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A perspective on color vision in platyrrhine monkeys.

Studies carried out over the past two decades show that many platyrrhine (New World) monkeys have polymorphic color vision. This condition results from the sorting of allelic versions of X-chromosome cone opsin genes at a single gene site, yielding a mixture of dichromatic and trichromatic phenotypes in the population. Two genera of platyrrhine monkey are known to deviate significantly from this pattern. Examination of color vision, photopigments, and photopigment genes of all of these monkeys have stimulated a renewed interest in understanding the evolution of primate color vision.

Animals↗

Relating color discrimination to photopigment genes in deutan observers.

Deutan observers are a heterogeneous group, varying nearly continuously from deuteranomalous trichromats with fine chromatic discrimination in the red/green range to deuteranopes who have none. We sought to relate chromatic discriminative ability among deutans measured psychophysically (phenotypes) to observers' separation between long-wave visual pigments inferred from visual pigment genes (genotypes). If middle-wave pigment genes are assumed not to be expressed in these deutan observers there is a clear relation between phenotype and genotype.

Color Perception↗

Effect of luminance on color perception of protanopes.

Small-field color-naming performance of two protanopes over a 4-log luminance range was impoverished in comparison with that of normal trichromats, and was more strongly affected by changes in luminance. At 200 cd/m2 responses to mid-spectral lights were dominated by 'yellow'; with lowering luminance, 'green' and 'red' were increasingly used. In the color spaces derived from these data the first two dimensions for trichromats are red-green and yellow-blue: those of the protanopes appear to be brightness and 'red-blue'. In the protanopes' color space the greater separation of stimuli at 0.2 cd/m2 suggests that with low luminance their color discrimination improves.

Adult↗

Macular pigment densities derived from central and peripheral spectral sensitivity differences.

Estimates of the density spectrum of the macular pigment (Wyszecki G, Stiles WS. Color Science: Concepts and Methods. Quantitative Data and Formulas. 1st ed. New York: Wiley, 1967); (Vos JJ. Literature review of human macular absorption in the visible and its consequences for the cone receptor primaries. Institute for Perception. Soesterberg, The Netherlands, 1972) are partially based on the difference between central and peripheral spectral sensitivities, measured under conditions chosen to isolate a single cone class (Stiles WS. Madrid: Union Internationale de Physique Pure et Appliquée, 1953;1:65-103). Such derivations assume that the isolated spectral sensitivity is the same at both retinal locations, save for the intervening macular pigment. If this is true, then the type of cone class mediating detection should not influence the calculated difference spectrum. To test this assumption, we measured central and peripheral spectral sensitivities in a deuteranope, a protanope and a normal trichromat observer: (a) for short-wave sensitive (S-) cone detection; and (b) for long-wave sensitive (L-) cone detection (deuteranope), for middle-wave sensitive (M-) cone detection (protanope) or for both L- and M-cone detection (normal trichromat). The difference spectra determined for L- or M-cone detection deviate significantly from those measured for S-cone detection, at wavelengths below 450 nm. A theoretical analysis suggests that the discrepancies are owing, in part, to regional variation in the optical density of the cone pigments; and that such receptor variation cannot be ignored when deriving the standard density spectrum of the macular pigment.

Color Vision Defects↗

Flicker cone electroretinogram in dichromats and trichromats.

To measure cone signal strengths in the flicker electroretinogram (ERG) of dichromats and trichromats, we developed a set of flickering stimuli (30 Hz), which excite the middle-wavelength-sensitive (M-) and long-wavelength-sensitive (L-) cones independently. ERG responses to eight different ratios of L- to M-cone contrasts were recorded from each subject. The short-wavelength-sensitive (S-) cone contrast was 0% in all measurements. The recordings were Fourier analyzed to determine the amplitude of the fundamental component. ERG threshold values for each subject resulted in ellipses when plotted in an L-/M-cone contrast space. As expected, the orientations of the threshold ellipses of the protanopes (N = 2) were parallel to the L-cone axis, whereas those of the deuteranopes (N = 2) were parallel to the M-cone axis. For the trichromats (N = 5), there was considerable interindividual variation in ellipse orientation.

Adult↗

Color vision in two observers with highly biased LWS/MWS cone ratios.

Two sisters, heterozygous carriers for congenital X-linked protanopia, were diagnosed as normal trichromats by the Rayleigh match on the anomaloscope. The heterozygous state was established by molecular analysis of their visual pigment genes. The normal color match establishes that the spectral sensitivities of their long-wavelength-sensitive (LWS) and middle-wavelength-sensitive (MWS) cone visual photopigments are within normal variability. Their FM 100-hue test error scores were low, demonstrating superior chromatic discrimination. Heterochromatic flicker photometric (HEP) spectral sensitivities were like those of protanopes. The estimated LWS/MWS cone ratios from the HFP data were 0.09/1 and 0.03/1, compared with ratios in the range of 0.6/1 to 10/1 for typical normal trichromats. Measurements of chromatic grating acuity on chromatically selective backgrounds were performed to study the cone mosaic. The data were consistent with a sparsity of LWS cones. Both protan carriers showed normal spectral sensitivities for all three cone types under cone isolating chromatic adaptation and normal three-peaked curves for increment thresholds on a white pedestal. Hue estimation, run on one carrier was normal. The equilibrium yellow locus was measured in the other carrier and was in the range of normal trichromats. The data indicate that normal color vision can occur even when the LWS/MWS cone ratio is quite abnormal.

Adaptation, Ocular↗

Colour vision in AIDS patients without HIV retinopathy.

Patients suffering from AIDS develop ocular complications, the most frequent being HIV retinopathy. It is however not clear, if functional visual impairments can be observed as early indicators of ocular complications, before clinical diagnosis of HIV retinopathy is made at fundus examination. To address this issue, we measured colour vision in a group of 49 AIDS subjects with normal clinical fundi using the 'two equation method'. This method, combining red-green Rayleigh and the blue-green Moreland metameric matches, enables more complete and quantitative assessments of colour vision than those based on pigmentary tests. Data were collected on our computer controlled colorimeter and compared to those of normal subjects. While most AIDS subjects without HIV retinopathy demonstrated normal colour vision, a significant portion of them had wider matches than normal subjects (11% for the Rayleigh equation and 16% for the Moreland equation). Furthermore, matching ranges of the Moreland equation were significantly correlated with CD4 lymphocyte counts. Patients with low CD4 values tended to produce larger matching ranges than the patients with high CD4 values. A within subject study on 17 patients confirmed this trend and showed that the patients who increased/decreased their CD4 blood counts generally improved/impaired their colour discrimination in the Moreland match. No such correlation was found between the matching ranges of the Rayleigh equation and the CD4 counts. These results show that colour discrimination is slightly reduced in some AIDS subjects, although there are no detectable ocular complications. They also suggest two different types of colour vision impairments in AIDS patients without retinopathy: one reversible process affecting colour discrimination in the blue-green range; and another irreversible process affecting colour discrimination in the red-green range.

Acquired Immunodeficiency Syndrome↗

On derivation of spectral sensitivities of the human cones from trichromatic colour matching functions.

Despite the recent advance made by using the direct methods of retinal densitometry, microspectrophotometry and suction electrophysiology, the psychophysical approach based on colour matching data still remains an important source of accurate information about the spectral sensitivity of the cone photoreceptors in the human visual system. However, the commonly used technique of estimating cone sensitivities, based on the assumption that dichromacy is caused by the lack of one of the three types of the cone photoreceptors, requires the colour matching functions not only from trichromatic observers but from dichromats as well. Here we evaluate an alternative approach, originally put forward by Bongard and Smimov, that derives cone spectral sensitivities from colour matching functions only; without resorting to colour deficiency or any other data. When applied to CIE standard colour matching functions, this method yields curves of spectral sensitivities that are close to the classical Smith-Pokorny fundamentals, though the long-wave cone is shifted towards the short-wave region of the spectrum by 5 nm, as compared with Smith and Pokorny's results.

Color Perception↗

Response phase of the flicker electroretinogram (ERG) is influenced by cone excitation strength.

We measured electroretinogram (ERG) response phases at different cone contrasts in trichromats and dichromats to investigate the dynamics of the long-wavelength-sensitive (L-) and middle-wavelength-sensitive (M-) cone pathways. ERG responses to stimuli, temporally modulated at 30 Hz, were recorded. The stimuli were generated on a computer controlled colour monitor. Thirty-two different combinations of L- and M-cone excitation strength, expressed as cone contrasts, were presented. The short-wavelength-sensitive (S-) cones were not stimulated (S-cone contrast = 0%). The response phase of the fundamental stimulus component was obtained from Fourier analysis. The ERG response phase lags decreased with increasing cone contrast. This was observed in all subjects with a normal appearing fundus. In dichromats and trichromats at low and intermediate contrasts, the phase lags to M-cone isolating conditions were smaller than those to L-cone isolating stimuli. In one dichromat with extreme myopia and cupping of the optic disc, the ERG phase lags increased with increasing cone contrast. The ERG response phase may be potentially useful for detecting retinal abnormalities.

Color Vision Defects↗

Human cone spectral sensitivities: a progress report.

The spectral sensitivities of the short (S-), middle (M-) and long (L-) wave-sensitive cones have been measured in normal trichromats and in dichromats and monochromats of known genotype. For the S-cone sensitivities, three blue-cone monochromats and five normals were used; for the M-cone sensitivities, nine protanopes (three with a single L1M2 gene, three with a single L2M3 gene, one with both an L1M2 and an M gene, and two with both an L2M3 and an M gene); and for the L-cone sensitivities, 22 deuteranopes (five with a single L(ala180) gene and 17 with a single L(ser180) gene). We compare existing cone spectral sensitivity estimates with these results and with tritanopic color matches. The new findings are more consistent with the cone fundamentals of Stockman et al. (JOSA 1993(A10), 2491) than with those of Smith and Pokorny (Vision Research 1975(15), 161). The discrepancies that we find, however, are sufficient to warrant the replacement of both sets.

Color Perception↗

Development of the spatio-chromatic visual evoked potential (VEP): a longitudinal study.

Most prior visual evoked potential (VEP) research on the development of color vision has employed pattern-reversing stimuli that are not optimal for producing chromatic responses. We measured infant VEPs using low spatial frequency, onset-offset stimuli, modulated along the three axes of a cone-based color space (Derrington et al. [J. Physiol 1984;357, 241-265.]). Three color-normal infants were tested in a longitudinal design over the first postnatal year. One red/green color-deficient infant was also tested at 197 days. We found that VEP responses to S-axis (tritan) stimuli have their initial onset later than responses to red/green (L-M) or achromatic stimuli, and that developmental changes in VEP waveforms are more complex and longer lasting for chromatic than for achromatic stimuli. Possible mechanisms underlying these changes are discussed.

Adult↗

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↗

Skewness and transformations of Farnsworth-Munsell 100-hue test scores.

In the past, suggested transformations of Farnsworth-Munsell 100-Hue Test (FM 100-Hue) test scores distributions have been limited to a square root transformation. In this study, the choice of transformations of total error scores (TES) are considered by identifying a possible source of skewness. Several distributions of FM100-Hue Test TES were assessed for skewness (third moment). The error score (ES) distributions for the 85 individual caps in each of the populations were also analysed for skewness (Figs. 3 and 4). There is no single transformation which will normalise all TES distributions. The single cap ES distributions with low mean ES (such as those achieved normals and, for some regions of the test, by anomalous trichromats and dichromats) are symmetrical because most subjects can organise the cap perfectly (and could do even better given smaller colour differences). The distributions of ESs where the mean ES is in the moderate range (such as those achieved by diabetics) are skewed because some ESs at the lower end of the range represent performance which could also be better than the test allows. ES distributions with a high mean (such as random distributions and some regions of the test by congenital dichromats) are symmetrical being unaffected by the limitations of the test. TES distributions of diabetics are asymmetrical and comprise skewed cap ES distributions. A suggestion for a transformation is made.

Color Perception Tests↗

Comparison of the D-15 and City University (second) color vision tests.

This study examines the equivalence of the City University color vision test (CUT) to the Farnsworth Munsell D-15 (D-15). This information is important because a number of occupations require individuals to pass either the D-15 of the CUT. Using two or more major crossings on the D-15 as failure, our results indicate that agreement between tests can be improved if the failing criterion for the CUT is either > or = 5 total errors, or > or = 2 errors on the large circles. Our preference is to use the criterion based on the large circles because there is a marginal improvement in agreement between the two tests for protanopes.

Adult↗

Analysis of red/green color discrimination in subjects with a single X-linked photopigment gene.

Many subjects despite having only a single X-linked pigment gene (single-L/M-gene subjects) are able to make chromatic discriminations by Rayleigh matching, especially when large fields are used. We used a combination of psychophysics (Rayleigh match), electroretinograms (ERG), and molecular genetic techniques to rule out several possible explanations of this phenomenon. Use of rods for chromatic discrimination was unlikely since strong adapting fields were employed and the large-field match results were not consistent with rod participation. A putative mid- to long-wavelength photopigment that escapes detection by current molecular genetic analysis was ruled out by finding only a single L/M photopigment in flicker ERGs from 16 single-L/M-gene subjects. Large-field match results were not consistent with participation of S cones. Amino acid sequence polymorphisms in the S-pigment gene that might have shifted the S cone spectrum towards longer wavelengths were not found on sequencing. The mechanism of chromatic discrimination in the presence of a single photopigment therefore remains unknown. Further possible explanations such as variations in cone pigment density and retinal inhomogeneities are discussed.

Adaptation, Ocular↗

S-cone ERGs elicited by a simple technique in normals and in tritanopes.

PURPOSE: To measure changes in the relative spectral sensitivities of the dark adapted and light adapted ERG and thus to establish the possible contribution of rods to the 'blue cone' ERG elicited by flashes of blue light. BACKGROUND: Short wavelength stimuli in the light-adapted eye evoke small rounded b-waves which have been considered to be S-cone responses. We have recorded such responses from tritanopes, which called the assumptions into question. METHODS: Small ERGs were recorded to blue and green flashes. The stimulus was a Ganzfeld which employed light emitting diodes. ERGs were obtained in both the dark-adapted eye and after light adaptation to intense orange light (peak wavelength 610 nm). The change in sensitivity with light adaptation and the relative spectral sensitivity was determined from the voltage/log light intensity functions, using a 10 microV criterion. RESULTS: (1) peak times and changes in sensitivity did not help distinguish light-adapted rod from possible S-cone responses; (2) analysis of the change in the ratio of blue:green sensitivity from darkness to 4.4 log Td. 610 nm background suggests that in seven normal subjects, 90% or more of the ERG evoked by 440 nm flashes is generated by S-cones; (3) three tritanopes have insignificantly reduced S-cone responses. CONCLUSIONS: (1) clinical techniques used to isolate S-cone ERGs are appropriate; (2) there are at least two types of tritanope and in those we investigated, functional S-cones are probably displaced into the retinal periphery.

Adaptation, Ocular↗

Development of brightness matching and colour vision deficits in juvenile diabetics.

We studied hue discrimination and brightness matching throughout the spectrum in ten juvenile patients suffering from diabetes mellitus (Type I) with no (eight patients) or mild (two patients) retinopathy. In addition, the FM 100-Hue test was performed. The data were collected once every year over 5 years. Over the 5 years, the diabetics show a continual change in the shape of their brightness matching function. Wavelength discrimination ability remains quite stable with time at the long end of the spectrum but is variable at short wavelengths. FM-100 error scores remain similar over the period tested, at a level slightly higher than that of a control group. Additional experiments show that the sensitivity of the S-cone in the diabetic group is similar to that of controls. The results can be explained by an early relative reduction in the sensitivity of post-receptoral processes in juvenile diabetics.

Adolescent↗

Morphology of transient VEPs to luminance and chromatic pattern onset and offset.

Characteristics of the visual evoked response to chromatic and luminance-modulated stimuli reflect the activity of underlying neural mechanisms, although selective neuronal activity depends upon stimulus parameters. In the present study, the behaviour of the transient visual evoked response to low spatial and temporal frequency chromatic stimuli is investigated at a range of colour luminance ratios. Our results show that the response to pattern-offset may be used in addition to the pattern-onset response as part of the signature of the evoked response to luminance-modulated or isoluminant chromatic stimuli.

Adult↗