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Fibronectin matrix assembly starts as cells emerge from the primitive streak and ingress in the unsegmented presomitic mesoderm (PSM). PSM cells undergo cyclic waves of segmentation clock gene expression, followed by Notch-dependent upregulation of meso1 in the rostral PSM which induces somite cleft formation. However, the relevance of the fibronectin matrix for these molecular processes remains unknown. Here, we assessed the role of the PSM fibronectin matrix in the spatio-temporal regulation of chick embryo somitogenesis by perturbing (1) extracellular fibronectin matrix assembly, (2) integrin\u2013fibronectin binding, (3) Rho-associated protein kinase (ROCK) activity and (4) non-muscle myosin II (NM II) function. We found that integrin\u2013fibronectin engagement and NM II activity are required for cell polarization in the nascent somite. All treatments resulted in defective somitic clefts and significantly perturbed meso1 and segmentation clock gene expression in the PSM. Importantly, inhibition of actomyosin-mediated contractility increased the period of hairy1\/hes4 oscillations from 90 to 120 min. Together, our work strongly suggests that the fibronectin\u2013integrin\u2013ROCK\u2013NM II axis regulates segmentation clock dynamics and dictates the spatio-temporal localization of somitic clefts.<\/jats:p>","DOI":"10.3390\/cells11132003","type":"journal-article","created":{"date-parts":[[2022,6,22]],"date-time":"2022-06-22T21:31:06Z","timestamp":1655933466000},"page":"2003","update-policy":"https:\/\/doi.org\/10.3390\/mdpi_crossmark_policy","source":"Crossref","is-referenced-by-count":1,"title":["Cell\u2013Fibronectin Interactions and Actomyosin Contractility Regulate the Segmentation Clock and Spatio-Temporal Somite Cleft Formation during Chick Embryo Somitogenesis"],"prefix":"10.3390","volume":"11","author":[{"ORCID":"https:\/\/orcid.org\/0000-0003-2369-2874","authenticated-orcid":false,"given":"Patr\u00edcia","family":"Gomes de Almeida","sequence":"first","affiliation":[{"name":"cE3c\u2014CHANGE, Departmento de Biologia Animal, Faculdade de Ci\u00eancias, Universidade de Lisboa, 1740-016 Lisboa, Portugal"},{"name":"ABC-RI, Algarve Biomedical Center Research Institute, 8005-139 Faro, Portugal"},{"name":"Faculdade de Medicina e Ci\u00eancias Biom\u00e9dicas (FMCB), Universidade do Algarve, Campus de Gambelas, 8005-139 Faro, Portugal"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-9963-3291","authenticated-orcid":false,"given":"Pedro","family":"Rifes","sequence":"additional","affiliation":[{"name":"cE3c\u2014CHANGE, Departmento de Biologia Animal, Faculdade de Ci\u00eancias, Universidade de Lisboa, 1740-016 Lisboa, Portugal"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-1598-6397","authenticated-orcid":false,"given":"Ana P.","family":"Martins-Jesus","sequence":"additional","affiliation":[{"name":"ABC-RI, Algarve Biomedical Center Research Institute, 8005-139 Faro, Portugal"},{"name":"Faculdade de Medicina e Ci\u00eancias Biom\u00e9dicas (FMCB), Universidade do Algarve, Campus de Gambelas, 8005-139 Faro, Portugal"}]},{"given":"Gon\u00e7alo G.","family":"Pinheiro","sequence":"additional","affiliation":[{"name":"cE3c\u2014CHANGE, Departmento de Biologia Animal, Faculdade de Ci\u00eancias, Universidade de Lisboa, 1740-016 Lisboa, Portugal"},{"name":"ABC-RI, Algarve Biomedical Center Research Institute, 8005-139 Faro, Portugal"},{"name":"Faculdade de Medicina e Ci\u00eancias Biom\u00e9dicas (FMCB), Universidade do Algarve, Campus de Gambelas, 8005-139 Faro, Portugal"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-0397-5917","authenticated-orcid":false,"given":"Raquel P.","family":"Andrade","sequence":"additional","affiliation":[{"name":"ABC-RI, Algarve Biomedical Center Research Institute, 8005-139 Faro, Portugal"},{"name":"Faculdade de Medicina e Ci\u00eancias Biom\u00e9dicas (FMCB), Universidade do Algarve, Campus de Gambelas, 8005-139 Faro, Portugal"},{"name":"Champalimaud Research Program, Champalimaud Center for the Unknown, 1400-038 Lisboa, Portugal"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-9018-3431","authenticated-orcid":false,"given":"S\u00f3lveig","family":"Thorsteinsd\u00f3ttir","sequence":"additional","affiliation":[{"name":"cE3c\u2014CHANGE, Departmento de Biologia Animal, Faculdade de Ci\u00eancias, Universidade de Lisboa, 1740-016 Lisboa, Portugal"}]}],"member":"1968","published-online":{"date-parts":[[2022,6,22]]},"reference":[{"key":"ref_1","doi-asserted-by":"crossref","first-page":"a005041","DOI":"10.1101\/cshperspect.a005041","article-title":"Fibronectins, their fibrillogenesis, and in vivo functions","volume":"3","author":"Schwarzbauer","year":"2011","journal-title":"Cold Spring Harb. Perspect. Biol."},{"key":"ref_2","doi-asserted-by":"crossref","first-page":"245","DOI":"10.1016\/bs.ctdb.2018.03.006","article-title":"Extracellular Matrix (ECM) and the Sculpting of Embryonic Tissues","volume":"130","author":"Dzamba","year":"2018","journal-title":"Curr. Top. Dev. Biol."},{"key":"ref_3","doi-asserted-by":"crossref","first-page":"1079","DOI":"10.1242\/dev.119.4.1079","article-title":"Defects in mesoderm, neural tube and vascular development in mouse embryos lacking fibronectin","volume":"119","author":"George","year":"1993","journal-title":"Development"},{"key":"ref_4","doi-asserted-by":"crossref","first-page":"833","DOI":"10.1016\/j.cub.2006.03.038","article-title":"Integrin \u03b15\u03b21 and Fibronectin Regulate Polarized Cell Protrusions Required for Xenopus Convergence and Extension","volume":"16","author":"Davidson","year":"2006","journal-title":"Curr. Biol."},{"key":"ref_5","doi-asserted-by":"crossref","first-page":"3635","DOI":"10.1242\/dev.128.18.3635","article-title":"Regulation of cell polarity, radial intercalation and epiboly in Xenopus: Novel roles for integrin and fibronectin","volume":"128","author":"Marsden","year":"2001","journal-title":"Development"},{"key":"ref_6","doi-asserted-by":"crossref","first-page":"3155","DOI":"10.1242\/dev.003665","article-title":"Redefining the role of ectoderm in somitogenesis: A player in the formation of the fibronectin matrix of presomitic mesoderm","volume":"134","author":"Rifes","year":"2007","journal-title":"Development"},{"key":"ref_7","doi-asserted-by":"crossref","first-page":"677","DOI":"10.1242\/dev.026211","article-title":"Actomyosin stiffens the vertebrate embryo during crucial stages of elongation and neural tube closure","volume":"136","author":"Zhou","year":"2009","journal-title":"Development"},{"key":"ref_8","doi-asserted-by":"crossref","first-page":"2830","DOI":"10.1111\/febs.14818","article-title":"The extracellular matrix as a multitasking player in disease","volume":"286","author":"Theocharis","year":"2019","journal-title":"FEBS J."},{"key":"ref_9","doi-asserted-by":"crossref","first-page":"389","DOI":"10.1016\/j.matbio.2005.06.008","article-title":"Fibronectin fibrillogenesis, a cell-mediated matrix assembly process","volume":"24","author":"Mao","year":"2005","journal-title":"Matrix Biol."},{"key":"ref_10","doi-asserted-by":"crossref","first-page":"397","DOI":"10.1146\/annurev-cellbio-100109-104020","article-title":"Assembly of fibronectin extracellular matrix","volume":"26","author":"Singh","year":"2010","journal-title":"Annu. Rev. Cell Dev. Biol."},{"key":"ref_11","doi-asserted-by":"crossref","unstructured":"Dalton, C.J., and Lemmon, C.A. (2021). Fibronectin: Molecular structure, fibrillar structure and mechanochemical signaling. Cells, 10.","DOI":"10.20944\/preprints202108.0010.v1"},{"key":"ref_12","doi-asserted-by":"crossref","first-page":"a004994","DOI":"10.1101\/cshperspect.a004994","article-title":"Integrin structure, activation, and interactions","volume":"3","author":"Campbell","year":"2011","journal-title":"Cold Spring Harb. Perspect. Biol."},{"key":"ref_13","doi-asserted-by":"crossref","first-page":"447","DOI":"10.1016\/j.devcel.2013.02.012","article-title":"Dynamic regulation of the structure and functions of integrin adhesions","volume":"24","author":"Wolfenson","year":"2013","journal-title":"Dev. Cell"},{"key":"ref_14","doi-asserted-by":"crossref","first-page":"4159","DOI":"10.1242\/jcs.192054","article-title":"The integrin adhesome network at a glance","volume":"129","author":"Horton","year":"2016","journal-title":"J. Cell Sci."},{"key":"ref_15","doi-asserted-by":"crossref","first-page":"2209","DOI":"10.1242\/jcs.170068","article-title":"The contractome\u2014A systems view of actomyosin contractility in non-muscle cells","volume":"128","author":"Zhenhuan","year":"2015","journal-title":"J. Cell Sci."},{"key":"ref_16","doi-asserted-by":"crossref","first-page":"373","DOI":"10.1242\/jcs.111559","article-title":"Cadherin adhesome at a glance","volume":"126","year":"2013","journal-title":"J. Cell Sci."},{"key":"ref_17","doi-asserted-by":"crossref","first-page":"628","DOI":"10.1016\/j.ceb.2012.07.003","article-title":"Spatial segregation between cell-cell and cell-matrix adhesions","volume":"24","author":"Burute","year":"2012","journal-title":"Curr. Opin. Cell Biol."},{"key":"ref_18","doi-asserted-by":"crossref","first-page":"1093","DOI":"10.1242\/jcs.183699","article-title":"The mechanical regulation of integrin-cadherin crosstalk organizes cells, signaling and forces","volume":"129","author":"Mui","year":"2016","journal-title":"J. Cell Sci."},{"key":"ref_19","doi-asserted-by":"crossref","first-page":"585","DOI":"10.1146\/annurev-physiol-021317-121245","article-title":"Steps in Mechanotransduction Pathways that Control Cell Morphology","volume":"81","author":"Wolfenson","year":"2019","journal-title":"Annu. Rev. Physiol."},{"key":"ref_20","doi-asserted-by":"crossref","unstructured":"Bailey, C., and Dale, K. (2015). Somitogenesis in Vertebrate Development. eLS, 1\u201315.","DOI":"10.1002\/9780470015902.a0003820.pub2"},{"key":"ref_21","doi-asserted-by":"crossref","first-page":"2382","DOI":"10.1002\/dvdy.21189","article-title":"Amniote somite derivatives","volume":"236","author":"Christ","year":"2007","journal-title":"Dev. Dyn."},{"key":"ref_22","doi-asserted-by":"crossref","first-page":"1595","DOI":"10.1126\/science.1133141","article-title":"A complex oscillating network of signaling genes underlies the mouse segmentation clock","volume":"314","author":"Glynn","year":"2006","journal-title":"Science"},{"key":"ref_23","doi-asserted-by":"crossref","first-page":"1313","DOI":"10.1073\/pnas.0508658103","article-title":"Real-time imaging of the somite segmentation clock: Revelation of unstable oscillators in the individual presomitic mesoderm cells","volume":"103","author":"Masamizu","year":"2006","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_24","doi-asserted-by":"crossref","first-page":"639","DOI":"10.1016\/S0092-8674(00)80451-1","article-title":"Avian hairy gene expression identifies a molecular clock linked to vertebrate segmentation and somitogenesis","volume":"91","author":"Palmeirim","year":"1997","journal-title":"Cell"},{"key":"ref_25","doi-asserted-by":"crossref","first-page":"354","DOI":"10.1038\/nature03591","article-title":"The Mesp2 transcription factor establishes segmental borders by suppressing Notch activity","volume":"435","author":"Morimoto","year":"2005","journal-title":"Nature"},{"key":"ref_26","doi-asserted-by":"crossref","first-page":"390","DOI":"10.1038\/78062","article-title":"Mesp2 initiates somite segmentation through the Notch signalling pathway","volume":"25","author":"Takahashi","year":"2000","journal-title":"Nat. Genet."},{"key":"ref_27","doi-asserted-by":"crossref","first-page":"a008201","DOI":"10.1101\/cshperspect.a008201","article-title":"The synchrony and cyclicity of developmental events","volume":"4","author":"Saga","year":"2012","journal-title":"Cold Spring Harb. Perspect. Biol."},{"key":"ref_28","doi-asserted-by":"crossref","first-page":"1571","DOI":"10.1016\/j.cub.2003.08.030","article-title":"Eph\/Ephrin signaling regulates the mesenchymalto-epithelial transition of the paraxial mesoderm during somite morphogenesis","volume":"13","author":"Barrios","year":"2003","journal-title":"Curr. Biol."},{"key":"ref_29","doi-asserted-by":"crossref","first-page":"2517","DOI":"10.1242\/dev.02422","article-title":"Identification of Epha4 enhancer required for segmental expression and the regulation by Mesp2","volume":"133","author":"Nakajima","year":"2006","journal-title":"Development"},{"key":"ref_30","doi-asserted-by":"crossref","first-page":"7467","DOI":"10.1073\/pnas.0902859106","article-title":"EphrinB2 coordinates the formation of a morphological boundary and cell epithelialization during somite segmentation","volume":"106","author":"Watanabe","year":"2009","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_31","doi-asserted-by":"crossref","unstructured":"Martins, G.G., Rifes, P., Ama\u00e2ndio, R., Rodrigues, G., Palmeirim, I., and Thorsteinsd\u00f3ttir, S. (2009). Dynamic 3D cell rearrangements guided by a fibronectin matrix underlie somitogenesis. PLoS ONE, 4.","DOI":"10.1371\/journal.pone.0007429"},{"key":"ref_32","doi-asserted-by":"crossref","first-page":"33","DOI":"10.1016\/j.devcel.2015.05.005","article-title":"Cross-Scale Integrin Regulation Organizes ECM and Tissue Topology","volume":"34","author":"Cobb","year":"2015","journal-title":"Dev. Cell"},{"key":"ref_33","doi-asserted-by":"crossref","first-page":"145","DOI":"10.1002\/(SICI)1097-0177(199610)207:2<145::AID-AJA3>3.0.CO;2-H","article-title":"Mesodermal development in mouse embryos mutant for fibronectin","volume":"207","author":"George","year":"1996","journal-title":"Dev. Dyn."},{"key":"ref_34","doi-asserted-by":"crossref","first-page":"4309","DOI":"10.1242\/dev.124.21.4309","article-title":"Mesodermal defects and cranial neural crest apoptosis in \u03b15 integrin-null embryos","volume":"124","author":"Goh","year":"1997","journal-title":"Development"},{"key":"ref_35","doi-asserted-by":"crossref","first-page":"587","DOI":"10.1016\/j.devcel.2005.03.006","article-title":"Integrin\u03b15-dependent fibronectin accumulation for maintenance of somite boundaries in zebrafish embryos","volume":"8","author":"Koshida","year":"2005","journal-title":"Dev. Cell"},{"key":"ref_36","doi-asserted-by":"crossref","first-page":"2713","DOI":"10.1002\/dvdy.21280","article-title":"Integrin \u03b15 is required for somite rotation and boundary formation in Xenopus","volume":"236","author":"Kragtorp","year":"2007","journal-title":"Dev. Dyn."},{"key":"ref_37","doi-asserted-by":"crossref","first-page":"370","DOI":"10.1016\/j.ydbio.2012.06.003","article-title":"Extracellular matrix assembly and 3D organization during paraxial mesoderm development in the chick embryo","volume":"368","author":"Rifes","year":"2012","journal-title":"Dev. Biol."},{"key":"ref_38","doi-asserted-by":"crossref","first-page":"248","DOI":"10.1038\/nature09151","article-title":"A random cell motility gradient downstream of FGF controls elongation of an amniote embryo","volume":"466","author":"Francois","year":"2010","journal-title":"Nature"},{"key":"ref_39","doi-asserted-by":"crossref","first-page":"573","DOI":"10.1242\/dev.090381","article-title":"Regulated tissue fluidity steers zebrafish body elongation","volume":"140","author":"Lawton","year":"2013","journal-title":"Development"},{"key":"ref_40","doi-asserted-by":"crossref","first-page":"401","DOI":"10.1038\/s41586-018-0479-2","article-title":"A fluid-to-solid jamming transition underlies vertebrate body axis elongation","volume":"561","author":"Mongera","year":"2018","journal-title":"Nature"},{"key":"ref_41","doi-asserted-by":"crossref","first-page":"524","DOI":"10.1016\/j.actbio.2019.07.056","article-title":"Micro-indentation and optical coherence tomography for the mechanical characterization of embryos: Experimental setup and measurements on chicken embryos","volume":"97","author":"Marrese","year":"2019","journal-title":"Acta Biomater."},{"key":"ref_42","doi-asserted-by":"crossref","first-page":"12269","DOI":"10.1096\/fj.202000896R","article-title":"In vivo characterization of chick embryo mesoderm by optical coherence tomography-assisted microindentation","volume":"34","author":"Marrese","year":"2020","journal-title":"FASEB J."},{"key":"ref_43","doi-asserted-by":"crossref","first-page":"2913","DOI":"10.1242\/dev.038935","article-title":"Control of extracellular matrix assembly along tissue boundaries via integrin and Eph\/Ephrin signaling","volume":"136","author":"Mould","year":"2009","journal-title":"Development"},{"key":"ref_44","doi-asserted-by":"crossref","first-page":"516","DOI":"10.1038\/s41586-022-04646-9","article-title":"Left\u2013right symmetry of zebrafish embryos requires somite surface tension","volume":"605","author":"Naganathan","year":"2022","journal-title":"Nature"},{"key":"ref_45","doi-asserted-by":"crossref","first-page":"231","DOI":"10.1002\/aja.1001950404","article-title":"A series of normal stages in the development of the chick embryo","volume":"195","author":"Hamburger","year":"1992","journal-title":"Dev. Dyn."},{"key":"ref_46","doi-asserted-by":"crossref","first-page":"619","DOI":"10.1016\/S1534-5807(01)00082-X","article-title":"A nomenclature for prospective somites and phases of cyclic gene expression in the presomitic mesoderm","volume":"1","author":"Tam","year":"2001","journal-title":"Dev. Cell"},{"key":"ref_47","doi-asserted-by":"crossref","first-page":"364","DOI":"10.1083\/jcb.100.2.364","article-title":"Interaction of the 70,000-mol-wt amino-terminal fragment of fibronectin with the matrix-assembly receptor of fibroblasts","volume":"100","author":"Mosher","year":"1985","journal-title":"J. Cell Biol."},{"key":"ref_48","doi-asserted-by":"crossref","first-page":"281","DOI":"10.1242\/dev.141259","article-title":"Basal filopodia and vascular mechanical stress organize fibronectin into pillars bridging the mesoderm-endoderm gap","volume":"144","author":"Sato","year":"2017","journal-title":"Development"},{"key":"ref_49","doi-asserted-by":"crossref","first-page":"2452","DOI":"10.1242\/jcs.033001","article-title":"Binding of soluble fibronectin to integrin \u03b15\u03b21\u2014Link to focal adhesion redistribution and contractile shape","volume":"121","author":"Huveneers","year":"2008","journal-title":"J. Cell Sci."},{"key":"ref_50","doi-asserted-by":"crossref","first-page":"30","DOI":"10.1038\/309030a0","article-title":"Cell attachment activity of fibronectin can be duplicated by small synthetic fragments of the molecule","volume":"309","author":"Pierschbacher","year":"1984","journal-title":"Nature"},{"key":"ref_51","doi-asserted-by":"crossref","first-page":"385","DOI":"10.1016\/j.chembiol.2005.01.015","article-title":"Screening for cell migration inhibitors via automated microscopy reveals a Rho-kinase inhibitor","volume":"12","author":"Yarrow","year":"2005","journal-title":"Chem. Biol."},{"key":"ref_52","doi-asserted-by":"crossref","first-page":"1743","DOI":"10.1126\/science.1081412","article-title":"Dissecting temporal and spatial control of cytokinesis with a myosin II inhibitor","volume":"299","author":"Straight","year":"2003","journal-title":"Science"},{"key":"ref_53","doi-asserted-by":"crossref","first-page":"284","DOI":"10.1002\/1097-0177(20010301)220:3<284::AID-DVDY1102>3.0.CO;2-5","article-title":"Improved method for chick whole-embryo culture using a filter paper carrier","volume":"220","author":"Chapman","year":"2001","journal-title":"Dev. Dyn."},{"key":"ref_54","doi-asserted-by":"crossref","first-page":"335","DOI":"10.1046\/j.1440-169X.1999.413437.x","article-title":"Efficient targeting of gene expression in chick embryos by microelectroporation","volume":"41","author":"Momose","year":"1999","journal-title":"Dev. Growth Differ."},{"key":"ref_55","doi-asserted-by":"crossref","first-page":"1071","DOI":"10.1083\/jcb.200205014","article-title":"The fibronectin-binding integrins \u03b15\u03b21 and \u03b1v\u03b23 differentially modulate RhoA-GTP loading, organization of cell matrix adhesions, and fibronectin fibrillogenesis","volume":"159","author":"Danen","year":"2002","journal-title":"J. Cell Biol."},{"key":"ref_56","doi-asserted-by":"crossref","first-page":"520","DOI":"10.1002\/dvdy.24391","article-title":"Fibronectin assembly during early embryo development: A versatile communication system between cells and tissues","volume":"245","author":"Pinheiro","year":"2016","journal-title":"Dev. Dyn."},{"key":"ref_57","first-page":"e52769","article-title":"Application of the DNA-specific stain methyl green in the fluorescent labeling of embryos","volume":"2015","author":"Prieto","year":"2015","journal-title":"J. Vis. Exp."},{"key":"ref_58","doi-asserted-by":"crossref","first-page":"787","DOI":"10.1038\/375787a0","article-title":"Expression of a Delta homologue in prospective neurons in the chick","volume":"375","author":"Henrique","year":"1995","journal-title":"Nature"},{"key":"ref_59","doi-asserted-by":"crossref","first-page":"201","DOI":"10.1006\/dbio.1998.8919","article-title":"cMeso-1, a novel bHLH transcription factor, is involved in somite formation in chicken embryos","volume":"199","author":"Buchberger","year":"1998","journal-title":"Dev. Biol."},{"key":"ref_60","doi-asserted-by":"crossref","first-page":"1421","DOI":"10.1242\/dev.127.7.1421","article-title":"Notch signalling is required for cyclic expression of the hairy-like gene HES1 in the presomitic mesoderm","volume":"127","author":"Jouve","year":"2000","journal-title":"Development"},{"key":"ref_61","doi-asserted-by":"crossref","first-page":"1463","DOI":"10.1093\/bioinformatics\/btp184","article-title":"Globally optimal stitching of tiled 3D microscopic image acquisitions","volume":"25","author":"Preibisch","year":"2009","journal-title":"Bioinformatics"},{"key":"ref_62","doi-asserted-by":"crossref","first-page":"6","DOI":"10.1016\/1047-8477(91)90025-R","article-title":"Image averaging of flexible fibrous macromolecules: The clathrin triskelion has an elastic proximal segment","volume":"107","author":"Kocsis","year":"1991","journal-title":"J. Struct. Biol."},{"key":"ref_63","doi-asserted-by":"crossref","first-page":"418","DOI":"10.1016\/0012-1606(91)90092-H","article-title":"Integrins play an essential role in somite adhesion to the embryonic axis","volume":"143","author":"Drake","year":"1991","journal-title":"Dev. Biol."},{"key":"ref_64","doi-asserted-by":"crossref","first-page":"327","DOI":"10.1016\/0012-1606(92)90288-R","article-title":"Perturbation of beta 1 integrin-mediated adhesions results in altered somite cell shape and behavior","volume":"149","author":"Drake","year":"1992","journal-title":"Dev. Biol."},{"key":"ref_65","doi-asserted-by":"crossref","unstructured":"Gir\u00f3s, A., Grgur, K., Gossler, A., and Costell, M. (2011). \u03b15\u03b21 integrin-mediated adhesion to fibronectin is required for axis elongation and somitogenesis in mice. PLoS ONE, 6.","DOI":"10.1371\/journal.pone.0022002"},{"key":"ref_66","doi-asserted-by":"crossref","first-page":"167","DOI":"10.1083\/jcb.200703021","article-title":"The RGD motif in fibronectin is essential for development but dispensable for fibril assembly","volume":"178","author":"Takahashi","year":"2007","journal-title":"J. Cell Biol."},{"key":"ref_67","doi-asserted-by":"crossref","first-page":"219","DOI":"10.1016\/S0092-8674(01)00437-8","article-title":"FGF signaling controls somite boundary position and regulates segmentation clock control of spatiotemporal Hox gene activation","volume":"106","author":"Dubrulle","year":"2001","journal-title":"Cell"},{"key":"ref_68","doi-asserted-by":"crossref","first-page":"3633","DOI":"10.1242\/dev.129.15.3633","article-title":"Morphological boundary forms by a novel inductive event mediated by Lunatic fringe and Notch during somitic segmentation","volume":"129","author":"Sato","year":"2002","journal-title":"Development"},{"key":"ref_69","doi-asserted-by":"crossref","first-page":"77","DOI":"10.1002\/(SICI)1520-6408(1998)23:1<77::AID-DVG8>3.0.CO;2-3","article-title":"Uncoupling segmentation and somitogenesis in the chick presomitic mesoderm","volume":"23","author":"Palmeirim","year":"1998","journal-title":"Dev. Genet."},{"key":"ref_70","first-page":"1989","article-title":"Mesodermal defect in late phase of gastrulation by a targeted mutation of focal adhesion kinase, FAK","volume":"11","author":"Furuta","year":"1995","journal-title":"Oncogene"},{"key":"ref_71","doi-asserted-by":"crossref","first-page":"685","DOI":"10.1242\/dev.02230","article-title":"Regulation of somitogenesis by Ena\/VASP proteins and FAK during Xenopus development","volume":"133","author":"Kragtorp","year":"2006","journal-title":"Development"},{"key":"ref_72","doi-asserted-by":"crossref","first-page":"474","DOI":"10.1006\/dbio.2001.0467","article-title":"Roles for zebrafish focal adhesion kinase in notochord and somite morphogenesis","volume":"240","author":"Henry","year":"2001","journal-title":"Dev. Biol."},{"key":"ref_73","first-page":"1495","article-title":"Non-Muscle myosin II in disease: Mechanisms and therapeutic opportunities","volume":"8","author":"Horwitz","year":"2015","journal-title":"DMM Dis. Model. Mech."},{"key":"ref_74","doi-asserted-by":"crossref","first-page":"150","DOI":"10.1016\/j.tcb.2018.09.006","article-title":"Principles of Actomyosin Regulation In Vivo","volume":"29","author":"Agarwal","year":"2019","journal-title":"Trends Cell Biol."},{"key":"ref_75","doi-asserted-by":"crossref","first-page":"435","DOI":"10.1016\/0040-8166(82)90038-6","article-title":"Calcium dependence and contraction in somite formation","volume":"14","author":"Chernoff","year":"1982","journal-title":"Tissue Cell"},{"key":"ref_76","doi-asserted-by":"crossref","first-page":"175","DOI":"10.1016\/j.cub.2014.11.043","article-title":"Rac1-dependent phosphorylation and focal adhesion recruitment of myosin IIA regulates migration and mechanosensing","volume":"25","author":"Pasapera","year":"2015","journal-title":"Curr. Biol."},{"key":"ref_77","doi-asserted-by":"crossref","first-page":"425","DOI":"10.1016\/j.devcel.2004.08.003","article-title":"Mesenchymal-epithelial transition during somitic segmentation is regulated by differential roles of Cdc42 and Rac1","volume":"7","author":"Nakaya","year":"2004","journal-title":"Dev. Cell"},{"key":"ref_78","doi-asserted-by":"crossref","first-page":"3591","DOI":"10.1242\/dev.050070","article-title":"Cell-autonomous integrin control of Wnt and Notch signalling during somitogenesis","volume":"137","author":"Rallis","year":"2010","journal-title":"Development"},{"key":"ref_79","doi-asserted-by":"crossref","first-page":"729","DOI":"10.1016\/j.devcel.2015.05.004","article-title":"Mechanical Allostery: Evidence for a Force Requirement in the Proteolytic Activation of Notch","volume":"33","author":"Gordon","year":"2015","journal-title":"Dev. Cell"},{"key":"ref_80","doi-asserted-by":"crossref","first-page":"1320","DOI":"10.1126\/science.aaf9739","article-title":"Notch-Jagged complex structure implicates a catch bond in tuning ligand sensitivity","volume":"355","author":"Luca","year":"2017","journal-title":"Science"},{"key":"ref_81","doi-asserted-by":"crossref","first-page":"1299","DOI":"10.1016\/j.devcel.2012.04.005","article-title":"Notch Ligand Endocytosis Generates Mechanical Pulling Force Dependent on Dynamin, Epsins, and Actin","volume":"22","author":"Shergill","year":"2012","journal-title":"Dev. Cell"},{"key":"ref_82","doi-asserted-by":"crossref","first-page":"579073","DOI":"10.3389\/fcell.2020.579073","article-title":"Protrusion-Mediated Signaling Regulates Patterning of the Developing Nervous System","volume":"8","author":"Moore","year":"2020","journal-title":"Front. Cell Dev. Biol."},{"key":"ref_83","doi-asserted-by":"crossref","first-page":"jcs250738","DOI":"10.1242\/jcs.250738","article-title":"Notch in mechanotransduction\u2014From molecular mechanosensitivity to tissue mechanostasis","volume":"133","author":"Stassen","year":"2021","journal-title":"J. Cell Sci."},{"key":"ref_84","doi-asserted-by":"crossref","first-page":"78","DOI":"10.1016\/j.devcel.2010.06.006","article-title":"Dynamic Filopodia Transmit Intermittent Delta-Notch Signaling to Drive Pattern Refinement during Lateral Inhibition","volume":"19","author":"Cohen","year":"2010","journal-title":"Dev. Cell"},{"key":"ref_85","doi-asserted-by":"crossref","unstructured":"Hunter, G.L., He, L., Perrimon, N., Charras, G., Giniger, E., and Baum, B. (2019). A role for actomyosin contractility in Notch signaling. BMC Biol., 17.","DOI":"10.1186\/s12915-019-0625-9"},{"key":"ref_86","doi-asserted-by":"crossref","unstructured":"Hardy, K.M., Yatskievych, T.A., Konieczka, J., Bobbs, A.S., and Antin, P.B. (2011). FGF signalling through RAS\/MAPK and PI3K pathways regulates cell movement and gene expression in the chicken primitive streak without affecting E-cadherin expression. BMC Dev. Biol., 11.","DOI":"10.1186\/1471-213X-11-20"},{"key":"ref_87","doi-asserted-by":"crossref","first-page":"3521","DOI":"10.1242\/dev.023416","article-title":"PDGF signalling controls the migration of mesoderm cells during chick gastrulation by regulating N-cadherin expression","volume":"135","author":"Yang","year":"2008","journal-title":"Development"},{"key":"ref_88","doi-asserted-by":"crossref","first-page":"719","DOI":"10.1038\/nature04861","article-title":"Noise-resistant and synchronized oscillation of the segmentation clock","volume":"441","author":"Horikawa","year":"2006","journal-title":"Nature"},{"key":"ref_89","doi-asserted-by":"crossref","first-page":"11861","DOI":"10.1038\/ncomms11861","article-title":"Faster embryonic segmentation through elevated Delta-Notch signalling","volume":"7","author":"Liao","year":"2016","journal-title":"Nat. Commun."},{"key":"ref_90","doi-asserted-by":"crossref","first-page":"1244","DOI":"10.1016\/j.cub.2010.06.034","article-title":"Intercellular coupling regulates the period of the segmentation clock","volume":"20","author":"Herrgen","year":"2010","journal-title":"Curr. Biol."},{"key":"ref_91","doi-asserted-by":"crossref","unstructured":"Tenin, G., Wright, D., Ferjentsik, Z., Bone, R., McGrew, M.J., and Maroto, M. (2010). The chick somitogenesis oscillator is arrested before all paraxial mesoderm is segmented into somites. BMC Dev. Biol., 10.","DOI":"10.1186\/1471-213X-10-24"},{"key":"ref_92","doi-asserted-by":"crossref","first-page":"303","DOI":"10.1016\/j.jmb.2007.01.089","article-title":"A Molecular Clock Operates During Chick Autopod Proximal-distal Outgrowth","volume":"368","author":"Pascoal","year":"2007","journal-title":"J. Mol. Biol."},{"key":"ref_93","doi-asserted-by":"crossref","first-page":"124","DOI":"10.1038\/s41586-020-2144-9","article-title":"Recapitulating the human segmentation clock with pluripotent stem cells","volume":"580","author":"Matsuda","year":"2020","journal-title":"Nature"},{"key":"ref_94","doi-asserted-by":"crossref","first-page":"eaba7667","DOI":"10.1126\/science.aba7667","article-title":"Species-specific pace of development is associated with differences in protein stability","volume":"369","author":"Rayon","year":"2020","journal-title":"Science"},{"key":"ref_95","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1016\/j.ceb.2020.08.007","article-title":"Mechanosensing in embryogenesis","volume":"68","author":"Agarwal","year":"2021","journal-title":"Curr. Opin. Cell Biol."},{"key":"ref_96","doi-asserted-by":"crossref","first-page":"dev193268","DOI":"10.1242\/dev.193268","article-title":"Building bridges between fields: Bringing together development and homeostasis","volume":"148","author":"Weterings","year":"2021","journal-title":"Development"}],"container-title":["Cells"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.mdpi.com\/2073-4409\/11\/13\/2003\/pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2025,10,10]],"date-time":"2025-10-10T23:38:01Z","timestamp":1760139481000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.mdpi.com\/2073-4409\/11\/13\/2003"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2022,6,22]]},"references-count":96,"journal-issue":{"issue":"13","published-online":{"date-parts":[[2022,7]]}},"alternative-id":["cells11132003"],"URL":"https:\/\/doi.org\/10.3390\/cells11132003","relation":{"has-preprint":[{"id-type":"doi","id":"10.1101\/808121","asserted-by":"object"}]},"ISSN":["2073-4409"],"issn-type":[{"value":"2073-4409","type":"electronic"}],"subject":[],"published":{"date-parts":[[2022,6,22]]}}}