{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,4,22]],"date-time":"2026-04-22T14:11:01Z","timestamp":1776867061754,"version":"3.51.2"},"reference-count":60,"publisher":"Frontiers Media SA","license":[{"start":{"date-parts":[[2025,10,15]],"date-time":"2025-10-15T00:00:00Z","timestamp":1760486400000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"DOI":"10.13039\/501100012688","name":"Universit\u00e4t Rostock","doi-asserted-by":"publisher","id":[{"id":"10.13039\/501100012688","id-type":"DOI","asserted-by":"publisher"}]}],"content-domain":{"domain":["frontiersin.org"],"crossmark-restriction":true},"short-container-title":["Front. Immunol."],"abstract":"<jats:sec><jats:title>Background<\/jats:title><jats:p>Autoimmune pancreatitis (AIP) is a multifactorial disease caused by both genetic and environmental factors. Previous studies have implicated Bach2, a key regulator of adaptive immunity, in the pathogenesis of this disease. However, direct experimental evidence is lacking. Here, we used C57BL\/6N mice with a targeted deletion of <jats:italic>Bach2<\/jats:italic> (<jats:italic>Bach2<\/jats:italic> knockout mice) to study their susceptibility to AIP under homeostatic conditions and in response to two AIP triggers: a high-fat diet (HFD) and polyinosinic:polycytidylic acid (poly I:C).<\/jats:p><\/jats:sec><jats:sec><jats:title>Methods<\/jats:title><jats:p>In this multicenter preclinical study, <jats:italic>Bach2<\/jats:italic> wild type and knockout mice were maintained under homeostatic conditions, challenged with a HFD for 3 months, or treated with poly I:C for 6 weeks. The pancreata were examined histopathologically. Additionally, RNA sequencing and PamGene multiplex kinase activation measurements were performed. To assess the effects of the HFD, the livers were evaluated for the presence of fatty liver disease.<\/jats:p><\/jats:sec><jats:sec><jats:title>Results<\/jats:title><jats:p>Consistent with the results of previous studies, <jats:italic>Bach2<\/jats:italic> knockout mice showed reduced growth and developed eosinophilic crystalline pneumonia, necessitating humane euthanasia at the age of 18 weeks. At 8 and 18 weeks of age, pancreatic infiltrates with lymphocytes typical of AIP were frequently detected in <jats:italic>Bach2<\/jats:italic> knockout mice but not in wild type animals without additional manipulations. RNA sequencing analyses and kinase activity assays revealed the activation of processes linked to adaptive immunity in the pancreatic tissues of <jats:italic>Bach2<\/jats:italic> knockout mice. Wild type mice treated with poly I:C showed lymphocytic infiltrates similar to those of untreated knockout mice, whereas HFD did not induce AIP. In <jats:italic>Bach2<\/jats:italic> knockout mice, HFD and poly I:C did not further enhance the disease. As expected, HFD-fed wild type mice developed fatty liver disease. Strikingly, the livers of <jats:italic>Bach2<\/jats:italic> knockout mice were almost free of fat and histological changes, such as hepatocyte ballooning and degeneration. The data obtained from the two project sites were highly consistent, indicating strong intersite reproducibility.<\/jats:p><\/jats:sec><jats:sec><jats:title>Conclusion<\/jats:title><jats:p><jats:italic>Bach2<\/jats:italic>-deficient C57BL\/6N mice were prone to spontaneous AIP development. This could be due to disturbed immune homeostasis with dysregulated activation of adaptive immune system cells. The protective effect of <jats:italic>Bach2<\/jats:italic> deficiency against the development of fatty liver disease warrants further investigation.<\/jats:p><\/jats:sec>","DOI":"10.3389\/fimmu.2025.1639622","type":"journal-article","created":{"date-parts":[[2025,10,15]],"date-time":"2025-10-15T05:43:58Z","timestamp":1760507038000},"update-policy":"https:\/\/doi.org\/10.3389\/crossmark-policy","source":"Crossref","is-referenced-by-count":1,"title":["Bach2-deficient mice are prone to autoimmune pancreatitis but protected from high-fat diet-induced fatty liver disease"],"prefix":"10.3389","volume":"16","author":[{"given":"Luise","family":"Ehlers","sequence":"first","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Anika","family":"Kasprick","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Ottavia","family":"Agrifoglio","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Natalie","family":"Gross","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Nancy","family":"Ernst","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Alanis","family":"Barbosa Gulde","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Colin","family":"Osterloh","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Noa Linn","family":"Brauckmann","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Nicole Y.","family":"Fischer","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Wendy","family":"Bergmann-Ewert","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Ralf J.","family":"Ludwig","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Katja","family":"Bieber","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Robert","family":"Jaster","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]}],"member":"1965","published-online":{"date-parts":[[2025,10,15]]},"reference":[{"key":"B1","doi-asserted-by":"publisher","DOI":"10.3748\/wjg.v30.i8.817","article-title":"Autoimmune pancreatitis: Cornerstones and future perspectives","volume":"30","author":"Gallo","year":"2024","journal-title":"World J Gastroenterol"},{"key":"B2","doi-asserted-by":"publisher","DOI":"10.3390\/ijms232012667","article-title":"Autoimmune pancreatitis: from pathogenesis to treatment","volume":"23","author":"Nista","year":"2022","journal-title":"Int J Mol Sci"},{"key":"B3","doi-asserted-by":"publisher","DOI":"10.3389\/fimmu.2025.1626082","article-title":"Autoimmune disease: genetic susceptibility, environmental triggers, and immune dysregulation. Where can we develop therapies","volume":"16","author":"Kumar","year":"2025","journal-title":"Front Immunol"},{"key":"B4","doi-asserted-by":"publisher","DOI":"10.1111\/jcmm.15540","article-title":"Impact of diet and genes on murine autoimmune pancreatitis","volume":"24","author":"Jaster","year":"2020","journal-title":"J Cell Mol Med"},{"key":"B5","doi-asserted-by":"publisher","first-page":"4097","DOI":"10.1038\/s41467-019-11952-w","article-title":"Gene-diet interactions associated with complex trait variation in an advanced intercross outbred mouse line","volume":"10","author":"Vorobyev","year":"2019","journal-title":"Nat Commun"},{"key":"B6","doi-asserted-by":"publisher","DOI":"10.1016\/j.pan.2022.03.015","article-title":"HLA-DRB1\u221716 and -DQB1\u221705 alleles are strongly associated with autoimmune pancreatitis in a cohort of hundred patients","volume":"22","author":"Goni","year":"2022","journal-title":"Pancreatology"},{"key":"B7","doi-asserted-by":"publisher","DOI":"10.1136\/gut.2006.095059","article-title":"Genetic association of Fc receptor-like 3 polymorphisms with autoimmune pancreatitis in Japanese patients","volume":"55","author":"Umemura","year":"2006","journal-title":"Gut"},{"key":"B8","doi-asserted-by":"publisher","DOI":"10.1373\/clinchem.2007.085951","article-title":"T-cell regulatory gene CTLA-4 polymorphism\/haplotype association with autoimmune pancreatitis","volume":"53","author":"Chang","year":"2007","journal-title":"Clin Chem"},{"key":"B9","doi-asserted-by":"publisher","DOI":"10.1111\/j.1572-0241.2007.01750.x","article-title":"Association of autoimmune pancreatitis with cytotoxic T-lymphocyte antigen 4 gene polymorphisms in Japanese patients","volume":"103","author":"Umemura","year":"2008","journal-title":"Am J Gastroenterol"},{"key":"B10","doi-asserted-by":"publisher","DOI":"10.1155\/2011\/264747","article-title":"Polymorphism in the KCNA3 gene is associated with susceptibility to autoimmune pancreatitis in the Japanese population","volume":"31","author":"Ota","year":"2011","journal-title":"Dis Markers"},{"key":"B11","doi-asserted-by":"publisher","first-page":"3332","DOI":"10.3748\/wjg.v19.i21.3332","article-title":"PRSS1 _p.Leu81Met mutation results in autoimmune pancreatitis","volume":"19","author":"Gao","year":"2013","journal-title":"World J Gastroenterol"},{"key":"B12","doi-asserted-by":"publisher","DOI":"10.2174\/1566524013666131118114432","article-title":"Identification of a novel frame-shift mutation in PRSS1 gene in Han patients with autoimmune pancreatitis","volume":"14","author":"Gao","year":"2014","journal-title":"CMM"},{"key":"B13","doi-asserted-by":"publisher","first-page":"68","DOI":"10.1111\/j.1365-2249.1992.tb06879.x","article-title":"Spontaneous development of pancreatitis in the MRL\/Mp strain of mice in autoimmune mechanism","volume":"89","author":"Kanno","year":"1992","journal-title":"Clin Exp Immunol"},{"key":"B14","doi-asserted-by":"publisher","first-page":"27","DOI":"10.1046\/j.1365-2249.2002.01881.x","article-title":"A novel autoimmune pancreatitis model in MRL mice treated with polyinosinic:polycytidylic acid","volume":"129","author":"Qu","year":"2002","journal-title":"Clin Exp Immunol"},{"key":"B15","doi-asserted-by":"publisher","DOI":"10.1136\/jmg.2011.089730","article-title":"Identification of quantitative trait loci for murine autoimmune pancreatitis","volume":"48","author":"Asghari","year":"2011","journal-title":"J Med Genet"},{"key":"B16","doi-asserted-by":"publisher","DOI":"10.3390\/biomedicines12112480","article-title":"Dexamethasone\u2019s clinical efficacy in experimental autoimmune pancreatitis correlates with a unique transcriptomic signature, whilst kinase inhibitors are not effective","volume":"12","author":"Agrifoglio","year":"2024","journal-title":"Biomedicines"},{"key":"B17","doi-asserted-by":"publisher","first-page":"e2412850","DOI":"10.1002\/advs.202412850","article-title":"The multifaceted roles of BACH1 in disease: implications for biological functions and therapeutic applications","volume":"12","author":"Wei","year":"2025","journal-title":"Adv Sci (Weinh)"},{"key":"B18","doi-asserted-by":"publisher","first-page":"189325","DOI":"10.1016\/j.bbcan.2025.189325","article-title":"BTB domain and CNC homolog 2: A master regulator that controls immune response and cancer progression","volume":"1880","author":"Li","year":"2025","journal-title":"Biochim Biophys Acta Rev Cancer"},{"key":"B19","doi-asserted-by":"publisher","first-page":"1347969","DOI":"10.1155\/2018\/1347969","article-title":"Bach1: function, regulation, and involvement in disease","volume":"2018","author":"Zhang","year":"2018","journal-title":"Oxid Med Cell Longev"},{"key":"B20","doi-asserted-by":"publisher","DOI":"10.1038\/nature14154","article-title":"Super-enhancers delineate disease-associated regulatory nodes in T cells","volume":"520","author":"Vahedi","year":"2015","journal-title":"Nature"},{"key":"B21","doi-asserted-by":"publisher","DOI":"10.1038\/ni.3753","article-title":"BACH2 immunodeficiency illustrates an association between super-enhancers and haploinsufficiency","volume":"18","author":"Afzali","year":"2017","journal-title":"Nat Immunol"},{"key":"B22","doi-asserted-by":"publisher","DOI":"10.1038\/ng.249","article-title":"Meta-analysis of genome-wide association study data identifies additional type 1 diabetes risk loci","volume":"40","author":"Cooper","year":"2008","journal-title":"Nat Genet"},{"key":"B23","doi-asserted-by":"publisher","DOI":"10.1038\/ng.717","article-title":"Genome-wide meta-analysis increases to 71 the number of confirmed Crohn\u2019s disease susceptibility loci","volume":"42","author":"Franke","year":"2010","journal-title":"Nat Genet"},{"key":"B24","doi-asserted-by":"publisher","first-page":"295","DOI":"10.1038\/ng.543","article-title":"Multiple common variants for celiac disease influencing immune gene expression","volume":"42","author":"Dubois","year":"2010","journal-title":"Nat Genet"},{"key":"B25","doi-asserted-by":"publisher","DOI":"10.1038\/nature10251","article-title":"Genetic risk and a primary role for cell-mediated immune mechanisms in multiple sclerosis","volume":"476","author":"Sawcer","year":"2011","journal-title":"Nature"},{"key":"B26","doi-asserted-by":"publisher","first-page":"1214","DOI":"10.1016\/j.cell.2020.08.008","article-title":"The polygenic and monogenic basis of blood traits and diseases","volume":"182","author":"Vuckovic","year":"2020","journal-title":"Cell"},{"key":"B27","doi-asserted-by":"publisher","first-page":"e1003225","DOI":"10.1371\/journal.pgen.1003225","article-title":"Loci associated with N-glycosylation of human immunoglobulin G show pleiotropy with autoimmune diseases and haematological cancers","volume":"9","author":"Lauc","year":"2013","journal-title":"PloS Genet"},{"key":"B28","doi-asserted-by":"publisher","DOI":"10.4049\/jimmunol.1801626","article-title":"Bach2 negatively regulates T follicular helper cell differentiation and is critical for CD4+ T cell memory","volume":"202","author":"Geng","year":"2019","journal-title":"J Immunol"},{"key":"B29","doi-asserted-by":"publisher","first-page":"e0136298","DOI":"10.1371\/journal.pone.0136298","article-title":"Quantitative trait locus analysis implicates CD4+\/CD44high memory T cells in the pathogenesis of murine autoimmune pancreatitis","volume":"10","author":"Bischof","year":"2015","journal-title":"PloS One"},{"key":"B30","doi-asserted-by":"publisher","DOI":"10.1177\/2050640617716596","article-title":"Bach2 repression mediates Th17 cell induced inflammation and associates with clinical features of advanced disease in chronic pancreatitis","volume":"6","author":"Sasikala","year":"2018","journal-title":"United Eur Gastroenterol J"},{"key":"B31","doi-asserted-by":"publisher","first-page":"494","DOI":"10.1136\/gutjnl-2012-303635","article-title":"Autoimmune pancreatitis in MRL\/Mp mice is a T cell-mediated disease responsive to cyclosporine A and rapamycin treatment","volume":"63","author":"Schwaiger","year":"2014","journal-title":"Gut"},{"key":"B32","doi-asserted-by":"publisher","first-page":"197","DOI":"10.7570\/jomes23052","article-title":"Metabolic dysfunction-associated steatotic liver disease (MASLD): A state-of-the-art review","volume":"32","author":"Chan","year":"2023","journal-title":"J Obes Metab Syndr"},{"key":"B33","doi-asserted-by":"publisher","DOI":"10.1093\/nar\/gku1193","article-title":"INFRAFRONTIER\u2013providing mutant mouse resources as research tools for the international scientific community","volume":"43","year":"2015","journal-title":"Nucleic Acids Res"},{"key":"B34","doi-asserted-by":"publisher","DOI":"10.1038\/nature10163","article-title":"A conditional knockout resource for the genome-wide study of mouse gene function","volume":"474","author":"Skarnes","year":"2011","journal-title":"Nature"},{"key":"B35","doi-asserted-by":"publisher","DOI":"10.1111\/jcmm.13537","article-title":"Adoptive transfer of CD3+ T cells and CD4+ CD44high memory T cells induces autoimmune pancreatitis in MRL\/MpJ mice","volume":"22","author":"Ehlers","year":"2018","journal-title":"J Cell Mol Med"},{"key":"B36","doi-asserted-by":"publisher","DOI":"10.1016\/j.hbpd.2019.10.007","article-title":"Different characteristics of chronic dibutyltin dichloride-induced pancreatitis and cholangitis in mouse and rat","volume":"19","author":"L\u00fctt","year":"2020","journal-title":"Hepatobiliary Pancreat Dis Int"},{"key":"B37","doi-asserted-by":"publisher","DOI":"10.1002\/hep.20701","article-title":"Design and validation of a histological scoring system for nonalcoholic fatty liver disease","volume":"41","author":"Kleiner","year":"2005","journal-title":"Hepatology"},{"key":"B38","doi-asserted-by":"publisher","first-page":"408","DOI":"10.2144\/97223bm07","article-title":"Tissue homogenization as a key step in extracting RNA from human and rat pancreatic tissue","volume":"22","author":"Sparmann","year":"1997","journal-title":"Biotechniques"},{"key":"B39","doi-asserted-by":"publisher","first-page":"97","DOI":"10.1016\/j.cellsig.2009.09.019","article-title":"Mathematical modelling of interferon-gamma signalling in pancreatic stellate cells reflects and predicts the dynamics of STAT1 pathway activity","volume":"22","author":"Rateitschak","year":"2010","journal-title":"Cell Signal"},{"key":"B40","doi-asserted-by":"publisher","DOI":"10.3389\/fmed.2021.713312","article-title":"Topical application of the PI3K\u03b2-selective small molecule inhibitor TGX-221 is an effective treatment option for experimental epidermolysis bullosa acquisita","volume":"8","author":"Zillikens","year":"2021","journal-title":"Front Med (Lausanne)"},{"key":"B41","doi-asserted-by":"publisher","DOI":"10.3389\/fimmu.2022.865241","article-title":"Cutaneous kinase activity correlates with treatment outcomes following PI3K delta inhibition in mice with experimental pemphigoid diseases","volume":"13","author":"Ghorbanalipoor","year":"2022","journal-title":"Front Immunol"},{"key":"B42","doi-asserted-by":"publisher","first-page":"306798","DOI":"10.6064\/2012\/306798","article-title":"Kinome profiling","volume":"2012","author":"Peppelenbosch","year":"2012","journal-title":"Scientifica (Cairo)"},{"key":"B43","doi-asserted-by":"publisher","DOI":"10.1073\/pnas.1314810111","article-title":"Visual account of protein investment in cellular functions","volume":"111","author":"Liebermeister","year":"2014","journal-title":"Proc Natl Acad Sci U.S.A"},{"key":"B44","doi-asserted-by":"publisher","first-page":"347","DOI":"10.1016\/j.cels.2018.07.001","article-title":"Coral: clear and customizable visualization of human kinome data","volume":"7","author":"Metz","year":"2018","journal-title":"Cell Syst"},{"key":"B45","doi-asserted-by":"publisher","DOI":"10.1093\/nar\/gkae1113","article-title":"The STRING database in 2025: protein networks with directionality of regulation","volume":"53","author":"Szklarczyk","year":"2025","journal-title":"Nucleic Acids Res"},{"key":"B46","doi-asserted-by":"publisher","DOI":"10.1084\/jem.20130028","article-title":"Transcription repressor Bach2 is required for pulmonary surfactant homeostasis and alveolar macrophage function","volume":"210","author":"Nakamura","year":"2013","journal-title":"J Exp Med"},{"key":"B47","doi-asserted-by":"publisher","DOI":"10.4049\/jimmunol.1302378","article-title":"Bach2 regulates homeostasis of Foxp3+ regulatory T cells and protects against fatal lung disease in mice","volume":"192","author":"Kim","year":"2014","journal-title":"J Immunol"},{"key":"B48","doi-asserted-by":"publisher","DOI":"10.1182\/blood-2010-07-296483","article-title":"Heme regulates B-cell differentiation, antibody class switch, and heme oxygenase-1 expression in B cells as a ligand of Bach2","volume":"117","author":"Watanabe-Matsui","year":"2011","journal-title":"Blood"},{"key":"B49","doi-asserted-by":"publisher","DOI":"10.3390\/ijms24129946","article-title":"Acinar-to-ductal metaplasia (ADM): on the road to pancreatic intraepithelial neoplasia (PanIN) and pancreatic cancer","volume":"24","author":"Marstrand-Dauc\u00e9","year":"2023","journal-title":"Int J Mol Sci"},{"key":"B50","doi-asserted-by":"publisher","DOI":"10.1111\/j.0019-2805.2004.01832.x","article-title":"Positive and negative adaptors in T-cell signalling","volume":"111","author":"Wilkinson","year":"2004","journal-title":"Immunology"},{"key":"B51","doi-asserted-by":"publisher","first-page":"412","DOI":"10.1038\/s41392-021-00823-w","article-title":"T cell receptor (TCR) signaling in health and disease","volume":"6","author":"Shah","year":"2021","journal-title":"Signal Transduct Target Ther"},{"key":"B52","doi-asserted-by":"publisher","first-page":"e2003693","DOI":"10.1371\/journal.pbio.2003693","article-title":"Reproducibility of preclinical animal research improves with heterogeneity of study samples","volume":"16","author":"Voelkl","year":"2018","journal-title":"PloS Biol"},{"key":"B53","doi-asserted-by":"publisher","first-page":"14","DOI":"10.1038\/s41572-025-00599-1","article-title":"Metabolic dysfunction-associated steatotic liver disease in adults","volume":"11","author":"Huang","year":"2025","journal-title":"Nat Rev Dis Primers"},{"key":"B54","doi-asserted-by":"publisher","DOI":"10.1093\/eurjpc\/zwae306","article-title":"Cardiovascular disease and metabolic dysfunction-associated steatotic liver disease: pathophysiology and diagnostic aspects","author":"M\u00f8ller","year":"2025","journal-title":"Eur J Prev Cardiol"},{"key":"B55","doi-asserted-by":"publisher","DOI":"10.2147\/IJGM.S413545","article-title":"Identification and validation of hub immune-related genes in non-alcoholic fatty liver disease","volume":"16","author":"Li","year":"2023","journal-title":"Int J Gen Med"},{"key":"B56","doi-asserted-by":"publisher","DOI":"10.1016\/j.jhep.2021.07.032","article-title":"Regulatory T-cell and neutrophil extracellular trap interaction contributes to carcinogenesis in non-alcoholic steatohepatitis","volume":"75","author":"Wang","year":"2021","journal-title":"J Hepatol"},{"key":"B57","doi-asserted-by":"publisher","DOI":"10.1038\/s41586-021-03233-8","article-title":"Auto-aggressive CXCR6+ CD8 T cells cause liver immune pathology in NASH","volume":"592","author":"Dudek","year":"2021","journal-title":"Nature"},{"key":"B58","doi-asserted-by":"publisher","DOI":"10.1016\/j.stemcr.2020.12.005","article-title":"Bach2 deficiency promotes intestinal epithelial regeneration by accelerating DNA repair in intestinal stem cells","volume":"16","author":"Li","year":"2021","journal-title":"Stem Cell Rep"},{"key":"B59","doi-asserted-by":"publisher","DOI":"10.1002\/ijc.35326","article-title":"Pancreatic stellate cell: Update on molecular investigations and clinical translation in pancreatic cancer","volume":"156","author":"Liu","year":"2025","journal-title":"Int J Cancer"},{"key":"B60","doi-asserted-by":"publisher","DOI":"10.1093\/nar\/30.1.207","article-title":"Gene Expression Omnibus: NCBI gene expression and hybridization array data repository","volume":"30","author":"Edgar","year":"2002","journal-title":"Nucleic Acids Res"}],"container-title":["Frontiers in Immunology"],"original-title":[],"link":[{"URL":"https:\/\/www.frontiersin.org\/articles\/10.3389\/fimmu.2025.1639622\/full","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2025,10,15]],"date-time":"2025-10-15T05:44:00Z","timestamp":1760507040000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.frontiersin.org\/articles\/10.3389\/fimmu.2025.1639622\/full"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2025,10,15]]},"references-count":60,"alternative-id":["10.3389\/fimmu.2025.1639622"],"URL":"https:\/\/doi.org\/10.3389\/fimmu.2025.1639622","relation":{},"ISSN":["1664-3224"],"issn-type":[{"value":"1664-3224","type":"electronic"}],"subject":[],"published":{"date-parts":[[2025,10,15]]},"article-number":"1639622"}}