{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2025,8,26]],"date-time":"2025-08-26T08:40:02Z","timestamp":1756197602439,"version":"3.44.0"},"reference-count":83,"publisher":"The Company of Biologists","issue":"16","license":[{"start":{"date-parts":[[2025,8,26]],"date-time":"2025-08-26T00:00:00Z","timestamp":1756166400000},"content-version":"vor","delay-in-days":11,"URL":"http:\/\/creativecommons.org\/licenses\/by\/4.0"}],"funder":[{"DOI":"10.13039\/501100000024","name":"Canadian Institutes of Health Research","doi-asserted-by":"crossref","award":["PJT-15614"],"award-info":[{"award-number":["PJT-15614"]}],"id":[{"id":"10.13039\/501100000024","id-type":"DOI","asserted-by":"crossref"}]},{"DOI":"10.13039\/501100000038","name":"Natural Sciences and Engineering Research Council of Canada","doi-asserted-by":"crossref","award":["RGPIN-2017-06667"],"award-info":[{"award-number":["RGPIN-2017-06667"]}],"id":[{"id":"10.13039\/501100000038","id-type":"DOI","asserted-by":"crossref"}]},{"DOI":"10.13039\/501100003579","name":"University of Toronto","doi-asserted-by":"crossref","id":[{"id":"10.13039\/501100003579","id-type":"DOI","asserted-by":"crossref"}]}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":[],"published-print":{"date-parts":[[2025,8,15]]},"abstract":"<jats:title>ABSTRACT<\/jats:title>\n               <jats:p>Pericellular matrix-mediated cell-cell adhesion in Xenopus gastrula tissues is characterized by a spectrum of narrow and wide cell contacts that alternate with the non-adhesive surfaces of the interstitial space. Here we show, first, that knockdown of a pericellular matrix adhesion molecule, fibronectin, diminishes contact abundance, and hence cell-packing density, without reducing adhesion strength. Second, we find that cell surfaces in gastrula tissues exhibit solid-like behavior in the form of buckling and crumpling, shape modifications that are typically seen in thin elastic films. We propose that both phenomena are explained by generic properties of the pericellular matrix: its compression and consequent stiffening by the interpenetration of matrix layers during adhesive contact formation. We argue that this renders part of the cell surface non-adhesive to form interstitial gaps, and both gap surfaces and contacts prone to buckling and crumpling in line with cell contractility fluctuations. In this elasto-capillary model of tissue cohesion, the size of the interstitial space is determined by the abundance of the pericellular matrix, not by adhesion strength.<\/jats:p>","DOI":"10.1242\/dev.204663","type":"journal-article","created":{"date-parts":[[2025,8,7]],"date-time":"2025-08-07T12:52:33Z","timestamp":1754571153000},"source":"Crossref","is-referenced-by-count":0,"title":["Adhesion strength, cell packing density and cell surface buckling in pericellular matrix-mediated tissue cohesion"],"prefix":"10.1242","volume":"152","author":[{"ORCID":"https:\/\/orcid.org\/0000-0002-0628-0897","authenticated-orcid":false,"given":"Rudolf","family":"Winklbauer","sequence":"first","affiliation":[{"name":"University of Toronto Department of Cell and Systems Biology , , Toronto M5S 3G5 ,","place":["Canada"]}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Olivia","family":"Luu","sequence":"additional","affiliation":[{"name":"University of Toronto Department of Cell and Systems Biology , , Toronto M5S 3G5 ,","place":["Canada"]}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Debanjan","family":"Barua","sequence":"additional","affiliation":[{"name":"University of Toronto Department of Cell and Systems Biology , , Toronto M5S 3G5 ,","place":["Canada"]}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Martina","family":"Nagel","sequence":"additional","affiliation":[{"name":"University of Toronto Department of Cell and Systems Biology , , Toronto M5S 3G5 ,","place":["Canada"]}],"role":[{"role":"author","vocabulary":"crossref"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-6488-5088","authenticated-orcid":false,"given":"Yunyun","family":"Huang","sequence":"additional","affiliation":[{"name":"University of Toronto Department of Cell and Systems Biology , , Toronto M5S 3G5 ,","place":["Canada"]}],"role":[{"role":"author","vocabulary":"crossref"}]}],"member":"237","published-online":{"date-parts":[[2025,8,26]]},"reference":[{"key":"2025082604154291500_DEV204663C1","doi-asserted-by":"publisher","first-page":"20150520","DOI":"10.1098\/rstb.2015.0520","article-title":"Vertex models: from cell mechanics to tissue morphogenesis","volume":"372","author":"Alt","year":"2017","journal-title":"Philos. Trans. R. Soc. Lond. B Biol. Sci."},{"key":"2025082604154291500_DEV204663C2","doi-asserted-by":"publisher","first-page":"73","DOI":"10.1016\/j.ydbio.2022.07.007","article-title":"Eph\/ephrin signaling controls cell contacts and formation of a structurally asymmetrical tissue boundary in the Xenopus gastrula","volume":"490","author":"Barua","year":"2022","journal-title":"Dev. Biol."},{"key":"2025082604154291500_DEV204663C3","doi-asserted-by":"publisher","first-page":"923","DOI":"10.1016\/j.bpj.2017.06.063","article-title":"The mechanics of fluid-filled interstitial gaps. II. Gap characteristics in Xenopus embryonic ectoderm","volume":"113","author":"Barua","year":"2017","journal-title":"Biophys. J."},{"key":"2025082604154291500_DEV204663C4","doi-asserted-by":"publisher","first-page":"e2107953118","DOI":"10.1073\/pnas.2107953118","article-title":"Cell-cell contact landscapes in Xenopus gastrula tissues","volume":"118","author":"Barua","year":"2021","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"2025082604154291500_DEV204663C5","doi-asserted-by":"publisher","first-page":"125","DOI":"10.1002\/ar.1091940109","article-title":"Fine structural differentiation of germ layers in the mouse at the time of mesoderm formation","volume":"194","author":"Batten","year":"1979","journal-title":"Anat. Rec."},{"key":"2025082604154291500_DEV204663C6","doi-asserted-by":"publisher","first-page":"845","DOI":"10.1016\/S0021-9290(00)00011-7","article-title":"The mechanics of cell sorting and envelopment","volume":"33","author":"Brodland","year":"2000","journal-title":"J. Biomech."},{"key":"2025082604154291500_DEV204663C7","doi-asserted-by":"publisher","first-page":"252","DOI":"10.1038\/s41580-023-00688-7","article-title":"Adherens junctions as molecular regulators of emergent tissue mechanics","volume":"25","author":"Campas","year":"2024","journal-title":"Nat. Rev. Mol. Cell Biol."},{"key":"2025082604154291500_DEV204663C8","doi-asserted-by":"publisher","first-page":"2528","DOI":"10.1016\/j.bpj.2016.04.034","article-title":"Actomyosin cortical mechanical properties in nonadherent cells determined by atomic force microscopy","volume":"110","author":"Cartagena-Rivera","year":"2016","journal-title":"Biophys. J."},{"key":"2025082604154291500_DEV204663C9","doi-asserted-by":"publisher","first-page":"074302","DOI":"10.1103\/PhysRevLett.90.074302","article-title":"Geometry and physics of wrinkling","volume":"90","author":"Cerda","year":"2003","journal-title":"Phys. Rev. Lett."},{"key":"2025082604154291500_DEV204663C10","doi-asserted-by":"publisher","first-page":"671","DOI":"10.1098\/rspa.2004.1371","article-title":"Confined developable elastic surfaces: cylinders, cones and the Elastica","volume":"461","author":"Cerda","year":"2005","journal-title":"Proc. R. Soc. A"},{"key":"2025082604154291500_DEV204663C11","doi-asserted-by":"publisher","first-page":"3132","DOI":"10.1016\/j.cub.2017.09.005","article-title":"Viscoelastic dissipation stabilizes cell shape changes during tissue morphogenesis","volume":"27","author":"Clement","year":"2017","journal-title":"Curr. Biol."},{"key":"2025082604154291500_DEV204663C12","doi-asserted-by":"publisher","first-page":"2729","DOI":"10.1016\/j.cub.2017.08.002","article-title":"Inter-adipocyte adhesion and signaling by collagen IV intercellular concentrations in Drosophila","volume":"27","author":"Dai","year":"2017","journal-title":"Curr. Biol."},{"key":"2025082604154291500_DEV204663C13","doi-asserted-by":"publisher","first-page":"319","DOI":"10.1016\/j.ydbio.2008.06.018","article-title":"Limb bud and flank mesoderm have distinct \u201cphysical phenotypes\u201d that may contribute to limb budding","volume":"321","author":"Damon","year":"2008","journal-title":"Dev. Biol."},{"key":"2025082604154291500_DEV204663C14","doi-asserted-by":"publisher","first-page":"3672","DOI":"10.1242\/dev.104315","article-title":"Tissue cohesion and the mechanics of cell rearrangement","volume":"141","author":"David","year":"2014","journal-title":"Development"},{"key":"2025082604154291500_DEV204663C15","doi-asserted-by":"publisher","first-page":"833","DOI":"10.1016\/j.cub.2006.03.038","article-title":"Integrin a5b1 and fibronectin regulate polarized cell protrusions required for xenopus convergence and extension","volume":"16","author":"Davidson","year":"2006","journal-title":"Curr. Biol."},{"key":"2025082604154291500_DEV204663C16","doi-asserted-by":"publisher","first-page":"136","DOI":"10.1006\/jcis.1997.5214","article-title":"Axisymmetric Drop Shape Analysis: computational methods for the measurement of interfacial properties from the shape and dimensions of pendant and sessile drops","volume":"196","author":"Del Rio","year":"1997","journal-title":"J. Colloid. Interface Sci."},{"key":"2025082604154291500_DEV204663C17","doi-asserted-by":"publisher","first-page":"465","DOI":"10.1126\/science.aaw7709","article-title":"Hyrdaulic fracturing and active coarsening position the lumen of the mouse blastocyst","volume":"365","author":"Dumortier","year":"2019","journal-title":"Science"},{"key":"2025082604154291500_DEV204663C18","doi-asserted-by":"publisher","first-page":"151","DOI":"10.1016\/S0006-3495(89)82660-8","article-title":"Apparent viscosity and cortical tension of blood granulocytes determined by micropipet aspiration","volume":"56","author":"Evans","year":"1989","journal-title":"Biophys. J."},{"key":"2025082604154291500_DEV204663C19","first-page":"2638","article-title":"Mammary collective cell migration involves transient loss of epithelial features and individual cell migration within the epithelium","volume":"125","author":"Ewald","year":"2012","journal-title":"J. Cell Sci."},{"key":"2025082604154291500_DEV204663C20","doi-asserted-by":"publisher","first-page":"589","DOI":"10.1016\/j.bpj.2016.06.008","article-title":"Rheology of the active cell cortex in mitosis","volume":"111","author":"Fischer-Friedrich","year":"2016","journal-title":"Biophys. J."},{"key":"2025082604154291500_DEV204663C21","doi-asserted-by":"publisher","first-page":"757","DOI":"10.1091\/mbc.e12-06-0485","article-title":"Analysis of turnover dynamics of the submembranous actin cortex","volume":"24","author":"Fritzsche","year":"2013","journal-title":"Mol. Biol. Cell"},{"key":"2025082604154291500_DEV204663C22","doi-asserted-by":"publisher","first-page":"3653","DOI":"10.1039\/D4SM00235K","article-title":"Softness matters: effects of compression on the behavior of adsorbed microgels at interfaces","volume":"20","author":"Gerelli","year":"2024","journal-title":"Soft Mat."},{"key":"2025082604154291500_DEV204663C23","doi-asserted-by":"publisher","first-page":"531","DOI":"10.1016\/j.ceb.2011.06.002","article-title":"Dynamics of actomyosin contractile activity during epithelial morphogenesis","volume":"23","author":"Gorfinkiel","year":"2011","journal-title":"Curr. Opin. Cell Biol."},{"key":"2025082604154291500_DEV204663C24","first-page":"445","article-title":"Can surface adhesion drive cell-rearrangement? Part I: biological cell-sorting","volume":"164","author":"Graner","year":"1993","journal-title":"J. Theor. Biol."},{"key":"2025082604154291500_DEV204663C25","doi-asserted-by":"publisher","first-page":"2013","DOI":"10.1103\/PhysRevLett.69.2013","article-title":"Simulation of biological cell sorting using a two-dimensional extended Potts model","volume":"69","author":"Graner","year":"1992","journal-title":"Phys. Rev. Lett."},{"key":"2025082604154291500_DEV204663C26","doi-asserted-by":"publisher","first-page":"563","DOI":"10.1016\/0012-1606(70)90141-7","article-title":"Cytoplasmic microtubules and the mechanism of avian gastrulation","volume":"23","author":"Granholm","year":"1970","journal-title":"Dev. Biol."},{"key":"2025082604154291500_DEV204663C27","doi-asserted-by":"publisher","first-page":"502","DOI":"10.1038\/nrm2927","article-title":"Adherens junctions: from molecules to morphogenesis","volume":"11","author":"Harris","year":"2010","journal-title":"Nat. Rev. Mol. Cell Biol."},{"key":"2025082604154291500_DEV204663C28","doi-asserted-by":"publisher","first-page":"e65390","DOI":"10.7554\/eLife.65390","article-title":"Mechanical heterogeneity along single cell-cell junctions is driven by lateral clustering of cadherins during vertebrate axis elongation","volume":"10","author":"Huebner","year":"2021","journal-title":"eLife"},{"key":"2025082604154291500_DEV204663C29","doi-asserted-by":"publisher","first-page":"582","DOI":"10.1016\/j.devcel.2023.02.016","article-title":"A hydraulic feedback loop between mesendoderm cell migration and interstitial fluid relocalization promotes embryonic axis formation in zebrafish","volume":"58","author":"Huljev","year":"2023","journal-title":"Dev. Cell"},{"year":"1996","author":"Hutchinson","key":"2025082604154291500_DEV204663C30"},{"key":"2025082604154291500_DEV204663C31","doi-asserted-by":"publisher","first-page":"062711","DOI":"10.1103\/PhysRevE.92.062711","article-title":"Wrinkling of a spherical lipid interface induced by actomyosin cortex","volume":"92","author":"Ito","year":"2015","journal-title":"Phys. Rev. E"},{"key":"2025082604154291500_DEV204663C32","doi-asserted-by":"publisher","first-page":"165","DOI":"10.1016\/j.ydbio.2014.11.002","article-title":"Quantitative methods for analyzing cell-cell adhesion in development","volume":"401","author":"Kashef","year":"2015","journal-title":"Dev. Biol."},{"key":"2025082604154291500_DEV204663C33","doi-asserted-by":"publisher","first-page":"635","DOI":"10.1242\/jcs.067579","article-title":"Punctuated actin contractions during convergent extension and their permissive regulation by the con-canonical Wnt-signaling pathway","volume":"124","author":"Kim","year":"2011","journal-title":"J. Cell Sci."},{"key":"2025082604154291500_DEV204663C34","doi-asserted-by":"publisher","first-page":"859","DOI":"10.1038\/s41567-021-01215-1","article-title":"Embryonic tissues as active foams","volume":"17","author":"Kim","year":"2021","journal-title":"Nat. Phys."},{"key":"2025082604154291500_DEV204663C35","doi-asserted-by":"publisher","first-page":"9716","DOI":"10.1073\/pnas.1201201109","article-title":"Elastic sheet on a liquid drop reveals wrinkling and crumpling as distinct symmetry-breaking instabilities","volume":"109","author":"King","year":"2012","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"2025082604154291500_DEV204663C36","doi-asserted-by":"publisher","first-page":"R1045","DOI":"10.1016\/j.cub.2024.08.043","article-title":"Actomyosin clusters as active units shaping living matter","volume":"34","author":"Kruse","year":"2024","journal-title":"Curr. Biol."},{"key":"2025082604154291500_DEV204663C37","doi-asserted-by":"publisher","first-page":"9647","DOI":"10.1039\/C9SM01902B","article-title":"Actin shells control buckling and wrinkling of biomembranes","volume":"15","author":"Kusters","year":"2019","journal-title":"Soft Mat."},{"key":"2025082604154291500_DEV204663C38","doi-asserted-by":"publisher","first-page":"1376","DOI":"10.1039\/C4SM02271H","article-title":"Membrane rigidity induced by grafted polymer brush","volume":"11","author":"Lei","year":"2015","journal-title":"Soft Mat."},{"key":"2025082604154291500_DEV204663C39","doi-asserted-by":"publisher","first-page":"202","DOI":"10.1016\/j.devcel.2020.12.018","article-title":"Cell junction mechanics beyond the bounds of adhesion and tension","volume":"56","author":"Lenne","year":"2021","journal-title":"Dev. Cell"},{"key":"2025082604154291500_DEV204663C40","doi-asserted-by":"publisher","first-page":"409","DOI":"10.1113\/expphysiol.1987.sp003085","article-title":"Flow through interstitium and other fibrous matrices","volume":"72","author":"Levick","year":"1987","journal-title":"Q. J. Exp. Physiol."},{"key":"2025082604154291500_DEV204663C41","doi-asserted-by":"publisher","first-page":"046604","DOI":"10.1103\/PhysRevE.73.046604","article-title":"Spontaneous curvature cancellation in forces thin sheets","volume":"73","author":"Liang","year":"2006","journal-title":"Phys. Rev. E"},{"key":"2025082604154291500_DEV204663C42","doi-asserted-by":"publisher","first-page":"17","DOI":"10.1080\/00268949108035656","article-title":"Adhesion of vesicles and membranes","volume":"202","author":"Lipowsky","year":"1991","journal-title":"Mol. Cryst. Liq. Cryst."},{"key":"2025082604154291500_DEV204663C43","doi-asserted-by":"publisher","first-page":"e2309125121","DOI":"10.1073\/pnas.2309125121","article-title":"Self-assembled active actomyosin gels spontaneously curve and wrinkle similar to biological cells and tissues","volume":"121","author":"Livne","year":"2024","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"2025082604154291500_DEV204663C44","doi-asserted-by":"publisher","first-page":"4000","DOI":"10.1073\/pnas.1010331108","article-title":"Large-scale mechanical properties of Xenopus embryonic epithelium","volume":"108","author":"Luu","year":"2011","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"2025082604154291500_DEV204663C45","doi-asserted-by":"publisher","first-page":"e0297420","DOI":"10.1371\/journal.pone.0297420","article-title":"Cell contacts and pericellular matrix in the Xenopus gastrula chordamesoderm","volume":"19","author":"Luu","year":"2024","journal-title":"PLoS ONE"},{"key":"2025082604154291500_DEV204663C46","doi-asserted-by":"publisher","first-page":"253","DOI":"10.1126\/science.1225399","article-title":"Adhesion functions in cell sorting by mechanically coupling the cortices of adhering cells","volume":"338","author":"Ma\u00eetre","year":"2012","journal-title":"Science"},{"key":"2025082604154291500_DEV204663C47","doi-asserted-by":"publisher","first-page":"12517","DOI":"10.1073\/pnas.1003743107","article-title":"Coaction of intercellular adhesion and cortical tension specifies tissue surface tension","volume":"107","author":"Manning","year":"2010","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"2025082604154291500_DEV204663C48","doi-asserted-by":"publisher","first-page":"495","DOI":"10.1209\/0295-5075\/9\/5\/015","article-title":"Buckling of Langmuir monolayers","volume":"9","author":"Milner","year":"1989","journal-title":"Europhys. Lett."},{"key":"2025082604154291500_DEV204663C49","doi-asserted-by":"publisher","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":"2025082604154291500_DEV204663C50","doi-asserted-by":"publisher","first-page":"354","DOI":"10.1016\/j.devcel.2017.01.010","article-title":"The physical basis of coordinated tissue spreading in zebrafish gastrulation","volume":"40","author":"Morita","year":"2017","journal-title":"Dev. Cell"},{"key":"2025082604154291500_DEV204663C51","doi-asserted-by":"publisher","first-page":"492","DOI":"10.1038\/ncb1399","article-title":"Syndecan-4 regulates non-canonical Wnt signalling and is essential for convergent and extension movements in Xenopus embryos","volume":"8","author":"Mu\u00f1oz","year":"2006","journal-title":"Nat. Cell Biol."},{"key":"2025082604154291500_DEV204663C52","doi-asserted-by":"publisher","first-page":"1735","DOI":"10.1016\/j.bpj.2020.09.030","article-title":"Morphologies of vesicle doublets: competition among bending elasticity, surface tension, and adhesion","volume":"119","author":"Murakami","year":"2020","journal-title":"Biophys. J."},{"key":"2025082604154291500_DEV204663C53","doi-asserted-by":"publisher","first-page":"20820","DOI":"10.1073\/pnas.1214753109","article-title":"F-actin buckling coordinates contractility and severing in a biomimetic actomyosin cortex","volume":"109","author":"Murrell","year":"2012","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"2025082604154291500_DEV204663C54","doi-asserted-by":"publisher","first-page":"dev162651","DOI":"10.1242\/dev.162651","article-title":"PDGF-A suppresses contact inhibition during directional collective cell migration","volume":"145","author":"Nagel","year":"2018","journal-title":"Development"},{"key":"2025082604154291500_DEV204663C55","doi-asserted-by":"publisher","first-page":"1877","DOI":"10.1242\/jcs.095315","article-title":"Cadherin-dependent differential cell adhesion in Xenopus causes cell sorting in vitro but not in the embryo","volume":"125","author":"Ninomiya","year":"2012","journal-title":"J. Cell Sci."},{"key":"2025082604154291500_DEV204663C56","doi-asserted-by":"publisher","first-page":"913","DOI":"10.1016\/j.bpj.2017.06.062","article-title":"The mechanics of fluid-filled interstitial gaps. I. Modeling gaps in a compact tissue","volume":"113","author":"Parent","year":"2017","journal-title":"Biophys. J."},{"key":"2025082604154291500_DEV204663C57","doi-asserted-by":"publisher","first-page":"141","DOI":"10.1016\/j.devcel.2023.11.014","article-title":"Two-phase kinetics and cell cortex elastic behavior in Xenopus gastrula cell-cell adhesion","volume":"59","author":"Parent","year":"2024","journal-title":"Dev. Cell"},{"key":"2025082604154291500_DEV204663C58","doi-asserted-by":"publisher","first-page":"169","DOI":"10.1038\/s41556-018-0247-4","article-title":"Fluidization-mediated tissue spreading by mitotic cell rounding and non-canonical Wnt signalling","volume":"21","author":"Petridou","year":"2019","journal-title":"Nat. Cell Biol."},{"key":"2025082604154291500_DEV204663C59","doi-asserted-by":"publisher","first-page":"5344","DOI":"10.1073\/pnas.1813255116","article-title":"Theory of mechanochemical patterning in biphasic biological tissues","volume":"116","author":"Recho","year":"2019","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"2025082604154291500_DEV204663C60","doi-asserted-by":"publisher","first-page":"dev192773","DOI":"10.1242\/dev.192773","article-title":"Inferring cell junction tension and pressure from cell geometry","volume":"148","author":"Roffay","year":"2021","journal-title":"Development"},{"key":"2025082604154291500_DEV204663C61","doi-asserted-by":"publisher","first-page":"1644","DOI":"10.1039\/c3sm51910d","article-title":"Physics of cell adhesion: some lessons from cell-mimetic systems","volume":"10","author":"Sackmann","year":"2014","journal-title":"Soft Mat."},{"key":"2025082604154291500_DEV204663C62","doi-asserted-by":"publisher","first-page":"1421","DOI":"10.1016\/j.bpj.2016.02.013","article-title":"Determining physical properties of the cell cortex","volume":"110","author":"Saha","year":"2016","journal-title":"Biophys. J."},{"key":"2025082604154291500_DEV204663C63","doi-asserted-by":"publisher","first-page":"536","DOI":"10.1016\/j.tcb.2012.07.001","article-title":"Actin cortex mechanics and cellular morphogenesis","volume":"22","author":"Salbreux","year":"2012","journal-title":"Trends Cell Biol.."},{"key":"2025082604154291500_DEV204663C64","doi-asserted-by":"publisher","first-page":"1669","DOI":"10.1038\/s41556-024-01501-z","article-title":"Inverse blebs operate as hydraulic pumps during mouse blastocyst formation","volume":"26","author":"Schliffka","year":"2024","journal-title":"Nat. Cell Biol."},{"key":"2025082604154291500_DEV204663C65","doi-asserted-by":"publisher","first-page":"93","DOI":"10.1007\/BF00493158","article-title":"The use of tannic acid for the ultrastructual visualization of hyaluronic acid","volume":"65","author":"Singley","year":"1980","journal-title":"Histochemistry"},{"key":"2025082604154291500_DEV204663C66","doi-asserted-by":"publisher","first-page":"89","DOI":"10.1016\/S0014-5793(98)00570-5","article-title":"Adhesion-related glycocalyx study: quantitative approach with imaging-spectrum in the energy filtering transmission electron microscope (EFTEM)","volume":"429","author":"Soler","year":"1998","journal-title":"FEBS Lett."},{"key":"2025082604154291500_DEV204663C67","doi-asserted-by":"publisher","first-page":"1577","DOI":"10.1073\/pnas.48.9.1577","article-title":"On the mechanism of tissue reconstruction by dissociated cells. I. Population kinetics, differential adhesiveness, and the absence of directed migration","volume":"48","author":"Steinberg","year":"1962","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"2025082604154291500_DEV204663C68","doi-asserted-by":"publisher","first-page":"401","DOI":"10.1126\/science.141.3579.401","article-title":"Reconstruction of tissues by dissociated cells","volume":"141","author":"Steinberg","year":"1963","journal-title":"Science"},{"key":"2025082604154291500_DEV204663C69","doi-asserted-by":"crossref","first-page":"99","DOI":"10.1007\/978-1-4899-7124-1_3","article-title":"Specific cell ligands and the differential adhesion hypothesis: how do they fit together?","volume-title":"Specificity of Embryological Interactions (Receptors and Recognition, Series B)","author":"Steinberg","year":"1978"},{"key":"2025082604154291500_DEV204663C70","doi-asserted-by":"publisher","first-page":"91","DOI":"10.1038\/35040042","article-title":"Cadherins in embryonic and neural morphogenesis","volume":"1","author":"Tepass","year":"2000","journal-title":"Nat. Rev. Mol. Cell Biol."},{"key":"2025082604154291500_DEV204663C71","first-page":"558","article-title":"An analysis of the condensation process during chondrogenesis in the embryonic chick hind limb","volume":"33","author":"Thorogood","year":"1975","journal-title":"J. Embryol. Exp. Morph."},{"key":"2025082604154291500_DEV204663C72","doi-asserted-by":"publisher","first-page":"021008","DOI":"10.1103\/PhysRevX.10.021008","article-title":"Crumples as a generic stress-focusing instability in confined sheets","volume":"10","author":"Timounay","year":"2020","journal-title":"Phys. Rev."},{"key":"2025082604154291500_DEV204663C73","doi-asserted-by":"publisher","first-page":"105981","DOI":"10.1016\/j.nbd.2022.105981","article-title":"Local diffusion in the extracellular space of the brain","volume":"177","author":"Tonnesen","year":"2023","journal-title":"Neurobiol. Dis."},{"key":"2025082604154291500_DEV204663C74","doi-asserted-by":"publisher","first-page":"374","DOI":"10.1016\/j.tcb.2022.08.007","article-title":"Adherens junction: the ensemble of specialized cadherin clusters","volume":"33","author":"Troyanovsky","year":"2023","journal-title":"Trends Cell Biol."},{"key":"2025082604154291500_DEV204663C75","doi-asserted-by":"publisher","first-page":"121","DOI":"10.1146\/annurev.bioeng.9.060906.151959","article-title":"The structure and function of the endothelial glycocalyx layer","volume":"9","author":"Weinbaum","year":"2007","journal-title":"Annu. Rev. Biomed. Eng."},{"key":"2025082604154291500_DEV204663C76","doi-asserted-by":"publisher","first-page":"335","DOI":"10.1002\/(SICI)1097-0177(199807)212:3&lt;335::AID-AJA1&gt;3.0.CO;2-I","article-title":"Conditions for fibronectin fibril formation in the early Xenopus embryo","volume":"212","author":"Winklbauer","year":"1998","journal-title":"Dev. Dyn."},{"key":"2025082604154291500_DEV204663C77","doi-asserted-by":"publisher","first-page":"3687","DOI":"10.1242\/jcs.174623","article-title":"Cell adhesion strength from cortical tension: an integration of concepts","volume":"128","author":"Winklbauer","year":"2015","journal-title":"J. Cell Sci."},{"key":"2025082604154291500_DEV204663C78","doi-asserted-by":"publisher","first-page":"jcs231597","DOI":"10.1242\/jcs.231597","article-title":"Dynamic cell-cell adhesion mediated by pericellular matrix interaction \u2013 a hypothesis","volume":"132","author":"Winklbauer","year":"2019","journal-title":"J. Cell Sci."},{"key":"2025082604154291500_DEV204663C79","doi-asserted-by":"publisher","first-page":"643","DOI":"10.1103\/RevModPhys.79.643","article-title":"Stress focusing in elastic sheets","volume":"79","author":"Witten","year":"2007","journal-title":"Rev. Mod. Phys."},{"key":"2025082604154291500_DEV204663C80","doi-asserted-by":"publisher","first-page":"e60","DOI":"10.1002\/mdp2.60","article-title":"Determination of residual stress and interface adhesion toughness of thin films by blisters","volume":"1","author":"Yuan","year":"2019","journal-title":"Mat. Design Process Comm."},{"key":"2025082604154291500_DEV204663C81","doi-asserted-by":"publisher","first-page":"187","DOI":"10.1016\/j.ydbio.2016.05.025","article-title":"Syndecan4 coordinates Wnt\/JNK and BMP signaling to regulate foregut progenitor development","volume":"416","author":"Zhang","year":"2016","journal-title":"Dev. Biol."},{"key":"2025082604154291500_DEV204663C82","doi-asserted-by":"publisher","first-page":"761","DOI":"10.1073\/pnas.0607633104","article-title":"Flat and sigmoidally curved contact zones in vesicle-vesicle adhesion","volume":"104","author":"Ziherl","year":"2007","journal-title":"Proc. natl. Acad. Sci. USA"},{"key":"2025082604154291500_DEV204663C83","doi-asserted-by":"publisher","first-page":"1937","DOI":"10.1039\/b802733a","article-title":"Membrane elasticity molds aggregates of simple cells","volume":"4","author":"Ziherl","year":"2008","journal-title":"Soft Mat."}],"container-title":["Development"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/journals.biologists.com\/dev\/article-pdf\/doi\/10.1242\/dev.204663\/3660593\/dev204663.pdf","content-type":"application\/pdf","content-version":"am","intended-application":"syndication"},{"URL":"https:\/\/journals.biologists.com\/dev\/article-pdf\/doi\/10.1242\/dev.204663\/3662402\/dev204663.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"syndication"},{"URL":"https:\/\/journals.biologists.com\/dev\/article-pdf\/doi\/10.1242\/dev.204663\/3662402\/dev204663.pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2025,8,26]],"date-time":"2025-08-26T08:15:48Z","timestamp":1756196148000},"score":1,"resource":{"primary":{"URL":"https:\/\/journals.biologists.com\/dev\/article\/152\/16\/dev204663\/368975\/Adhesion-strength-cell-packing-density-and-cell"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2025,8,15]]},"references-count":83,"journal-issue":{"issue":"16","published-print":{"date-parts":[[2025,8,15]]}},"URL":"https:\/\/doi.org\/10.1242\/dev.204663","relation":{},"ISSN":["0950-1991","1477-9129"],"issn-type":[{"type":"print","value":"0950-1991"},{"type":"electronic","value":"1477-9129"}],"subject":[],"published":{"date-parts":[[2025,8,15]]},"article-number":"dev204663"}}