{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,6,4]],"date-time":"2026-06-04T22:13:49Z","timestamp":1780611229765,"version":"3.54.1"},"reference-count":63,"publisher":"The Company of Biologists","issue":"10","license":[{"start":{"date-parts":[[2026,5,15]],"date-time":"2026-05-15T00:00:00Z","timestamp":1778803200000},"content-version":"vor","delay-in-days":0,"URL":"http:\/\/www.biologists.com\/user-licence-1-2\/"}],"funder":[{"DOI":"10.13039\/501100001691","name":"Japan Society for the Promotion of Science","doi-asserted-by":"crossref","award":["21H03107"],"award-info":[{"award-number":["21H03107"]}],"id":[{"id":"10.13039\/501100001691","id-type":"DOI","asserted-by":"crossref"}]},{"DOI":"10.13039\/501100001691","name":"Japan Society for the Promotion of Science","doi-asserted-by":"crossref","award":["25KJ1878"],"award-info":[{"award-number":["25KJ1878"]}],"id":[{"id":"10.13039\/501100001691","id-type":"DOI","asserted-by":"crossref"}]},{"DOI":"10.13039\/501100001691","name":"Japan Society for the Promotion of Science","doi-asserted-by":"crossref","award":["24K02076"],"award-info":[{"award-number":["24K02076"]}],"id":[{"id":"10.13039\/501100001691","id-type":"DOI","asserted-by":"crossref"}]},{"DOI":"10.13039\/501100001691","name":"Japan Society for the Promotion of Science","doi-asserted-by":"crossref","award":["24K02611"],"award-info":[{"award-number":["24K02611"]}],"id":[{"id":"10.13039\/501100001691","id-type":"DOI","asserted-by":"crossref"}]},{"DOI":"10.13039\/501100001691","name":"Japan Society for the Promotion of Science","doi-asserted-by":"crossref","award":["21KK0161"],"award-info":[{"award-number":["21KK0161"]}],"id":[{"id":"10.13039\/501100001691","id-type":"DOI","asserted-by":"crossref"}]},{"DOI":"10.13039\/501100002241","name":"Japan Science and Technology Agency","doi-asserted-by":"crossref","award":["JPMJSP2132"],"award-info":[{"award-number":["JPMJSP2132"]}],"id":[{"id":"10.13039\/501100002241","id-type":"DOI","asserted-by":"crossref"}]},{"DOI":"10.13039\/100016845","name":"Institute for Frontier Life and Medical Sciences, Kyoto University","doi-asserted-by":"crossref","id":[{"id":"10.13039\/100016845","id-type":"DOI","asserted-by":"crossref"}]}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":[],"published-print":{"date-parts":[[2026,5,15]]},"abstract":"<jats:title>ABSTRACT<\/jats:title>\n                  <jats:p>Scleraxis (Scx) is a basic helix-loop-helix transcription factor predominantly expressed in tendons, ligaments, and their attachment sites, where it regulates maturation. Here, we report a novel Scx enhancer that drives tissue-specific expression. In transgenic reporter mice, a 5.3\u2005kb downstream Scx enhancer (dSE) conferred robust, stable, and faithful reporter activity reproducing the endogenous Scx expression pattern. Within the dSE, we identified a 343\u2005bp Scx enhancer (CSE), conserved from lobe-finned fishes to tetrapods, that recapitulates Scx expression in limbs. In developing limbs, CSE-deficient mice exhibited a marked reduction of endogenous Scx expression and a complete loss of the deltoid tuberosity (DT) supporting forelimb leverage, accompanied by attenuated BMP and TGF\u03b2 signaling, as in Scx-deficient mice. Precise enhancer activation within a particular developmental window was indispensable for DT formation. In contrast, tendon\/ligament maturation retained prolonged plasticity, with Scx expression partially recovering later, likely through additional elements within the dSE. These findings uncover an essential role of the CSE in DT morphogenesis and reveal distinct temporal enhancer requirements for DT versus tendon\/ligament formation, with additional enhancer dependence confined to tendons\/ligaments.<\/jats:p>","DOI":"10.1242\/dev.205325","type":"journal-article","created":{"date-parts":[[2026,4,24]],"date-time":"2026-04-24T07:41:36Z","timestamp":1777016496000},"source":"Crossref","is-referenced-by-count":1,"title":["Divergent temporal control of deltoid tuberosity and limb tendon development by an evolutionarily conserved scleraxis enhancer"],"prefix":"10.1242","volume":"153","author":[{"given":"Shinsei","family":"Yambe","sequence":"first","affiliation":[{"name":"Biomedical Sciences Major, Graduate School of Biomedical and Health Sciences, Hiroshima University 1 Department of Molecular Biology and Biochemistry , , Hiroshima 734-8553 ,","place":["Japan"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-8785-5439","authenticated-orcid":false,"given":"Kenta","family":"Sumiyama","sequence":"additional","affiliation":[{"name":"Graduate School of Bioagricultural Sciences, Nagoya University 2 Department of Animal Sciences , , 464-8601 Nagoya ,","place":["Japan"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Hitomi","family":"Watanabe","sequence":"additional","affiliation":[{"name":"Institute for Life and Medical Sciences, Kyoto University 3 Laboratory of Integrative Biological Science , , Kyoto 606-8507 ,","place":["Japan"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-4963-1693","authenticated-orcid":false,"given":"Aki","family":"Takimoto","sequence":"additional","affiliation":[{"name":"Institute for Frontier Medical Sciences, Kyoto University 4 Department of Cellular Differentiation , , Kyoto 606-8507 ,","place":["Japan"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-0514-9931","authenticated-orcid":false,"given":"Takako","family":"Sasaki","sequence":"additional","affiliation":[{"name":"Oita University 5 Department of Pharmacology, Faculty of Medicine , , Oita 879-5593 ,","place":["Japan"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Gen","family":"Kondoh","sequence":"additional","affiliation":[{"name":"Institute for Life and Medical Sciences, Kyoto University 3 Laboratory of Integrative Biological Science , , Kyoto 606-8507 ,","place":["Japan"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-7588-1290","authenticated-orcid":false,"given":"Denitsa","family":"Docheva","sequence":"additional","affiliation":[{"name":"Orthopaedic Hospital K\u00f6nig-Ludwig-Haus, University of Wuerzburg 6 Department of Musculoskeletal Tissue Regeneration , , 97076 W\u00fcrzburg ,","place":["Germany"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-5616-6761","authenticated-orcid":false,"given":"Chisa","family":"Shukunami","sequence":"additional","affiliation":[{"name":"Biomedical Sciences Major, Graduate School of Biomedical and Health Sciences, Hiroshima University 1 Department of Molecular Biology and Biochemistry , , Hiroshima 734-8553 ,","place":["Japan"]}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"237","published-online":{"date-parts":[[2026,6,1]]},"reference":[{"key":"2026060111291923400_DEV205325C61","doi-asserted-by":"publisher","first-page":"4806","DOI":"10.1074\/jbc.M109.036681","article-title":"Estrogen down-regulation of the Scx gene is mediated by the opposing strand-overlapping gene Bop1","volume":"285","author":"Arao","year":"2010","journal-title":"J Biol Chem"},{"key":"2026060111291923400_DEV205325C1","doi-asserted-by":"publisher","first-page":"363","DOI":"10.1016\/j.ydbio.2006.06.002","article-title":"Tgfbr2 regulates the maintenance of boundaries in the axial skeleton","volume":"296","author":"Baffi","year":"2006","journal-title":"Dev. Biol."},{"key":"2026060111291923400_DEV205325C2","doi-asserted-by":"publisher","first-page":"861","DOI":"10.1016\/j.devcel.2009.10.010","article-title":"Bone ridge patterning during musculoskeletal assembly is mediated through SCX regulation of Bmp4 at the tendon-skeleton junction","volume":"17","author":"Blitz","year":"2009","journal-title":"Dev. Cell"},{"key":"2026060111291923400_DEV205325C3","doi-asserted-by":"publisher","first-page":"2680","DOI":"10.1242\/dev.093906","article-title":"Tendon-bone attachment unit is formed modularly by a distinct pool of Scx- and Sox9-positive progenitors","volume":"140","author":"Blitz","year":"2013","journal-title":"Development"},{"key":"2026060111291923400_DEV205325C4","doi-asserted-by":"publisher","first-page":"825","DOI":"10.1016\/j.molcel.2013.01.038","article-title":"Modification of enhancer chromatin: what, how, and why?","volume":"49","author":"Calo","year":"2013","journal-title":"Mol. Cell"},{"key":"2026060111291923400_DEV205325C5","doi-asserted-by":"publisher","first-page":"156","DOI":"10.1016\/0003-2697(87)90021-2","article-title":"Single-step method of RNA isolation by acid guanidinium thiocyanate-phenol-chloroform extraction","volume":"162","author":"Chomczynski","year":"1987","journal-title":"Anal. Biochem."},{"key":"2026060111291923400_DEV205325C6","doi-asserted-by":"publisher","first-page":"1099","DOI":"10.1242\/dev.121.4.1099","article-title":"Scleraxis: a basic helix-loop-helix protein that prefigures skeletal formation during mouse embryogenesis","volume":"121","author":"Cserjesi","year":"1995","journal-title":"Development"},{"key":"2026060111291923400_DEV205325C7","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1016\/j.gene.2016.04.051","article-title":"Tenogenic modulating insider factor: Systematic assessment on the functions of tenomodulin gene","volume":"587","author":"Dex","year":"2016","journal-title":"Gene"},{"key":"2026060111291923400_DEV205325C8","doi-asserted-by":"publisher","first-page":"179","DOI":"10.1038\/nature10137","article-title":"Role of YAP\/TAZ in mechanotransduction","volume":"474","author":"Dupont","year":"2011","journal-title":"Nature"},{"key":"2026060111291923400_DEV205325C9","doi-asserted-by":"publisher","first-page":"699","DOI":"10.1038\/s41586-020-2493-4","article-title":"Expanded encyclopaedias of DNA elements in the human and mouse genomes","volume":"583","author":"ENCODE Project Consortium","year":"2020","journal-title":"Nature"},{"key":"2026060111291923400_DEV205325C10","doi-asserted-by":"publisher","first-page":"dev165381","DOI":"10.1242\/dev.165381","article-title":"Development of migrating tendon-bone attachments involves replacement of progenitor populations","volume":"145","author":"Felsenthal","year":"2018","journal-title":"Development"},{"key":"2026060111291923400_DEV205325C11","doi-asserted-by":"publisher","first-page":"84","DOI":"10.1016\/j.bbrc.2023.07.039","article-title":"Potential function of Scx+\/Sox9+ cells as progenitor cells in rotator cuff tear repair in rats","volume":"676","author":"Fukuma","year":"2023","journal-title":"Biochem. Biophys. Res. Commun."},{"key":"2026060111291923400_DEV205325C12","doi-asserted-by":"publisher","first-page":"744","DOI":"10.1038\/s41586-020-2093-3","article-title":"An atlas of dynamic chromatin landscapes in mouse fetal development","volume":"583","author":"Gorkin","year":"2020","journal-title":"Nature"},{"key":"2026060111291923400_DEV205325C13","doi-asserted-by":"publisher","first-page":"e138295","DOI":"10.1172\/jci.insight.138295","article-title":"Scleraxis is required for the growth of adult tendons in response to mechanical loading","volume":"5","author":"Gumucio","year":"2020","journal-title":"JCI Insight"},{"key":"2026060111291923400_DEV205325C14","volume-title":"Manipulating the Mouse Embryo: A Laboratory Manual","author":"Hogan","year":"1994","edition":"2nd edn."},{"key":"2026060111291923400_DEV205325C15","doi-asserted-by":"publisher","first-page":"45238","DOI":"10.1038\/srep45238","article-title":"Novel model of tendon regeneration reveals distinct cell mechanisms underlying regenerative and fibrotic tendon healing","volume":"7","author":"Howell","year":"2017","journal-title":"Sci. Rep."},{"key":"2026060111291923400_DEV205325C16","doi-asserted-by":"publisher","first-page":"dev182782","DOI":"10.1242\/dev.182782","article-title":"Requirement for scleraxis in the recruitment of mesenchymal progenitors during embryonic tendon elongation","volume":"146","author":"Huang","year":"2019","journal-title":"Development"},{"key":"2026060111291923400_DEV205325C17","doi-asserted-by":"publisher","first-page":"275","DOI":"10.1093\/bioinformatics\/8.3.275","article-title":"The rapid generation of mutation data matrices from protein sequences","volume":"8","author":"Jones","year":"1992","journal-title":"Comput. Appl. Biosci."},{"key":"2026060111291923400_DEV205325C18","doi-asserted-by":"publisher","first-page":"123","DOI":"10.1679\/aohc.66.123","article-title":"Use of a new adhesive film for the preparation of multi-purpose fresh-frozen sections from hard tissues, whole-animals, insects and plants","volume":"66","author":"Kawamoto","year":"2003","journal-title":"Arch. Histol. Cytol."},{"key":"2026060111291923400_DEV205325C19","doi-asserted-by":"crossref","first-page":"259","DOI":"10.1007\/978-1-0716-1028-2_15","article-title":"Preparation of thin frozen sections from nonfixed and undecalcified hard tissues using Kawamoto's film method (2020)","volume-title":"Skeletal Development and Repair","author":"Kawamoto","year":"2021"},{"key":"2026060111291923400_DEV205325C20","doi-asserted-by":"publisher","first-page":"115969","DOI":"10.1016\/j.bone.2021.115969","article-title":"Scleraxis upregulated by transforming growth factor-beta1 signaling inhibits tension-induced osteoblast differentiation of priodontal ligament cells via ephrin A2","volume":"149","author":"Kawatsu","year":"2021","journal-title":"Bone"},{"key":"2026060111291923400_DEV205325C62","doi-asserted-by":"publisher","first-page":"1258","DOI":"10.1016\/j.joca.2012.07.013","article-title":"The extent of degeneration of cruciate ligament is associated with chondrogenic differentiation in patients with osteoarthritis of the knee","volume":"20","author":"Kumagai","year":"2012","journal-title":"Osteoarthritis Cartilage"},{"key":"2026060111291923400_DEV205325C21","doi-asserted-by":"publisher","first-page":"633","DOI":"10.1016\/j.cell.2016.09.028","article-title":"Progressive loss of function in a limb enhancer during snake evolution","volume":"167","author":"Kvon","year":"2016","journal-title":"Cell"},{"key":"2026060111291923400_DEV205325C22","doi-asserted-by":"publisher","first-page":"2336","DOI":"10.1128\/MCB.17.4.2336","article-title":"SOX9 is a potent activator of the chondrocyte-specific enhancer of the pro\u03b11(II) collagen gene","volume":"17","author":"Lefebvre","year":"1997","journal-title":"Mol. Cell. Biol."},{"key":"2026060111291923400_DEV205325C23","doi-asserted-by":"publisher","first-page":"654397","DOI":"10.3389\/fcell.2021.654397","article-title":"The scleraxis transcription factor directly regulates multiple distinct molecular and cellular processes during early tendon cell differentiation","volume":"9","author":"Liu","year":"2021","journal-title":"Front. Cell Dev. Biol."},{"key":"2026060111291923400_DEV205325C24","doi-asserted-by":"publisher","first-page":"22","DOI":"10.1186\/s13059-014-0560-6","article-title":"Gateways to the FANTOM5 promoter level mammalian expression atlas","volume":"16","author":"Lizio","year":"2015","journal-title":"Genome Biol."},{"key":"2026060111291923400_DEV205325C25","doi-asserted-by":"publisher","first-page":"D752","DOI":"10.1093\/nar\/gky1099","article-title":"Update of the FANTOM web resource: expansion to provide additional transcriptome atlases","volume":"47","author":"Lizio","year":"2019","journal-title":"Nucleic Acids Res."},{"key":"2026060111291923400_DEV205325C26","doi-asserted-by":"publisher","first-page":"933","DOI":"10.1016\/j.cub.2011.04.007","article-title":"Conversion of mechanical force into TGF-beta-mediated biochemical signals","volume":"21","author":"Maeda","year":"2011","journal-title":"Curr. Biol."},{"key":"2026060111291923400_DEV205325C27","doi-asserted-by":"publisher","first-page":"565","DOI":"10.1016\/j.devcel.2023.02.013","article-title":"A single-cell census of mouse limb development identifies complex spatiotemporal dynamics of skeleton formation","volume":"58","author":"Markman","year":"2023","journal-title":"Dev. Cell"},{"key":"2026060111291923400_DEV205325C28","doi-asserted-by":"publisher","first-page":"2697","DOI":"10.1242\/dev.001933","article-title":"Regulation of tendon differentiation by scleraxis distinguishes force-transmitting tendons from muscle-anchoring tendons","volume":"134","author":"Murchison","year":"2007","journal-title":"Development"},{"key":"2026060111291923400_DEV205325C29","doi-asserted-by":"publisher","first-page":"dev201101","DOI":"10.1242\/dev.201101","article-title":"Scleraxis-lineage cells are required for correct muscle patterning","volume":"150","author":"Ono","year":"2023","journal-title":"Development"},{"key":"2026060111291923400_DEV205325C30","doi-asserted-by":"publisher","first-page":"239","DOI":"10.1038\/nature25461","article-title":"Enhancer redundancy provides phenotypic robustness in mammalian development","volume":"554","author":"Osterwalder","year":"2018","journal-title":"Nature"},{"key":"2026060111291923400_DEV205325C31","doi-asserted-by":"publisher","first-page":"499","DOI":"10.1038\/nature05295","article-title":"In vivo enhancer analysis of human conserved non-coding sequences","volume":"444","author":"Pennacchio","year":"2006","journal-title":"Nature"},{"key":"2026060111291923400_DEV205325C32","doi-asserted-by":"publisher","first-page":"1153","DOI":"10.1016\/j.mod.2003.08.003","article-title":"Scleraxis (Scx) directs lacZ expression in tendon of transgenic mice","volume":"120","author":"Perez","year":"2003","journal-title":"Mech. Dev."},{"key":"2026060111291923400_DEV205325C33","doi-asserted-by":"publisher","first-page":"1677","DOI":"10.1002\/dvdy.21179","article-title":"Generation of transgenic tendon reporters, ScxGFP and ScxAP, using regulatory elements of the scleraxis gene","volume":"236","author":"Pryce","year":"2007","journal-title":"Dev. Dyn."},{"key":"2026060111291923400_DEV205325C34","doi-asserted-by":"publisher","first-page":"1351","DOI":"10.1242\/dev.027342","article-title":"Recruitment and maintenance of tendon progenitors by TGFbeta signaling are essential for tendon formation","volume":"136","author":"Pryce","year":"2009","journal-title":"Development"},{"key":"2026060111291923400_DEV205325C35","doi-asserted-by":"publisher","first-page":"2030","DOI":"10.1038\/s41467-024-46396-4","article-title":"Dynamic enhancer landscapes in human craniofacial development","volume":"15","author":"Rajderkar","year":"2024","journal-title":"Nat. Commun."},{"key":"2026060111291923400_DEV205325C36","doi-asserted-by":"publisher","first-page":"113","DOI":"10.1007\/978-1-62703-989-5_9","article-title":"Whole-mount skeletal staining","volume":"1130","author":"Rigueur","year":"2014","journal-title":"Methods Mol. Biol."},{"key":"2026060111291923400_DEV205325C37","doi-asserted-by":"publisher","first-page":"2659","DOI":"10.1242\/dev.124.13.2659","article-title":"TGFbeta2 knockout mice have multiple developmental defects that are non-overlapping with other TGFbeta knockout phenotypes","volume":"124","author":"Sanford","year":"1997","journal-title":"Development"},{"key":"2026060111291923400_DEV205325C38","doi-asserted-by":"publisher","first-page":"59","DOI":"10.1046\/j.1365-2443.1996.04004.x","article-title":"Enhancer analysis of the mouse HNF-3 beta gene: regulatory elements for node\/notochord and floor plate are independent and consist of multiple sub-elements","volume":"1","author":"Sasaki","year":"1996","journal-title":"Genes Cells"},{"key":"2026060111291923400_DEV205325C39","doi-asserted-by":"publisher","first-page":"3855","DOI":"10.1242\/dev.128.19.3855","article-title":"Analysis of the tendon cell fate using Scleraxis, a specific marker for tendons and ligaments","volume":"128","author":"Schweitzer","year":"2001","journal-title":"Development"},{"key":"2026060111291923400_DEV205325C40","doi-asserted-by":"publisher","first-page":"304","DOI":"10.1016\/j.ydbio.2007.07.040","article-title":"Deletion of Tgfbr2 in Prx1-cre expressing mesenchyme results in defects in development of the long bones and joints","volume":"310","author":"Seo","year":"2007","journal-title":"Dev. Biol."},{"key":"2026060111291923400_DEV205325C41","doi-asserted-by":"publisher","first-page":"234","DOI":"10.1016\/j.ydbio.2006.06.036","article-title":"Scleraxis positively regulates the expression of tenomodulin, a differentiation marker of tenocytes","volume":"298","author":"Shukunami","year":"2006","journal-title":"Dev. Biol."},{"key":"2026060111291923400_DEV205325C42","doi-asserted-by":"publisher","first-page":"3155","DOI":"10.1038\/s41598-018-21194-3","article-title":"Scleraxis is a transcriptional activator that regulates the expression of Tenomodulin, a marker of mature tenocytes and ligamentocytes","volume":"8","author":"Shukunami","year":"2018","journal-title":"Sci. Rep."},{"key":"2026060111291923400_DEV205325C43","doi-asserted-by":"publisher","first-page":"449","DOI":"10.1146\/annurev.biochem.72.121801.161520","article-title":"The RNA polymerase II core promoter","volume":"72","author":"Smale","year":"2003","journal-title":"Annu. Rev. Biochem."},{"key":"2026060111291923400_DEV205325C44","doi-asserted-by":"publisher","first-page":"e23568","DOI":"10.1096\/fj.202301452R","article-title":"A role for TGFbeta signaling in Gli1+ tendon and enthesis cells","volume":"38","author":"Song","year":"2024","journal-title":"FASEB J."},{"key":"2026060111291923400_DEV205325C45","doi-asserted-by":"publisher","first-page":"1105","DOI":"10.1083\/jcb.200611031","article-title":"TGF-beta signaling is essential for joint morphogenesis","volume":"177","author":"Spagnoli","year":"2007","journal-title":"J. Cell Biol."},{"key":"2026060111291923400_DEV205325C46","doi-asserted-by":"publisher","first-page":"1237","DOI":"10.1093\/molbev\/msz312","article-title":"Molecular evolutionary genetics analysis (MEGA) for macOS","volume":"37","author":"Stecher","year":"2020","journal-title":"Mol. Biol. Evol."},{"key":"2026060111291923400_DEV205325C47","doi-asserted-by":"publisher","first-page":"2280","DOI":"10.1242\/dev.096354","article-title":"Scx+\/Sox9+ progenitors contribute to the establishment of the junction between cartilage and tendon\/ligament","volume":"140","author":"Sugimoto","year":"2013","journal-title":"Development"},{"key":"2026060111291923400_DEV205325C48","doi-asserted-by":"publisher","first-page":"275","DOI":"10.1002\/dvg.22372","article-title":"Generation and characterization of ScxCre transgenic mice","volume":"51","author":"Sugimoto","year":"2013","journal-title":"Genesis"},{"key":"2026060111291923400_DEV205325C49","doi-asserted-by":"publisher","first-page":"306","DOI":"10.1016\/j.ygeno.2010.02.006","article-title":"A simple and highly efficient transgenesis method in mice with the Tol2 transposon system and cytoplasmic microinjection","volume":"95","author":"Sumiyama","year":"2010","journal-title":"Genomics"},{"key":"2026060111291923400_DEV205325C50","doi-asserted-by":"publisher","first-page":"787","DOI":"10.1242\/dev.116228","article-title":"Scleraxis and osterix antagonistically regulate tensile force-responsive remodeling of the periodontal ligament and alveolar bone","volume":"142","author":"Takimoto","year":"2015","journal-title":"Development"},{"key":"2026060111291923400_DEV205325C51","doi-asserted-by":"publisher","first-page":"3022","DOI":"10.1093\/molbev\/msab120","article-title":"MEGA11: Molecular Evolutionary Genetics Analysis Version 11","volume":"38","author":"Tamura","year":"2021","journal-title":"Mol. Biol. Evol."},{"key":"2026060111291923400_DEV205325C52","doi-asserted-by":"publisher","first-page":"e52695","DOI":"10.7554\/eLife.52695","article-title":"Tgfbeta signaling is critical for maintenance of the tendon cell fate","volume":"9","author":"Tan","year":"2020","journal-title":"eLife"},{"key":"2026060111291923400_DEV205325C53","doi-asserted-by":"publisher","first-page":"2411","DOI":"10.1177\/0363546515597488","article-title":"FGF-2 stimulates the growth of tenogenic progenitor cells to facilitate the generation of tenomodulin-positive tenocytes in a rat rotator cuff healing model","volume":"43","author":"Tokunaga","year":"2015","journal-title":"Am. J. Sports Med."},{"key":"2026060111291923400_DEV205325C54","doi-asserted-by":"publisher","first-page":"52","DOI":"10.2144\/00291bm09","article-title":"Preparation of PCR-quality mouse genomic DNA with hot sodium hydroxide and tris (HotSHOT)","volume":"29","author":"Truett","year":"2000","journal-title":"BioTechniques"},{"key":"2026060111291923400_DEV205325C55","doi-asserted-by":"publisher","first-page":"554","DOI":"10.1016\/j.cell.2015.01.006","article-title":"Enhancer evolution across 20 mammalian species","volume":"160","author":"Villar","year":"2015","journal-title":"Cell"},{"key":"2026060111291923400_DEV205325C56","doi-asserted-by":"publisher","first-page":"307","DOI":"10.1016\/j.cell.2013.03.035","article-title":"Master transcription factors and mediator establish super-enhancers at key cell identity genes","volume":"153","author":"Whyte","year":"2013","journal-title":"Cell"},{"key":"2026060111291923400_DEV205325C57","doi-asserted-by":"publisher","first-page":"1360041","DOI":"10.3389\/fcell.2024.1360041","article-title":"Sclerostin modulates mineralization degree and stiffness profile in the fibrocartilaginous enthesis for mechanical tissue integrity","volume":"12","author":"Yambe","year":"2024","journal-title":"Front. Cell Dev. Biol."},{"key":"2026060111291923400_DEV205325C58","doi-asserted-by":"publisher","first-page":"5394","DOI":"10.1093\/nar\/gkv426","article-title":"The SOX9 upstream region prone to chromosomal aberrations causing campomelic dysplasia contains multiple cartilage enhancers","volume":"43","author":"Yao","year":"2015","journal-title":"Nucleic Acids Res."},{"key":"2026060111291923400_DEV205325C59","doi-asserted-by":"publisher","first-page":"45010","DOI":"10.1038\/srep45010","article-title":"Scleraxis is required for maturation of tissue domains for proper integration of the musculoskeletal system","volume":"7","author":"Yoshimoto","year":"2017","journal-title":"Sci. Rep."},{"key":"2026060111291923400_DEV205325C63","doi-asserted-by":"publisher","first-page":"780038","DOI":"10.3389\/fcell.2022.780038","article-title":"Tenogenic induction from induced pluripotent stem cells unveils the trajectory towards tenocyte differentiation","volume":"10","author":"Yoshimoto","year":"2022","journal-title":"Front. Cell. Dev. Biol."},{"key":"2026060111291923400_DEV205325C60","doi-asserted-by":"publisher","first-page":"dev204512","DOI":"10.1242\/dev.204512","article-title":"Dynamic interactions between cartilaginous and tendinous\/ligamentous primordia during musculoskeletal integration","volume":"152","author":"Yu","year":"2025","journal-title":"Development"}],"container-title":["Development"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/journals.biologists.com\/dev\/article-pdf\/doi\/10.1242\/dev.205325\/3695638\/dev205325.pdf","content-type":"application\/pdf","content-version":"am","intended-application":"syndication"},{"URL":"https:\/\/journals.biologists.com\/dev\/article-pdf\/doi\/10.1242\/dev.205325\/3701673\/dev205325.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"syndication"},{"URL":"https:\/\/journals.biologists.com\/dev\/article-pdf\/doi\/10.1242\/dev.205325\/3701673\/dev205325.pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,6,1]],"date-time":"2026-06-01T15:29:28Z","timestamp":1780327768000},"score":1,"resource":{"primary":{"URL":"https:\/\/journals.biologists.com\/dev\/article\/153\/10\/dev205325\/371861\/Divergent-temporal-control-of-deltoid-tuberosity"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,5,15]]},"references-count":63,"journal-issue":{"issue":"10","published-print":{"date-parts":[[2026,5,15]]}},"URL":"https:\/\/doi.org\/10.1242\/dev.205325","relation":{},"ISSN":["0950-1991","1477-9129"],"issn-type":[{"value":"0950-1991","type":"print"},{"value":"1477-9129","type":"electronic"}],"subject":[],"published":{"date-parts":[[2026,5,15]]},"article-number":"dev205325"}}