Color Vision
Eye 1998;12 ( Pt 3b):541-7
Evolution of colour vision in vertebrates.
Bowmaker JK
Department of Visual Science, University College London, UK. j.bowmaker@ucl.ac.uk
The expression of five major families of visual pigments occurred early in vertebrae evolution, probably about 350-400 million years ago, before the separation of the major vertebrate classes. Phylogenetic analysis of opsin gene sequences suggests that the ancestral pigments were cone pigments, with rod pigments evolving last. Modern teleosts, reptiles and birds have genera that possess rods and four spectral classes of cone each representing one of the five visual pigment families. The complement of four spectrally distinct cone classes endows these species with the potential for tetrachromatic colour vision. In contrast, probably because of their nocturnal ancestry, mammals have rod-dominated retinas with colour vision reduced to a basic dichromatic system subserved by only two spectral classes of cone. It is only within primates, about 35 millions years ago, that mammals 're-evolved' a higher level of colour vision: trichromacy. This was achieved by a gene duplication within the longer-wave cone class to produce two spectrally distinct members of the same visual pigment family which, in conjunction with a short-wavelength pigment, provide the three spectral classes of cone necessary to subserve trichromacy.
Synaptic feedback, depolarization, and color opponency in cone photoreceptors.
Burkhardt DA
Department of Psychology, University of Minnesota, Minneapolis 55455.
For some 20 years, synaptic feedback from horizontal cells to cones has often been invoked, more or less convincingly, in discussions of retinal action and vision. However, feedback in cones has proved to be rather complex and difficult to study experimentally. The mechanisms and consequences of feedback are therefore still only partly understood. This review attempts to assess the knowns and unknowns. The limitations of the evidence for feedback are reviewed to support the position that unequivocal evidence still largely rests on intracellular recording from cones. Of the three distinct types of depolarization observed in cones, the graded depolarization is taken as the fundamental manifestation of feedback. The evidence for the hypothesis that GABA is the neurotransmitter for feedback appears reasonably strong but several complications will have to be resolved to make the hypothesis more secure. There is evidence that feedback contributes to aspects of light adaptation and spatiotemporal processing of visual information. The contributions seem modest in magnitude. The role of feedback in shaping the color-opponent responses of retinal neurons is evaluated with particular emphasis on pharmacological studies, spatial and temporal aspects of the response of chromatic horizontal cells, and the enigmatic nature of depolarizations in blue- and green-sensitive cones. On this and other evidence, it is suggested that feedback may impress some detectable wavelength dependency in some cones but the dominant mechanisms for color opponency probably reside beyond the photoreceptors.
The distribution and nature of colour vision among the mammals.
Jacobs GH
Department of Psychology, University of California, Santa Barbara 93106.
1. An oft-cited view, derived principally from the writings of Gordon L. Walls, is that relatively few mammalian species have a capacity for colour vision. This review has evaluated that proposition in the light of recent research on colour vision and its mechanisms in mammals. 2. To yield colour vision a retina must contain two or more spectrally discrete types of photopigment. While this is a necessary condition, it is not a sufficient one. This means, in particular, that inferences about the presence of colour vision drawn from studies of photopigments, the precursors of photopigments, or from nervous system signals must be accepted with due caution. 3. Conjoint signals from rods and cones may be exploited by mammalian nervous systems to yield behavioural discriminations consistent with the formal definition of colour vision. Many mammalian retinas are relatively cone-poor, and thus there are abundant opportunities for such rod/cone interactions. Several instances were cited in which animals having (apparently) only one type of cone photopigment succeed at colour discriminations using such a mechanism. it is suggested that the exploitation of such a mechanism may not be uncommon among mammals. 4. Based on ideas drawn from natural history, Walls (1942) proposed that the receptors and photopigments necessary to support colour vision were lost during the nocturnal phase of mammalian history and then re-acquired during the subsequent mammalian radiations. Contemporary examination of photopigment genes along with the utilization of better techniques for identifying rods and cones suggest a different view, that the earliest mammals had retinas containing some cones and two types of cone photopigment. Thus the baseline mammalian colour vision is argued to be dichromacy. 5. A consideration of the broad range of mammalian niches and activity cycles suggests that many mammals are active during photic periods that would make a colour vision capacity potentially useful. 6. A systematic survey was presented that summarized the evidence for colour vision in mammals. Indications of the presence and nature of colour vision were drawn both from direct studies of colour vision and from studies of those retinal mechanisms that are most closely associated with the possession of colour vision. Information about colour vision can be adduced for species drawn from nine mammalian orders.
Anatomical pathways for color vision in the human retina.
Kolb H
Physiology Department, University of Utah School of Medicine, Salt Lake City, UT 84108.
The major neurons and neural circuits that are involved in the transmission of color signals through the human retina to produce the color and spatially opponent P cell or midget ganglion cell responses are described. The older findings of single cone to midget bipolar connectivity is reviewed, and the single midget bipolar cell to midget ganglion cell connectivity as revealed by a recent serial section electron microscope study is described in detail. Our present knowledge concerning the discrimination of the blue-cone subtype from the other longer wavelength cones in the human at the outer plexiform layer is summarized, and our most recent findings concerning horizontal cell connectivity to the different spectral types of cones are discussed. Finally, a hypothetical pathway is proposed for color-opponent surrounds of midget ganglion cells using both horizontal cells at the outer plexiform layer and amacrine cell pathways at the inner plexiform layer.