{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,10,14]],"date-time":"2026-10-14T21:28:19Z","timestamp":1792013299671,"version":"4.4.0"},"reference-count":201,"publisher":"Frontiers Media SA","license":[{"start":{"date-parts":[[2026,7,15]],"date-time":"2026-07-15T00:00:00Z","timestamp":1784073600000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"content-domain":{"domain":["frontiersin.org"],"crossmark-restriction":true},"short-container-title":["Front. Immunol."],"abstract":"<jats:p>Rheumatoid arthritis is a chronic systemic autoimmune disease in which the earliest breaks in immune tolerance may arise at mucosal surfaces before the clinical onset of synovitis. Among these sites, the gut has emerged as a particularly compelling candidate because it integrates microbial, epithelial, metabolic, and immune pathways with the potential to shape systemic inflammation. In this review, we examine the biological foundations of the gut\u2013joint axis in rheumatoid arthritis, focusing on intestinal barrier structure, microbiome alterations, mucosal immune crosstalk, and mechanisms of barrier dysfunction. Current human evidence links rheumatoid arthritis to heterogeneous shifts in gut microbial composition, depletion of beneficial metabolite-producing commensals, altered immune\u2013metabolic signaling, and biomarker patterns consistent with epithelial injury and microbial-product translocation. At the same time, available data do not support the existence of a single, universal microbial or permeability signature that defines the disease across populations. Recent longitudinal studies further challenge the concept of stable, long-standing dysbiosis and instead suggest a late, transient phase of ecological instability arising close to symptom onset. Experimental models provide stronger mechanistic support, showing that dysbiotic microbial communities, impaired barrier integrity, and strain-specific host\u2013microbe interactions can promote T helper 17-skewed immunity and aggravate arthritis. Collectively, these findings support a context-dependent contribution of the gut to rheumatoid arthritis pathogenesis while underscoring the need for longitudinal, strain-resolved, and multi-omic human studies to clarify causality, refine disease models, and identify clinically meaningful windows for intervention.<\/jats:p>","DOI":"10.3389\/fimmu.2026.1887782","type":"journal-article","created":{"date-parts":[[2026,7,15]],"date-time":"2026-07-15T06:10:55Z","timestamp":1784095855000},"update-policy":"https:\/\/doi.org\/10.3389\/crossmark-policy","source":"Crossref","is-referenced-by-count":2,"title":["Gut microbiota and intestinal permeability in rheumatoid arthritis: pathogenic mechanisms"],"prefix":"10.3389","volume":"17","author":[{"given":"Jan","family":"Bilski","sequence":"first","affiliation":[{"name":"Department of Biomechanics and Kinesiology, Chair of Biomedical Sciences, Faculty of Health Sciences, Institute of Physiotherapy, Jagiellonian University Medical College","place":["Krakow, Poland"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Agata I.","family":"Schramm-Luc","sequence":"additional","affiliation":[{"name":"Department of Internal and Agricultural Medicine, Faculty of Medicine, Jagiellonian University Medical College","place":["Krakow, Poland"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Marian","family":"Szczepanik","sequence":"additional","affiliation":[{"name":"Chair of Biomedical Sciences, Institute of Physiotherapy, Faculty of Health Sciences, Jagiellonian University Medical College","place":["Krakow, Poland"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Piotr","family":"Pierzchalski","sequence":"additional","affiliation":[{"name":"Department of Medical Physiology, Chair of Biomedical Sciences, Faculty of Health Sciences, Institute of Physiotherapy, Jagiellonian University Medical College","place":["Krakow, Poland"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Agnieszka","family":"Krawczyk","sequence":"additional","affiliation":[{"name":"Department of Biomechanics and Kinesiology, Chair of Biomedical Sciences, Faculty of Health Sciences, Institute of Physiotherapy, Jagiellonian University Medical College","place":["Krakow, Poland"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Kevin","family":"Luc","sequence":"additional","affiliation":[{"name":"Department of Internal and Agricultural Medicine, Faculty of Medicine, Jagiellonian University Medical College","place":["Krakow, Poland"]}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"1965","published-online":{"date-parts":[[2026,7,15]]},"reference":[{"key":"B1","doi-asserted-by":"publisher","DOI":"10.3389\/fimmu.2025.1547763","article-title":"Global, regional, and national burden and trends of rheumatoid arthritis among the elderly population: An analysis based on the 2021 global burden of disease study","volume":"16","author":"Wei","year":"2025","journal-title":"Front Immunol"},{"key":"B2","doi-asserted-by":"publisher","DOI":"10.1016\/j.immuni.2022.11.009","article-title":"The pathogenesis of rheumatoid arthritis","volume":"55","author":"Alivernini","year":"2022","journal-title":"Immunity"},{"key":"B3","doi-asserted-by":"publisher","DOI":"10.5603\/RF.2022.0024","article-title":"Intestinal dysbiosis and increased intestinal permeability as a potential risk factor for the development and progression of rheumatoid arthritis","volume":"8","author":"W\u0105\u017c","year":"2022","journal-title":"Rheumatol Forum"},{"key":"B4","doi-asserted-by":"publisher","DOI":"10.3390\/ijms22020686","article-title":"From rheumatoid factor to anti-citrullinated protein antibodies and anti-carbamylated protein antibodies for diagnosis and prognosis prediction in patients with rheumatoid arthritis","volume":"22","author":"Wu","year":"2021","journal-title":"Int J Mol Sci"},{"key":"B5","doi-asserted-by":"publisher","DOI":"10.1155\/2019\/7592851","article-title":"An overview of the intrinsic role of citrullination in autoimmune disorders","volume":"2019","author":"Alghamdi","year":"2019","journal-title":"J Immunol Res"},{"key":"B6","doi-asserted-by":"publisher","DOI":"10.3389\/fimmu.2022.980805","article-title":"Host and bacterial factors linking periodontitis and rheumatoid arthritis","volume":"13","author":"Krutyholowa","year":"2022","journal-title":"Front Immunol"},{"key":"B7","doi-asserted-by":"publisher","first-page":"369ra176","DOI":"10.1126\/scitranslmed.aaj1921","article-title":"Aggregatibacter actinomycetemcomitans-induced hypercitrullination links periodontal infection to autoimmunity in rheumatoid arthritis","volume":"8","author":"Konig","year":"2016","journal-title":"Sci Transl Med"},{"key":"B8","doi-asserted-by":"publisher","DOI":"10.3389\/fimmu.2025.1503474","article-title":"Linking microbial communities to rheumatoid arthritis: Focus on gut, oral microbiome and their extracellular vesicles","volume":"16","author":"Lu","year":"2025","journal-title":"Front Immunol"},{"key":"B9","doi-asserted-by":"publisher","DOI":"10.1097\/BOR.0000000000000008","article-title":"Microbiome and mucosal inflammation as extra-articular triggers for rheumatoid arthritis and autoimmunity","volume":"26","author":"Brusca","year":"2014","journal-title":"Curr Opin Rheumatol"},{"key":"B10","doi-asserted-by":"publisher","first-page":"18","DOI":"10.1007\/s40588-023-00213-6","article-title":"Gut microbiota and autoimmune diseases: Mechanisms, treatment, challenges, and future recommendations","volume":"11","author":"Sadeghpour Heravi","year":"2024","journal-title":"Curr Clin Microbiol Rep"},{"key":"B11","doi-asserted-by":"publisher","DOI":"10.3389\/fimmu.2017.00598","article-title":"Leaky gut as a danger signal for autoimmune diseases","volume":"8","author":"Mu","year":"2017","journal-title":"Front Immunol"},{"key":"B12","doi-asserted-by":"publisher","DOI":"10.1007\/s11739-023-03374-w","article-title":"Gut microbiota, intestinal permeability, and systemic inflammation: A narrative review","volume":"19","author":"Di Vincenzo","year":"2024","journal-title":"Intern Emerg Med"},{"key":"B13","doi-asserted-by":"publisher","DOI":"10.3389\/fmed.2018.00349","article-title":"Diet, microbiota, and gut permeability-the unknown triad in rheumatoid arthritis","volume":"5","author":"Guerreiro","year":"2018","journal-title":"Front Med (Lausanne)"},{"key":"B14","doi-asserted-by":"publisher","DOI":"10.1242\/jcs.261468","article-title":"Muc13 negatively regulates tight junction proteins and intestinal epithelial barrier integrity via protein kinase C","volume":"137","author":"Segui-Perez","year":"2024","journal-title":"J Cell Sci"},{"key":"B15","doi-asserted-by":"publisher","DOI":"10.3389\/fimmu.2020.00323","article-title":"The crosstalk between hippo-yap pathway and innate immunity","volume":"11","author":"Wang","year":"2020","journal-title":"Front Immunol"},{"key":"B16","doi-asserted-by":"publisher","DOI":"10.3390\/cells9020475","article-title":"The impact of cigarette smoking on risk of rheumatoid arthritis: A narrative review","volume":"9","author":"Ishikawa","year":"2020","journal-title":"Cells"},{"key":"B17","doi-asserted-by":"publisher","DOI":"10.1186\/s13075-023-03118-3","article-title":"Faecalibacterium prausnitzii alleviates inflammatory arthritis and regulates il-17 production, short chain fatty acids, and the intestinal microbial flora in experimental mouse model for rheumatoid arthritis","volume":"25","author":"Moon","year":"2023","journal-title":"Arthritis Res Ther"},{"key":"B18","doi-asserted-by":"publisher","DOI":"10.3390\/biom14121653","article-title":"Gut microbiota modulation: A novel strategy for rheumatoid arthritis therapy","volume":"14","author":"Chasov","year":"2024","journal-title":"Biomolecules"},{"key":"B19","doi-asserted-by":"publisher","first-page":"895","DOI":"10.1038\/nm.3914","article-title":"The oral and gut microbiomes are perturbed in rheumatoid arthritis and partly normalized after treatment","volume":"21","author":"Zhang","year":"2015","journal-title":"Nat Med"},{"key":"B20","doi-asserted-by":"publisher","DOI":"10.1136\/ard-2022-222881","article-title":"Genomic repertoires linked with pathogenic potency of arthritogenic prevotella copri isolated from the gut of patients with rheumatoid arthritis","volume":"82","author":"Nii","year":"2023","journal-title":"Ann Rheum Dis"},{"key":"B21","doi-asserted-by":"publisher","DOI":"10.1002\/rai2.12043","article-title":"Mechanism-driven strategies for prevention of rheumatoid arthritis","volume":"2","author":"Holers","year":"2022","journal-title":"Rheumatol Autoimmun"},{"key":"B22","doi-asserted-by":"publisher","DOI":"10.37349\/emed.2024.00251","article-title":"Gut microbiota dysbiosis in rheumatoid arthritis: A systematic review and meta-analysis","volume":"5","author":"Su","year":"2024","journal-title":"Explor Med"},{"key":"B23","doi-asserted-by":"publisher","DOI":"10.3389\/fcimb.2021.763507","article-title":"The gut microbiome and metabolites are altered and interrelated in patients with rheumatoid arthritis","volume":"11","author":"Yu","year":"2021","journal-title":"Front Cell Infect Microbiol"},{"key":"B24","doi-asserted-by":"publisher","DOI":"10.1136\/rmdopen-2023-003589","article-title":"Gut microbiome and intestinal inflammation in preclinical stages of rheumatoid arthritis","volume":"10","author":"Gilbert","year":"2024","journal-title":"RMD Open"},{"key":"B25","doi-asserted-by":"publisher","DOI":"10.1136\/ard-2024-226362","article-title":"Dynamics of the gut microbiome in individuals at risk of rheumatoid arthritis: A cross-sectional and longitudinal observational study","author":"Rooney","year":"2024","journal-title":"Ann Rheum Dis"},{"key":"B26","doi-asserted-by":"publisher","DOI":"10.1172\/JCI195374","article-title":"Microbial threads in the tapestry of rheumatoid arthritis","volume":"135","author":"Li","year":"2025","journal-title":"J Clin Invest"},{"key":"B27","doi-asserted-by":"publisher","DOI":"10.3389\/fmicb.2024.1414000","article-title":"Temporal dynamics and species-level complexity of prevotella spp. in the human gut microbiota: Implications for enterotypes and health","volume":"15","author":"Han","year":"2024","journal-title":"Front Microbiol"},{"key":"B28","doi-asserted-by":"publisher","DOI":"10.3389\/fimmu.2022.1007165","article-title":"Gut microbiota and rheumatoid arthritis: From pathogenesis to novel therapeutic opportunities","volume":"13","author":"Zhao","year":"2022","journal-title":"Front Immunol"},{"key":"B29","doi-asserted-by":"publisher","DOI":"10.3390\/ijms222010922","article-title":"Cytokine networks in the pathogenesis of rheumatoid arthritis","volume":"22","author":"Kondo","year":"2021","journal-title":"Int J Mol Sci"},{"key":"B30","doi-asserted-by":"publisher","DOI":"10.3389\/fimmu.2022.930112","article-title":"How ra associated hla-dr molecules contribute to the development of antibodies to citrullinated proteins: The hapten carrier model","volume":"13","author":"Roudier","year":"2022","journal-title":"Front Immunol"},{"key":"B31","doi-asserted-by":"publisher","DOI":"10.1186\/s12876-014-0189-7","article-title":"Intestinal permeability--a new target for disease prevention and therapy","volume":"14","author":"Bischoff","year":"2014","journal-title":"BMC Gastroenterol"},{"key":"B32","doi-asserted-by":"publisher","DOI":"10.3390\/ijerph182312836","article-title":"Intestinal barrier in human health and disease","volume":"18","author":"Di Tommaso","year":"2021","journal-title":"Int J Environ Res Public Health"},{"key":"B33","doi-asserted-by":"publisher","DOI":"10.1016\/j.aninu.2025.03.004","article-title":"The role of glycosylated mucins in maintaining intestinal homeostasis and gut health","volume":"21","author":"Cheng","year":"2025","journal-title":"Anim Nutr"},{"key":"B34","doi-asserted-by":"publisher","first-page":"8","DOI":"10.1111\/imr.12182","article-title":"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","volume":"260","author":"Pelaseyed","year":"2014","journal-title":"Immunol Rev"},{"key":"B35","doi-asserted-by":"publisher","DOI":"10.3390\/ijms24044227","article-title":"View from the biological property: Insight into the functional diversity and complexity of the gut mucus","volume":"24","author":"He","year":"2023","journal-title":"Int J Mol Sci"},{"key":"B36","doi-asserted-by":"publisher","DOI":"10.1038\/s12276-023-00960-y","article-title":"Intestinal mucus components and secretion mechanisms: What we do and do not know","volume":"55","author":"Song","year":"2023","journal-title":"Exp Mol Med"},{"key":"B37","doi-asserted-by":"publisher","DOI":"10.1073\/pnas.0705984104","article-title":"Gel-forming mucins appeared early in metazoan evolution","volume":"104","author":"Lang","year":"2007","journal-title":"Proc Natl Acad Sci USA"},{"key":"B38","doi-asserted-by":"publisher","DOI":"10.1080\/21688370.2022.2077069","article-title":"The diet-microbiota axis: A key regulator of intestinal permeability in human health and disease","volume":"11","author":"Lama Tamang","year":"2023","journal-title":"Tissue Barriers"},{"key":"B39","doi-asserted-by":"publisher","DOI":"10.1080\/19490976.2022.2052699","article-title":"The barrier and beyond: Roles of intestinal mucus and mucin-type o-glycosylation in resistance and tolerance defense strategies guiding host-microbe symbiosis","volume":"14","author":"Bergstrom","year":"2022","journal-title":"Gut Microbes"},{"key":"B40","doi-asserted-by":"publisher","first-page":"202","DOI":"10.1186\/s12876-024-03287-8","article-title":"B3gnt7 regulates mucin o-glycosylation to alleviate colonic inflammation","volume":"24","author":"Wang","year":"2024","journal-title":"BMC Gastroenterol"},{"key":"B41","doi-asserted-by":"publisher","first-page":"5999","DOI":"10.1093\/emboj\/20.21.5999","article-title":"Cytoplasmic o-glycosylation prevents cell surface transport of e-cadherin during apoptosis","volume":"20","author":"Zhu","year":"2001","journal-title":"EMBO J"},{"key":"B42","doi-asserted-by":"publisher","first-page":"597","DOI":"10.1038\/s41575-020-0331-7","article-title":"Intestinal epithelial glycosylation in homeostasis and gut microbiota interactions in ibd","volume":"17","author":"Kudelka","year":"2020","journal-title":"Nat Rev Gastroenterol Hepatol"},{"key":"B43","doi-asserted-by":"publisher","DOI":"10.1038\/s42003-019-0735-5","article-title":"Nanoarchitecture and dynamics of the mouse enteric glycocalyx examined by freeze-etching electron tomography and intravital microscopy","volume":"3","author":"Sun","year":"2020","journal-title":"Commun Biol"},{"key":"B44","doi-asserted-by":"publisher","DOI":"10.1016\/j.bcp.2013.10.021","article-title":"Modulation of n-glycosylation by mesalamine facilitates membranous e-cadherin expression in colon epithelial cells","volume":"87","author":"Khare","year":"2014","journal-title":"Biochem Pharmacol"},{"key":"B45","doi-asserted-by":"publisher","DOI":"10.1016\/j.ejcb.2021.151186","article-title":"Advances in understanding n-glycosylation structure, function, and regulation in health and disease","volume":"100","author":"Esmail","year":"2021","journal-title":"Eur J Cell Biol"},{"key":"B46","doi-asserted-by":"publisher","DOI":"10.1074\/jbc.M704713200","article-title":"Inverse correlation between the extent of n-glycan branching and intercellular adhesion in epithelia. Contribution of the na,k-atpase beta1 subunit","volume":"283","author":"Vagin","year":"2008","journal-title":"J Biol Chem"},{"key":"B47","doi-asserted-by":"publisher","DOI":"10.1111\/j.1742-4658.2008.06346.x","article-title":"Branched n-glycans regulate the biological functions of integrins and cadherins","volume":"275","author":"Zhao","year":"2008","journal-title":"FEBS J"},{"key":"B48","doi-asserted-by":"publisher","DOI":"10.1091\/mbc.E14-12-1604","article-title":"N-glycosylation controls the function of junctional adhesion molecule-a","volume":"26","author":"Scott","year":"2015","journal-title":"Mol Biol Cell"},{"key":"B49","doi-asserted-by":"publisher","DOI":"10.3389\/fcimb.2022.856962","article-title":"Mucus and mucins: The underappreciated host defence system","volume":"12","author":"Sheng","year":"2022","journal-title":"Front Cell Infect Microbiol"},{"key":"B50","doi-asserted-by":"publisher","DOI":"10.1007\/BF00702345","article-title":"Colonic mucins in ulcerative colitis: Evidence for loss of sulfation","volume":"13","author":"Corfield","year":"1996","journal-title":"Glycoconj J"},{"key":"B51","doi-asserted-by":"publisher","DOI":"10.1002\/ibd.21625","article-title":"Altered O-glycosylation profile of Muc2 mucin occurs in active ulcerative colitis and is associated with increased inflammation","volume":"17","author":"Larsson","year":"2011","journal-title":"Inflammation Bowel Dis"},{"key":"B52","doi-asserted-by":"publisher","DOI":"10.1038\/s41385-021-00466-8","article-title":"Finding the sweet spot: Glycosylation mediated regulation of intestinal inflammation","volume":"15","author":"Brazil","year":"2022","journal-title":"Mucosal Immunol"},{"key":"B53","doi-asserted-by":"publisher","DOI":"10.3390\/ijms221910224","article-title":"Gut microbiota and dietary factors as modulators of the mucus layer in inflammatory bowel disease","volume":"22","author":"Fernandez-Tome","year":"2021","journal-title":"Int J Mol Sci"},{"key":"B54","doi-asserted-by":"publisher","DOI":"10.1016\/j.tim.2013.02.001","article-title":"The intestinal microbiota and host immune interactions in the critically ill","volume":"21","author":"Schuijt","year":"2013","journal-title":"Trends Microbiol"},{"key":"B55","doi-asserted-by":"publisher","DOI":"10.3390\/nu12041185","article-title":"Intestinal permeability, inflammation and the role of nutrients","volume":"12","author":"Farre","year":"2020","journal-title":"Nutrients"},{"key":"B56","doi-asserted-by":"publisher","DOI":"10.1016\/j.it.2018.04.002","article-title":"The intestinal epithelium: Central coordinator of mucosal immunity","volume":"39","author":"Allaire","year":"2018","journal-title":"Trends Immunol"},{"key":"B57","doi-asserted-by":"publisher","DOI":"10.1007\/s00109-017-1557-x","article-title":"Layered defense: How mucus and tight junctions seal the intestinal barrier","volume":"95","author":"Capaldo","year":"2017","journal-title":"J Mol Med (Berl)"},{"key":"B58","doi-asserted-by":"publisher","DOI":"10.1080\/17474124.2017.1343143","article-title":"The intestinal barrier: A fundamental role in health and disease","volume":"11","author":"Vancamelbeke","year":"2017","journal-title":"Expert Rev Gastroenterol Hepatol"},{"key":"B59","doi-asserted-by":"publisher","DOI":"10.1080\/21688370.2016.1251384","article-title":"Zonulin, a regulator of epithelial and endothelial barrier functions, and its involvement in chronic inflammatory diseases","volume":"4","author":"Sturgeon","year":"2016","journal-title":"Tissue Barriers"},{"key":"B60","doi-asserted-by":"publisher","DOI":"10.3390\/cells9081909","article-title":"The intestinal barrier and current techniques for the assessment of gut permeability","volume":"9","author":"Schoultz","year":"2020","journal-title":"Cells"},{"key":"B61","doi-asserted-by":"publisher","DOI":"10.1126\/science.aad0135","article-title":"A gut-vascular barrier controls the systemic dissemination of bacteria","volume":"350","author":"Spadoni","year":"2015","journal-title":"Science"},{"key":"B62","doi-asserted-by":"publisher","DOI":"10.1016\/j.jhep.2019.08.005","article-title":"Microbiota-driven gut vascular barrier disruption is a prerequisite for non-alcoholic steatohepatitis development","volume":"71","author":"Mouries","year":"2019","journal-title":"J Hepatol"},{"key":"B63","doi-asserted-by":"publisher","DOI":"10.1016\/j.clim.2023.109683","article-title":"Gut vascular barrier in the pathogenesis and resolution of Crohn's disease: A novel link from origination to therapy","volume":"253","author":"Jingjie","year":"2023","journal-title":"Clin Immunol"},{"key":"B64","doi-asserted-by":"publisher","DOI":"10.1053\/j.gastro.2006.04.020","article-title":"Muc2-deficient mice spontaneously develop colitis, indicating that Muc2 is critical for colonic protection","volume":"131","author":"Van der Sluis","year":"2006","journal-title":"Gastroenterology"},{"key":"B65","doi-asserted-by":"publisher","DOI":"10.3389\/fnut.2021.751010","article-title":"Antimicrobial peptides in gut health: A review","volume":"8","author":"Gong","year":"2021","journal-title":"Front Nutr"},{"key":"B66","doi-asserted-by":"publisher","DOI":"10.1038\/mi.2013.30","article-title":"Microfold (M) cells: Important immunosurveillance posts in the intestinal epithelium","volume":"6","author":"Mabbott","year":"2013","journal-title":"Mucosal Immunol"},{"key":"B67","doi-asserted-by":"publisher","DOI":"10.4049\/jimmunol.2200204","article-title":"Migrating type 2 dendritic cells prime mucosal Th17 cells specific to small intestinal commensal bacteria","volume":"209","author":"Ngoi","year":"2022","journal-title":"J Immunol"},{"key":"B68","doi-asserted-by":"publisher","first-page":"1995","DOI":"10.1038\/s41467-020-15831-7","article-title":"Targeting zonulin and intestinal epithelial barrier function to prevent onset of arthritis","volume":"11","author":"Tajik","year":"2020","journal-title":"Nat Commun"},{"key":"B69","doi-asserted-by":"publisher","DOI":"10.1038\/mi.2011.39","article-title":"Structure and function relationships in Iga","volume":"4","author":"Woof","year":"2011","journal-title":"Mucosal Immunol"},{"key":"B70","doi-asserted-by":"publisher","DOI":"10.3389\/fimmu.2021.791907","article-title":"Yap\/Taz: Key players for rheumatoid arthritis severity by driving fibroblast like synoviocytes phenotype and fibro-inflammatory response","volume":"12","author":"Caire","year":"2021","journal-title":"Front Immunol"},{"key":"B71","doi-asserted-by":"publisher","DOI":"10.2147\/JIR.S444758","article-title":"The Hippo-Yap signaling pathway in osteoarthritis and rheumatoid arthritis","volume":"17","author":"Li","year":"2024","journal-title":"J Inflammation Res"},{"key":"B72","doi-asserted-by":"publisher","DOI":"10.1073\/pnas.1103345108","article-title":"E-cadherin mediates contact inhibition of proliferation through Hippo signaling-pathway components","volume":"108","author":"Kim","year":"2011","journal-title":"Proc Natl Acad Sci USA"},{"key":"B73","doi-asserted-by":"publisher","first-page":"79","DOI":"10.1038\/s41419-021-03395-3","article-title":"The role of the Hippo pathway in the pathogenesis of inflammatory bowel disease","volume":"12","author":"Xie","year":"2021","journal-title":"Cell Death Dis"},{"key":"B74","doi-asserted-by":"publisher","first-page":"294","DOI":"10.1111\/j.1749-6632.2009.04062.x","article-title":"Epithelial tight junctions in intestinal inflammation","volume":"1165","author":"Schulzke","year":"2009","journal-title":"Ann N Y Acad Sci"},{"key":"B75","doi-asserted-by":"publisher","DOI":"10.1101\/gad.1978810","article-title":"The Hippo signaling pathway restricts the oncogenic potential of an intestinal regeneration program","volume":"24","author":"Cai","year":"2010","journal-title":"Genes Dev"},{"key":"B76","doi-asserted-by":"publisher","first-page":"35","DOI":"10.1016\/j.stem.2017.11.001","article-title":"Yap\/Taz-dependent reprogramming of colonic epithelium links Ecm remodeling to tissue regeneration","volume":"22","author":"Yui","year":"2018","journal-title":"Cell Stem Cell"},{"key":"B77","doi-asserted-by":"publisher","DOI":"10.1016\/j.devcel.2014.06.003","article-title":"The Hippo pathway effector Yap controls patterning and differentiation of airway epithelial progenitors","volume":"30","author":"Mahoney","year":"2014","journal-title":"Dev Cell"},{"key":"B78","doi-asserted-by":"publisher","DOI":"10.1016\/j.cell.2014.06.013","article-title":"Yap\/Taz incorporation in the beta-catenin destruction complex orchestrates the Wnt response","volume":"158","author":"Azzolin","year":"2014","journal-title":"Cell"},{"key":"B79","doi-asserted-by":"publisher","DOI":"10.3389\/fcell.2022.894737","article-title":"The Hippo-Yap\/Taz signaling pathway in intestinal self-renewal and regeneration after injury","volume":"10","author":"Deng","year":"2022","journal-title":"Front Cell Dev Biol"},{"key":"B80","doi-asserted-by":"publisher","DOI":"10.1016\/j.isci.2022.104316","article-title":"Jam-a signals through the Hippo pathway to regulate intestinal epithelial proliferation","volume":"25","author":"Fan","year":"2022","journal-title":"iScience"},{"key":"B81","doi-asserted-by":"publisher","first-page":"3","DOI":"10.20517\/mrr.2025.69","article-title":"An overview of the role of the gut microbiota in rheumatoid arthritis","volume":"5","author":"Poznyak","year":"2026","journal-title":"Microbiome Res Rep"},{"key":"B82","doi-asserted-by":"publisher","DOI":"10.1080\/07853890.2026.2613484","article-title":"Gut microbiota in early and established stages of rheumatoid arthritis: From pathogenesis to promising prevention and treatment","volume":"58","author":"Chen","year":"2026","journal-title":"Ann Med"},{"key":"B83","doi-asserted-by":"publisher","first-page":"43","DOI":"10.1186\/s13073-016-0299-7","article-title":"An expansion of rare lineage intestinal microbes characterizes rheumatoid arthritis","volume":"8","author":"Chen","year":"2016","journal-title":"Genome Med"},{"key":"B84","doi-asserted-by":"publisher","DOI":"10.3390\/ijms19102938","article-title":"Analysis of gut microbiota in rheumatoid arthritis patients: Disease-related dysbiosis and modifications induced by etanercept","volume":"19","author":"Picchianti-Diamanti","year":"2018","journal-title":"Int J Mol Sci"},{"key":"B85","doi-asserted-by":"publisher","first-page":"221","DOI":"10.1186\/s40168-018-0603-4","article-title":"A comparative study of the gut microbiota in immune-mediated inflammatory diseases-does a common dysbiosis exist","volume":"6","author":"Forbes","year":"2018","journal-title":"Microbiome"},{"key":"B86","doi-asserted-by":"publisher","DOI":"10.3389\/fcimb.2019.00440","article-title":"Molecular characterization of circulating microbiome signatures in rheumatoid arthritis","volume":"9","author":"Hammad","year":"2019","journal-title":"Front Cell Infect Microbiol"},{"key":"B87","doi-asserted-by":"publisher","DOI":"10.1016\/j.ard.2025.08.029","article-title":"Short-chain fatty acids and their gut microbial pathways distinguish rheumatoid arthritis in discordant monozygotic twins","volume":"85","author":"Blank","year":"2026","journal-title":"Ann Rheum Dis"},{"key":"B88","doi-asserted-by":"publisher","DOI":"10.1007\/s00284-013-0338-1","article-title":"Analysis of fecal Lactobacillus community structure in patients with early rheumatoid arthritis","volume":"67","author":"Liu","year":"2013","journal-title":"Curr Microbiol"},{"key":"B89","doi-asserted-by":"publisher","DOI":"10.7554\/eLife.01202","article-title":"Expansion of intestinal Prevotella copri correlates with enhanced susceptibility to arthritis","volume":"2","author":"Scher","year":"2013","journal-title":"Elife"},{"key":"B90","doi-asserted-by":"publisher","DOI":"10.3389\/fcimb.2019.00369","article-title":"Characteristics of gut microbiota in patients with rheumatoid arthritis in Shanghai, China","volume":"9","author":"Sun","year":"2019","journal-title":"Front Cell Infect Microbiol"},{"key":"B91","doi-asserted-by":"publisher","DOI":"10.1126\/scitranslmed.abn5166","article-title":"Clonal Iga and Igg autoantibodies from individuals at risk for rheumatoid arthritis identify an arthritogenic strain of Subdoligranulum","volume":"14","author":"Chriswell","year":"2022","journal-title":"Sci Transl Med"},{"key":"B92","doi-asserted-by":"publisher","DOI":"10.3389\/fimmu.2024.1418717","article-title":"Gut Subdoligranulum variabile ameliorates rheumatoid arthritis by promoting Tsg-6 synthesis from joint cells","volume":"15","author":"Li","year":"2024","journal-title":"Front Immunol"},{"key":"B93","doi-asserted-by":"publisher","DOI":"10.1172\/JCI93450","article-title":"Two rheumatoid arthritis-specific autoantigens correlate microbial immunity with autoimmune responses in joints","volume":"127","author":"Pianta","year":"2017","journal-title":"J Clin Invest"},{"key":"B94","doi-asserted-by":"publisher","DOI":"10.1002\/art.40003","article-title":"Evidence of the immune relevance of Prevotella copri, a gut microbe, in patients with rheumatoid arthritis","volume":"69","author":"Pianta","year":"2017","journal-title":"Arthritis Rheumatol"},{"key":"B95","doi-asserted-by":"publisher","DOI":"10.1002\/art.42370","article-title":"Association of antibodies to Prevotella copri in anti-cyclic citrullinated peptide-positive individuals at risk of developing rheumatoid arthritis and in patients with early or established rheumatoid arthritis","volume":"75","author":"Seifert","year":"2023","journal-title":"Arthritis Rheumatol"},{"key":"B96","doi-asserted-by":"publisher","first-page":"5","DOI":"10.1002\/rai2.12025","article-title":"Identification of lipopolysaccharide\u2010binding protein as a novel citrullinated autoantigen in rheumatoid arthritis","volume":"2","author":"Zhao","year":"2022","journal-title":"Rheumatol Autoimmun"},{"key":"B97","doi-asserted-by":"publisher","first-page":"32","DOI":"10.1186\/s13075-023-03013-x","article-title":"Factors associated with the composition of the gut microbiome in patients with established rheumatoid arthritis and its value for predicting treatment responses","volume":"25","author":"Koh","year":"2023","journal-title":"Arthritis Res Ther"},{"key":"B98","doi-asserted-by":"publisher","first-page":"95","DOI":"10.1186\/s13075-023-03069-9","article-title":"Increased gut permeability and intestinal inflammation precede arthritis onset in the adjuvant-induced model of arthritis","volume":"25","author":"Hecquet","year":"2023","journal-title":"Arthritis Res Ther"},{"key":"B99","doi-asserted-by":"publisher","DOI":"10.1038\/s41579-025-01205-7","article-title":"Lactobacilli biology, applications and host interactions","volume":"24","author":"Mej\u00eda-Caballero","year":"2026","journal-title":"Nat Rev Microbiol"},{"key":"B100","doi-asserted-by":"publisher","first-page":"5029","DOI":"10.1038\/s41467-019-13036-1","article-title":"Evaluation of 16s rrna gene sequencing for species and strain-level microbiome analysis","volume":"10","author":"Johnson","year":"2019","journal-title":"Nat Commun"},{"key":"B101","doi-asserted-by":"publisher","first-page":"16816","DOI":"10.1038\/s41598-024-67499-4","article-title":"Optimised human stool sample collection for multi-omic microbiota analysis","volume":"14","author":"Gemmell","year":"2024","journal-title":"Sci Rep"},{"key":"B102","doi-asserted-by":"publisher","first-page":"7668","DOI":"10.1038\/s41598-023-33489-1","article-title":"Shallow shotgun sequencing reduces technical variation in microbiome analysis","volume":"13","author":"La Reau","year":"2023","journal-title":"Sci Rep"},{"key":"B103","doi-asserted-by":"publisher","DOI":"10.1097\/MD.0000000000042596","article-title":"Causal associations between gut microbiota and rheumatoid arthritis: a two-sample mendelian randomization study","volume":"104","author":"Wei","year":"2025","journal-title":"Med (Baltimore)"},{"key":"B104","doi-asserted-by":"publisher","DOI":"10.1002\/jmv.29781","article-title":"Multikingdom characterization of gut microbiota in patients with rheumatoid arthritis and rheumatoid arthritis-associated interstitial lung disease","volume":"96","author":"Xing","year":"2024","journal-title":"J Med Virol"},{"key":"B105","doi-asserted-by":"publisher","DOI":"10.7717\/peerj.13037","article-title":"Alterations of gut fungal microbiota in patients with rheumatoid arthritis","volume":"10","author":"Sun","year":"2022","journal-title":"PeerJ"},{"key":"B106","doi-asserted-by":"publisher","DOI":"10.1136\/annrheumdis-2018-214514","article-title":"Prevotella copri in individuals at risk for rheumatoid arthritis","volume":"78","author":"Alpizar-Rodriguez","year":"2019","journal-title":"Ann Rheum Dis"},{"key":"B107","doi-asserted-by":"publisher","DOI":"10.1016\/j.crmicr.2025.100366","article-title":"Participation of gut microbiota and bacterial translocation in chronic systemic inflammation in recently diagnosed rheumatoid arthritis patients","volume":"8","author":"Dunyach-Remy","year":"2025","journal-title":"Curr Res Microb Sci"},{"key":"B108","doi-asserted-by":"publisher","DOI":"10.3389\/fendo.2018.00022","article-title":"Widely used commercial elisa does not detect precursor of haptoglobin2, but recognizes properdin as a potential second member of the zonulin family","volume":"9","author":"Scheffler","year":"2018","journal-title":"Front Endocrinol (Lausanne)"},{"key":"B109","doi-asserted-by":"publisher","DOI":"10.1371\/journal.pone.0210728","article-title":"Serum zonulin as a marker of intestinal mucosal barrier function: may not be what it seems","volume":"14","author":"Ajamian","year":"2019","journal-title":"PloS One"},{"key":"B110","doi-asserted-by":"publisher","DOI":"10.3390\/nu16183215","article-title":"Gut dysbiosis and dietary interventions in rheumatoid arthritis-a narrative review","volume":"16","author":"Bakinowska","year":"2024","journal-title":"Nutrients"},{"key":"B111","doi-asserted-by":"publisher","DOI":"10.3389\/fimmu.2025.1584023","article-title":"Dual regulatory effects of gut microbiota and their metabolites in rheumatoid arthritis: balancing pathogenic and protective mechanisms","volume":"16","author":"Xie","year":"2025","journal-title":"Front Immunol"},{"key":"B112","doi-asserted-by":"publisher","DOI":"10.3389\/fimmu.2025.1717671","article-title":"Neutrophil extracellular traps in rheumatoid arthritis: pathogenic mechanisms and therapeutic potential","volume":"16","author":"Mao","year":"2025","journal-title":"Front Immunol"},{"key":"B113","doi-asserted-by":"publisher","DOI":"10.3389\/fimmu.2022.1007610","article-title":"The bridge of the gut-joint axis: gut microbial metabolites in rheumatoid arthritis","volume":"13","author":"Xu","year":"2022","journal-title":"Front Immunol"},{"key":"B114","doi-asserted-by":"publisher","DOI":"10.1016\/j.biopha.2023.114985","article-title":"Immunological mechanisms of inflammatory diseases caused by gut microbiota dysbiosis: a review","volume":"164","author":"Zhao","year":"2023","journal-title":"BioMed Pharmacother"},{"key":"B115","doi-asserted-by":"publisher","DOI":"10.14336\/AD.2022.0104","article-title":"Functions of gut microbiota metabolites, current status and future perspectives","volume":"13","author":"Liu","year":"2022","journal-title":"Aging Dis"},{"key":"B116","doi-asserted-by":"publisher","first-page":"499","DOI":"10.1046\/j.1365-2036.1998.00337.x","article-title":"Review article: short chain fatty acids in health and disease","volume":"12","author":"Cook","year":"1998","journal-title":"Aliment Pharmacol Ther"},{"key":"B117","doi-asserted-by":"publisher","DOI":"10.1093\/advances\/nmz127","article-title":"Tryptophan metabolism: a link between the gut microbiota and brain","volume":"11","author":"Gao","year":"2020","journal-title":"Adv Nutr"},{"key":"B118","doi-asserted-by":"publisher","DOI":"10.1128\/mBio.02781-18","article-title":"Gut microbiota metabolite indole propionic acid targets tryptophan biosynthesis in Mycobacterium tuberculosis","volume":"10","author":"Negatu","year":"2019","journal-title":"mBio"},{"key":"B119","doi-asserted-by":"publisher","DOI":"10.1080\/19490976.2024.2393270","article-title":"The interplay between gut microbiota, short-chain fatty acids, and implications for host health and disease","volume":"16","author":"Hays","year":"2024","journal-title":"Gut Microbes"},{"key":"B120","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1007\/s00535-016-1242-9","article-title":"Microbiota metabolite short chain fatty acids, gpcr, and inflammatory bowel diseases","volume":"52","author":"Sun","year":"2017","journal-title":"J Gastroenterol"},{"key":"B121","doi-asserted-by":"publisher","DOI":"10.3390\/life14050559","article-title":"Short-chain fatty acids and human health: from metabolic pathways to current therapeutic implications","volume":"14","author":"Facchin","year":"2024","journal-title":"Life (Basel)"},{"key":"B122","doi-asserted-by":"publisher","DOI":"10.3390\/nu15122760","article-title":"Butyrate protects barrier integrity and suppresses immune activation in a Caco-2\/PBMC co-culture model while HDAC inhibition mimics butyrate in restoring cytokine-induced barrier disruption","volume":"15","author":"Korsten","year":"2023","journal-title":"Nutrients"},{"key":"B123","doi-asserted-by":"publisher","DOI":"10.3390\/cells10071775","article-title":"Butyrate and the intestinal epithelium: modulation of proliferation and inflammation in homeostasis and disease","volume":"10","author":"Salvi","year":"2021","journal-title":"Cells"},{"key":"B124","doi-asserted-by":"publisher","first-page":"31","DOI":"10.1038\/s41413-024-00336-6","article-title":"Gut microbial metabolite targets HDAC3-FOxK1-interferon axis in fibroblast-like synoviocytes to ameliorate rheumatoid arthritis","volume":"12","author":"Chen","year":"2024","journal-title":"Bone Res"},{"key":"B125","doi-asserted-by":"publisher","DOI":"10.3389\/fimmu.2025.1610254","article-title":"Immunomodulatory mechanisms of the gut microbiota and metabolites on regulatory T cells in rheumatoid arthritis","volume":"16","author":"Xu","year":"2025","journal-title":"Front Immunol"},{"key":"B126","doi-asserted-by":"publisher","DOI":"10.1172\/JCI167671","article-title":"Microbiota-dependent indole production stimulates the development of collagen-induced arthritis in mice","volume":"134","author":"Seymour","year":"2023","journal-title":"J Clin Invest"},{"key":"B127","doi-asserted-by":"publisher","DOI":"10.3390\/nu15102411","article-title":"Crosstalk between gut microbiota and bile acids in cholestatic liver disease","volume":"15","author":"Shi","year":"2023","journal-title":"Nutrients"},{"key":"B128","doi-asserted-by":"publisher","DOI":"10.3390\/ijms27052315","article-title":"A potential central hub of histamine in the microbiota-gut-joint axis in rheumatoid arthritis: mechanisms and translational implications","volume":"27","author":"Kong","year":"2026","journal-title":"Int J Mol Sci"},{"key":"B129","doi-asserted-by":"publisher","DOI":"10.3390\/cancers14194758","article-title":"Bacteriocins as potential therapeutic approaches in the treatment of various cancers: a review of in vitro studies","volume":"14","author":"Molujin","year":"2022","journal-title":"Cancers (Basel)"},{"key":"B130","doi-asserted-by":"publisher","DOI":"10.3390\/antibiotics9080462","article-title":"Short chain fatty acids modulate the growth and virulence of pathosymbiont Escherichia coli and host response","volume":"9","author":"Zhang","year":"2020","journal-title":"Antibiotics (Basel)"},{"key":"B131","doi-asserted-by":"publisher","DOI":"10.1038\/s12276-023-01088-9","article-title":"Unconventional immune cells in the gut mucosal barrier: regulation by symbiotic microbiota","volume":"55","author":"Yoo","year":"2023","journal-title":"Exp Mol Med"},{"key":"B132","doi-asserted-by":"publisher","DOI":"10.3390\/microorganisms8101587","article-title":"Gut microbiota and immune system interactions","volume":"8","author":"Yoo","year":"2020","journal-title":"Microorganisms"},{"key":"B133","doi-asserted-by":"publisher","DOI":"10.1016\/j.chom.2022.08.009","article-title":"Human gut microbiota stimulate defined innate immune responses that vary from phylum to strain","volume":"30","author":"Spindler","year":"2022","journal-title":"Cell Host Microbe"},{"key":"B134","doi-asserted-by":"publisher","DOI":"10.1136\/ard.2010.143768","article-title":"Effects of interleukin (IL)-17a and IL-17f in human rheumatoid arthritis synoviocytes","volume":"70","author":"Hot","year":"2011","journal-title":"Ann Rheum Dis"},{"key":"B135","doi-asserted-by":"publisher","DOI":"10.1084\/jem.20192048","article-title":"Interleukin-17 in rheumatoid arthritis: trials and tribulations","volume":"217","author":"Taams","year":"2020","journal-title":"J Exp Med"},{"key":"B136","doi-asserted-by":"publisher","DOI":"10.3389\/fimmu.2024.1413485","article-title":"Crosstalk between gut microbiota and host immune system and its response to traumatic injury","volume":"15","author":"Ullah","year":"2024","journal-title":"Front Immunol"},{"key":"B137","doi-asserted-by":"publisher","first-page":"499","DOI":"10.1016\/j.it.2023.05.002","article-title":"Conditioning of the immune system by the microbiome","volume":"44","author":"Graham","year":"2023","journal-title":"Trends Immunol"},{"key":"B138","doi-asserted-by":"publisher","first-page":"33","DOI":"10.1186\/s13045-024-01541-w","article-title":"Critical role of the gut microbiota in immune responses and cancer immunotherapy","volume":"17","author":"Li","year":"2024","journal-title":"J Hematol Oncol"},{"key":"B139","doi-asserted-by":"publisher","first-page":"492","DOI":"10.1038\/s41422-020-0332-7","article-title":"Interaction between microbiota and immunity in health and disease","volume":"30","author":"Zheng","year":"2020","journal-title":"Cell Res"},{"key":"B140","doi-asserted-by":"publisher","DOI":"10.3389\/fimmu.2024.1312919","article-title":"Augmenting regulatory T cells: new therapeutic strategy for rheumatoid arthritis","volume":"15","author":"Zhang","year":"2024","journal-title":"Front Immunol"},{"key":"B141","doi-asserted-by":"publisher","first-page":"3","DOI":"10.1136\/ard-2022-223356","article-title":"EULAR recommendations for the management of rheumatoid arthritis with synthetic and biological disease-modifying antirheumatic drugs: 2022 update","volume":"82","author":"Smolen","year":"2023","journal-title":"Ann Rheum Dis"},{"key":"B142","doi-asserted-by":"publisher","first-page":"495","DOI":"10.3899\/jrheum.150117","article-title":"Efficacy and safety of subcutaneous and intravenous loading dose regimens of secukinumab in patients with active rheumatoid arthritis: results from a randomized phase II study","volume":"43","author":"Tlustochowicz","year":"2016","journal-title":"J Rheumatol"},{"key":"B143","doi-asserted-by":"publisher","DOI":"10.3389\/fcell.2020.538130","article-title":"The role of the microbiome in driving RA-related autoimmunity","volume":"8","author":"Rooney","year":"2020","journal-title":"Front Cell Dev Biol"},{"key":"B144","doi-asserted-by":"publisher","DOI":"10.1136\/gutjnl-2025-336451","article-title":"Pueraria lobata-derived exosome-like nanovesicles alleviate rheumatoid arthritis via targeting Ruminococcus gnavus phenylethylamine production","volume":"75","author":"Han","year":"2026","journal-title":"Gut"},{"key":"B145","doi-asserted-by":"publisher","first-page":"376","DOI":"10.1186\/s12951-025-03450-7","article-title":"An orally-administered nanotherapeutics with gold nanospheres supplying for rheumatoid arthritis therapy by re-shaping gut microbial tryptophan metabolism","volume":"23","author":"Yang","year":"2025","journal-title":"J Nanobiotechnology"},{"key":"B146","doi-asserted-by":"publisher","DOI":"10.1039\/d5fo01457c","article-title":"A biotin-free diet attenuates the incidence of collagen-induced arthritis and alleviates microbial dysbiosis","volume":"16","author":"Su","year":"2025","journal-title":"Food Funct"},{"key":"B147","doi-asserted-by":"publisher","DOI":"10.1002\/art.42616","article-title":"Alteration of gut microbiota in individuals at high-risk for rheumatoid arthritis associated with disturbed metabolome and the initiation of arthritis through the triggering of mucosal immunity imbalance","volume":"75","author":"Luo","year":"2023","journal-title":"Arthritis Rheumatol"},{"key":"B148","doi-asserted-by":"publisher","DOI":"10.1016\/j.pharep.2013.09.007","article-title":"Partial depletion of natural gut flora by antibiotic aggravates collagen induced arthritis (CIA) in mice","volume":"66","author":"Dorozynska","year":"2014","journal-title":"Pharmacol Rep"},{"key":"B149","doi-asserted-by":"publisher","DOI":"10.1002\/art.40490","article-title":"Modulation of inflammatory arthritis in mice by gut microbiota through mucosal inflammation and autoantibody generation","volume":"70","author":"Jubair","year":"2018","journal-title":"Arthritis Rheumatol"},{"key":"B150","doi-asserted-by":"publisher","first-page":"15613","DOI":"10.1038\/s41598-017-15802-x","article-title":"Alteration of the intestinal microbiome characterizes preclinical inflammatory arthritis in mice and its modulation attenuates established arthritis","volume":"7","author":"Rogier","year":"2017","journal-title":"Sci Rep"},{"key":"B151","doi-asserted-by":"publisher","DOI":"10.1016\/j.immuni.2010.06.001","article-title":"Gut-residing segmented filamentous bacteria drive autoimmune arthritis via T helper 17 cells","volume":"32","author":"Wu","year":"2010","journal-title":"Immunity"},{"key":"B152","doi-asserted-by":"publisher","DOI":"10.1172\/JCI32639","article-title":"Stimulation of Tlr2 and Tlr4 differentially skews the balance of T cells in a mouse model of arthritis","volume":"118","author":"Abdollahi-Roodsaz","year":"2008","journal-title":"J Clin Invest"},{"key":"B153","doi-asserted-by":"publisher","DOI":"10.1002\/art.38773","article-title":"Zap-70 genotype disrupts the relationship between microbiota and host, leading to spondyloarthritis and ileitis in Skg mice","volume":"66","author":"Rehaume","year":"2014","journal-title":"Arthritis Rheumatol"},{"key":"B154","doi-asserted-by":"publisher","DOI":"10.1002\/art.39783","article-title":"Dysbiosis contributes to arthritis development via activation of autoreactive T cells in the intestine","volume":"68","author":"Maeda","year":"2016","journal-title":"Arthritis Rheumatol"},{"key":"B155","doi-asserted-by":"publisher","DOI":"10.3390\/ijms26115099","article-title":"Investigating the role of gut microbiota in the pathogenesis and progression of rheumatoid arthritis in a collagen-induced arthritis mouse model","volume":"26","author":"Belvoncikova","year":"2025","journal-title":"Int J Mol Sci"},{"key":"B156","doi-asserted-by":"publisher","first-page":"1554","DOI":"10.1038\/s41467-019-09361-0","article-title":"The parasitic worm product Es-62 normalises the gut microbiota bone marrow axis in inflammatory arthritis","volume":"10","author":"Doonan","year":"2019","journal-title":"Nat Commun"},{"key":"B157","doi-asserted-by":"publisher","first-page":"73","DOI":"10.1038\/s41401-019-0279-8","article-title":"Rebalancing of the gut flora and microbial metabolism is responsible for the anti-arthritis effect of kaempferol","volume":"41","author":"Aa","year":"2020","journal-title":"Acta Pharmacol Sin"},{"key":"B158","doi-asserted-by":"publisher","DOI":"10.1136\/ard-2023-224491","article-title":"Expression of Hif1alpha in intestinal epithelium restricts arthritis inflammation by inhibiting Ripk3-induced cell death machinery","volume":"83","author":"Lyu","year":"2024","journal-title":"Ann Rheum Dis"},{"key":"B159","doi-asserted-by":"publisher","DOI":"10.3389\/fimmu.2023.1286387","article-title":"Peptoniphilus gorbachii alleviates collagen-induced arthritis in mice by improving intestinal homeostasis and immune regulation","volume":"14","author":"Kim","year":"2023","journal-title":"Front Immunol"},{"key":"B160","doi-asserted-by":"publisher","DOI":"10.3390\/nu11040785","article-title":"Lactobacillus fermentum Pc1 has the capacity to attenuate joint inflammation in collagen-induced arthritis in Dba\/1 mice","volume":"11","author":"Esvaran","year":"2019","journal-title":"Nutrients"},{"key":"B161","doi-asserted-by":"publisher","DOI":"10.1016\/j.micinf.2024.105411","article-title":"Microbiota transfer early after birth modulates genetic susceptibility to chronic arthritis in mice","volume":"27","author":"Borrego","year":"2025","journal-title":"Microbes Infect"},{"key":"B162","doi-asserted-by":"publisher","DOI":"10.1016\/j.semarthrit.2021.04.015","article-title":"Intestinal permeability in spondyloarthritis and rheumatoid arthritis: A systematic review of the literature","volume":"51","author":"Hecquet","year":"2021","journal-title":"Semin Arthritis Rheum"},{"key":"B163","doi-asserted-by":"publisher","DOI":"10.1016\/j.medj.2021.04.013","article-title":"Intestinal barrier dysfunction plays an integral role in arthritis pathology and can be targeted to ameliorate disease","volume":"2","author":"Matei","year":"2021","journal-title":"Med"},{"key":"B164","doi-asserted-by":"publisher","DOI":"10.1152\/physrev.00003.2008","article-title":"Zonulin and its regulation of intestinal barrier function: The biological door to inflammation, autoimmunity, and cancer","volume":"91","author":"Fasano","year":"2011","journal-title":"Physiol Rev"},{"key":"B165","doi-asserted-by":"publisher","DOI":"10.3899\/jrheum.2023-0160","article-title":"Elevated serum levels of zonulin family peptides in anticitrullinated protein antibody-positive at-risk individuals without arthritis","volume":"51","author":"Hemgren","year":"2024","journal-title":"J Rheumatol"},{"key":"B166","doi-asserted-by":"publisher","DOI":"10.1093\/rheumatology\/keac454","article-title":"Rheumatoid arthritis is associated with increased gut permeability and bacterial translocation that are reversed by inflammation control","volume":"62","author":"Audo","year":"2023","journal-title":"Rheumatology"},{"key":"B167","doi-asserted-by":"publisher","DOI":"10.3390\/nu15102386","article-title":"Assessment of intestinal permeability and inflammation bio-markers in patients with rheumatoid arthritis","volume":"15","author":"Heidt","year":"2023","journal-title":"Nutrients"},{"key":"B168","doi-asserted-by":"publisher","DOI":"10.3389\/fimmu.2026.1756621","article-title":"Association between intestinal permeability, systemic inflammation, and response to anti-Tnf therapy in patients with rheumatoid arthritis: A prospective controlled study","volume":"17","author":"Garcia-Studer","year":"2026","journal-title":"Front Immunol"},{"key":"B169","doi-asserted-by":"publisher","DOI":"10.1136\/rmdopen-2024-004752","article-title":"Biomarkers reflecting disturbed gut barrier under treatment with Tnf inhibitors in radiographic axial spondyloarthritis","volume":"10","author":"Rademacher","year":"2024","journal-title":"RMD Open"},{"key":"B170","doi-asserted-by":"publisher","DOI":"10.3389\/fcimb.2023.1092118","article-title":"Gut-joint axis: Gut dysbiosis can contribute to the onset of rheumatoid arthritis via multiple pathways","volume":"13","author":"Romero-Figueroa","year":"2023","journal-title":"Front Cell Infect Microbiol"},{"key":"B171","doi-asserted-by":"publisher","first-page":"59","DOI":"10.1080\/27697061.2023.2211129","article-title":"Microbial imbalance and intestinal permeability in the pathogenesis of rheumatoid arthritis: A mechanism review with a focus on bacterial translocation, citrullination, and probiotic intervention","volume":"43","author":"Blenkinsopp","year":"2024","journal-title":"J Am Nutr Assoc"},{"key":"B172","doi-asserted-by":"publisher","DOI":"10.3389\/fimmu.2023.1117742","article-title":"Brief report: Assessment of mucosal barrier integrity using serological biomarkers in preclinical stages of rheumatoid arthritis","volume":"14","author":"Gilbert","year":"2023","journal-title":"Front Immunol"},{"key":"B173","article-title":"Lipopolysaccharide-binding protein is a sensitive disease activity biomarker for rheumatoid arthritis","volume":"36","author":"Wen","year":"2018","journal-title":"Clin Exp Rheumatol"},{"key":"B174","doi-asserted-by":"publisher","DOI":"10.3389\/fimmu.2025.1577590","article-title":"Profiling of fecal analytes as a potential biomarker in rheumatoid arthritis","volume":"16","author":"Wang","year":"2025","journal-title":"Front Immunol"},{"key":"B175","doi-asserted-by":"publisher","first-page":"14305","DOI":"10.1038\/s41598-018-32675-w","article-title":"Detection and characterization of bacterial nucleic acids in culture-negative synovial tissue and fluid samples from rheumatoid arthritis or osteoarthritis patients","volume":"8","author":"Zhao","year":"2018","journal-title":"Sci Rep"},{"key":"B176","doi-asserted-by":"publisher","DOI":"10.1111\/imm.12950","article-title":"Segmented filamentous bacteria-induced immune responses: A balancing act between host protection and autoimmunity","volume":"154","author":"Flannigan","year":"2018","journal-title":"Immunology"},{"key":"B177","doi-asserted-by":"publisher","DOI":"10.3389\/fimmu.2019.02750","article-title":"Induction of intestinal Th17 cells by flagellins from segmented filamentous bacteria","volume":"10","author":"Wang","year":"2019","journal-title":"Front Immunol"},{"key":"B178","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1038\/s12276-019-0283-6","article-title":"Host-microbiota interactions in rheumatoid arthritis","volume":"51","author":"Maeda","year":"2019","journal-title":"Exp Mol Med"},{"key":"B179","doi-asserted-by":"publisher","DOI":"10.1002\/art.43430","article-title":"Gut microbiota-derived tryptophan indole metabolites ameliorate collagen-induced arthritis in mice via aryl hydrocarbon receptor activation","volume":"78","author":"Wu","year":"2026","journal-title":"Arthritis Rheumatol"},{"key":"B180","doi-asserted-by":"publisher","DOI":"10.1038\/s41584-025-01282-1","article-title":"More than a leaky gut: How gut priming shapes arthritis","volume":"21","author":"Kuhn","year":"2025","journal-title":"Nat Rev Rheumatol"},{"key":"B181","doi-asserted-by":"publisher","DOI":"10.1152\/ajpgi.00113.2021","article-title":"Biomarkers for assessment of intestinal permeability in clinical practice","volume":"321","author":"Seethaler","year":"2021","journal-title":"Am J Physiology-Gastrointestinal Liver Physiol"},{"key":"B182","doi-asserted-by":"publisher","first-page":"34","DOI":"10.1016\/j.jim.2014.02.009","article-title":"Development of monoclonal antibodies to pre-haptoglobin 2 and their use in an enzyme-linked immunosorbent assay (Elisa)","volume":"406","author":"Flanagan","year":"2014","journal-title":"J Immunol Methods"},{"key":"B183","article-title":"Gastrointestinal involvement in rheumatoid arthritis: A biopsy study","volume":"6","author":"Marcolongo","year":"1979","journal-title":"J Rheumatol"},{"key":"B184","doi-asserted-by":"publisher","DOI":"10.1136\/ard.2003.011304","article-title":"Immune activation in the small intestine in patients with rheumatoid arthritis","volume":"63","author":"Nissinen","year":"2004","journal-title":"Ann Rheum Dis"},{"key":"B185","doi-asserted-by":"publisher","first-page":"798","DOI":"10.1002\/eji.202048876","article-title":"Crossing the barriers: Revisiting the gut feeling in rheumatoid arthritis","volume":"51","author":"Brandl","year":"2021","journal-title":"Eur J Immunol"},{"key":"B186","doi-asserted-by":"publisher","DOI":"10.1371\/journal.pone.0338053","article-title":"Evidence of an allostatic response by intestinal tissues following induction of joint inflammation","volume":"21","author":"Moran","year":"2026","journal-title":"PloS One"},{"key":"B187","doi-asserted-by":"publisher","DOI":"10.1016\/j.bbamem.2008.08.027","article-title":"Cytokine regulation of tight junctions","volume":"1788","author":"Capaldo","year":"2009","journal-title":"Biochim Biophys Acta"},{"key":"B188","doi-asserted-by":"publisher","first-page":"135","DOI":"10.1186\/s13075-024-03378-7","article-title":"Stabilizing the integrity of intestinal barrier to control arthritis","volume":"26","author":"Kalliolias","year":"2024","journal-title":"Arthritis Res Ther"},{"key":"B189","doi-asserted-by":"publisher","DOI":"10.1016\/j.celrep.2025.116755","article-title":"Mucosal innate immune activation as the trigger to Prevotella species-induced arthritis in genetically resistant mice","volume":"45","author":"Maeda","year":"2026","journal-title":"Cell Rep"},{"key":"B190","doi-asserted-by":"publisher","DOI":"10.3389\/fmed.2018.00364","article-title":"Il-17 in rheumatoid arthritis and precision medicine: From synovitis expression to circulating bioactive levels","volume":"5","author":"Robert","year":"2018","journal-title":"Front Med (Lausanne)"},{"key":"B191","doi-asserted-by":"publisher","DOI":"10.2139\/ssrn.5472943","article-title":"Association between intestinal permeability, systemic inflammation, and response to anti-Tnf therapy: A prospective controlled study","author":"Garcia Studer","year":"2025"},{"key":"B192","doi-asserted-by":"publisher","DOI":"10.3390\/jcm13010055","article-title":"Tailoring rheumatoid arthritis treatment through a sex and gender lens","volume":"13","author":"Carmona","year":"2023","journal-title":"J Clin Med"},{"key":"B193","doi-asserted-by":"publisher","first-page":"9067","DOI":"10.1038\/s41598-024-59667-3","article-title":"The overlooked evolutionary dynamics of 16s Rrna revises its role as the \"gold standard\" for bacterial species identification","volume":"14","author":"Bartos","year":"2024","journal-title":"Sci Rep"},{"key":"B194","doi-asserted-by":"publisher","DOI":"10.1099\/mgen.0.001203","article-title":"On the limits of 16s Rrna gene-based metagenome prediction and functional profiling","volume":"10","author":"Matchado","year":"2024","journal-title":"Microb Genom"},{"key":"B195","doi-asserted-by":"publisher","DOI":"10.1038\/ajg.2012.211","article-title":"A randomized, double-blind study of larazotide acetate to prevent the activation of celiac disease during gluten challenge","volume":"107","author":"Leffler","year":"2012","journal-title":"Am J Gastroenterol"},{"key":"B196","doi-asserted-by":"publisher","DOI":"10.1136\/annrheumdis-2019-215743","article-title":"Metagenome-wide association study of gut microbiome revealed novel aetiology of rheumatoid arthritis in the Japanese population","volume":"79","author":"Kishikawa","year":"2020","journal-title":"Ann Rheum Dis"},{"key":"B197","doi-asserted-by":"publisher","DOI":"10.18632\/aging.203641","article-title":"Analysis of gut microbiota and metabolites in patients with rheumatoid arthritis and identification of potential biomarkers","volume":"13","author":"Chen","year":"2021","journal-title":"Aging (Albany NY)"},{"key":"B198","doi-asserted-by":"publisher","first-page":"21","DOI":"10.1186\/s13075-023-03260-y","article-title":"Impaired immune tolerance mediated by reduced Tfr cells in rheumatoid arthritis linked to gut microbiota dysbiosis and altered metabolites","volume":"26","author":"Wu","year":"2024","journal-title":"Arthritis Res Ther"},{"key":"B199","doi-asserted-by":"publisher","DOI":"10.1002\/art.39785","article-title":"Suppression of inflammatory arthritis by human gut-derived Prevotella histicola in humanized mice","volume":"68","author":"Marietta","year":"2016","journal-title":"Arthritis Rheumatol"},{"key":"B200","doi-asserted-by":"publisher","DOI":"10.1016\/j.jaut.2020.102564","article-title":"Resistant starch intake alleviates collagen-induced arthritis in mice by modulating gut microbiota and promoting concomitant propionate production","volume":"116","author":"Bai","year":"2021","journal-title":"J Autoimmun"},{"key":"B201","doi-asserted-by":"publisher","first-page":"44","DOI":"10.1186\/s42358-024-00382-y","article-title":"High fat diet increases the severity of collagen-induced arthritis in mice by altering the gut microbial community","volume":"64","author":"Zhang","year":"2024","journal-title":"Adv Rheumatol"}],"container-title":["Frontiers in Immunology"],"original-title":[],"link":[{"URL":"https:\/\/www.frontiersin.org\/articles\/10.3389\/fimmu.2026.1887782\/full","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,7,15]],"date-time":"2026-07-15T06:10:57Z","timestamp":1784095857000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.frontiersin.org\/articles\/10.3389\/fimmu.2026.1887782\/full"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,7,15]]},"references-count":201,"alternative-id":["10.3389\/fimmu.2026.1887782"],"URL":"https:\/\/doi.org\/10.3389\/fimmu.2026.1887782","relation":{},"ISSN":["1664-3224"],"issn-type":[{"value":"1664-3224","type":"electronic"}],"subject":[],"published":{"date-parts":[[2026,7,15]]},"article-number":"1887782"}}