{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,8,19]],"date-time":"2026-08-19T17:37:15Z","timestamp":1787161035817,"version":"build-2736575974"},"reference-count":60,"publisher":"Royal Society of Chemistry (RSC)","issue":"32","license":[{"start":{"date-parts":[[2026,7,29]],"date-time":"2026-07-29T00:00:00Z","timestamp":1785283200000},"content-version":"vor","delay-in-days":209,"URL":"http:\/\/creativecommons.org\/licenses\/by-nc\/3.0\/"}],"funder":[{"DOI":"10.13039\/501100008530","name":"European Regional Development Fund","doi-asserted-by":"publisher","award":["PID2021-126121NB-I00"],"award-info":[{"award-number":["PID2021-126121NB-I00"]}],"id":[{"id":"10.13039\/501100008530","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100008530","name":"European Regional Development Fund","doi-asserted-by":"publisher","award":["PID2024- 161591NB-I00"],"award-info":[{"award-number":["PID2024- 161591NB-I00"]}],"id":[{"id":"10.13039\/501100008530","id-type":"DOI","asserted-by":"publisher"}]}],"content-domain":{"domain":["rsc.org"],"crossmark-restriction":true},"short-container-title":["Soft Matter"],"abstract":"<jats:p>We reveal how internal mobility mimicking nuclear pseudostratification dictates phase behaviour in dense cellular assemblies. This physical mechanism naturally yields the structural diversity of biological epithelia.<\/jats:p>","DOI":"10.1039\/d6sm00469e","type":"journal-article","created":{"date-parts":[[2026,7,29]],"date-time":"2026-07-29T09:03:43Z","timestamp":1785315823000},"page":"5377-5390","update-policy":"https:\/\/doi.org\/10.1039\/rsc_crossmark_policy","source":"Crossref","is-referenced-by-count":0,"title":["Emergent symmetries and phase phenomenology in an individual-based model of cell-like particles with internal mobility"],"prefix":"10.1039","volume":"22","author":[{"ORCID":"https:\/\/orcid.org\/0000-0002-5314-4206","authenticated-orcid":false,"given":"Germ\u00e1n","family":"Luque-Caballero","sequence":"first","affiliation":[{"name":"Department of Applied Physics I, Universidad de Sevilla, Sevilla, Spain"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0009-0001-9102-8249","authenticated-orcid":false,"given":"Antonio J.","family":"D\u00edaz","sequence":"additional","affiliation":[{"name":"Department of Multicellular Dynamics, CABD (Andalusian Centre for Developmental Biology), CSIC\/Universidad Pablo de Olavide\/Junta de Andaluc\u00eda, Sevilla, Spain"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-2181-3858","authenticated-orcid":false,"given":"Fernando","family":"Casares","sequence":"additional","affiliation":[{"name":"Department of Multicellular Dynamics, CABD (Andalusian Centre for Developmental Biology), CSIC\/Universidad Pablo de Olavide\/Junta de Andaluc\u00eda, Sevilla, Spain"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-2170-0535","authenticated-orcid":false,"given":"Alejandro","family":"Cuetos","sequence":"additional","affiliation":[{"name":"Department of Physical, Chemical and Natural Systems, Universidad Pablo de Olavide, Sevilla, Spain"},{"name":"Center for Nanoscience and Sustainable Technologies (CNATS), Universidad Pablo de Olavide, Sevilla, Spain"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"292","published-online":{"date-parts":[[2026]]},"reference":[{"key":"D6SM00469E\/cit1\/1","doi-asserted-by":"crossref","first-page":"954","DOI":"10.1016\/j.copbio.2013.04.006","volume":"24","author":"Gleghorn","year":"2013","journal-title":"Curr. Opin. Biotechnol."},{"key":"D6SM00469E\/cit2\/1","doi-asserted-by":"crossref","first-page":"R175","DOI":"10.1530\/REP-12-0081","volume":"145","author":"Vasieva","year":"2013","journal-title":"Reproduction"},{"key":"D6SM00469E\/cit3\/1","doi-asserted-by":"crossref","first-page":"dev204894","DOI":"10.1242\/dev.204894","volume":"152","author":"Claussen","year":"2025","journal-title":"Development"},{"key":"D6SM00469E\/cit4\/1","doi-asserted-by":"crossref","DOI":"10.1017\/CBO9781107589070","volume-title":"On Growth and Form","author":"Thompson","year":"2014","unstructured":"D. W.Thompson ,  On Growth and Form ,  Cambridge University Press ,  Cambridge, UK ,  2014"},{"key":"D6SM00469E\/cit5\/1","doi-asserted-by":"crossref","first-page":"e66750","DOI":"10.7554\/eLife.66750","volume":"10","author":"Alba","year":"2021","journal-title":"eLife"},{"key":"D6SM00469E\/cit6\/1","doi-asserted-by":"crossref","first-page":"401","DOI":"10.1146\/annurev-biophys-081624-030543","volume":"54","author":"Russo","year":"2025","journal-title":"Annu. Rev. Biophys."},{"key":"D6SM00469E\/cit7\/1","doi-asserted-by":"crossref","first-page":"887","DOI":"10.1038\/s41586-024-07934-8","volume":"633","author":"Caldarelli","year":"2024","journal-title":"Nature"},{"key":"D6SM00469E\/cit8\/1","doi-asserted-by":"crossref","first-page":"23","DOI":"10.1299\/jbse.23-00339","volume":"18","author":"Okajima","year":"2023","journal-title":"J. Biomech. Sci. Eng."},{"key":"D6SM00469E\/cit9\/1","doi-asserted-by":"crossref","first-page":"101867","DOI":"10.1016\/j.jbc.2022.101867","volume":"298","author":"Zancla","year":"2022","journal-title":"J. Biol. Chem."},{"key":"D6SM00469E\/cit10\/1","doi-asserted-by":"crossref","first-page":"1514","DOI":"10.1038\/s41556-025-01632-x","volume":"27","author":"Villeneuve","year":"2025","journal-title":"Nat. Cell Biol."},{"key":"D6SM00469E\/cit11\/1","doi-asserted-by":"crossref","first-page":"681933","DOI":"10.3389\/fcell.2021.681933","volume":"9","author":"Lobo-Cabrera","year":"2021","journal-title":"Front. Cell Dev. Biol."},{"key":"D6SM00469E\/cit12\/1","doi-asserted-by":"crossref","first-page":"1009","DOI":"10.1016\/j.ssc.2010.01.021","volume":"150","author":"Wang","year":"2010","journal-title":"Solid State Commun."},{"key":"D6SM00469E\/cit13\/1","doi-asserted-by":"crossref","first-page":"401","DOI":"10.1007\/s40571-015-0082-3","volume":"2","author":"Van Liedekerke","year":"2015","journal-title":"Comput. Part. Mech."},{"key":"D6SM00469E\/cit14\/1","doi-asserted-by":"crossref","first-page":"633","DOI":"10.1038\/nrm2222","volume":"8","author":"Lecuit","year":"2007","journal-title":"Nat. Rev. Mol. Cell Biol."},{"key":"D6SM00469E\/cit15\/1","doi-asserted-by":"crossref","first-page":"225","DOI":"10.1038\/nrm3775","volume":"15","author":"Rodriguez-Boulan","year":"2014","journal-title":"Nat. Rev. Mol. Cell Biol."},{"key":"D6SM00469E\/cit16\/1","doi-asserted-by":"crossref","first-page":"1307","DOI":"10.1161\/CIRCRESAHA.108.173989","volume":"102","author":"Dahl","year":"2008","journal-title":"Circ. Res."},{"key":"D6SM00469E\/cit17\/1","doi-asserted-by":"crossref","first-page":"1240104","DOI":"10.1126\/science.1240104","volume":"341","author":"Swift","year":"2013","journal-title":"Science"},{"key":"D6SM00469E\/cit18\/1","doi-asserted-by":"crossref","first-page":"377","DOI":"10.1002\/cne.900620207","volume":"62","author":"Sauer","year":"1935","journal-title":"J. Comp. Neurol."},{"key":"D6SM00469E\/cit19\/1","doi-asserted-by":"crossref","first-page":"306","DOI":"10.1111\/j.1440-169X.2012.01342.x","volume":"54","author":"Spear","year":"2012","journal-title":"Dev., Growth Differ."},{"key":"D6SM00469E\/cit20\/1","doi-asserted-by":"crossref","first-page":"5497","DOI":"10.1073\/pnas.1312076111","volume":"111","author":"Hannezo","year":"2014","journal-title":"Proc. Natl. Acad. Sci. U. S. A."},{"key":"D6SM00469E\/cit21\/1","doi-asserted-by":"crossref","first-page":"1195","DOI":"10.1016\/j.cell.2009.06.032","volume":"138","author":"Norden","year":"2009","journal-title":"Cell"},{"key":"D6SM00469E\/cit22\/1","doi-asserted-by":"crossref","first-page":"2727","DOI":"10.1007\/s00018-012-0952-2","volume":"69","author":"Kosodo","year":"2012","journal-title":"Cell. Mol. Life Sci."},{"key":"D6SM00469E\/cit23\/1","doi-asserted-by":"crossref","first-page":"179","DOI":"10.1038\/nature10137","volume":"474","author":"Dupont","year":"2011","journal-title":"Nature"},{"key":"D6SM00469E\/cit24\/1","doi-asserted-by":"crossref","first-page":"948","DOI":"10.1016\/j.cell.2013.05.008","volume":"153","author":"Heisenberg","year":"2013","journal-title":"Cell"},{"key":"D6SM00469E\/cit25\/1","doi-asserted-by":"crossref","first-page":"581","DOI":"10.1038\/ncb3524","volume":"19","author":"Petridou","year":"2017","journal-title":"Nat. Cell Biol."},{"key":"D6SM00469E\/cit26\/1","doi-asserted-by":"crossref","first-page":"613","DOI":"10.1038\/s41567-018-0089-9","volume":"14","author":"Atia","year":"2018","journal-title":"Nat. Phys."},{"key":"D6SM00469E\/cit27\/1","doi-asserted-by":"crossref","first-page":"1592","DOI":"10.1038\/s41563-024-01972-3","volume":"23","author":"Kim","year":"2024","journal-title":"Nat. Mater."},{"key":"D6SM00469E\/cit28\/1","doi-asserted-by":"crossref","first-page":"2177","DOI":"10.1039\/b716393b","volume":"18","author":"Yang","year":"2008","journal-title":"J. Mater. Chem."},{"key":"D6SM00469E\/cit29\/1","doi-asserted-by":"crossref","first-page":"1688","DOI":"10.1038\/ncomms2694","volume":"4","author":"Sacanna","year":"2013","journal-title":"Nat. Commun."},{"key":"D6SM00469E\/cit30\/1","doi-asserted-by":"crossref","first-page":"1053","DOI":"10.1038\/s41578-021-00323-x","volume":"6","author":"Hueckel","year":"2021","journal-title":"Nat. Rev. Mater."},{"key":"D6SM00469E\/cit31\/1","doi-asserted-by":"crossref","first-page":"433","DOI":"10.1038\/s41570-024-00603-4","volume":"8","author":"Kamp","year":"2024","journal-title":"Nat. Rev. Chem."},{"key":"D6SM00469E\/cit32\/1","doi-asserted-by":"crossref","first-page":"55","DOI":"10.1016\/0097-8485(94)80023-5","volume":"18","author":"Vega","year":"1994","journal-title":"Comput. Chem."},{"key":"D6SM00469E\/cit33\/1","doi-asserted-by":"crossref","first-page":"1196","DOI":"10.1103\/PhysRevLett.45.1196","volume":"45","author":"Parrinello","year":"1980","journal-title":"Phys. Rev. Lett."},{"key":"D6SM00469E\/cit34\/1","doi-asserted-by":"crossref","first-page":"7182","DOI":"10.1063\/1.328693","volume":"52","author":"Parrinello","year":"1981","journal-title":"J. Appl. Phys."},{"key":"D6SM00469E\/cit35\/1","volume-title":"Understanding Molecular Simulation: From Algorithms to Applications","author":"Frenkel","year":"2002","unstructured":"D.Frenkel  and  B.Smit ,  Understanding Molecular Simulation: From Algorithms to Applications ,  Academic Press ,  San Diego , 2nd edn,  2002"},{"key":"D6SM00469E\/cit36\/1","doi-asserted-by":"crossref","first-page":"27059","DOI":"10.1039\/C8CP05248D","volume":"20","author":"Eslami","year":"2018","journal-title":"Phys. Chem. Chem. Phys."},{"key":"D6SM00469E\/cit37\/1","doi-asserted-by":"crossref","first-page":"095101","DOI":"10.1063\/1.4999709","volume":"147","author":"Pantelopulos","year":"2017","journal-title":"J. Chem. Phys."},{"key":"D6SM00469E\/cit38\/1","doi-asserted-by":"crossref","first-page":"102252","DOI":"10.1016\/j.cis.2020.102252","volume":"284","author":"Lotito","year":"2020","journal-title":"Adv. Colloid Interface Sci."},{"key":"D6SM00469E\/cit39\/1","doi-asserted-by":"crossref","first-page":"2026","DOI":"10.1039\/D4SM01377H","volume":"21","author":"L\u00f6ffler","year":"2025","journal-title":"Soft Matter"},{"key":"D6SM00469E\/cit40\/1","first-page":"012103","volume":"653","author":"Koss","year":"2015","journal-title":"J. Phys.: Conf. Ser."},{"key":"D6SM00469E\/cit41\/1","doi-asserted-by":"crossref","first-page":"197","DOI":"10.1016\/j.devcel.2019.05.027","volume":"50","author":"Ambrosini","year":"2019","journal-title":"Dev. Cell"},{"key":"D6SM00469E\/cit42\/1","doi-asserted-by":"crossref","first-page":"676","DOI":"10.1038\/nmeth.2019","volume":"9","author":"Schindelin","year":"2012","journal-title":"Nat. Methods"},{"key":"D6SM00469E\/cit43\/1","doi-asserted-by":"crossref","first-page":"e1905897","DOI":"10.1080\/00268976.2021.1905897","volume":"119","author":"Liu","year":"2021","journal-title":"Mol. Phys."},{"key":"D6SM00469E\/cit44\/1","doi-asserted-by":"crossref","first-page":"578","DOI":"10.1016\/0009-2614(74)80109-0","volume":"28","author":"Torrie","year":"1974","journal-title":"Chem. Phys. Lett."},{"key":"D6SM00469E\/cit45\/1","doi-asserted-by":"crossref","first-page":"187","DOI":"10.1016\/0021-9991(77)90121-8","volume":"23","author":"Torrie","year":"1977","journal-title":"J. Comput. Phys."},{"key":"D6SM00469E\/cit46\/1","doi-asserted-by":"crossref","first-page":"1020","DOI":"10.1038\/35059035","volume":"409","author":"Auer","year":"2001","journal-title":"Nature"},{"key":"D6SM00469E\/cit47\/1","doi-asserted-by":"crossref","first-page":"292","DOI":"10.1016\/j.jcis.2014.10.044","volume":"440","author":"Puertas","year":"2015","journal-title":"J. Colloid Interface Sci."},{"key":"D6SM00469E\/cit48\/1","doi-asserted-by":"crossref","first-page":"244115","DOI":"10.1063\/1.3506838","volume":"133","author":"Filion","year":"2010","journal-title":"J. Chem. Phys."},{"key":"D6SM00469E\/cit49\/1","doi-asserted-by":"crossref","first-page":"144104","DOI":"10.1063\/1.2052648","volume":"123","author":"K\u00e4stner","year":"2005","journal-title":"J. Chem.\nPhys."},{"key":"D6SM00469E\/cit50\/1","doi-asserted-by":"crossref","first-page":"L124","DOI":"10.1088\/0022-3719\/5\/11\/002","volume":"5","author":"Kosterlitz","year":"1972","journal-title":"J. Phys. C-Solid State Phys."},{"key":"D6SM00469E\/cit51\/1","doi-asserted-by":"crossref","first-page":"1181","DOI":"10.1088\/0022-3719\/6\/7\/010","volume":"6","author":"Kosterlitz","year":"1973","journal-title":"J. Phys. C-Solid State Phys."},{"key":"D6SM00469E\/cit52\/1","doi-asserted-by":"crossref","first-page":"1201","DOI":"10.1103\/PhysRevLett.39.1201","volume":"39","author":"Nelson","year":"1977","journal-title":"Phys. Rev. Lett."},{"key":"D6SM00469E\/cit53\/1","doi-asserted-by":"crossref","first-page":"2457","DOI":"10.1103\/PhysRevB.19.2457","volume":"19","author":"Nelson","year":"1979","journal-title":"Phys. Rev. B:Condens. Matter Mater. Phys."},{"key":"D6SM00469E\/cit54\/1","doi-asserted-by":"crossref","first-page":"121","DOI":"10.1103\/PhysRevLett.41.121","volume":"41","author":"Halperin","year":"1978","journal-title":"Phys. Rev. Lett."},{"key":"D6SM00469E\/cit55\/1","doi-asserted-by":"crossref","first-page":"1855","DOI":"10.1103\/PhysRevB.19.1855","volume":"19","author":"Young","year":"1979","journal-title":"Phys. Rev. B:Condens. Matter Mater. Phys."},{"key":"D6SM00469E\/cit56\/1","doi-asserted-by":"crossref","first-page":"235","DOI":"10.12921\/cmst.2004.10.02.235-255","volume":"10","author":"Wojciechowski","year":"2004","journal-title":"Comput. Methods Sci. Technol."},{"key":"D6SM00469E\/cit57\/1","first-page":"L371","volume":"18","author":"Fortini","year":"2006","journal-title":"J. Phys.: Condens. Matter"},{"key":"D6SM00469E\/cit58\/1","doi-asserted-by":"crossref","first-page":"347","DOI":"10.1146\/annurev-conmatphys-070909-104045","volume":"1","author":"Liu","year":"2010","journal-title":"Annu. Rev. Condens. Matter Phys."},{"key":"D6SM00469E\/cit59\/1","doi-asserted-by":"crossref","first-page":"105045","DOI":"10.1016\/j.biosystems.2023.105045","volume":"234","author":"Pajic-Lijakovic","year":"2023","journal-title":"BioSystems"},{"key":"D6SM00469E\/cit60\/1","doi-asserted-by":"crossref","first-page":"8171","DOI":"10.1063\/1.459295","volume":"93","author":"Vega","year":"1990","journal-title":"J. Chem. Phys."}],"container-title":["Soft Matter"],"original-title":[],"language":"en","link":[{"URL":"http:\/\/pubs.rsc.org\/SM\/article-pdf\/doi\/10.1039\/D6SM00469E\/0\/D6SM00469E.pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,8,19]],"date-time":"2026-08-19T16:42:58Z","timestamp":1787157778000},"score":1,"resource":{"primary":{"URL":"http:\/\/pubs.rsc.org\/SM\/article-lookup\/doi\/10.1039\/D6SM00469E"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026]]},"references-count":60,"journal-issue":{"issue":"32","published-print":{"date-parts":[[2026,8,19]]}},"URL":"https:\/\/doi.org\/10.1039\/d6sm00469e","relation":{"has-review":[{"id-type":"doi","id":"10.1039\/D6SM00469E\/v3\/decision1","asserted-by":"object"},{"id-type":"doi","id":"10.1039\/D6SM00469E\/v2\/review1","asserted-by":"object"},{"id-type":"doi","id":"10.1039\/D6SM00469E\/v3\/response1","asserted-by":"object"},{"id-type":"doi","id":"10.1039\/D6SM00469E\/v2\/review2","asserted-by":"object"},{"id-type":"doi","id":"10.1039\/D6SM00469E\/v1\/review1","asserted-by":"object"},{"id-type":"doi","id":"10.1039\/D6SM00469E\/v2\/decision1","asserted-by":"object"},{"id-type":"doi","id":"10.1039\/D6SM00469E\/v1\/review2","asserted-by":"object"},{"id-type":"doi","id":"10.1039\/D6SM00469E\/v2\/response1","asserted-by":"object"},{"id-type":"doi","id":"10.1039\/D6SM00469E\/v1\/decision1","asserted-by":"object"}]},"ISSN":["1744-683X","1744-6848"],"issn-type":[{"value":"1744-683X","type":"print"},{"value":"1744-6848","type":"electronic"}],"subject":[],"published":{"date-parts":[[2026]]}}}