The International Colour Vision Society awarded the 2003 Verriest Medal to André Roth, Honorary Professor of Ophthalmology at the University of Geneva. This award is bestowed by the Society to honor long-term contributions to the field of color vision. André Roth created the Roth 28 Hue test for ophthalmological examination, and developed and standardized a sophisticated diagnostic set of tests based on color metrics for acquired color vision deficiencies. He developed an anomaloscope specially for the investigation of acquired and inherited color vision deficiencies in ophthalmology. As director of the Geneva University clinic, he has studied most eye diseases in which acquired color deficiencies play a significant role. Work for the IRGCVD and later ICVS was a significant part of his professional life. Together with Guy Verriest (the first President of the Society, for whom the medal is named), André belonged to the clinically oriented group, which recommended a separation of the IRGCVD from the AIC, to give ophthalmologists, physiologists and other clinicians a scientific home and a connection to color science. Soon after the death of Guy Verriest he took over the presidency of the Society from Wolfgang Jaeger and carefully led its further development, with expanded emphasis of genetic, molecular biological, physiological, and psychophysical research.

The normal X-chromosome-linked color-vision gene array is composed of a single long-wave-sensitive (L-) pigment gene followed by one or more middle-wave-sensitive (M-) pigment genes. The expression of these genes to form L- or M-cones is controlled by the proximal promoter and by the locus control region. The high degree of homology between the L- and M-pigment genes predisposed them to unequal recombination, leading to gene deletion or the formation of L/M hybrid genes that explain the majority of the common red–green color-vision deficiencies. Hybrid genes encode a variety of L-like or M-like pigments. Analysis of the gene order in arrays of normal and deutan subjects indicates that only the two most proximal genes of the array contribute to the color-vision phenotype. This is supported by the observation that only the first two genes of the array are expressed in the human retina. The severity of the color-vision defect is roughly related to the difference in absorption maxima (λmax) between the photopigments encoded by the first two genes of the array. A single amino acid polymorphism (Ser180Ala) in the L pigment accounts for the subtle difference in normal color vision and influences the severity of red–green color-vision deficiency.

Blue-cone monochromacy is a rare disorder that involves absence of L- and M-cone function. It is caused either by deletion of a critical region that regulates expression of the L/M gene array, or by mutations that inactivate the L- and M-pigment genes. Total color blindness is another rare disease that involves complete absence of all cone function. A number of mutants in the genes encoding the cone-specific α- and β-subunits of the cGMP-gated cation channel as well as in the α-subunit of transducin have been implicated in this disorder.

X-linked incomplete achromatopsia (XIA), also called blue-cone monochromacy (BCM), is a rare cone disorder that most commonly results either from one of two conditions.. The first condition is a deletion of the locus control region (LCR) which is a critical DNA element that lies upstream of the L and M photopigment gene array on the X-chromosome and is necessary for expression of the photopigment genes. The second condition is an inactivating point mutation within the coding sequence of the remaining photopigment gene in an array from which all but one gene has been deleted. Many previous studies have concluded that affected individuals either have only rods and S-cones (Blackwell & Blackwell, 1957, 1961; Daw & Enoch, 1973; Hess et al., 1989) or have rods, S-cones, and another cone type that contains the rod pigment (Pokorny et al., 1970; Alpern et al., 1971). However, Smith et al. (1983) described individuals with XIA who had residual L-cone function. Here we report results for a subject with XIA who appears to have residual M-cone function. Genetic analysis revealed that he had apparently normal genes for M-cone photopigment thus leaving open the possibility that he has a contribution to vision based on expression of these genes at a very low level.

The hypothesis that dichromatic behavior on a clinical anomaloscope can be explained by the complement and arrangement of the long- (L) and middle-wavelength (M) pigment genes was tested. It was predicted that dichromacy is associated with an X-chromosome pigment gene array capable of producing only a single functional pigment type. The simplest case of this is when deletion has left only a single X-chromosome pigment gene. The production of a single L or M pigment type can also result from rearrangements in which multiple genes remain. Often, only the two genes at the 5′ end of the array are expressed; thus, dichromacy is also predicted to occur if one of these is defective or encodes a defective pigment, or if both of them encode pigments with identical spectral sensitivities. Subjects were 128 males who accepted the full range of admixtures of the two primary lights as matching the comparison light on a Neitz or Nagel anomaloscope. Strikingly, examination of the L and M pigment genes revealed a potential cause for a color-vision defect in all 128 dichromats. This indicates that the major component of color-vision deficiency could be attributed to alterations of the pigment genes or their regulatory regions in all cases, and the variety of gene arrangements associated with dichromacy is cataloged here. However, a fraction of the dichromats (17 out of 128; 13%) had genes predicted to encode pigments that would result in two populations of cones with different spectral sensitivities. Nine of the 17 were predicted to have two pigments with slightly different spectral peaks (usually ≤ 2.5 nm) and eight had genes which specified pigments identical in peak absorption, but different in amino acid positions previously associated with optical density differences. In other subjects, reported previously, the same small spectral differences were associated with anomalous trichromacy rather than dichromacy. It appears that when the spectral difference specified by the genes is very small, the amount of residual red–green color vision measured varies; some individuals test as dichromats, others test as anomalous trichromats. The discrepancy is probably partly attributable to testing method differences and partly to a difference in performance not perception, but it seems there must also be cases in which other factors, for example, cone ratio, contribute to a person's ability to extract a color signal from a small spectral difference.

Most platyrrhine monkeys have a triallelic M/L opsin gene polymorphism that underlies significant individual variations in color vision. A survey of the frequencies of these polymorphic genes suggests that the three alleles occur with equal frequency among squirrel monkeys (subfamily Cebinae), but are not equally frequent in a number of species from the subfamily Callitrichinae. This departure from equal frequency in the Callitrichids should slightly increase the ratio of dichromats to trichromats in the population and significantly alter the relative representation of the three possible dichromatic and trichromatic phenotypes. A particular feature of the inequality is that it leads to a relative increase in the number of trichromats whose M/L pigments have the largest possible spectral separation. To assess whether these trichromatic phenotypes are equally well equipped to make relevant visual discriminations, psychophysical experiments were run on human observers. A technique involving the functional substitution of photopigments was used to simulate the discrimination between fruits among a background of leaves. The goal of the simulation was to reproduce in the cones of human observers excitations equivalent to those produced in monkey cones as the animals view fruit. Three different viewing conditions were examined involving variations in the relative luminances of fruit and leaves and the spectrum of the illuminant. In all cases, performance was best for simulated trichromacies including M/L pigments with the largest spectral separation. Thus, the inequality of opsin gene frequency in Callitrichid monkeys may reflect adaptive pressures.

Studies of color vision in marsupial mammals have been very limited. Two photoreceptor genes have been characterized from the tammar wallaby, but a third cone pigment was suggested by microspectrophotometric measurements on cone photoreceptors in two other species, including the fat-tailed dunnart, Sminthopsis crassicaudata. To determine the sequence and infer absorption maxima of the cone photoreceptor pigments of S. crassicaudata and the related stripe-faced dunnart (Sminthopsis macroura), we have used evolutionarily conserved sequences of the cone pigments of other species, including the tammar wallaby, to design primers to amplify the S. macroura and S. crassicaudata pigment sequences by the polymerase chain reaction (PCR) using genomic DNA or retinal cDNA as a template. These primers will be useful for amplifying cone opsin coding sequences from a variety of vertebrates. Amplified products were directly sequenced to determine gene structure and coding sequences. The inferred amino acid sequences of the cone visual pigments indicated that both species have middle-wave-sensitive (MWS) pigments with a predicted absorption maximum (λmax) at 530 nm, and ultraviolet-sensitive (UVS) pigments with a predicted λmax at 360 nm. The MWS pigments of the two species differ by two, and UVS by three amino acid positions. No evidence was obtained for a third cone pigment in either species.

It is implicit in many cone-specific ERG studies that the amplitude is proportional to the numbers of cones stimulated. The objective of these experiments was to test this idea by comparing ERGs obtained from different areas of the retina with histological data on cone-density distributions. The histology (Curcio et al., 1990) shows that the cumulative number of cones in the human retina increases exponentially with stimulus diameter between 0- and 40-deg eccentricity. L-, M-, and (L+M) cone-driven 30-Hz ERGs were obtained from a series of stimuli with one of the following configurations: (1) Circular stimuli of different angular subtense up to 70-deg diameter. (2) Annuli with 70-deg outer diameter but variable inner diameter. (3) Annuli of constant area but increasing eccentricity. Cone contrasts were equalized for each stimulus condition. The modulated and nonmodulated regions of the screen had the same mean hue and luminance. The data suggest that the L+M cone ERG amplitude increases with stimulus diameter in direct proportion to the estimated number of cones stimulated. Furthermore, the total L+M responses appear to be predicted from individual L and M responses by simple linear summation for both the disc and annular stimuli.

To investigate the effect of foveal inhomogeneities on sensitivity to chromatic stimuli, we measured simple reaction times (RTs) and detection thresholds to temporally and spatially blurred isoluminant stimuli at retinal eccentricities from 0 deg to 8 deg. Three color-normal subjects participated. Contrast gain was derived from the slope of the RT versus contrast function. With a Gaussian spatial distribution (S.D. = 0.5 deg) and modulation between white (CIE x,y,L = 0.31, 0.316, 12.5 cd.m−2) and blue (MBDKL 90 deg), gain was maximal at about 2-deg eccentricity and declined by approximately 1 log unit towards the center and the periphery. The red (0 deg) and green (180 deg) cardinal axes showed maximum gain in the center, whilst the yellow (270 deg) data were intermediate. Although the spatial extent of the Gaussian spot was much larger than the S-cone free zone, we wished to determine whether foveal tritanopia was responsible for the marked drop in sensitivity to the 90-deg stimulus. To align the color vector along a tritan line, we used a smaller disk (0.3 deg) with a blurred edge and measured detection threshold, rotating the vector until minimum central sensitivity was obtained. Other workers have used transient tritanopia or minimally distinct border to similar effect. By repeating this at different locations in color space, a group of vectors were obtained. These converged near to the S-cone co-punctal point, evidence that they lay along tritan confusion lines. These threshold findings were then confirmed using the RT-derived contrast gain function. The tritan vectors were less pronounced as stimulus size increased. With the vector optimized to produce foveal tritanopia, the RT gain versus eccentricity functions for the 90-deg and 270-deg stimuli both fell markedly in the center and periphery, and sensitivity peaked at about 3-deg eccentricity. There are some similarities between these findings and the underlying photoreceptor distributions. As a result, there is a greater difference in gain between red–green and blue–yellow systems in the center than in the near periphery. We conclude that the RT versus contrast function is a sensitive index of foveal opponency.

Shapiro et al. (2004) introduced a new visual effect (the induced contrast asynchrony) that demonstrates a perceptual separation between the response to a modulated light and the response to contrast of the light relative to background. The effect is composed of two physically identical disks, one surrounded by a dark annulus and the other by a light annulus. The luminance levels of both central disks were modulated in time, producing a stimulus with in-phase luminance modulation and antiphase contrast modulation. Observers primarily perceived the disks to be modulating asynchronously (i.e. they perceived the contrast), but at low temporal frequencies could also track the luminance level. Here we document that the induced contrast asynchrony disappears when the surrounds are achromatic and the center lights are modulated near the equiluminant axis. Observers viewed 1-deg-diameter disks embedded 2-deg-diameter achromatic surrounds. The chromaticity of the disks was modulated in time (1 Hz) along lines in an S versus Luminance cardinal color plane and an L-M versus Luminance cardinal color plane; observers responded as to whether the modulation appeared in phase. For all observers and both color planes, the lights appeared in phase most frequently at angles near the standard observer's equiluminant line and out of phase at angles further away from that line. Observers differed in the range of angles that produce the appearance of in-phase modulation. The results suggest that induced contrast asynchronies may be useful as a technique for equating luminance of disparate lights.

The aim of this study was to obtain information about single cone class driven activity in the inner and outer retina in humans. We examined outer retinal activity with the multifocal electroretinogram (mfERG) and inner retinal activity using multifocal oscillatory potentials (mfOPs). A standard (black-white) stimulus was used, as well as stimuli aimed at isolating a single photoreceptor class. The results of 10 trichromats were compared to those of 2 protanopes and 2 deuteranopes. At both retinal layers we find that trichromats show cone isolating response amplitudes that reflect the expected number of cones and that single- gene dichromats have a similar total number of functioning cones as trichromats. The ratio of the responses of the L- and M-cones is slightly smaller for the mfOPs than for the mfERGs. The results indicate that there are major changes in the gain of retinal signals after the inner plexiform layer.

This study tests the generality of previously demonstrated rod hue biases (red and blue biases at shorter wavelengths and a green bias at longer wavelengths) that cause the loci of the three spectral unique hues to shift to longer wavelengths. We found rod hue biases for 2-deg targets to be generally similar in magnitude and light-level dependence to those observed for 7.4-deg targets (the size most often studied) when measured at 7-deg eccentricity. The largest effects for both test sizes occurred at the lowest light levels tested, 1 log scotopic troland. All three rod hue biases were found with 0.6-deg targets, but were not reliably measurable at the lowest light levels and were reduced in magnitude and consistency across observers. The largest rod hue biases all occurred at the same scotopic light level, which corresponds to different photopic light levels for the three hue biases, because of differences in photopic and scotopic spectral sensitivity. This suggests that no single photopic light level will produce such large effects for all three rod hue biases. Finally, when the rod influence on a specific unique-hue locus was measured using photopically (rather than scotopically) constant stimuli, rod hue biases were still found but were more variable in magnitude and incidence across observers. We conclude that the rod hue biases we have previously described can be found with smaller stimuli, at somewhat higher light levels, and under photopically constant conditions, although our prior conditions tend to produce larger, more reliable rod hue biases.

This report describes a second-generation photostimulator with four primary lights that allows independent control of the stimulation of the four receptor types in the human eye. The new design uses LEDs (with light levels controlled by eight drivers that include voltage-to-frequency converters that provide 1-μs pulses at frequencies up to 250 kHz), with four center channels being combined by use of a fiber optic assembly, and likewise for four surround channels. Four fiber optic bundles are merged into a single bundle whose output is fed into a spatial homogenizer terminated by a diffuser. An interference filter is sandwiched between each LED and the fiber optic bundle. Two camera lenses collimate light from the diffusers, one for center and one for surround. The center-surround field configuration is formed by a photometric cube with a mirrored ellipse on the hypotenuse. A field lens places images of the diffusers in the plane of an artificial pupil. The fields are highly uniform. Following alignment and calibration, the center and surround fields are indistinguishable. An observer calibration procedure, designed to compensate for prereceptoral filtering, is shown by calculation to correct also for normal observer receptoral spectral sensitivity variation. With the instrument calibrated for the individual observer, a peripherally fixated 200-ms 40% contrast rod center field pulse, highly conspicuous under dark adaptation, is invisible following light adaptation.

A novel noise-masking technique was used to test D'Zmura and Knoblauch's (1998) idea that subjects employ off-channel looking in detecting chromatic test stimuli embedded in spatiotemporal chromatic noise. Detection thresholds were obtained for stationary, isoluminant, Gaussian-windowed (σx = σy = 2.25 deg; σt = 0.25 s), 135 deg (yellow/blue) or 160 deg (orange/blue–green), sinusoidal test gratings (11 deg × 11 deg; 0.75 cycle/deg) superimposed on each of a series of dynamic, random-check chromatic noise masks varying in azimuth in DKL space. Thresholds for detecting the test in the presence of these variable masks were again measured in the presence of an additional (auxiliary) noise mask created from colors falling on azimuths of 0 deg or 90 deg (135-deg test) or 0 deg or 135 deg (160-deg test). The effectiveness, kvar, of the variable noise masks in elevating grating detection thresholds was determined by fitting the detection data to the Pelli-Legge equation relating test detection energy to variable noise-mask energy: Et = K + kvarNvar. Differences in the calculated values of kvar for detection data obtained with and without the auxiliary masks were consistent with off-channel looking and were well accounted for by a simple model based on the idea that subjects possess a multichannel array of linear chromatic detectors spanning the isoluminant plane of DKL space, and they can choose the channel that has the highest signal-to-noise ratio.

We have studied the influence of chromatic adaptation upon the perceived visual position of a test stimulus using a Vernier alignment task. Maximum and minimum offsets in spatial position are generated when the adapting and test stimuli lie on the same and orthogonal axes in MBDKL color space, respectively. When the test stimuli lie on intermediate color axes, the measured positional shifts decrease as a function of the angular separation in color space (φ) from the adapting stimulus. At low stimulus contrasts, these shifts follow a sinusoidal function of φ and exhibit broad chromatic tuning and can be accounted for by a model in which the centroid is extracted from the linear combination of after-image, formed by the adapting stimulus, and the test stimulus. Such linear, broadband behavior is consistent with the response properties of chromatic neurons in the precortical visual pathway. At high contrast, and when adaptation gets closer to the S/(L+M) axis, the tuning functions become narrower and require sinusoids raised to increasingly higher exponents in order to describe the data. This narrowing of chromatic tuning is consistent with the tuning properties of chromatic neurons in the striate cortex, and implies the operation of a nonlinear mechanism in the combination of cone outputs.

We measured the regions of the equiluminant plane that are exploited by observers during a Yes/No detection task. The signal was a 640-ms Gaussian modulation (σt = 160 ms) of a Gaussian spatial patch (σs = 2.4 deg) presented in chromatically bivariate uniform noise. One component of the noise was along the direction axial with the signal in color space, the other perpendicular. Four signal directions were tested: along cardinal LM and S axes and two intermediate directions to which the cardinal axes were equally sensitive. The distribution of noise chromaticities from each trial was correlated with the observers' responses and the presence and absence of the signal to build a classification image of the distribution of chromaticities on which the decision of the observer was based. The images show a narrowly selective peak in the signal direction flanked by regions with a broader selectivity. These results raise the possibility that detection judgments are mediated by both linear and nonlinear mechanisms with peak sensitivities between the cardinal directions.

The spectral properties of chromatic-detection mechanisms were investigated using a noise-masking paradigm. Contrast-detection thresholds were measured for a signal with a Gaussian spatial profile, modulated in the equiluminant plane in the presence of spatial chromatic noise. The noise was distributed within a sector in the equiluminant plane, centered on the signal direction. Each stimulus consisted of two adjacent fields, one of which contained the signal, separated horizontally by a gap with the same average chromaticity as the uniform background. Observers were asked to judge on which side of the central fixation point the signal was displayed via a two-alternative, forced-choice (2AFC) paradigm. Contrast thresholds were measured for four color directions and three sector widths at increasing levels of the average energy of the axial component of the noise. Results show that contrast thresholds are unaffected by the width of the noise sector, as previously found for temporally modulated stimuli (D'Zmura & Knoblauch, 1998). The results are consistent with the existence of spectrally broadband linear-detection mechanisms tuned to the signal color direction and support the hypothesis of the existence of higher-order color mechanisms with sensitivities tuned to intermediate directions in color space.

Previous work (Nagy & Thomas, 2003) showed that signals in different Cardinal color mechanisms could be combined to facilitate search for a color target. Further investigation (Nagy et al., 2003) suggested that signals in one Cardinal color mechanism were used to select a subset of stimuli to be attended, while signals in second Cardinal mechanism were used to discriminate the stimuli within the selected subset. In the studies described below, we asked if observers could use color mechanisms tuned to directions other than the Cardinal directions to select and discriminate stimuli. Observers searched for a single target stimulus that differed in chromaticity from nine distractor stimuli. A two-alternative forced-choice procedure was used to estimate thresholds. Results were consistent with the hypothesis that color mechanisms tuned to many different directions in color space mediate discrimination, but suggest that only signals in Cardinal mechanisms can be used to select stimuli for attention.

Results imply that the selection of stimuli for attention on the basis of color may be mediated at the level of the lateral geniculate nucleus (LGN).

The optical density of the human crystalline lens progressively increases with age, the greatest increase in the visible spectrum being at short wavelengths. This produces a gradual shift in the spectral distribution of the light reaching the retina, yet color appearance remains relatively stable across the life span, implying that the visual system adapts to compensate for changes in spectral sensitivity. We explored properties of this adaptive renormalization by measuring changes in color appearance following cataract surgery. When the lens is removed, cataract patients often report a large perceptual shift in color appearance that can last for months. This change in color appearance was quantified for four cataract patients (63–84 years) by determining the chromaticity of stimuli that appeared achromatic before surgery, and at various intervals after surgery for up to 1 year. Stimuli were presented on a calibrated CRT as 9.5-deg spots, with 3-s duration and 3-s interstimulus intervals (ISIs). Chromaticity was adjusted by the subjects in CIE L*a*b* color space with luminance fixed at 32 cd/m2, on a dark background. We also estimated the optical density of the cataractous lens by comparing absolute scotopic thresholds from 410 nm to 600 nm before and after surgery. The results demonstrated that immediately following surgery there is a large increase in the short-wave light reaching the retina, mainly below 500 nm. The achromatic settings generally showed an initial large shift in the “yellow” direction after surgery that gradually (but never fully) returned to the original achromatic point before surgery. The shifts in the achromatic point occur over a number of months and appear to occur independently of the fellow eye.

If the relative phase of red and green modulated lights is changed, at low temporal frequencies the response of cells of the magnocellular (MC) pathway has been found to be minimal not to counterphase, chromatic modulation (as expected of a luminance mechanism) but shifted to some phase intermediate between luminance and chromatic modulation. The results could only be modeled by assuming interaction between achromatic and chromatic inputs to MC cells. The ‘phase shift’ resembled that seen with psychophysical threshold measurements using the same stimuli. Psychophysical results also showed that the phase shift is dependent on the chromaticity of a background. The results reported here show that the direction of the phase shift in MC cells is reversed by changing the background from long to short wavelengths and is consistent with psychophysical observations. Cell behavior was again modeled by assuming vector summation of achromatic and chromatic inputs. The reversal of phase-shift direction requires a reversal in polarity of the chromatic input. The underlying physiological mechanism may involve summation of chromatic signals of opposite polarity; if the relative size of these signals depends on the background, this may determine the direction of phase shift.