Calcium and light adaptation in retinal rods and cones

Retinal rods and cones respond to light with a membrane hyperpolarization. This hyperpolarization is mediated by an ionic conductance (the light-regulated conductance) that is kept open in darkness by cyclic GMP acting as a ligand, and which closes in the light as a result of an increase in cGMP hydrolysis triggered by illumination. Calcium ions appear to have a role in this phototransduction process: they provide negative feedback between the conductance, which is permeable to Ca2+ (refs 4, 5), and the concentration of cGMP, which is sensitive to Ca2+ (refs 6-8). This feedback down-regulates the sensitivity to light of a photoreceptor and probably contributes to the important phenomenon of light adaptation in vision. It is still not clear, however, how much of the light adaptation is actually attributable to this Ca2+ feedback. We have examined the responses of amphibian rods and cones to light with the Ca2+ feedback removed. Normally, the response of a cell to a step of light rises transiently to a peak, but rapidly relaxes to a lower level, indicative of light adaptation. When the feedback is removed, however, the relaxation of the response is completely absent; furthermore, the steady response levels at different light-step intensities are well predicted by a statistical superposition of invariant single-photon responses. We therefore conclude that the Ca2+ feedback underlies essentially all light adaptation in rods and cones.