Arp2/3 Complex and Actin Depolymerizing Factor/Cofilin in Dendritic Organization and Treadmilling of Actin Filament Array in Lamellipodia

doi:10.1083/jcb.145.5.1009

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© The Rockefeller University Press, 0021-9525/1999//1009 $5.00
The Journal of Cell Biology, Volume 145, Number 5, , 1999 1009-1026

Regular Articles

Arp2/3 Complex and Actin Depolymerizing Factor/Cofilin in Dendritic Organization and Treadmilling of Actin Filament Array in Lamellipodia

Tatyana M. Svitkina and Gary G. Borisy

Laboratory of Molecular Biology, University of Wisconsin, Madison, Wisconsin 53706

The leading edge ( $~$ 1 µm) of lamellipodia in Xenopus laeviskeratocytes and fibroblasts was shown to have an extensivelybranched organization of actin filaments, which we term thedendritic brush. Pointed ends of individual filaments werelocated at Y-junctions, where the Arp2/3 complex was also localized,suggesting a role of the Arp2/3 complex in branch formation. Differential depolymerization experiments suggested that theArp2/3 complex also provided protection of pointed ends fromdepolymerization. Actin depolymerizing factor (ADF)/cofilinwas excluded from the distal 0.4 µm of the lamellipodialnetwork of keratocytes and in fibroblasts it was located withinthe depolymerization-resistant zone. These results suggest thatADF/cofilin, per se, is not sufficient for actin brush depolymerizationand a regulatory step is required. Our evidence supports adendritic nucleation model (Mullins, R.D., J.A. Heuser, andT.D. Pollard. 1998. Proc. Natl. Acad. Sci. USA. 95:6181–6186)for lamellipodial protrusion, which involves treadmilling ofa branched actin array instead of treadmilling of individualfilaments. In this model, Arp2/3 complex and ADF/cofilin haveantagonistic activities. Arp2/3 complex is responsible for integrationof nascent actin filaments into the actin network at the cellfront and stabilizing pointed ends from depolymerization, whileADF/cofilin promotes filament disassembly at the rear of thebrush, presumably by pointed end depolymerization after dissociationof the Arp2/3 complex.

Key Words: actin • Arp2/3 complex • actin depolymerizing factor/cofilin • locomotion • treadmilling

Abbreviations used in this paper: ADF, actin depolymerizing factor; CD, cytochalasin D; LA, latrunculin A; PEG, polyethylene glycol; XAC, Xenopus ADF/cofilin.

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