Cells crawl in response to external stimuli by extending and remodelin
g peripheral elastic lamellae in the direction of locomotion. The remo
deling requires vectorial assembly of actin subunits into linear polym
ers at the lamella's leading edge and the crosslinking of the filament
s by bifunctional gelation proteins. The disassembly of the crosslinke
d filaments into short fragments or monomeric subunits away from the l
eading edge supplies components for the actin assembly reactions that
drive protrusion. Cellular proteins that respond to lipid and ionic si
gnals elicited by sensory cues escort actin through this cycle in whic
h filaments are assembled, crosslinked, and disassembled. One class of
myosin molecules may contribute to crawling by guiding sensory recept
ors to the cell surface, and another class may contribute by imposing
contractile forces on actin networks in the lamellae.