Understanding Pattern Formation in Embryos: Experiment, Theory, and Simulation.

Twenty-five years ago, Lewis Wolpert, the eminent developmental biologist, asked the question, "Do We Understand Development?" He concluded that such rapid progress had been made in the preceding two decades that "It is not unreasonable to think that enough will eventually be known to program a computer and simulate some aspects of development." This prediction has been fulfilled, at least partially, with data-driven simulations of several different developmental processes being developed in the intervening years. Nevertheless, the question remains of whether we "understand" development and if simulations are sufficient to provide an explanation of development. While in silico replications and models are undoubtedly an important tool in the investigation and dissection of developmental processes, which complement traditional experimental methods, these need to be supplemented by theory that identifies principles and provides coherent explanations. Here, I use the example of pattern formation in the vertebrate neural tube to illustrate this idea.