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Here, we uncover a role for the actin regulator WASP in the 3D migration of neutrophils. We show that WASP responds to substrate topology by enriching to sites of inward, substrate-induced membrane deformation. Superresolution imaging reveals that WASP preferentially enriches to the necks of these substrate-induced invaginations, a distribution that could support substrate pinching. WASP facilitates recruitment of the Arp2\/3 complex to these sites, stimulating local actin assembly that couples substrate features with the cytoskeleton. Surprisingly, WASP only enriches to membrane deformations in the front half of the cell, within a permissive zone set by WASP\u2019s front-biased regulator Cdc42. While WASP KO cells exhibit relatively normal migration on flat substrates, they are defective at topology-directed migration. Our data suggest that WASP integrates substrate topology with cell polarity by selectively polymerizing actin around substrate-induced membrane deformations in the front half of the cell.<\/jats:p>","DOI":"10.1083\/jcb.202104046","type":"journal-article","created":{"date-parts":[[2021,12,11]],"date-time":"2021-12-11T02:22:48Z","timestamp":1639189368000},"update-policy":"https:\/\/doi.org\/10.1083\/jcb.crossmarkpolicy","source":"Crossref","is-referenced-by-count":54,"title":["WASP integrates substrate topology and cell polarity to guide neutrophil migration"],"prefix":"10.1083","volume":"221","author":[{"ORCID":"https:\/\/orcid.org\/0000-0003-4821-3788","authenticated-orcid":true,"given":"Rachel M.","family":"Brunetti","sequence":"first","affiliation":[{"name":"Cardiovascular Research Institute, University of California, San Francisco, San Francisco, CA 1"},{"name":"Department of Biochemistry and Biophysics, University of California, San Francisco, San Francisco, CA 2"},{"name":"Center 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