{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,11,4]],"date-time":"2026-11-04T06:49:42Z","timestamp":1793774982639,"version":"4.6.1"},"reference-count":57,"publisher":"Springer Science and Business Media LLC","issue":"1","license":[{"start":{"date-parts":[[2026,5,14]],"date-time":"2026-05-14T00:00:00Z","timestamp":1778716800000},"content-version":"tdm","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0"},{"start":{"date-parts":[[2026,5,14]],"date-time":"2026-05-14T00:00:00Z","timestamp":1778716800000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0"}],"funder":[{"DOI":"10.13039\/501100001809","name":"National Natural Science Foundation of China","doi-asserted-by":"publisher","award":["82271017"],"award-info":[{"award-number":["82271017"]}],"id":[{"id":"10.13039\/501100001809","id-type":"DOI","asserted-by":"publisher"}]}],"content-domain":{"domain":["link.springer.com"],"crossmark-restriction":false},"short-container-title":["Bone Res"],"abstract":"<jats:title>Abstract<\/jats:title>\n                  <jats:p>Orthodontically induced inflammatory root resorption (OIIRR) is a prevalent complication driven by excessive mechanical force, yet the underlying mechanisms linking mechanotransduction to osteoclast activation remain elusive. Here, we identify a novel signaling axis wherein sphingosine kinase 1 (SphK1) in cementocytes translates heavy orthodontic force into a pro-osteoclastogenic signal via mitophagy-mediated mitochondrial transfer. In vivo, heavy force induced OIIRR and upregulated mitophagy markers in cementocytes. In vitro, heavy compression force triggered SphK1-dependent mitophagy in IDG-CM6 cementocytes, as evidenced by increased mitophagosome formation, co-localization of mitochondria with lysosomes, and elevated PINK1\/PARKIN signaling. Inhibition of SphK1, either pharmacologically or genetically, suppressed this mitophagic response. Conditioned media from force-loaded cementocytes enhanced osteoclast differentiation and glycolytic metabolism, effects that were abolished by SphK1 inhibition and rescued by a mitophagy agonist. Crucially, we demonstrated that heavy force promotes the transfer of mitochondria from cementocytes to osteoclast precursors, a process dependent on mitophagy. This transferred mitochondrial cargo functioned as a metabolic subsidy, boosting osteoclast bioenergetics and resorptive activity. Our findings unveil the SphK1-mitophagy-mitochondrial transfer axis as a fundamental mechanism of cementocyte-osteoclast communication, positioning SphK1 as a promising therapeutic target to prevent OIIRR.<\/jats:p>","DOI":"10.1038\/s41413-026-00538-0","type":"journal-article","created":{"date-parts":[[2026,5,14]],"date-time":"2026-05-14T07:04:49Z","timestamp":1778742289000},"update-policy":"https:\/\/doi.org\/10.1007\/springer_crossmark_policy","source":"Crossref","is-referenced-by-count":3,"title":["SphK1\/mitophagy axis in cementocytes drives orthodontic root resorption via mitochondrial transfer to osteoclasts"],"prefix":"10.1038","volume":"14","author":[{"given":"Han","family":"Wang","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Sihang","family":"Chen","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Shuo","family":"Chen","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Chengchen","family":"Duan","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Li","family":"Zhu","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Hengyi","family":"Lin","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Shujuan","family":"Zou","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Yu","family":"Li","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Peipei","family":"Duan","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"297","published-online":{"date-parts":[[2026,5,14]]},"reference":[{"key":"538_CR1","doi-asserted-by":"publisher","first-page":"149","DOI":"10.1148\/radiol.2019190070","volume":"294","author":"X Miao","year":"2020","unstructured":"Miao, X. et al. Whole-brain functional and diffusion tensor MRI in human participants with metallic orthodontic braces. Radiology 294, 149\u2013157 (2020).","journal-title":"Radiology"},{"key":"538_CR2","doi-asserted-by":"publisher","first-page":"238","DOI":"10.1016\/j.ajodo.2021.03.023","volume":"162","author":"C Mann","year":"2022","unstructured":"Mann, C. et al. Physical properties of root cementum: part 28. Effects of high and low water fluoridation on orthodontic root resorption: a microcomputed tomography study. Am. J. Orthod. Dentofac. Orthop. 162, 238\u2013246 (2022).","journal-title":"Am. J. Orthod. Dentofac. Orthop."},{"key":"538_CR3","doi-asserted-by":"publisher","first-page":"86","DOI":"10.1111\/adj.12486","volume":"62","author":"M Wishney","year":"2017","unstructured":"Wishney, M. Potential risks of orthodontic therapy: a critical review and conceptual framework. Aust. Dent. J. 62, 86\u201396 (2017).","journal-title":"Aust. Dent. J."},{"key":"538_CR4","doi-asserted-by":"publisher","DOI":"10.1186\/s12903-019-0778-5","volume":"19","author":"N Okamoto","year":"2019","unstructured":"Okamoto, N., Amano, N., Nakamura, T. & Yanagi, M. Relationship between tooth loss, low masticatory ability, and nutritional indices in the elderly: a cross-sectional study. BMC Oral. Health 19, 110 (2019).","journal-title":"BMC Oral. Health"},{"key":"538_CR5","doi-asserted-by":"crossref","unstructured":"Sameshima, G. T. & Iglesias-Linares, A. Orthodontic root resorption. J. World Fed. Orthod. 10, 135\u2013143 (2021).","DOI":"10.1016\/j.ejwf.2021.09.003"},{"key":"538_CR6","doi-asserted-by":"crossref","unstructured":"Yamaguchi, M. & Fukasawa, S. Is inflammation a friend or foe for orthodontic treatment? Inflammation in orthodontically induced inflammatory root resorption and accelerating tooth movement. Int. J. Mol. Sci. 22, 2388 (2021).","DOI":"10.3390\/ijms22052388"},{"key":"538_CR7","doi-asserted-by":"publisher","first-page":"1376","DOI":"10.1177\/00220345231195765","volume":"102","author":"H Wang","year":"2023","unstructured":"Wang, H. et al. Force-loaded cementocytes regulate osteoclastogenesis via S1P\/S1PR1\/Rac1 axis. J. Dent. Res. 102, 1376\u20131386 (2023).","journal-title":"J. Dent. Res."},{"key":"538_CR8","doi-asserted-by":"publisher","first-page":"145","DOI":"10.1177\/0022034516677539","volume":"96","author":"A Iglesias-Linares","year":"2017","unstructured":"Iglesias-Linares, A. & Hartsfield, J. K. Jr Cellular and molecular pathways leading to external root resorption. J. Dent. Res. 96, 145\u2013152 (2017).","journal-title":"J. Dent. Res."},{"key":"538_CR9","doi-asserted-by":"publisher","first-page":"860","DOI":"10.1177\/0022034513500306","volume":"92","author":"BF Boyce","year":"2013","unstructured":"Boyce, B. F. Advances in the regulation of osteoclasts and osteoclast functions. J. Dent. Res. 92, 860\u2013867 (2013).","journal-title":"J. Dent. Res."},{"key":"538_CR10","doi-asserted-by":"publisher","first-page":"5840","DOI":"10.1038\/sj.emboj.7601430","volume":"25","author":"J Ryu","year":"2006","unstructured":"Ryu, J. et al. Sphingosine 1-phosphate as a regulator of osteoclast differentiation and osteoclast-osteoblast coupling. EMBO J. 25, 5840\u20135851 (2006).","journal-title":"EMBO J."},{"key":"538_CR11","doi-asserted-by":"publisher","first-page":"474","DOI":"10.1111\/jcpe.13914","volume":"51","author":"X Li","year":"2024","unstructured":"Li, X. et al. NLRP3-mediated periodontal ligament cell pyroptosis promotes root resorption. J. Clin. Periodontol. 51, 474\u2013486 (2024).","journal-title":"J. Clin. Periodontol."},{"key":"538_CR12","doi-asserted-by":"publisher","first-page":"804","DOI":"10.2319\/072418-536.1","volume":"89","author":"Q Li","year":"2019","unstructured":"Li, Q., Zhang, J., Liu, D., Liu, Y. & Zhou, Y. Force-induced decline of FOXM1 in human periodontal ligament cells contributes to osteoclast differentiation. Angle Orthod. 89, 804\u2013811 (2019).","journal-title":"Angle Orthod."},{"key":"538_CR13","doi-asserted-by":"crossref","unstructured":"Wang, H. et al. Mechanisms of sphingosine-1-phosphate (S1P) signaling on excessive stress-induced root resorption during orthodontic molar intrusion. Clin. Oral Investig. 26, 1003\u20131016 (2021).","DOI":"10.1007\/s00784-021-04084-3"},{"key":"538_CR14","doi-asserted-by":"publisher","first-page":"21140","DOI":"10.3390\/ijms141021140","volume":"14","author":"X Bao","year":"2013","unstructured":"Bao, X., Liu, Y., Han, G., Zuo, Z. & Hu, M. The effect on proliferation and differentiation of cementoblast by using sclerostin as inhibitor. Int. J. Mol. Sci. 14, 21140\u201321152 (2013).","journal-title":"Int. J. Mol. Sci."},{"key":"538_CR15","doi-asserted-by":"crossref","unstructured":"De Rossi, A. et al. Cementocytes express receptor activator of the nuclear factor kappa-B ligand in response to endodontic infection in mice. J Endod. 42, 1251\u20131257 (2016).","DOI":"10.1016\/j.joen.2016.05.009"},{"key":"538_CR16","doi-asserted-by":"crossref","unstructured":"Wei, T., Shan, Z., Wen, X., Zhao, N. & Shen, G. Dynamic alternations of RANKL\/OPG ratio expressed by cementocytes in response to orthodontic\u2011induced external apical root resorption in a rat model. Mol. Med. Rep. 26, 228 (2022).","DOI":"10.3892\/mmr.2022.12744"},{"key":"538_CR17","doi-asserted-by":"publisher","DOI":"10.1038\/s41467-024-49159-3","volume":"15","author":"P Ding","year":"2024","unstructured":"Ding, P. et al. Mitochondria from osteolineage cells regulate myeloid cell-mediated bone resorption. Nat. Commun. 15, 5094 (2024).","journal-title":"Nat. Commun."},{"key":"538_CR18","doi-asserted-by":"publisher","DOI":"10.1002\/advs.202204871","volume":"10","author":"W Cai","year":"2023","unstructured":"Cai, W. et al. Mitochondrial transfer regulates cell fate through metabolic remodeling in osteoporosis. Adv. Sci. 10, e2204871 (2023).","journal-title":"Adv. Sci."},{"key":"538_CR19","doi-asserted-by":"publisher","DOI":"10.1038\/s41419-022-05343-1","volume":"13","author":"J Sarkar","year":"2022","unstructured":"Sarkar, J. et al. Epigallocatechin-3-gallate inhibits osteoclastic differentiation by modulating mitophagy and mitochondrial functions. Cell Death Dis. 13, 908 (2022).","journal-title":"Cell Death Dis."},{"key":"538_CR20","doi-asserted-by":"publisher","first-page":"335","DOI":"10.1002\/jbmr.4756","volume":"38","author":"L Zhu","year":"2023","unstructured":"Zhu, L. et al. STAT3\/mitophagy axis coordinates macrophage NLRP3 inflammasome activation and inflammatory bone loss. J. Bone Miner. Res. 38, 335\u2013353 (2023).","journal-title":"J. Bone Miner. Res."},{"key":"538_CR21","doi-asserted-by":"publisher","first-page":"1472","DOI":"10.1038\/s41418-023-01146-9","volume":"30","author":"L Sanchez","year":"2023","unstructured":"Sanchez, L. et al. SIM2s directed Parkin-mediated mitophagy promotes mammary epithelial cell differentiation. Cell Death Differ. 30, 1472\u20131487 (2023).","journal-title":"Cell Death Differ."},{"key":"538_CR22","doi-asserted-by":"publisher","DOI":"10.1016\/j.redox.2023.103023","volume":"69","author":"JS Jang","year":"2024","unstructured":"Jang, J. S. et al. PINK1 restrains periodontitis-induced bone loss by preventing osteoclast mitophagy impairment. Redox Biol. 69, 103023 (2024).","journal-title":"Redox Biol."},{"key":"538_CR23","doi-asserted-by":"publisher","DOI":"10.1038\/ncomms9472","volume":"6","author":"DG Phinney","year":"2015","unstructured":"Phinney, D. G. et al. Mesenchymal stem cells use extracellular vesicles to outsource mitophagy and shuttle microRNAs. Nat. Commun. 6, 8472 (2015).","journal-title":"Nat. Commun."},{"key":"538_CR24","doi-asserted-by":"publisher","first-page":"1620","DOI":"10.1016\/j.cmet.2022.10.008","volume":"34","author":"M Picard","year":"2022","unstructured":"Picard, M. & Shirihai, O. S. Mitochondrial signal transduction. Cell Metab. 34, 1620\u20131653 (2022).","journal-title":"Cell Metab."},{"key":"538_CR25","doi-asserted-by":"publisher","DOI":"10.1126\/sciadv.aaw7215","volume":"5","author":"J Gao","year":"2019","unstructured":"Gao, J. et al. Endoplasmic reticulum mediates mitochondrial transfer within the osteocyte dendritic network. Sci. Adv. 5, eaaw7215 (2019).","journal-title":"Sci. Adv."},{"key":"538_CR26","doi-asserted-by":"publisher","DOI":"10.1126\/sciadv.adi4298","volume":"10","author":"H Zhou","year":"2024","unstructured":"Zhou, H. et al. Osteocyte mitochondria inhibit tumor development via STING-dependent antitumor immunity. Sci. Adv. 10, eadi4298 (2024).","journal-title":"Sci. Adv."},{"key":"538_CR27","doi-asserted-by":"publisher","DOI":"10.1038\/s41467-024-46095-0","volume":"15","author":"P Liao","year":"2024","unstructured":"Liao, P. et al. Osteocyte mitochondria regulate angiogenesis of transcortical vessels. Nat. Commun. 15, 2529 (2024).","journal-title":"Nat. Commun."},{"key":"538_CR28","doi-asserted-by":"publisher","first-page":"430","DOI":"10.1002\/jbmr.2690","volume":"31","author":"N Zhao","year":"2016","unstructured":"Zhao, N. et al. Isolation and functional analysis of an immortalized murine cementocyte cell line, IDG-CM6. J. Bone Miner. Res. 31, 430\u2013442 (2016).","journal-title":"J. Bone Miner. Res."},{"key":"538_CR29","doi-asserted-by":"publisher","first-page":"1290","DOI":"10.1126\/science.1188635","volume":"328","author":"P P","year":"2010","unstructured":"P, P. et al. SphK1 regulates proinflammatory responses associated with endotoxin and polymicrobial sepsis. Science 328, 1290\u20131294 (2010).","journal-title":"Science"},{"key":"538_CR30","doi-asserted-by":"publisher","first-page":"2264","DOI":"10.1002\/jbmr.3844","volume":"34","author":"A El Jamal","year":"2019","unstructured":"El Jamal, A. et al. Cytokine-induced and stretch-induced sphingosine 1-phosphate production by enthesis cells could favor abnormal ossification in spondyloarthritis. J. Bone Miner. Res. 34, 2264\u20132276 (2019).","journal-title":"J. Bone Miner. Res."},{"key":"538_CR31","doi-asserted-by":"publisher","first-page":"1090","DOI":"10.1002\/jbmr.3396","volume":"33","author":"L Xiao","year":"2018","unstructured":"Xiao, L. et al. SPHK1-S1PR1-RANKL axis regulates the interactions between macrophages and BMSCs in inflammatory bone loss. J. Bone Miner. Res. 33, 1090\u20131104 (2018).","journal-title":"J. Bone Miner. Res."},{"key":"538_CR32","doi-asserted-by":"publisher","DOI":"10.1096\/fj.202403234RR","volume":"39","author":"H Wang","year":"2025","unstructured":"Wang, H. et al. Compression force regulates cementoblast mineralization via S1PR1\/mitophagy axis. FASEB J. 39, e70446 (2025).","journal-title":"FASEB J."},{"key":"538_CR33","doi-asserted-by":"crossref","unstructured":"Li, N. et al. PPP2\/PP2A-mediated dephosphorylation of LC3B links PINK1-PRKN\/Parkin-mediated mitophagy to SCA12 pathogenesis. Autophagy. 21, 1\u201313 (2025).","DOI":"10.1080\/15548627.2025.2572528"},{"key":"538_CR34","doi-asserted-by":"publisher","DOI":"10.1038\/s41419-018-0755-y","volume":"9","author":"SH Su","year":"2018","unstructured":"Su, S. H., Wu, Y. F., Wang, D. P. & Hai, J. Inhibition of excessive autophagy and mitophagy mediates neuroprotective effects of URB597 against chronic cerebral hypoperfusion. Cell Death Dis. 9, 733 (2018).","journal-title":"Cell Death Dis."},{"key":"538_CR35","doi-asserted-by":"publisher","DOI":"10.1016\/j.celrep.2023.112883","volume":"42","author":"DC Maddison","year":"2023","unstructured":"Maddison, D. C. et al. COPI-regulated mitochondria-ER contact site formation maintains axonal integrity. Cell Rep. 42, 112883 (2023).","journal-title":"Cell Rep."},{"key":"538_CR36","doi-asserted-by":"publisher","first-page":"595","DOI":"10.1016\/j.bbrc.2020.01.003","volume":"523","author":"H Wang","year":"2020","unstructured":"Wang, H. et al. The role of sphingosine-1-phosphate signaling pathway in cementocyte mechanotransduction. Biochem. Biophys. Res. Commun. 523, 595\u2013601 (2020).","journal-title":"Biochem. Biophys. Res. Commun."},{"key":"538_CR37","doi-asserted-by":"publisher","DOI":"10.1111\/cpr.13604","volume":"57","author":"H Wang","year":"2024","unstructured":"Wang, H. et al. Long non-coding RNA LncTUG1 regulates favourable compression force-induced cementocytes mineralization via PU.1\/TLR4\/SphK1 signalling. Cell Prolif. 57, e13604 (2024).","journal-title":"Cell Prolif."},{"key":"538_CR38","doi-asserted-by":"publisher","first-page":"71","DOI":"10.1016\/j.freeradbiomed.2022.07.001","volume":"189","author":"S Arora","year":"2022","unstructured":"Arora, S., Singh, P., Tabassum, G., Dohare, R. & Syed, M. A. miR-495-3p regulates sphingolipid metabolic reprogramming to induce Sphk1\/ceramide mediated mitophagy and apoptosis in NSCLC. Free Radic. Biol. Med. 189, 71\u201384 (2022).","journal-title":"Free Radic. Biol. Med."},{"key":"538_CR39","doi-asserted-by":"publisher","first-page":"1924","DOI":"10.1038\/s41418-019-0471-x","volume":"27","author":"L Li","year":"2020","unstructured":"Li, L. et al. SPHK1 deficiency protects mice from acetaminophen-induced ER stress and mitochondrial permeability transition. Cell Death Differ. 27, 1924\u20131937 (2020).","journal-title":"Cell Death Differ."},{"key":"538_CR40","doi-asserted-by":"publisher","first-page":"660","DOI":"10.1007\/s11914-023-00825-3","volume":"21","author":"MG Ledesma-Colunga","year":"2023","unstructured":"Ledesma-Colunga, M. G., Passin, V., Lademann, F., Hofbauer, L. C. & Rauner, M. Novel insights into osteoclast energy metabolism. Curr. Osteoporos. Rep. 21, 660\u2013669 (2023).","journal-title":"Curr. Osteoporos. Rep."},{"key":"538_CR41","doi-asserted-by":"publisher","first-page":"948","DOI":"10.1002\/art.42074","volume":"74","author":"K Doi","year":"2022","unstructured":"Doi, K. et al. Role of lysine-specific demethylase 1 in metabolically integrating osteoclast differentiation and inflammatory bone resorption through hypoxia-inducible factor 1\u03b1 and E2F1. Arthritis Rheumatol. 74, 948\u2013960 (2022).","journal-title":"Arthritis Rheumatol."},{"key":"538_CR42","doi-asserted-by":"publisher","DOI":"10.1016\/j.phrs.2023.107056","volume":"200","author":"R Liu","year":"2024","unstructured":"Liu, R. et al. Low-dose Olaparib improves septic cardiac function by reducing ferroptosis via accelerated mitophagy flux. Pharmacol. Res. 200, 107056 (2024).","journal-title":"Pharmacol. Res."},{"key":"538_CR43","first-page":"3174","volume":"20","author":"T Wei","year":"2020","unstructured":"Wei, T., Xie, Y., Wen, X., Zhao, N. & Shen, G. Establishment of in vitro three-dimensional cementocyte differentiation scaffolds to study orthodontic root resorption. Exp. Ther. Med. 20, 3174\u20133184 (2020).","journal-title":"Exp. Ther. Med."},{"key":"538_CR44","doi-asserted-by":"publisher","first-page":"2675","DOI":"10.7150\/ijbs.46627","volume":"16","author":"S Wang","year":"2020","unstructured":"Wang, S. et al. The role of autophagy and mitophagy in bone metabolic disorders. Int. J. Biol. Sci. 16, 2675\u20132691 (2020).","journal-title":"Int. J. Biol. Sci."},{"key":"538_CR45","doi-asserted-by":"publisher","first-page":"859","DOI":"10.1111\/cns.13140","volume":"25","author":"Y Wang","year":"2019","unstructured":"Wang, Y., Liu, N. & Lu, B. Mechanisms and roles of mitophagy in neurodegenerative diseases. CNS Neurosci. Ther. 25, 859\u2013875 (2019).","journal-title":"CNS Neurosci. Ther."},{"key":"538_CR46","doi-asserted-by":"publisher","first-page":"31","DOI":"10.1038\/cdd.2012.81","volume":"20","author":"G Ashrafi","year":"2013","unstructured":"Ashrafi, G. & Schwarz, T. L. The pathways of mitophagy for quality control and clearance of mitochondria. Cell Death Differ. 20, 31\u201342 (2013).","journal-title":"Cell Death Differ."},{"key":"538_CR47","doi-asserted-by":"publisher","first-page":"80","DOI":"10.1016\/j.jmb.2019.07.016","volume":"432","author":"T Johansen","year":"2020","unstructured":"Johansen, T. & Lamark, T. Selective autophagy: ATG8 family proteins, LIR motifs and cargo receptors. J. Mol. Biol. 432, 80\u2013103 (2020).","journal-title":"J. Mol. Biol."},{"key":"538_CR48","doi-asserted-by":"publisher","DOI":"10.1096\/fj.202401280R","volume":"38","author":"T Yan","year":"2024","unstructured":"Yan, T., Li, H., Yan, J., Ma, S. & Tan, J. Age-related mitophagy regulates orthodontic tooth movement by affecting PDLSCs mitochondrial function and RANKL\/OPG. FASEB J. 38, e23865 (2024).","journal-title":"FASEB J."},{"key":"538_CR49","doi-asserted-by":"publisher","DOI":"10.3389\/fphar.2021.703847","volume":"12","author":"Y He","year":"2021","unstructured":"He, Y., Yocum, L., Alexander, P. G., Jurczak, M. J. & Lin, H. Urolithin A protects chondrocytes from mechanical overloading-induced injuries. Front. Pharmacol. 12, 703847 (2021).","journal-title":"Front. Pharmacol."},{"key":"538_CR50","doi-asserted-by":"publisher","first-page":"57","DOI":"10.1186\/s11658-023-00453-w","volume":"28","author":"Z Zhang","year":"2023","unstructured":"Zhang, Z. et al. Mechanical force induces mitophagy-mediated anaerobic oxidation in periodontal ligament stem cells. Cell. Mol. Biol. Lett. 28, 57 (2023).","journal-title":"Cell. Mol. Biol. Lett."},{"key":"538_CR51","doi-asserted-by":"crossref","unstructured":"Coon, B. G. et al. A mitochondrial contribution to anti-inflammatory shear stress signaling in vascular endothelial cells. J. Cell Biol. 221, e202109144 (2022).","DOI":"10.1083\/jcb.202109144"},{"key":"538_CR52","doi-asserted-by":"publisher","first-page":"210","DOI":"10.1359\/jbmr.2002.17.2.210","volume":"17","author":"H Kanzaki","year":"2002","unstructured":"Kanzaki, H., Chiba, M., Shimizu, Y. & Mitani, H. Periodontal ligament cells under mechanical stress induce osteoclastogenesis by receptor activator of nuclear factor kappaB ligand up-regulation via prostaglandin E2 synthesis. J. Bone Miner. Res. 17, 210\u2013220 (2002).","journal-title":"J. Bone Miner. Res."},{"key":"538_CR53","doi-asserted-by":"publisher","first-page":"1235","DOI":"10.1038\/s41423-022-00921-x","volume":"19","author":"L Tian","year":"2022","unstructured":"Tian, L. et al. S1P\/S1PR1 signaling differentially regulates the allogeneic response of CD4 and CD8 T cells by modulating mitochondrial fission. Cell. Mol. Immunol. 19, 1235\u20131250 (2022).","journal-title":"Cell. Mol. Immunol."},{"key":"538_CR54","doi-asserted-by":"publisher","first-page":"47","DOI":"10.1016\/j.exphem.2019.01.004","volume":"72","author":"C Yang","year":"2019","unstructured":"Yang, C., Hashimoto, M., Lin, Q. X. X., Tan, D. Q. & Suda, T. Sphingosine-1-phosphate signaling modulates terminal erythroid differentiation through the regulation of mitophagy. Exp. Hematol. 72, 47\u201359.e41 (2019).","journal-title":"Exp. Hematol."},{"key":"538_CR55","doi-asserted-by":"crossref","unstructured":"Kong, J. et al. Mitochondrial extracellular vesicles: a novel approach to mitochondrial quality control. Biomolecules 15, 1145 (2025).","DOI":"10.3390\/biom15081145"},{"key":"538_CR56","first-page":"773","volume":"53","author":"M Zhang","year":"2025","unstructured":"Zhang, M. et al. Autophagic-active nanosystem for senile bone regeneration by in-situ mitochondrial biogenesis and intercellular transfer. Bioact. Mater. 53, 773\u2013788 (2025).","journal-title":"Bioact. Mater."},{"key":"538_CR57","doi-asserted-by":"publisher","DOI":"10.1016\/j.jhazmat.2025.140486","volume":"500","author":"W Yu","year":"2025","unstructured":"Yu, W., Zhang, B., Yang, M., Li, Y. & Cao, Z. Deoxynivalenol induces liver injury by inhibiting the p62-Keap1-Nrf2 signaling pathway via overactivation of PINK1\/Parkin-mediated mitophagy. J. Hazard. Mater. 500, 140486 (2025).","journal-title":"J. Hazard. Mater."}],"container-title":["Bone Research"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.nature.com\/articles\/s41413-026-00538-0.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/www.nature.com\/articles\/s41413-026-00538-0","content-type":"text\/html","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/www.nature.com\/articles\/s41413-026-00538-0.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,5,14]],"date-time":"2026-05-14T09:06:04Z","timestamp":1778749564000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.nature.com\/articles\/s41413-026-00538-0"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,5,14]]},"references-count":57,"journal-issue":{"issue":"1","published-online":{"date-parts":[[2026,12]]}},"alternative-id":["538"],"URL":"https:\/\/doi.org\/10.1038\/s41413-026-00538-0","relation":{},"ISSN":["2095-6231"],"issn-type":[{"value":"2095-6231","type":"electronic"}],"subject":[],"published":{"date-parts":[[2026,5,14]]},"assertion":[{"value":"11 December 2025","order":1,"name":"received","label":"Received","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"14 February 2026","order":2,"name":"revised","label":"Revised","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"7 March 2026","order":3,"name":"accepted","label":"Accepted","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"14 May 2026","order":4,"name":"first_online","label":"First Online","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"The authors declare no competing interests.","order":1,"name":"Ethics","group":{"name":"EthicsHeading","label":"Competing interests"}}],"article-number":"52"}}