Orientation of Cell Polarity by Chemical Gradients

Accurate decoding of spatial chemical landscapes is critical for many cell functions. Eukaryotic cells decode local chemical gradients to orient growth or movement in productive directions. Recent work on yeast model systems, whose gradient sensing pathways display much less complexity than those in animal cells, has suggested new paradigms for how these very small cells successfully exploit information in noisy and dynamic pheromone gradients to identify their mates. Pheromone receptors regulate a polarity circuit centered on the conserved Rho-family GTPase, Cdc42. The polarity circuit contains both positive and negative feedback pathways, allowing spontaneous symmetry breaking and also polarity site disassembly and relocation. Cdc42 orients the actin cytoskeleton, leading to focused vesicle traffic that promotes movement of the polarity site and also reshapes the cortical distribution of receptors at the cell surface. In this article, we review the advances from work on yeasts and compare them with the excitable signaling pathways that have been revealed in chemotactic animal cells.