The Limulus ventral photoreceptor: light response and the role of calcium in a classic preparation

The ventral nerve photoreceptor of the horseshoe crab Limulus polyphemus has been used for many years to investigate basic mechanisms of invertebrate phototransduction. The activation of rhodopsin leads in visual cells of invertebrates to an enzyme cascade at the end of which ion channels in the plasma membrane are transiently opened. This allows an influx of cations resulting in a depolarization of the photoreceptor cell. The receptor current of the Limulus ventral photoreceptor consists of three components which differ in several aspects, such as the time course of activation, the time course of recovery from light adaptation, and the reversal potential. Each component is influenced in a different, characteristic way by various pharmacological manipulations. In addition, at least two types of single photon-evoked events (bumps) and three elementary channel conductances are observed in this photoreceptor cell. These findings suggest that the receptor current components are controlled by three different light-activated enzymatic pathways using three different ligands to increase membrane conductance. Probably one of these ligands is cyclic GMP, another one is activated via the IP3-cascade and calcium, the third one might be cyclic AMP. Calcium ions are very important for the excitation and adaptation of visual cells in invertebrates. The extracellular and intracellular calcium concentrations determine the functional state of the visual cell. A rise in the cytosolic calcium concentration appears to be an essential step in the excitatory transduction cascade. Cytosolic calcium is the major intracellular mediator of adaptation. If the cytosolic calcium level exceeds a certain threshold value after exposure to light it causes the desensitization of the visual cell. On the other hand, from a slight rise in cytosolic calcium facilitation results, i.e. increased sensitivity of the photoreceptor.