{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,8,10]],"date-time":"2026-08-10T15:10:54Z","timestamp":1786374654128,"version":"build-2736575974"},"reference-count":62,"publisher":"Springer Science and Business Media LLC","issue":"1","license":[{"start":{"date-parts":[[2026,8,10]],"date-time":"2026-08-10T00:00:00Z","timestamp":1786320000000},"content-version":"tdm","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0"},{"start":{"date-parts":[[2026,8,10]],"date-time":"2026-08-10T00:00:00Z","timestamp":1786320000000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0"}],"funder":[{"DOI":"10.13039\/100014440","name":"Ministerio de Ciencia, Innovaci\u00f3n y Universidades","doi-asserted-by":"publisher","award":["PID2023-151388OB-I00"],"award-info":[{"award-number":["PID2023-151388OB-I00"]}],"id":[{"id":"10.13039\/100014440","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/100011941","name":"Gobierno del Principado de Asturias","doi-asserted-by":"publisher","award":["IDE\/2024\/000709"],"award-info":[{"award-number":["IDE\/2024\/000709"]}],"id":[{"id":"10.13039\/100011941","id-type":"DOI","asserted-by":"publisher"}]}],"content-domain":{"domain":["link.springer.com"],"crossmark-restriction":false},"short-container-title":["Mol Biomed"],"abstract":"<jats:title>Abstract<\/jats:title>\n                  <jats:p>miR-210-3p is a well-established hypoxia-induced microRNA that is commonly upregulated in a wide range of solid tumors, traditionally linked to mitochondrial repression and hypoxia-inducible factor (HIF) signaling. However, its functional role in cancer remains complex and highly context dependent. Here we perform a comprehensive pan-cancer transcriptomic analysis together with functional assays, revealing that miR-210-3p not only mediates classical hypoxic responses but also amplifies mitotic gene expression through activation of FOXM1. Mechanistically, this effect is shown to be dependent on HIF1\u03b1 but not on HIF2\u03b1 and, importantly, it has not been recapitulated by hypoxia alone. Notably, activation of the mitotic program is observed in breast cancer cells but not in head and neck squamous carcinoma models, highlighting a strong degree of context dependency across tumor types. In breast cancer cells, miR-210-3p overexpression enhances FOXM1 phosphorylation, upregulates kinetochore regulators, and induces mitotic defects, correlating with poor prognosis in aggressive tumors. Together, these findings position miR-210-3p as a molecular integrator linking pseudohypoxia to mitotic dysregulation, contributing to tumor aggressiveness by sustaining HIF activity and promoting mitotic stress. This dual functionality reconciles its paradoxical effects on proliferation and highlights its potential as a therapeutic target in cancers characterized by pseudohypoxia and mitotic abnormalities.<\/jats:p>","DOI":"10.1186\/s43556-026-00530-4","type":"journal-article","created":{"date-parts":[[2026,8,10]],"date-time":"2026-08-10T01:25:17Z","timestamp":1786325117000},"update-policy":"https:\/\/doi.org\/10.1007\/springer_crossmark_policy","source":"Crossref","is-referenced-by-count":0,"title":["A dual role for miR-210-3p in hypoxia-driven signaling and mitotic regulation in cancer"],"prefix":"10.1186","volume":"7","author":[{"given":"Jaime","family":"San-Juan-Guardado","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Mar\u00eda","family":"Turienzo-Dur\u00e1n","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"\u00c1lvaro","family":"Su\u00e1rez-Priede","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Nerea","family":"G\u00f3mez-Su\u00e1rez","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Candelaria","family":"Aguilar-Garc\u00eda","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Tamara","family":"Cubiella","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-1112-1583","authenticated-orcid":false,"given":"Mar\u00eda-Dolores","family":"Chiara","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"297","published-online":{"date-parts":[[2026,8,10]]},"reference":[{"issue":"12","key":"530_CR1","doi-asserted-by":"publisher","first-page":"758","DOI":"10.1016\/j.trecan.2016.10.016","volume":"2","author":"L Schito","year":"2016","unstructured":"Schito L, Semenza GL. Hypoxia-inducible factors: master regulators of cancer progression. Trends Cancer. 2016;2(12):758\u201370. https:\/\/doi.org\/10.1016\/j.trecan.2016.10.016.","journal-title":"Trends Cancer"},{"issue":"14","key":"530_CR2","doi-asserted-by":"publisher","first-page":"47","DOI":"10.1146\/annurev-pathol-012513-104720","volume":"9","author":"GL Semenza","year":"2014","unstructured":"Semenza GL. Oxygen sensing, hypoxia-inducible factors, and disease pathophysiology. Annu Rev Pathol. 2014;9(14):47\u201371. https:\/\/doi.org\/10.1146\/annurev-pathol-012513-104720.","journal-title":"Annu Rev Pathol"},{"issue":"7627","key":"530_CR3","doi-asserted-by":"publisher","first-page":"112","DOI":"10.1038\/nature19796","volume":"539","author":"W Chen","year":"2016","unstructured":"Chen W, Hill H, Christie A, Kim MS, Holloman E, Pavia-Jimenez A, et al. Targeting renal cell carcinoma with a HIF-2 antagonist. Nature. 2016;539(7627):112\u20137. https:\/\/doi.org\/10.1038\/nature19796.","journal-title":"Nature"},{"issue":"13","key":"530_CR4","doi-asserted-by":"publisher","first-page":"5675","DOI":"10.1128\/MCB.25.13.5675-5686.2005","volume":"25","author":"RR Raval","year":"2005","unstructured":"Raval RR, Lau KW, Tran MGB, Sowter HM, Mandriota SJ, Li J-L, et al. Contrasting properties of hypoxia-inducible factor 1 (HIF-1) and HIF-2 in von Hippel-Lindau-associated renal cell carcinoma. Mol Cell Biol. 2005;25(13):5675\u201386. https:\/\/doi.org\/10.1128\/MCB.25.13.5675-5686.2005.","journal-title":"Mol Cell Biol"},{"issue":"11","key":"530_CR5","doi-asserted-by":"publisher","first-page":"4593","DOI":"10.1210\/jc.2006-0920","volume":"91","author":"PJ Pollard","year":"2006","unstructured":"Pollard PJ, El-Bahrawy M, Poulsom R, Elia G, Killick P, Kelly G, et al. Expression of HIF-1alpha, HIF-2alpha (EPAS1), and their target genes in paraganglioma and pheochromocytoma with VHL and SDH mutations. J Clin Endocrinol Metab. 2006;91(11):4593\u20138. https:\/\/doi.org\/10.1210\/jc.2006-0920.","journal-title":"J Clin Endocrinol Metab"},{"issue":"20","key":"530_CR6","doi-asserted-by":"publisher","first-page":"2012","DOI":"10.1056\/NEJMoa2504964","volume":"393","author":"C Jimenez","year":"2025","unstructured":"Jimenez C, Andreassen M, Durand A, Moog S, Hendifar A, Welin S, et al. Belzutifan for advanced pheochromocytoma or paraganglioma. N Engl J Med. 2025;393(20):2012\u201322. https:\/\/doi.org\/10.1056\/NEJMoa2504964.","journal-title":"N Engl J Med"},{"issue":"12","key":"530_CR7","doi-asserted-by":"publisher","first-page":"2986","DOI":"10.3390\/cancers14122986","volume":"14","author":"L Celada","year":"2022","unstructured":"Celada L, Cubiella T, San-Juan-Guardado J, San Jos\u00e9 Mart\u00ednez A, Vald\u00e9s N, Jim\u00e9nez-Fonseca P, et al. Differential HIF2\u03b1 protein expression in human carotid body and adrenal medulla under physiologic and tumorigenic conditions. Cancers (Basel). 2022;14(12):2986. https:\/\/doi.org\/10.3390\/cancers14122986.","journal-title":"Cancers (Basel)"},{"issue":"6","key":"530_CR8","doi-asserted-by":"publisher","first-page":"1072","DOI":"10.4161\/cc.9.6.11006","volume":"9","author":"SY Chan","year":"2010","unstructured":"Chan SY, Loscalzo J. MicroRNA-210: a unique and pleiotropic hypoxamir. Cell Cycle. 2010;9(6):1072\u201383. https:\/\/doi.org\/10.4161\/cc.9.6.11006.","journal-title":"Cell Cycle"},{"issue":"10-12","key":"530_CR9","doi-asserted-by":"publisher","first-page":"685","DOI":"10.1089\/ars.2021.0200","volume":"36","author":"G Zaccagnini","year":"2022","unstructured":"Zaccagnini G, Greco S, Voellenkle C, Gaetano C, Martelli F. MiR-210 hypoxamiR in angiogenesis and diabetes. Antioxid Redox Signal. 2022;36(10\u201312):685\u2013706. https:\/\/doi.org\/10.1089\/ars.2021.0200.","journal-title":"Antioxid Redox Signal"},{"issue":"30","key":"530_CR10","doi-asserted-by":"publisher","first-page":"4362","DOI":"10.1038\/onc.2010.193","volume":"29","author":"Z Chen","year":"2010","unstructured":"Chen Z, Li Y, Zhang H, Huang P, Luthra R. Hypoxia-regulated microRNA-210 modulates mitochondrial function and decreases ISCU and COX10 expression. Oncogene. 2010;29(30):4362\u20138. https:\/\/doi.org\/10.1038\/onc.2010.193.","journal-title":"Oncogene"},{"issue":"8","key":"530_CR11","doi-asserted-by":"publisher","first-page":"13730","DOI":"10.18632\/oncotarget.14629","volume":"8","author":"I S\u00e1enz-de-Santa-Mar\u00eda","year":"2017","unstructured":"S\u00e1enz-de-Santa-Mar\u00eda I, Bernardo-Casti\u00f1eira C, Secades P, Bernaldo-de-Quir\u00f3s S, Rodrigo JP, Astudillo A, et al. Clinically relevant HIF-1\u03b1-dependent metabolic reprogramming in oropharyngeal squamous cell carcinomas includes coordinated activation of CAIX and the miR-210\/ISCU signaling axis, but not MCT1 and MCT4 upregulation. Oncotarget. 2017;8(8):13730\u201346. https:\/\/doi.org\/10.18632\/oncotarget.14629.","journal-title":"Oncotarget"},{"issue":"4","key":"530_CR12","doi-asserted-by":"publisher","first-page":"e10345","DOI":"10.1371\/journal.pone.0010345","volume":"5","author":"E Favaro","year":"2010","unstructured":"Favaro E, Ramachandran A, McCormick R, Gee H, Blancher C, Crosby M, et al. MicroRNA-210 regulates mitochondrial free radical response to hypoxia and krebs cycle in cancer cells by targeting iron sulfur cluster protein ISCU. PLoS ONE. 2010;5(4):e10345. https:\/\/doi.org\/10.1371\/journal.pone.0010345.","journal-title":"PLoS ONE"},{"issue":"3","key":"530_CR13","doi-asserted-by":"publisher","first-page":"1221","DOI":"10.1158\/0008-5472.CAN-08-2516","volume":"69","author":"ME Crosby","year":"2009","unstructured":"Crosby ME, Kulshreshtha R, Ivan M, Glazer PM. MicroRNA regulation of DNA repair gene expression in hypoxic stress. Cancer Res. 2009;69(3):1221\u20139. https:\/\/doi.org\/10.1158\/0008-5472.CAN-08-2516.","journal-title":"Cancer Res"},{"issue":"3","key":"530_CR14","doi-asserted-by":"publisher","first-page":"182","DOI":"10.1159\/000331054","volume":"35","author":"F Liu","year":"2012","unstructured":"Liu F, Lou Y-L, Wu J, Ruan Q-F, Xie A, Guo F, et al. Upregulation of microRNA-210 regulates renal angiogenesis mediated by activation of VEGF signaling pathway under ischemia\/perfusion injury in vivo and in vitro. Kidney Blood Press Res. 2012;35(3):182\u201391. https:\/\/doi.org\/10.1159\/000331054.","journal-title":"Kidney Blood Press Res"},{"key":"530_CR15","doi-asserted-by":"publisher","first-page":"7752277","DOI":"10.1155\/2022\/7752277","volume":"2022","author":"G Wu","year":"2022","unstructured":"Wu G, Ding X, Quan G, Xiong J, Li Q, Li Z, et al. Hypoxia-Induced miR-210 Promotes Endothelial Cell Permeability and Angiogenesis via Exosomes in Pancreatic Ductal Adenocarcinoma. Biochem Res Int. 2022;2022:7752277. https:\/\/doi.org\/10.1155\/2022\/7752277.","journal-title":"Biochem Res Int"},{"issue":"5","key":"530_CR16","doi-asserted-by":"publisher","first-page":"645","DOI":"10.1007\/s10517-022-05605-0","volume":"173","author":"D Suwei","year":"2022","unstructured":"Suwei D, Zhen L, Zhimin L, Mei L, Jianping K, Zhuohui P, et al. Hypoxia Modulates Melanoma Cells Proliferation and Apoptosis via miRNA-210\/ISCU\/ROS Signaling. Bull Exp Biol Med. 2022;173(5):645\u201350. https:\/\/doi.org\/10.1007\/s10517-022-05605-0.","journal-title":"Bull Exp Biol Med"},{"key":"530_CR17","doi-asserted-by":"publisher","first-page":"3687","DOI":"10.2147\/OTT.S288788","volume":"14","author":"Q Chen","year":"2021","unstructured":"Chen Q, Zhang H, Zhang J, Shen L, Yang J, Wang Y, et al. miR-210-3p Promotes Lung Cancer Development and Progression by Modulating USF1 and PCGF3. Onco Targets Ther. 2021;14:3687\u2013700. https:\/\/doi.org\/10.2147\/OTT.S288788.","journal-title":"Onco Targets Ther"},{"issue":"8","key":"530_CR18","doi-asserted-by":"publisher","first-page":"14050","DOI":"10.1002\/jcp.28093","volume":"234","author":"S Xie","year":"2019","unstructured":"Xie S, Liu G, Huang J, Hu H-B, Jiang W. MiR-210 promotes lung adenocarcinoma proliferation, migration, and invasion by targeting lysyl oxidase-like 4. J Cell Physiol. 2019;234(8):14050\u20137. https:\/\/doi.org\/10.1002\/jcp.28093.","journal-title":"J Cell Physiol"},{"key":"530_CR19","doi-asserted-by":"publisher","unstructured":"Tang T, Yang Z, Zhu Q, Wu Y, Sun K, Alahdal M, et al. Up-regulation of miR-210 induced by a hypoxic microenvironment promotes breast cancer stem cells metastasis, proliferation, and self-renewal by targeting E-cadherin. FASEB J. 2018;fj201801013R. https:\/\/doi.org\/10.1096\/fj.201801013R.","DOI":"10.1096\/fj.201801013R"},{"issue":"11","key":"530_CR20","doi-asserted-by":"publisher","first-page":"992","DOI":"10.1038\/s41419-020-03150-0","volume":"11","author":"P Wang","year":"2020","unstructured":"Wang P, Yan Q, Liao B, Zhao L, Xiong S, Wang J, et al. The HIF1\u03b1\/HIF2\u03b1-miR210-3p network regulates glioblastoma cell proliferation, dedifferentiation and chemoresistance through EGF under hypoxic conditions. Cell Death Dis. 2020;11(11):992. https:\/\/doi.org\/10.1038\/s41419-020-03150-0.","journal-title":"Cell Death Dis"},{"issue":"8","key":"530_CR21","doi-asserted-by":"publisher","first-page":"1813","DOI":"10.1007\/s11064-020-03043-w","volume":"45","author":"H He","year":"2020","unstructured":"He H, Liu J, Li W, Yao X, Ren Q, Shen B, et al. MiR-210-3p Inhibits Proliferation and Migration of C6 Cells by Targeting Iscu. Neurochem Res. 2020;45(8):1813\u201324. https:\/\/doi.org\/10.1007\/s11064-020-03043-w.","journal-title":"Neurochem Res"},{"issue":"8","key":"530_CR22","doi-asserted-by":"publisher","first-page":"1738","DOI":"10.3390\/cimb47040286","volume":"7","author":"X Yang","year":"2017","unstructured":"Yang X, Shi L, Yi C, Yang Y, Chang L, Song D. MiR-210-3p inhibits the tumor growth and metastasis of bladder cancer via targeting fibroblast growth factor receptor-like 1. Am J Cancer Res. 2017;7(8):1738\u201353. https:\/\/doi.org\/10.3390\/cimb47040286.","journal-title":"Am J Cancer Res"},{"issue":"46","key":"530_CR23","doi-asserted-by":"publisher","first-page":"13030","DOI":"10.3748\/wjg.v21.i46.13030","volume":"21","author":"W Tan","year":"2015","unstructured":"Tan W, Lim S-G, Tan TMC. Up-regulation of microRNA-210 inhibits proliferation of hepatocellular carcinoma cells by targeting YES1. World J Gastroenterol. 2015;21(46):13030\u201341. https:\/\/doi.org\/10.3748\/wjg.v21.i46.13030.","journal-title":"World J Gastroenterol"},{"key":"530_CR24","doi-asserted-by":"publisher","first-page":"4497","DOI":"10.1038\/ncomms5497","volume":"5","author":"K Kiga","year":"2014","unstructured":"Kiga K, Mimuro H, Suzuki M, Shinozaki-Ushiku A, Kobayashi T, Sanada T, et al. Epigenetic silencing of miR-210 increases the proliferation of gastric epithelium during chronic Helicobacter pylori infection. Nat Commun. 2014;5:4497. https:\/\/doi.org\/10.1038\/ncomms5497.","journal-title":"Nat Commun"},{"issue":"1","key":"530_CR25","doi-asserted-by":"publisher","first-page":"50","DOI":"10.1186\/1756-9966-33-50","volume":"33","author":"Q Qin","year":"2014","unstructured":"Qin Q, Furong W, Baosheng L. Multiple functions of hypoxia-regulated miR-210 in cancer. J Exp Clin Cancer Res. 2014;33(1):50. https:\/\/doi.org\/10.1186\/1756-9966-33-50.","journal-title":"J Exp Clin Cancer Res"},{"key":"530_CR26","doi-asserted-by":"publisher","first-page":"154244","DOI":"10.1016\/j.prp.2022.154244","volume":"241","author":"S Khalilian","year":"2023","unstructured":"Khalilian S, Bijanvand A, Abedinlou H, Ghafouri-Fard S. A review on the role of miR-210 in human disorders. Pathol Res Pract. 2023;241:154244. https:\/\/doi.org\/10.1016\/j.prp.2022.154244.","journal-title":"Pathol Res Pract"},{"issue":"6","key":"530_CR27","doi-asserted-by":"publisher","first-page":"1129","DOI":"10.32604\/or.2024.047524","volume":"32","author":"J Yi","year":"2024","unstructured":"Yi J, Li B, Yin X, Liu L, Song C, Zhao Y, et al. CircMYBL2 facilitates hepatocellular carcinoma progression by regulating E2F1 expression. Oncol Res. 2024;32(6):1129\u201339. https:\/\/doi.org\/10.32604\/or.2024.047524.","journal-title":"Oncol Res"},{"issue":"25","key":"530_CR28","doi-asserted-by":"publisher","first-page":"01014","DOI":"10.1016\/j.jare.2025.12.023","volume":"S2090\u20131232","author":"X Li","year":"2025","unstructured":"Li X, Han M, Zhang L, Xie X, Li C, Zhang H, et al. Gymconopin C exhibits anti-non-small cell lung cancer effect by regulating miR-6777-5p\/ADRB2 pathway to promote mitophagy. J Adv Res. 2025;S2090\u20131232(25):01014\u20138. https:\/\/doi.org\/10.1016\/j.jare.2025.12.023.","journal-title":"J Adv Res"},{"issue":"9","key":"530_CR29","doi-asserted-by":"publisher","first-page":"731","DOI":"10.1038\/s41419-020-02952-6","volume":"11","author":"Y Du","year":"2020","unstructured":"Du Y, Wei N, Ma R, Jiang S, Song D. A miR-210-3p regulon that controls the Warburg effect by modulating HIF-1\u03b1 and p53 activity in triple-negative breast cancer. Cell Death Dis. 2020;11(9):731. https:\/\/doi.org\/10.1038\/s41419-020-02952-6.","journal-title":"Cell Death Dis"},{"issue":"35","key":"530_CR30","doi-asserted-by":"publisher","first-page":"13021","DOI":"10.1073\/pnas.0803304105","volume":"105","author":"JA Foekens","year":"2008","unstructured":"Foekens JA, Sieuwerts AM, Smid M, Look MP, de Weerd V, Boersma AWM, et al. Four miRNAs associated with aggressiveness of lymph node-negative, estrogen receptor-positive human breast cancer. Proc Natl Acad Sci U S A. 2008;105(35):13021\u20136. https:\/\/doi.org\/10.1073\/pnas.0803304105.","journal-title":"Proc Natl Acad Sci U S A"},{"issue":"1","key":"530_CR31","doi-asserted-by":"publisher","first-page":"38","DOI":"10.1186\/s13578-022-00772-z","volume":"12","author":"W Shi","year":"2022","unstructured":"Shi W, Tang Y, Lu J, Zhuang Y, Wang J. MIR210HG promotes breast cancer progression by IGF2BP1 mediated m6A modification. Cell Biosci. 2022;12(1):38. https:\/\/doi.org\/10.1186\/s13578-022-00772-z.","journal-title":"Cell Biosci"},{"issue":"2","key":"530_CR32","doi-asserted-by":"publisher","first-page":"255","DOI":"10.4161\/cbt.7.2.5297","volume":"7","author":"A Giannakakis","year":"2008","unstructured":"Giannakakis A, Sandaltzopoulos R, Greshock J, Liang S, Huang J, Hasegawa K, et al. MiR-210 links hypoxia with cell cycle regulation and is deleted in human epithelial ovarian cancer. Cancer Biol Ther. 2008;7(2):255\u201364. https:\/\/doi.org\/10.4161\/cbt.7.2.5297.","journal-title":"Cancer Biol Ther"},{"issue":"2","key":"530_CR33","doi-asserted-by":"publisher","first-page":"280","DOI":"10.1002\/path.2860","volume":"224","author":"C Nakada","year":"2011","unstructured":"Nakada C, Tsukamoto Y, Matsuura K, Nguyen TL, Hijiya N, Uchida T, et al. Overexpression of miR-210, a downstream target of HIF1\u03b1, causes centrosome amplification in renal carcinoma cells. J Pathol. 2011;224(2):280\u20138. https:\/\/doi.org\/10.1002\/path.2860.","journal-title":"J Pathol"},{"issue":"1","key":"530_CR34","doi-asserted-by":"publisher","first-page":"498","DOI":"10.1093\/nar\/gks995","volume":"41","author":"J He","year":"2013","unstructured":"He J, Wu J, Xu N, Xie W, Li M, Li J, et al. MiR-210 disturbs mitotic progression through regulating a group of mitosis-related genes. Nucleic Acids Res. 2013;41(1):498\u2013508. https:\/\/doi.org\/10.1093\/nar\/gks995.","journal-title":"Nucleic Acids Res"},{"key":"530_CR35","doi-asserted-by":"publisher","unstructured":"Bavelloni A, Ramazzotti G, Poli A, Piazzi M, Focaccia E, Blalock W, et al. MiRNA-210: A Current Overview. Anticancer Res. 2017;37(12):6511\u201321. https:\/\/doi.org\/10.21873\/anticanres.12107.","DOI":"10.21873\/anticanres.12107"},{"issue":"16","key":"530_CR36","doi-asserted-by":"publisher","first-page":"4667","DOI":"10.1158\/1078-0432.CCR-06-2510","volume":"13","author":"G Bratslavsky","year":"2007","unstructured":"Bratslavsky G, Sudarshan S, Neckers L, Linehan WM. Pseudohypoxic pathways in renal cell carcinoma. Clin Cancer Res. 2007;13(16):4667\u201371. https:\/\/doi.org\/10.1158\/1078-0432.CCR-06-2510.","journal-title":"Clin Cancer Res"},{"issue":"6","key":"530_CR37","doi-asserted-by":"publisher","first-page":"957","DOI":"10.1016\/j.beem.2010.10.004","volume":"24","author":"J Favier","year":"2010","unstructured":"Favier J, Gimenez-Roqueplo A-P. Pheochromocytomas: the (pseudo)-hypoxia hypothesis. Best Pract Res Clin Endocrinol Metab. 2010;24(6):957\u201368. https:\/\/doi.org\/10.1016\/j.beem.2010.10.004.","journal-title":"Best Pract Res Clin Endocrinol Metab"},{"issue":"4","key":"530_CR38","doi-asserted-by":"publisher","first-page":"273","DOI":"10.1016\/j.cmet.2009.08.015","volume":"10","author":"SY Chan","year":"2009","unstructured":"Chan SY, Zhang Y-Y, Hemann C, Mahoney CE, Zweier JL, Loscalzo J. MicroRNA-210 controls mitochondrial metabolism during hypoxia by repressing the iron-sulfur cluster assembly proteins ISCU1\/2. Cell Metab. 2009;10(4):273\u201384. https:\/\/doi.org\/10.1016\/j.cmet.2009.08.015.","journal-title":"Cell Metab"},{"issue":"7","key":"530_CR39","doi-asserted-by":"publisher","first-page":"111652","DOI":"10.1016\/j.celrep.2022.111652","volume":"41","author":"O Lombardi","year":"2022","unstructured":"Lombardi O, Li R, Halim S, Choudhry H, Ratcliffe PJ, Mole DR. Pan-cancer analysis of tissue and single-cell HIF-pathway activation using a conserved gene signature. Cell Rep. 2022;41(7):111652. https:\/\/doi.org\/10.1016\/j.celrep.2022.111652.","journal-title":"Cell Rep"},{"issue":"13","key":"530_CR40","doi-asserted-by":"publisher","first-page":"1983","DOI":"10.1091\/mbc.E14-03-0837","volume":"25","author":"P Thiru","year":"2014","unstructured":"Thiru P, Kern DM, McKinley KL, Monda JK, Rago F, Su K-C, et al. Kinetochore genes are coordinately up-regulated in human tumors as part of a FoxM1-related cell division program. Mol Biol Cell. 2014;25(13):1983\u201394. https:\/\/doi.org\/10.1091\/mbc.E14-03-0837.","journal-title":"Mol Biol Cell"},{"issue":"1","key":"530_CR41","doi-asserted-by":"publisher","first-page":"257","DOI":"10.3892\/ijo.2019.4805","volume":"55","author":"L Shan","year":"2019","unstructured":"Shan L, Zhao M, Lu Y, Ning H, Yang S, Song Y, et al. CENPE promotes lung adenocarcinoma proliferation and is directly regulated by FOXM1. Int J Oncol. 2019;55(1):257\u201366. https:\/\/doi.org\/10.3892\/ijo.2019.4805.","journal-title":"Int J Oncol"},{"issue":"22","key":"530_CR42","doi-asserted-by":"publisher","first-page":"5889","DOI":"10.1158\/0008-5472.CAN-12-1991","volume":"72","author":"X Wan","year":"2012","unstructured":"Wan X, Yeung C, Kim SY, Dolan JG, Ngo VN, Burkett S, et al. Identification of FoxM1\/Bub1b signaling pathway as a required component for growth and survival of rhabdomyosarcoma. Cancer Res. 2012;72(22):5889\u201399. https:\/\/doi.org\/10.1158\/0008-5472.CAN-12-1991.","journal-title":"Cancer Res"},{"issue":"8","key":"530_CR43","doi-asserted-by":"publisher","first-page":"543","DOI":"10.1038\/s41580-023-00593-z","volume":"24","author":"AD McAinsh","year":"2023","unstructured":"McAinsh AD, Kops GJPL. Principles and dynamics of spindle assembly checkpoint signalling. Nat Rev Mol Cell Biol. 2023;24(8):543\u201359. https:\/\/doi.org\/10.1038\/s41580-023-00593-z.","journal-title":"Nat Rev Mol Cell Biol"},{"issue":"2","key":"530_CR44","doi-asserted-by":"publisher","first-page":"313","DOI":"10.1042\/EBC20190073","volume":"64","author":"B Craske","year":"2020","unstructured":"Craske B, Welburn JPI. Leaving no-one behind: how CENP-E facilitates chromosome alignment. Essays Biochem. 2020;64(2):313\u201324. https:\/\/doi.org\/10.1042\/EBC20190073.","journal-title":"Essays Biochem"},{"key":"530_CR45","doi-asserted-by":"publisher","first-page":"6894684","DOI":"10.1155\/2020\/6894684","volume":"2020","author":"F Tian","year":"2020","unstructured":"Tian F, Tang P, Sun Z, Zhang R, Zhu D, He J, et al. miR-210 in Exosomes Derived from Macrophages under High Glucose Promotes Mouse Diabetic Obesity Pathogenesis by Suppressing NDUFA4 Expression. J Diabetes Res. 2020;2020:6894684. https:\/\/doi.org\/10.1155\/2020\/6894684.","journal-title":"J Diabetes Res"},{"issue":"39","key":"530_CR46","doi-asserted-by":"publisher","first-page":"e2106947118","DOI":"10.1073\/pnas.2106947118","volume":"118","author":"RK Aggarwal","year":"2021","unstructured":"Aggarwal RK, Luchtel RA, Machha V, Tischer A, Zou Y, Pradhan K, et al. Functional succinate dehydrogenase deficiency is a common adverse feature of clear cell renal cancer. Proc Natl Acad Sci U S A. 2021;118(39):e2106947118. https:\/\/doi.org\/10.1073\/pnas.2106947118.","journal-title":"Proc Natl Acad Sci U S A"},{"issue":"12","key":"530_CR47","doi-asserted-by":"publisher","first-page":"2181","DOI":"10.1002\/mc.23107","volume":"58","author":"E Pranzini","year":"2019","unstructured":"Pranzini E, Leo A, Rapizzi E, Ramazzotti M, Magherini F, Giovannelli L, et al. miR-210-3p mediates metabolic adaptation and sustains DNA damage repair of resistant colon cancer cells to treatment with 5-fluorouracil. Mol Carcinog. 2019;58(12):2181\u201392. https:\/\/doi.org\/10.1002\/mc.23107.","journal-title":"Mol Carcinog"},{"issue":"2","key":"530_CR48","doi-asserted-by":"publisher","first-page":"110820","DOI":"10.1016\/j.ygeno.2024.110820","volume":"116","author":"TA van Gelderen","year":"2024","unstructured":"van Gelderen TA, Ribas L. MiR-210 promotes immune- and suppresses oocyte meiosis-related genes in the zebrafish ovarian cells. Genomics. 2024;116(2):110820. https:\/\/doi.org\/10.1016\/j.ygeno.2024.110820.","journal-title":"Genomics"},{"issue":"6","key":"530_CR49","doi-asserted-by":"publisher","first-page":"2064","DOI":"10.1002\/hep.24614","volume":"54","author":"Q Ying","year":"2011","unstructured":"Ying Q, Liang L, Guo W, Zha R, Tian Q, Huang S, et al. Hypoxia-inducible microRNA-210 augments the metastatic potential of tumor cells by targeting vacuole membrane protein 1 in hepatocellular carcinoma. Hepatology. 2011;54(6):2064\u201375. https:\/\/doi.org\/10.1002\/hep.24614.","journal-title":"Hepatology"},{"key":"530_CR50","doi-asserted-by":"publisher","DOI":"10.3389\/fphar.2012.00134","volume":"3","author":"H Cui","year":"2012","unstructured":"Cui H, Grosso S, Schelter F, Mari B, Kr\u00fcger A. On the pro-metastatic stress response to cancer therapies: evidence for a positive co-operation between TIMP-1, HIF-1\u03b1, and miR-210. Front Pharmacol. 2012;3:134. https:\/\/doi.org\/10.3389\/fphar.2012.00134.","journal-title":"Front Pharmacol"},{"issue":"1","key":"530_CR51","doi-asserted-by":"publisher","first-page":"117","DOI":"10.1186\/s12943-017-0688-6","volume":"16","author":"D Ren","year":"2017","unstructured":"Ren D, Yang Q, Dai Y, Guo W, Du H, Song L, et al. Oncogenic miR-210-3p promotes prostate cancer cell EMT and bone metastasis via NF-\u03baB signaling pathway. Mol Cancer. 2017;16(1):117. https:\/\/doi.org\/10.1186\/s12943-017-0688-6.","journal-title":"Mol Cancer"},{"issue":"18","key":"530_CR52","doi-asserted-by":"publisher","first-page":"3191","DOI":"10.3390\/cancers16183191","volume":"16","author":"P Rida","year":"2024","unstructured":"Rida P, Baker S, Saidykhan A, Bown I, Jinna N. FOXM1 Transcriptionally Co-Upregulates Centrosome Amplification and Clustering Genes and Is a Biomarker for Poor Prognosis in Androgen Receptor-Low Triple-Negative Breast Cancer. Cancers (Basel). 2024;16(18):3191. https:\/\/doi.org\/10.3390\/cancers16183191.","journal-title":"Cancers (Basel)"},{"issue":"13","key":"530_CR53","doi-asserted-by":"publisher","first-page":"2696","DOI":"10.1128\/MCB.01242-10","volume":"31","author":"TJ Kelly","year":"2011","unstructured":"Kelly TJ, Souza AL, Clish CB, Puigserver P. A hypoxia-induced positive feedback loop promotes hypoxia-inducible factor 1alpha stability through miR-210 suppression of glycerol-3-phosphate dehydrogenase 1-like. Mol Cell Biol. 2011;31(13):2696\u2013706. https:\/\/doi.org\/10.1128\/MCB.01242-10.","journal-title":"Mol Cell Biol"},{"issue":"11","key":"530_CR54","doi-asserted-by":"publisher","first-page":"E2194","DOI":"10.1210\/jc.2012-2410","volume":"97","author":"A Merlo","year":"2012","unstructured":"Merlo A, de Quiros SB, Secades P, Zambrano I, Balb\u00edn M, Astudillo A, et al. Identification of a Signaling Axis HIF-1\u03b1\/MicroRNA-210\/ISCU Independent of SDH Mutation That Defines a Subgroup of Head and Neck Paragangliomas. J Clin Endocrinol Metab. 2012;97(11):E2194\u2013200. https:\/\/doi.org\/10.1210\/jc.2012-2410.","journal-title":"J Clin Endocrinol Metab"},{"issue":"9","key":"530_CR55","doi-asserted-by":"publisher","first-page":"2813","DOI":"10.1002\/jcb.24178","volume":"113","author":"AD Walters","year":"2012","unstructured":"Walters AD, Bommakanti A, Cohen-Fix O. Shaping the nucleus: factors and forces. J Cell Biochem. 2012;113(9):2813\u201321. https:\/\/doi.org\/10.1002\/jcb.24178.","journal-title":"J Cell Biochem"},{"key":"530_CR56","doi-asserted-by":"publisher","DOI":"10.1016\/j.biopha.2019.108608","volume":"112","author":"Y Lu","year":"2019","unstructured":"Lu Y, Li Y, Wang Z, Xie S, Wang Q, Lei X, et al. Downregulation of RGMA by HIF-1A\/miR-210-3p axis promotes cell proliferation in oral squamous cell carcinoma. Biomed Pharmacother. 2019;112:108608. https:\/\/doi.org\/10.1016\/j.biopha.2019.108608.","journal-title":"Biomed Pharmacother"},{"issue":"5","key":"530_CR57","doi-asserted-by":"publisher","first-page":"755","DOI":"10.1016\/j.jocn.2013.06.024","volume":"21","author":"N-S Lai","year":"2014","unstructured":"Lai N-S, Dong Q-S, Ding H, Miao Z-L, Lin Y-C. MicroRNA-210 overexpression predicts poorer prognosis in glioma patients. J Clin Neurosci. 2014;21(5):755\u201360. https:\/\/doi.org\/10.1016\/j.jocn.2013.06.024.","journal-title":"J Clin Neurosci"},{"issue":"9","key":"530_CR58","first-page":"15283","volume":"8","author":"X Xie","year":"2015","unstructured":"Xie X, Wu W, Liang L, Han S, Chen T, Pan S, et al. Prognostic role of microRNA-210 in various carcinomas: a meta-analysis. Int J Clin Exp Med. 2015;8(9):15283\u20139.","journal-title":"Int J Clin Exp Med"},{"issue":"5","key":"530_CR59","doi-asserted-by":"publisher","DOI":"10.3390\/ijms22052622","volume":"22","author":"\u00c1 Bartha","year":"2021","unstructured":"Bartha \u00c1, Gy\u0151rffy B. TNMplot.com: a web tool for the comparison of gene expression in normal, tumor and metastatic tissues. Int J Mol Sci. 2021;22(5):2622. https:\/\/doi.org\/10.3390\/ijms22052622.","journal-title":"Int J Mol Sci"},{"issue":"23","key":"530_CR60","doi-asserted-by":"publisher","first-page":"5734","DOI":"10.1111\/bph.70215","volume":"182","author":"M Posta","year":"2025","unstructured":"Posta M, Gy\u0151rffy B. Pathway-level mutational signatures predict breast cancer outcomes and reveal therapeutic targets. Br J Pharmacol. 2025;182(23):5734\u201347. https:\/\/doi.org\/10.1111\/bph.70215.","journal-title":"Br J Pharmacol"},{"issue":"7403","key":"530_CR61","doi-asserted-by":"publisher","first-page":"346","DOI":"10.1038\/nature10983","volume":"486","author":"C Curtis","year":"2012","unstructured":"Curtis C, Shah SP, Chin S-F, Turashvili G, Rueda OM, Dunning MJ, et al. The genomic and transcriptomic architecture of 2,000 breast tumours reveals novel subgroups. Nature. 2012;486(7403):346\u201352. https:\/\/doi.org\/10.1038\/nature10983.","journal-title":"Nature"},{"key":"530_CR62","doi-asserted-by":"publisher","first-page":"4101","DOI":"10.1016\/j.csbj.2021.07.014","volume":"19","author":"B Gy\u0151rffy","year":"2021","unstructured":"Gy\u0151rffy B. Survival analysis across the entire transcriptome identifies biomarkers with the highest prognostic power in breast cancer. Comput Struct Biotechnol J. 2021;19:4101\u20139. https:\/\/doi.org\/10.1016\/j.csbj.2021.07.014.","journal-title":"Comput Struct Biotechnol J"}],"container-title":["Molecular Biomedicine"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/link.springer.com\/content\/pdf\/10.1186\/s43556-026-00530-4.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/link.springer.com\/article\/10.1186\/s43556-026-00530-4","content-type":"text\/html","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/link.springer.com\/content\/pdf\/10.1186\/s43556-026-00530-4.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,8,10]],"date-time":"2026-08-10T14:03:26Z","timestamp":1786370606000},"score":1,"resource":{"primary":{"URL":"https:\/\/link.springer.com\/10.1186\/s43556-026-00530-4"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,8,10]]},"references-count":62,"journal-issue":{"issue":"1","published-online":{"date-parts":[[2026,12]]}},"alternative-id":["530"],"URL":"https:\/\/doi.org\/10.1186\/s43556-026-00530-4","relation":{},"ISSN":["2662-8651"],"issn-type":[{"value":"2662-8651","type":"electronic"}],"subject":[],"published":{"date-parts":[[2026,8,10]]},"assertion":[{"value":"5 December 2025","order":1,"name":"received","label":"Received","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"23 July 2026","order":2,"name":"revised","label":"Revised","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"24 July 2026","order":3,"name":"accepted","label":"Accepted","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"10 August 2026","order":4,"name":"first_online","label":"First Online","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"Not applicable.","order":1,"name":"Ethics","label":"Ethics approval and consent to participate","group":{"name":"EthicsHeading","label":"Declarations"}},{"value":"Not applicable.","order":2,"name":"Ethics","label":"Consent for publication","group":{"name":"EthicsHeading","label":"Declarations"}},{"value":"The authors declare that they have no competing interests.","order":3,"name":"Ethics","label":"Competing interest","group":{"name":"EthicsHeading","label":"Declarations"}}],"article-number":"134"}}