The ecology of visual pigments.

The visual systems of vertebrates have adapted to function in photic environments ranging from the broad spectrum of full sunlight to almost total darkness, including the restricted spectral ranges found in different coloured aquatic environments. Such adaptations are immediately obvious at the level of retinal photoreceptors. The basic vertebrate photoreceptor pattern consists of rods and four spectrally distinct classes of cone that span the spectrum from the near ultraviolet to the far red. This arrangement is found in many diurnal species including shallow-living teleosts, reptiles and birds, but is noticeably absent in mammals. In freshwater teleosts the visual pigments may be porphyropsins which have maximum sensitivities displaced to longer wavelengths than their equivalent rhodopsins. Water acts as a monochromator, so that with increasing depth the spectral range of the ambient light is restricted, primarily at long wavelengths. Therefore, at depth the down-welling daylight is not only attenuated in intensity, but is restricted to a narrow spectral band centred around 470 nm. Closely related species that live at increasing depths show a loss of long-wave-sensitive cones and a displacement of the maximum sensitivities of middle-wave-sensitive cones and rods to shorter wavelengths. Such species offer a natural model for determining specific amino acids in opsin responsible for the spectral tuning of these middle-wave-sensitive pigments.