Background adaptation by Xenopus laevis: a model for studying neuronal information processing in the pituitary pars intermedia.

This review is concerned with recent literature on the neural control of the pituitary pars intermedia of the amphibian Xenopus laevis. This aquatic toad adapts skin colour to the light intensity of its environment, by releasing the proopiomelanocortin (POMC)-derived peptide alpha-MSH (alpha-melanophore-stimulating hormone) from melanotrope cells. The activity of these cells is controlled by brain centers of which the hypothalamic suprachiasmatic and magnocellular nuclei, respectively, inhibit and stimulate both biosynthesis and release of alpha-MSH. The suprachiasmatic nucleus secretes dopamine, GABA, and NPY from synaptic terminals on the melanotropes. The structure of the synapses depends on the adaptation state of the animal. The inhibitory transmitters act via cAMP. Under inhibition conditions, melanotropes actively export cAMP, which might have a first messenger action. The magnocellular nucleus produces CRH and TRH. CRH, acting via cAMP, and TRH stimulate POMC-biosynthesis and POMC-peptide release. ACh is produced by the melanotrope cell and acts in an autoexcitatory feedback on melanotrope M1 muscarinic receptors to activate secretory activity. POMC-peptide secretion is driven by oscillations of the [Ca2+]i, which are initiated by receptor-mediated stimulation of Ca2+ influx via N-type calcium channels. The hypothalamic neurotransmitters and ACh control Ca2+ oscillatory activity. The structural and functional aspects of the various neural and endocrine steps in the regulation of skin colour adaptation by Xenopus reveal a high degree of plasticity, enabling the animal to respond optimally to the external demands for physiological adaptation.