{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,10,14]],"date-time":"2026-10-14T03:07:51Z","timestamp":1791947271973,"version":"4.3.4"},"reference-count":94,"publisher":"Frontiers Media SA","license":[{"start":{"date-parts":[[2023,8,17]],"date-time":"2023-08-17T00:00:00Z","timestamp":1692230400000},"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. Mol. Biosci."],"abstract":"<jats:p>SOX9, a member of the SRY-related HMG-box transcription factors, has been reported to critically regulate fetal development and stem cell homeostasis. Wnt signalling is a highly conserved signalling pathway that controls stem cell fate decision and stemness maintenance throughout embryonic development and adult life. Many studies have shown that the interactions between SOX9 and the canonical Wnt signalling pathway are involved in many of the physiological and pathological processes of stem cells, including organ development, the proliferation, differentiation and stemness maintenance of stem cells, and tumorigenesis. In this review, we summarize the already-known molecular mechanism of cross-interactions between SOX9 and the canonical Wnt signalling pathway, outline its regulatory effects on the maintenance of homeostasis in different types of stem cells, and explore its potential in translational stem cell therapy.<\/jats:p>","DOI":"10.3389\/fmolb.2023.1250530","type":"journal-article","created":{"date-parts":[[2023,8,18]],"date-time":"2023-08-18T04:49:39Z","timestamp":1692334179000},"update-policy":"https:\/\/doi.org\/10.3389\/crossmark-policy","source":"Crossref","is-referenced-by-count":12,"title":["Cross-regulation between SOX9 and the canonical Wnt signalling pathway in stem cells"],"prefix":"10.3389","volume":"10","author":[{"given":"Jiajia","family":"Wang","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Xichen","family":"Wan","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Qihua","family":"Le","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"1965","published-online":{"date-parts":[[2023,8,17]]},"reference":[{"key":"B1","doi-asserted-by":"publisher","first-page":"2493","DOI":"10.1038\/onc.2010.621","article-title":"MiniSOX9, a dominant-negative variant in colon cancer cells","volume":"30","author":"Abdel-Samad","year":"2011","journal-title":"Oncogene"},{"key":"B2","doi-asserted-by":"publisher","first-page":"288","DOI":"10.1038\/cr.2012.11","article-title":"Neural crest stem cells: discovery, properties and potential for therapy","volume":"22","author":"Achilleos","year":"2012","journal-title":"Cell. Res."},{"key":"B3","doi-asserted-by":"publisher","first-page":"1072","DOI":"10.1101\/gad.1171104","article-title":"Interactions between Sox9 and beta-catenin control chondrocyte differentiation","volume":"18","author":"Akiyama","year":"2004","journal-title":"Genes. Dev."},{"key":"B4","doi-asserted-by":"publisher","first-page":"3923","DOI":"10.1038\/ncomms4923","article-title":"Two nested developmental waves demarcate a compartment boundary in the mouse lung","volume":"5","author":"Alanis","year":"2014","journal-title":"Nat. Commun."},{"key":"B5","doi-asserted-by":"publisher","first-page":"e2208623119","DOI":"10.1073\/pnas.2208623119","article-title":"Hypomorphic and dominant-negative impact of truncated SOX9 dysregulates Hedgehog-Wnt signaling, causing campomelia","volume":"120","author":"Au","year":"2023","journal-title":"Proc. Natl. Acad. Sci. U. S. A."},{"key":"B6","doi-asserted-by":"publisher","first-page":"433","DOI":"10.1016\/j.biocel.2009.07.015","article-title":"SOX E genes: SOX9 and SOX8 in mammalian testis development","volume":"42","author":"Barrionuevo","year":"2010","journal-title":"Int. J. Biochem. Cell. Biol."},{"key":"B7","doi-asserted-by":"publisher","first-page":"635","DOI":"10.1083\/jcb.200704152","article-title":"Sox9 regulates cell proliferation and is required for Paneth cell differentiation in the intestinal epithelium","volume":"178","author":"Bastide","year":"2007","journal-title":"J. Cell. Biol."},{"key":"B8","doi-asserted-by":"publisher","first-page":"179","DOI":"10.1038\/ncb0303-179","article-title":"Armadillo\/beta-catenin signals in the nucleus-proof beyond a reasonable doubt?","volume":"5","author":"Bienz","year":"2003","journal-title":"Nat. Cell. Biol."},{"key":"B9","doi-asserted-by":"publisher","first-page":"37","DOI":"10.1083\/jcb.200311021","article-title":"SOX9 is an intestine crypt transcription factor, is regulated by the Wnt pathway, and represses the CDX2 and MUC2 genes","volume":"166","author":"Blache","year":"2004","journal-title":"J. Cell. Biol."},{"key":"B10","doi-asserted-by":"publisher","first-page":"239","DOI":"10.1006\/dbio.2000.9883","article-title":"Phylogeny of the SOX family of developmental transcription factors based on sequence and structural indicators","volume":"227","author":"Bowles","year":"2000","journal-title":"Dev. Biol."},{"key":"B11","doi-asserted-by":"publisher","first-page":"5434","DOI":"10.7150\/thno.73426","article-title":"Transient upregulation of EGR1 signaling enhances kidney repair by activating SOX9+ renal tubular cells","volume":"12","author":"Chen","year":"2022","journal-title":"Theranostics"},{"key":"B12","doi-asserted-by":"publisher","first-page":"2680","DOI":"10.1002\/jbmr.174","article-title":"SOX9 determines RUNX2 transactivity by directing intracellular degradation","volume":"25","author":"Cheng","year":"2010","journal-title":"J. Bone Mineral Res."},{"key":"B13","doi-asserted-by":"publisher","first-page":"823","DOI":"10.1007\/s13770-022-00456-2","article-title":"Distinctive roles of Wnt signaling in chondrogenic differentiation of BMSCs under coupling of pressure and platelet-rich fibrin","volume":"19","author":"Cheng","year":"2022","journal-title":"Tissue Eng. Regen. Med."},{"key":"B14","doi-asserted-by":"publisher","first-page":"485","DOI":"10.1016\/s0168-9525(97)01305-x","article-title":"TCF\/LEF factor earn their wings","volume":"13","author":"Clevers","year":"1997","journal-title":"Trends Genet."},{"key":"B15","doi-asserted-by":"publisher","first-page":"406","DOI":"10.1177\/0022034520969109","article-title":"Med23 regulates Sox9 expression during craniofacial development","volume":"100","author":"Dash","year":"2021","journal-title":"J. Dent. Res."},{"key":"B16","doi-asserted-by":"publisher","first-page":"115409","DOI":"10.1016\/j.bone.2020.115409","article-title":"The development, patterning and evolution of neural crest cell differentiation into cartilage and bone","volume":"137","author":"Dash","year":"2020","journal-title":"Bone"},{"key":"B17","doi-asserted-by":"publisher","first-page":"2193","DOI":"10.1007\/s12015-021-10232-9","article-title":"New hPSC SOX9 and INS reporter cell lines facilitate the observation and optimization of differentiation into insulin-producing cells","volume":"17","author":"Dettmer","year":"2021","journal-title":"Stem Cell. Rev. Rep."},{"key":"B18","doi-asserted-by":"publisher","first-page":"597","DOI":"10.1016\/j.devcel.2011.12.024","article-title":"Sox9 directs hypertrophic maturation and blocks osteoblast differentiation of growth plate chondrocytes","volume":"22","author":"Dy","year":"2012","journal-title":"Dev. Cell."},{"key":"B19","doi-asserted-by":"publisher","first-page":"233","DOI":"10.1016\/s0955-0674(99)80031-3","article-title":"Regulation of LEF-1\/TCF transcription factors by Wnt and other signals","volume":"11","author":"Eastman","year":"1999","journal-title":"Curr. Opin. Cell. Biol."},{"key":"B20","doi-asserted-by":"publisher","first-page":"G1108","DOI":"10.1152\/ajpgi.00004.2009","article-title":"Distinct SOX9 levels differentially mark stem\/progenitor populations and enteroendocrine cells of the small intestine epithelium","volume":"296","author":"Formeister","year":"2009","journal-title":"Am. J. Physiology-Gastrointestinal Liver Physiology"},{"key":"B21","doi-asserted-by":"publisher","first-page":"347","DOI":"10.3390\/cells8040347","article-title":"YAP, \u0394Np63, and \u03b2-catenin signaling pathways are involved in the modulation of corneal epithelial stem cell phenotype induced by substrate stiffness","volume":"8","author":"Gouveia","year":"2019","journal-title":"Cells"},{"key":"B22","doi-asserted-by":"publisher","first-page":"245","DOI":"10.1038\/346245a0","article-title":"A gene mapping to the sex-determining region of the mouse Y chromosome is a member of a novel family of embryonically expressed genes","volume":"346","author":"Gubbay","year":"1990","journal-title":"Nature"},{"key":"B23","doi-asserted-by":"publisher","first-page":"1155","DOI":"10.1007\/s10616-018-0206-8","article-title":"The role of Sox9 in maintaining the characteristics and pluripotency of Arbas Cashmere goat hair follicle stem cells","volume":"70","author":"He","year":"2018","journal-title":"Cytotechnology"},{"key":"B24","doi-asserted-by":"publisher","first-page":"1598","DOI":"10.1016\/j.devcel.2022.05.010","article-title":"R-SPONDIN2(+) mesenchymal cells form the bud tip progenitor niche during human lung development","volume":"57","author":"Hein","year":"2022","journal-title":"Dev. Cell."},{"key":"B25","doi-asserted-by":"publisher","first-page":"727","DOI":"10.1016\/j.devcel.2005.02.013","article-title":"Canonical Wnt\/beta-catenin signaling prevents osteoblasts from differentiating into chondrocytes","volume":"8","author":"Hill","year":"2005","journal-title":"Dev. Cell."},{"key":"B26","doi-asserted-by":"publisher","first-page":"662754","DOI":"10.3389\/fcell.2021.662754","article-title":"Between fate choice and self-renewal-heterogeneity of adult neural crest-derived stem cells","volume":"9","author":"Hoving","year":"2021","journal-title":"Front. Cell. Dev. Biol."},{"key":"B27","doi-asserted-by":"publisher","first-page":"27702","DOI":"10.1074\/jbc.M110.108498","article-title":"Stage-specific control of connective tissue growth factor (CTGF\/CCN2) expression in chondrocytes by Sox9 and beta-catenin","volume":"285","author":"Huang","year":"2010","journal-title":"J. Biol. Chem."},{"key":"B28","doi-asserted-by":"publisher","first-page":"60","DOI":"10.1186\/s13287-017-0507-4","article-title":"Sox9 transcriptionally regulates Wnt signaling in intestinal epithelial stem cells in hypomethylated crypts in the diabetic state","volume":"8","author":"Huang","year":"2017","journal-title":"Stem Cell. Res. Ther."},{"key":"B29","doi-asserted-by":"publisher","first-page":"143","DOI":"10.1186\/s12967-019-1895-2","article-title":"SOX9 drives the epithelial-mesenchymal transition in non-small-cell lung cancer through the Wnt\/-catenin pathway","volume":"17","author":"Huang","year":"2019","journal-title":"J. Transl. Med."},{"key":"B30","doi-asserted-by":"publisher","first-page":"10398","DOI":"10.1038\/srep10398","article-title":"SOXE transcription factors form selective dimers on non-compact DNA motifs through multifaceted interactions between dimerization and high-mobility group domains","volume":"5","author":"Huang","year":"2015","journal-title":"Sci. Rep."},{"key":"B31","doi-asserted-by":"publisher","first-page":"778","DOI":"10.1093\/stmcls\/sxac038","article-title":"Sox9 and Lef1 regulate the fate and behavior of airway glandular progenitors in response to injury","volume":"40","author":"Ievlev","year":"2022","journal-title":"Stem Cells"},{"key":"B32","doi-asserted-by":"publisher","first-page":"113789","DOI":"10.1016\/j.bcp.2019.113789","article-title":"SOX9: the master regulator of cell fate in breast cancer","volume":"174","author":"Jana","year":"2020","journal-title":"Biochem. Pharmacol."},{"key":"B33","doi-asserted-by":"crossref","first-page":"353","DOI":"10.1016\/S1016-8478(23)17431-0","article-title":"BMP-2-enhanced chondrogenesis involves p38 MAPK-mediated down-regulation of Wnt-7a pathway","volume":"22","author":"Jin","year":"2006","journal-title":"Mol. Cells"},{"key":"B34","doi-asserted-by":"publisher","first-page":"149","DOI":"10.1016\/j.gendis.2014.09.004","article-title":"The versatile functions of Sox9 in development, stem cells, and human diseases","volume":"1","author":"Jo","year":"2014","journal-title":"Genes. Dis."},{"key":"B35","doi-asserted-by":"publisher","first-page":"583","DOI":"10.1038\/s41586-021-03819-2","article-title":"Highly accurate protein structure prediction with AlphaFold","volume":"596","author":"Jumper","year":"2021","journal-title":"Nature"},{"key":"B36","doi-asserted-by":"publisher","first-page":"4129","DOI":"10.1242\/dev.091793","article-title":"Sox proteins: regulators of cell fate specification and differentiation","volume":"140","author":"Kamachi","year":"2013","journal-title":"Development"},{"key":"B37","doi-asserted-by":"publisher","first-page":"30489","DOI":"10.1038\/srep30489","article-title":"SOX9 is a novel cancer stem cell marker surrogated by osteopontin in human hepatocellular carcinoma","volume":"6","author":"Kawai","year":"2016","journal-title":"Sci. Rep."},{"key":"B38","doi-asserted-by":"publisher","first-page":"3960","DOI":"10.1038\/s41467-022-31697-3","article-title":"Mast4 determines the cell fate of MSCs for bone and cartilage development","volume":"13","author":"Kim","year":"2022","journal-title":"Nat. Commun."},{"key":"B39","doi-asserted-by":"publisher","first-page":"56","DOI":"10.1002\/dvdy.22046","article-title":"Interactions between SOX factors and Wnt\/beta-catenin signaling in development and disease","volume":"239","author":"Kormish","year":"2010","journal-title":"Dev. Dyn."},{"key":"B40","doi-asserted-by":"publisher","first-page":"1419","DOI":"10.1016\/j.celrep.2014.07.038","article-title":"Fibroblast growth factor maintains chondrogenic potential of limb bud mesenchymal cells by modulating DNMT3A recruitment","volume":"8","author":"Kumar","year":"2014","journal-title":"Cell. Rep."},{"key":"B41","doi-asserted-by":"publisher","first-page":"1015","DOI":"10.1016\/j.stemcr.2023.02.004","article-title":"Wnt signaling from Gli1-expressing apical stem\/progenitor cells is essential for\u00a0the coordination of tooth root development","volume":"18","author":"Lav","year":"2023","journal-title":"Stem Cell. Rep."},{"key":"B42","doi-asserted-by":"publisher","first-page":"93","DOI":"10.1016\/j.ydbio.2004.07.036","article-title":"Early requirement of the transcriptional activator Sox9 for neural crest specification in Xenopus","volume":"275","author":"Lee","year":"2004","journal-title":"Dev. Biol."},{"key":"B43","doi-asserted-by":"publisher","first-page":"32","DOI":"10.1007\/s11914-016-0296-1","article-title":"SOXC genes and the control of skeletogenesis","volume":"14","author":"Lefebvre","year":"2016","journal-title":"Curr. Osteoporos. Rep."},{"key":"B44","doi-asserted-by":"publisher","first-page":"29371","DOI":"10.18632\/oncotarget.8835","article-title":"Sox9 confers stemness properties in hepatocellular carcinoma through Frizzled-7 mediated Wnt\/\u03b2-catenin signaling","volume":"7","author":"Leung","year":"2016","journal-title":"Oncotarget"},{"key":"B45","doi-asserted-by":"publisher","first-page":"1860","DOI":"10.7150\/thno.83353","article-title":"Histone H3K27 methyltransferase EZH2 regulates apoptotic and inflammatory responses in sepsis-induced AKI","volume":"13","author":"Li","year":"2023","journal-title":"Theranostics"},{"key":"B46","doi-asserted-by":"publisher","first-page":"209","DOI":"10.1053\/j.gastro.2021.09.044","article-title":"An enhancer-driven stem cell-like program mediated by SOX9 blocks intestinal differentiation in colorectal cancer","volume":"162","author":"Liang","year":"2022","journal-title":"Gastroenterology"},{"key":"B47","doi-asserted-by":"publisher","first-page":"2882","DOI":"10.1073\/pnas.1211747110","article-title":"Phosphorylation of Sox9 is required for neural crest delamination and is regulated downstream of BMP and canonical Wnt signaling","volume":"110","author":"Liu","year":"2013","journal-title":"Proc. Natl. Acad. Sci. U. S. A."},{"key":"B48","doi-asserted-by":"publisher","first-page":"123","DOI":"10.1007\/s11010-014-1963-0","article-title":"Overexpression of Wnt11 promotes chondrogenic differentiation of bone marrow-derived mesenchymal stem cells in synergism with TGF-\u03b2","volume":"390","author":"Liu","year":"2014","journal-title":"Mol. Cell. Biochem."},{"key":"B49","doi-asserted-by":"publisher","first-page":"2885","DOI":"10.1038\/s41388-022-02313-x","article-title":"Wnt-TCF7-SOX9 axis promotes cholangiocarcinoma proliferation and pemigatinib resistance in a FGF7-FGFR2 autocrine pathway","volume":"41","author":"Liu","year":"2022","journal-title":"Oncogene"},{"key":"B50","doi-asserted-by":"publisher","first-page":"199","DOI":"10.1007\/s00432-003-0431-0","article-title":"The Wnt signaling pathway and its role in tumor development","volume":"129","author":"Lustig","year":"2003","journal-title":"J. Cancer Res. Clin. Oncol."},{"key":"B51","doi-asserted-by":"publisher","first-page":"1745","DOI":"10.1172\/JCI78815","article-title":"SOX9 drives WNT pathway activation in prostate cancer","volume":"126","author":"Ma","year":"2016","journal-title":"J. Clin. Investigation"},{"key":"B52","doi-asserted-by":"publisher","first-page":"466","DOI":"10.1038\/embor.2010.61","article-title":"SOX5 controls cell cycle progression in neural progenitors by interfering with the WNT-beta-catenin pathway","volume":"11","author":"Martinez-Morales","year":"2010","journal-title":"EMBO Rep."},{"key":"B53","doi-asserted-by":"publisher","first-page":"72","DOI":"10.1016\/j.biocel.2011.09.008","article-title":"SOX9 regulates endocrine cell differentiation during human fetal pancreas development","volume":"44","author":"Mcdonald","year":"2012","journal-title":"Int. J. Biochem. Cell. Biol."},{"key":"B54","doi-asserted-by":"publisher","first-page":"2197054","DOI":"10.1080\/08941939.2023.2197054","article-title":"SOX9 inhibits the progression of osteonecrosis of the femoral head via the activation of the wnt\/beta-catenin pathway","volume":"36","author":"Meng","year":"2023","journal-title":"J. Investig. Surg."},{"key":"B55","doi-asserted-by":"publisher","first-page":"10268","DOI":"10.1038\/s41598-018-28596-3","article-title":"Transcription factor profiling identifies Sox9 as regulator of proliferation and differentiation in corneal epithelial stem\/progenitor cells","volume":"8","author":"Menzel-Severing","year":"2018","journal-title":"Sci. Rep."},{"key":"B56","doi-asserted-by":"publisher","first-page":"110200","DOI":"10.1016\/j.cellsig.2021.110200","article-title":"ZRANB1 enhances stem-cell-like features and accelerates tumor progression by regulating Sox9-mediated USP22\/Wnt\/\u03b2-catenin pathway in colorectal cancer","volume":"90","author":"Miao","year":"2022","journal-title":"Cell. Signal"},{"key":"B57","doi-asserted-by":"publisher","first-page":"3221","DOI":"10.1002\/jcp.25802","article-title":"The cross-regulation between SOX15 and Wnt signaling pathway","volume":"232","author":"Moradi","year":"2017","journal-title":"J. Cell. Physiol."},{"key":"B58","doi-asserted-by":"publisher","first-page":"539","DOI":"10.1053\/j.gastro.2007.05.020","article-title":"SOX9 is required for the differentiation of paneth cells in the intestinal epithelium","volume":"133","author":"Mori-Akiyama","year":"2007","journal-title":"Gastroenterology"},{"key":"B59","doi-asserted-by":"publisher","first-page":"4734","DOI":"10.1167\/iovs.10-6486","article-title":"Wnt\/\u03b2-catenin signaling regulates proliferation of human cornea epithelial stem\/progenitor cells","volume":"52","author":"Nakatsu","year":"2011","journal-title":"Investig. Ophthalmol. Vis. Sci."},{"key":"B61","doi-asserted-by":"publisher","DOI":"10.1242\/dmm.046573","article-title":"Disruption of a Hedgehog-Foxf1-Rspo2 signaling axis leads to tracheomalacia and a loss of Sox9+ tracheal chondrocytes","volume":"14","author":"Nasr","year":"2021","journal-title":"Dis. Models Mech."},{"key":"B62","doi-asserted-by":"publisher","first-page":"985","DOI":"10.1016\/j.cell.2017.05.016","article-title":"Wnt\/\u03b2-Catenin signaling, disease, and emerging therapeutic modalities","volume":"169","author":"Nusse","year":"2017","journal-title":"Cell."},{"key":"B63","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1242\/dev.160788","article-title":"beta-Catenin maintains lung epithelial progenitors after lung specification","volume":"145","author":"Ostrin","year":"2018","journal-title":"Development"},{"key":"B64","doi-asserted-by":"publisher","first-page":"156","DOI":"10.1016\/j.cell.2015.11.058","article-title":"WNT-SHH antagonism specifies and expands stem cells prior to niche formation","volume":"164","author":"Ouspenskaia","year":"2016","journal-title":"Cell."},{"key":"B65","doi-asserted-by":"publisher","first-page":"1642","DOI":"10.1002\/stem.1962","article-title":"microRNA-103\/107 family regulates multiple epithelial stem cell characteristics","volume":"33","author":"Peng","year":"2015","journal-title":"Stem Cells"},{"key":"B66","doi-asserted-by":"publisher","first-page":"82228","DOI":"10.18632\/oncotarget.10573","article-title":"SOX9 is an atypical intestinal tumor suppressor controlling the oncogenic Wnt\/\u00df-catenin signaling","volume":"7","author":"Prevostel","year":"2016","journal-title":"Oncotarget"},{"key":"B67","doi-asserted-by":"publisher","first-page":"2563","DOI":"10.1101\/gad.227785.113","article-title":"VHL-mediated disruption of Sox9 activity compromises beta-cell identity and results in diabetes mellitus","volume":"27","author":"Puri","year":"2013","journal-title":"Genes. & Dev."},{"key":"B68","doi-asserted-by":"publisher","first-page":"1254","DOI":"10.1038\/s41374-021-00611-8","article-title":"Adipose mesenchymal stem cell-derived exosomes accelerate skin wound healing via the lncRNA H19\/miR-19b\/SOX9 axis","volume":"101","author":"Qian","year":"2021","journal-title":"Lab. Investig."},{"key":"B69","doi-asserted-by":"publisher","first-page":"102735","DOI":"10.1016\/j.jbc.2022.102735","article-title":"SOX9 and TCF transcription factors associate to mediate Wnt\/\u03b2-catenin target gene activation in colorectal cancer","volume":"299","author":"Ramakrishnan","year":"2023","journal-title":"J. Biol. Chem."},{"key":"B70","doi-asserted-by":"publisher","first-page":"571","DOI":"10.1146\/annurev-biochem-040320-103615","article-title":"The Wnt pathway: from signaling mechanisms to synthetic modulators","volume":"91","author":"Rim","year":"2022","journal-title":"Annu. Rev. Biochem."},{"key":"B71","doi-asserted-by":"publisher","first-page":"E4456","DOI":"10.1073\/pnas.1311847110","article-title":"Sox9 plays multiple roles in the lung epithelium during branching morphogenesis","volume":"110","author":"Rockich","year":"2013","journal-title":"Proc. Natl. Acad. Sci. U. S. A."},{"key":"B72","doi-asserted-by":"publisher","first-page":"167","DOI":"10.1016\/s1534-5807(02)00223-x","article-title":"Twenty pairs of sox: extent, homology, and nomenclature of the mouse and human sox transcription factor gene families","volume":"3","author":"Schepers","year":"2002","journal-title":"Dev. Cell."},{"key":"B73","doi-asserted-by":"publisher","first-page":"1666","DOI":"10.1016\/j.bbamcr.2012.06.027","article-title":"KLF4 and SOX9 transcription factors antagonize \u03b2-catenin and inhibit TCF-activity in cancer cells","volume":"1823","author":"Sellak","year":"2012","journal-title":"Biochim. Biophys. Acta"},{"key":"B74","doi-asserted-by":"publisher","first-page":"557","DOI":"10.1002\/stem.2511","article-title":"Neurotrophic factor-alpha 1: a key wnt-beta-catenin dependent anti-proliferation factor and ERK-sox9 activated inducer of embryonic neural stem cell differentiation to astrocytes in neurodevelopment","volume":"35","author":"Selvaraj","year":"2017","journal-title":"Stem Cells"},{"key":"B75","doi-asserted-by":"publisher","first-page":"858782","DOI":"10.3389\/fonc.2022.858782","article-title":"Phosphorylation-dependent regulation of WNT\/Beta-Catenin signaling","volume":"12","author":"Shah","year":"2022","journal-title":"Front. Oncol."},{"key":"B76","doi-asserted-by":"publisher","first-page":"547","DOI":"10.1016\/j.ejcb.2015.08.002","article-title":"SOX family transcription factors involved in diverse cellular events during development","volume":"94","author":"She","year":"2015","journal-title":"Eur. J. Cell. Biol."},{"key":"B77","doi-asserted-by":"publisher","first-page":"240","DOI":"10.1038\/346240a0","article-title":"A gene from the human sex-determining region encodes a protein with homology to a conserved DNA-binding motif","volume":"346","author":"Sinclair","year":"1990","journal-title":"Nature"},{"key":"B78","doi-asserted-by":"publisher","first-page":"eabe0849","DOI":"10.1126\/sciadv.abe0849","article-title":"Repression of Wnt\/\u03b2-catenin signaling by SOX9 and Mastermind-like transcriptional coactivator 2","volume":"7","author":"Sinha","year":"2021","journal-title":"Sci. Adv."},{"key":"B79","doi-asserted-by":"publisher","first-page":"56","DOI":"10.1016\/j.ydbio.2015.06.009","article-title":"Endodermal Wnt signaling is required for tracheal cartilage formation","volume":"405","author":"Snowball","year":"2015","journal-title":"Dev. Biol."},{"key":"B80","doi-asserted-by":"publisher","first-page":"243","DOI":"10.1002\/bdrc.20195","article-title":"Noncanonical Wnt signaling in vertebrate development, stem cells, and diseases","volume":"90","author":"Sugimura","year":"2010","journal-title":"Birth Defects Res. C Embryo Today"},{"key":"B81","doi-asserted-by":"publisher","first-page":"e111338","DOI":"10.15252\/embj.2022111338","article-title":"SOX9 maintains human foetal lung tip progenitor state by enhancing WNT and RTK signalling","volume":"41","author":"Sun","year":"2022","journal-title":"Embo J."},{"key":"B82","doi-asserted-by":"publisher","first-page":"3323","DOI":"10.1074\/jbc.M808048200","article-title":"Sox9 inhibits Wnt signaling by promoting beta-catenin phosphorylation in the nucleus","volume":"284","author":"Topol","year":"2009","journal-title":"J. Biol. Chem."},{"key":"B83","doi-asserted-by":"publisher","first-page":"590","DOI":"10.1002\/dvdy.24393","article-title":"\u03b2-catenin and Kras\/Foxm1 signaling pathway are critical to restrict Sox9 in basal cells during pulmonary branching morphogenesis","volume":"245","author":"Ustiyan","year":"2016","journal-title":"Dev. Dyn."},{"key":"B84","doi-asserted-by":"publisher","first-page":"D439","DOI":"10.1093\/nar\/gkab1061","article-title":"AlphaFold protein structure database: massively expanding the structural coverage of protein-sequence space with high-accuracy models","volume":"50","author":"Varadi","year":"2022","journal-title":"Nucleic Acids Res."},{"key":"B85","doi-asserted-by":"publisher","first-page":"22","DOI":"10.1186\/scrt113","article-title":"SOX9 gene transfer via safe, stable, replication-defective recombinant adeno-associated virus vectors as a novel, powerful tool to enhance the chondrogenic potential of human mesenchymal stem cells","volume":"3","author":"Venkatesan","year":"2012","journal-title":"Stem Cell. Res. Ther."},{"key":"B86","volume-title":"Corneal development: role of the periocular mesenchyme and bi-directional signalling","author":"Walker","year":"2020"},{"key":"B87","doi-asserted-by":"publisher","first-page":"6478","DOI":"10.1074\/jbc.M112.419184","article-title":"SOX9 regulates low density lipoprotein receptor-related protein 6 (LRP6) and T-cell factor 4 (TCF4) expression and Wnt\/\u03b2-catenin activation in breast cancer","volume":"288","author":"Wang","year":"2013","journal-title":"J. Biol. Chem."},{"key":"B88","doi-asserted-by":"publisher","first-page":"15008","DOI":"10.1038\/ncomms15008","article-title":"HMGA1 amplifies Wnt signalling and expands the intestinal stem cell compartment and Paneth cell niche","volume":"8","author":"Xian","year":"2017","journal-title":"Nat. Commun."},{"key":"B89","doi-asserted-by":"publisher","first-page":"230","DOI":"10.1016\/j.jgg.2021.09.008","article-title":"Sox9+ cells are required for salivary gland regeneration after radiation damage via the Wnt\/\u03b2-catenin pathway","volume":"49","author":"Xu","year":"2022","journal-title":"J. Genet. Genomics"},{"key":"B90","doi-asserted-by":"publisher","first-page":"59","DOI":"10.1186\/s13287-019-1154-8","article-title":"microRNA-690 regulates induced pluripotent stem cells (iPSCs) differentiation into insulin-producing cells by targeting Sox9","volume":"10","author":"Xu","year":"2019","journal-title":"Stem Cell. Res. Ther."},{"key":"B91","doi-asserted-by":"publisher","first-page":"e10567","DOI":"10.7554\/eLife.10567","article-title":"Embryonic attenuated Wnt\/\u03b2-catenin signaling defines niche location and long-term stem cell fate in hair follicle","volume":"4","author":"Xu","year":"2015","journal-title":"Elife"},{"key":"B92","doi-asserted-by":"publisher","first-page":"166","DOI":"10.1016\/j.ydbio.2012.09.006","article-title":"FGF signaling transforms non-neural ectoderm into neural crest","volume":"372","author":"Yardley","year":"2012","journal-title":"Dev. Biol."},{"key":"B93","doi-asserted-by":"publisher","first-page":"1020","DOI":"10.1002\/stem.3192","article-title":"Wnt7b-induced Sox11 functions enhance self-renewal and osteogenic commitment of bone marrow mesenchymal stem cells","volume":"38","author":"Yu","year":"2020","journal-title":"Stem Cells"},{"key":"B94","doi-asserted-by":"publisher","first-page":"2194","DOI":"10.1002\/1878-0261.12560","article-title":"A SOX9-AS1\/miR-5590-3p\/SOX9 positive feedback loop drives tumor growth and metastasis in hepatocellular carcinoma through the Wnt\/\u03b2-catenin pathway","volume":"13","author":"Zhang","year":"2019","journal-title":"Mol. Oncol."},{"key":"B95","doi-asserted-by":"publisher","first-page":"eabn9009","DOI":"10.1126\/scisignal.abn9009","article-title":"Yap and Taz promote osteogenesis and prevent chondrogenesis in neural crest cells in vitro and in vivo","volume":"15","author":"Zhao","year":"2022","journal-title":"Sci. Signal"}],"container-title":["Frontiers in Molecular Biosciences"],"original-title":[],"link":[{"URL":"https:\/\/www.frontiersin.org\/articles\/10.3389\/fmolb.2023.1250530\/full","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2024,10,26]],"date-time":"2024-10-26T09:27:01Z","timestamp":1729934821000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.frontiersin.org\/articles\/10.3389\/fmolb.2023.1250530\/full"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2023,8,17]]},"references-count":94,"alternative-id":["10.3389\/fmolb.2023.1250530"],"URL":"https:\/\/doi.org\/10.3389\/fmolb.2023.1250530","relation":{},"ISSN":["2296-889X"],"issn-type":[{"value":"2296-889X","type":"electronic"}],"subject":[],"published":{"date-parts":[[2023,8,17]]},"article-number":"1250530"}}