{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,7,24]],"date-time":"2026-07-24T04:40:06Z","timestamp":1784868006512,"version":"3.55.0"},"reference-count":47,"publisher":"Springer Science and Business Media LLC","issue":"1","license":[{"start":{"date-parts":[[2025,4,28]],"date-time":"2025-04-28T00:00:00Z","timestamp":1745798400000},"content-version":"tdm","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0"},{"start":{"date-parts":[[2025,4,28]],"date-time":"2025-04-28T00:00:00Z","timestamp":1745798400000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0"}],"funder":[{"DOI":"10.13039\/501100002428","name":"Austrian Science Fund","doi-asserted-by":"publisher","award":["P31199-B27"],"award-info":[{"award-number":["P31199-B27"]}],"id":[{"id":"10.13039\/501100002428","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100002428","name":"Austrian Science Fund","doi-asserted-by":"publisher","award":["P31199-B27"],"award-info":[{"award-number":["P31199-B27"]}],"id":[{"id":"10.13039\/501100002428","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100002428","name":"Austrian Science Fund","doi-asserted-by":"publisher","award":["P31199-B27"],"award-info":[{"award-number":["P31199-B27"]}],"id":[{"id":"10.13039\/501100002428","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100002428","name":"Austrian Science Fund","doi-asserted-by":"publisher","award":["P31199-B27"],"award-info":[{"award-number":["P31199-B27"]}],"id":[{"id":"10.13039\/501100002428","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100005204","name":"\u00d6sterreichische Austauschdienst","doi-asserted-by":"publisher","award":["Marietta-Blau"],"award-info":[{"award-number":["Marietta-Blau"]}],"id":[{"id":"10.13039\/501100005204","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100001711","name":"Schweizerischer Nationalfonds zur F\u00f6rderung der Wissenschaftlichen Forschung","doi-asserted-by":"publisher","award":["PCEFP3_181102"],"award-info":[{"award-number":["PCEFP3_181102"]}],"id":[{"id":"10.13039\/501100001711","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100003246","name":"Nederlandse Organisatie voor Wetenschappelijk Onderzoek","doi-asserted-by":"publisher","award":["016.Vidi.188.014"],"award-info":[{"award-number":["016.Vidi.188.014"]}],"id":[{"id":"10.13039\/501100003246","id-type":"DOI","asserted-by":"publisher"}]}],"content-domain":{"domain":["link.springer.com"],"crossmark-restriction":false},"short-container-title":["Sci Rep"],"abstract":"<jats:title>Abstract<\/jats:title>\n                  <jats:p>The neuromuscular junction (NMJ) is the unique interface between lower motor neurons and skeletal muscle fibers and is indispensable for muscle function. Tight control of its localized formation at the center of every muscle fiber, and maintenance throughout lifetime are sustained by muscle-specific kinase (MuSK). MuSK acts as central regulator of acetylcholine receptor clustering at the postsynapse. Localized and temporally controlled signaling of MuSK is primarily achieved by tyrosine autophosphorylation and inhibition thereof. Previous investigations suggested serine phosphorylation of the activation domain as an additional modulator of MuSK activation. Here we identified calcium\/calmodulin dependent protein kinase II (CaMK2) and in particular CaMK2\u03b2 as novel catalyst of MuSK activation and confirmed its capability to phosphorylate MuSK in heterologous cells. However, whereas CaMK2\u03b2 absence in muscle cells reduced AChR clustering, MuSK phosphorylation was unchanged. Accordingly, we ruled out MuSK phosphorylation as the cause of synapse fragmentation in a mouse model for myotonic dystrophy type 1, in which the muscle-specific splice-variant of CaMK2\u03b2 is missing, or as the cause of ataxia or delayed muscle development in CaMK2\u03b2 knockout animals. Histological characterization of muscles of CaMK2\u03b2 knockout mice indicated specific roles of CaMK2\u03b2 in fast glycolytic versus slow oxidative muscle. Taken together, our data shows that MuSK can be phosphorylated by CaMK2\u03b2, but loss of CaMK2\u03b2 is likely compensated by other CaMK2 paralogs at the NMJ.<\/jats:p>","DOI":"10.1038\/s41598-025-95053-3","type":"journal-article","created":{"date-parts":[[2025,4,28]],"date-time":"2025-04-28T09:29:54Z","timestamp":1745832594000},"update-policy":"https:\/\/doi.org\/10.1007\/springer_crossmark_policy","source":"Crossref","is-referenced-by-count":4,"title":["MuSK is a substrate for CaMK2\u03b2 but this interaction is dispensable for MuSK activation in vivo"],"prefix":"10.1038","volume":"15","author":[{"given":"Jakob J.","family":"Pr\u00f6mer","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Sara","family":"Wolske","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Perrine","family":"Castets","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Geeske M.","family":"van Woerden","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Cinzia","family":"Barresi","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Kevin C.","family":"O\u2019Connor","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Ruth","family":"Herbst","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"297","published-online":{"date-parts":[[2025,4,28]]},"reference":[{"key":"95053_CR1","doi-asserted-by":"publisher","first-page":"501","DOI":"10.1016\/s0092-8674(00)81251-9","volume":"85","author":"TM DeChiara","year":"1996","unstructured":"DeChiara, T. M. et al. The receptor tyrosine kinase musk is required for neuromuscular junction formation in vivo. Cell 85, 501\u2013512. https:\/\/doi.org\/10.1016\/s0092-8674(00)81251-9 (1996).","journal-title":"Cell"},{"key":"95053_CR2","doi-asserted-by":"publisher","first-page":"525","DOI":"10.1016\/s0092-8674(00)81253-2","volume":"85","author":"M Gautam","year":"1996","unstructured":"Gautam, M. et al. Defective neuromuscular synaptogenesis in agrin-deficient mutant mice. Cell 85, 525\u2013535. https:\/\/doi.org\/10.1016\/s0092-8674(00)81253-2 (1996).","journal-title":"Cell"},{"key":"95053_CR3","doi-asserted-by":"publisher","first-page":"4993","DOI":"10.1242\/dev.02696","volume":"133","author":"SD Weatherbee","year":"2006","unstructured":"Weatherbee, S. D., Anderson, K. V. & Niswander, L. A. LDL-receptor-related protein 4 is crucial for formation of the neuromuscular junction. Dev. (Cambridge England). 133, 4993\u20135000. https:\/\/doi.org\/10.1242\/dev.02696 (2006).","journal-title":"Dev. (Cambridge England)"},{"key":"95053_CR4","doi-asserted-by":"publisher","first-page":"247","DOI":"10.1101\/gad.180885.111","volume":"26","author":"Y Zong","year":"2012","unstructured":"Zong, Y. et al. Structural basis of agrin-LRP4-MuSK signaling. Genes Dev. 26, 247\u2013258. https:\/\/doi.org\/10.1101\/gad.180885.111 (2012).","journal-title":"Genes Dev."},{"key":"95053_CR5","doi-asserted-by":"publisher","first-page":"67","DOI":"10.1093\/emboj\/19.1.67","volume":"19","author":"R Herbst","year":"2000","unstructured":"Herbst, R. & Burden, S. J. The juxtamembrane region of musk has a critical role in agrin-mediated signaling. EMBO J. 19, 67\u201377. https:\/\/doi.org\/10.1093\/emboj\/19.1.67 (2000).","journal-title":"EMBO J."},{"key":"95053_CR6","doi-asserted-by":"publisher","first-page":"1802","DOI":"10.1126\/science.1127142","volume":"312","author":"K Okada","year":"2006","unstructured":"Okada, K. et al. The muscle protein Dok-7 is essential for neuromuscular synaptogenesis. Science 312, 1802\u20131805. https:\/\/doi.org\/10.1126\/science.1127142 (2006).","journal-title":"Science"},{"key":"95053_CR7","doi-asserted-by":"publisher","first-page":"2451","DOI":"10.1101\/gad.1977710","volume":"24","author":"PT Hallock","year":"2010","unstructured":"Hallock, P. T. et al. Dok-7 regulates neuromuscular synapse formation by recruiting Crk and Crk-L. Genes Dev. 24, 2451\u20132461. https:\/\/doi.org\/10.1101\/gad.1977710 (2010).","journal-title":"Genes Dev."},{"key":"95053_CR8","doi-asserted-by":"publisher","first-page":"110584","DOI":"10.1016\/j.cellsig.2022.110584","volume":"104","author":"J Pr\u00f6mer","year":"2023","unstructured":"Pr\u00f6mer, J., Barresi, C. & Herbst, R. From phosphorylation to phenotype - Recent key findings on kinase regulation, downstream signaling and disease surrounding the receptor tyrosine kinase musk. Cell. Signal. 104, 110584. https:\/\/doi.org\/10.1016\/j.cellsig.2022.110584 (2023).","journal-title":"Cell. Signal."},{"key":"95053_CR9","doi-asserted-by":"publisher","first-page":"100221","DOI":"10.1016\/j.mcpro.2022.100221","volume":"21","author":"HG Budayeva","year":"2022","unstructured":"Budayeva, H. G. et al. Phosphoproteome profiling of the receptor tyrosine kinase musk identifies tyrosine phosphorylation of Rab GTPases. Mol. Cell. Proteom. 21, 100221. https:\/\/doi.org\/10.1016\/j.mcpro.2022.100221 (2022).","journal-title":"Mol. Cell. Proteom."},{"key":"95053_CR10","doi-asserted-by":"publisher","first-page":"1993","DOI":"10.1074\/mcp.M113.036087","volume":"13","author":"G D\u00fcrnberger","year":"2014","unstructured":"D\u00fcrnberger, G. et al. Global analysis of muscle-specific kinase signaling by quantitative phosphoproteomics. Mol. Cell. Proteom. 13, 1993\u20132003. https:\/\/doi.org\/10.1074\/mcp.M113.036087 (2014).","journal-title":"Mol. Cell. Proteom."},{"key":"95053_CR11","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1038\/s41598-016-0001-8","volume":"6","author":"B Camurdanoglu","year":"2016","unstructured":"Camurdanoglu, B. et al. Musk kinase activity is modulated by a Serine phosphorylation site in the kinase loop. Sci. Rep. 6, 1\u201315 (2016).","journal-title":"Sci. Rep."},{"key":"95053_CR12","doi-asserted-by":"publisher","first-page":"1978","DOI":"10.1126\/science.1088583","volume":"302","author":"RN Kanadia","year":"2003","unstructured":"Kanadia, R. N. et al. A muscleblind knockout model for myotonic dystrophy. Science 302, 1978\u20131980. https:\/\/doi.org\/10.1126\/science.1088583 (2003).","journal-title":"Science"},{"key":"95053_CR13","doi-asserted-by":"publisher","first-page":"11","DOI":"10.1186\/s13395-024-00345-3","volume":"14","author":"D Falcetta","year":"2024","unstructured":"Falcetta, D. et al. CaMKIIbeta deregulation contributes to neuromuscular junction destabilization in myotonic dystrophy type I. Skelet. Muscle. 14, 11. https:\/\/doi.org\/10.1186\/s13395-024-00345-3 (2024).","journal-title":"Skelet. Muscle"},{"key":"95053_CR14","doi-asserted-by":"publisher","first-page":"4656","DOI":"10.1091\/mbc.e06-03-0252","volume":"17","author":"H O\u2019Leary","year":"2006","unstructured":"O\u2019Leary, H., Lasda, E. & Bayer, K. U. CaMKII\u03b2 association with the actin cytoskeleton is regulated by alternative splicing. Mol. Biol. Cell. 17, 4656\u20134665. https:\/\/doi.org\/10.1091\/mbc.e06-03-0252 (2006).","journal-title":"Mol. Biol. Cell"},{"key":"95053_CR15","doi-asserted-by":"publisher","first-page":"759","DOI":"10.1038\/s41586-022-05575-3","volume":"613","author":"JL Johnson","year":"2023","unstructured":"Johnson, J. L. et al. An atlas of substrate specificities for the human serine\/threonine Kinome. Nature 613, 759\u2013766. https:\/\/doi.org\/10.1038\/s41586-022-05575-3 (2023).","journal-title":"Nature"},{"key":"95053_CR16","doi-asserted-by":"publisher","first-page":"111064","DOI":"10.1016\/j.celrep.2022.111064","volume":"40","author":"C \u00d6zden","year":"2022","unstructured":"\u00d6zden, C. et al. CaMKII binds both substrates and activators at the active site. Cell. Rep. 40, 111064. https:\/\/doi.org\/10.1016\/j.celrep.2022.111064 (2022).","journal-title":"Cell. Rep."},{"key":"95053_CR17","doi-asserted-by":"publisher","first-page":"943","DOI":"10.1016\/0896-6273(94)90306-9","volume":"12","author":"PI Hanson","year":"1994","unstructured":"Hanson, P. I., Meyer, T., Stryer, L. & Schulman, H. Dual role of calmodulin in autophosphorylation of multifunctional cam kinase May underlie decoding of calcium signals. Neuron 12, 943\u2013956. https:\/\/doi.org\/10.1016\/0896-6273(94)90306-9 (1994).","journal-title":"Neuron"},{"key":"95053_CR18","doi-asserted-by":"publisher","first-page":"209","DOI":"10.1016\/j.brainresbull.2021.03.014","volume":"171","author":"M Proietti Onori","year":"2021","unstructured":"Proietti Onori, M. & van Woerden, G. M. Role of calcium\/calmodulin-dependent kinase 2 in neurodevelopmental disorders. Brain Res. Bull. 171, 209\u2013220. https:\/\/doi.org\/10.1016\/j.brainresbull.2021.03.014 (2021).","journal-title":"Brain Res. Bull."},{"key":"95053_CR19","doi-asserted-by":"publisher","first-page":"e105191","DOI":"10.1371\/journal.pone.0105191","volume":"9","author":"AD Bachstetter","year":"2014","unstructured":"Bachstetter, A. D. et al. Generation and behavior characterization of CaMKIIbeta knockout mice. PLoS One. 9, e105191. https:\/\/doi.org\/10.1371\/journal.pone.0105191 (2014).","journal-title":"PLoS One"},{"key":"95053_CR20","doi-asserted-by":"publisher","first-page":"26989","DOI":"10.1038\/srep26989","volume":"6","author":"MJ Kool","year":"2016","unstructured":"Kool, M. J., van de Bree, J. E., Bodde, H. E., Elgersma, Y. & van Woerden, G. M. The molecular, Temporal and region-specific requirements of the beta isoform of Calcium\/Calmodulin-dependent protein kinase type 2 (CAMK2B) in mouse locomotion. Sci. Rep. 6, 26989. https:\/\/doi.org\/10.1038\/srep26989 (2016).","journal-title":"Sci. Rep."},{"key":"95053_CR21","doi-asserted-by":"publisher","first-page":"12455","DOI":"10.1523\/JNEUROSCI.3309-10.2010","volume":"30","author":"I Martinez-Pena y Valenzuela","year":"2010","unstructured":"Martinez-Pena y Valenzuela, I., Mouslim, C. & Akaaboune, M. Calcium\/calmodulin kinase II-dependent acetylcholine receptor cycling at the mammalian neuromuscular junction in vivo. J. Neurosci. 30, 12455\u201312465. https:\/\/doi.org\/10.1523\/JNEUROSCI.3309-10.2010 (2010).","journal-title":"J. Neurosci."},{"key":"95053_CR22","doi-asserted-by":"publisher","first-page":"5598","DOI":"10.1093\/emboj\/17.19.5598","volume":"17","author":"KU Bayer","year":"1998","unstructured":"Bayer, K. U., Harbers, K. & Schulman, H. AlphaKAP is an anchoring protein for a novel cam kinase II isoform in skeletal muscle. EMBO J. 17, 5598\u20135605. https:\/\/doi.org\/10.1093\/emboj\/17.19.5598 (1998).","journal-title":"EMBO J."},{"key":"95053_CR23","doi-asserted-by":"publisher","first-page":"823","DOI":"10.1038\/nn.2329","volume":"12","author":"GM van Woerden","year":"2009","unstructured":"van Woerden, G. M. et al. BetaCaMKII controls the direction of plasticity at parallel fiber-Purkinje cell synapses. Nat. Neurosci. 12, 823\u2013825. https:\/\/doi.org\/10.1038\/nn.2329 (2009).","journal-title":"Nat. Neurosci."},{"key":"95053_CR24","doi-asserted-by":"publisher","first-page":"1645","DOI":"10.1002\/cphy.c130009","volume":"3","author":"B Blaauw","year":"2013","unstructured":"Blaauw, B., Schiaffino, S. & Reggiani, C. Mechanisms modulating skeletal muscle phenotype. Compr. Physiol. 3, 1645\u20131687. https:\/\/doi.org\/10.1002\/cphy.c130009 (2013).","journal-title":"Compr. Physiol."},{"key":"95053_CR25","doi-asserted-by":"publisher","first-page":"91","DOI":"10.1111\/sms.13828","volume":"31","author":"M Martinez-Canton","year":"2021","unstructured":"Martinez-Canton, M. et al. Role of camkii and Sarcolipin in muscle adaptations to strength training with different levels of fatigue in the set. Scand. J. Med. Sci. Sports. 31, 91\u2013103. https:\/\/doi.org\/10.1111\/sms.13828 (2021).","journal-title":"Scand. J. Med. Sci. Sports"},{"key":"95053_CR26","doi-asserted-by":"publisher","first-page":"15","DOI":"10.3389\/fphar.2014.00015","volume":"5","author":"CB Gray","year":"2014","unstructured":"Gray, C. B. & Heller Brown, J. CaMKIIdelta subtypes: localization and function. Front. Pharmacol. 5, 15. https:\/\/doi.org\/10.3389\/fphar.2014.00015 (2014).","journal-title":"Front. Pharmacol."},{"key":"95053_CR27","doi-asserted-by":"publisher","first-page":"1853","DOI":"10.1172\/jci27438","volume":"116","author":"J Backs","year":"2006","unstructured":"Backs, J., Song, K., Bezprozvannaya, S., Chang, S. & Olson, E. N. CaM kinase II selectively signals to histone deacetylase 4 during cardiomyocyte hypertrophy. J. Clin. Invest. 116, 1853\u20131864. https:\/\/doi.org\/10.1172\/jci27438 (2006).","journal-title":"J. Clin. Invest."},{"key":"95053_CR28","doi-asserted-by":"publisher","first-page":"144567","DOI":"10.1016\/j.gene.2020.144567","volume":"742","author":"SC Rothschild","year":"2020","unstructured":"Rothschild, S. C. et al. Genetic compensation of \u03b3 CaMKII, an evolutionarily conserved gene. Gene 742, 144567. https:\/\/doi.org\/10.1016\/j.gene.2020.144567 (2020). https:\/\/doi.org\/https:\/\/doi.org\/","journal-title":"Gene"},{"key":"95053_CR29","doi-asserted-by":"publisher","first-page":"206","DOI":"10.1126\/science.1321493","volume":"257","author":"AJ Silva","year":"1992","unstructured":"Silva, A. J., Paylor, R., Wehner, J. M. & Tonegawa, S. Impaired Spatial learning in \u03b1-calcium-calmodulin kinase II mutant mice. Science 257, 206\u2013211 (1992).","journal-title":"Science"},{"key":"95053_CR30","doi-asserted-by":"publisher","first-page":"5424","DOI":"10.1523\/JNEUROSCI.1341-18.2019","volume":"39","author":"MJ Kool","year":"2019","unstructured":"Kool, M. J. et al. CAMK2-Dependent signaling in neurons is essential for survival. J. Neurosci. 39, 5424\u20135439. https:\/\/doi.org\/10.1523\/JNEUROSCI.1341-18.2019 (2019).","journal-title":"J. Neurosci."},{"key":"95053_CR31","doi-asserted-by":"publisher","first-page":"10141","DOI":"10.1523\/JNEUROSCI.5105-10.2011","volume":"31","author":"NZ Borgesius","year":"2011","unstructured":"Borgesius, N. Z. et al. BetaCaMKII plays a nonenzymatic role in hippocampal synaptic plasticity and learning by targeting alphacamkii to synapses. J. Neurosci. 31, 10141\u201310148. https:\/\/doi.org\/10.1523\/JNEUROSCI.5105-10.2011 (2011).","journal-title":"J. Neurosci."},{"key":"95053_CR32","doi-asserted-by":"publisher","first-page":"283","DOI":"10.1016\/s0896-6273(03)00428-8","volume":"39","author":"CC Fink","year":"2003","unstructured":"Fink, C. C. et al. Selective regulation of neurite extension and synapse formation by the beta but not the alpha isoform of camkii. Neuron 39, 283\u2013297. https:\/\/doi.org\/10.1016\/s0896-6273(03)00428-8 (2003).","journal-title":"Neuron"},{"key":"95053_CR33","doi-asserted-by":"publisher","first-page":"15638","DOI":"10.1074\/jbc.M109864200","volume":"277","author":"P Macpherson","year":"2002","unstructured":"Macpherson, P., Kostrominova, T., Tang, H. & Goldman, D. Protein kinase C and calcium\/calmodulin-activated protein kinase II (CaMK II) suppress nicotinic acetylcholine receptor gene expression in mammalian muscle. A specific role for CaMK II in activity-dependent gene expression. J. Biol. Chem. 277, 15638\u201315646. https:\/\/doi.org\/10.1074\/jbc.M109864200 (2002).","journal-title":"J. Biol. Chem."},{"key":"95053_CR34","doi-asserted-by":"publisher","first-page":"1120","DOI":"10.1091\/mbc.e08-07-0759","volume":"20","author":"H Tang","year":"2009","unstructured":"Tang, H. et al. A histone deacetylase 4\/myogenin positive feedback loop coordinates denervation-dependent gene induction and suppression. Mol. Biol. Cell. 20, 1120\u20131131. https:\/\/doi.org\/10.1091\/mbc.e08-07-0759 (2009).","journal-title":"Mol. Biol. Cell."},{"key":"95053_CR35","doi-asserted-by":"publisher","first-page":"3943","DOI":"10.1074\/jbc.M511317200","volume":"281","author":"H Tang","year":"2006","unstructured":"Tang, H., Veldman, M. B. & Goldman, D. Characterization of a muscle-specific enhancer in human musk promoter reveals the essential role of Myogenin in controlling activity-dependent gene regulation. J. Biol. Chem. 281, 3943\u20133953. https:\/\/doi.org\/10.1074\/jbc.M511317200 (2006).","journal-title":"J. Biol. Chem."},{"key":"95053_CR36","doi-asserted-by":"publisher","first-page":"7","DOI":"10.1186\/s12899-014-0007-z","volume":"14","author":"W Eilers","year":"2014","unstructured":"Eilers, W. et al. CaMKII content affects contractile, but not mitochondrial, characteristics in regenerating skeletal muscle. BMC Physiol. 14, 7. https:\/\/doi.org\/10.1186\/s12899-014-0007-z (2014).","journal-title":"BMC Physiol."},{"key":"95053_CR37","doi-asserted-by":"publisher","first-page":"5099","DOI":"10.1093\/emboj\/18.18.5099","volume":"18","author":"EA Miska","year":"1999","unstructured":"Miska, E. A. et al. HDAC4 deacetylase associates with and represses the MEF2 transcription factor. EMBO J. 18, 5099\u20135107. https:\/\/doi.org\/10.1093\/emboj\/18.18.5099 (1999).","journal-title":"EMBO J."},{"key":"95053_CR38","doi-asserted-by":"publisher","first-page":"2459","DOI":"10.1172\/JCI31960","volume":"117","author":"MJ Potthoff","year":"2007","unstructured":"Potthoff, M. J. et al. Histone deacetylase degradation and MEF2 activation promote the formation of slow-twitch myofibers. J. Clin. Invest. 117, 2459\u20132467. https:\/\/doi.org\/10.1172\/JCI31960 (2007).","journal-title":"J. Clin. Invest."},{"key":"95053_CR39","doi-asserted-by":"publisher","unstructured":"Eigler, T. et al. ERK1\/2 Inhibition promotes robust myotube growth via camkii activation resulting in myoblast-to-myotube fusion. Dev. Cell. 56 (3349\u20133363 e3346). https:\/\/doi.org\/10.1016\/j.devcel.2021.11.022 (2021).","DOI":"10.1016\/j.devcel.2021.11.022"},{"key":"95053_CR40","doi-asserted-by":"publisher","unstructured":"Takata, K. et al. Characterization of pathogenic monoclonal autoantibodies derived from muscle-specific kinase myasthenia Gravis patients. JCI Insight. 4 https:\/\/doi.org\/10.1172\/jci.insight.127167 (2019).","DOI":"10.1172\/jci.insight.127167"},{"key":"95053_CR41","doi-asserted-by":"publisher","first-page":"2281","DOI":"10.1038\/nprot.2013.143","volume":"8","author":"FA Ran","year":"2013","unstructured":"Ran, F. A. et al. Genome engineering using the CRISPR-Cas9 system. Nat. Protoc. 8, 2281\u20132308. https:\/\/doi.org\/10.1038\/nprot.2013.143 (2013).","journal-title":"Nat. Protoc."},{"key":"95053_CR42","doi-asserted-by":"publisher","first-page":"768","DOI":"10.1016\/j.ajhg.2017.10.003","volume":"101","author":"S K\u00fcry","year":"2017","unstructured":"K\u00fcry, S. et al. De Novo mutations in protein kinase genes CAMK2A and CAMK2B cause intellectual disability. Am. J. Hum. Genet. 101, 768\u2013788. https:\/\/doi.org\/10.1016\/j.ajhg.2017.10.003 (2017).","journal-title":"Am. J. Hum. Genet."},{"key":"95053_CR43","doi-asserted-by":"publisher","first-page":"780659","DOI":"10.3389\/fnmol.2022.780659","volume":"15","author":"A Gemza","year":"2022","unstructured":"Gemza, A. et al. Internalization of Muscle-Specific kinase is increased by Agrin and independent of kinase-Activity, Lrp4 and dynamin. Front. Mol. Neurosci. 15, 780659. https:\/\/doi.org\/10.3389\/fnmol.2022.780659 (2022).","journal-title":"Front. Mol. Neurosci."},{"key":"95053_CR44","doi-asserted-by":"publisher","first-page":"619","DOI":"10.1002\/humu.23949","volume":"41","author":"PM Rodriguez Cruz","year":"2020","unstructured":"Rodriguez Cruz, P. M. et al. Congenital myasthenic syndrome due to mutations in MUSK suggests that the level of musk phosphorylation is crucial for governing synaptic structure. Hum. Mutat. 41, 619\u2013631. https:\/\/doi.org\/10.1002\/humu.23949 (2020).","journal-title":"Hum. Mutat."},{"key":"95053_CR45","doi-asserted-by":"publisher","first-page":"676","DOI":"10.1038\/nmeth.2019","volume":"9","author":"J Schindelin","year":"2012","unstructured":"Schindelin, J. et al. Fiji: an open-source platform for biological-image analysis. Nat. Methods. 9, 676\u2013682. https:\/\/doi.org\/10.1038\/nmeth.2019 (2012).","journal-title":"Nat. Methods"},{"key":"95053_CR46","doi-asserted-by":"publisher","first-page":"200128","DOI":"10.1098\/rsos.200128","volume":"7","author":"G Minty","year":"2020","unstructured":"Minty, G. et al. aNMJ-morph: a simple macro for rapid analysis of neuromuscular junction morphology. R Soc. Open. Sci. 7, 200128. https:\/\/doi.org\/10.1098\/rsos.200128 (2020).","journal-title":"R Soc. Open. Sci."},{"key":"95053_CR47","doi-asserted-by":"publisher","first-page":"4510","DOI":"10.1038\/s41467-020-18140-1","volume":"11","author":"DJ Ham","year":"2020","unstructured":"Ham, D. J. et al. The neuromuscular junction is a focal point of mTORC1 signaling in sarcopenia. Nat. Commun. 11, 4510. https:\/\/doi.org\/10.1038\/s41467-020-18140-1 (2020).","journal-title":"Nat. Commun."}],"container-title":["Scientific Reports"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.nature.com\/articles\/s41598-025-95053-3.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/www.nature.com\/articles\/s41598-025-95053-3","content-type":"text\/html","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/www.nature.com\/articles\/s41598-025-95053-3.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2025,5,2]],"date-time":"2025-05-02T21:02:28Z","timestamp":1746219748000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.nature.com\/articles\/s41598-025-95053-3"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2025,4,28]]},"references-count":47,"journal-issue":{"issue":"1","published-online":{"date-parts":[[2025,12]]}},"alternative-id":["95053"],"URL":"https:\/\/doi.org\/10.1038\/s41598-025-95053-3","relation":{"has-preprint":[{"id-type":"doi","id":"10.21203\/rs.3.rs-5371568\/v1","asserted-by":"object"}]},"ISSN":["2045-2322"],"issn-type":[{"value":"2045-2322","type":"electronic"}],"subject":[],"published":{"date-parts":[[2025,4,28]]},"assertion":[{"value":"1 November 2024","order":1,"name":"received","label":"Received","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"18 March 2025","order":2,"name":"accepted","label":"Accepted","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"28 April 2025","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 no competing interests.","order":2,"name":"Ethics","group":{"name":"EthicsHeading","label":"Competing interests"}}],"article-number":"14865"}}