Morphological color changes in fish: regulation of pigment cell density and morphology

Pigment cells enable fish to change their coloration. It has been recognized that fish color changes can be divided into two categories; one is a physiological color change, which is attributed to rapid motile responses of chromatophores, and the other is a morphological color change, which results from changes in the morphology and density of chromatophores. Long-term adaptation of fish to a certain background can be a general cue to morphological color changes, and has been studied from the beginning of the 19th century. Although the motile mechanism and its control in fish chromatophores are now being elucidated, it is not yet clear how chromatophores change their density and what controls morphological color changes. In recent years, chromatophores, especially melanophores, have been shown to differentiate and to die by apoptosis under the influence of factors that regulate motile responses. Those factors are likely to utilize common intracellular signaling pathways used in part to regulate both types of color changes. In this article, after briefly reviewing the history of early studies, recent findings are discussed relevant to increases or decreases in chromatophores, and changes in their morphology. Finally, morphological color changes are discussed as physiological phenomena involved in the balance between differentiation and apoptosis of chromatophores.