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Competition between nonspecific repulsion and specific bonding.","volume":"45","author":"GI Bell","year":"1984","journal-title":"Biophys J"},{"key":"ref48","doi-asserted-by":"crossref","first-page":"1465","DOI":"10.1103\/PhysRevB.14.1465","article-title":"Microscopic fracture studies in the two-dimensional triangular lattice.","volume":"14","author":"WT Ashurst","year":"1976","journal-title":"Phys Rev B"},{"key":"ref49","doi-asserted-by":"crossref","first-page":"82","DOI":"10.1016\/S0927-0256(96)00064-X","article-title":"Spring network models in elasticity and fracture composites and polycrystals.","volume":"7","author":"M Ostoja-Starzewski","year":"1996","journal-title":"Computational Materials Science"},{"key":"ref50","doi-asserted-by":"crossref","first-page":"302","DOI":"10.1038\/sj.emboj.7600960","article-title":"Dynamic study of the transition from hyaluronan- to integrin-mediated adhesion in chondrocytes.","volume":"25","author":"M Cohen","year":"2006","journal-title":"Embo 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Biol"},{"key":"ref54","doi-asserted-by":"crossref","first-page":"883","DOI":"10.1126\/science.3277283","article-title":"Cortical flow in animal cells.","volume":"239","author":"D Bray","year":"1988","journal-title":"Science"},{"key":"ref55","doi-asserted-by":"crossref","first-page":"10259","DOI":"10.1073\/pnas.0510348103","article-title":"The consensus mechanics of cultured mammalian cells.","volume":"103","author":"BD Hoffman","year":"2006","journal-title":"Proc Natl Acad Sci U S A"},{"key":"ref56","doi-asserted-by":"crossref","first-page":"15848","DOI":"10.1021\/jp074514x","article-title":"The role of glycocalyx in nanocarrier-cell adhesion investigated using a thermodynamic model and monte carlo simulations.","volume":"111","author":"NJ Agrawal","year":"2007","journal-title":"J Phys Chem C"},{"key":"ref57","doi-asserted-by":"crossref","first-page":"3271","DOI":"10.1103\/PhysRevE.55.3271","article-title":"Deformation and failure in cellular materials.","volume":"55","author":"JC Ladd","year":"1997","journal-title":"Phys Rev E"},{"key":"ref58","doi-asserted-by":"crossref","first-page":"349","DOI":"10.1016\/S0378-4371(97)00158-1","article-title":"Elastic constants of cellular structures.","volume":"240","author":"JC Ladd","year":"1997","journal-title":"Physica A"},{"key":"ref59","doi-asserted-by":"crossref","first-page":"10988","DOI":"10.1074\/jbc.273.18.10988","article-title":"Force required to break alpha5beta1 integrin-fibronectin bonds in intact adherent cells is sensitive to integrin activation state.","volume":"273","author":"AJ Garcia","year":"1998","journal-title":"J Biol Chem"},{"key":"ref60","doi-asserted-by":"crossref","first-page":"785","DOI":"10.1091\/mbc.10.3.785","article-title":"Modulation of cell proliferation and differentiation through substrate-dependent changes in fibronectin conformation.","volume":"10","author":"AJ Garcia","year":"1999","journal-title":"Mol Biol Cell"},{"key":"ref61","doi-asserted-by":"crossref","first-page":"579","DOI":"10.1016\/j.ceb.2006.08.005","article-title":"Integrin structures and conformational signaling.","volume":"18","author":"BH Luo","year":"2006","journal-title":"Curr Opin Cell Biol"},{"key":"ref62","doi-asserted-by":"crossref","first-page":"27466","DOI":"10.1074\/jbc.M404354200","article-title":"Allosteric beta1 integrin antibodies that stabilize the low affinity state by preventing the swing-out of the hybrid domain.","volume":"279","author":"BH Luo","year":"2004","journal-title":"J Biol Chem"},{"key":"ref63","doi-asserted-by":"crossref","first-page":"412","DOI":"10.1038\/87569","article-title":"C-terminal opening mimics \u2018inside-out\u2019 activation of integrin alpha5beta1.","volume":"8","author":"J Takagi","year":"2001","journal-title":"Nat Struct Biol"},{"key":"ref64","doi-asserted-by":"crossref","first-page":"599","DOI":"10.1016\/S0092-8674(02)00935-2","article-title":"Global conformational 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Chem"},{"key":"ref68","doi-asserted-by":"crossref","first-page":"1252","DOI":"10.1016\/S0006-3495(03)74940-6","article-title":"Force measurements of the alpha5beta1 integrin-fibronectin interaction.","volume":"84","author":"F Li","year":"2003","journal-title":"Biophys J"},{"key":"ref69","doi-asserted-by":"crossref","first-page":"131","DOI":"10.1002\/cm.20029","article-title":"Dynamic change in the distribution of alpha5beta1 integrin on isolated ventral membrane: effect of divalent cation species.","volume":"59","author":"H Hirata","year":"2004","journal-title":"Cell Motil Cytoskeleton"},{"key":"ref70","doi-asserted-by":"crossref","first-page":"936","DOI":"10.1016\/S0006-3495(93)81456-5","article-title":"A theoretical analysis for the effect of focal contact formation on cell-substrate attachment strength.","volume":"64","author":"MD Ward","year":"1993","journal-title":"Biophys J"},{"key":"ref71","article-title":"Mechanics of motor proteins and the cytoskeleton","author":"J Howard","year":"2001"},{"key":"ref72","doi-asserted-by":"crossref","first-page":"1836","DOI":"10.1529\/biophysj.107.113605","article-title":"Life and times of a cellular bleb.","volume":"94","author":"GT Charras","year":"2008","journal-title":"Biophys J"},{"key":"ref73","doi-asserted-by":"crossref","first-page":"1338","DOI":"10.1529\/biophysj.103.037044","article-title":"Microviscoelasticity of the apical cell surface of human umbilical vein endothelial cells (HUVEC) within confluent monolayers.","volume":"87","author":"W Feneberg","year":"2004","journal-title":"Biophys J"},{"key":"ref74","doi-asserted-by":"crossref","first-page":"1139","DOI":"10.1126\/science.1116995","article-title":"Tissue cells feel and respond to the stiffness of their substrate.","volume":"310","author":"DE Discher","year":"2005","journal-title":"Science"},{"key":"ref75","doi-asserted-by":"crossref","first-page":"368","DOI":"10.1111\/j.2517-6161.1979.tb01091.x","article-title":"Test of randomness for spatial point patterns.","volume":"B41","author":"BD Ripley","year":"1979","journal-title":"J R Statist Soc"},{"key":"ref76","doi-asserted-by":"crossref","first-page":"172","DOI":"10.1111\/j.2517-6161.1977.tb01615.x","article-title":"Modelling Spatial Patterns.","volume":"B39","author":"BD Ripley","year":"1977","journal-title":"J R Statist Soc"},{"key":"ref77","first-page":"193","article-title":"Contribution to the discussion of Dr. Ripley&apos;s paper.","volume":"B39","author":"JE Besag","year":"1977","journal-title":"J R Statist Soc"},{"key":"ref78","doi-asserted-by":"crossref","first-page":"1634","DOI":"10.1091\/mbc.E06-09-0777","article-title":"Revealing early steps of alpha2beta1 integrin-mediated adhesion to collagen type I by using single-cell force spectroscopy.","volume":"18","author":"A Taubenberger","year":"2007","journal-title":"Mol Biol Cell"},{"key":"ref79","doi-asserted-by":"crossref","first-page":"398","DOI":"10.1021\/nl052168u","article-title":"Cellular unbinding forces of initial adhesion processes on nanopatterned surfaces probed with magnetic tweezers.","volume":"6","author":"N Walter","year":"2006","journal-title":"Nano Lett"},{"key":"ref80","doi-asserted-by":"crossref","first-page":"795","DOI":"10.1126\/science.1079441","article-title":"Activation of integrin alphaIIbbeta3 by modulation of transmembrane helix associations.","volume":"300","author":"R Li","year":"2003","journal-title":"Science"},{"key":"ref81","doi-asserted-by":"crossref","first-page":"7395","DOI":"10.1021\/bi9702187","article-title":"Integrin alphaIIb beta3 reconstituted into lipid bilayers is nonclustered in its activated state but clusters after fibrinogen binding.","volume":"36","author":"EM Erb","year":"1997","journal-title":"Biochemistry"},{"key":"ref82","doi-asserted-by":"crossref","first-page":"547","DOI":"10.1016\/j.ceb.2003.08.003","article-title":"Integrin avidity regulation: are changes in affinity and conformation underemphasized?","volume":"15","author":"CV Carman","year":"2003","journal-title":"Curr Opin Cell Biol"},{"key":"ref83","doi-asserted-by":"crossref","first-page":"339","DOI":"10.1023\/A:1011379725811","article-title":"MUC1, the renaissance molecule.","volume":"6","author":"SJ Gendler","year":"2001","journal-title":"J Mammary Gland Biol Neoplasia"},{"key":"ref84","doi-asserted-by":"crossref","first-page":"24","DOI":"10.1002\/cm.20041","article-title":"Effects of substrate stiffness on cell morphology, cytoskeletal structure, and adhesion.","volume":"60","author":"T Yeung","year":"2005","journal-title":"Cell Motil Cytoskeleton"},{"key":"ref85","doi-asserted-by":"crossref","first-page":"275","DOI":"10.1146\/annurev.bioeng.6.040803.140040","article-title":"Mechanotransduction at cell-matrix and cell-cell contacts.","volume":"6","author":"CS Chen","year":"2004","journal-title":"Annu Rev Biomed Eng"},{"key":"ref86","doi-asserted-by":"crossref","first-page":"3583","DOI":"10.1242\/jcs.114.20.3583","article-title":"Molecular complexity and dynamics of cell-matrix adhesions.","volume":"114","author":"E Zamir","year":"2001","journal-title":"J Cell Sci"},{"key":"ref87","article-title":"Interferometric fluorescent super-resolution microscopy resolves 3D cellular ultrastructure.","author":"G Shtengel","year":"2009","journal-title":"Proc Natl Acad Sci U S A"},{"key":"ref88","doi-asserted-by":"crossref","first-page":"359","DOI":"10.1016\/0014-4827(71)90420-4","article-title":"The locomotion of fibroblasts in culture. IV. Electron microscopy of the leading lamella.","volume":"67","author":"M Abercrombie","year":"1971","journal-title":"Exp Cell Res"},{"key":"ref89","doi-asserted-by":"crossref","first-page":"129","DOI":"10.1242\/jcs.21.1.129","article-title":"Cell-to-substrate contacts in living fibroblasts: an interference reflexion study with an evaluation of the technique.","volume":"21","author":"CS Izzard","year":"1976","journal-title":"J Cell Sci"},{"key":"ref90","doi-asserted-by":"crossref","first-page":"12293","DOI":"10.1021\/la701824q","article-title":"Adhesion of giant vesicles mediated by weak binding of sialyl-LewisX to E-selectin in the presence of repelling poly(ethylene glycol) molecules.","volume":"23","author":"BG Lorz","year":"2007","journal-title":"Langmuir"},{"key":"ref91","doi-asserted-by":"crossref","first-page":"2311","DOI":"10.1016\/S0006-3495(99)77070-0","article-title":"Adhesion-induced receptor segregation and adhesion plaque formation: A model membrane study.","volume":"77","author":"A Kloboucek","year":"1999","journal-title":"Biophys J"},{"key":"ref92","doi-asserted-by":"crossref","first-page":"2669","DOI":"10.1016\/S0006-3495(97)78910-0","article-title":"Energy of dissociation of lipid bilayer from the membrane skeleton of red blood cells.","volume":"72","author":"WC Hwang","year":"1997","journal-title":"Biophys J"},{"key":"ref93","first-page":"249","article-title":"What does a protein molecule look like.","volume":"6","author":"VN Morozov","year":"1990","journal-title":"Com Mol Cell Biophys"},{"key":"ref94","doi-asserted-by":"crossref","first-page":"4093","DOI":"10.1002\/j.1460-2075.1988.tb03303.x","article-title":"Electron microscopy and structural model of human fibronectin receptor.","volume":"7","author":"MV Nermut","year":"1988","journal-title":"Embo J"},{"key":"ref95","doi-asserted-by":"crossref","first-page":"5146","DOI":"10.1021\/bi992770x","article-title":"Statistical conformation of human plasma fibronectin.","volume":"39","author":"J Pelta","year":"2000","journal-title":"Biochemistry"},{"key":"ref96","doi-asserted-by":"crossref","first-page":"945","DOI":"10.1007\/s11538-005-9049-5","article-title":"Integrin activation\u2013the importance of a positive feedback.","volume":"68","author":"D Iber","year":"2006","journal-title":"Bull Math Biol"},{"key":"ref97","doi-asserted-by":"crossref","first-page":"100","DOI":"10.1016\/j.bbagen.2005.01.014","article-title":"Mobility of integrin alpha5beta1 measured on the isolated ventral membranes of human skin fibroblasts.","volume":"1723","author":"H Hirata","year":"2005","journal-title":"Biochim Biophys Acta"},{"key":"ref98","doi-asserted-by":"crossref","first-page":"5521","DOI":"10.1242\/jcs.01416","article-title":"Spatial mapping of integrin interactions and dynamics during cell migration by image correlation microscopy.","volume":"117","author":"PW Wiseman","year":"2004","journal-title":"J Cell Sci"},{"key":"ref99","doi-asserted-by":"crossref","first-page":"27119","DOI":"10.1074\/jbc.274.38.27119","article-title":"Quantitative relationship among integrin-ligand binding, adhesion, and signaling via focal adhesion kinase and extracellular signal-regulated kinase 2.","volume":"274","author":"AR Asthagiri","year":"1999","journal-title":"J Biol Chem"}],"container-title":["PLoS Computational Biology"],"original-title":[],"language":"en","link":[{"URL":"http:\/\/dx.plos.org\/10.1371\/journal.pcbi.1000604","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2025,2,13]],"date-time":"2025-02-13T19:30:04Z","timestamp":1739475004000},"score":1,"resource":{"primary":{"URL":"https:\/\/dx.plos.org\/10.1371\/journal.pcbi.1000604"}},"subtitle":[],"editor":[{"given":"Douglas","family":"Lauffenberger","sequence":"first","affiliation":[]}],"short-title":[],"issued":{"date-parts":[[2009,12,11]]},"references-count":99,"journal-issue":{"issue":"12","published-online":{"date-parts":[[2009,12,11]]}},"URL":"https:\/\/doi.org\/10.1371\/journal.pcbi.1000604","relation":{},"ISSN":["1553-7358"],"issn-type":[{"value":"1553-7358","type":"electronic"}],"subject":[],"published":{"date-parts":[[2009,12,11]]}}}