Simple Photoreceptors In Some Invertebrates: Physiological Properties Of A New Photosensory Modality

Simple photoreceptors, namely photoresponsive neurons without microvilli and/or cilia have long been known in the central ganglion of crayfish, Aplysia, Onchidium and Helix. Recently, similar simple photoreceptors, ipRGCs were discovered in the mammalian retinas. A characteristic common to all of their photoreceptor potentials shows a slow kinetics and little adaptation, contrasting with the fast and adaptive photoresponses in eye photoreceptors. Furthermore, these simple photoreceptors are not only first-order photosensory cells, but also second-order interneurons. Such characteristics suggested that simple photoreceptors function as a new sensory modality, non-image-forming vision, which is different from the image-forming vision of eye photoreceptors. The Onchidium simple photoreceptors A-P-1 and Es-1 respond to light with a depolarizing receptor potential, caused by closing of light-dependent, cGMP-gated K+ channels, as in vertebrate cGMP cascade mediated by Gt-type G-protein. The same simple photoreceptors Ip-2 and Ip-1 are hyperpolarized by light, owing to opening of the same K+ channels. This shows the first demonstration of a new type of cGMP cascade, in which Ip-2/Ip-1 are hyperpolarized when light activates guanylate cyclase (GC) through a Go-type G-protein. The ipRGCs, as involved in non-imaging function of ipRGCs, contribute to pupillary light reflex and circadian clocks. However, their function as interneurons has not been ascertained. In Onchidium simple photoreceptors, A-P-1/Es-1 and Ip-2/Ip-1 cells the photoreceptor potentials play a role in LTP-like long-lasting potentiation (LLP) of the non-imaging functions, e.g., excitatory tactile or inhibitory pressure synaptic transmission and the subsequent behavioral responses. It was also shown that this LLP is effective, even if their photoresponse is subthreshold. (C) 2008 Elsevier B.V. All rights reserved.