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Role of the blue mechanism in wavelength discrimination.

The role of blue cones as well as the pathways they supply (collectively called the "blue mechanism") is evaluated by comparing ordinary wavelength discrimination functions with those obtained using two methods designed to inhibit the blue mechanism selectively. These methods use a just-noticeable-border criterion (JNB), instead of the usual one of just-noticeable-difference, and a yellow preadapting field to induce transient tritanopia. Without transient tritanopia, the data obtained using the just-noticeable-border criterion reveal a small contribution of the blue mechanism to wavelength discrimination. Transient tritanopia, with JNB, produces an additional selective loss of wavelength discrimination in a spectral region flanking 460 nm, which yields a function resembling those for tritanopes previously examined.

Color Perception↗

Red-green mixture thresholds in congenital and acquired color defects.

A color television display was used to measure thresholds for mixtures of red and green on a white background; red and green components could be either incremental, decremental or zero. Ellipses are fitted to a plot of green contrast as a function of red contrast, and it is argued that the length of the ellipse is a measure of red-green color discrimination and the width of the ellipse is a measure of luminance discrimination. It is shown that the technique reliably distinguishes normals from congenital color defectives and also protan from deutan subjects. For some cases of acquired color defects (e.g. optic neuritis), there is a roughly equal loss of color and luminance discrimination whereas, in other cases (e.g., hereditary optic atrophies), the loss of color discrimination is much greater than the loss of luminance discrimination.

Adult↗

Equilibrium hue judgements of dichromats.

It is generally held that protanopes and deuteranopes see only regions of blues and yellows in the visible spectrum, with an achromatic point, called the neutral point, separating these regions. Considerations of a zone model of color vision for the dichromatic observer led us to predict that a reduced form of red/green discrimination would allow equilibrium blue judgements to be made by protanopes. We show that protanopes can make equilibrium blue determinations with as much reliability as they make neutral point settings. Our results indicate that protanopes but not deuteranopes are able to rely on a reduced form of red/green discrimination in the short wavelength part of the spectrum. Protanopes describe wavelengths longer than the neutral point as yellow. Between the neutral point and equilibrium blue, different wavelengths are described as having varying aspects of blue and green; and short of equilibrium blue they appear reddish blue. For dueteranopes, the spectrum longer than the neutral point appears yellow, and short of it, blue. The results of our experiments showing that the protanopic equilibrium blue is invariant with intensity variations, as it is in the trichromat, add support to the idea of a reduced form of red/green discrimination for protanopes. Our results also allow the evaluation of various models of protanopia.

Color Perception↗

Color discrimination and neural coding in color deficients.

Rayleigh color match ranges obtained from color deficient observers varied considerably as a function of spatial and temporal parameters of stimulus presentation. The results suggest that color discrimination losses in color deficients result from abnormalities in the spatial and temporal properties of neural coding in addition to cone photopigment abnormalities.

Color Perception↗

Differences in the responses of heterozygous carriers of colorblindness and normal controls to briefly presented stimuli.

Two experiments were conducted in order to investigate the possible effects of X-inactivation (Lyon, 1961) on female carriers of colorblindness. The results of the first experiment, like those of Grützner et al. (1976), were consistent with the prediction of the Lyon (1961) hypothesis that the retinas of female carriers are composed of mosaic patches of colorblind and normal areas. In this first experiment, rows and columns of colored spots were presented tachistoscopically, and subjects were asked to identify the colors of the spots. In the second experiment, plates from the Ishihara test of colorblindness were presented tachistoscopically and subjects were asked to identify the number which was embedded in the pattern of colored dots. Both experiments support the Lyon hypothesis in that female carriers were found to have more difficulty in perceiving patterns of colored stimuli than did control subjects, and they suggest that the amount of time that a carrier has to scan colored stimuli plays an important role in her ability to accurately perceive them.

Color Perception↗

Discrimination of luminance and chromaticity differences by dichromatic and trichromatic monkeys.

Dichromatic and trichromatic representatives from two genera of platyrrhine monkeys that show widespread color vision polymorphism (Saguinus--tamarins, Saimiri--squirrel monkeys) were tested for their abilities to make increment-threshold and flicker discriminations based on luminance and chromaticity differences. The details of these tests were arranged to attempt to emphasize the relative contributions to visual behavior of non-opponent and spectrally-opponent neural mechanisms. The results indicate that dichromatic and trichromatic monkeys differ only trivially on tests where performance is based on the contributions of non-opponent mechanisms, that the contribution of spectrally opponent mechanisms to the "brightness signal" is very similar in trichromatic and dichromatic monkeys, and that in increment-threshold discriminations where there are both chromaticity and luminance cues some test wavelengths yield superior performance for trichromats while others appear to favor the dichromat.

Animals↗

Color vision following intense green light exposure: data and a model.

Hue discrimination, spectral sensitivity, and mathematical models of both are presented for a rhesus monkey which was exposed to intense green light. One of the monkey's eyes was blue-blinded in a previous experimental procedure and the other was color normal. The results of green light exposure showed a loss of sensitivity on both measures, with greater loss in the blue-blinded eye. Although there was considerable loss of hue-discrimination in the blue-green spectral regions, hue-discrimination at the point of best discrimination, 590 nm, remained unaffected. This pattern of results poses difficulties for models of hue discrimination, and has resulted in the proposed model employing three opponent color channels. The number of free-parameters are minimized and the integration between spectral sensitivity and hue discrimination enhanced by deriving parameters used in modeling hue discrimination from spectral sensitivity or vice versa.

Animals↗

Induced color blindness in goldfish: a behavioral and electrophysiological study.

To answer the question whether, like man, ethambutol treated fish would become color-blind, wavelength discrimination was measured behaviorally in goldfish, preceding, during and after ethambutol treatment. The results are that of the three high discrimination abilities at around 400, 500 and 600 nm, ethambutol affected the latter one. Red-green discrimination is lost reversibly leaving the discriminations in the blue-green and violet range unaffected. This red-green discrimination deficiency cannot be accounted for by a loss of long wavelength cones since the ERG and luminosity functions remain unaffected. Intracellular horizontal cell recordings in goldfish show that ethambutol hyperpolarizes all three types of cone driven horizontal cells and changes their color coding such that their spectral characteristics become cone-like as is the case in dark adapted retina. So, the initial effect induced by ethambutol seems to be an adaptation deficiency in color vision related tasks. Human wavelength discrimination and increment threshold spectral sensitivity functions obtained at low luminance levels are compared to behavioral functions in ethambutol treated goldfish. The high similarity between the ethambutol effects in man and goldfish, and the effects observed in the horizontal cell responses in goldfish are highly indicative that horizontal cells play a key role in color vision. So far their function has been puzzling.

Animals↗

Bicuculline produces reversible red-green color blindness in goldfish, as revealed by monocular behavioral testing.

The effect of the GABAA receptor antagonist bicuculline (methiodide) on goldfish wavelength discrimination was studied. This was done using a behavioral two alternative forced choice procedure at training wavelengths 500 and 600 nm, where goldfish wavelength discrimination is the best. During the experiments the goldfish could use only the eye that was intravitreally injected with bicuculline; the other eye was covered. In control experiments, to exclude systemic effects, the covering of the eyes was reversed. Bicuculline induced loss of wavelength discrimination ability around 600 nm, while this ability was not affected around 500 nm. The effect was reversible, since discrimination around 600 nm returned to normal within a day after injection. The results indicate that GABAA receptor mediated processes, like horizontal cell to cone feedback, play an important role in wavelength discrimination at the long wavelength part of the spectrum.

Animals↗

Brightness, discriminability and the "crispening effect".

Subjects adjusted the luminances, L, of 16 or 25 circles, all visible at the same time on a computer monitor, to make equal-interval brightness series. The background was black, white or grey. The luminance steps between adjacent circles behaved like the luminance discrimination thresholds of Whittle, P. [(1986) Vision Research, 26, 1677-1691)]. They showed a sharp minimum at the background luminance, Lb: the "Crispening Effect". They followed Weber's Law with respect to L when L was small, but with respect to delta L (= magnitude of L-Lb) near Lb. The Crispening Effect was abolished by a thin outline or a hue difference between circles and background.

Color Perception↗

Retinal and cortical activity in human subjects during color flicker fusion.

Pattern electroretinograms (PERG) and cortical visually evoked potentials (VEP) were simultaneously recorded from 7 visually normal and 1 protanopic subjects. Stimuli were color checkerboards (0.5 degrees check size), phase-reversing at 17 Hz (i.e. 34 reversals/sec). Using a stepwise sweep procedure, the luminance of the red (lambda peak = 550 nm) and green (lambda peak = 630 nm) checks varied in 11 steps in opposite directions from 0 to 30 cd/m2, embracing the subjective equiluminance point. For normal subjects at subjective equiluminance, the VEP amplitude dropped sharply down to 13 +/- 2% of the value at pure luminance contrast. The PERG, however, was only reduced to 56 +/- 10% at this point, an attenuation 4 times less than that of the VEP. In contrast to normal subjects, in the protanopic subject the PERG was sharply reduced at equiluminance, parallel to the VEP. This would be expected when L-cones are missing. Assuming that the PERG reflects the activity of the retinal ganglion cells, our findings suggest that human retinal ganglion cells respond well under the condition of equiluminant flicker fusion, which is in agreement with recent single-cell studies in the monkey. Consequently, the temporal low-pass filter, which mediates color-flicker fusion, would seem to lie central to the retinal ganglion cells.

Adult↗

Serine/alanine amino acid polymorphism of the L and M cone pigments: effects on Rayleigh matches among deuteranopes, protanopes and color normal observers.

In a first experiment, groups of deuteranopes and protanopes were characterized psychophysically by the slopes of regression lines fitted to yellow intensity settings from their Rayleigh matches. In a second experiment, color normal male subjects were characterized by their 2 and 8 deg Rayleigh match points. All subjects had been previously characterized genetically by the presence of the amino acid serine or alanine at position 180 on their L cone or L/M hybrid opsins. Dichromats and color normal subjects with serine had greater sensitivity to the red primary than did those with alanine. Calculations based on psychophysical results suggest that the substitution of serine by alanine in the L cone opsin or L/M hybrid opsin produces a shift in lambda max of the expressed pigment toward shorter wavelengths by an amount varying between 2.6 and 4.3 nm, with the shifts in lambda max for the dichromats being larger than those for the color normal subjects.

Adult↗

Does chromatic sensitivity develop more slowly than luminance sensitivity?

Chromatic sensitivity is very low in humans during the first few months of life. We examined whether low chromatic sensitivity reflects a deficiency among chromatic mechanisms or whether it is simply a manifestation of poor visual sensitivity in general. The sweep VEP was used to measure contrast sensitivity to gratings varying in the mixture of red and green components. For infants from 2 to 8 weeks of age, sensitivity to all mixtures was lower than color-normal adults' sensitivity, but infant and adult ratios of luminance/chromatic sensitivity were similar. This finding is consistent with the hypothesis that infants have functional MWS and LWS cones and the requisite post-receptor chromatic mechanisms to compare their signals.

Age Factors↗

Horizontal cells function normally in ethambutol-treated goldfish.

Ethambutol, a tuberculostatic drug, induces red-green colour vision defects in man and goldfish. The ethambutol-induced red-green colour vision defect in goldfish was argued to originate in the retina because after ethambutol application: (1) inhibitive interactions in red-green (double) opponent ganglion cells are lost [Van Dijk & Spekreijse, 1982 (Investigative Ophthalmology and Visual Science, 24, 128-133); Wietsma & Spekreijse, 1992 (Investigative Ophthalmology and Visual Science Suppl., 33, 1032)] and (2) the depolarizing responses to red light in the biphasic horizontal cells are reduced. To account for these findings Spekreijse, Wietsma and Neumeyer [(1991) Vision Research, 31, 551-562] suggested that ethambutol induced dark adaptation in the retina. In this paper the dark adaptation hypothesis is tested with the following results: (1) ethambutol changes only transiently the receptive field size and spectral sensitivity of horizontal cells; (2) the spectral characteristics of horizontal cells do not change in long-term ethambutol-treated goldfish; (3) formation of spinules on horizontal cell dendrites in cone terminals, a parameter for light adaptation, remains unaffected. Therefore we conclude that ethambutol does not induce functional dark adaptation of horizontal cells and that the ethambutol-induced red-green colour vision deficiency does not originate in the horizontal cell layers.

Action Potentials↗

Peripheral cone contrast sensitivity in glaucoma.

Colour vision tests for detection of glaucomatous damage frequently suffer from two problems: most tests are confined to foveal vision, whereas defects tend to appear first extrafoveally; and the modulation directions in colour space are not optimal. This paper deals with peripheral testing à la Yu, Falcao-Reis, Spileers and Arden [(1991) Investigative Ophthalmology and Visual Science, 32, 2779-2789], and investigates whether there are modulation directions that show preferential sensitivity reduction in glaucoma. In 14 eyes with early glaucoma, 17 risk eyes and 10 normals, 12 deg peripheral colour contrast thresholds were determined for L, M, S, L-M and L+M test directions. Threshold elevations were correlated in all test directions, with S modulation yielding the largest elevations.

Aged↗

Spectral sensitivity of dichromats: role of postreceptoral processes.

Increment spectral sensitivity functions were determined for dichromatic subjects; these functions were interpreted in the context of parvo and magno pathways. Increments of either 200 or 10 msec in duration were presented on a spatially coincident, 1000 td white background. When compared to the control trichromatic data, the 200 msec functions of dichromats manifest a reduction in sensitivity at middle and long wavelengths. These data are consistent with the notion that 200 msec threshold increments reveal the sensitivity of the parvo pathway; the sensitivity of this pathway is reduced in dichromacy due to pigment replacement with a resultant loss of spectral opponency. The 10 msec functions of dichromats do not show a comparable reduction in sensitivity. Therefore, it is concluded that 10 msec increments reveal the sensitivity of a pathway whose spectral-opponent status is not substantially altered in dichromacy, presumably the magno pathway.

Adult↗

Surface color naming in dichromats.

In previous experiments, Montag and Boynton [(1987) Vision Research, 27, 2153-2162] found that many dichromats can categorize colors using color naming in fair agreement with color-normal subjects. The contribution of rods to color vision was suspected as underlying this ability. Here we follow up on these experiments by having dichromats name colors under various conditions. When the stimuli are limited to a brief presentation time (60 msec) the dichromats' categorization in the three dimensions of the OSA color space is impaired. Using high light levels so that the rods are saturated does not impair performance. The dichromats named colors during the period of the cone plateau following a rod bleach. Contrary to Montag and Boynton (1987) there was no deficit. These results suggest that an anomalous third cone pigment is responsible for the categorization in three dimensions. It is concluded that the receptors containing the anomalous pigment require greater temporal and spatial summation in order to contribute to the dichromats' color categorization.

Color Perception↗