{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,3,18]],"date-time":"2026-03-18T09:19:53Z","timestamp":1773825593166,"version":"3.50.1"},"update-to":[{"DOI":"10.1371\/journal.pcbi.1009156","type":"new_version","label":"New version","source":"publisher","updated":{"date-parts":[[2022,2,25]],"date-time":"2022-02-25T00:00:00Z","timestamp":1645747200000}}],"reference-count":69,"publisher":"Public Library of Science (PLoS)","issue":"2","license":[{"start":{"date-parts":[[2022,2,14]],"date-time":"2022-02-14T00:00:00Z","timestamp":1644796800000},"content-version":"vor","delay-in-days":0,"URL":"http:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"name":"SURF Cooperative","award":["2019.001"],"award-info":[{"award-number":["2019.001"]}]},{"name":"French Embassy in the Netherlands","award":["Bourse d\u2019Excellence Descartes de stage"],"award-info":[{"award-number":["Bourse d\u2019Excellence Descartes de stage"]}]},{"name":"Human Frontiers Science Program grant","award":["RGP0053\/2020"],"award-info":[{"award-number":["RGP0053\/2020"]}]},{"DOI":"10.13039\/501100003246","name":"Nederlandse Organisatie voor Wetenschappelijk Onderzoek","doi-asserted-by":"publisher","award":["ENW-VICI 865.17.004"],"award-info":[{"award-number":["ENW-VICI 865.17.004"]}],"id":[{"id":"10.13039\/501100003246","id-type":"DOI","asserted-by":"publisher"}]}],"content-domain":{"domain":["www.ploscompbiol.org"],"crossmark-restriction":false},"short-container-title":["PLoS Comput Biol"],"abstract":"<jats:p>\n                    Lymphocytes have been described to perform different motility patterns such as Brownian random walks, persistent random walks, and L\u00e9vy walks. Depending on the conditions, such as confinement or the distribution of target cells, either Brownian or L\u00e9vy walks lead to more efficient interaction with the targets. The diversity of these motility patterns may be explained by an adaptive response to the surrounding extracellular matrix (ECM). Indeed, depending on the ECM composition, lymphocytes either display a floating motility without attaching to the ECM, or sliding and stepping motility with respectively continuous or discontinuous attachment to the ECM, or pivoting behaviour with sustained attachment to the ECM. Moreover, on the long term, lymphocytes either perform a persistent random walk or a Brownian-like movement depending on the ECM composition. How the ECM affects cell motility is still incompletely understood. Here, we integrate essential mechanistic details of the lymphocyte-matrix adhesions and lymphocyte intrinsic cytoskeletal induced cell propulsion into a Cellular Potts model (CPM). We show that the combination of\n                    <jats:italic>de novo<\/jats:italic>\n                    cell-matrix adhesion formation, adhesion growth and shrinkage, adhesion rupture, and feedback of adhesions onto cell propulsion recapitulates multiple lymphocyte behaviours, for different lymphocyte subsets and various substrates. With an increasing attachment area and increased adhesion strength, the cells\u2019 speed and persistence decreases. Additionally, the model predicts random walks with short-term persistent but long-term subdiffusive properties resulting in a pivoting type of motility. For small adhesion areas, the spatial distribution of adhesions emerges as a key factor influencing cell motility. Small adhesions at the front allow for more persistent motility than larger clusters at the back, despite a similar total adhesion area. In conclusion, we present an integrated framework to simulate the effects of ECM proteins on cell-matrix adhesion dynamics. The model reveals a sufficient set of principles explaining the plasticity of lymphocyte motility.\n                  <\/jats:p>","DOI":"10.1371\/journal.pcbi.1009156","type":"journal-article","created":{"date-parts":[[2022,2,14]],"date-time":"2022-02-14T14:02:13Z","timestamp":1644847333000},"page":"e1009156","update-policy":"https:\/\/doi.org\/10.1371\/journal.pcbi.corrections_policy","source":"Crossref","is-referenced-by-count":13,"title":["Computational modelling of cell motility modes emerging from cell-matrix adhesion dynamics"],"prefix":"10.1371","volume":"18","author":[{"ORCID":"https:\/\/orcid.org\/0000-0002-3282-5571","authenticated-orcid":true,"given":"Leonie","family":"van Steijn","sequence":"first","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-3362-5229","authenticated-orcid":true,"given":"Inge M. 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Tejedor","year":"2012","journal-title":"Phys Rev Lett"},{"issue":"43","key":"pcbi.1009156.ref004","doi-asserted-by":"crossref","first-page":"11350","DOI":"10.1073\/pnas.1711371114","article-title":"The topography of the environment alters the optimal search strategy for active particles","volume":"114","author":"G Volpe","year":"2017","journal-title":"P Natl Acad Sci USA"},{"issue":"1","key":"pcbi.1009156.ref005","doi-asserted-by":"crossref","DOI":"10.1529\/biophysj.107.117044","article-title":"Sampling the cell with anomalous diffusion\u2014The discovery of slowness","volume":"94","author":"G Guigas","year":"2008","journal-title":"Biophys J"},{"issue":"7","key":"pcbi.1009156.ref006","first-page":"1","article-title":"Heterogeneous, delayed-onset killing by multiple-hitting T cells: Stochastic simulations to assess methods for analysis of imaging data","volume":"16","author":"RJ Beck","year":"2020","journal-title":"PLOS Comput 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quantitative limits of intravital two-photon lymphocyte tracking","volume":"108","author":"J Textor","year":"2011","journal-title":"P Natl Acad Sci USA"},{"issue":"7404","key":"pcbi.1009156.ref011","doi-asserted-by":"crossref","first-page":"545","DOI":"10.1038\/nature11098","article-title":"Generalized L\u00e9vy walks and the role of chemokines in migration of effector CD8+ T cells","volume":"486","author":"TH Harris","year":"2012","journal-title":"Nature"},{"issue":"MAY","key":"pcbi.1009156.ref012","doi-asserted-by":"crossref","first-page":"1156","DOI":"10.3389\/fimmu.2018.01156","article-title":"Integrin \u03b21 optimizes diabetogenic T cell migration and function in the pancreas","volume":"9","author":"G Espinosa-Carrasco","year":"2018","journal-title":"Front Immunol"},{"issue":"070601","key":"pcbi.1009156.ref013","article-title":"Optimal Non-Markovian Search Strategies with n-Step Memory","volume":"127","author":"H Meyer","year":"2021","journal-title":"Phys Rev 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E"},{"key":"pcbi.1009156.ref026","first-page":"1","volume-title":"Mathematical Systems Theory in Biology, Communications, Computation, and Finance","author":"MS Alber","year":"2003"},{"issue":"5","key":"pcbi.1009156.ref027","doi-asserted-by":"crossref","first-page":"881","DOI":"10.1083\/jcb.200206043","article-title":"Recruitment of the Arp2\/3 complex to vinculin: coupling membrane protrusion to matrix adhesion","volume":"159","author":"KA DeMali","year":"2002","journal-title":"J Cell Biol"},{"issue":"3","key":"pcbi.1009156.ref028","doi-asserted-by":"crossref","first-page":"561","DOI":"10.1016\/j.cell.2006.12.039","article-title":"Lamellipodial Actin Mechanically Links Myosin Activity with Adhesion-Site Formation","volume":"128","author":"G Giannone","year":"2007","journal-title":"Cell"},{"issue":"1","key":"pcbi.1009156.ref029","doi-asserted-by":"crossref","first-page":"62","DOI":"10.1038\/sj.emboj.7601959","article-title":"Intermediate-affinity LFA-1 binds \u03b1-actinin-1 to control migration at the leading edge of the T cell","volume":"27","author":"P Stanley","year":"2008","journal-title":"EMBO J"},{"issue":"6","key":"pcbi.1009156.ref030","doi-asserted-by":"crossref","first-page":"061902","DOI":"10.1103\/PhysRevE.71.061902","article-title":"Effective adhesion strength of specifically bound vesicles","volume":"71","author":"AS Smith","year":"2005","journal-title":"Phys Rev E"},{"key":"pcbi.1009156.ref031","unstructured":"Eden M. A two-dimensional growth process. In: Berkeley Symposium on Mathematical Statistics and Probability; 1961. p. 223\u2013239."},{"key":"pcbi.1009156.ref032","doi-asserted-by":"crossref","unstructured":"Daub JT, Merks RMH. Cell-Based Computational Modeling of Vascular Morphogenesis Using Tissue Simulation Toolkit. In: Ribatti D, editor. Vascular Morphogenesis. vol. 1214 of Methods Mol. Biol. link.springer.com; 2014. p. 67\u2014127. Available from: http:\/\/dx.doi.org\/10.1007\/978-1-4939-1462-3_6.","DOI":"10.1007\/978-1-4939-1462-3_6"},{"key":"pcbi.1009156.ref033","doi-asserted-by":"crossref","first-page":"e61288","DOI":"10.7554\/eLife.61288","article-title":"Artistoo, a library to build, share, and explore simulations of cells and tissues in the web browser","volume":"10","author":"IM Wortel","year":"2021","journal-title":"eLife"},{"issue":"3","key":"pcbi.1009156.ref034","doi-asserted-by":"crossref","first-page":"244","DOI":"10.1007\/BF01328731","article-title":"Die Brownsche Bewegung bei Ber\u00fccksichtigung einer Persistenz der Bewegungsrichtung. Mit Anwendungen auf die Bewegung lebender Infusorien","volume":"2","author":"R F\u00fcrth","year":"1920","journal-title":"Z Phys"},{"issue":"2","key":"pcbi.1009156.ref035","doi-asserted-by":"crossref","first-page":"912","DOI":"10.1529\/biophysj.105.061150","article-title":"Cell motility as persistent random motion: Theories from experiments","volume":"89","author":"D Selmeczi","year":"2005","journal-title":"Biophys J"},{"issue":"2","key":"pcbi.1009156.ref036","doi-asserted-by":"crossref","first-page":"23","DOI":"10.1140\/epje\/i2017-11510-0","article-title":"Active Brownian particles moving in a random Lorentz gas","volume":"40","author":"M Zeitz","year":"2017","journal-title":"Eur Phys J E"},{"issue":"2","key":"pcbi.1009156.ref037","doi-asserted-by":"crossref","first-page":"459","DOI":"10.1073\/pnas.0707603105","article-title":"Anomalous dynamics of cell migration","volume":"105","author":"P Dieterich","year":"2008","journal-title":"P Natl Acad Sci USA"},{"issue":"16","key":"pcbi.1009156.ref038","doi-asserted-by":"crossref","first-page":"3866","DOI":"10.1021\/jp993491m","article-title":"Fractional kramers equation","volume":"104","author":"E Barkai","year":"2000","journal-title":"Journal of Physical Chemistry B"},{"issue":"2","key":"pcbi.1009156.ref039","article-title":"Fractional Brownian motion and motion governed by the fractional Langevin equation in confined geometries","volume":"81","author":"JH Jeon","year":"2010","journal-title":"Phys Rev E"},{"issue":"5908","key":"pcbi.1009156.ref040","doi-asserted-by":"crossref","first-page":"1687","DOI":"10.1126\/science.1163595","article-title":"Traction dynamics of filopodia on compliant substrates","volume":"322","author":"CE Chan","year":"2008","journal-title":"Science"},{"issue":"5808","key":"pcbi.1009156.ref041","doi-asserted-by":"crossref","first-page":"111","DOI":"10.1126\/science.1135085","article-title":"Differential transmission of actin motion within focal adhesions","volume":"315","author":"K Hu","year":"2007","journal-title":"Science"},{"issue":"2","key":"pcbi.1009156.ref042","doi-asserted-by":"crossref","first-page":"507","DOI":"10.1091\/mbc.e04-10-0860","article-title":"Slipping or Gripping? Fluorescent Speckle Microscopy in Fish Keratocytes Reveals Two Different Mechanisms for Generating a Retrograde Flow of Actin","volume":"16","author":"C Jurado","year":"2005","journal-title":"Mol Biol Cell"},{"issue":"2","key":"pcbi.1009156.ref043","doi-asserted-by":"crossref","first-page":"374","DOI":"10.1016\/j.cell.2015.01.056","article-title":"Actin flows mediate a universal coupling between cell speed and cell persistence","volume":"161","author":"P Maiuri","year":"2015","journal-title":"Cell"},{"issue":"6","key":"pcbi.1009156.ref044","doi-asserted-by":"crossref","first-page":"625","DOI":"10.1038\/ncb2747","article-title":"\u03b21- and \u03b1v-class integrins cooperate to regulate myosin II during rigidity sensing of fibronectin-based microenvironments","volume":"15","author":"HB Schiller","year":"2013","journal-title":"Nat Cell Biol"},{"issue":"31","key":"pcbi.1009156.ref045","doi-asserted-by":"crossref","first-page":"11549","DOI":"10.1073\/pnas.0601909103","article-title":"Cellular asymmetry and individuality in directional sensing","volume":"103","author":"A Samadani","year":"2006","journal-title":"P Natl Acad Sci USA"},{"issue":"3","key":"pcbi.1009156.ref046","doi-asserted-by":"crossref","first-page":"1442","DOI":"10.4049\/jimmunol.180.3.1442","article-title":"Th17 Cells Exhibit a Distinct Calcium Profile from Th1 and Th2 Cells and Have Th1-Like Motility and NF-AT Nuclear Localization","volume":"180","author":"KS Weber","year":"2008","journal-title":"J Immunol"},{"issue":"2","key":"pcbi.1009156.ref047","doi-asserted-by":"crossref","first-page":"298","DOI":"10.1016\/j.immuni.2019.06.026","article-title":"Programming of Distinct Chemokine-Dependent and -Independent Search Strategies for Th1 and Th2 Cells Optimizes Function at Inflamed Sites","volume":"51","author":"A Gaylo-Moynihan","year":"2019","journal-title":"Immunity"},{"issue":"6616","key":"pcbi.1009156.ref048","doi-asserted-by":"crossref","first-page":"537","DOI":"10.1038\/385537a0","article-title":"Integrin-ligand binding properties govern cell migration speed through cell-substratum adhesiveness","volume":"385","author":"SP Palecek","year":"1997","journal-title":"Nature"},{"issue":"9","key":"pcbi.1009156.ref049","doi-asserted-by":"crossref","first-page":"4329","DOI":"10.1091\/mbc.e05-02-0170","article-title":"Cell adhesion strengthening: Contributions of adhesive area, integrin binding, and focal adhesion assembly","volume":"16","author":"ND Gallant","year":"2005","journal-title":"Mol Biol Cell"},{"issue":"5","key":"pcbi.1009156.ref050","doi-asserted-by":"crossref","first-page":"1634","DOI":"10.1091\/mbc.e06-09-0777","article-title":"Revealing Early Steps of \u03b12 \u03b21 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"},{"issue":"3","key":"pcbi.1009156.ref051","first-page":"645","article-title":"Single integrin molecule adhesion forces in intact cells measured by atomic force microscopy","volume":"259","author":"PP Lehenkari","year":"1999","journal-title":"Biochem Biophys"},{"issue":"8","key":"pcbi.1009156.ref052","doi-asserted-by":"crossref","first-page":"1306","DOI":"10.1177\/153537020623100804","article-title":"Dynamic adhesion of T lymphocytes to endothelial cells revealed by atomic force microscopy","volume":"231","author":"X Zhang","year":"2006","journal-title":"Exp Biol Med"},{"issue":"35","key":"pcbi.1009156.ref053","doi-asserted-by":"crossref","DOI":"10.1039\/C7SM00439G","article-title":"Complex self-propelled rings: A minimal model for cell motility","volume":"13","author":"C Abaurrea Velasco","year":"2017","journal-title":"Soft Matter"},{"issue":"4","key":"pcbi.1009156.ref054","doi-asserted-by":"crossref","first-page":"821","DOI":"10.1016\/j.bpj.2014.11.3478","article-title":"Three-dimensional balance of cortical tension and axial contractility enables fast amoeboid migration","volume":"108","author":"B \u00c1lvarez-Gonz\u00e1lez","year":"2015","journal-title":"Biophys J"},{"issue":"10","key":"pcbi.1009156.ref055","doi-asserted-by":"crossref","first-page":"953","DOI":"10.1038\/ni.1936","article-title":"Confinement-optimized three-dimensional T cell amoeboid motility is modulated via myosin IIA-regulated adhesions","volume":"11","author":"J Jacobelli","year":"2010","journal-title":"Nat Immunol"},{"issue":"7544","key":"pcbi.1009156.ref056","doi-asserted-by":"crossref","first-page":"425","DOI":"10.1038\/nature14323","article-title":"MAP4K4 regulates integrin-FERM binding to control endothelial cell motility","volume":"519","author":"P Vitorino","year":"2015","journal-title":"Nature"},{"issue":"18","key":"pcbi.1009156.ref057","doi-asserted-by":"crossref","first-page":"6851","DOI":"10.1073\/pnas.1203252109","article-title":"Coupling actin flow, adhesion, and morphology in a computational cell motility model","volume":"109","author":"D Shao","year":"2012","journal-title":"P Natl Acad Sci USA"},{"issue":"3","key":"pcbi.1009156.ref058","doi-asserted-by":"crossref","first-page":"E390","DOI":"10.1073\/pnas.1717230115","article-title":"Computational modeling of three-dimensional ECM-rigidity sensing to guide directed cell migration","volume":"155","author":"MC Kim","year":"2018","journal-title":"P Natl Acad Sci USA"},{"issue":"8","key":"pcbi.1009156.ref059","doi-asserted-by":"crossref","first-page":"e1003774","DOI":"10.1371\/journal.pcbi.1003774","article-title":"Mechanical Cell-Matrix Feedback Explains Pairwise and Collective Endothelial Cell Behavior In Vitro","volume":"10","author":"RFM Van Oers","year":"2014","journal-title":"PLOS Comp Biol"},{"issue":"12","key":"pcbi.1009156.ref060","doi-asserted-by":"crossref","first-page":"e1007459","DOI":"10.1371\/journal.pcbi.1007459","article-title":"From energy to cellular forces in the Cellular Potts Model: An algorithmic approach","volume":"15","author":"EG Rens","year":"2019","journal-title":"PLOS Comp Biol"},{"issue":"4","key":"pcbi.1009156.ref061","doi-asserted-by":"crossref","first-page":"755","DOI":"10.1016\/j.bpj.2016.12.012","article-title":"Cell Contractility Facilitates Alignment of Cells and Tissues to Static Uniaxial Stretch","volume":"112","author":"EG Rens","year":"2017","journal-title":"Biophys J"},{"issue":"9","key":"pcbi.1009156.ref062","doi-asserted-by":"crossref","first-page":"101488","DOI":"10.1016\/j.isci.2020.101488","article-title":"Cell Shape and Durotaxis Explained from Cell-Extracellular Matrix Forces and Focal Adhesion Dynamics","volume":"23","author":"EG Rens","year":"2020","journal-title":"iScience"},{"issue":"2","key":"pcbi.1009156.ref063","doi-asserted-by":"crossref","first-page":"746","DOI":"10.1016\/S0006-3495(94)80535-1","article-title":"Self-assembly of membrane junctions","volume":"67","author":"R Bruinsma","year":"1994","journal-title":"Biophys J"},{"issue":"3","key":"pcbi.1009156.ref064","doi-asserted-by":"crossref","first-page":"262","DOI":"10.1002\/1439-7641(20020315)3:3<262::AID-CPHC262>3.0.CO;2-U","article-title":"Cell Adhesion as Wetting Transition?","volume":"3","author":"E Sackmann","year":"2002","journal-title":"ChemPhysChem"},{"issue":"15","key":"pcbi.1009156.ref065","doi-asserted-by":"crossref","first-page":"2841","DOI":"10.1039\/b822956b","article-title":"Physics puzzles on membrane domains posed by cell biology","volume":"5","author":"PF Lenne","year":"2009","journal-title":"Soft 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