{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,9,21]],"date-time":"2026-09-21T11:34:56Z","timestamp":1789990496215,"version":"4.0.1"},"reference-count":57,"publisher":"Oxford University Press (OUP)","issue":"2","license":[{"start":{"date-parts":[[2014,8,22]],"date-time":"2014-08-22T00:00:00Z","timestamp":1408665600000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/academic.oup.com\/journals\/pages\/open_access\/funder_policies\/chorus\/standard_publication_model"}],"funder":[{"name":"Grants-in-Aid from the Ministry of Education, Culture, Sports, Science and Technology (MEXT)"}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":[],"published-print":{"date-parts":[[2015,2,1]]},"abstract":"<jats:title>ABSTRACT<\/jats:title>\n               <jats:sec>\n                  <jats:title>\u2003<\/jats:title>\n                  <jats:p>Transforming growth factor (TGF)-\u03b2 signaling plays critical roles during skeletal development and its excessive signaling causes genetic diseases of connective tissues including Marfan syndrome and acromelic dysplasia. However, the mechanisms underlying prevention of excessive TGF-\u03b2 signaling in skeletogenesis remain unclear. We previously reported that Dullard\/Ctdnep1 encoding a small phosphatase is required for nephron maintenance after birth through suppression of bone morphogenetic protein (BMP) signaling. Unexpectedly, we found that Dullard is involved in suppression of TGF-\u03b2 signaling during endochondral ossification. Conditional Dullard-deficient mice in the limb and sternum mesenchyme by Prx1-Cre displayed the impaired growth and ossification of skeletal elements leading to postnatal lethality. Dullard was expressed in early cartilage condensations and later in growth plate chondrocytes. The tibia growth plate of newborn Dullard mutant mice showed reduction of the proliferative and hypertrophic chondrocyte layers. The sternum showed deformity of cartilage primordia and delayed hypertrophy. Micromass culture experiments revealed that Dullard deficiency enhanced early cartilage condensation and differentiation, but suppressed mineralized hypertrophic chondrocyte differentiation, which was reversed by treatment with TGF-\u03b2 type I receptor kinase blocker LY-364947. Dullard deficiency induced upregulation of protein levels of both phospho-Smad2\/3 and total Smad2\/3 in micromass cultures without increase of Smad2\/3 mRNA levels, suggesting that Dullard may affect Smad2\/3 protein stability. The phospho-Smad2\/3 level was also upregulated in perichondrium and hypertrophic chondrocytes in Dullard-deficient embryos. Response to TGF-\u03b2 signaling was enhanced in Dullard-deficient primary chondrocyte cultures at late, but not early, time point. Moreover, perinatal administration of LY-364947 ameliorated the sternum deformity in vivo. Thus, we identified Dullard as a new negative regulator of TGF-\u03b2 signaling in endochondral ossification. \u00a9 2014 American Society for Bone and Mineral Research.<\/jats:p>\n               <\/jats:sec>","DOI":"10.1002\/jbmr.2343","type":"journal-article","created":{"date-parts":[[2014,8,22]],"date-time":"2014-08-22T21:33:50Z","timestamp":1408743230000},"page":"318-329","source":"Crossref","is-referenced-by-count":19,"title":["<i>Dullard<\/i>\/<i>Ctdnep1<\/i> Regulates Endochondral Ossification via Suppression of TGF-\u03b2 Signaling"],"prefix":"10.1093","volume":"30","author":[{"given":"Tadayoshi","family":"Hayata","sequence":"first","affiliation":[{"name":"Department of Molecular PharmacologyMedical Research InstituteTokyo Medical and Dental UniversityTokyoJapan"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"family":"Yoichi","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"family":"Ezura","sequence":"additional","affiliation":[{"name":"Department of Molecular PharmacologyMedical Research InstituteTokyo Medical and Dental UniversityTokyoJapan"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Makoto","family":"Asashima","sequence":"additional","affiliation":[{"name":"Research Center for Stem Cell EngineeringNational Institute of Advanced Industrial Science and Technology (AIST)IbarakiJapan"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Ryuichi","family":"Nishinakamura","sequence":"additional","affiliation":[{"name":"Department of Kidney Development, Institute of Molecular Embryology and GeneticsKumamoto UniversityKumamotoJapan"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Masaki","family":"Noda","sequence":"additional","affiliation":[{"name":"Global Centers of Excellence (COE) ProgramTokyo Medical and Dental UniversityTokyoJapan"},{"name":"Department of Molecular PharmacologyMedical Research InstituteTokyo Medical and Dental UniversityTokyoJapan"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"286","published-online":{"date-parts":[[2014,8,22]]},"reference":[{"issue":"6937","key":"2024020319195389200_jbmr2343-bib-0001","doi-asserted-by":"crossref","first-page":"332","DOI":"10.1038\/nature01657","article-title":"Developmental regulation of the growth plate","volume":"423","author":"Kronenberg","year":"2003","journal-title":"Nature."},{"issue":"1\u20132","key":"2024020319195389200_jbmr2343-bib-0002","first-page":"73","article-title":"Tgfbeta2\u2013\/\u2013 Tgfbeta3\u2013\/\u2013 double knockout mice display severe midline fusion defects and early embryonic lethality","volume":"206","author":"D\u00fcnker","year":"2002","journal-title":"Anat Embryol (Berl)."},{"issue":"2","key":"2024020319195389200_jbmr2343-bib-0003","doi-asserted-by":"crossref","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."},{"issue":"6","key":"2024020319195389200_jbmr2343-bib-0004","doi-asserted-by":"crossref","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."},{"issue":"2","key":"2024020319195389200_jbmr2343-bib-0005","doi-asserted-by":"crossref","first-page":"325","DOI":"10.1016\/j.ydbio.2009.06.002","article-title":"Critical roles of the TGF-beta type I receptor ALK5 in perichondrial formation and function, cartilage integrity, and osteoblast differentiation during growth plate development","volume":"332","author":"Matsunobu","year":"2009","journal-title":"Dev Biol."},{"issue":"6","key":"2024020319195389200_jbmr2343-bib-0006","doi-asserted-by":"crossref","first-page":"352","DOI":"10.1016\/j.matbio.2012.07.002","article-title":"Conditional deletion of Tgfbr2 in hypertrophic chondrocytes delays terminal chondrocyte differentiation","volume":"31","author":"Sueyoshi","year":"2012","journal-title":"Matrix Biol."},{"issue":"6333","key":"2024020319195389200_jbmr2343-bib-0007","doi-asserted-by":"crossref","first-page":"337","DOI":"10.1038\/352337a0","article-title":"Marfan syndrome caused by a recurrent de novo missense mutation in the fibrillin gene","volume":"352","author":"Dietz","year":"1991","journal-title":"Nature."},{"issue":"3","key":"2024020319195389200_jbmr2343-bib-0008","doi-asserted-by":"crossref","first-page":"275","DOI":"10.1038\/ng1511","article-title":"A syndrome of altered cardiovascular, craniofacial, neurocognitive and skeletal development caused by mutations in TGFBR1 or TGFBR2","volume":"37","author":"Loeys","year":"2005","journal-title":"Nat Genet."},{"issue":"2","key":"2024020319195389200_jbmr2343-bib-0009","doi-asserted-by":"crossref","first-page":"121","DOI":"10.1038\/ng.744","article-title":"Mutations in SMAD3 cause a syndromic form of aortic aneurysms and dissections with early-onset osteoarthritis","volume":"43","author":"van de Laar","year":"2011","journal-title":"Nat Genet."},{"issue":"8","key":"2024020319195389200_jbmr2343-bib-0010","doi-asserted-by":"crossref","first-page":"922","DOI":"10.1038\/ng.2349","article-title":"Loss-of-function mutations in TGFB2 cause a syndromic presentation of thoracic aortic aneurysm","volume":"44","author":"Lindsay","year":"2012","journal-title":"Nat Genet."},{"issue":"11","key":"2024020319195389200_jbmr2343-bib-0011","doi-asserted-by":"crossref","first-page":"1249","DOI":"10.1038\/ng.2421","article-title":"Mutations in the TGF-\u03b2 repressor SKI cause Shprintzen-Goldberg syndrome with aortic aneurysm","volume":"44","author":"Doyle","year":"2012","journal-title":"Nat Genet."},{"issue":"1","key":"2024020319195389200_jbmr2343-bib-0012","doi-asserted-by":"crossref","first-page":"34","DOI":"10.1136\/jmg.40.1.34","article-title":"In frame fibrillin-1 gene deletion in autosomal dominant Weill-Marchesani syndrome","volume":"40","author":"Faivre","year":"2003","journal-title":"J Med Genet."},{"issue":"5","key":"2024020319195389200_jbmr2343-bib-0013","doi-asserted-by":"crossref","first-page":"801","DOI":"10.1086\/425231","article-title":"ADAMTS10 mutations in autosomal recessive Weill-Marchesani syndrome","volume":"75","author":"Dagoneau","year":"2004","journal-title":"Am J Hum Genet."},{"issue":"9","key":"2024020319195389200_jbmr2343-bib-0014","doi-asserted-by":"crossref","first-page":"1119","DOI":"10.1038\/ng.199","article-title":"ADAMTSL2 mutations in geleophysic dysplasia demonstrate a role for ADAMTS-like proteins in TGF-beta bioavailability regulation","volume":"40","author":"Le Goff","year":"2008","journal-title":"Nat Genet."},{"issue":"1","key":"2024020319195389200_jbmr2343-bib-0015","doi-asserted-by":"crossref","first-page":"7","DOI":"10.1016\/j.ajhg.2011.05.012","article-title":"Mutations in the TGF\u03b2 binding-protein-like domain 5 of FBN1 are responsible for acromicric and geleophysic dysplasias","volume":"89","author":"Le Goff","year":"2011","journal-title":"Am J Hum Genet."},{"issue":"1","key":"2024020319195389200_jbmr2343-bib-0016","doi-asserted-by":"crossref","first-page":"85","DOI":"10.1038\/ng.1016","article-title":"Mutations at a single codon in Mad homology 2 domain of SMAD4 cause Myhre syndrome","volume":"44","author":"Le Goff","year":"2012","journal-title":"Nat Genet."},{"issue":"3","key":"2024020319195389200_jbmr2343-bib-0017","doi-asserted-by":"crossref","first-page":"145","DOI":"10.1002\/ajmg.c.31337","article-title":"From tall to short: the role of TGF\u03b2 signaling in growth and its disorders","volume":"160C","author":"Le Goff","year":"2012","journal-title":"Am J Med Genet C Semin Med Genet."},{"issue":"1","key":"2024020319195389200_jbmr2343-bib-0018","doi-asserted-by":"crossref","first-page":"85","DOI":"10.1016\/S0006-291X(02)00641-1","article-title":"Molecular cloning and characterization of dullard: a novel gene required for neural development","volume":"295","author":"Satow","year":"2002","journal-title":"Biochem Biophys Res Commun."},{"issue":"6","key":"2024020319195389200_jbmr2343-bib-0019","doi-asserted-by":"crossref","first-page":"763","DOI":"10.1016\/j.devcel.2006.10.001","article-title":"Dullard promotes degradation and dephosphorylation of BMP receptors and is required for neural induction","volume":"11","author":"Satow","year":"2006","journal-title":"Dev Cell."},{"issue":"16","key":"2024020319195389200_jbmr2343-bib-0020","doi-asserted-by":"crossref","first-page":"6596","DOI":"10.1073\/pnas.0702099104","article-title":"A conserved phosphatase cascade that regulates nuclear membrane biogenesis","volume":"104","author":"Kim","year":"2007","journal-title":"Proc Natl Acad Sci U S A."},{"issue":"15","key":"2024020319195389200_jbmr2343-bib-0021","doi-asserted-by":"crossref","first-page":"3045","DOI":"10.1021\/bi200336b","article-title":"Homo sapiens dullard protein phosphatase shows a preference for the insulin-dependent phosphorylation site of lipin1","volume":"50","author":"Wu","year":"2011","journal-title":"Biochemistry."},{"issue":"3","key":"2024020319195389200_jbmr2343-bib-0022","doi-asserted-by":"crossref","first-page":"e57428","DOI":"10.1371\/journal.pone.0057428","article-title":"Dullard\/Ctdnep1 modulates WNT signalling activity for the formation of primordial germ cells in the mouse embryo","volume":"8","author":"Tanaka","year":"2013","journal-title":"PLoS One."},{"key":"2024020319195389200_jbmr2343-bib-0023","doi-asserted-by":"crossref","first-page":"1398","DOI":"10.1038\/ncomms2408","article-title":"The phosphatase Dullard negatively regulates BMP signalling and is essential for nephron maintenance after birth","volume":"4","author":"Sakaguchi","year":"2013","journal-title":"Nat Commun."},{"issue":"7409","key":"2024020319195389200_jbmr2343-bib-0024","doi-asserted-by":"crossref","first-page":"100","DOI":"10.1038\/nature11284","article-title":"Dissecting the genomic complexity underlying medulloblastoma","volume":"488","author":"Jones","year":"2012","journal-title":"Nature."},{"issue":"7409","key":"2024020319195389200_jbmr2343-bib-0025","doi-asserted-by":"crossref","first-page":"106","DOI":"10.1038\/nature11329","article-title":"Medulloblastoma exome sequencing uncovers subtype-specific somatic mutations","volume":"488","author":"Pugh","year":"2012","journal-title":"Nature."},{"issue":"5368","key":"2024020319195389200_jbmr2343-bib-0026","doi-asserted-by":"crossref","first-page":"1455","DOI":"10.1126\/science.280.5368.1455","article-title":"Noggin, cartilage morphogenesis, and joint formation in the mammalian skeleton","volume":"280","author":"Brunet","year":"1998","journal-title":"Science."},{"issue":"14","key":"2024020319195389200_jbmr2343-bib-0027","doi-asserted-by":"crossref","first-page":"5062","DOI":"10.1073\/pnas.0500031102","article-title":"Bmpr1a and Bmpr1b have overlapping functions and are essential for chondrogenesis in vivo","volume":"102","author":"Yoon","year":"2005","journal-title":"Proc Natl Acad Sci U S A."},{"issue":"10","key":"2024020319195389200_jbmr2343-bib-0028","doi-asserted-by":"crossref","first-page":"2498","DOI":"10.1002\/jbmr.443","article-title":"Smad6 is essential to limit BMP signaling during cartilage development","volume":"26","author":"Estrada","year":"2011","journal-title":"J Bone Miner Res."},{"issue":"2","key":"2024020319195389200_jbmr2343-bib-0029","doi-asserted-by":"crossref","first-page":"77","DOI":"10.1002\/gene.10092","article-title":"Expression of Cre Recombinase in the developing mouse limb bud driven by a Prxl enhancer","volume":"33","author":"Logan","year":"2002","journal-title":"Genesis."},{"key":"2024020319195389200_jbmr2343-bib-0030","volume-title":"Manipulating the mouse embryo. A laboratory manual","author":"Hogan","year":"1994","edition":"2nd ed"},{"issue":"3","key":"2024020319195389200_jbmr2343-bib-0031","doi-asserted-by":"crossref","first-page":"191","DOI":"10.1007\/s00774-005-0671-y","article-title":"Involvement of Notch signaling in initiation of prechondrogenic condensation and nodule formation in limb bud micromass cultures","volume":"24","author":"Fujimaki","year":"2006","journal-title":"J Bone Miner Metab."},{"issue":"8","key":"2024020319195389200_jbmr2343-bib-0032","doi-asserted-by":"crossref","first-page":"1253","DOI":"10.1038\/nprot.2008.95","article-title":"Primary culture and phenotyping of murine chondrocytes","volume":"3","author":"Gosset","year":"2008","journal-title":"Nat Protoc."},{"issue":"11","key":"2024020319195389200_jbmr2343-bib-0033","doi-asserted-by":"crossref","first-page":"3091","DOI":"10.1093\/emboj\/17.11.3091","article-title":"Direct binding of Smad3 and Smad4 to critical TGF beta-inducible elements in the promoter of human plasminogen activator inhibitor-type 1 gene","volume":"17","author":"Dennler","year":"1998","journal-title":"EMBO J."},{"issue":"2","key":"2024020319195389200_jbmr2343-bib-0034","doi-asserted-by":"crossref","first-page":"210","DOI":"10.1016\/j.ydbio.2005.02.014","article-title":"Phylogenetic footprinting and genome scanning identify vertebrate BMP response elements and new target genes","volume":"281","author":"von Bubnoff","year":"2005","journal-title":"Dev Biol."},{"issue":"40","key":"2024020319195389200_jbmr2343-bib-0035","doi-asserted-by":"crossref","first-page":"33545","DOI":"10.1074\/jbc.M111.329938","article-title":"Profilin1 regulates sternum development and endochondral bone formation","volume":"287","author":"Miyajima","year":"2012","journal-title":"J Biol Chem."},{"issue":"2","key":"2024020319195389200_jbmr2343-bib-0036","doi-asserted-by":"crossref","first-page":"103","DOI":"10.1002\/aja.1002000203","article-title":"Transforming growth factor-beta and bone morphogenetic protein-2 act by distinct mechanisms to promote chick limb cartilage differentiation in vitro","volume":"200","author":"Roark","year":"1994","journal-title":"Dev Dyn."},{"issue":"2","key":"2024020319195389200_jbmr2343-bib-0037","doi-asserted-by":"crossref","first-page":"195","DOI":"10.1002\/dvdy.10151","article-title":"Analysis of N-cadherin function in limb mesenchymal chondrogenesis in vitro","volume":"225","author":"Delise","year":"2002","journal-title":"Dev Dyn."},{"issue":"1","key":"2024020319195389200_jbmr2343-bib-0038","doi-asserted-by":"crossref","first-page":"35","DOI":"10.1083\/jcb.153.1.35","article-title":"TGF-\u03b2\/Smad3 signals repress chondrocyte hypertrophic differentiation and are required for maintaining articular cartilage","volume":"153","author":"Yang","year":"2001","journal-title":"J Cell Biol."},{"issue":"9","key":"2024020319195389200_jbmr2343-bib-0039","doi-asserted-by":"crossref","first-page":"e24546","DOI":"10.1371\/journal.pone.0024546","article-title":"Defining the earliest transcriptional steps of chondrogenic progenitor specification during the formation of the digits in the embryonic limb","volume":"6","author":"Lorda-Diez","year":"2011","journal-title":"PLoS One."},{"issue":"16","key":"2024020319195389200_jbmr2343-bib-0040","doi-asserted-by":"crossref","first-page":"2072","DOI":"10.1101\/gad.13.16.2072","article-title":"Indian hedgehog signaling regulates proliferation and differentiation of chondrocytes and is essential for bone formation","volume":"13","author":"St-Jacques","year":"1999","journal-title":"Genes Dev."},{"issue":"21","key":"2024020319195389200_jbmr2343-bib-0041","doi-asserted-by":"crossref","first-page":"2813","DOI":"10.1101\/gad.1017802","article-title":"The transcription factor Sox9 has essential roles in successive steps of the chondrocyte differentiation pathway and is required for expression of Sox5 and Sox6","volume":"16","author":"Akiyama","year":"2002","journal-title":"Genes Dev."},{"issue":"12","key":"2024020319195389200_jbmr2343-bib-0042","doi-asserted-by":"crossref","first-page":"2779","DOI":"10.1242\/dev.00505","article-title":"Connective tissue growth factor coordinates chondrogenesis and angiogenesis during skeletal development","volume":"130","author":"Ivkovic","year":"2003","journal-title":"Development."},{"issue":"1","key":"2024020319195389200_jbmr2343-bib-0043","doi-asserted-by":"crossref","first-page":"206","DOI":"10.1006\/bbrc.1997.6528","article-title":"Cloning of a mRNA preferentially expressed in chondrocytes by differential display-PCR from a human chondrocytic cell line that is identical with connective tissue growth factor (CTGF) mRNA","volume":"234","author":"Nakanishi","year":"1997","journal-title":"Biochem Biophys Res Commun."},{"issue":"9","key":"2024020319195389200_jbmr2343-bib-0044","doi-asserted-by":"crossref","first-page":"2161","DOI":"10.1242\/dev.01053","article-title":"VEGFA is necessary for chondrocyte survival during bone development","volume":"131","author":"Zelzer","year":"2004","journal-title":"Development."},{"issue":"Pt 1","key":"2024020319195389200_jbmr2343-bib-0045","doi-asserted-by":"crossref","first-page":"59","DOI":"10.1242\/jcs.113.1.59","article-title":"Vascular endothelial growth factor (VEGF) in cartilage neovascularization and chondrocyte differentiation: auto-paracrine role during endochondral bone formation","volume":"113","author":"Carlevaro","year":"2000","journal-title":"J Cell Sci."},{"issue":"1","key":"2024020319195389200_jbmr2343-bib-0046","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1002\/(SICI)1097-4644(199607)62:1<1::AID-JCB1>3.0.CO;2-X","article-title":"Expression of bone-specific genes by hypertrophic chondrocytes: implication of the complex functions of the hypertrophic chondrocyte during endochondral bone development","volume":"62","author":"Gerstenfeld","year":"1996","journal-title":"J Cell Biochem."},{"issue":"1","key":"2024020319195389200_jbmr2343-bib-0047","doi-asserted-by":"crossref","first-page":"17","DOI":"10.1016\/S0092-8674(01)00622-5","article-title":"The novel zinc finger-containing transcription factor osterix is required for osteoblast differentiation and bone formation","volume":"108","author":"Nakashima","year":"2002","journal-title":"Cell."},{"issue":"12","key":"2024020319195389200_jbmr2343-bib-0048","doi-asserted-by":"crossref","first-page":"8695","DOI":"10.1074\/jbc.275.12.8695","article-title":"Cbfa1 is a positive regulatory factor in chondrocyte maturation","volume":"275","author":"Enomoto","year":"2000","journal-title":"J Biol Chem."},{"issue":"6","key":"2024020319195389200_jbmr2343-bib-0049","doi-asserted-by":"crossref","first-page":"822","DOI":"10.1111\/j.1440-169X.2011.01289.x","article-title":"Negative modulation of bone morphogenetic protein signaling by Dullard during wing vein formation in Drosophila","volume":"53","author":"Liu","year":"2011","journal-title":"Dev Growth Differ."},{"issue":"1","key":"2024020319195389200_jbmr2343-bib-0050","doi-asserted-by":"crossref","first-page":"4","DOI":"10.1359\/JBMR.050911","article-title":"Smad3-deficient chondrocytes have enhanced BMP signaling and accelerated differentiation","volume":"21","author":"Li","year":"2006","journal-title":"J Bone Miner Res."},{"issue":"14","key":"2024020319195389200_jbmr2343-bib-0051","doi-asserted-by":"crossref","first-page":"2904","DOI":"10.1128\/MCB.00231-12","article-title":"Transforming growth factor \u03b2 inhibits bone morphogenetic protein-induced transcription through novel phosphorylated Smad1\/5-Smad3 complexes","volume":"32","author":"Gr\u00f6nroos","year":"2012","journal-title":"Mol Cell Biol."},{"issue":"22","key":"2024020319195389200_jbmr2343-bib-0052","doi-asserted-by":"crossref","first-page":"4467","DOI":"10.1242\/dev.124.22.4467","article-title":"Cellular interpretation of multiple TGF-beta signals: intracellular antagonism between activin\/BVg1 and BMP-2\/4 signaling mediated by Smads","volume":"124","author":"Candia","year":"1997","journal-title":"Development."},{"issue":"3","key":"2024020319195389200_jbmr2343-bib-0053","doi-asserted-by":"crossref","first-page":"552","DOI":"10.1038\/sj.emboj.7600067","article-title":"Endogenous TGF-beta signaling suppresses maturation of osteoblastic mesenchymal cells","volume":"23","author":"Maeda","year":"2004","journal-title":"EMBO J."},{"issue":"6","key":"2024020319195389200_jbmr2343-bib-0054","doi-asserted-by":"crossref","first-page":"1476","DOI":"10.1634\/stemcells.2005-0299","article-title":"Activin A maintains self-renewal and regulates fibroblast growth factor, Wnt, and bone morphogenic protein pathways in human embryonic stem cells","volume":"24","author":"Xiao","year":"2006","journal-title":"Stem Cells."},{"issue":"5","key":"2024020319195389200_jbmr2343-bib-0055","doi-asserted-by":"crossref","first-page":"949","DOI":"10.1016\/S1097-2765(01)00249-0","article-title":"Nodal signals to Smads through Cripto-dependent and Cripto-independent mechanisms","volume":"7","author":"Yeo","year":"2001","journal-title":"Mol Cell."},{"issue":"3","key":"2024020319195389200_jbmr2343-bib-0056","doi-asserted-by":"crossref","first-page":"457","DOI":"10.1016\/j.molcel.2009.09.043","article-title":"Ubiquitin ligase Nedd4L targets activated Smad2\/3 to limit TGF-beta signaling","volume":"36","author":"Gao","year":"2009","journal-title":"Mol Cell."},{"issue":"3","key":"2024020319195389200_jbmr2343-bib-0057","doi-asserted-by":"crossref","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."}],"container-title":["Journal of Bone and Mineral Research"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/api.wiley.com\/onlinelibrary\/tdm\/v1\/articles\/10.1002%2Fjbmr.2343","content-type":"unspecified","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/academic.oup.com\/jbmr\/article-pdf\/30\/2\/318\/56569115\/jbmr2343.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"syndication"},{"URL":"https:\/\/academic.oup.com\/jbmr\/article-pdf\/30\/2\/318\/56569115\/jbmr2343.pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2024,3,13]],"date-time":"2024-03-13T14:52:49Z","timestamp":1710341569000},"score":1,"resource":{"primary":{"URL":"https:\/\/academic.oup.com\/jbmr\/article\/30\/2\/318-329\/7599346"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2014,8,22]]},"references-count":57,"journal-issue":{"issue":"2","published-online":{"date-parts":[[2014,8,22]]},"published-print":{"date-parts":[[2015,2,1]]}},"URL":"https:\/\/doi.org\/10.1002\/jbmr.2343","relation":{},"ISSN":["0884-0431","1523-4681"],"issn-type":[{"value":"0884-0431","type":"print"},{"value":"1523-4681","type":"electronic"}],"subject":[],"published-other":{"date-parts":[[2015,2,1]]},"published":{"date-parts":[[2014,8,22]]}}}