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Colour vision in blue-cone 'monochromacy'.

1. Atypical (blue cone) monochromats show two action spectra when tested by the increment threshold method of Stiles with ;central' fixation. One spectrum peaks near 450 nm and has the spectral characteristics of normal blue cones. The other resembles rhodopsin (pi(0)) modified slightly by photostable macular pigment.2. Under some circumstances such observers are dichromats. There is a neutral point (matched to Illuminant ;C') in the neighbourhood of 460-470 nm.3. The spectral colour matching functions using two primaries have been measured on three such subjects. They can be fit reasonably well, although imperfectly, by linear combinations of pi(0) and pi(1). The chromaticity co-ordinates have been calculated according to the convention of W. D. Wright and compared to the results predicted from pi(0) and pi(1). The comparison suggests that part of the imperfections of the colour matching function fit is due to prereceptor differences (e.g. macular pigment) between the blue-cone monochromats and the hypothetical pi(0), pi(1) observer.4. Colour matches made at high light levels continue to hold when the intensity of the field is reduced below the cone threshold.5. Therefore one of the visual pigments participating in the colour matches has an action spectrum which is not measurably different from that of the rod pigment rhodopsin.6. Increment threshold measurements show that the mechanism which has the rhodopsin action spectrum has the directional sensitivity of cones, not rods.7. Blue test threshold during dark adaptation after a full bleach follow a bipartite exponential recovery curve. The first exponential has a time constant of 1 min, the second 2 min. By comparing these curves to the increment thresholds it is possible to relate the first to the pi(1), the second to the pi(0) cones.8. Using a broad band blue gelatin filter in the measuring light of the retinal densitometer and studying the same retinal region tested in 7 it is possible to follow the regeneration of a pigment after a full bleach which has an exponential recovery with a time constant of 1.0 min. With a yellow green filter in the measuring light the exponential recovery observed after a full bleach has a time constant of 2.0 min.9. One of the two visual pigments participating in the colour matches resides in receptors which have the action spectrum, the directional sensitivity and probably the dark adaptation curve of normal blue cones.10. The other resides in receptors which have the action spectrum of normal rods but the directional sensitivity and the dark adaptation curve of normal red and green cones.

Color Perception↗

Psychophysical estimates of visual pigment densities in red-green dichromats.

1. The spectral sensitivity of red-green dichromats was determined using heterochromatic flicker photometric matches (25-30 c/s) on the fovea. These matches are upset after a bright bleach and consequently the spectral sensitivity is altered.2. Preliminary experiments indicate that under the conditions in which these experiments were performed, the blue cone mechanism of deuteranopes and protanopes cannot follow 20 c/s flicker. If dichromats lack one of the normal pigments then the upset of these matches monitors the change in spectral sensitivity of a single mechanism.3. After a bleach which removes all the cone pigments, the spectral sensitivity recovers with the time course of pigment kinetics as measured by densitometry.4. An intense background also changes the relative spectral sensitivity of the dichromats. On real equilibrium backgrounds, the changes in spectral sensitivity follow those predicted by the pigment changes measured by densitometry. The predicted changes are obtained by modifying the Rushton equilibrium equation to take into account the density of pigment.5. The relationship of these changes to the luminance of the background is independent of the colour of the background light.6. In contradistinction the effect is dependent on the colour of the lights which were flickered. These experiments indicate that a narrowing of the spectral sensitivity curves takes place on both sides of the dichromats' lambda(max).7. The change in relative spectral sensitivity as a function of background intensity was also determined by increment threshold measurements. These changes can be expressed in terms of deviations from Weber's law (DeltaI/I = const.) if DeltaI and I represent the number of chromophores destroyed by the test and background.8. The relative spectral sensitivity of the dichromat was changed by decentering the point of pupil entry. This upset was abolished by bleaching. The size of the upset was correlated with the magnitude of the S-C I effect.9. Given the hypothesis of pigment density (self-screening), the results of expts. (3)-(8) are consistent and allow the calculation of a maximum optical density for those pigments which underlie the dichromats' long-wave mechanism. For the deuteranope a D(lambdamax) of 0.5-0.6 is calculated and for the protanope a D(lambdamax) of 0.4-0.5 is obtained.

Color Perception↗

Electrical responses and photopigments of twin cones in the retina of the walleye.

1. The properties of twin and single cones in the retina of the walleye (Stizostedion vitreum vitreum) were studied by intracellular recording, dye injection and microspectrophotometry. 2. Twin cones generate hyperpolarizing responses to central illumination, can receive depolarizing influences (feed-back) from the receptive field surround, and show no detectable dye coupling when injected with Procion yellow. In seventeen of eighteen dye-injected cones, fluorescence was intense in the inner segment and undetectable or weak in the cone pedicle. 3. Both members of the twin cone contain the same photopigment in their outer segments. It absorbs maximally at about 605 nm. 4. A 533 nm green-sensitive photopigment was found in single cones. No blue-sensitive cones have been found. 5. With the exception of a modest discrepancy in the violet, the absorptance spectrum of the 605 nm photopigment of twin cones agrees closely with the action spectrum measured by intracellular recording. 6. The spectral properties established by the twin cone's photopigment are not detectably altered by the hyperpolarizing influences arising from nearby cones or by the depolarizing influences arising from the receptive field surround. 7. The twin cones of the walleye retina are thus "identical twins', both photochemically and physiologically, and seem designed to function as long-wave, spectrally univariant receptor units for colour vision. 8. The available evidence suggests that identical twin cones differ functionally from double cones and non-identical twin cones. 9. Although they outnumber single cones by about three to one in adults, identifiable twin cones were rarely observed in the cone population of retinas examined 3-5 days after birth. If walleye twin cones develop by fusion of single cones this process apparently occurs only for cones containing the 605 nm photopigment.

Animals↗

Spectral sensitivity functions of post-receptoral responses in human vision.

1. Increment threshold illumination levels, It, for visual detection of a one-dimensional, spatially periodic test pattern (i.e. a grating) presented to the right eye have been measured following a period of adaptation to high contrast patterns. The right (test) eye was adapted to a grating, matched in bar-width and orientation to the test grating, and the left eye was adapted to a two-dimensional matrix array of circular spots. It is shown that the value of It is dependent on the spatial and spectral parameters of the spot matrix. 2. The variation of It with change in matrix wavelength, lambda, is greatest for matrix spots of some 10-15 min of arc diameter. 3. Values of It were determined as a function of the matrix wavelength, lambda. Three kinds of spectral response have been observed, each associated with a different stimulus geometry, and each characterized by the wavelength, lambda, for which It is minimum. With an adaptation matrix consisting of coloured spots on a dark surround, It is minimum for values of lambda around 600 nm, whereas with a matrix of dark spots on a coloured surround, the minimum occurs for lambda around 520 nm. The third spectral response is found when a blue test grating is superimposed on a yellow background, conditions under which detection is mediated by the blue-sensitive, pi 1-increment threshold mechanism. In this case, It is minimum for lambda equal to about 450 nm. 4. Three subjects with normal colour vision each give similar spectral responses, but the data for two deuteranopic red-green dichromats deviate significantly from the normal response pattern. 5. The binocular interaction mechanism through which the spot matrix influences threshold detection of the test grating is discussed in relation to electrophysiological and other psychophysical data.

Adaptation, Ocular↗

Color defect and color theory; studies of normal and colorblind persons, including a subject color-blind in one eye but not in the other.

It is important to find answers to two questions concerning the visual discriminations of dichromatic persons, especially deuteranopes: (i) Do such persons show a loss of sensitivity to various wavelengths of the spectrum as compared with normal subjects? (ii) What colors do they see? A number of experiments were performed on the first question. First, luminosity curves were determined on three groups of subjects, consisting respectively of five protanopes, six deuteranopes, and seven normal individuals. As compared with normal subjects, protanopes show a loss of luminosity in the red, whereas deuteranopes show a loss in the blue-to-green region of the spectrum (See 10). Second, we examined the luminosity curves of a subject whose right eye is classifiable (on the basis of color-mixture determinations) as normal and whose left eye is classifiable as dichromatic. (The hue discrimination curve for her dichromatic eye seemed comparable to the curve of the usual deuteranope except in the violet, where it manifested relatively good discrimination.) The luminosity function for this subject's dichromatic eye, determined by data on threshold and flicker, exhibits the same type of luminosity loss in the blue and green regions of the spectrum as was shown by our group of six deuteranopes. Only unilaterally dichromatic subjects can tell us how colors seen by a dichromatic eye appear to a normal eye. In the color-blind eye, our unilaterally dichromatic subject sees wavelengths below and above her neutral ("grey") point (which occurs at 502 mmicro) as, respectively, a blue equivalent to about 470 mmicro and a yellow equivalent to about 570 mmicro in her normal eye. The results on (i) luminosity loss and (ii) the seeing of wavelengths above 502 mmicro as yellow are considered theoretically. The seeing of yellow by deuteranopes and protanopes may be accounted for by an idea based on Leber-Fick transmation theory. It is proposed that the characteristic sensitivities of the red and green receptors become similar while no change takes place in their central brain connections. Losses may be introduced into the transformed sensitivity curves to indicate appropriate degrees of luminosity deficit for deuteranopes and protanopes.

Color↗

Changes in the perceived color of very bright stimuli.

When very intense stimuli in the long-wavelength region of the visual spectrum are viewed continuously, they change in hue from red, through yellow, to green. The relation of the time course of these changes to the intensity of the stimulus is reported.

Biological Phenomena↗

Visual pigment gene structure and the severity of color vision defects.

Rearrangements of the visual pigment genes are associated with defective color vision and with differences between types of red-green color blindness. Among individuals within the most common category of defective color vision, deuteranomaly, there is a large variation in the severity of color vision loss. An examination of specific photopigment gene sites responsible for tuning photopigment absorption spectra revealed differences that predict these variations in the color defect. The results indicate that the severity of the defect in deuteranomalous color vision depends on the degree of similarity among the residual photopigments that serve vision in the color-anomalous eye.

Blotting, Southern↗

Molecular genetics of inherited variation in human color vision.

The hypothesis that red-green "color blindness" is caused by alterations in the genes encoding red and green visual pigments has been tested and shown to be correct. Genomic DNA's from 25 males with various red-green color vision deficiencies were analyzed by Southern blot hybridization with the cloned red and green pigment genes as probes. The observed genotypes appear to result from unequal recombination or gene conversion (or both). Together with chromosome mapping experiments, these data identify each of the cloned human visual pigment genes.

Animals↗

Visual sensitivity: significant within-species variations in a nonhuman primate.

Among squirrel monkeys (Saimiri sciureus) there are significant sex-related differences in visual sensitivity. As measured behaviorally in an increment-thershold task, a sample of males was found to be substantially less sensitive to long-wavelength (640-nanometer) light than a group of females tested in the same way, although the two groups showed no significant differences in sensitivity to a middle-wavelength (540-nanometer) light. The two group also differed on a test designed to measure the effects of chromatic adaptation.

Animals↗

Psychophysical evidence for more than two kinds of cone in dichromatic color blindness.

Psychophysical evidence shows that at least some classically diagnosed dichromats have three cone types rather than two. The anomalous cones, previously thought to be absent, are less sensitive than normal cones to both spectral and temporal variations, and have spectral sensitivities like those of the abnormal cones of anomalous trichromats. These results are not consistent with either loss or replacement models of X-linked recessive color-vision defects, since some dichromats apparently have the same three photopigments as anomalous trichromats.

Color Perception↗