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Control of the cytoskeleton and mechanical contacts with the extracellularenvironment are essential component of motility in eukaryotic cells. In theabsence of signals, cells continuously rebuild the cytoskeleton andperiodically extend pseudopods or other protrusions at random membranelocations. Extracellular signals bias the direction of movement by biasing theextension of protrusions, but this involves another layer of biochemicalnetworks for signal detection, transduction, and control of the rebuilding ofthe cytoskeleton. Here we develop a model for the latter processes that centerson a Ras-based module that adapts to constant extracellular signals andcontrols the downstream PI3K-PIP3-based module responsible for amplifying aspatial gradient of the signal. The resulting spatial gradient can lead topolarization, which enables cells to move in the preferred direction (upgradient for attractants and down-gradient for repellents). We show that themodel can replicate many of the observed characteristics of the responses tocAMP stimulation for Dictyostelium, and analyze how cell geometry and signalinginteract to produce the observed localization of some of the key components ofthe amplification module. We show how polarization can emerge withoutdirectional cues, and how it interacts with directional signals and leads todirectional persistence. Since other cells such as neutrophils use similarpathways, the model is a generic one for a large class of eukaryotic cells.

Signal transduction and directional sensing in eukaryotes