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How precisely localized actomyosin regulators drive apical constriction remains poorly understood. Caenorhabditis elegans gastrulation provides a valuable model to address this question. The Arp2\/3 complex is essential in C. elegans gastrulation. To understand how Arp2\/3 is locally regulated, we imaged embryos with endogenously tagged Arp2\/3 and its nucleation-promoting factors (NPFs). The three NPFs\u2014WAVE, WASP, and WASH\u2014controlled Arp2\/3 localization at distinct subcellular locations. We exploited this finding to study distinct populations of Arp2\/3 and found that only WAVE depletion caused penetrant gastrulation defects. WAVE localized basolaterally with Arp2\/3 and controlled F-actin levels near cell\u2013cell contacts. WAVE and Arp2\/3 localization depended on CED-10\/Rac. Establishing ectopic cell contacts recruited WAVE and Arp2\/3, identifying the contact as a symmetry-breaking cue for localization of these proteins. These results suggest that cell\u2013cell signaling via Rac activates WAVE and Arp2\/3 basolaterally and that basolateral Arp2\/3 makes an important contribution to apical constriction.<\/jats:p>","DOI":"10.1083\/jcb.202409133","type":"journal-article","created":{"date-parts":[[2025,2,12]],"date-time":"2025-02-12T02:21:03Z","timestamp":1739326863000},"update-policy":"https:\/\/doi.org\/10.1083\/jcb.crossmarkpolicy","source":"Crossref","is-referenced-by-count":4,"title":["Cell signaling facilitates apical constriction by basolaterally recruiting Arp2\/3 via Rac and WAVE"],"prefix":"10.1083","volume":"224","author":[{"ORCID":"https:\/\/orcid.org\/0000-0003-4174-8202","authenticated-orcid":true,"given":"Pu","family":"Zhang","sequence":"first","affiliation":[{"id":[{"id":"https:\/\/ror.org\/0130frc33","id-type":"ROR","asserted-by":"publisher"}],"name":"University of North Carolina at Chapel Hill 1 Biology Department, , Chapel Hill, NC,","place":["USA"]},{"id":[{"id":"https:\/\/ror.org\/0130frc33","id-type":"ROR","asserted-by":"publisher"}],"name":"University of North Carolina at Chapel Hill 2 Curriculum in Cell Biology and Physiology, , Chapel Hill, NC,","place":["USA"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-7989-3291","authenticated-orcid":true,"given":"Taylor N.","family":"Medwig-Kinney","sequence":"additional","affiliation":[{"id":[{"id":"https:\/\/ror.org\/0130frc33","id-type":"ROR","asserted-by":"publisher"}],"name":"University of North Carolina at Chapel Hill 1 Biology Department, , Chapel Hill, NC,","place":["USA"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0009-0002-5476-5319","authenticated-orcid":true,"given":"Eleanor A.","family":"Breiner","sequence":"additional","affiliation":[{"id":[{"id":"https:\/\/ror.org\/0130frc33","id-type":"ROR","asserted-by":"publisher"}],"name":"University of North Carolina at Chapel Hill 1 Biology Department, , Chapel Hill, NC,","place":["USA"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0009-0003-6987-1322","authenticated-orcid":true,"given":"Jadyn M.","family":"Perez","sequence":"additional","affiliation":[{"id":[{"id":"https:\/\/ror.org\/0130frc33","id-type":"ROR","asserted-by":"publisher"}],"name":"University of North Carolina at Chapel Hill 1 Biology Department, , Chapel Hill, NC,","place":["USA"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0009-0000-2748-6379","authenticated-orcid":true,"given":"April N.","family":"Song","sequence":"additional","affiliation":[{"id":[{"id":"https:\/\/ror.org\/0130frc33","id-type":"ROR","asserted-by":"publisher"}],"name":"University of North Carolina at Chapel Hill 1 Biology Department, , Chapel Hill, NC,","place":["USA"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-6961-675X","authenticated-orcid":true,"given":"Bob","family":"Goldstein","sequence":"additional","affiliation":[{"id":[{"id":"https:\/\/ror.org\/0130frc33","id-type":"ROR","asserted-by":"publisher"}],"name":"University of North Carolina at Chapel Hill 1 Biology Department, , Chapel Hill, NC,","place":["USA"]},{"id":[{"id":"https:\/\/ror.org\/0130frc33","id-type":"ROR","asserted-by":"publisher"}],"name":"University of North Carolina at Chapel Hill 2 Curriculum in Cell Biology and Physiology, , Chapel Hill, NC,","place":["USA"]},{"id":[{"id":"https:\/\/ror.org\/0130frc33","id-type":"ROR","asserted-by":"publisher"}],"name":"Lineberger Comprehensive Cancer Center, University of North Carolina at Chapel Hill 3 , Chapel Hill, NC,","place":["USA"]}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"291","published-online":{"date-parts":[[2025,3,5]]},"reference":[{"key":"2025042817504404800_bib1","doi-asserted-by":"publisher","first-page":"286","DOI":"10.1016\/j.cels.2020.08.008","article-title":"Cell mechanics at the rear act to steer the direction of cell migration","volume":"11","author":"Allen","year":"2020","journal-title":"Cell Syst."},{"key":"2025042817504404800_bib2","doi-asserted-by":"publisher","first-page":"3103","DOI":"10.1242\/jcs.02444","article-title":"N-WASP deficiency impairs EGF internalization and actin assembly at clathrin-coated pits","volume":"118","author":"Benesch","year":"2005","journal-title":"J. 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