How animals get their skin patterns: fish pigment pattern as a live Turing wave

It is more than fifty years since Alan Turing first presented the reaction-diffusion (RD) model, to account for the mechanism of biological pattern formation. In the paper entitled The chemical basis of morphogenesis, Turing concluded that spatial patterns autonomously made in the embryo are generated as the stationary wave of the chemical (cellular) reactions. Although this novel idea was paid little attention by experimental biologists, recent experimental data are suggesting that the RD mechanism really functions in some of the course of animal development. Among the phenomena in which involvement of the RD mechanism is suspected, the striped pigment pattern of zebrafish has been highlighted as an ideal model system for the following reasons: the stationary wave made by the RD mechanism stays alive and can be observed only in the fish skin; and in zebrafish, we can utilize genomic information and molecular genetic techniques to clarify the molecular basis of pattern formation. In this review, we summarize recent progresses in the study of zebrafish pigment pattern formation that is uncovering how the RD wave is made and maintained in the skin.