Color vision: opsins and options.
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Animals↗
Biomedical subjects
J D Mollon
Publications and source records attributed to J D Mollon.
The evolution of trichromatic color vision by opsin gene duplication in New World and Old World primates.
Trichromacy in all Old World primates is dependent on separate X-linked MW and LW opsin genes that are organized into a head-to-tail tandem array flanked on the upstream side by a locus control region (LCR). The 5' regions of these two genes show homology for only the first 236 bp, although within this region, the differences are conserved in humans, chimpanzees, and two species of cercopithecoid monkeys. In contrast, most New World primates have only a single polymorphic X-linked opsin gene; all males are dichromats and trichromacy is achieved only in those females that possess a different form of this gene on each X chromosome. By sequencing the upstream region of this gene in a New World monkey, the marmoset, we have been able to demonstrate the presence of an LCR in an equivalent position to that in Old World primates. Moreover, the marmoset sequence shows extensive homology from the coding region to the LCR with the upstream sequence of the human LW gene, a distance of >3 kb, whereas homology with the human MW gene is again limited to the first 236 bp, indicating that the divergent MW sequence identifies the site of insertion of the duplicated gene. This is further supported by the presence of an incomplete Alu element on the upstream side of this insertion point in the MW gene of both humans and a cercopithecoid monkey, with additional Alu elements present further upstream. Therefore, these Alu elements may have been involved in the initial gene duplication and may also be responsible for the high frequency of gene loss and gene duplication within the opsin gene array. Full trichromacy is present in one species of New World monkey, the howler monkey, in which separate MW and LW genes are again present. In contrast to the separate genes in humans, however, the upstream sequences of the two howler genes show homology with the marmoset for at least 600 bp, which is well beyond the point of divergence of the human MW and LW genes, and each sequence is associated with a different LCR, indicating that the duplication in the howler monkey involved the entire upstream region. [The sequence data described in this paper have been submitted to GenBank under accession nos. AF155218, AF156715, and AF156716.]
Animals↗
Colour discrimination thresholds in Parkinson's disease: results obtained with a rapid computer-controlled colour vision test.
A dysfunction of dopaminergic retinal neurons is thought to occur in Parkinson's disease, manifesting itself in impaired performance on various visual discrimination tasks. We have investigated whether differences in colour discrimination could readily be detected between a normal group and a Parkinsonian group, using a computer-controlled test of colour vision. Although some individual Parkinsonian patients showed an abnormal elevation of colour discrimination thresholds, there was no significant difference between the normal group and the Parkinsonian group.
Age Factors↗
Shift in Rayleigh matches after adaptation to monochromatic light of various intensities.
The action spectrum for inducing a long-lasting protan shift in colour matches was investigated. Rayleigh matches were measured before and after 30 min adaptation to monochromatic light of 520, 550, 580 or 620 nm. For each wavelength, seven retinal illuminances, ranging from 3.0 to 5.0 log td, were chosen in random order. Results for one colour-normal observer show that the shift in Rayleigh match after adaptation increases monotonically as a function of the luminance of the adapting light. The dynamic response range is from 3.3 to 4.7 log td. The wavelength of the adapting light had no systematic influence on the form of the response function. The results imply that the effect is triggered by light absorbed in the photopigments themselves.
Adaptation, Ocular↗
Molecular evolution of trichromacy in primates.
Although trichromacy in Old and New World primates is based on three visual pigments with spectral peaks in the violet (SW, shortwave), green (MW, middlewave) and yellow-green (LW, longwave) regions of the spectrum, the underlying genetic mechanisms differ. The SW pigment is encoded in both cases by an autosomal gene and, in Old World primates, the MW and LW pigments by separate genes on the X chromosome. In contrast, there is a single polymorphic X-linked gene in most New World primates with three alleles coding for spectrally distinct pigments. The one reported exception to this rule is the New World howler monkey that follows the Old World system of separate LW and MW genes. A comparison of gene sequences in these different genetic systems indicates that the duplication that gave rise to the separate MW and LW genes of Old World primates is more ancient than that in the howler monkey. In addition, the amino acid sequences of the two howler monkey pigments show similarities to the pigments encoded by the polymorphic gene of other New World primates. It would appear therefore that the howler monkey gene duplication arose after the split between New and Old World primates and was generated by an unequal crossover that placed two different forms of the New World polymorphic gene on to a single chromosome. In contrast, the lack of identity at variable sites within the New and Old World systems argues for the origin of the separate genes in Old World primates by the duplication of a single form of the gene followed by divergence to give spectrally distinct LW and MW pigments. In contrast, the similarity in amino acid variation across the tri-allelic system of New World primates indicates that this polymorphism had a single origin in New World primates. A striking feature of all these pigments is the use of a common set of substitutions at three amino acid sites to achieve the spectral shift from MW at around 530 nm to LW at around 560 nm. The separate origin of the trichromacy in New and Old World primates would indicate that the selection of these three sites is the result of convergent evolution, perhaps as a consequence of visual adaptation in both cases to foraging for yellow and orange fruits against a green foliage.
Alouatta↗
Frugivory and colour vision in Alouatta seniculus, a trichromatic platyrrhine monkey.
It is a long-standing hypothesis that primate trichromacy evolved to help fruit-eating primates find fruits amongst leaves. We measured the reflectance spectra of fruits eaten by a trichromatic primate, Alouatta seniculus, in the rainforest of French Guiana, as well as those of the leaves that form the natural background to fruits. We develop a method of specifying these natural colour signals in a chromaticity diagram appropriate for A. seniculus. By treating the task facing frugivorous monkeys as a signal detection task, we show that the spectral tuning of the L and M cone pigments in A. seniculus is optimal for detecting fruits amongst leaves.
Alouatta↗
Colour discrimination ellipses in patients with dominant optic atrophy.
Many colour tests require a visual acuity of at least 0.1, making them unsuitable for low vision patients. To assess colour vision in patients with sub-normal acuity, we re-designed a previously described test so that its spatial details would be coarse enough to be resolvable by subjects with severe visual impairment. The test measures chromatic discrimination along 20 axes evenly spaced in CIE 1976 L*u*v* colour space. We detail the results for this test in a group of patients with dominant optic atrophy. Despite the lack of evidence for genetic heterogeneity in dominant optic atrophy, we observed phenotypic variation both between and within families.
Adult↗
What is the function of the cone-rich rim of the retina?
Although there is good histological evidence for a rim of cones extending round the margin of the human retina at the ora serrata, the function of these cones is unknown, and indeed it is not known whether they are functional at all. Four possibilities are discussed here: (i) the cones of the ora serrata may alert us to sudden movements in the far peripheral field, (ii) their signal may be used in estimating optic flow during locomotion, (iii) they may integrate light scattered within the globe of the eye or passing through the sclera, for purposes of colour constancy, or (iv) they may drive circadian rhythms. We report two experiments designed to detect psychophysical correlates of the cone rim. Under the conditions we have used, neither flicker detection nor colour naming show, near the limit of vision, a discontinuity that would correspond to the cone-rich rim.
Adult↗
The number of triplet repeats in five brain-expressed loci with CAG repeats is not associated with schizophrenia.
We have previously shown that large expansions of CAG (CTG) triplets are associated both with schizophrenia itself and with an early age-at-onset of the disease. However, the repeat expansion detection (RED) method used did not provide a chromosomal location for the expanded region(s) (Morris et al., 1995). In a further study of our schizophrenic and control patients, we have now examined the length of the repeated sequence in five loci that are expressed in brain and are known to contain CAG repeat regions (Li et al., 1993). No enlarged repeat regions were identified; it is unlikely therefore that expansions at any of these five loci can account for expansions of up to 136 triplets identified by the RED method.
Age of Onset↗
Adaptation of colour vision to sunlight.
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Adaptation, Ocular↗
Motion minima for different directions in color space.
We have used the minimum-motion stimulus of Cavanagh, MacLeod & Anstis [(1987) Journal of the Optical Society of America A, 4, 1428-1438] to examine how signals along different directions in color space interact in motion perception. Stimuli were pairs of counterphasing gratings combined 90 deg out of phase in both space and time and modulated along different color-luminance axes. The axis for one of the gratings was fixed, while the axis for the second was varied so as to null perceived motion in the stimulus. The motion nulls show that observers are sensitive to motion signals carried by each of the cardinal directions of color space [an achromatic axis and L-M and S-(L+M) chromatic axes], but that signals along different cardinal axes are not combined to yield a net direction of motion. Pairing an achromatic and chromatic grating resulted in a motion null regardless of the relative or overall contrast of the two gratings, while the null directions for intermediate axes shifted depending on contrast. This result points to the special status of the luminance and chromatic axes. However, our results do not reveal a special pair of axes within the equiluminant plane. When contrasts along the cardinal axes are scaled for equal multiples of their respective detection thresholds, the L-M and S chromatic contrasts contribute roughly equally to the perceived motion, but are many times weaker than luminance contrast. Moreover, sensitivity to luminance motion is little affected by the presence of chromatic contrast, whereas sensitivity to chromatic motion is strongly masked by either luminance or chromatic contrast. These asymmetric interactions suggest that the motion of the luminance and chromatic components is encoded in qualitatively different ways.
Color Perception↗
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.
Adaptation, Ocular↗
"... On the basis of velocity clues alone": some perceptual themes 1946-1996.
Three factors that have transformed perceptual research in the last fifty years are the digital computer, single-unit electrophysiology, and molecular biology. Amongst the developments in which members of the Experimental Psychology Society have been central are: the recognition of the role of optic flow in spatial vision; the demonstration that our perceptual systems contain parallel pathways extracting different information from the sensory array; the identification of specific detectors that can be selectively adapted in psychophysical experiments; and the transfer of the concepts of fourier analysis from audition to vision. The history of Opponent Process Theory offers an example where experimental psychologists have been misled by too simple an interpretation of physiological recordings.
Animals↗
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 dichromat's 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 another's sensations, the simulation provides a means of quantifying and illustrating the residual color information available to dichromats in any digitized image.
Algorithms↗
Assessment of association of D3 dopamine receptor MscI polymorphism with schizophrenia: analysis of symptom ratings, family history, age at onset, and movement disorders.
Several studies have reported an association between schizophrenia and homozygosity for the MscI restriction site in exon 1 of the D3 dopamine receptor gene, but other studies have failed to find this association. Recent reports have suggested that the association is most salient in male patients with a family history of schizophrenia. We examined this restriction site in a group of schizophrenic patients (n = 84) and in normal controls (n = 77). Patients were subdivided according to demographic and clinical features, particular attention being paid to movement disorders. No significant difference in allelic or genotypic distribution was seen between the two groups. No association was seen between homozygosity and a positive family history, age at onset of illness, clinical subtype, negative symptom score, or movement disorder scores.
Chi-Square Distribution↗
Errors of judgement at Greenwich in 1796.
The origins of experimental psychology can be traced back to 1796, when the then Astronomer Royal dismissed his assistant for making some seemingly inaccurate measurements. But there is more to the story than meets the eye.
Astronomy↗
Measurements of human sensitivity to comb-filtered spectra.
Using a novel stimulator that incorporates a liquid crystal display, the spectral modulation sensitivity function of the colour vision system was derived by measuring discrimination thresholds for comb-filtered spectra. This function shows a peak of sensitivity at 0.97 c/300 nm with a plateau that extends to 1.67 c/300 nm. Extrapolation of the curve gives an estimated high-frequency cut-off at 5 c/300 nm. The thresholds are also transformed to the CIE (1931) diagram and the elliptical isosensitivity contours thus obtained are compared with the classical discrimination ellipse of MacAdam for the corresponding region of colour space.
Color Perception↗
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.
Humans↗