Cellular signalling in vertebrate pigment cells.

Chromatophores are specialized integumental stellate cells that synthesize and store pigments. Pigment granules are translocated within chromatophores of poikilothermic vertebrates and crustaceans in response to photic, thermal and/or neurohormonal stimuli, allowing the animal to rapidly change color for thermoregulation, adaptation to light and background, and social behavior display. Birds and mammals do not show color changes, but may present slow long-term responses, such as melanocyte proliferation, melanin synthesis and melanin granule translocation into feathers, hair and surrounding keratinocytes. Pigment translocation in lower vertebrates as well as pigment production in all vertebrates are modulated by a variety of hormones and neurotransmitters acting on transmembrane receptors located on the cell surface. Alpha-melanocyte-stimulating hormone (alpha-MSH), melanin-concentrating hormone (MCHA), melatonin and catecholamines are the most important pigment cell agonist in vertebrates. The major signalling pathway leading to pigment dispersion and melanin synthesis appears to be involve stimulation of adenylate cyclase followed by an increase in the cAMP level and activation of cAMP-dependent protein kinases (PKAs). Another melanogenesis-related intracellular pathway involves the activation of protein kinase C (PKC) by diacylglycerol, and the increase in cytosolic Ca2+ by inositol triphosphate. Growth factors such as basic fibroblast growth factor (bFGF), hepatocyte growth factor (HGF) and mast cell growth factor (MGF or KIT ligand), and UV radiation modulate the melanogenic and mitogenic processes in vertebrate melanocytes as well.