Getting the actin filaments straight: nucleation-release or treadmilling?

The dynamic turnover of actin filaments plays a central role in the locomotion of metazoan cells. Based on results obtained with actin labelled with a caged fluorescent probe, Theriot and Mitchison proposed a 'nucleation-release' model for the fast-moving fish keratocyte, which predicts the existence of short non-oriented filaments in the motile lamellipodium. More recent structural data on keratocyte cytoskeletons do not support this model, but are consistent with the treadmilling of long actin filaments of graded length. Taken together with Theriot and Mitchison's demonstration that the cytoskeleton remains stationary relative to the substrate in the moving keratocyte, the structural data raise the possibility that a lateral flow of filaments plays a role in lamellipodia motility.