science Foundation of China","doi-asserted-by":"publisher","award":["32301201"],"award-info":[{"award-number":["32301201"]}],"id":[{"id":"10.13039\/501100001809","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100001809","name":"National Natural Science Foundation of China","doi-asserted-by":"publisher","award":["U24A20788"],"award-info":[{"award-number":["U24A20788"]}],"id":[{"id":"10.13039\/501100001809","id-type":"DOI","asserted-by":"publisher"}]}],"content-domain":{"domain":["link.springer.com"],"crossmark-restriction":false},"short-container-title":["Cell Death Dis"],"abstract":"<jats:title>Abstract<\/jats:title>\n                  <jats:p>The impaired repair of intestinal mucosal damage is an important pathological feature of ulcerative colitis (UC). The critical role of intestinal epithelial cells (IECs) proliferation and migration in the repair of damaged mucosal epithelium has been well established. However, the molecular circuitry that decodes IECs sense intestinal mucosal damage signals to initiate and drive repair program remains elusive. Here, we identify a tryptophan (Trp) metabolic gatekeeping mechanism wherein G protein-coupled receptor 35 (GPR35) senses intestinal mucosal damage through monitoring Trp-kynurenine (KYN)-kynurenic acid (KA) axis metabolism with a unique \u201csandwich\u201d structural binding mode. We delineate a GPR35-Kruppel-like factor 5 (KLF5) regulatory circuit in which KLF5 serves as the central effector, translating GPR35-mediated KA sensing into repair programming through PI3K-AKT-mTOR signaling cascade. This circuitry precisely orchestrates IECs proliferation and migration by regulating KLF5-dependent gene expression networks that essential for restoring damaged mucosa. Once this metabolic gatekeeping system is disrupted, either through impaired GPR35-mediated KA sensing or defective signal transduction, compromises damage signal decoding, leading to inadequate repair responses. Such dysregulation results in delayed intestinal mucosal repair and exacerbation of tissue damage. Our findings highlight GPR35 as a surveillant of abnormal Trp-KYN-KA axis metabolism, enabling IECs to detect intestinal mucosal damage and orchestrate repair through KLF5 response. This provides important implications for UC prevention and treatment by targeting GPR35-KLF5 circuit.<\/jats:p>","DOI":"10.1038\/s41419-025-08237-0","type":"journal-article","created":{"date-parts":[[2026,1,9]],"date-time":"2026-01-09T14:43:03Z","timestamp":1767969783000},"update-policy":"https:\/\/doi.org\/10.1007\/springer_crossmark_policy","source":"Crossref","is-referenced-by-count":7,"title":["Tryptophan metabolic gatekeeping in epithelial repair: GPR35-KLF5 circuitry decodes mucosal damage signals for repair programming"],"prefix":"10.1038","volume":"17","author":[{"given":"Biao","family":"Xie","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-1175-8410","authenticated-orcid":false,"given":"Meimei","family":"Wang","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Yaping","family":"Xiao","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Xin","family":"Zhang","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Meng","family":"Liu","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Jie","family":"Miao","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Yunfei","family":"Mo","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Hongxin","family":"Liu","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Jihui","family":"Wang","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Fengguo","family":"Xu","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-3940-6177","authenticated-orcid":false,"given":"Di","family":"Wang","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"297","published-online":{"date-parts":[[2026,1,9]]},"reference":[{"key":"8237_CR1","doi-asserted-by":"publisher","first-page":"571","DOI":"10.1016\/S0140-6736(23)00966-2","volume":"402","author":"C Le Berre","year":"2023","unstructured":"Le Berre C, Honap S, Peyrin-Biroulet L. Ulcerative colitis. Lancet. 2023;402:571\u201384.","journal-title":"Lancet"},{"key":"8237_CR2","doi-asserted-by":"publisher","first-page":"74","DOI":"10.1038\/s41572-020-0205-x","volume":"6","author":"T Kobayashi","year":"2020","unstructured":"Kobayashi T, Siegmund B, Le Berre C, Wei SC, Ferrante M, Shen B, et al. Ulcerative colitis. Nat Rev Dis Prim. 2020;6:74.","journal-title":"Nat Rev Dis Prim"},{"key":"8237_CR3","doi-asserted-by":"publisher","first-page":"3053","DOI":"10.1007\/s11033-020-05318-5","volume":"47","author":"N Younis","year":"2020","unstructured":"Younis N, Zarif R, Mahfouz R. Inflammatory bowel disease: between genetics and microbiota. Mol Biol Rep. 2020;47:3053\u201363.","journal-title":"Mol Biol Rep"},{"key":"8237_CR4","doi-asserted-by":"publisher","first-page":"95","DOI":"10.1038\/s41418-020-00665-z","volume":"28","author":"J Sprangers","year":"2020","unstructured":"Sprangers J, Zaalberg IC, Maurice MM. Organoid-based modeling of intestinal development, regeneration, and repair. Cell Death Differ. 2020;28:95\u2013107.","journal-title":"Cell Death Differ"},{"key":"8237_CR5","doi-asserted-by":"publisher","first-page":"39","DOI":"10.1038\/s41580-020-0278-0","volume":"22","author":"J Beumer","year":"2020","unstructured":"Beumer J, Clevers H. Cell fate specification and differentiation in the adult mammalian intestine. Nat Rev Mol Cell Biol. 2020;22:39\u201353.","journal-title":"Nat Rev Mol Cell Biol"},{"key":"8237_CR6","doi-asserted-by":"publisher","first-page":"543","DOI":"10.1038\/s41575-020-0326-4","volume":"17","author":"JV Patankar","year":"2020","unstructured":"Patankar JV, Becker C. Cell death in the gut epithelium and implications for chronic inflammation. Nat Rev Gastroenterol Hepatol. 2020;17:543\u201356.","journal-title":"Nat Rev Gastroenterol Hepatol"},{"key":"8237_CR7","doi-asserted-by":"publisher","first-page":"263","DOI":"10.1038\/s41575-019-0261-4","volume":"17","author":"JF Burgue\u00f1o","year":"2020","unstructured":"Burgue\u00f1o JF, Abreu MT. Epithelial Toll-like receptors and their role in gut homeostasis and disease. Nat Rev Gastroenterol Hepatol. 2020;17:263\u201378.","journal-title":"Nat Rev Gastroenterol Hepatol"},{"key":"8237_CR8","doi-asserted-by":"publisher","first-page":"593","DOI":"10.1146\/annurev-physiol-022516-034356","volume":"79","author":"KM Vannella","year":"2017","unstructured":"Vannella KM, Wynn TA. Mechanisms of organ injury and repair by macrophages. Annu Rev Physiol. 2017;79:593\u2013617.","journal-title":"Annu Rev Physiol"},{"key":"8237_CR9","doi-asserted-by":"publisher","first-page":"355","DOI":"10.1038\/s41580-022-00572-w","volume":"24","author":"SA Baker","year":"2023","unstructured":"Baker SA, Rutter J. Metabolites as signalling molecules. Nat Rev Mol Cell Biol. 2023;24:355\u201374.","journal-title":"Nat Rev Mol Cell Biol"},{"key":"8237_CR10","doi-asserted-by":"publisher","first-page":"184","DOI":"10.1038\/s41577-019-0125-8","volume":"19","author":"V Rothhammer","year":"2019","unstructured":"Rothhammer V, Quintana FJ. The aryl hydrocarbon receptor: an environmental sensor integrating immune responses in health and disease. Nat Rev Immunol. 2019;19:184\u201397.","journal-title":"Nat Rev Immunol"},{"key":"8237_CR11","doi-asserted-by":"publisher","first-page":"9614","DOI":"10.3390\/ijms21249614","volume":"21","author":"IA Murray","year":"2020","unstructured":"Murray IA, Perdew GH. How Ah receptor ligand specificity became important in understanding its physiological function. Int J Mol Sci. 2020;21:9614.","journal-title":"Int J Mol Sci"},{"key":"8237_CR12","doi-asserted-by":"publisher","first-page":"450","DOI":"10.3390\/cells8050450","volume":"8","author":"H Melhem","year":"2019","unstructured":"Melhem H, Kaya B, Ayata CK, Hruz P, Niess JH. Metabolite-Sensing G protein-coupled receptors connect the diet-microbiota-metabolites axis to inflammatory bowel disease. Cells. 2019;8:450.","journal-title":"Cells"},{"key":"8237_CR13","doi-asserted-by":"publisher","first-page":"371","DOI":"10.1146\/annurev-immunol-051116-052235","volume":"35","author":"JK Tan","year":"2017","unstructured":"Tan JK, McKenzie C, Marino E, Macia L, Mackay CR. Metabolite-sensing G protein-coupled receptors-facilitators of diet-related immune regulation. Annu Rev Immunol. 2017;35:371\u2013402.","journal-title":"Annu Rev Immunol"},{"key":"8237_CR14","doi-asserted-by":"publisher","first-page":"263","DOI":"10.1016\/j.tips.2023.03.001","volume":"44","author":"G Milligan","year":"2023","unstructured":"Milligan G. GPR35: from enigma to therapeutic target. Trends Pharm Sci. 2023;44:263\u201373.","journal-title":"Trends Pharm Sci"},{"key":"8237_CR15","doi-asserted-by":"publisher","DOI":"10.3389\/fimmu.2021.717392","volume":"12","author":"B Kaya","year":"2021","unstructured":"Kaya B, Melhem H, Niess JH. GPR35 in intestinal diseases: from risk gene to function. Front Immunol. 2021;12:717392.","journal-title":"Front Immunol"},{"key":"8237_CR16","doi-asserted-by":"publisher","first-page":"763","DOI":"10.1016\/j.apsb.2020.07.017","volume":"11","author":"D Wang","year":"2021","unstructured":"Wang D, Li D, Zhang Y, Chen J, Zhang Y, Liao C, et al. Functional metabolomics reveal the role of AHR\/GPR35 mediated kynurenic acid gradient sensing in chemotherapy-induced intestinal damage. Acta Pharm Sin B. 2021;11:763\u201380.","journal-title":"Acta Pharm Sin B"},{"key":"8237_CR17","doi-asserted-by":"publisher","DOI":"10.1177\/17562848221138160","volume":"15","author":"N Aslam","year":"2022","unstructured":"Aslam N, Lo SW, Sikafi R, Barnes T, Segal J, Smith PJ, et al. A review of the therapeutic management of ulcerative colitis. Therap Adv Gastroenterol. 2022;15:17562848221138160.","journal-title":"Therap Adv Gastroenterol"},{"key":"8237_CR18","doi-asserted-by":"publisher","first-page":"1157","DOI":"10.3748\/wjg.v29.i7.1157","volume":"29","author":"ML Otte","year":"2023","unstructured":"Otte ML, Lama Tamang R, Papapanagiotou J, Ahmad R, Dhawan P, Singh AB. Mucosal healing and inflammatory bowel disease: therapeutic implications and new targets. World J Gastroenterol. 2023;29:1157\u201372.","journal-title":"World J Gastroenterol"},{"key":"8237_CR19","doi-asserted-by":"publisher","first-page":"88","DOI":"10.1038\/s41392-024-01803-6","volume":"9","author":"M Zhang","year":"2024","unstructured":"Zhang M, Chen T, Lu X, Lan X, Chen Z, Lu S. G protein-coupled receptors (GPCRs): advances in structures, mechanisms, and drug discovery. Signal Transduct Target Ther. 2024;9:88.","journal-title":"Signal Transduct Target Ther"},{"key":"8237_CR20","doi-asserted-by":"publisher","first-page":"621","DOI":"10.1126\/science.abm1638","volume":"377","author":"GA Wyant","year":"2022","unstructured":"Wyant GA, Yu W, Doulamis IP, Nomoto RS, Saeed MY, Duignan T, et al. Mitochondrial remodeling and ischemic protection by G protein\u2013coupled receptor 35 agonists. Science. 2022;377:621\u201329.","journal-title":"Science"},{"key":"8237_CR21","doi-asserted-by":"publisher","first-page":"730","DOI":"10.1177\/1087057105277968","volume":"10","author":"RIW Osmond","year":"2005","unstructured":"Osmond RIW, Sheehan A, Borowicz R, Barnett E, Harvey G, Turner C, et al. GPCR Screening via ERK 1\/2: a novel platform for screening g protein\u2013coupled receptors. SLAS Discov. 2005;10:730\u201337.","journal-title":"SLAS Discov"},{"key":"8237_CR22","doi-asserted-by":"publisher","first-page":"105218","DOI":"10.1016\/j.jbc.2023.105218","volume":"299","author":"A Ganguly","year":"2023","unstructured":"Ganguly A, Quon T, Jenkins L, Joseph B, Al-awar R, Chevigne A, et al. G protein\u2013receptor kinases 5\/6 are the key regulators of G protein\u2013coupled receptor 35\u2013arrestin interactions. J Biol Chem. 2023;299:105218.","journal-title":"J Biol Chem"},{"key":"8237_CR23","doi-asserted-by":"publisher","first-page":"2211","DOI":"10.1016\/j.cgh.2022.06.030","volume":"21","author":"MJ Buie","year":"2023","unstructured":"Buie MJ, Quan J, Windsor JW, Coward S, Hansen TM, King JA, et al. Global hospitalization trends for Crohn\u2019s disease and ulcerative colitis in the 21st Century: a systematic review with temporal analyses. Clin Gastroenterol Hepatol. 2023;21:2211\u201321.","journal-title":"Clin Gastroenterol Hepatol"},{"key":"8237_CR24","doi-asserted-by":"publisher","first-page":"5551","DOI":"10.3390\/jcm10235551","volume":"10","author":"P Wetwittayakhlang","year":"2021","unstructured":"Wetwittayakhlang P, Lontai L, Gonczi L, Golovics PA, Hahn GD, Bessissow T, et al. Treatment targets in ulcerative colitis: is it time for all in, including histology?. J Clin Med. 2021;10:5551.","journal-title":"J Clin Med"},{"key":"8237_CR25","doi-asserted-by":"publisher","DOI":"10.1126\/science.add6220","volume":"380","author":"C Mao","year":"2023","unstructured":"Mao C, Xiao P, Tao XN, Qin J, He QT, Zhang C, et al. Unsaturated bond recognition leads to biased signal in a fatty acid receptor. Science. 2023;380:eadd6220.","journal-title":"Science"},{"key":"8237_CR26","doi-asserted-by":"publisher","first-page":"815","DOI":"10.1016\/j.cell.2022.01.010","volume":"185","author":"M De Giovanni","year":"2022","unstructured":"De Giovanni M, Tam H, Valet C, Xu Y, Looney MR, Cyster JG. GPR35 promotes neutrophil recruitment in response to serotonin metabolite 5-HIAA. Cell. 2022;185:815\u201330.","journal-title":"Cell"},{"key":"8237_CR27","doi-asserted-by":"publisher","DOI":"10.1038\/s41467-020-14737-8","volume":"11","author":"B Zhang","year":"2020","unstructured":"Zhang B, Ci X, Tao R, Ni JJ, Xuan X, King JL, et al. Klf5 acetylation regulates luminal differentiation of basal progenitors in prostate development and regeneration. Nat Commun. 2020;11:997.","journal-title":"Nat Commun"},{"key":"8237_CR28","doi-asserted-by":"publisher","first-page":"79","DOI":"10.1093\/jmcb\/mjaa080","volume":"13","author":"Z Xie","year":"2021","unstructured":"Xie Z, Chen J, Wang C, Zhang J, Wu Y, Yan X. Current knowledge of Kruppel-like factor 5 and vascular remodeling: providing insights for therapeutic strategies. J Mol Cell Biol. 2021;13:79\u201390.","journal-title":"J Mol Cell Biol"},{"key":"8237_CR29","doi-asserted-by":"publisher","first-page":"587","DOI":"10.1016\/j.jcmgh.2019.11.009","volume":"9","author":"CK Kim","year":"2020","unstructured":"Kim CK, Saxena M, Maharjan K, Song JJ, Shroyer KR, Bialkowska AB, et al. Kruppel-like Factor 5 regulates stemness, lineage specification, and regeneration of intestinal epithelial stem cells. Cell Mol Gastroenterol Hepatol. 2020;9:587\u2013609.","journal-title":"Cell Mol Gastroenterol Hepatol"},{"key":"8237_CR30","doi-asserted-by":"publisher","first-page":"G478","DOI":"10.1152\/ajpgi.00172.2017","volume":"313","author":"Y Liu","year":"2017","unstructured":"Liu Y, Chidgey M, Yang VW, Bialkowska AB. Kr\u00fcppel-like factor 5 is essential for maintenance of barrier function in mouse colon. Am J Physiol Gastrointest Liver Physiol. 2017;313:G478\u2013G91.","journal-title":"Am J Physiol Gastrointest Liver Physiol"},{"key":"8237_CR31","doi-asserted-by":"publisher","first-page":"1139","DOI":"10.1161\/CIRCULATIONAHA.120.047420","volume":"143","author":"M Hoffman","year":"2021","unstructured":"Hoffman M, Palioura D, Kyriazis ID, Cimini M, Badolia R, Rajan S, et al. Cardiomyocyte Kruppel-Like Factor 5 promotes De Novo ceramide biosynthesis and contributes to eccentric remodeling in ischemic cardiomyopathy. Circulation. 2021;143:1139\u201356.","journal-title":"Circulation"},{"key":"8237_CR32","doi-asserted-by":"publisher","first-page":"549","DOI":"10.1002\/bies.20581","volume":"29","author":"BB McConnell","year":"2007","unstructured":"McConnell BB, Ghaleb AM, Nandan MO, Yang VW. The diverse functions of Kr\u00fcppel-like factors 4 and 5 in epithelial biology and pathobiology. Bioessays. 2007;29:549\u201357.","journal-title":"Bioessays"},{"key":"8237_CR33","doi-asserted-by":"publisher","first-page":"6304","DOI":"10.1039\/D2RA07005G","volume":"13","author":"M Chang","year":"2023","unstructured":"Chang M. Matrix metalloproteinase profiling and their roles in disease. RSC Adv. 2023;13:6304\u201316.","journal-title":"RSC Adv"},{"key":"8237_CR34","doi-asserted-by":"publisher","first-page":"61","DOI":"10.1038\/s41392-024-01764-w","volume":"9","author":"Z Deng","year":"2024","unstructured":"Deng Z, Fan T, Xiao C, Tian H, Zheng Y, Li C, et al. TGF-beta signaling in health, disease, and therapeutics. Signal Transduct Target Ther. 2024;9:61.","journal-title":"Signal Transduct Target Ther"},{"key":"8237_CR35","doi-asserted-by":"publisher","DOI":"10.1016\/j.cellsig.2022.110470","volume":"100","author":"Z Lin","year":"2022","unstructured":"Lin Z, Wan X, Zhang T, Huo H, Zhang X, Li K, et al. Trefoil factor 3: new highlights in chronic kidney disease research. Cell Signal. 2022;100:110470.","journal-title":"Cell Signal"},{"key":"8237_CR36","doi-asserted-by":"crossref","unstructured":"Yun CC, Han Y, McConnell B. Lysophosphatidic acid signaling in the GI system. Cell Mol Gastroenterol Hepatol. 2024;18:101398.","DOI":"10.1016\/j.jcmgh.2024.101398"},{"key":"8237_CR37","doi-asserted-by":"publisher","first-page":"781","DOI":"10.3390\/cells13090781","volume":"13","author":"T Zhao","year":"2024","unstructured":"Zhao T, Fan J, Abu-Zaid A, Burley SK, Zheng XFS. Nuclear mTOR signaling orchestrates transcriptional programs underlying cellular growth and metabolism. Cells. 2024;13:781.","journal-title":"Cells"},{"key":"8237_CR38","doi-asserted-by":"publisher","first-page":"3049","DOI":"10.1016\/j.apsb.2024.04.017","volume":"14","author":"J Yang","year":"2024","unstructured":"Yang J, Xiao Y, Zhao N, Pei G, Sun Y, Sun X, et al. PIM1-HDAC2 axis modulates intestinal homeostasis through epigenetic modification. Acta Pharm Sin B. 2024;14:3049\u201367.","journal-title":"Acta Pharm Sin B"},{"key":"8237_CR39","doi-asserted-by":"publisher","first-page":"31","DOI":"10.1007\/978-1-4939-1050-2_3","volume":"1180","author":"AT Feldman","year":"2014","unstructured":"Feldman AT, Wolfe D. Tissue processing and hematoxylin and eosin staining. Methods Mol Biol. 2014;1180:31\u201343.","journal-title":"Methods Mol Biol"},{"key":"8237_CR40","doi-asserted-by":"publisher","first-page":"85","DOI":"10.1007\/978-1-0716-2675-7_7","volume":"2566","author":"D Sanchez-Porras","year":"2023","unstructured":"Sanchez-Porras D, Varas J, Godoy-Guzman C, Bermejo-Casares F, San Martin S, Carriel V. Histochemical and immunohistochemical methods for the identification of proteoglycans. Methods Mol Biol. 2023;2566:85\u201398.","journal-title":"Methods Mol Biol"}],"container-title":["Cell Death &amp; Disease"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.nature.com\/articles\/s41419-025-08237-0.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/www.nature.com\/articles\/s41419-025-08237-0","content-type":"text\/html","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/www.nature.com\/articles\/s41419-025-08237-0.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,6,8]],"date-time":"2026-06-08T09:55:13Z","timestamp":1780912513000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.nature.com\/articles\/s41419-025-08237-0"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,1,9]]},"references-count":40,"journal-issue":{"issue":"1","published-online":{"date-parts":[[2026,12]]}},"alternative-id":["8237"],"URL":"https:\/\/doi.org\/10.1038\/s41419-025-08237-0","relation":{},"ISSN":["2041-4889"],"issn-type":[{"value":"2041-4889","type":"electronic"}],"subject":[],"published":{"date-parts":[[2026,1,9]]},"assertion":[{"value":"4 August 2025","order":1,"name":"received","label":"Received","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"16 October 2025","order":2,"name":"revised","label":"Revised","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"3 November 2025","order":3,"name":"accepted","label":"Accepted","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"9 January 2026","order":4,"name":"first_online","label":"First Online","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"5 June 2026","order":6,"name":"change_date","label":"Change Date","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"Update","order":7,"name":"change_type","label":"Change Type","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"Due to our negligence, the affiliation information for one of the co-authors, Biao Xie, is incomplete in the published version, only part of his institutional affiliation is listed. To ensure the accuracy and integrity of the published work, and out of responsibility to all authors and affiliations, we kindly request that the affiliation for Biao Xie be updated to reflect his full and correct institutional details. All authors agree to this correction of the affiliation and further declare that the correction does not involve any research content in the main text of our paper, nor does it affect the rigor and scientificity of the paper.","order":8,"name":"change_details","label":"Change Details","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"The authors declare no competing interests.","order":1,"name":"Ethics","group":{"name":"EthicsHeading","label":"Competing interests"}}],"article-number":"25"}}