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Fat2polarizestheWAVEcomplexintranstoalign1cellprotrusionsforcollectivemigration2AudreyM.Williams1,SethDonoughe1,EdwinMunro1,2,3,SallyHorne-Badovinac1,2,∗34Abstract5Foragroupofcellstomigratetogether,eachcellmustcouplethepolarityofitsmigratorymachinerywiththat6oftheothercellsinthecohort.Althoughcollectivecellmigrationsarecommoninanimaldevelopment,littleis7knownabouthowprotrusionsarecoherentlypolarizedamonggroupsofmigratingepithelialcells.Weaddress8thisprobleminthecollectivemigrationofthefollicularepithelialcellsinDrosophilamelanogaster.Inthisepithe-9lium,thecadherinFat2localizestothetrailingedgeofeachcellandpromotestheformationoflamellipodiaat10theleadingedgeofthecellbehind.WeshowthatFat2performsthisfunctionbyactingintranstorestrictWAVE11complexactivitytoonelong-livedregionalongeachcell’sleadingedge.WithoutFat2,theWAVEcomplexdistri-12butionexpandsaroundthecellperimeterandfluctuatesovertime,resultinginreduced,unpolarizedprotrusive13activity.WefurthershowthatFat2’sinfluenceisverylocal,with...