{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,8,18]],"date-time":"2026-08-18T18:29:28Z","timestamp":1787077768450,"version":"build-2736575974"},"reference-count":57,"publisher":"National Academy of Sciences","issue":"34","license":[{"start":{"date-parts":[[2026,8,18]],"date-time":"2026-08-18T00:00:00Z","timestamp":1787011200000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by-nc-nd\/4.0\/"}],"funder":[{"DOI":"10.13039\/100000057","name":"HHS | NIH | National Institute of General Medical Sciences","doi-asserted-by":"publisher","award":["R35GM148485"],"award-info":[{"award-number":["R35GM148485"]}],"id":[{"id":"10.13039\/100000057","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/100000057","name":"HHS | NIH | National Institute of General Medical Sciences","doi-asserted-by":"publisher","award":["R01GM126047"],"award-info":[{"award-number":["R01GM126047"]}],"id":[{"id":"10.13039\/100000057","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/100000057","name":"HHS | NIH | National Institute of General Medical Sciences","doi-asserted-by":"publisher","award":["T32GM007183"],"award-info":[{"award-number":["T32GM007183"]}],"id":[{"id":"10.13039\/100000057","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/100000057","name":"HHS | NIH | National Institute of General Medical Sciences","doi-asserted-by":"publisher","award":["R35GM156889"],"award-info":[{"award-number":["R35GM156889"]}],"id":[{"id":"10.13039\/100000057","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/100009633","name":"HHS | NIH | Eunice Kennedy Shriver National Institute of Child Health and Human Development","doi-asserted-by":"publisher","award":["T32HD055164"],"award-info":[{"award-number":["T32HD055164"]}],"id":[{"id":"10.13039\/100009633","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/100000001","name":"National Science Foundation","doi-asserted-by":"publisher","award":["PHY-2413073"],"award-info":[{"award-number":["PHY-2413073"]}],"id":[{"id":"10.13039\/100000001","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/100000001","name":"National Science Foundation","doi-asserted-by":"publisher","award":["PHY-2443851"],"award-info":[{"award-number":["PHY-2443851"]}],"id":[{"id":"10.13039\/100000001","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100000854","name":"Human Frontier Science Program","doi-asserted-by":"publisher","award":["RGEC31\/2024"],"award-info":[{"award-number":["RGEC31\/2024"]}],"id":[{"id":"10.13039\/501100000854","id-type":"DOI","asserted-by":"publisher"}]}],"content-domain":{"domain":["www.pnas.org"],"crossmark-restriction":true},"short-container-title":["Proc. Natl. Acad. Sci. U.S.A."],"published-print":{"date-parts":[[2026,8,25]]},"abstract":"<jats:p>\n                    Collective migration of epithelial cells drives diverse tissue remodeling processes. In many cases, a free tissue edge works alone or in combination with other external cues to align the cells for collective movement, but how edge-free or closed epithelia become polarized for directed migration without these cues is unclear. Here, we use the rotational migration of the follicular epithelial cells in the\n                    <jats:italic toggle=\"yes\">Drosophila<\/jats:italic>\n                    egg chamber to explore how cells in an edgeless epithelium initiate rotational collective migration, and how the rotational axis is specified. By employing methods for long-term live imaging and delaying the onset of rotation, we show that symmetry breaking can occur at multiple developmental stages and that the atypical cadherin Fat2 promotes local motility at the basal epithelial surface before rotation begins. We then combine experiments with theoretical modeling to identify a positive feedback loop in which planar polarization of Fat2 aligns the front\u2013rear axes of the individual cells in a common direction and the resulting tissue motion leads to the planar polarization of Fat2. This mechanosensitive feedback, coupled with rigid-body dynamics of the egg chamber, can break chiral symmetry and produce sustained rotation in silico. We further propose that mechanical constraints arising from intertissue interactions and tissue geometry ensure that rotation occurs around the anterior\u2013posterior axis. Our findings suggest a biophysical mechanism\u2014combining Fat2-mediated velocity\u2013polarity alignment, rigid-body dynamics, and tissue geometry\u2014by which a closed epithelial tissue can self-organize into persistent, large-scale rotational migration in vivo, expanding current flocking theories.\n                  <\/jats:p>","DOI":"10.1073\/pnas.2528342123","type":"journal-article","created":{"date-parts":[[2026,8,18]],"date-time":"2026-08-18T17:59:27Z","timestamp":1787075967000},"update-policy":"https:\/\/doi.org\/10.1073\/pnas.cm10313","source":"Crossref","is-referenced-by-count":0,"title":["Initiation of rotational collective migration in\n                    <i>Drosophila<\/i>\n                    through tissue geometry and mechanochemical feedback"],"prefix":"10.1073","volume":"123","author":[{"given":"Sierra","family":"Schwabach","sequence":"first","affiliation":[{"id":[{"id":"https:\/\/ror.org\/024mw5h28","id-type":"ROR","asserted-by":"publisher"}],"name":"Department of Molecular Genetics and Cell Biology, The University of Chicago","place":["Chicago"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-5278-6277","authenticated-orcid":false,"given":"Sreejith","family":"Santhosh","sequence":"additional","affiliation":[{"id":[{"id":"https:\/\/ror.org\/0168r3w48","id-type":"ROR","asserted-by":"publisher"}],"name":"Department of Physics, University of California","place":["San Diego"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-6170-3365","authenticated-orcid":false,"given":"Audrey Miller","family":"Williams","sequence":"additional","affiliation":[{"id":[{"id":"https:\/\/ror.org\/024mw5h28","id-type":"ROR","asserted-by":"publisher"}],"name":"Department of Molecular Genetics and Cell Biology, The University of Chicago","place":["Chicago"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-7551-6560","authenticated-orcid":false,"given":"Maureen","family":"Cetera","sequence":"additional","affiliation":[{"id":[{"id":"https:\/\/ror.org\/024mw5h28","id-type":"ROR","asserted-by":"publisher"}],"name":"Committee on Development, Regeneration, and Stem Cell Biology, The University of Chicago","place":["Chicago"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-6007-5878","authenticated-orcid":false,"given":"Mattia","family":"Serra","sequence":"additional","affiliation":[{"id":[{"id":"https:\/\/ror.org\/0168r3w48","id-type":"ROR","asserted-by":"publisher"}],"name":"Department of Physics, University of California","place":["San Diego"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-0473-7451","authenticated-orcid":false,"given":"Sally","family":"Horne-Badovinac","sequence":"additional","affiliation":[{"id":[{"id":"https:\/\/ror.org\/024mw5h28","id-type":"ROR","asserted-by":"publisher"}],"name":"Department of Molecular Genetics and Cell Biology, The University of Chicago","place":["Chicago"]},{"id":[{"id":"https:\/\/ror.org\/024mw5h28","id-type":"ROR","asserted-by":"publisher"}],"name":"Committee on Development, Regeneration, and Stem Cell Biology, The University of Chicago","place":["Chicago"]}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"341","published-online":{"date-parts":[[2026,8,18]]},"reference":[{"key":"e_1_3_4_1_2","doi-asserted-by":"publisher","DOI":"10.1038\/s41580-025-00858-9"},{"key":"e_1_3_4_2_2","doi-asserted-by":"publisher","DOI":"10.1007\/s00018-012-1251-7"},{"key":"e_1_3_4_3_2","doi-asserted-by":"publisher","DOI":"10.1016\/j.ceb.2016.04.001"},{"key":"e_1_3_4_4_2","doi-asserted-by":"publisher","DOI":"10.1126\/science.aau3429"},{"key":"e_1_3_4_5_2","doi-asserted-by":"publisher","DOI":"10.1038\/nrm.2015.14"},{"key":"e_1_3_4_6_2","doi-asserted-by":"publisher","DOI":"10.1073\/pnas.0705062104"},{"key":"e_1_3_4_7_2","doi-asserted-by":"publisher","DOI":"10.1016\/j.ceb.2021.04.006"},{"key":"e_1_3_4_8_2","doi-asserted-by":"publisher","DOI":"10.1038\/s41580-021-00366-6"},{"key":"e_1_3_4_9_2","doi-asserted-by":"publisher","DOI":"10.1002\/cm.20077"},{"key":"e_1_3_4_10_2","doi-asserted-by":"publisher","DOI":"10.1039\/c3ib40054a"},{"key":"e_1_3_4_11_2","doi-asserted-by":"publisher","DOI":"10.1038\/s41567-020-0875-z"},{"key":"e_1_3_4_12_2","doi-asserted-by":"publisher","DOI":"10.1038\/s41567-023-02295-x"},{"key":"e_1_3_4_13_2","doi-asserted-by":"publisher","DOI":"10.1103\/PhysRevLett.114.228102"},{"key":"e_1_3_4_14_2","doi-asserted-by":"publisher","DOI":"10.1073\/pnas.1119578109"},{"key":"e_1_3_4_15_2","doi-asserted-by":"publisher","DOI":"10.1073\/pnas.1201141110"},{"key":"e_1_3_4_16_2","doi-asserted-by":"publisher","DOI":"10.1073\/pnas.1805932115"},{"key":"e_1_3_4_17_2","doi-asserted-by":"publisher","DOI":"10.1103\/PRXLife.2.033006"},{"key":"e_1_3_4_18_2","doi-asserted-by":"publisher","DOI":"10.1038\/s41467-023-37054-2"},{"key":"e_1_3_4_19_2","doi-asserted-by":"publisher","DOI":"10.1038\/s42003-025-09136-y"},{"key":"e_1_3_4_20_2","doi-asserted-by":"publisher","DOI":"10.1016\/j.devcel.2022.08.008"},{"key":"e_1_3_4_21_2","doi-asserted-by":"publisher","DOI":"10.1111\/cas.13816"},{"key":"e_1_3_4_22_2","doi-asserted-by":"publisher","DOI":"10.1126\/sciadv.abn5406"},{"key":"e_1_3_4_23_2","doi-asserted-by":"publisher","DOI":"10.1038\/s41567-021-01336-7"},{"key":"e_1_3_4_24_2","doi-asserted-by":"publisher","DOI":"10.1038\/embor.2012.149"},{"key":"e_1_3_4_25_2","doi-asserted-by":"publisher","DOI":"10.1002\/dvdy.20286"},{"key":"e_1_3_4_26_2","doi-asserted-by":"publisher","DOI":"10.1093\/genetics\/iyad183"},{"key":"e_1_3_4_27_2","doi-asserted-by":"publisher","DOI":"10.1126\/science.1199424"},{"key":"e_1_3_4_28_2","doi-asserted-by":"publisher","DOI":"10.1038\/ncomms6511"},{"key":"e_1_3_4_29_2","doi-asserted-by":"publisher","DOI":"10.1016\/j.devcel.2016.06.009"},{"key":"e_1_3_4_30_2","doi-asserted-by":"publisher","DOI":"10.7554\/eLife.24958"},{"key":"e_1_3_4_31_2","doi-asserted-by":"publisher","DOI":"10.7554\/eLife.32943"},{"key":"e_1_3_4_32_2","doi-asserted-by":"publisher","DOI":"10.1083\/jcb.201508081"},{"key":"e_1_3_4_33_2","doi-asserted-by":"publisher","DOI":"10.1016\/j.devcel.2017.02.003"},{"key":"e_1_3_4_34_2","doi-asserted-by":"publisher","DOI":"10.7554\/eLife.78343"},{"key":"e_1_3_4_35_2","doi-asserted-by":"publisher","DOI":"10.1016\/j.cub.2013.06.014"},{"key":"e_1_3_4_36_2","doi-asserted-by":"publisher","DOI":"10.1242\/jcs.261173"},{"key":"e_1_3_4_37_2","doi-asserted-by":"publisher","DOI":"10.1371\/journal.pgen.1007107"},{"key":"e_1_3_4_38_2","doi-asserted-by":"publisher","DOI":"10.1016\/j.celrep.2016.04.014"},{"key":"e_1_3_4_39_2","doi-asserted-by":"publisher","DOI":"10.1016\/j.xpro.2021.100371"},{"key":"e_1_3_4_40_2","doi-asserted-by":"publisher","DOI":"10.1534\/genetics.106.065961"},{"key":"e_1_3_4_41_2","doi-asserted-by":"publisher","DOI":"10.1242\/dev.02800"},{"key":"e_1_3_4_42_2","doi-asserted-by":"publisher","DOI":"10.1016\/j.cub.2019.01.049"},{"key":"e_1_3_4_43_2","doi-asserted-by":"publisher","DOI":"10.1038\/ncb1058"},{"key":"e_1_3_4_44_2","doi-asserted-by":"publisher","DOI":"10.1083\/jcb.201007142"},{"key":"e_1_3_4_45_2","unstructured":"A. C. Spradling \u201cDevelopmental genetics of oogenesis\u201d in The Development of Drosophila melanogaster M. Bate A. Martinez Arias Eds. (Cold Spring Harbor Laboratory Press Cold Spring Harbor NY 1993) vol. I chap. 1."},{"key":"e_1_3_4_46_2","doi-asserted-by":"publisher","DOI":"10.1242\/dev.200456"},{"key":"e_1_3_4_47_2","doi-asserted-by":"publisher","DOI":"10.1038\/s41556-022-00844-9"},{"key":"e_1_3_4_48_2","doi-asserted-by":"publisher","DOI":"10.1242\/dev.104.2.245"},{"key":"e_1_3_4_49_2","doi-asserted-by":"publisher","DOI":"10.1242\/dev.065508"},{"key":"e_1_3_4_50_2","doi-asserted-by":"publisher","DOI":"10.1038\/s41467-019-10874-x"},{"key":"e_1_3_4_51_2","doi-asserted-by":"publisher","DOI":"10.1146\/annurev-conmatphys-031218-013516"},{"key":"e_1_3_4_52_2","doi-asserted-by":"publisher","DOI":"10.1016\/j.celrep.2017.06.031"},{"key":"e_1_3_4_53_2","doi-asserted-by":"publisher","DOI":"10.1038\/nprot.2007.363"},{"key":"e_1_3_4_54_2","doi-asserted-by":"publisher","DOI":"10.1007\/978-1-4939-6371-3_12"},{"key":"e_1_3_4_55_2","doi-asserted-by":"publisher","DOI":"10.1038\/s41592-020-01018-x"},{"key":"e_1_3_4_56_2","unstructured":"S. Santhosh Code and data from \u201cInitiation of rotational collective migration in Drosophila through tissue geometry and mechanochemical feedback.\u201d Github. https:\/\/github.com\/SreejithSanthosh\/egg-chamber-rotation. Deposited 5 September 2025."},{"key":"e_1_3_4_57_2","unstructured":"S. Schwabach Data from \u201cInitiation of rotational collective migration in Drosophila through tissue geometry and mechanochemical feedback.\u201d Figshare. https:\/\/doi.org\/10.6084\/m9.figshare.32562264.v1. Deposited 2 June 2026."}],"container-title":["Proceedings of the National Academy of Sciences"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/pnas.org\/doi\/pdf\/10.1073\/pnas.2528342123","content-type":"application\/pdf","content-version":"vor","intended-application":"syndication"},{"URL":"https:\/\/pnas.org\/doi\/pdf\/10.1073\/pnas.2528342123","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,8,18]],"date-time":"2026-08-18T18:00:15Z","timestamp":1787076015000},"score":1,"resource":{"primary":{"URL":"https:\/\/pnas.org\/doi\/10.1073\/pnas.2528342123"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,8,18]]},"references-count":57,"journal-issue":{"issue":"34","published-print":{"date-parts":[[2026,8,25]]}},"alternative-id":["10.1073\/pnas.2528342123"],"URL":"https:\/\/doi.org\/10.1073\/pnas.2528342123","relation":{},"ISSN":["0027-8424","1091-6490"],"issn-type":[{"value":"0027-8424","type":"print"},{"value":"1091-6490","type":"electronic"}],"subject":[],"published":{"date-parts":[[2026,8,18]]},"assertion":[{"value":"2025-10-13","order":0,"name":"received","label":"Received","group":{"name":"publication_history","label":"Publication History"}},{"value":"2026-07-10","order":2,"name":"accepted","label":"Accepted","group":{"name":"publication_history","label":"Publication History"}},{"value":"2026-08-18","order":3,"name":"published","label":"Published","group":{"name":"publication_history","label":"Publication History"}}],"article-number":"e2528342123"}}