J. D. Mollon

Human visual pigments: microspectrophotometric results from the eyes of seven persons

The material for this work was obtained from seven eyes removed because of malignant growths. Foveal and parafoveal samples of the retinas were taken and transverse measurements were made of the absorbance spectra of the outer segments of the rods and cones, using a Liebman microspectrophotometer. Four kinds of spectra were obtained with absorbance peaks at the following wavelengths: rods, 496.3 ± 2.3 nm (n = 39); red cones, 558.4 ± 5.2 nm (n = 58); green cones, 530.8 ± 3.5 nm (n = 45); blue cones, 419.0 ± 3.6 nm (n = 5). The distribution of the peaks was unimodal for the rods. For the red and green cones, however, there was evidence for bimodal distributions, with sub-population maxima at 563.2 ± 3.1 nm (n = 27) and 554.2 ± 2.3 nm (n = 31) for the reds and at 533.7 ± 2.1 nm (n = 23) and 527.8 ± 1.8 nm (n = 22) for the greens. A substantial difference in mean spectral location of the red cones was observed between patient 1 (561 nm) and patient 4 (553 nm). Both patients were classified as normal trichromats by all clinical tests of colour vision but there was a clear difference in their relative sensitivities to long-wave fields. In both direction and magnitude, this difference proved to be that required by the microspectrophotometric results.

Variations of Colour Vision in a New World Primate Can be Explained by Polymorphism of Retinal Photopigments

The squirrel monkey (Saimiri sciureus) exhibits a polymorphism of colour vision: some animals are dichromatic, some trichromatic, and within each of these classes there are subtypes that resemble the protan and deutan variants of human colour vision. For each of ten individual monkeys we have obtained (i) behavioural measurements of colour vision and (ii) microspectrophotometric measurements of retinal photopigments. The behavioural tests, carried out in Santa Barbara, included wavelength discrimination, Rayleigh matches, and increment sensitivity at 540 and 640 nm. The microspectrophotometric measurements were made in London, using samples of fresh retinal tissue and a modified Liebman microspectrophotometer: the absorbance spectra for single retinal cells were obtained by passing a monochromatic measuring beam through the outer segments of individual rods and cones. The two types of data, behavioural and microspectrophotometric, were obtained independently and were handed to a third party before being interchanged between experimenters. From all ten animals, a rod pigment was recorded with λmax (wavelength of peak absorbance) close to 500 nm. In several animals, receptors were found that contained a short-wave pigment (mean λmax = 433.5 nm): these violet-sensitive receptors were rare, as in man and other primate species. In the middle- to long-wave part of the spectrum, there appear to be at least three possible Saimiri photopigments (with λmax values at about 537, 550 and 565 nm) and individual animals draw either one or two pigments from this set, giving dichromatic or trichromatic colour vision. Thus, those animals that behaviourally resembled human protanopes exhibited only one pigment in the red-green range, with λmax = 537 nm ; other behaviourally dichromatic animals had single pigments lying at longer wavelengths and these were the animals that behaviourally had higher sensitivity to long wavelengths. Four of the monkeys were behaviourally judged to be trichromatic. None of the latter animals exhibited the two widely separated pigments (close to 535 and 567 nm) that are found in the middle- and long-wave cones of macaque monkeys. But the spread of λmax values for individual cones was greater in the trichromatic squirrel monkeys than in the dichromats; and in the case of three, behaviourally deuteranomalous, trichromats there wasclear evidence that the distribution of λmax values was bimodal, suggesting photopigments at approximately 552 and 565 nm. The fourth, behaviourally protanomalous, trichrom at exhibited a spread of individual λmax values that ranged between 530 and 550 nm. Good quantitative agreement was found when the microspectrophoto-metrically measured absorbance spectra were used to predict the behavioural sensitivity of individual animals to long wavelengths. The concordance of the two sets of measurements places beyond question the existence of a polymorphism of colour vision in Saimiri sciureus and suggests that the behavioural variation arises from variation in the retinal photopigments. Heterozygous advantage may explain the polymorphism.

Luminance noise and the rapid determination of discrimination ellipses in colour deficiency

A computer-controlled test of colour vision is described, in which luminance noise and masking contours are used to ensure that the subjects responses depend on chromatic signals. The test avoids the need--common to most computer-controlled tests--to define equiluminance for the individual subject before the colour test itself can be administered. The test achieves a good separation of protan and deutan subjects and reveals the large range of chromatic sensibilities among anomalous trichromats. As a population, dichromats had higher thresholds on the tritan axis of the test than did normals. In an extension of the test, full discrimination ellipses were measured for normal and colour-deficient observers. The nature of anomalous trichromacy is discussed and the possibility is raised that hybrid genes, resulting from genetic recombination, may code for incorrectly labelled or functionally impaired molecules.

The influence of contrast adaptation on color appearance

Most models of color vision assume that signals from the three classes of cone receptor are recoded into only three independent post-receptoral channels: one that encodes luminance and two that encode color. Stimuli that are equated for their effects on two of the channels should be discriminable only to the remaining channel, and are thus assumed to isolate the responses of single channels. We used an asymmetric matching task to examine whether such models can account for changes in color appearance following adaptation to contrast--to temporal variations in luminance and chromaticity around a fixed mean luminance and chromaticity. The experiments extend to suprathreshold color appearance the threshold adaptation paradigm of Krauskopf, Williams and Heeley [(1982) Vision Research, 32, 1123-1131]. Adaptation changes the perceived color of chromatic test stimuli both by reducing their saturation (contrast) and by changing their hue (direction within the equiluminant plane). The saturation losses are largest for test stimuli that lie along the chromatic axis defining the adapting modulation, while the hue changes are rotations away from the adapting direction and toward an orthogonal direction within the S and L-M plane. Similar selective changes in both perceived color and perceived lightness occur following adaptation to stimuli that covary in luminance and chromaticity. The selectivity of the aftereffects for multiple directions within color-luminance space is inconsistent with sensitivity changes in only three independent channels. These aftereffects suggest instead that color appearance depends on channels that can be selectively tuned to any color-luminance direction, and that there are no directions that invariably isolate responses in only a single channel. We use the perceived color changes to examine the spectral sensitivities of the chromatic channels and to estimate the distribution of channels. We also examine how adaptation alters the contrast-response function, how it affects reaction times for luminance and chromatic contrast, the extent to which the aftereffects exhibit interocular transfer, and the way in which the perceived color changes differ from those induced by conventional light adaptation.

Computerized simulation of color appearance for dichromats

We propose an algorithm that transforms a digitized color image so as to simulate for normal observers the appearance of the image for people who have dichromatic forms of color blindness. The dichromats color confusions are deduced from colorimetry, and the residual hues in the transformed image are derived from the reports of unilateral dichromats described in the literature. We represent color stimuli as vectors in a three-dimensional LMS space, and the simulation algorithm is expressed in terms of transformations of this space. The algorithm replaces each stimulus by its projection onto a reduced stimulus surface. This surface is defined by a neutral axis and by the LMS locations of those monochromatic stimuli that are perceived as the same hue by normal trichromats and a given type of dichromat. These monochromatic stimuli were a yellow of 575 nm and a blue of 475 nm for the protan and deutan simulations, and a red of 660 nm and a blue-green of 485 nm for the tritan simulation. The operation of the algorithm is demonstrated with a mosaic of square color patches. A protanope and a deuteranope accepted the match between the original and the appropriate image, confirming that the reduction is colorimetrically accurate. Although we can never be certain of anothers sensations, the simulation provides a means of quantifying and illustrating the residual color information available to dichromats in any digitized image.

Dichromats Detect Colour-Camouflaged Objects that are not Detected by Trichromats

To explain the surprisingly high frequency of congenital red–green colour blindness, the suggestion has been made that dichromats might be at an advantage in breaking certain kinds of colour camouflage. We have compared the performance of dichromats and normal observers in a task in which texture is camouflaged by colour. The texture elements in a target area differed in either orientation or size from the background elements. In one condition, the texture elements were all of the same colour; in the camouflage condition they were randomly coloured red or green. For trichromats, it proved to be more difficult to detect the target region in the camouflage condition, even though colour was completely irrelevant to the task. Dichromats (n = 7) did not show this effect, and indeed performed better than trichromats in the camouflage condition. We conclude that colour can interfere with segregation based upon texture, and that dichromats are less susceptible to such interference.

Adaptation and the color statistics of natural images.

Color perception depends profoundly on adaptation processes that adjust sensitivity in response to the prevailing pattern of stimulation. We examined how color sensitivity and appearance might be influenced by adaptation to the color distributions characteristic of natural images. Color distributions were measured for natural scenes by sampling an array of locations within each scene with a spectroradiometer, or by recording each scene with a digital camera successively through 31 interference filters. The images were used to reconstruct the L, M and S cone excitation at each spatial location, and the contrasts along three post-receptoral axes [L + M, L - M or S - (L + M)]. Individual scenes varied substantially in their mean chromaticity and luminance, in the principal color-luminance axes of their distributions, and in the range of contrasts in their distributions. Chromatic contrasts were biased along a relatively narrow range of bluish to yellowish-green angles, lying roughly between the S - (L + M) axis (which was more characteristic of scenes with lush vegetation and little sky) and a unique blue-yellow axis (which was more typical of arid scenes). For many scenes L - M and S - (L + M) signals were highly correlated, with weaker correlations between luminance and chromaticity. We use a two-stage model (von Kries scaling followed by decorrelation) to show how the appearance of colors may be altered by light adaptation to the mean of the distributions and by contrast adaptation to the contrast range and principal axes of the distributions; and we show that such adjustments are qualitatively consistent with empirical measurements of asymmetric color matches obtained after adaptation to successive random samples drawn from natural distributions of chromaticities and lightnesses. Such adaptation effects define the natural range of operating states of the visual system.

Three remarks on perceptual learning

This essay makes three points. (1) From the failure of perceptual learning to transfer when stimulus parameters are changed, it cannot necessarily be concluded that the site of learning is distal: rather, the learning may be central and the specificity may lie in what is learnt. (2) Mere exposure to a stimulus may not be sufficient for learning: even in the absence of explicit feedback, other sources of information are often available in a perceptual learning task. (3) Procedural learning may sometimes be apparent only after a delay: this phenomenon has a long history and is known as reminiscence.

REACTION TIME AS A MEASURE OF THE TEMPORAL RESPONSE PROPERTIES OF INDIVIDUAL COLOUR MECHANISMS

Abstract The old question of whether visual latency varies with wavelength has been reexamined by isolating individual π mechanisms. Reaction times were recorded to monochromatic increments presented on monochromatic adapting fields and it was found that the time constants of each colour mechanism vary independently according to the sensitivity of the mechanism concerned and do not depend upon the adaptive state of the retina as a whole. This independence may partially explain discrepant results in heterochromatic photometry and distortions of hue that arise from brief or intermittent stimulation. A correlation between the present results and the corresponding critical durations demonstrates the value of reaction times in the study of visual processes.

The relationship between cone pigments and behavioural sensitivity in a new world monkey (Callithrix jacchus jacchus)

Microspectrophotometric measurements of visual pigments and behavioural measurements of spectral sensitivity are reported for individual marmosets from 3 family groups. The sex differences and polymorphism that characterise the long-wave cone pigments in this species are well reflected by variations in the behavioural sensitivities. With one exception, the pattern of inheritance is compatible with a genetic model in which the long-wave pigment is specified by a single polymorphic locus on the X-chromosome. Measurements are also reported for the spectral absorbance of the marmoset lens, and these are used to reconstruct short-wave behavioural sensitivity from the microspectrophotometric measurements of the short-wave cones.