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ell crawling is thought to be the result of three co- ordinated motility behaviors: ( a ) protrusion and adhesion of the front end, probably driven by ac- tin assembly; ( b ) traction force that leads to the advance of the nucleus and bulk cytoplasm; and ( c ) release and retrac- tion of the tail in most cell types. (In neurons, the last step is highly modified, and an axon elaborates from behind the advancing growth cone.) As Mitchison and Cramer (15) point out, it is the traction step that is least understood and seems most central to productive locomotion. Two papers in JCB now shed new light, both mechanical and molecu- lar, on the traction step of cell crawling. A report in this is- sue of JCB on growth cone movements from the Forscher group (19) and a paper from the Borisy lab (20) on kerato- cyte motility provide new insights into the role of myosin in traction. Specifically, the two papers clarify the mechan- ical connections to the motor that produce the motile out- put of various animal cells. These papers also provide in- sight into how a conserved mechanism for animal cell crawling could underlie visually different crawling behav- ior. Indeed, the cell types used in the new studies are at two extremes of cell crawling behavior. Neuronal growth cones change shape markedly during their inconstant, stop-and-go advance, which is accompanied by a variety of seemingly futile motile behaviors such as waving of, ex-

Cell Crawling: First the Motor, Now the Transmission