{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,9,22]],"date-time":"2026-09-22T01:16:05Z","timestamp":1790039765464,"version":"4.0.1"},"reference-count":97,"publisher":"MDPI AG","issue":"17","license":[{"start":{"date-parts":[[2021,9,4]],"date-time":"2021-09-04T00:00:00Z","timestamp":1630713600000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"DOI":"10.13039\/501100000925","name":"National Health and Medical Research Council","doi-asserted-by":"publisher","award":["GNT1162652"],"award-info":[{"award-number":["GNT1162652"]}],"id":[{"id":"10.13039\/501100000925","id-type":"DOI","asserted-by":"publisher"}]}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":["IJMS"],"abstract":"<jats:p>Primary cilia are non-motile, cell cycle-associated organelles that can be found on most vertebrate cell types. Comprised of microtubule bundles organised into an axoneme and anchored by a mature centriole or basal body, primary cilia are dynamic signalling platforms that are intimately involved in cellular responses to their extracellular milieu. Defects in ciliogenesis or dysfunction in cilia signalling underlie a host of developmental disorders collectively referred to as ciliopathies, reinforcing important roles for cilia in human health. Whilst primary cilia have long been recognised to be present in striated muscle, their role in muscle is not well understood. However, recent studies indicate important contributions, particularly in skeletal muscle, that have to date remained underappreciated. Here, we explore recent revelations that the sensory and signalling functions of cilia on muscle progenitors regulate cell cycle progression, trigger differentiation and maintain a commitment to myogenesis. Cilia disassembly is initiated during myoblast fusion. However, the remnants of primary cilia persist in multi-nucleated myotubes, and we discuss their potential role in late-stage differentiation and myofiber formation. Reciprocal interactions between cilia and the extracellular matrix (ECM) microenvironment described for other tissues may also inform on parallel interactions in skeletal muscle. We also discuss emerging evidence that cilia on fibroblasts\/fibro\u2013adipogenic progenitors and myofibroblasts may influence cell fate in both a cell autonomous and non-autonomous manner with critical consequences for skeletal muscle ageing and repair in response to injury and disease. This review addresses the enigmatic but emerging role of primary cilia in satellite cells in myoblasts and myofibers during myogenesis, as well as the wider tissue microenvironment required for skeletal muscle formation and homeostasis.<\/jats:p>","DOI":"10.3390\/ijms22179605","type":"journal-article","created":{"date-parts":[[2021,9,6]],"date-time":"2021-09-06T13:15:56Z","timestamp":1630934156000},"page":"9605","update-policy":"https:\/\/doi.org\/10.3390\/mdpi_crossmark_policy","source":"Crossref","is-referenced-by-count":31,"title":["Cilia, Centrosomes and Skeletal Muscle"],"prefix":"10.3390","volume":"22","author":[{"given":"Dominic C. H.","family":"Ng","sequence":"first","affiliation":[{"name":"School of Biomedical Science, Faculty of Medicine, University of Queensland, St Lucia, Brisbane, QLD 4072, Australia"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Uda Y.","family":"Ho","sequence":"additional","affiliation":[{"name":"School of Biomedical Science, Faculty of Medicine, University of Queensland, St Lucia, Brisbane, QLD 4072, Australia"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Miranda D.","family":"Grounds","sequence":"additional","affiliation":[{"name":"School of Human Sciences, Faculty of Medicine, University of Western Australia, Perth, WA 6009, Australia"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"1968","published-online":{"date-parts":[[2021,9,4]]},"reference":[{"key":"ref_1","doi-asserted-by":"crossref","first-page":"17","DOI":"10.1016\/j.jbiomech.2011.08.008","article-title":"The mechanics of the primary cilium: An intricate structure with complex function","volume":"45","author":"Hoey","year":"2012","journal-title":"J. Biomech."},{"key":"ref_2","doi-asserted-by":"crossref","first-page":"663","DOI":"10.1016\/j.cell.2009.10.036","article-title":"Centrioles, centrosomes, and cilia in health and disease","volume":"139","author":"Nigg","year":"2009","journal-title":"Cell"},{"key":"ref_3","doi-asserted-by":"crossref","first-page":"506","DOI":"10.1016\/j.ceb.2013.04.011","article-title":"Assembling a primary cilium","volume":"25","author":"Kim","year":"2013","journal-title":"Curr. Opin. Cell Biol."},{"key":"ref_4","doi-asserted-by":"crossref","first-page":"285","DOI":"10.1007\/BF00220126","article-title":"Fine structure of the avian muscle spindle capsule","volume":"166","author":"Ovalle","year":"1976","journal-title":"Cell Tissue Res."},{"key":"ref_5","doi-asserted-by":"crossref","first-page":"89","DOI":"10.1016\/j.semcdb.2020.05.029","article-title":"Developmental and regenerative paradigms of cilia regulated hedgehog signaling","volume":"110","author":"Kopinke","year":"2021","journal-title":"Semin. Cell Dev. Biol."},{"key":"ref_6","doi-asserted-by":"crossref","first-page":"935","DOI":"10.1002\/jcp.25683","article-title":"The multifaceted roles of primary cilia in the regulation of stem cell properties and functions","volume":"232","author":"Lyu","year":"2017","journal-title":"J. Cell Physiol."},{"key":"ref_7","doi-asserted-by":"crossref","first-page":"817","DOI":"10.1038\/s42255-020-0251-4","article-title":"Skeletal muscle energy metabolism during exercise","volume":"2","author":"Hargreaves","year":"2020","journal-title":"Nat. Metab."},{"key":"ref_8","doi-asserted-by":"crossref","first-page":"aad9969","DOI":"10.1126\/science.aad9969","article-title":"Asymmetric division of clonal muscle stem cells coordinates muscle regeneration in vivo","volume":"353","author":"Gurevich","year":"2016","journal-title":"Science"},{"key":"ref_9","doi-asserted-by":"crossref","first-page":"999","DOI":"10.1016\/j.cell.2007.03.044","article-title":"Asymmetric self-renewal and commitment of satellite stem cells in muscle","volume":"129","author":"Kuang","year":"2007","journal-title":"Cell"},{"key":"ref_10","doi-asserted-by":"crossref","first-page":"798","DOI":"10.1016\/j.stemcr.2016.04.004","article-title":"Asymmetric distribution of primary cilia allocates satellite cells for self-renewal","volume":"6","author":"Marican","year":"2016","journal-title":"Stem Cell Rep."},{"key":"ref_11","doi-asserted-by":"crossref","first-page":"340","DOI":"10.1016\/j.cell.2017.06.035","article-title":"Ciliary hedgehog signaling restricts injury-induced adipogenesis","volume":"170","author":"Kopinke","year":"2017","journal-title":"Cell"},{"key":"ref_12","doi-asserted-by":"crossref","first-page":"214","DOI":"10.1083\/jcb.49.1.214","article-title":"Occurrence of centrioles during skeletal and cardiac myogenesis","volume":"49","author":"Przybylski","year":"1971","journal-title":"J. Cell Biol."},{"key":"ref_13","doi-asserted-by":"crossref","first-page":"433","DOI":"10.1097\/00005072-197207000-00003","article-title":"Ultrastructural development of explanted human skeletal muscle in tissue culture","volume":"31","author":"Mendell","year":"1972","journal-title":"J. Neuropathol. Exp. Neurol."},{"key":"ref_14","doi-asserted-by":"crossref","first-page":"170","DOI":"10.1016\/1047-8477(90)90111-O","article-title":"Fusion between myogenic cells in vivo: An ultrastructural study in regenerating murine skeletal muscle","volume":"105","author":"Robertson","year":"1990","journal-title":"J. Struct. Biol."},{"key":"ref_15","doi-asserted-by":"crossref","first-page":"9151","DOI":"10.1073\/pnas.1323265111","article-title":"Primary cilia control hedgehog signaling during muscle differentiation and are deregulated in rhabdomyosarcoma","volume":"111","author":"Fu","year":"2014","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_16","doi-asserted-by":"crossref","first-page":"691","DOI":"10.1146\/annurev-biochem-013118-111153","article-title":"Mechanism and regulation of centriole and cilium biogenesis","volume":"88","author":"Breslow","year":"2019","journal-title":"Ann. Rev. Biochem."},{"key":"ref_17","doi-asserted-by":"crossref","first-page":"2658","DOI":"10.1091\/mbc.e11-12-1056","article-title":"Calmodulin activation of Aurora-A kinase (AURKA) is required during ciliary disassembly and in mitosis","volume":"23","author":"Plotnikova","year":"2012","journal-title":"Mol. Biol. Cell"},{"key":"ref_18","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1002\/cm.20042","article-title":"Reorganization of microtubule nucleation during muscle differentiation","volume":"60","author":"Bugnard","year":"2005","journal-title":"Cell Motil. Cytoskelet."},{"key":"ref_19","doi-asserted-by":"crossref","first-page":"199","DOI":"10.1038\/s41581-019-0116-9","article-title":"Cellular signalling by primary cilia in development, organ function and disease","volume":"15","author":"Anvarian","year":"2019","journal-title":"Nat. Rev. Nephrol."},{"key":"ref_20","doi-asserted-by":"crossref","first-page":"707","DOI":"10.1083\/jcb.200503053","article-title":"Cilia and the cell cycle?","volume":"169","author":"Quarmby","year":"2005","journal-title":"J. Cell Biol."},{"key":"ref_21","doi-asserted-by":"crossref","first-page":"628","DOI":"10.1242\/jcs.03366","article-title":"The intraflagellar transport component IFT88\/polaris is a centrosomal protein regulating G1-S transition in non-ciliated cells","volume":"120","author":"Robert","year":"2007","journal-title":"J. Cell Sci."},{"key":"ref_22","doi-asserted-by":"crossref","first-page":"511","DOI":"10.1038\/s41568-018-0023-6","article-title":"Ciliary signalling in cancer","volume":"18","author":"Liu","year":"2018","journal-title":"Nat. Rev. Cancer"},{"key":"ref_23","doi-asserted-by":"crossref","first-page":"2058","DOI":"10.1158\/0008-5472.CAN-07-5838","article-title":"Cell cycle-dependent ciliogenesis and cancer","volume":"68","author":"Plotnikova","year":"2008","journal-title":"Cancer Res."},{"key":"ref_24","doi-asserted-by":"crossref","first-page":"137","DOI":"10.1016\/S0091-679X(08)94007-3","article-title":"Primary cilia and the cell cycle","volume":"94","author":"Plotnikova","year":"2009","journal-title":"Methods Cell Biol."},{"key":"ref_25","doi-asserted-by":"crossref","first-page":"264","DOI":"10.1016\/j.cell.2016.12.032","article-title":"Dynamic remodeling of membrane composition drives cell cycle through primary cilia excision","volume":"168","author":"Phua","year":"2017","journal-title":"Cell"},{"key":"ref_26","doi-asserted-by":"crossref","unstructured":"Mirvis, M., Siemers, K.A., Nelson, W.J., and Stearns, T.P. (2019). Primary cilium loss in mammalian cells occurs predominantly by whole-cilium shedding. PLoS Biol., 17.","DOI":"10.1371\/journal.pbio.3000381"},{"key":"ref_27","doi-asserted-by":"crossref","first-page":"3893","DOI":"10.1007\/s00018-013-1302-8","article-title":"Cell cycle progression by the repression of primary cilia formation in proliferating cells","volume":"70","author":"Goto","year":"2013","journal-title":"Cell Mol. Life Sci."},{"key":"ref_28","doi-asserted-by":"crossref","first-page":"1351","DOI":"10.1016\/j.cell.2007.04.035","article-title":"HEF1-dependent Aurora A activation induces disassembly of the primary cilium","volume":"129","author":"Pugacheva","year":"2007","journal-title":"Cell"},{"key":"ref_29","doi-asserted-by":"crossref","unstructured":"Venugopal, N., Ghosh, A., Gala, H., Aloysius, A., Vyas, N., and Dhawan, J. (2020). The primary cilium dampens proliferative signaling and represses a G2\/M transcriptional network in quiescent myoblasts. BMC Mol. Cell Biol., 21.","DOI":"10.1186\/s12860-020-00266-1"},{"key":"ref_30","doi-asserted-by":"crossref","first-page":"15248","DOI":"10.1038\/s41598-017-15649-2","article-title":"The primary cilium is necessary for the differentiation and the maintenance of human adipose progenitors into myofibroblasts","volume":"7","author":"Arrighi","year":"2017","journal-title":"Sci. Rep."},{"key":"ref_31","doi-asserted-by":"crossref","first-page":"4053","DOI":"10.1242\/dev.126.18.4053","article-title":"Sonic hedgehog controls epaxial muscle determination through Myf5 activation","volume":"126","author":"Borycki","year":"1999","journal-title":"Development"},{"key":"ref_32","doi-asserted-by":"crossref","first-page":"372","DOI":"10.1126\/science.1139740","article-title":"Patched1 regulates hedgehog signaling at the primary cilium","volume":"317","author":"Rohatgi","year":"2007","journal-title":"Science"},{"key":"ref_33","doi-asserted-by":"crossref","first-page":"225","DOI":"10.1016\/j.devcel.2013.12.020","article-title":"Gene regulatory networks and transcriptional mechanisms that control myogenesis","volume":"28","author":"Buckingham","year":"2014","journal-title":"Dev. Cell"},{"key":"ref_34","doi-asserted-by":"crossref","first-page":"1863","DOI":"10.1007\/s00018-005-5072-9","article-title":"Pleiotropic effects of sonic hedgehog on muscle satellite cells","volume":"62","author":"Koleva","year":"2005","journal-title":"Cell Mol. Life Sci."},{"key":"ref_35","doi-asserted-by":"crossref","first-page":"259","DOI":"10.1038\/ng1953","article-title":"Zebrafish miR-214 modulates Hedgehog signaling to specify muscle cell fate","volume":"39","author":"Flynt","year":"2007","journal-title":"Nat. Genet."},{"key":"ref_36","doi-asserted-by":"crossref","unstructured":"Li, X., Blagden, C.S., Bildsoe, H., Bonnin, M.A., Duprez, D., and Hughes, S.M. (2004). Hedgehog can drive terminal differentiation of amniote slow skeletal muscle. BMC Dev. Biol., 4.","DOI":"10.1186\/1471-213X-4-9"},{"key":"ref_37","doi-asserted-by":"crossref","first-page":"1498","DOI":"10.1016\/j.cub.2009.07.034","article-title":"Centriole age underlies asynchronous primary cilium growth in mammalian cells","volume":"19","author":"Anderson","year":"2009","journal-title":"Curr. Biol."},{"key":"ref_38","doi-asserted-by":"crossref","first-page":"397","DOI":"10.1016\/S1534-5807(02)00254-X","article-title":"The regulation of Notch signaling controls satellite cell activation and cell fate determination in postnatal myogenesis","volume":"3","author":"Conboy","year":"2002","journal-title":"Dev. Cell."},{"key":"ref_39","doi-asserted-by":"crossref","first-page":"541","DOI":"10.1016\/j.stem.2012.05.025","article-title":"Coordination of satellite cell activation and self-renewal by Par-complex-dependent asymmetric activation of p38alpha\/beta MAPK","volume":"11","author":"Troy","year":"2012","journal-title":"Cell Stem Cell"},{"key":"ref_40","doi-asserted-by":"crossref","first-page":"333","DOI":"10.1016\/j.cell.2013.08.060","article-title":"Asymmetric inheritance of centrosome-associated primary cilium membrane directs ciliogenesis after cell division","volume":"155","author":"Paridaen","year":"2013","journal-title":"Cell"},{"key":"ref_41","doi-asserted-by":"crossref","first-page":"2103","DOI":"10.1101\/gad.187807.112","article-title":"Sonic hedgehog acts cell-autonomously on muscle precursor cells to generate limb muscle diversity","volume":"26","author":"Anderson","year":"2012","journal-title":"Genes Dev."},{"key":"ref_42","doi-asserted-by":"crossref","unstructured":"Cruz-Migoni, S.B., Imran, K.M., Wahid, A., Rahman, O., Briscoe, J., and Borycki, A.-G. (2019). A switch in cilia-mediated Hedgehog signaling controls muscle stem cell quiescence and cell cycle progression. bioRxiv.","DOI":"10.1101\/2019.12.21.884601"},{"key":"ref_43","doi-asserted-by":"crossref","unstructured":"Brun, C., Sincennes, M.-C., Lin, A.Y.T., Hall, D., Jarassier, W., Feige, P., Ritso, M., Le Grand, F., and Rudnicki, M.A. (2020). GLI3 processing by the primary cilium regulates muscle stem cell entry into GAlert. bioRxiv.","DOI":"10.1101\/2020.12.07.415273"},{"key":"ref_44","doi-asserted-by":"crossref","first-page":"4389","DOI":"10.1074\/jbc.M112.400184","article-title":"Hedgehog signaling regulates MyoD expression and activity","volume":"288","author":"Voronova","year":"2013","journal-title":"J. Biol. Chem."},{"key":"ref_45","doi-asserted-by":"crossref","unstructured":"Palla, A.R., Hilgendorf, K.I., Yang, A.V., Kerr, J.P., Hinken, A.C., Demeter, J., Kraft, P., Mooney, N.A., Yucel, N., and Jackson, P.K. (2020). Ciliation of muscle stem cells is critical to maintain regenerative capacity and is lost during aging. bioRxiv.","DOI":"10.1101\/2020.03.20.000943"},{"key":"ref_46","doi-asserted-by":"crossref","first-page":"81","DOI":"10.4161\/bioa.29668","article-title":"Therapies for sarcopenia and regeneration of old skeletal muscles: More a case of old tissue architecture than old stem cells","volume":"4","author":"Grounds","year":"2014","journal-title":"Bioarchitecture"},{"key":"ref_47","doi-asserted-by":"crossref","first-page":"e13411","DOI":"10.1111\/acel.13411","article-title":"Human muscle stem cells are refractory to aging","volume":"20","author":"Novak","year":"2021","journal-title":"Aging Cell"},{"key":"ref_48","doi-asserted-by":"crossref","first-page":"16","DOI":"10.1186\/s13395-021-00265-6","article-title":"Origins, potency, and heterogeneity of skeletal muscle fibro-adipogenic progenitors-time for new definitions","volume":"11","author":"Contreras","year":"2021","journal-title":"Skelet. Muscle"},{"key":"ref_49","doi-asserted-by":"crossref","first-page":"153","DOI":"10.1038\/ncb2015","article-title":"Muscle injury activates resident fibro\/adipogenic progenitors that facilitate myogenesis","volume":"12","author":"Joe","year":"2010","journal-title":"Nat. Cell Biol."},{"key":"ref_50","doi-asserted-by":"crossref","first-page":"786","DOI":"10.1038\/nm.3869","article-title":"Nilotinib reduces muscle fibrosis in chronic muscle injury by promoting TNF-mediated apoptosis of fibro\/adipogenic progenitors","volume":"21","author":"Lemos","year":"2015","journal-title":"Nat. Med."},{"key":"ref_51","doi-asserted-by":"crossref","first-page":"143","DOI":"10.1038\/ncb2014","article-title":"Mesenchymal progenitors distinct from satellite cells contribute to ectopic fat cell formation in skeletal muscle","volume":"12","author":"Uezumi","year":"2010","journal-title":"Nat. Cell Biol."},{"key":"ref_52","doi-asserted-by":"crossref","first-page":"171","DOI":"10.1159\/000132462","article-title":"Non-random distribution and sensory functions of primary cilia in vascular smooth muscle cells","volume":"31","author":"Lu","year":"2008","journal-title":"Kidney Blood Press Res."},{"key":"ref_53","doi-asserted-by":"crossref","first-page":"e12086","DOI":"10.1002\/jev2.12086","article-title":"Ciliary extracellular vesicles are distinct from the cytosolic extracellular vesicles","volume":"10","author":"Mohieldin","year":"2021","journal-title":"J. Extracell. Vesicles"},{"key":"ref_54","doi-asserted-by":"crossref","first-page":"S78","DOI":"10.1016\/j.diff.2011.10.006","article-title":"Hedgehog signaling regulates myelination in the peripheral nervous system through primary cilia","volume":"83","author":"Yoshimura","year":"2012","journal-title":"Differentiation"},{"key":"ref_55","doi-asserted-by":"crossref","first-page":"jcs239871","DOI":"10.1242\/jcs.239871","article-title":"Ciliary proteins specify the cell inflammatory response by tuning NFkappaB signalling, independently of primary cilia","volume":"133","author":"Koneva","year":"2020","journal-title":"J. Cell Sci."},{"key":"ref_56","doi-asserted-by":"crossref","first-page":"3654","DOI":"10.1242\/jcs.086629","article-title":"Fibrosis and adipogenesis originate from a common mesenchymal progenitor in skeletal muscle","volume":"124","author":"Uezumi","year":"2011","journal-title":"J. Cell Sci."},{"key":"ref_57","doi-asserted-by":"crossref","first-page":"103549","DOI":"10.1016\/j.mcn.2020.103549","article-title":"Dystrophic Dmd(mdx) rats show early neuronal changes (increased S100beta and Tau5) at 8 months, supporting severe dystropathology in this rodent model of Duchenne muscular dystrophy","volume":"108","author":"Krishnan","year":"2020","journal-title":"Mol. Cell Neurosci."},{"key":"ref_58","doi-asserted-by":"crossref","first-page":"F1000","DOI":"10.12688\/f1000research.7247.1","article-title":"Axonal maintenance, glia, exosomes, and heat shock proteins","volume":"5","author":"Tytell","year":"2016","journal-title":"F1000Research"},{"key":"ref_59","doi-asserted-by":"crossref","first-page":"108","DOI":"10.1016\/j.ccell.2017.12.001","article-title":"Hedgehog pathway drives fusion-negative rhabdomyosarcoma initiated from non-myogenic endothelial progenitors","volume":"33","author":"Drummond","year":"2018","journal-title":"Cancer Cell"},{"key":"ref_60","doi-asserted-by":"crossref","first-page":"426","DOI":"10.1038\/nrc3961","article-title":"Probing for a deeper understanding of rhabdomyosarcoma: Insights from complementary model systems","volume":"15","author":"Kashi","year":"2015","journal-title":"Nat. Rev. Cancer"},{"key":"ref_61","doi-asserted-by":"crossref","first-page":"641","DOI":"10.1242\/dev.068353","article-title":"Myoblast fusion: Lessons from flies and mice","volume":"139","author":"Abmayr","year":"2012","journal-title":"Development"},{"key":"ref_62","doi-asserted-by":"crossref","first-page":"101","DOI":"10.1007\/978-1-4684-5865-7_13","article-title":"The proliferation and fusion of myoblasts in vivo","volume":"280","author":"Grounds","year":"1990","journal-title":"Adv. Exp. Med. Biol."},{"key":"ref_63","doi-asserted-by":"crossref","first-page":"3","DOI":"10.1186\/s13395-017-0149-3","article-title":"Myoblast fusion confusion: The resolution begins","volume":"8","author":"Sampath","year":"2018","journal-title":"Skelet. Muscle"},{"key":"ref_64","doi-asserted-by":"crossref","first-page":"323","DOI":"10.1126\/science.aam9361","article-title":"Control of muscle formation by the fusogenic micropeptide myomixer","volume":"356","author":"Bi","year":"2017","journal-title":"Science"},{"key":"ref_65","doi-asserted-by":"crossref","first-page":"15665","DOI":"10.1038\/ncomms15665","article-title":"Myomerger induces fusion of non-fusogenic cells and is required for skeletal muscle development","volume":"8","author":"Quinn","year":"2017","journal-title":"Nat. Commun."},{"key":"ref_66","doi-asserted-by":"crossref","first-page":"re2","DOI":"10.1126\/scisignal.2003832","article-title":"Signaling mechanisms in mammalian myoblast fusion","volume":"6","author":"Hindi","year":"2013","journal-title":"Sci. Signal."},{"key":"ref_67","doi-asserted-by":"crossref","first-page":"82","DOI":"10.1016\/j.yexcr.2004.02.012","article-title":"A novel role for non-muscle gamma-actin in skeletal muscle sarcomere assembly","volume":"297","author":"Lloyd","year":"2004","journal-title":"Exp. Cell Res."},{"key":"ref_68","doi-asserted-by":"crossref","first-page":"1048","DOI":"10.1038\/nature08895","article-title":"Functional genomic screen for modulators of ciliogenesis and cilium length","volume":"464","author":"Kim","year":"2010","journal-title":"Nature"},{"key":"ref_69","doi-asserted-by":"crossref","first-page":"e42144","DOI":"10.7554\/eLife.42144","article-title":"Muscle-specific stress fibers give rise to sarcomeres in cardiomyocytes","volume":"7","author":"Fenix","year":"2018","journal-title":"Elife."},{"key":"ref_70","doi-asserted-by":"crossref","first-page":"271","DOI":"10.1016\/j.cell.2013.09.027","article-title":"Remembrance of cilia past","volume":"155","author":"Hoerner","year":"2013","journal-title":"Cell"},{"key":"ref_71","unstructured":"Schiaffino, S., and Partridge, T. (2008). Complexity of extracellular matrix and skeletal muscle regeneration. Skeletal Muscle Repair and Regeneration, Springer."},{"key":"ref_72","doi-asserted-by":"crossref","unstructured":"Collins, I., and Wann, A.K.T. (2020). Regulation of the extracellular matrix by ciliary machinery. Cells, 9.","DOI":"10.3390\/cells9020278"},{"key":"ref_73","doi-asserted-by":"crossref","first-page":"652","DOI":"10.1016\/j.ceb.2012.06.002","article-title":"The extracellular matrix and ciliary signaling","volume":"24","author":"Golemis","year":"2012","journal-title":"Curr. Opin. Cell Biol."},{"key":"ref_74","doi-asserted-by":"crossref","first-page":"2111","DOI":"10.1101\/gad.1689908","article-title":"Laminin-511 is an epithelial message promoting dermal papilla development and function during early hair morphogenesis","volume":"22","author":"Gao","year":"2008","journal-title":"Genes Dev."},{"key":"ref_75","doi-asserted-by":"crossref","first-page":"8","DOI":"10.1016\/j.jbiosc.2017.07.012","article-title":"Type I collagen promotes primary cilia growth through down-regulating HDAC6-mediated autophagy in confluent mouse embryo fibroblast 3T3-L1 cells","volume":"125","author":"Xu","year":"2018","journal-title":"J. Biosci. Bioeng."},{"key":"ref_76","doi-asserted-by":"crossref","first-page":"78","DOI":"10.1111\/nyas.12463","article-title":"Function and regulation of primary cilia and intraflagellar transport proteins in the skeleton","volume":"1335","author":"Yuan","year":"2015","journal-title":"Ann. N. Y. Acad. Sci."},{"key":"ref_77","doi-asserted-by":"crossref","first-page":"731","DOI":"10.1016\/j.cub.2010.02.048","article-title":"Primary cilia regulate branching morphogenesis during mammary gland development","volume":"20","author":"McDermott","year":"2010","journal-title":"Curr. Biol."},{"key":"ref_78","doi-asserted-by":"crossref","first-page":"147","DOI":"10.1016\/j.ygcen.2008.07.003","article-title":"Molecular characterization of follicle stimulating hormone receptor (FSHR) gene in the Indian river buffalo (Bubalus bubalis)","volume":"158","author":"Minj","year":"2008","journal-title":"Gen. Comp. Endocrinol."},{"key":"ref_79","doi-asserted-by":"crossref","first-page":"1964","DOI":"10.1038\/ncomms2964","article-title":"Collagen VI regulates satellite cell self-renewal and muscle regeneration","volume":"4","author":"Urciuolo","year":"2013","journal-title":"Nat. Commun."},{"key":"ref_80","doi-asserted-by":"crossref","first-page":"897","DOI":"10.1038\/nm.4126","article-title":"Loss of fibronectin from the aged stem cell niche affects the regenerative capacity of skeletal muscle in mice","volume":"22","author":"Lukjanenko","year":"2016","journal-title":"Nat. Med."},{"key":"ref_81","doi-asserted-by":"crossref","first-page":"56","DOI":"10.1016\/j.biocel.2014.09.010","article-title":"The need to more precisely define aspects of skeletal muscle regeneration","volume":"56","author":"Grounds","year":"2014","journal-title":"Int. J. Biochem. Cell Biol."},{"key":"ref_82","doi-asserted-by":"crossref","first-page":"518","DOI":"10.1111\/acel.12578","article-title":"Aging of the skeletal muscle extracellular matrix drives a stem cell fibrogenic conversion","volume":"16","author":"Beezhold","year":"2017","journal-title":"Aging Cell"},{"key":"ref_83","doi-asserted-by":"crossref","first-page":"1734","DOI":"10.1172\/JCI24397","article-title":"Indian hedgehog stimulates periarticular chondrocyte differentiation to regulate growth plate length independently of PTHrP","volume":"115","author":"Kobayashi","year":"2005","journal-title":"J. Clin. Investig."},{"key":"ref_84","doi-asserted-by":"crossref","first-page":"234","DOI":"10.1016\/j.matbio.2006.12.003","article-title":"Articular cartilage and growth plate defects are associated with chondrocyte cytoskeletal abnormalities in Tg737orpk mice lacking the primary cilia protein polaris","volume":"26","author":"McGlashan","year":"2007","journal-title":"Matrix Biol."},{"key":"ref_85","doi-asserted-by":"crossref","first-page":"202","DOI":"10.1016\/j.ydbio.2007.02.003","article-title":"Development of the post-natal growth plate requires intraflagellar transport proteins","volume":"305","author":"Song","year":"2007","journal-title":"Dev. Biol."},{"key":"ref_86","doi-asserted-by":"crossref","unstructured":"Ali, S.A., Niu, B., Cheah, K.S.E., and Alman, B. (2019). Unique and overlapping GLI1 and GLI2 transcriptional targets in neoplastic chondrocytes. PLoS ONE, 14.","DOI":"10.1371\/journal.pone.0211333"},{"key":"ref_87","doi-asserted-by":"crossref","first-page":"2584","DOI":"10.1016\/j.celrep.2018.02.043","article-title":"CEP128 localizes to the subdistal appendages of the mother centriole and regulates TGF-beta\/BMP signaling at the primary cilium","volume":"22","author":"Monnich","year":"2018","journal-title":"Cell Rep."},{"key":"ref_88","doi-asserted-by":"crossref","first-page":"333","DOI":"10.33594\/000000223","article-title":"Matrix metalloproteinase 13 from satellite cells is required for efficient muscle growth and regeneration","volume":"54","author":"Smith","year":"2020","journal-title":"Cell Physiol. Biochem."},{"key":"ref_89","doi-asserted-by":"crossref","first-page":"91","DOI":"10.1016\/S0070-2153(01)51003-0","article-title":"Myofibroblasts: Molecular crossdressers","volume":"51","author":"Walker","year":"2001","journal-title":"Curr. Top Dev. Biol."},{"key":"ref_90","doi-asserted-by":"crossref","first-page":"609653","DOI":"10.3389\/fbioe.2020.609653","article-title":"Mechanical and physical regulation of fibroblast-myofibroblast transition: From cellular mechanoresponse to tissue pathology","volume":"8","author":"Kurniawan","year":"2020","journal-title":"Front. Bioeng. Biotechnol."},{"key":"ref_91","doi-asserted-by":"crossref","first-page":"56","DOI":"10.1016\/j.stem.2016.09.010","article-title":"Myogenic progenitor cells control extracellular matrix production by fibroblasts during skeletal muscle hypertrophy","volume":"20","author":"Fry","year":"2017","journal-title":"Cell Stem. Cell"},{"key":"ref_92","doi-asserted-by":"crossref","first-page":"354","DOI":"10.1242\/dmm.003194","article-title":"The ADPKD genes pkd1a\/b and pkd2 regulate extracellular matrix formation","volume":"3","author":"Mangos","year":"2010","journal-title":"Dis. Model Mech."},{"key":"ref_93","doi-asserted-by":"crossref","first-page":"323","DOI":"10.1007\/978-3-319-51436-9_12","article-title":"Inflammation and fibrosis in polycystic kidney disease","volume":"60","author":"Song","year":"2017","journal-title":"Results Probl. Cell Differ."},{"key":"ref_94","doi-asserted-by":"crossref","first-page":"681","DOI":"10.1056\/NEJM197303292881310","article-title":"Cilia and cystic fibrosis","volume":"288","author":"Waring","year":"1973","journal-title":"N. Engl. J. Med."},{"key":"ref_95","doi-asserted-by":"crossref","first-page":"29","DOI":"10.1007\/s11926-019-0822-0","article-title":"The primary cilium: Emerging role as a key player in fibrosis","volume":"21","author":"Teves","year":"2019","journal-title":"Curr. Rheumatol. Rep."},{"key":"ref_96","doi-asserted-by":"crossref","first-page":"L978","DOI":"10.1152\/ajplung.00184.2012","article-title":"Role of Sonic Hedgehog in idiopathic pulmonary fibrosis","volume":"303","author":"Bolanos","year":"2012","journal-title":"Am. J. Physiol. Lung Cell Mol. Physiol."},{"key":"ref_97","doi-asserted-by":"crossref","first-page":"2342","DOI":"10.1161\/CIRCULATIONAHA.117.028752","article-title":"Fibroblast primary cilia are required for cardiac fibrosis","volume":"139","author":"Villalobos","year":"2019","journal-title":"Circulation"}],"container-title":["International Journal of Molecular Sciences"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.mdpi.com\/1422-0067\/22\/17\/9605\/pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2025,10,11]],"date-time":"2025-10-11T06:56:25Z","timestamp":1760165785000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.mdpi.com\/1422-0067\/22\/17\/9605"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2021,9,4]]},"references-count":97,"journal-issue":{"issue":"17","published-online":{"date-parts":[[2021,9]]}},"alternative-id":["ijms22179605"],"URL":"https:\/\/doi.org\/10.3390\/ijms22179605","relation":{},"ISSN":["1422-0067"],"issn-type":[{"value":"1422-0067","type":"electronic"}],"subject":[],"published":{"date-parts":[[2021,9,4]]}}}