Experiments, measurements, and mathematical modeling to decipher time signals in development.

Good progress has been made in identifying key signaling molecules and explaining how they are used to generate spatial patterns during embryonic development. In contrast, little is known about the control of timing or how cells use time signals in the developing embryo. In this review, I describe how direct measurements from the embryo combined with mathematical modeling could bring new insights. To illustrate this point, I discuss three examples: the Dpp gradient during growth of the Drosophila wing imaginal disc; the Polycomb-based epigenetic silencing during vernalization in plants; and the Notch-dependent somite segmentation clock.