{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,10,14]],"date-time":"2026-10-14T23:13:28Z","timestamp":1792019608137,"version":"4.4.0"},"reference-count":60,"publisher":"Springer Science and Business Media LLC","issue":"1","license":[{"start":{"date-parts":[[2026,1,16]],"date-time":"2026-01-16T00:00:00Z","timestamp":1768521600000},"content-version":"tdm","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by-nc-nd\/4.0"},{"start":{"date-parts":[[2026,2,6]],"date-time":"2026-02-06T00:00:00Z","timestamp":1770336000000},"content-version":"vor","delay-in-days":21,"URL":"https:\/\/creativecommons.org\/licenses\/by-nc-nd\/4.0"}],"content-domain":{"domain":["link.springer.com"],"crossmark-restriction":false},"short-container-title":["Sci Rep"],"DOI":"10.1038\/s41598-026-35002-w","type":"journal-article","created":{"date-parts":[[2026,1,16]],"date-time":"2026-01-16T08:16:59Z","timestamp":1768551419000},"update-policy":"https:\/\/doi.org\/10.1007\/springer_crossmark_policy","source":"Crossref","is-referenced-by-count":3,"title":["Transcriptomic characterization of the aberrant alternative splicing in skeletal muscles of sarcopenia patients"],"prefix":"10.1038","volume":"16","author":[{"given":"Yuanyuan","family":"Li","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Xiaoxuan","family":"Guo","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Sujun","family":"Li","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Xiaowen","family":"Wei","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Tianyi","family":"Li","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"297","published-online":{"date-parts":[[2026,1,16]]},"reference":[{"issue":"10191","key":"35002_CR1","doi-asserted-by":"publisher","first-page":"2636","DOI":"10.1016\/S0140-6736(19)31138-9","volume":"393","author":"AJ Cruz-Jentoft","year":"2019","unstructured":"Cruz-Jentoft, A. J. & Sayer, A. A. Sarcopenia. Lancet 393(10191), 2636\u20132646. https:\/\/doi.org\/10.1016\/S0140-6736(19)31138-9 (2019).","journal-title":"Lancet"},{"issue":"1","key":"35002_CR2","doi-asserted-by":"publisher","first-page":"86","DOI":"10.1002\/jcsm.12783","volume":"13","author":"F Petermann-Rocha","year":"2022","unstructured":"Petermann-Rocha, F. et al. Global prevalence of sarcopenia and severe sarcopenia: A systematic review and meta-analysis. J. Cachexia Sarcopenia Muscle 13(1), 86\u201399. https:\/\/doi.org\/10.1002\/jcsm.12783 (2022).","journal-title":"J. Cachexia Sarcopenia Muscle"},{"key":"35002_CR3","doi-asserted-by":"publisher","DOI":"10.3390\/nu13010183","author":"A Izzo","year":"2021","unstructured":"Izzo, A. et al. A narrative review on sarcopenia in type 2 diabetes mellitus: Prevalence and associated factors. Nutrients https:\/\/doi.org\/10.3390\/nu13010183 (2021).","journal-title":"Nutrients"},{"issue":"20","key":"35002_CR4","doi-asserted-by":"publisher","first-page":"1534","DOI":"10.1161\/circulationaha.123.064071","volume":"147","author":"AA Damluji","year":"2023","unstructured":"Damluji, A. A. et al. Sarcopenia and cardiovascular diseases. Circulation 147(20), 1534\u20131553. https:\/\/doi.org\/10.1161\/circulationaha.123.064071 (2023).","journal-title":"Circulation"},{"issue":"3","key":"35002_CR5","doi-asserted-by":"publisher","first-page":"221","DOI":"10.1002\/agm2.12168","volume":"4","author":"Y Feike","year":"2021","unstructured":"Feike, Y., Zhijie, L. & Wei, C. Advances in research on pharmacotherapy of sarcopenia. Aging Med (Milton) 4(3), 221\u2013233. https:\/\/doi.org\/10.1002\/agm2.12168 (2021).","journal-title":"Aging Med (Milton)"},{"issue":"4","key":"35002_CR6","doi-asserted-by":"publisher","first-page":"242","DOI":"10.1038\/s41580-022-00545-z","volume":"24","author":"LE Marasco","year":"2023","unstructured":"Marasco, L. E. & Kornblihtt, A. R. The physiology of alternative splicing. Nat. Rev. Mol. Cell Biol. 24(4), 242\u2013254. https:\/\/doi.org\/10.1038\/s41580-022-00545-z (2023).","journal-title":"Nat. Rev. Mol. Cell Biol."},{"issue":"4","key":"35002_CR7","doi-asserted-by":"publisher","first-page":"441","DOI":"10.4161\/rna.7.4.12258","volume":"7","author":"M Pistoni","year":"2010","unstructured":"Pistoni, M., Ghigna, C. & Gabellini, D. Alternative splicing and muscular dystrophy. RNA Biol. 7(4), 441\u2013452. https:\/\/doi.org\/10.4161\/rna.7.4.12258 (2010).","journal-title":"RNA Biol."},{"key":"35002_CR8","doi-asserted-by":"publisher","DOI":"10.3390\/genes13030401","author":"Q Liu","year":"2022","unstructured":"Liu, Q., Fang, L. & Wu, C. Alternative splicing and isoforms: From mechanisms to diseases. Genes https:\/\/doi.org\/10.3390\/genes13030401 (2022).","journal-title":"Genes"},{"issue":"8","key":"35002_CR9","doi-asserted-by":"publisher","first-page":"457","DOI":"10.1038\/s41583-023-00717-6","volume":"24","author":"D Nikom","year":"2023","unstructured":"Nikom, D. & Zheng, S. Alternative splicing in neurodegenerative disease and the promise of RNA therapies. Nat. Rev. Neurosci. 24(8), 457\u2013473. https:\/\/doi.org\/10.1038\/s41583-023-00717-6 (2023).","journal-title":"Nat. Rev. Neurosci."},{"key":"35002_CR10","doi-asserted-by":"publisher","DOI":"10.3390\/genes12091346","author":"J Lejman","year":"2021","unstructured":"Lejman, J. et al. Alternative splicing role in new therapies of spinal muscular atrophy. Genes https:\/\/doi.org\/10.3390\/genes12091346 (2021).","journal-title":"Genes"},{"issue":"2","key":"35002_CR11","doi-asserted-by":"publisher","DOI":"10.4081\/ejh.2013.e15","volume":"57","author":"M Malatesta","year":"2013","unstructured":"Malatesta, M. et al. Muscleblind-like1 undergoes ectopic relocation in the nuclei of skeletal muscles in myotonic dystrophy and sarcopenia. Eur. J. Histochem. 57(2), e15. https:\/\/doi.org\/10.4081\/ejh.2013.e15 (2013).","journal-title":"Eur. J. Histochem."},{"key":"35002_CR12","doi-asserted-by":"publisher","DOI":"10.1016\/j.mad.2021.111510","volume":"197","author":"EM Solovyeva","year":"2021","unstructured":"Solovyeva, E. M. et al. New insights into molecular changes in skeletal muscle aging and disease: Differential alternative splicing and senescence. Mech. Ageing Dev. 197, 111510. https:\/\/doi.org\/10.1016\/j.mad.2021.111510 (2021).","journal-title":"Mech. Ageing Dev."},{"key":"35002_CR13","doi-asserted-by":"publisher","DOI":"10.1016\/j.archger.2022.104920","volume":"108","author":"QQ Sun","year":"2023","unstructured":"Sun, Q. Q. et al. RNA analysis of diet-induced sarcopenic obesity in rats. Arch. Gerontol. Geriatr. 108, 104920. https:\/\/doi.org\/10.1016\/j.archger.2022.104920 (2023).","journal-title":"Arch. Gerontol. Geriatr."},{"issue":"1","key":"35002_CR14","doi-asserted-by":"publisher","first-page":"5808","DOI":"10.1038\/s41467-019-13694-1","volume":"10","author":"E Migliavacca","year":"2019","unstructured":"Migliavacca, E. et al. Mitochondrial oxidative capacity and NAD(+) biosynthesis are reduced in human sarcopenia across ethnicities. Nat. Commun. 10(1), 5808. https:\/\/doi.org\/10.1038\/s41467-019-13694-1 (2019).","journal-title":"Nat. Commun."},{"issue":"11","key":"35002_CR15","doi-asserted-by":"publisher","first-page":"8276","DOI":"10.1109\/tpami.2021.3103980","volume":"44","author":"B Yu","year":"2022","unstructured":"Yu, B. & Tao, D. Heatmap regression via randomized rounding. IEEE Trans. Pattern Anal. Mach. Intell. 44(11), 8276\u20138289. https:\/\/doi.org\/10.1109\/tpami.2021.3103980 (2022).","journal-title":"IEEE Trans. Pattern Anal. Mach. Intell."},{"issue":"11","key":"35002_CR16","doi-asserted-by":"publisher","first-page":"3393","DOI":"10.1021\/acssynbio.3c00423","volume":"12","author":"WM Shaw","year":"2023","unstructured":"Shaw, W. M., Khalil, A. S. & Ellis, T. A multiplex MoClo toolkit for extensive and flexible engineering of saccharomyces cerevisiae. ACS Synth. Biol. 12(11), 3393\u20133405. https:\/\/doi.org\/10.1021\/acssynbio.3c00423 (2023).","journal-title":"ACS Synth. Biol."},{"issue":"4","key":"35002_CR17","doi-asserted-by":"publisher","first-page":"R36","DOI":"10.1186\/gb-2013-14-4-r36","volume":"14","author":"D Kim","year":"2013","unstructured":"Kim, D. et al. TopHat2: Accurate alignment of transcriptomes in the presence of insertions, deletions and gene fusions. Genome Biol. 14(4), R36. https:\/\/doi.org\/10.1186\/gb-2013-14-4-r36 (2013).","journal-title":"Genome Biol."},{"issue":"1","key":"35002_CR18","doi-asserted-by":"publisher","first-page":"139","DOI":"10.1093\/bioinformatics\/btp616","volume":"26","author":"MD Robinson","year":"2010","unstructured":"Robinson, M. D., McCarthy, D. J. & Smyth, G. K. edgeR: A Bioconductor package for differential expression analysis of digital gene expression data. Bioinformatics 26(1), 139\u2013140. https:\/\/doi.org\/10.1093\/bioinformatics\/btp616 (2010).","journal-title":"Bioinformatics"},{"issue":"9","key":"35002_CR19","first-page":"15137","volume":"8","author":"HW Wang","year":"2015","unstructured":"Wang, H. W. et al. Saikosaponin-d attenuates ventilator-induced lung injury in rats. Int. J. Clin. Exp. Med. 8(9), 15137\u201315145 (2015).","journal-title":"Int. J. Clin. Exp. Med."},{"key":"35002_CR20","doi-asserted-by":"publisher","unstructured":"Maag J. L. V.(2018) gganatogram: An R package for modular visualisation of anatograms and tissues based on ggplot2 . F1000Res, 7: 1576. https:\/\/doi.org\/10.12688\/f1000research.16409.2.","DOI":"10.12688\/f1000research.16409.2"},{"key":"35002_CR21","doi-asserted-by":"publisher","first-page":"911","DOI":"10.1016\/j.bbagrm.2017.07.004","volume":"9","author":"H Xia","year":"1860","unstructured":"Xia, H. et al. (2017) CELF1 preferentially binds to exon-intron boundary and regulates alternative splicing in HeLa cells. Biochim. Biophys. Acta Gene Regul. Mech. 9, 911\u2013921. https:\/\/doi.org\/10.1016\/j.bbagrm.2017.07.004 (1860).","journal-title":"Biochim. Biophys. Acta Gene Regul. Mech."},{"key":"35002_CR22","doi-asserted-by":"publisher","DOI":"10.1016\/j.neo.2022.100858","volume":"35","author":"E Beebe","year":"2023","unstructured":"Beebe, E. et al. Proteomic profiling of canine fibrosarcoma and adjacent peritumoral tissue. Neoplasia 35, 100858. https:\/\/doi.org\/10.1016\/j.neo.2022.100858 (2023).","journal-title":"Neoplasia"},{"issue":"17","key":"35002_CR23","doi-asserted-by":"publisher","first-page":"2912","DOI":"10.1093\/bioinformatics\/btv300","volume":"31","author":"W Walter","year":"2015","unstructured":"Walter, W., S\u00e1nchez-Cabo, F. & Ricote, M. GOplot: An R package for visually combining expression data with functional analysis. Bioinformatics 31(17), 2912\u20132914. https:\/\/doi.org\/10.1093\/bioinformatics\/btv300 (2015).","journal-title":"Bioinformatics"},{"issue":"3","key":"35002_CR24","doi-asserted-by":"publisher","first-page":"538","DOI":"10.1002\/jcsm.12709","volume":"12","author":"WQ Xie","year":"2021","unstructured":"Xie, W. Q. et al. Mouse models of sarcopenia: Classification and evaluation. J. Cachexia Sarcopenia Muscle 12(3), 538\u2013554. https:\/\/doi.org\/10.1002\/jcsm.12709 (2021).","journal-title":"J. Cachexia Sarcopenia Muscle"},{"issue":"4","key":"35002_CR25","doi-asserted-by":"publisher","first-page":"402","DOI":"10.1006\/meth.2001.1262","volume":"25","author":"KJ Livak","year":"2001","unstructured":"Livak, K. J. & Schmittgen, T. D. Analysis of relative gene expression data using real-time quantitative PCR and the 2(-Delta Delta C(T)) method. Methods 25(4), 402\u2013408. https:\/\/doi.org\/10.1006\/meth.2001.1262 (2001).","journal-title":"Methods"},{"key":"35002_CR26","doi-asserted-by":"publisher","first-page":"136","DOI":"10.1016\/j.coph.2021.10.001","volume":"61","author":"PJ Liu","year":"2021","unstructured":"Liu, P. J. et al. Nutrient weight against sarcopenia: regulation of the IGF-1\/PI3K\/Akt\/FOXO pathway in quinoa metabolites. Curr. Opin. Pharmacol. 61, 136\u2013141. https:\/\/doi.org\/10.1016\/j.coph.2021.10.001 (2021).","journal-title":"Curr. Opin. Pharmacol."},{"key":"35002_CR27","doi-asserted-by":"publisher","DOI":"10.3390\/cells12242787","author":"B Gellhaus","year":"2023","unstructured":"Gellhaus, B. et al. Therapeutic consequences of targeting the IGF-1\/PI3K\/AKT\/FOXO3 axis in sarcopenia: A narrative review. Cells https:\/\/doi.org\/10.3390\/cells12242787 (2023).","journal-title":"Cells"},{"key":"35002_CR28","doi-asserted-by":"publisher","DOI":"10.3389\/fphys.2020.583478","volume":"11","author":"Z Zeng","year":"2020","unstructured":"Zeng, Z. et al. Exercise-induced autophagy suppresses sarcopenia through Akt\/mTOR and Akt\/FoxO3a signal pathways and AMPK-mediated mitochondrial quality control. Front. Physiol. 11, 583478. https:\/\/doi.org\/10.3389\/fphys.2020.583478 (2020).","journal-title":"Front. Physiol."},{"key":"35002_CR29","doi-asserted-by":"publisher","DOI":"10.3390\/metabo11050323","author":"J Liang","year":"2021","unstructured":"Liang, J. et al. Lifelong aerobic exercise alleviates sarcopenia by activating autophagy and inhibiting protein degradation via the AMPK\/PGC-1\u03b1 signaling pathway. Metabolites https:\/\/doi.org\/10.3390\/metabo11050323 (2021).","journal-title":"Metabolites"},{"issue":"1","key":"35002_CR30","doi-asserted-by":"publisher","first-page":"542","DOI":"10.1186\/s12877-021-02481-7","volume":"21","author":"Y Du","year":"2021","unstructured":"Du, Y. et al. Serum concentrations of oxytocin, DHEA and follistatin are associated with osteoporosis or sarcopenia in community-dwelling postmenopausal women. BMC Geriatr 21(1), 542. https:\/\/doi.org\/10.1186\/s12877-021-02481-7 (2021).","journal-title":"BMC Geriatr"},{"key":"35002_CR31","doi-asserted-by":"publisher","first-page":"133","DOI":"10.1093\/geroni\/igaa057.439","volume":"4","author":"T Liu","year":"2020","unstructured":"Liu, T. et al. Mitochondrial dysfunction and cellular senescence due to aging contributes to prostatic fibrosis. Innov. Aging 4, 133\u2013134. https:\/\/doi.org\/10.1093\/geroni\/igaa057.439 (2020).","journal-title":"Innov. Aging"},{"issue":"12","key":"35002_CR32","doi-asserted-by":"publisher","first-page":"967","DOI":"10.1016\/j.exger.2011.08.012","volume":"46","author":"F Sanchis-Gomar","year":"2011","unstructured":"Sanchis-Gomar, F., G\u00f3mez-Cabrera, M. C. & Vi\u00f1a, J. The loss of muscle mass and sarcopenia: Non hormonal intervention. Exp. Gerontol. 46(12), 967\u2013969. https:\/\/doi.org\/10.1016\/j.exger.2011.08.012 (2011).","journal-title":"Exp. Gerontol."},{"issue":"1","key":"35002_CR33","doi-asserted-by":"publisher","first-page":"5655","DOI":"10.1038\/s41467-022-33008-2","volume":"13","author":"T Sasako","year":"2022","unstructured":"Sasako, T. et al. Deletion of skeletal muscle Akt1\/2 causes osteosarcopenia and reduces lifespan in mice. Nat. Commun. 13(1), 5655. https:\/\/doi.org\/10.1038\/s41467-022-33008-2 (2022).","journal-title":"Nat. Commun."},{"issue":"Pt 3","key":"35002_CR34","doi-asserted-by":"publisher","DOI":"10.1016\/j.ijbiomac.2024.136326","volume":"281","author":"J Zhu","year":"2024","unstructured":"Zhu, J. et al. Single-nucleus transcriptional profiling reveals TCF7L2 as a key regulator in adipogenesis in goat skeletal muscle development. Int. J. Biol. Macromol. 281(Pt 3), 136326. https:\/\/doi.org\/10.1016\/j.ijbiomac.2024.136326 (2024).","journal-title":"Int. J. Biol. Macromol."},{"key":"35002_CR35","doi-asserted-by":"publisher","DOI":"10.1242\/jcs.242297","author":"O Contreras","year":"2020","unstructured":"Contreras, O. et al. TGF-\u03b2-driven downregulation of the transcription factor TCF7L2 affects Wnt\/\u03b2-catenin signaling in PDGFR\u03b1(+) fibroblasts. J. Cell. Sci. https:\/\/doi.org\/10.1242\/jcs.242297 (2020).","journal-title":"J. Cell. Sci."},{"issue":"10","key":"35002_CR36","doi-asserted-by":"publisher","first-page":"7183","DOI":"10.1002\/jcp.29617","volume":"235","author":"S Li","year":"2020","unstructured":"Li, S. et al. TCP11L2 promotes bovine skeletal muscle-derived satellite cell migration and differentiation via FMNL2. J. Cell. Physiol. 235(10), 7183\u20137193. https:\/\/doi.org\/10.1002\/jcp.29617 (2020).","journal-title":"J. Cell. Physiol."},{"key":"35002_CR37","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1016\/j.gene.2018.07.013","volume":"677","author":"F Zhang","year":"2018","unstructured":"Zhang, F. et al. Characterization of the Ubiquitin Specific Protease (USP) family members in the fast and slow muscle fibers from Chinese perch (Siniperca chuatsi). Gene 677, 1\u20139. https:\/\/doi.org\/10.1016\/j.gene.2018.07.013 (2018).","journal-title":"Gene"},{"key":"35002_CR38","doi-asserted-by":"publisher","first-page":"159","DOI":"10.1146\/annurev-physiol-022516-034255","volume":"80","author":"L Li","year":"2018","unstructured":"Li, L., Xiong, W. C. & Mei, L. Neuromuscular junction formation, aging, and disorders. Ann. Rev. Physiol. 80, 159\u2013188. https:\/\/doi.org\/10.1146\/annurev-physiol-022516-034255 (2018).","journal-title":"Ann. Rev. Physiol."},{"issue":"2","key":"35002_CR39","doi-asserted-by":"publisher","first-page":"176","DOI":"10.1016\/s1474-4422(21)00297-0","volume":"21","author":"AR Punga","year":"2022","unstructured":"Punga, A. R. et al. Epidemiology, diagnostics, and biomarkers of autoimmune neuromuscular junction disorders. Lancet Neurol. 21(2), 176\u2013188. https:\/\/doi.org\/10.1016\/s1474-4422(21)00297-0 (2022).","journal-title":"Lancet Neurol."},{"key":"35002_CR40","doi-asserted-by":"publisher","DOI":"10.1016\/j.arr.2023.101966","volume":"89","author":"WD Arnold","year":"2023","unstructured":"Arnold, W. D. & Clark, B. C. Neuromuscular junction transmission failure in aging and sarcopenia: the nexus of the neurological and muscular systems. Ageing Res. Rev. 89, 101966. https:\/\/doi.org\/10.1016\/j.arr.2023.101966 (2023).","journal-title":"Ageing Res. Rev."},{"issue":"22","key":"35002_CR41","doi-asserted-by":"publisher","first-page":"11185","DOI":"10.3390\/ijms262211185","volume":"26","author":"D Kupriyanova","year":"2025","unstructured":"Kupriyanova, D. et al. Neural cues and genomic clues: NGS insights into neurogenic sarcopenia and muscle atrophy. Int. J. Mol. Sci. 26(22), 11185 (2025).","journal-title":"Int. J. Mol. Sci."},{"issue":"1\u20132","key":"35002_CR42","doi-asserted-by":"publisher","first-page":"129","DOI":"10.1016\/j.physbeh.2007.05.040","volume":"92","author":"E Edstr\u00f6m","year":"2007","unstructured":"Edstr\u00f6m, E. et al. Factors contributing to neuromuscular impairment and sarcopenia during aging. Physiol. Behav. 92(1\u20132), 129\u2013135. https:\/\/doi.org\/10.1016\/j.physbeh.2007.05.040 (2007).","journal-title":"Physiol. Behav."},{"issue":"2","key":"35002_CR43","doi-asserted-by":"publisher","first-page":"C317","DOI":"10.1152\/ajpcell.00174.2021","volume":"321","author":"C Soendenbroe","year":"2021","unstructured":"Soendenbroe, C., Andersen, J. L. & Mackey, A. L. Muscle-nerve communication and the molecular assessment of human skeletal muscle denervation with aging. Am. J. Physiol. Cell Physiol. 321(2), C317-c329. https:\/\/doi.org\/10.1152\/ajpcell.00174.2021 (2021).","journal-title":"Am. J. Physiol. Cell Physiol."},{"key":"35002_CR44","doi-asserted-by":"publisher","DOI":"10.3390\/ijms24032976","author":"N Ryczek","year":"2023","unstructured":"Ryczek, N., \u0141y\u015b, A. & Maka\u0142owska, I. The functional meaning of 5\u2019UTR in protein-coding genes. Int. J. Mol. Sci. https:\/\/doi.org\/10.3390\/ijms24032976 (2023).","journal-title":"Int. J. Mol. Sci."},{"issue":"1","key":"35002_CR45","doi-asserted-by":"publisher","first-page":"101","DOI":"10.1007\/s00421-017-3749-z","volume":"118","author":"RB Zak","year":"2018","unstructured":"Zak, R. B. et al. Impact of local heating and cooling on skeletal muscle transcriptional response related to myogenesis and proteolysis. Eur. J. Appl. Physiol. 118(1), 101\u2013109. https:\/\/doi.org\/10.1007\/s00421-017-3749-z (2018).","journal-title":"Eur. J. Appl. Physiol."},{"issue":"7","key":"35002_CR46","doi-asserted-by":"publisher","first-page":"1261","DOI":"10.1016\/j.cell.2007.06.009","volume":"129","author":"BD Manning","year":"2007","unstructured":"Manning, B. D. & Cantley, L. C. AKT\/PKB signaling: Navigating downstream. Cell 129(7), 1261\u20131274. https:\/\/doi.org\/10.1016\/j.cell.2007.06.009 (2007).","journal-title":"Cell"},{"issue":"11","key":"35002_CR47","doi-asserted-by":"publisher","first-page":"1352","DOI":"10.1101\/gad.1089403","volume":"17","author":"XD Peng","year":"2003","unstructured":"Peng, X. D. et al. Dwarfism, impaired skin development, skeletal muscle atrophy, delayed bone development, and impeded adipogenesis in mice lacking Akt1 and Akt2. Genes Dev. 17(11), 1352\u20131365. https:\/\/doi.org\/10.1101\/gad.1089403 (2003).","journal-title":"Genes Dev."},{"key":"35002_CR48","doi-asserted-by":"publisher","DOI":"10.1016\/j.phymed.2022.154450","volume":"107","author":"L Zhang","year":"2022","unstructured":"Zhang, L. et al. Euonymine inhibits in-stent restenosis through enhancing contractile phenotype of vascular smooth muscle cells via modulating the PTEN\/AKT\/mTOR signaling pathway. Phytomedicine 107, 154450. https:\/\/doi.org\/10.1016\/j.phymed.2022.154450 (2022).","journal-title":"Phytomedicine"},{"key":"35002_CR49","doi-asserted-by":"publisher","DOI":"10.3892\/mmr.2021.11893","author":"J Dong","year":"2021","unstructured":"Dong, J. et al. Critical implication of the PTEN\/PI3K\/AKT pathway during BMP2-induced heterotopic ossification. Mol. Med. Rep. https:\/\/doi.org\/10.3892\/mmr.2021.11893 (2021).","journal-title":"Mol. Med. Rep."},{"issue":"Pt 5","key":"35002_CR50","doi-asserted-by":"publisher","first-page":"716","DOI":"10.1242\/jcs.042770","volume":"122","author":"A Mukherjee","year":"2009","unstructured":"Mukherjee, A. & Rotwein, P. Akt promotes BMP2-mediated osteoblast differentiation and bone development. J Cell Sci 122(Pt 5), 716\u2013726. https:\/\/doi.org\/10.1242\/jcs.042770 (2009).","journal-title":"J Cell Sci"},{"key":"35002_CR51","doi-asserted-by":"publisher","first-page":"55","DOI":"10.1186\/s13098-019-0449-3","volume":"11","author":"HH Wu","year":"2019","unstructured":"Wu, H. H. et al. TCF7L2 regulates pancreatic \u03b2-cell function through PI3K\/AKT signal pathway. Diabetol. Metab. Syndr. 11, 55. https:\/\/doi.org\/10.1186\/s13098-019-0449-3 (2019).","journal-title":"Diabetol. Metab. Syndr."},{"key":"35002_CR52","doi-asserted-by":"publisher","DOI":"10.3390\/biom12111639","author":"Y Sun","year":"2022","unstructured":"Sun, Y. et al. FGF9 promotes expression of HAS2 in palatal elevation via the Wnt\/\u03b2-Catenin\/TCF7L2 pathway. Biomolecules https:\/\/doi.org\/10.3390\/biom12111639 (2022).","journal-title":"Biomolecules"},{"issue":"2","key":"35002_CR53","doi-asserted-by":"publisher","first-page":"371","DOI":"10.1242\/dev.057463","volume":"138","author":"SJ Mathew","year":"2011","unstructured":"Mathew, S. J. et al. Connective tissue fibroblasts and Tcf4 regulate myogenesis. Development 138(2), 371\u2013384. https:\/\/doi.org\/10.1242\/dev.057463 (2011).","journal-title":"Development"},{"key":"35002_CR54","doi-asserted-by":"publisher","first-page":"55","DOI":"10.1186\/1471-2121-11-55","volume":"11","author":"M Gardberg","year":"2010","unstructured":"Gardberg, M. et al. Characterization of Diaphanous-related formin FMNL2 in human tissues. BMC Cell Biol. 11, 55. https:\/\/doi.org\/10.1186\/1471-2121-11-55 (2010).","journal-title":"BMC Cell Biol."},{"key":"35002_CR55","doi-asserted-by":"publisher","DOI":"10.1083\/jcb.201906111","author":"K Pfisterer","year":"2020","unstructured":"Pfisterer, K. et al. FMNL2 regulates dynamics of fascin in filopodia. J. Cell. Biol. https:\/\/doi.org\/10.1083\/jcb.201906111 (2020).","journal-title":"J. Cell. Biol."},{"issue":"6","key":"35002_CR56","doi-asserted-by":"publisher","first-page":"811","DOI":"10.1091\/mbc.E13-08-0443","volume":"25","author":"M Rosado","year":"2014","unstructured":"Rosado, M. et al. Critical roles for multiple formins during cardiac myofibril development and repair. Mol. Biol. Cell 25(6), 811\u2013827. https:\/\/doi.org\/10.1091\/mbc.E13-08-0443 (2014).","journal-title":"Mol. Biol. Cell"},{"issue":"13\u201314","key":"35002_CR57","doi-asserted-by":"publisher","first-page":"955","DOI":"10.1016\/j.cca.2010.03.013","volume":"411","author":"YR Wu","year":"2010","unstructured":"Wu, Y. R. et al. Ubiquitin specific proteases USP24 and USP40 and ubiquitin thiolesterase UCHL1 polymorphisms have synergic effect on the risk of Parkinson\u2019s disease among Taiwanese. Clin. Chim. Acta 411(13\u201314), 955\u2013958. https:\/\/doi.org\/10.1016\/j.cca.2010.03.013 (2010).","journal-title":"Clin. Chim. Acta"},{"issue":"7","key":"35002_CR58","doi-asserted-by":"publisher","DOI":"10.1096\/fj.202200246R","volume":"36","author":"Z Zhang","year":"2022","unstructured":"Zhang, Z. et al. MicroRNA profiling reveals miR-145-5p inhibits goat myoblast differentiation by targeting the coding domain sequence of USP13. Faseb J. 36(7), e22370. https:\/\/doi.org\/10.1096\/fj.202200246R (2022).","journal-title":"Faseb J."},{"key":"35002_CR59","doi-asserted-by":"publisher","DOI":"10.3390\/ijms221910289","author":"S Chaugule","year":"2021","unstructured":"Chaugule, S. et al. Deubiquitinating enzyme USP8 Is essential for skeletogenesis by regulating Wnt signaling. Int. J. Mol. Sci. https:\/\/doi.org\/10.3390\/ijms221910289 (2021).","journal-title":"Int. J. Mol. Sci."},{"issue":"21","key":"35002_CR60","doi-asserted-by":"publisher","first-page":"4659","DOI":"10.1111\/febs.15269","volume":"287","author":"E de la Vega","year":"2020","unstructured":"de la Vega, E. et al. USP7-dependent control of myogenin stability is required for terminal differentiation in skeletal muscle progenitors. Febs J. 287(21), 4659\u20134677. https:\/\/doi.org\/10.1111\/febs.15269 (2020).","journal-title":"Febs J."}],"container-title":["Scientific Reports"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.nature.com\/articles\/s41598-026-35002-w","content-type":"text\/html","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/www.nature.com\/articles\/s41598-026-35002-w.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/www.nature.com\/articles\/s41598-026-35002-w.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,2,6]],"date-time":"2026-02-06T22:03:36Z","timestamp":1770415416000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.nature.com\/articles\/s41598-026-35002-w"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,1,16]]},"references-count":60,"journal-issue":{"issue":"1","published-online":{"date-parts":[[2026,12]]}},"alternative-id":["35002"],"URL":"https:\/\/doi.org\/10.1038\/s41598-026-35002-w","relation":{},"ISSN":["2045-2322"],"issn-type":[{"value":"2045-2322","type":"electronic"}],"subject":[],"published":{"date-parts":[[2026,1,16]]},"assertion":[{"value":"17 September 2025","order":1,"name":"received","label":"Received","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"1 January 2026","order":2,"name":"accepted","label":"Accepted","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"16 January 2026","order":3,"name":"first_online","label":"First Online","group":{"name":"ArticleHistory","label":"Article History"}},{"order":1,"name":"Ethics","group":{"name":"EthicsHeading","label":"Declarations"}},{"value":"The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper.","order":2,"name":"Ethics","group":{"name":"EthicsHeading","label":"Competing interests"}},{"value":"The animal experiments were approved by the Ethics Committee of the First Affiliated Hospital of Zhengzhou University (2021-KS-HNSR093).","order":3,"name":"Ethics","group":{"name":"EthicsHeading","label":"Ethical approval"}}],"article-number":"5334"}}