Light adaptation and sensitivity controlling mechanisms in vertebrate photoreceptors.

The human visual system can discriminate increment and decrement light stimuli over a wide range of ambient illumination; from moonlight to bright sunlight. Several mechanisms contribute to this property but the major ones reside in the retina and more specifically within the photoreceptors themselves. Numerous studies in retinae from cold- and warm-blooded vertebrates have demonstrated the ability of the photoreceptors to respond in a graded manner to light increments and decrements even if these are applied during a background illumination that is expected to saturate the cells. In all photoreceptors regardless of type and species, three cellular mechanisms have been identified that contribute to background desensitization and light adaptation. These gain controlling mechanisms include; response-compression due to the non-linearity of the intensity-response function, biochemical modulation of the phototransduction process and pigment bleaching. The overall ability of a photoreceptor to adapt to background lights reflects the relative contribution of each of these mechanisms and the light intensity range over which they operate. In rods of most species, response-compression tends to dominate these mechanisms at light levels too weak to cause significant pigment bleaching and therefore, rods exhibit saturation. In contrast, cones are characterized by powerful background-induced modulation of the phototransduction process at moderate to bright background intensities where pigment bleaching becomes significant.Therefore, cones do not exhibit saturation even when the level of ambient illumination is raised by 6-7 log units.