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Here, we show that WASP\u2010family proteins also function as polymerases that accelerate elongation of uncapped actin filaments. When clustered on a surface, WASP\u2010family proteins can drive branched actin networks to grow much faster than they could by direct incorporation of soluble monomers. This polymerase activity arises from the coordinated action of two regulatory sequences: (i) a WASP homology 2 (WH2) domain that binds actin, and (ii) a proline\u2010rich sequence that binds profilin\u2013actin complexes. In the absence of profilin, WH2 domains are sufficient to accelerate filament elongation, but in the presence of profilin, proline\u2010rich sequences are required to support polymerase activity by (i) bringing polymerization\u2010competent actin monomers in proximity to growing filament ends, and (ii) promoting shuttling of actin monomers from profilin\u2013actin complexes onto nearby WH2 domains. Unoccupied WH2 domains transiently associate with free filament ends, preventing their growth and dynamically tethering the branched actin network to the WASP\u2010family proteins that create it. Collaboration between WH2 and proline\u2010rich sequences thus strikes a balance between filament growth and tethering. Our work expands the number of critical roles that WASP\u2010family proteins play in the assembly of branched actin networks to at least three: (i) promoting dendritic nucleation; (ii) linking actin networks to membranes; and (iii) accelerating filament elongation.<\/jats:p>","DOI":"10.15252\/embj.201797039","type":"journal-article","created":{"date-parts":[[2017,11,15]],"date-time":"2017-11-15T20:10:22Z","timestamp":1510776622000},"page":"102-121","update-policy":"https:\/\/doi.org\/10.1007\/springer_crossmark_policy","source":"Crossref","is-referenced-by-count":99,"title":["WH2 and proline\u2010rich domains of WASP\u2010family proteins collaborate to accelerate actin filament elongation"],"prefix":"10.1038","volume":"37","author":[{"ORCID":"https:\/\/orcid.org\/0000-0002-7458-4358","authenticated-orcid":false,"given":"Peter","family":"Bieling","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Scott D","family":"Hansen","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Orkun","family":"Akin","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Tai\u2010De","family":"Li","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Carl C","family":"Hayden","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-1890-5364","authenticated-orcid":false,"given":"Daniel A","family":"Fletcher","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-0871-5479","authenticated-orcid":false,"given":"R Dyche","family":"Mullins","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"297","published-online":{"date-parts":[[2017,11,15]]},"reference":[{"key":"BFEMBJ201797039_CR1","doi-asserted-by":"crossref","first-page":"841","DOI":"10.1016\/j.cell.2008.04.011","volume":"133","author":"O Akin","year":"2008","unstructured":"Akin O, Mullins RD (2008) Capping protein increases the rate of actin\u2010based motility by promoting filament nucleation by the Arp2\/3 complex. 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