The crawling movement of a cell involves protrusion of its leading edge coordinated with translocation of its cell body and depends upon a cytoplasmic machinery able to respond to signals from the environment. Protrusion is now understood to be driven by actin polymerization and signalling from membrane receptors to actin has been shown to be mediated by the Rho family of GTPases. However, a major gap in our understanding of regulated motility has been to connect the signalling pathway to the motile machinery itself. Recent structural, biochemical and genetic studies have identified some of the missing links and provided a strong working model for the pathways and mechanism by which the signals are interpreted and implemented.