Beginning and ending an actin filament: control at the barbed end.

Dynamic actin filaments contribute to cell migration, organelle movements, memory, and gene regulation. These dynamic processes are often regulated by extracellular and?or cell cycle signals. Regulation targets, not actin itself, but the factors that determine it's dynamic properties. Thus, filament nucleation, rate and duration of elongation, and depolymerization are each controlled with regard to time and?or space. Two mechanisms exist for nucleating filaments de novo, the Arp23 complex and the formins; multiple pathways regulate each. A new filament elongates rapidly but transiently before its barbed end is capped. Rapid capping allows the cell to maintain fine temporal and spatial control over F-actin distribution. Modulation of capping protein activity and its access to barbed ends is emerging as a site of local regulation. Finally, to maintain a steady state filaments must depolymerize. Depolymerization can limit the rate of new filament nucleation and elongation. The activity of ADF?cofilin, which facilitates depolymerization, is also regulated by multiple inputs. This chapter describes (1) mechanism and regulation of new filament formation, (2) mechanism of enhancing elongation at barbed ends, (3) capping proteins and their regulators, and (4) recycling of actin monomers from filamentous actin (F-actin) back to globular actin (G-actin).