{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,10,12]],"date-time":"2026-10-12T16:58:45Z","timestamp":1791824325894,"version":"4.3.4"},"reference-count":43,"publisher":"Elsevier BV","license":[{"start":{"date-parts":[[2021,1,1]],"date-time":"2021-01-01T00:00:00Z","timestamp":1609459200000},"content-version":"tdm","delay-in-days":0,"URL":"https:\/\/www.elsevier.com\/tdm\/userlicense\/1.0\/"},{"start":{"date-parts":[[2021,1,1]],"date-time":"2021-01-01T00:00:00Z","timestamp":1609459200000},"content-version":"tdm","delay-in-days":0,"URL":"https:\/\/www.elsevier.com\/legal\/tdmrep-license"}],"funder":[{"DOI":"10.13039\/100014103","name":"Key Technology Research and Development Program of Shandong","doi-asserted-by":"publisher","id":[{"id":"10.13039\/100014103","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100014857","name":"National Natural Science Foundation of China-Guangdong Joint Fund","doi-asserted-by":"publisher","id":[{"id":"10.13039\/501100014857","id-type":"DOI","asserted-by":"publisher"}]}],"content-domain":{"domain":["clinicalkey.com","clinicalkey.com.au","clinicalkey.es","clinicalkey.fr","clinicalkey.jp","elsevier.com","sciencedirect.com"],"crossmark-restriction":true},"short-container-title":["Bone"],"published-print":{"date-parts":[[2021,1]]},"DOI":"10.1016\/j.bone.2020.115686","type":"journal-article","created":{"date-parts":[[2020,10,12]],"date-time":"2020-10-12T12:38:36Z","timestamp":1602506316000},"page":"115686","update-policy":"https:\/\/doi.org\/10.1016\/elsevier_cm_policy","source":"Crossref","is-referenced-by-count":24,"special_numbering":"C","title":["A SMOC2 variant inhibits BMP signaling by competitively binding to BMPR1B and causes growth plate defects"],"prefix":"10.1016","volume":"142","author":[{"given":"Feng","family":"Long","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Hongbiao","family":"Shi","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Pengyu","family":"Li","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Shaoqiang","family":"Guo","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Yuer","family":"Ma","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Shijun","family":"Wei","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Yan","family":"Li","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Fei","family":"Gao","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Shang","family":"Gao","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Meitian","family":"Wang","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Ruonan","family":"Duan","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Xiaojing","family":"Wang","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Kun","family":"Yang","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Wenjie","family":"Sun","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Xi","family":"Li","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Jiangxia","family":"Li","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Qiji","family":"Liu","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"78","reference":[{"key":"10.1016\/j.bone.2020.115686_bb0005","doi-asserted-by":"crossref","first-page":"191","DOI":"10.1146\/annurev.cellbio.16.1.191","article-title":"Bone development","volume":"16","author":"Olsen","year":"2000","journal-title":"Annu. Rev. Cell Dev. Biol."},{"issue":"14","key":"10.1016\/j.bone.2020.115686_bb0010","doi-asserted-by":"crossref","first-page":"1728","DOI":"10.1093\/hmg\/ddm121","article-title":"Decreased chondrocyte proliferation and dysregulated apoptosis in the cartilage growth plate are key features of a murine model of epiphyseal dysplasia caused by a matn3 mutation","volume":"16","author":"Leighton","year":"2007","journal-title":"Hum. Mol. Genet."},{"issue":"12","key":"10.1016\/j.bone.2020.115686_bb0015","doi-asserted-by":"crossref","first-page":"1730","DOI":"10.5966\/sctm.2015-0321","article-title":"Spontaneous in vivo chondrogenesis of bone marrow-derived mesenchymal progenitor cells by blocking vascular endothelial growth factor signaling","volume":"5","author":"Marsano","year":"2016","journal-title":"Stem Cells Transl. Med."},{"issue":"11","key":"10.1016\/j.bone.2020.115686_bb0020","doi-asserted-by":"crossref","first-page":"2478","DOI":"10.3390\/ijms18112478","article-title":"Vascular endothelial growth factor sequestration enhances in vivo cartilage formation","volume":"18","author":"Medeiros Da Cunha","year":"2017","journal-title":"Int J Mol Sci"},{"issue":"2","key":"10.1016\/j.bone.2020.115686_bb0025","doi-asserted-by":"crossref","first-page":"304","DOI":"10.1002\/stem.792","article-title":"The crosstalk between transforming growth factor-beta1 and delta like-1 mediates early chondrogenesis during embryonic endochondral ossification","volume":"30","author":"Taipaleenmaki","year":"2012","journal-title":"Stem Cells"},{"issue":"23\u201324","key":"10.1016\/j.bone.2020.115686_bb0030","doi-asserted-by":"crossref","first-page":"2638","DOI":"10.1089\/ten.tea.2013.0047","article-title":"Induction of chondrogenesis and expression of superficial zone protein in synovial explants with TGF-beta1 and BMP-7","volume":"19","author":"Iwakura","year":"2013","journal-title":"Tissue Eng Part A"},{"issue":"9","key":"10.1016\/j.bone.2020.115686_bb0035","doi-asserted-by":"crossref","first-page":"1081","DOI":"10.1016\/j.joca.2011.05.003","article-title":"The relationship between fibrogenic TGFbeta1 signaling in the joint and cartilage degradation in post-injury osteoarthritis","volume":"19","author":"Plaas","year":"2011","journal-title":"Osteoarthr. Cartil."},{"issue":"2","key":"10.1016\/j.bone.2020.115686_bb0040","doi-asserted-by":"crossref","first-page":"186","DOI":"10.1038\/labinvest.2015.142","article-title":"Wnt signaling in cartilage development and diseases: lessons from animal studies","volume":"96","author":"Usami","year":"2016","journal-title":"Lab. Investig."},{"issue":"7","key":"10.1016\/j.bone.2020.115686_bb0045","doi-asserted-by":"crossref","first-page":"485","DOI":"10.5483\/BMBRep.2008.41.7.485","article-title":"Wnt signaling in cartilage development and degeneration","volume":"41","author":"Chun","year":"2008","journal-title":"BMB Rep."},{"issue":"1","key":"10.1016\/j.bone.2020.115686_bb0050","doi-asserted-by":"crossref","first-page":"93","DOI":"10.1002\/jcb.20211","article-title":"Multiple functions of BMPs in chondrogenesis","volume":"93","author":"Yoon","year":"2004","journal-title":"J. Cell. Biochem."},{"issue":"7","key":"10.1016\/j.bone.2020.115686_bb0055","doi-asserted-by":"crossref","first-page":"1093","DOI":"10.1242\/dev.029926","article-title":"BMP canonical Smad signaling through Smad1 and Smad5 is required for endochondral bone formation","volume":"136","author":"Retting","year":"2009","journal-title":"Development"},{"issue":"1","key":"10.1016\/j.bone.2020.115686_bb0060","doi-asserted-by":"crossref","DOI":"10.1371\/journal.pone.0016421","article-title":"Interaction of TGFbeta and BMP signaling pathways during chondrogenesis","volume":"6","author":"Keller","year":"2011","journal-title":"PLoS One"},{"issue":"2","key":"10.1016\/j.bone.2020.115686_bb0065","doi-asserted-by":"crossref","first-page":"237","DOI":"10.1006\/dbio.2000.9610","article-title":"BMPs are required at two steps of limb chondrogenesis: formation of prechondrogenic condensations and their differentiation into chondrocytes","volume":"219","author":"Pizette","year":"2000","journal-title":"Dev. Biol."},{"issue":"22","key":"10.1016\/j.bone.2020.115686_bb0070","doi-asserted-by":"crossref","first-page":"4523","DOI":"10.1242\/dev.128.22.4523","article-title":"BMP and Ihh\/PTHrP signaling interact to coordinate chondrocyte proliferation and differentiation","volume":"128","author":"Minina","year":"2001","journal-title":"Development"},{"issue":"1","key":"10.1016\/j.bone.2020.115686_bb0075","doi-asserted-by":"crossref","first-page":"430","DOI":"10.1210\/endo.142.1.7901","article-title":"Regulation of growth plate chondrogenesis by bone morphogenetic protein-2","volume":"142","author":"De Luca","year":"2001","journal-title":"Endocrinology"},{"issue":"2","key":"10.1016\/j.bone.2020.115686_bb0080","doi-asserted-by":"crossref","first-page":"239","DOI":"10.1006\/dbio.1998.9181","article-title":"Interaction of Ihh and BMP\/noggin signaling during cartilage differentiation","volume":"209","author":"Pathi","year":"1999","journal-title":"Dev. Biol."},{"issue":"1","key":"10.1016\/j.bone.2020.115686_bb0085","doi-asserted-by":"crossref","first-page":"9","DOI":"10.1002\/path.5170","article-title":"Bone morphogenetic protein receptor signal transduction in human disease","volume":"247","author":"Gomez-Puerto","year":"2019","journal-title":"J. Pathol."},{"key":"10.1016\/j.bone.2020.115686_bb0090","doi-asserted-by":"crossref","first-page":"28","DOI":"10.1016\/j.pneurobio.2013.07.002","article-title":"BMP-Smad 1\/5\/8 signalling in the development of the nervous system","volume":"109","author":"Hegarty","year":"2013","journal-title":"Prog. Neurobiol."},{"issue":"4","key":"10.1016\/j.bone.2020.115686_bb0095","doi-asserted-by":"crossref","first-page":"203","DOI":"10.1038\/nrendo.2016.12","article-title":"BMP signalling in skeletal development, disease and repair","volume":"12","author":"Salazar","year":"2016","journal-title":"Nat Rev Endocrinol"},{"key":"10.1016\/j.bone.2020.115686_bb0100","doi-asserted-by":"crossref","first-page":"129","DOI":"10.1016\/j.cytogfr.2015.11.004","article-title":"Common mechanisms in development and disease: BMP signaling in craniofacial development","volume":"27","author":"Graf","year":"2016","journal-title":"Cytokine Growth Factor Rev."},{"issue":"25","key":"10.1016\/j.bone.2020.115686_bb0105","doi-asserted-by":"crossref","first-page":"16985","DOI":"10.1016\/S0021-9258(17)32506-1","article-title":"Identification of type I receptors for osteogenic protein-1 and bone morphogenetic protein-4","volume":"269","author":"Ten Dijke","year":"1994","journal-title":"J. Biol. Chem."},{"issue":"23","key":"10.1016\/j.bone.2020.115686_bb0110","doi-asserted-by":"crossref","first-page":"4667","DOI":"10.1242\/dev.02680","article-title":"BMPs regulate multiple aspects of growth-plate chondrogenesis through opposing actions on FGF pathways","volume":"133","author":"Yoon","year":"2006","journal-title":"Development"},{"issue":"Pt 3","key":"10.1016\/j.bone.2020.115686_bb0115","doi-asserted-by":"crossref","first-page":"805","DOI":"10.1042\/bj20030532","article-title":"Characterization of SMOC-2, a modular extracellular calcium-binding protein","volume":"373","author":"Vannahme","year":"2003","journal-title":"Biochem. J."},{"issue":"41","key":"10.1016\/j.bone.2020.115686_bb0120","doi-asserted-by":"crossref","first-page":"37977","DOI":"10.1074\/jbc.M203830200","article-title":"Characterization of SMOC-1, a novel modular calcium-binding protein in basement membranes","volume":"277","author":"Vannahme","year":"2002","journal-title":"J. Biol. Chem."},{"issue":"7","key":"10.1016\/j.bone.2020.115686_bb0125","doi-asserted-by":"crossref","DOI":"10.1371\/journal.pgen.1002114","article-title":"Loss of the BMP antagonist, SMOC-1, causes Ophthalmo-acromelic (Waardenburg Anophthalmia) syndrome in humans and mice","volume":"7","author":"Rainger","year":"2011","journal-title":"PLoS Genet."},{"issue":"11","key":"10.1016\/j.bone.2020.115686_bb0130","doi-asserted-by":"crossref","first-page":"1375","DOI":"10.1002\/dvdy.24164","article-title":"Smoc2 modulates embryonic myelopoiesis during zebrafish development","volume":"243","author":"Mommaerts","year":"2014","journal-title":"Dev. Dyn."},{"issue":"28","key":"10.1016\/j.bone.2020.115686_bb0135","doi-asserted-by":"crossref","first-page":"18994","DOI":"10.1074\/jbc.M807759200","article-title":"Xenopus SMOC-1 inhibits bone morphogenetic protein signaling downstream of receptor binding and is essential for postgastrulation development in Xenopus","volume":"284","author":"Thomas","year":"2009","journal-title":"J. Biol. Chem."},{"key":"10.1016\/j.bone.2020.115686_bb0140","doi-asserted-by":"crossref","DOI":"10.7554\/eLife.17935","article-title":"SMOC can act as both an antagonist and an expander of BMP signaling","volume":"6","author":"Thomas","year":"2017","journal-title":"Elife"},{"issue":"9","key":"10.1016\/j.bone.2020.115686_bb0145","doi-asserted-by":"crossref","first-page":"1105","DOI":"10.1038\/ejhg.2014.7","article-title":"Exome sequencing reveals a heterozygous DLX5 mutation in a Chinese family with autosomal-dominant split-hand\/foot malformation","volume":"22","author":"Wang","year":"2014","journal-title":"Eur. J. Hum. Genet."},{"key":"10.1016\/j.bone.2020.115686_bb0150","doi-asserted-by":"crossref","first-page":"17194","DOI":"10.1038\/srep17194","article-title":"ORMDL3 contributes to the risk of atherosclerosis in Chinese Han population and mediates oxidized low-density lipoprotein-induced autophagy in endothelial cells","volume":"5","author":"Ma","year":"2015","journal-title":"Sci. Rep."},{"issue":"6","key":"10.1016\/j.bone.2020.115686_bb0155","doi-asserted-by":"crossref","first-page":"1625","DOI":"10.1093\/emboj\/17.6.1625","article-title":"Crystal structure and mapping by site-directed mutagenesis of the collagen-binding epitope of an activated form of BM-40\/SPARC\/osteonectin","volume":"17","author":"Sasaki","year":"1998","journal-title":"EMBO J."},{"issue":"2","key":"10.1016\/j.bone.2020.115686_bb0160","doi-asserted-by":"crossref","first-page":"1120","DOI":"10.18632\/oncotarget.6683","article-title":"The protein kinase LKB1 negatively regulates bone morphogenetic protein receptor signaling","volume":"7","author":"Raja","year":"2016","journal-title":"Oncotarget"},{"issue":"1","key":"10.1016\/j.bone.2020.115686_bb0165","doi-asserted-by":"crossref","first-page":"248","DOI":"10.1091\/mbc.e07-05-0510","article-title":"The SPARC-related factor SMOC-2 promotes growth factor-induced cyclin D1 expression and DNA synthesis via integrin-linked kinase","volume":"19","author":"Liu","year":"2008","journal-title":"Mol. Biol. Cell"},{"issue":"5","key":"10.1016\/j.bone.2020.115686_bb0170","doi-asserted-by":"crossref","first-page":"549","DOI":"10.1038\/onc.2015.127","article-title":"Induction of the intestinal stem cell signature gene SMOC-2 is required for L1-mediated colon cancer progression","volume":"35","author":"Shvab","year":"2016","journal-title":"Oncogene"},{"issue":"32","key":"10.1016\/j.bone.2020.115686_bb0175","doi-asserted-by":"crossref","first-page":"22855","DOI":"10.1074\/jbc.M513463200","article-title":"The novel SPARC family member SMOC-2 potentiates angiogenic growth factor activity","volume":"281","author":"Rocnik","year":"2006","journal-title":"J. Biol. Chem."},{"key":"10.1016\/j.bone.2020.115686_bb0180","doi-asserted-by":"crossref","first-page":"841","DOI":"10.1016\/j.biopha.2018.03.058","article-title":"Suppression of SMOC2 reduces bleomycin (BLM)-induced pulmonary fibrosis by inhibition of TGF-beta1\/SMADs pathway","volume":"105","author":"Luo","year":"2018","journal-title":"Biomed. Pharmacother."},{"issue":"1","key":"10.1016\/j.bone.2020.115686_bb0185","doi-asserted-by":"crossref","first-page":"48","DOI":"10.1016\/j.bbrc.2018.12.006","article-title":"Secreted modular calcium-binding protein 2 promotes high fat diet (HFD)-induced hepatic steatosis through enhancing lipid deposition, fibrosis and inflammation via targeting TGF-beta1","volume":"509","author":"Yuting","year":"2019","journal-title":"Biochem. Biophys. Res. Commun."},{"key":"10.1016\/j.bone.2020.115686_bb0190","doi-asserted-by":"crossref","first-page":"276","DOI":"10.1016\/j.ebiom.2018.12.044","article-title":"Targeting cancer stem cell signature gene SMOC-2 overcomes chemoresistance and inhibits cell proliferation of endometrial carcinoma","volume":"40","author":"Lu","year":"2019","journal-title":"EBioMedicine"},{"issue":"2","key":"10.1016\/j.bone.2020.115686_bb0195","doi-asserted-by":"crossref","first-page":"683","DOI":"10.1534\/genetics.118.301805","article-title":"The Caenorhabditis elegans SMOC-1 protein acts cell nonautonomously to promote bone morphogenetic protein signaling","volume":"211","author":"Degroot","year":"2019","journal-title":"Genetics"},{"issue":"7","key":"10.1016\/j.bone.2020.115686_bb0200","doi-asserted-by":"crossref","first-page":"817","DOI":"10.1002\/dvdy.23969","article-title":"A morpholino-based screen to identify novel genes involved in craniofacial morphogenesis","volume":"242","author":"Melvin","year":"2013","journal-title":"Dev. Dyn."},{"issue":"6","key":"10.1016\/j.bone.2020.115686_bb0205","doi-asserted-by":"crossref","first-page":"773","DOI":"10.1016\/j.ajhg.2011.11.002","article-title":"Homozygosity mapping and candidate prioritization identify mutations, missed by whole-exome sequencing, in SMOC2, causing major dental developmental defects","volume":"89","author":"Bloch-Zupan","year":"2011","journal-title":"Am. J. Hum. Genet."},{"issue":"11","key":"10.1016\/j.bone.2020.115686_bb0210","doi-asserted-by":"crossref","first-page":"1573","DOI":"10.1016\/j.cub.2017.04.057","article-title":"Canine brachycephaly is associated with a retrotransposon-mediated missplicing of SMOC2","volume":"27","author":"Marchant","year":"2017","journal-title":"Curr. Biol."},{"issue":"1","key":"10.1016\/j.bone.2020.115686_bb0215","doi-asserted-by":"crossref","first-page":"14817","DOI":"10.1038\/s41598-020-71749-6","article-title":"Deficiency of the SMOC2 matricellular protein impairs bone healing and produces age-dependent bone loss","volume":"10","author":"Morkmued","year":"2020","journal-title":"Sci. Rep."}],"container-title":["Bone"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/api.elsevier.com\/content\/article\/PII:S875632822030466X?httpAccept=text\/xml","content-type":"text\/xml","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/api.elsevier.com\/content\/article\/PII:S875632822030466X?httpAccept=text\/plain","content-type":"text\/plain","content-version":"vor","intended-application":"text-mining"}],"deposited":{"date-parts":[[2025,9,11]],"date-time":"2025-09-11T17:51:11Z","timestamp":1757613071000},"score":1,"resource":{"primary":{"URL":"https:\/\/linkinghub.elsevier.com\/retrieve\/pii\/S875632822030466X"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2021,1]]},"references-count":43,"alternative-id":["S875632822030466X"],"URL":"https:\/\/doi.org\/10.1016\/j.bone.2020.115686","relation":{},"ISSN":["8756-3282"],"issn-type":[{"value":"8756-3282","type":"print"}],"subject":[],"published":{"date-parts":[[2021,1]]},"assertion":[{"value":"Elsevier","name":"publisher","label":"This article is maintained by"},{"value":"A SMOC2 variant inhibits BMP signaling by competitively binding to BMPR1B and causes growth plate defects","name":"articletitle","label":"Article Title"},{"value":"Bone","name":"journaltitle","label":"Journal Title"},{"value":"https:\/\/doi.org\/10.1016\/j.bone.2020.115686","name":"articlelink","label":"CrossRef DOI link to publisher maintained version"},{"value":"article","name":"content_type","label":"Content Type"},{"value":"\u00a9 2020 Elsevier Inc. All rights reserved.","name":"copyright","label":"Copyright"}],"article-number":"115686"}}