Mesopic state: cellular mechanisms involved in pre- and post-synaptic mixing of rod and cone signals

At least twice daily our retinas move between a light adapted, cone-dominated (photopic) state and a dark-adapted, color-blind and highly light-sensitive rod-dominated (scotopic) state. In between is a rather ill-defined transitional state called the mesopic state in which retinal circuits express both rod and cone signals. The mesopic state is characterized by its dynamic and fluid nature: the rod and cone signals flowing through retinal networks are continually changing. Consequently, in the mesopic state the retinal output to the brain contained in the firing patterns of the ganglion cells consists of information derived from both rod and cone signals. Morphology, physiology, and psychophysics all contributed to an understanding that the two systems are not independent but interact extensively via both pooling and mutual inhibition. This review lays down a rationale for such rod-cone interactions in the vertebrate retinas. It suggests that the important functional role of rod-cone interactions is that they shorten the duration of the mesopic state. As a result, the retina is maintained in either in the (rod-dominated) high sensitivity photon counting mode or in the second mode, which emphasizes temporal transients and spatial resolution (the cone-dominated photopic state). Experimental evidence for pre- and postsynaptic mixing of rod and cone signals in the retina of the clawed frog, Xenopus, is shown together with the preeminent neuromodulatory role of both light and dopamine in controlling interactions between rod and cone signals. Dopamine is shown to be both necessary and sufficient to mediate light adaptation in the amphibian retina.