{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,11,9]],"date-time":"2026-11-09T12:23:55Z","timestamp":1794227035949,"version":"4.9.0"},"reference-count":62,"publisher":"Springer Science and Business Media LLC","issue":"4","license":[{"start":{"date-parts":[[2022,2,10]],"date-time":"2022-02-10T00:00:00Z","timestamp":1644451200000},"content-version":"tdm","delay-in-days":0,"URL":"https:\/\/www.springernature.com\/gp\/researchers\/text-and-data-mining"},{"start":{"date-parts":[[2022,2,10]],"date-time":"2022-02-10T00:00:00Z","timestamp":1644451200000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/www.springernature.com\/gp\/researchers\/text-and-data-mining"}],"content-domain":{"domain":["link.springer.com"],"crossmark-restriction":false},"short-container-title":["Nat Aging"],"DOI":"10.1038\/s43587-021-00165-w","type":"journal-article","created":{"date-parts":[[2022,2,10]],"date-time":"2022-02-10T12:02:31Z","timestamp":1644494551000},"page":"332-347","update-policy":"https:\/\/doi.org\/10.1007\/springer_crossmark_policy","source":"Crossref","is-referenced-by-count":98,"title":["Kindlin-2 preserves integrity of the articular cartilage to protect against osteoarthritis"],"prefix":"10.1038","volume":"2","author":[{"given":"Xiaohao","family":"Wu","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Yumei","family":"Lai","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-1770-5018","authenticated-orcid":false,"given":"Sheng","family":"Chen","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Chunlei","family":"Zhou","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Chu","family":"Tao","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-8208-9939","authenticated-orcid":false,"given":"Xuekun","family":"Fu","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Jun","family":"Li","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Wei","family":"Tong","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Hongtao","family":"Tian","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Zengwu","family":"Shao","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Chuanju","family":"Liu","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Di","family":"Chen","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-9631-4781","authenticated-orcid":false,"given":"Xiaochun","family":"Bai","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-7728-1774","authenticated-orcid":false,"given":"Huiling","family":"Cao","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-4269-2450","authenticated-orcid":false,"given":"Guozhi","family":"Xiao","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"297","published-online":{"date-parts":[[2022,2,10]]},"reference":[{"key":"165_CR1","doi-asserted-by":"publisher","first-page":"1745","DOI":"10.1016\/S0140-6736(19)30417-9","volume":"393","author":"DJ Hunter","year":"2019","unstructured":"Hunter, D. J. & Bierma-Zeinstra, S. Osteoarthritis. Lancet 393, 1745\u20131759 (2019).","journal-title":"Lancet"},{"key":"165_CR2","doi-asserted-by":"publisher","unstructured":"He, Y. et al. Pathogenesis of osteoarthritis: risk factors, regulatory pathways in chondrocytes, and experimental models. Biology https:\/\/doi.org\/10.3390\/biology9080194 (2020).","DOI":"10.3390\/biology9080194"},{"key":"165_CR3","doi-asserted-by":"publisher","first-page":"74","DOI":"10.1002\/bdrc.21062","volume":"102","author":"MM Sun","year":"2014","unstructured":"Sun, M. M. & Beier, F. Chondrocyte hypertrophy in skeletal development, growth, and disease. Birth Defects Res. C Embryo Today 102, 74\u201382 (2014).","journal-title":"Birth Defects Res. C Embryo Today"},{"key":"165_CR4","doi-asserted-by":"publisher","first-page":"16044","DOI":"10.1038\/boneres.2016.44","volume":"5","author":"D Chen","year":"2017","unstructured":"Chen, D. et al. Osteoarthritis: toward a comprehensive understanding of pathological mechanism. Bone Res. 5, 16044 (2017).","journal-title":"Bone Res."},{"key":"165_CR5","doi-asserted-by":"publisher","unstructured":"Song, E. K. et al. ITGBL1 modulates integrin activity to promote cartilage formation and protect against arthritis. Sci. Transl. Med. https:\/\/doi.org\/10.1126\/scitranslmed.aam7486 (2018).","DOI":"10.1126\/scitranslmed.aam7486"},{"key":"165_CR6","doi-asserted-by":"publisher","first-page":"35","DOI":"10.1038\/s41413-020-00110-4","volume":"8","author":"E Charlier","year":"2020","unstructured":"Charlier, E. et al. Toward diagnostic relevance of the \u03b1V\u03b25, \u03b1V\u03b23, and \u03b1V\u03b26 integrins in OA: expression within human cartilage and spinal osteophytes. Bone Res. 8, 35 (2020).","journal-title":"Bone Res."},{"key":"165_CR7","doi-asserted-by":"publisher","first-page":"e128616","DOI":"10.1172\/jci.insight.128616","volume":"4","author":"Q Wang","year":"2019","unstructured":"Wang, Q. et al. Dysregulated integrin \u03b1V\u03b23 and CD47 signaling promotes joint inflammation, cartilage breakdown, and progression of osteoarthritis. JCI Insight 4, e128616 (2019).","journal-title":"JCI Insight"},{"key":"165_CR8","doi-asserted-by":"publisher","first-page":"833","DOI":"10.1083\/jcb.200211089","volume":"162","author":"Q Zheng","year":"2003","unstructured":"Zheng, Q. et al. Type X collagen gene regulation by Runx2 contributes directly to its hypertrophic chondrocyte-specific expression in vivo. J. Cell Biol. 162, 833\u2013842 (2003).","journal-title":"J. Cell Biol."},{"key":"165_CR9","doi-asserted-by":"publisher","first-page":"2899","DOI":"10.1002\/jbmr.504","volume":"26","author":"F Li","year":"2011","unstructured":"Li, F. et al. Runx2 contributes to murine Col10a1 gene regulation through direct interaction with its cis-enhancer. J. Bone Miner. Res. 26, 2899\u20132910 (2011).","journal-title":"J. Bone Miner. Res."},{"key":"165_CR10","doi-asserted-by":"publisher","DOI":"10.1038\/s41598-017-02490-w","volume":"7","author":"L Liao","year":"2017","unstructured":"Liao, L. et al. Deletion of Runx2 in articular chondrocytes decelerates the progression of DMM-induced osteoarthritis in adult mice. Sci. Rep. 7, 2371 (2017).","journal-title":"Sci. Rep."},{"key":"165_CR11","doi-asserted-by":"publisher","DOI":"10.1038\/s41467-019-10753-5","volume":"10","author":"J Huang","year":"2019","unstructured":"Huang, J. et al. The microRNAs miR-204 and miR-211 maintain joint homeostasis and protect against osteoarthritis progression. Nat. Commun. 10, 2876 (2019).","journal-title":"Nat. Commun."},{"key":"165_CR12","doi-asserted-by":"publisher","first-page":"1676","DOI":"10.1002\/jbmr.3737","volume":"34","author":"SE Catheline","year":"2019","unstructured":"Catheline, S. E. et al. Chondrocyte-specific RUNX2 overexpression accelerates post-traumatic osteoarthritis progression in adult mice. J. Bone Miner. Res. 34, 1676\u20131689 (2019).","journal-title":"J. Bone Miner. Res."},{"key":"165_CR13","doi-asserted-by":"publisher","first-page":"3464","DOI":"10.1093\/hmg\/ddy257","volume":"27","author":"SJ Rice","year":"2018","unstructured":"Rice, S. J. et al. Identification of a novel, methylation-dependent, RUNX2 regulatory region associated with osteoarthritis risk. Hum. Mol. Genet. 27, 3464\u20133474 (2018).","journal-title":"Hum. Mol. Genet."},{"key":"165_CR14","doi-asserted-by":"publisher","first-page":"132","DOI":"10.1016\/j.jot.2019.11.008","volume":"23","author":"D Chen","year":"2020","unstructured":"Chen, D., Kim, D. J., Shen, J., Zou, Z. & O\u2019Keefe, R. J. Runx2 plays a central role in osteoarthritis development. J. Orthop. Translat. 23, 132\u2013139 (2020).","journal-title":"J. Orthop. Translat."},{"key":"165_CR15","doi-asserted-by":"publisher","first-page":"748","DOI":"10.1136\/annrheumdis-2016-209757","volume":"76","author":"A Latourte","year":"2017","unstructured":"Latourte, A. et al. Systemic inhibition of IL-6\/Stat3 signalling protects against experimental osteoarthritis. Ann. Rheum. Dis. 76, 748\u2013755 (2017).","journal-title":"Ann. Rheum. Dis."},{"key":"165_CR16","doi-asserted-by":"publisher","first-page":"1032","DOI":"10.3892\/ijo.2014.2259","volume":"44","author":"SW Wang","year":"2014","unstructured":"Wang, S. W. & Sun, Y. M. The IL-6\/JAK\/STAT3 pathway: potential therapeutic strategies in treating colorectal cancer (review). Int. J. Oncol. 44, 1032\u20131040 (2014).","journal-title":"Int. J. Oncol."},{"key":"165_CR17","first-page":"13","volume":"13","author":"S Miscia","year":"2002","unstructured":"Miscia, S. et al. Tumor necrosis factor \u03b1 (TNF-\u03b1) activates Jak1\/Stat3-Stat5B signaling through TNFR-1 in human B cells. Cell Growth Differ. 13, 13\u201318 (2002).","journal-title":"Cell Growth Differ"},{"key":"165_CR18","doi-asserted-by":"publisher","first-page":"183206","DOI":"10.1016\/j.bbamem.2020.183206","volume":"1862","author":"YA Kadry","year":"2020","unstructured":"Kadry, Y. A. & Calderwood, D. A. Chapter 22: structural and signaling functions of integrins. Biochim. Biophys. Acta Biomembr. 1862, 183206 (2020).","journal-title":"Biochim. Biophys. Acta Biomembr."},{"key":"165_CR19","doi-asserted-by":"publisher","first-page":"25","DOI":"10.1038\/s41556-018-0234-9","volume":"21","author":"Z Sun","year":"2019","unstructured":"Sun, Z., Costell, M. & Fassler, R. Integrin activation by talin, kindlin and mechanical forces. Nat. Cell Biol. 21, 25\u201331 (2019).","journal-title":"Nat. Cell Biol."},{"key":"165_CR20","doi-asserted-by":"publisher","first-page":"31","DOI":"10.1016\/j.ceb.2019.12.008","volume":"63","author":"M Michael","year":"2020","unstructured":"Michael, M. & Parsons, M. New perspectives on integrin-dependent adhesions. Curr. Opin. Cell Biol. 63, 31\u201337 (2020).","journal-title":"Curr. Opin. Cell Biol."},{"key":"165_CR21","doi-asserted-by":"publisher","first-page":"17","DOI":"10.1242\/jcs.161190","volume":"129","author":"E Rognoni","year":"2016","unstructured":"Rognoni, E., Ruppert, R. & Fassler, R. The kindlin family: functions, signaling properties and implications for human disease. J. Cell Sci. 129, 17\u201327 (2016).","journal-title":"J. Cell Sci."},{"key":"165_CR22","doi-asserted-by":"publisher","first-page":"202","DOI":"10.1161\/CIRCRESAHA.118.314362","volume":"124","author":"EF Plow","year":"2019","unstructured":"Plow, E. F. & Qin, J. The kindlin family of adapter proteins. Circ. Res. 124, 202\u2013204 (2019).","journal-title":"Circ. Res."},{"key":"165_CR23","doi-asserted-by":"publisher","first-page":"306","DOI":"10.1038\/nm.1931","volume":"15","author":"L Svensson","year":"2009","unstructured":"Svensson, L. et al. Leukocyte adhesion deficiency-III is caused by mutations in KINDLIN3 affecting integrin activation. Nat. Med. 15, 306\u2013312 (2009).","journal-title":"Nat. Med."},{"key":"165_CR24","doi-asserted-by":"publisher","first-page":"325","DOI":"10.1038\/nm1722","volume":"14","author":"M Moser","year":"2008","unstructured":"Moser, M., Nieswandt, B., Ussar, S., Pozgajova, M. & Fassler, R. Kindlin-3 is essential for integrin activation and platelet aggregation. Nat. Med. 14, 325\u2013330 (2008).","journal-title":"Nat. Med."},{"key":"165_CR25","doi-asserted-by":"publisher","first-page":"883","DOI":"10.1083\/jcb.201007141","volume":"192","author":"S Schmidt","year":"2011","unstructured":"Schmidt, S. et al. Kindlin-3-mediated signaling from multiple integrin classes is required for osteoclast-mediated bone resorption. J. Cell Biol. 192, 883\u2013897 (2011).","journal-title":"J. Cell Biol."},{"key":"165_CR26","doi-asserted-by":"publisher","first-page":"313","DOI":"10.1038\/nm.1917","volume":"15","author":"NL Malinin","year":"2009","unstructured":"Malinin, N. L. et al. A point mutation in KINDLIN3 ablates activation of three integrin subfamilies in humans. Nat. Med. 15, 313\u2013318 (2009).","journal-title":"Nat. Med."},{"key":"165_CR27","doi-asserted-by":"publisher","first-page":"1325","DOI":"10.1101\/gad.469408","volume":"22","author":"E Montanez","year":"2008","unstructured":"Montanez, E. et al. Kindlin-2 controls bidirectional signaling of integrins. Genes Dev. 22, 1325\u20131330 (2008).","journal-title":"Genes Dev."},{"key":"165_CR28","doi-asserted-by":"publisher","first-page":"93","DOI":"10.1016\/j.biocel.2018.03.008","volume":"98","author":"J Zhan","year":"2018","unstructured":"Zhan, J. & Zhang, H. Kindlins: roles in development and cancer progression. Int. J. Biochem. Cell Biol. 98, 93\u2013103 (2018).","journal-title":"Int. J. Biochem. Cell Biol."},{"key":"165_CR29","doi-asserted-by":"publisher","unstructured":"Wei, X. et al. Kindlin-2 mediates activation of TGF-\u03b2\/Smad signaling and renal fibrosis. J. Am. Soc. Nephrol. https:\/\/doi.org\/10.1681\/ASN.2012101041 (2013).","DOI":"10.1681\/ASN.2012101041"},{"key":"165_CR30","doi-asserted-by":"publisher","first-page":"3545","DOI":"10.1681\/ASN.2016091021","volume":"28","author":"Y Sun","year":"2017","unstructured":"Sun, Y. et al. Kindlin-2 association with Rho GDP-dissociation inhibitor \u03b1 suppresses Rac1 activation and podocyte injury. J. Am. Soc. Nephrol. 28, 3545\u20133562 (2017).","journal-title":"J. Am. Soc. Nephrol."},{"key":"165_CR31","doi-asserted-by":"publisher","DOI":"10.1038\/s41467-019-14186-y","volume":"11","author":"K Zhu","year":"2020","unstructured":"Zhu, K. et al. Kindlin-2 modulates MafA and \u03b2-catenin expression to regulate \u03b2-cell function and mass in mice. Nat. Commun. 11, 484 (2020).","journal-title":"Nat. Commun."},{"key":"165_CR32","doi-asserted-by":"publisher","first-page":"4971","DOI":"10.1002\/cam4.3130","volume":"9","author":"X Xue","year":"2020","unstructured":"Xue, X., Xue, S., Wan, W., Li, J. & Shi, H. HIF-1\u03b1 interacts with kindlin-2 and influences breast cancer elasticity: a study based on shear wave elastography imaging. Cancer Med. 9, 4971\u20134979 (2020).","journal-title":"Cancer Med."},{"key":"165_CR33","doi-asserted-by":"publisher","first-page":"271","DOI":"10.1016\/j.canlet.2015.03.011","volume":"361","author":"B Guo","year":"2015","unstructured":"Guo, B., Gao, J., Zhan, J. & Zhang, H. Kindlin-2 interacts with and stabilizes EGFR and is required for EGF-induced breast cancer cell migration. Cancer Lett. 361, 271\u2013281 (2015).","journal-title":"Cancer Lett."},{"key":"165_CR34","doi-asserted-by":"publisher","DOI":"10.1038\/s41467-019-08772-3","volume":"10","author":"L Guo","year":"2019","unstructured":"Guo, L. et al. Kindlin-2 links mechano-environment to proline synthesis and tumor growth. Nat. Commun. 10, 845 (2019).","journal-title":"Nat. Commun."},{"key":"165_CR35","doi-asserted-by":"publisher","DOI":"10.1038\/s41467-020-18753-6","volume":"11","author":"L Guo","year":"2020","unstructured":"Guo, L. et al. PINCH-1 regulates mitochondrial dynamics to promote proline synthesis and tumor growth. Nat. Commun. 11, 4913 (2020).","journal-title":"Nat. Commun."},{"key":"165_CR36","doi-asserted-by":"publisher","first-page":"23","DOI":"10.1038\/s41413-020-0099-y","volume":"8","author":"L Qin","year":"2020","unstructured":"Qin, L., Liu, W., Cao, H. & Xiao, G. Molecular mechanosensors in osteocytes. Bone Res. 8, 23 (2020).","journal-title":"Bone Res."},{"key":"165_CR37","doi-asserted-by":"publisher","first-page":"6182","DOI":"10.7150\/thno.46553","volume":"10","author":"X He","year":"2020","unstructured":"He, X. et al. Kindlin-2 deficiency induces fatal intestinal obstruction in mice. Theranostics 10, 6182\u20136200 (2020).","journal-title":"Theranostics"},{"key":"165_CR38","doi-asserted-by":"publisher","DOI":"10.1038\/s41419-019-2121-0","volume":"10","author":"L Qi","year":"2019","unstructured":"Qi, L. et al. Kindlin-2 suppresses transcription factor GATA4 through interaction with SUV39H1 to attenuate hypertrophy. Cell Death Dis. 10, 890 (2019).","journal-title":"Cell Death Dis."},{"key":"165_CR39","doi-asserted-by":"publisher","first-page":"e128405","DOI":"10.1172\/jci.insight.128405","volume":"4","author":"H Gao","year":"2019","unstructured":"Gao, H. et al. Lipoatrophy and metabolic disturbance in mice with adipose-specific deletion of kindlin-2. JCI Insight 4, e128405 (2019).","journal-title":"JCI Insight"},{"key":"165_CR40","doi-asserted-by":"publisher","first-page":"1554","DOI":"10.1161\/CIRCULATIONAHA.118.038383","volume":"139","author":"Z Zhang","year":"2019","unstructured":"Zhang, Z. et al. Kindlin-2 is essential for preserving integrity of the developing heart and preventing ventricular rupture. Circulation 139, 1554\u20131556 (2019).","journal-title":"Circulation"},{"key":"165_CR41","doi-asserted-by":"publisher","DOI":"10.1038\/ncomms8531","volume":"6","author":"C Wu","year":"2015","unstructured":"Wu, C. et al. Kindlin-2 controls TGF-\u03b2 signalling and Sox9 expression to regulate chondrogenesis. Nat. Commun. 6, 7531 (2015).","journal-title":"Nat. Commun."},{"key":"165_CR42","doi-asserted-by":"publisher","first-page":"2","DOI":"10.1038\/s41413-019-0073-8","volume":"8","author":"H Cao","year":"2020","unstructured":"Cao, H. et al. Focal adhesion protein kindlin-2 regulates bone homeostasis in mice. Bone Res. 8, 2 (2020).","journal-title":"Bone Res."},{"key":"165_CR43","doi-asserted-by":"publisher","first-page":"402","DOI":"10.1038\/s42003-021-01950-4","volume":"4","author":"L Qin","year":"2021","unstructured":"Qin, L. et al. Kindlin-2 mediates mechanotransduction in bone by regulating expression of sclerostin in osteocytes. Commun. Biol. 4, 402 (2021).","journal-title":"Commun. Biol."},{"key":"165_CR44","doi-asserted-by":"publisher","first-page":"297","DOI":"10.1038\/s41392-020-00328-y","volume":"5","author":"X Fu","year":"2020","unstructured":"Fu, X. et al. Kindlin-2 regulates skeletal homeostasis by modulating PTH1R in mice. Signal Transduct. Target Ther. 5, 297 (2020).","journal-title":"Signal Transduct. Target Ther."},{"key":"165_CR45","doi-asserted-by":"publisher","first-page":"921","DOI":"10.1007\/s00018-016-2394-8","volume":"74","author":"G Dalagiorgou","year":"2017","unstructured":"Dalagiorgou, G. et al. Mechanosensor polycystin-1 potentiates differentiation of human osteoblastic cells by upregulating Runx2 expression via induction of JAK2\/STAT3 signaling axis. Cell. Mol. Life Sci. 74, 921\u2013936 (2017).","journal-title":"Cell. Mol. Life Sci."},{"key":"165_CR46","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1042\/BJ20081386","volume":"417","author":"MP Murphy","year":"2009","unstructured":"Murphy, M. P. How mitochondria produce reactive oxygen species. Biochem. J. 417, 1\u201313 (2009).","journal-title":"Biochem. J"},{"key":"165_CR47","doi-asserted-by":"publisher","first-page":"3974","DOI":"10.1111\/jcmm.14283","volume":"23","author":"S Perez-Garcia","year":"2019","unstructured":"Perez-Garcia, S. et al. Wnt and RUNX2 mediate cartilage breakdown by osteoarthritis synovial fibroblast-derived ADAMTS-7 and -12. J. Cell. Mol. Med. 23, 3974\u20133983 (2019).","journal-title":"J. Cell. Mol. Med."},{"key":"165_CR48","doi-asserted-by":"publisher","first-page":"3186","DOI":"10.1111\/j.1582-4934.2008.00678.x","volume":"13","author":"T Orfanidou","year":"2009","unstructured":"Orfanidou, T., Iliopoulos, D., Malizos, K. N. & Tsezou, A. Involvement of SOX-9 and FGF-23 in RUNX-2 regulation in osteoarthritic chondrocytes. J. Cell. Mol. Med. 13, 3186\u20133194 (2009).","journal-title":"J. Cell. Mol. Med."},{"key":"165_CR49","doi-asserted-by":"publisher","first-page":"486","DOI":"10.1097\/MAJ.0b013e3182579dc6","volume":"344","author":"D Li","year":"2012","unstructured":"Li, D., Xie, G. & Wang, W. Reactive oxygen species: the 2-edged sword of osteoarthritis. Am. J. Med. Sci. 344, 486\u2013490 (2012).","journal-title":"Am. J. Med. Sci."},{"key":"165_CR50","doi-asserted-by":"publisher","first-page":"805","DOI":"10.1002\/dvg.20564","volume":"47","author":"SP Henry","year":"2009","unstructured":"Henry, S. P. et al. Generation of aggrecan-CreERT2 knockin mice for inducible Cre activity in adult cartilage. Genesis 47, 805\u2013814 (2009).","journal-title":"Genesis"},{"key":"165_CR51","doi-asserted-by":"publisher","first-page":"2465","DOI":"10.1101\/gad.277003","volume":"17","author":"A Aszodi","year":"2003","unstructured":"Aszodi, A., Hunziker, E. B., Brakebusch, C. & Fassler, R. \u03b21 integrins regulate chondrocyte rotation, G1 progression, and cytokinesis. Genes Dev. 17, 2465\u20132479 (2003).","journal-title":"Genes Dev."},{"key":"165_CR52","doi-asserted-by":"publisher","first-page":"23780","DOI":"10.1074\/jbc.M109.039347","volume":"284","author":"A Raducanu","year":"2009","unstructured":"Raducanu, A., Hunziker, E. B., Drosse, I. & Aszodi, A. \u03b21 integrin deficiency results in multiple abnormalities of the knee joint. J. Biol. Chem. 284, 23780\u201323792 (2009).","journal-title":"J. Biol. Chem."},{"key":"165_CR53","doi-asserted-by":"publisher","first-page":"18938","DOI":"10.1016\/S0021-9258(18)37372-1","volume":"263","author":"RT Franceschi","year":"1988","unstructured":"Franceschi, R. T., Romano, P. R. & Park, K. Y. Regulation of type I collagen synthesis by 1,25-dihydroxyvitamin D3 in human osteosarcoma cells. J. Biol. Chem. 263, 18938\u201318945 (1988).","journal-title":"J. Biol. Chem."},{"key":"165_CR54","doi-asserted-by":"publisher","first-page":"6679","DOI":"10.1002\/jcp.27410","volume":"234","author":"L Liao","year":"2019","unstructured":"Liao, L. et al. Runx2 is required for postnatal intervertebral disc tissue growth and development. J. Cell. Physiol. 234, 6679\u20136687 (2019).","journal-title":"J. Cell. Physiol."},{"key":"165_CR55","doi-asserted-by":"publisher","first-page":"1018","DOI":"10.1038\/labinvest.3700630","volume":"87","author":"A Moh","year":"2007","unstructured":"Moh, A. et al. Role of STAT3 in liver regeneration: survival, DNA synthesis, inflammatory reaction and liver mass recovery. Lab. Invest. 87, 1018\u20131028 (2007).","journal-title":"Lab. Invest."},{"key":"165_CR56","doi-asserted-by":"publisher","first-page":"635","DOI":"10.1136\/annrheumdis-2019-216713","volume":"79","author":"J Li","year":"2020","unstructured":"Li, J. et al. Metformin limits osteoarthritis development and progression through activation of AMPK signalling. Ann. Rheum. Dis. 79, 635\u2013645 (2020).","journal-title":"Ann. Rheum. Dis."},{"key":"165_CR57","doi-asserted-by":"publisher","first-page":"55","DOI":"10.1016\/0165-0270(94)90144-9","volume":"53","author":"SR Chaplan","year":"1994","unstructured":"Chaplan, S. R., Bach, F. W., Pogrel, J. W., Chung, J. M. & Yaksh, T. L. Quantitative assessment of tactile allodynia in the rat paw. J. Neurosci. Methods 53, 55\u201363 (1994).","journal-title":"J. Neurosci. Methods"},{"key":"165_CR58","doi-asserted-by":"publisher","first-page":"1253","DOI":"10.1038\/nprot.2008.95","volume":"3","author":"M Gosset","year":"2008","unstructured":"Gosset, M., Berenbaum, F., Thirion, S. & Jacques, C. Primary culture and phenotyping of murine chondrocytes. Nat. Protoc. 3, 1253\u20131260 (2008).","journal-title":"Nat. Protoc."},{"key":"165_CR59","doi-asserted-by":"publisher","first-page":"374","DOI":"10.1093\/cvr\/cvv279","volume":"109","author":"L Valls-Lacalle","year":"2016","unstructured":"Valls-Lacalle, L. et al. Succinate dehydrogenase inhibition with malonate during reperfusion reduces infarct size by preventing mitochondrial permeability transition. Cardiovasc. Res. 109, 374\u2013384 (2016).","journal-title":"Cardiovasc. Res."},{"key":"165_CR60","doi-asserted-by":"publisher","DOI":"10.1186\/s12891-017-1621-2","volume":"18","author":"C Vaamonde-Garcia","year":"2017","unstructured":"Vaamonde-Garcia, C. et al. The mitochondrial inhibitor oligomycin induces an inflammatory response in the rat knee joint. BMC Musculoskelet. Disord. 18, 254 (2017).","journal-title":"BMC Musculoskelet. Disord."},{"key":"165_CR61","doi-asserted-by":"publisher","first-page":"30689","DOI":"10.1074\/jbc.M500750200","volume":"280","author":"G Xiao","year":"2005","unstructured":"Xiao, G. et al. cooperative interactions between activating transcription Factor 4 and Runx2\/Cbfa1 stimulate osteoblast-specific osteocalcin gene expression. J. Biol. Chem. 280, 30689\u201330696 (2005).","journal-title":"J. Biol. Chem."},{"key":"165_CR62","doi-asserted-by":"publisher","first-page":"37","DOI":"10.1038\/s41413-020-00108-y","volume":"8","author":"Y Lei","year":"2020","unstructured":"Lei, Y. et al. LIM domain proteins Pinch1\/2 regulate chondrogenesis and bone mass in mice. Bone Res. 8, 37 (2020).","journal-title":"Bone Res."}],"container-title":["Nature Aging"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.nature.com\/articles\/s43587-021-00165-w.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/www.nature.com\/articles\/s43587-021-00165-w","content-type":"text\/html","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/www.nature.com\/articles\/s43587-021-00165-w.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,11,9]],"date-time":"2026-11-09T11:48:44Z","timestamp":1794224924000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.nature.com\/articles\/s43587-021-00165-w"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2022,2,10]]},"references-count":62,"journal-issue":{"issue":"4","published-online":{"date-parts":[[2022,4]]}},"alternative-id":["165"],"URL":"https:\/\/doi.org\/10.1038\/s43587-021-00165-w","relation":{"has-preprint":[{"id-type":"doi","id":"10.1101\/2021.08.11.456023","asserted-by":"object"}]},"ISSN":["2662-8465"],"issn-type":[{"value":"2662-8465","type":"electronic"}],"subject":[],"published":{"date-parts":[[2022,2,10]]},"assertion":[{"value":"14 January 2021","order":1,"name":"received","label":"Received","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"21 December 2021","order":2,"name":"accepted","label":"Accepted","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"10 February 2022","order":3,"name":"first_online","label":"First Online","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"9 November 2026","order":5,"name":"change_date","label":"Change Date","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"Update","order":6,"name":"change_type","label":"Change Type","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"In the version of the article initially published, in the Fig. 3t source data, the values in the Stat3 and Runx2 tables were inadvertently duplicated from the Kindlin-2 (K2) dataset. The published graphs in Fig. 3t were generated using the correct data, and the error does not affect the reported results or the scientific conclusions of the article. The corrected Source Data file is now available online.","order":7,"name":"change_details","label":"Change Details","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"The authors declare no competing interests.","order":1,"name":"Ethics","label":"Competing interests","group":{"name":"EthicsHeading","label":"Ethics"}}]}}