The International Colour Vision Society awarded the 2005 Verriest Medal to John D. Mollon, Professor of Visual Neuroscience at the University of Cambridge, UK. This award is bestowed by the Society to honor long-term contributions to the field of color vision. If the field of color vision were itself a rainbow, then Professor Mollon's contributions cover nearly its full spectrum, including the isolation and elucidation of basic chromatic coding mechanisms and the constraints that they impose on human (and more generally primate) visual performance, the genetic basis of spectral coding mechanisms, the ecological influences on and evolutionary origins of chromatic discrimination. He has been instrumental in the design of several new color vision tests and has extensively exploited abnormal models, both congenital and acquired, to further our understanding of normal mechanisms. He is especially appreciated for his keen and profound sense of the history of science, in particular with respect to the field of color vision. He has been a member of the society for over 25 years and is currently serving on its board of directors. He organized the 2001 ICVS meeting in Cambridge, celebrating the bicentennial of Thomas Young's lecture on color vision.

The International Colour Vision Society (ICVS) held its 18th biennial meeting in Lyon, France in July 2005. The ICVS, originally founded in 1967 as the International Research Group in Colour Vision Deficiencies and renamed in 1997, brings together vision scientists and clinicians with a common interest in color vision and color vision deficiencies. With significant technological advances that have permitted new and deeper questions about color vision to be addressed, the subject matter of recent meetings has expanded to include greater contributions from such areas as molecular genetics and evolution, retinal and cerebral imaging studies and computational modeling. The peer-reviewed papers in this volume span these newer and the more traditional topics of interest to the society, covering both applied and fundamental topics.

In 1789, when neither the physical basis of hue nor the retinal basis of color perception was established, the mathematician Gaspard Monte stated firmly that our color perceptions do not depend on the absolute value of the physical variable, but are influenced by the context and in particular by our estimate of the illuminant. He used this insight to explain color contrast effects and the Paradox of Monge (the desaturation of red objects seen through a red filter). He proposed that we can estimate the chromaticity of the illuminant in any scene because all surfaces reflect to us varying mixtures of (i) the body color and (ii) a specular component that represents the illuminant. He also realized that white objects have a special property: Provided that they are illuminated by a single illuminant, such objects exhibit no variation in chromaticity across their surface. Thus at least one of the unique hues exists as an external reference on which observers can agree. It is suggested that other unique hues may also have a basis in the external world.

Researchers studying surface color perception have typically used stimuli that consist of a small number of matte patches (real or simulated) embedded in a plane perpendicular to the line of sight (a “Mondrian,” Land & McCann, 1971). Reliable estimation of the color of a matte surface is a difficult if not impossible computational problem in such limited scenes (Maloney, 1999). In more realistic, three-dimensional scenes the difficulty of the problem increases, in part, because the effective illumination incident on the surface (the light field) now depends on surface orientation and location. We review recent work in multiple laboratories that examines (1) the degree to which the human visual system discounts the light field in judging matte surface lightness and color and (2) what illuminant cues the visual system uses in estimating the flow of light in a scene.

The role of chromatic information in 3-D shape perception is controversial. We resolve this controversy by showing that chromatic orientation flows are sufficient for accurate perception of 3-D shape. Chromatic flows required less cone contrast to convey shape than did achromatic flows, thus ruling out luminance artifacts as a problem. Luminance artifacts were also ruled out by a protanope's inability to see 3-D shape from chromatic flows. Since chromatic orientation flows can only be extracted from retinal images by neurons that are responsive to color modulations and selective for orientation, the psychophysical results also resolve the controversy over the existence of such neurons. In addition, we show that identification of 3-D shapes from chromatic flows can be masked by luminance modulations, indicating that it is subserved by orientation-tuned neurons sensitive to both chromatic and luminance modulations.

Psychophysical studies suggest that different colors have different perceptual status: red and blue for example are thought of as elementary sensations whereas yellowish green is not. The dominant account for such perceptual asymmetries attributes them to specificities of the neuronal representation of colors. Alternative accounts involve cultural or linguistic arguments. What these accounts have in common is the idea that there are no asymmetries in the physics of light and surfaces that could underlie the perceptual structure of colors, and this is why neuronal or cultural processes must be invoked as the essential underlying mechanisms that structure color perception. Here, we suggest a biological approach for surface reflection properties that takes into account only the information about light that is accessible to an organism given the photopigments it possesses, and we show that now asymmetries appear in the behavior of surfaces with respect to light. These asymmetries provide a classification of surface properties that turns out to be identical to the one observed in linguistic color categorization across numerous cultures, as pinned down by cross cultural studies. Further, we show that data from psychophysical studies about unique hues and hue cancellation are consistent with the viewpoint that stimuli reported by observers as special are those associated with this singularity-based categorization of surfaces under a standard illuminant. The approach predicts that unique blue and unique yellow should be aligned in chromatic space while unique red and unique green should not, a fact usually conjectured to result from nonlinearities in chromatic pathways.

To what extent do observers' judgments of surface color with natural scenes depend on global image statistics? To address this question, a psychophysical experiment was performed in which images of natural scenes under two successive daylights were presented on a computer-controlled high-resolution color monitor. Observers reported whether there was a change in reflectance of a test surface in the scene. The scenes were obtained with a hyperspectral imaging system and included variously trees, shrubs, grasses, ferns, flowers, rocks, and buildings. Discrimination performance, quantified on a scale of 0 to 1 with a color-constancy index, varied from 0.69 to 0.97 over 21 scenes and two illuminant changes, from a correlated color temperature of 25,000 K to 6700 K and from 4000 K to 6700 K. The best account of these effects was provided by receptor-based rather than colorimetric properties of the images. Thus, in a linear regression, 43% of the variance in constancy index was explained by the log of the mean relative deviation in spatial cone-excitation ratios evaluated globally across the two images of a scene. A further 20% was explained by including the mean chroma of the first image and its difference from that of the second image and a further 7% by the mean difference in hue. Together, all four global color properties accounted for 70% of the variance and provided a good fit to the effects of scene and of illuminant change on color constancy, and, additionally, of changing test-surface position. By contrast, a spatial-frequency analysis of the images showed that the gradient of the luminance amplitude spectrum accounted for only 5% of the variance.

Observers can generally make reliable judgments of surface color in natural scenes despite changes in an illuminant that is out of view. This ability has sometimes been attributed to observers' estimating the spectral properties of the illuminant in order to compensate for its effects. To test this hypothesis, two surface-color-matching experiments were performed with images of natural scenes obtained from high-resolution hyperspectral images. In the first experiment, the sky illuminating the scene was directly visible to the observer, and its color was manipulated. In the second experiment, a large gray sphere was introduced into the scene so that its illumination by the sun and sky was also directly visible to the observer, and the color of that illumination was manipulated. Although the degree of color constancy varied across this and other variations of the images, there was no reliable effect of illuminant color. Even when the sky was eliminated from view, color constancy did not worsen. Judging surface color in natural scenes seems to be independent of an explicit illuminant cue.

In normal viewing, the visual system effortlessly assigns approximately constant attributes of color and shape to perceived objects. A fundamental component of this process is the compensation for illuminant variations and intervening media to recover reflectance properties of natural surfaces. We exploited the phenomenon of transparency perception to explore the cortical regions implicated in such processes, using fMRI. By manipulating the coherence of local color differences around a region in an image, we interfered with their global perceptual integration and thereby modified whether the region appeared transparent or not. We found the major cortical activation due to global integration of local color differences to be in the anterior part of the parahippocampal gyrus. Regions differentially activated by chromatic versus achromatic geometric patterns showed no significant differential response related to the coherence/incoherence of local color differences. The results link the integration of local color differences in the extraction of a transparent layer with sites activated by object-related properties of an image.

Platyrrhine monkeys typically have only a single X-chromosome opsin gene. Alleles of this gene code for multiple versions of middle- to long-wavelength cone photopigments. X-chromosome inactivation provides heterozygous females with a retinal mosaic of cones containing either of two types of M and L pigment, thus establishing the photopigment basis for trichromatic color vision. This study examined the proportions of L and M cones created by this process. For that purpose, electroretinogram flicker photometry was used to obtain complete spectral sensitivity functions from 60 heterozygous female monkeys drawn from seven genera of platyrrhine monkeys. To obtain estimates of cone proportions, these functions were subsequently fit with linear combinations of L and M cone fundamentals that were derived from similar recordings made on conspecific animals having only one type of M/L pigment. Consistent with a random X-chromosome inactivation process, the average L:M cone weighting across the sample was close to unity. At the same time, there were significant individual variations in L:M cone proportions. The genesis of this variation and its implications for seeing are discussed.

The key steps in the evolution of full trichromatic color vision in primates include duplication of the ancestral pigment gene to form the L and M pigment gene array on the X chromosome, mutually exclusive expression of the L and M pigment genes in cone photoreceptors, and formation of a retinal mosaic with randomly distributed L and M cones. Previous work using transgenic mice has indicated that a locus control region adjacent to this array of genes plays an important role in their mutually exclusive expression in respective cone cells (Smallwood et al., 2002). However, the mechanism by which this is accomplished is unknown. We searched for a cellular model system to investigate the mechanism of this mutually exclusive expression. We previously showed that the undifferentiated human retinoblastoma cell line WERI expresses L and M cone opsin but not rod opsin genes. We now show that WERI cells express the L and M pigment genes in a mutually exclusive manner, in that either L or M pigment mRNA is expressed in a single cell. Importantly, clonal analysis showed that single WERI cells that express either L or M generate, upon cell division produce, a mixed population of L- or M-expressing cells. These results indicate, first, that cell division resets L or M pigment gene expression, most likely due to disassembly and reassembly of LCR-promoter DNA-protein complexes during cell division. Second, a retinal mosaic with near-random distribution of L and M cones may have been generated automatically after duplication of the ancestral gene to form the L and M pigment genes. Third, determination of L and M cone identity may not require external molecular cues during differentiation, and is consistent with the idea that L and M cones are not intrinsically different.

The topographical distributions of the relative ratio of long- (L) and middle- (M) wavelength sensitive cone opsin messenger RNA (mRNA) in human and baboon retinas were mapped using real-time polymerase chain reaction. The L:M mRNA ratio increased in a central-to-peripheral gradient in both species, being quite pronounced for humans.

A model of the development of L/M cone ratios in the Old World primate retina is presented. It is supposed that during gestation, the cone cycles randomly between states in which it transcribes either L or M opsin. The current state determines and increases the probability that it will transcribe the same opsin in future cycles. These assumptions are sufficient to formalize the process as a Markov chain that can be modeled as an urn containing two types of balls, L and M. Drawing one ball results in the increase of its species and the decrease of the other. Over the long run, the urn will become populated with a single type of ball. This state corresponds to the photoreceptor adopting a fixed identity for its lifetime. We investigate the effect of the number of states and the rule that regulates the advantage of transition toward one cone type or another on the relation between fetal and adult L/M cone ratios. In the range of 100 to 1000 states, small variations of the initial L/M ratio or the transition advantage can each generate large changes in the final L/M ratio, in qualitative accord with the variation seen in human adult retinas. The time course to attain stable L/M ratios also varies with these parameters. If it is believed that the cycling follows a circadian rhythm, then final L/M cone ratios would be expected to stabilize shortly after birth in the human being and the macaque.

Congenital achromatopsia is a stationary retinal disorder with autosomal recessive inheritance that is characterized by loss of color discrimination, low visual acuity, photophobia, and nystagmus. This disorder has been shown to be associated with CNGA3, CNGB3, and GNAT2 mutations, and the frequency of mutations in the CNGA3 gene (encoding α subunit of the cone-specific cGMP-gated cation channel) was 23–33% in European populations. The aim of this study was to test the hypothesis that CNGA3 mutations are also responsible for congenital achromatopsia in Japanese patients. DNA from venous blood samples from a total of 14 patients from 13 Japanese pedigrees was prepared. Mutation screening of the CNGA3 gene was performed using direct sequencing and PCR-single-strand conformation polymorphism analysis. Compound heterozygous missense mutations (p.R436W and p.L633P, the latter of which was novel) were identified in one patient only, a 22-year-old female. Neither of these two mutations was found in 150 Japanese control individuals. The patient's parents and sister carried one of these mutations each but were not affected. No mutations in the CNGB3 or GNAT2 genes were identified in the patient. Clinically, best-corrected visual acuity was 0.1 in both eyes. No specific findings were obtained in funduscopy. Optical coherence topography revealed a normal foveal thickness but a 20% decrease in parafoveal thickness. Ganzfeld full-field electroretinograms (ERGs) showed normal responses in rod and mixed rod-plus-cone ERGs but no response in cone or 30-Hz flicker ERGs. Spectral sensitivity on a white background revealed a curve with only one peak at around 500 nm, which fits the absorption spectrum of human rhodopsin. L633, conserved among vertebrate orthologs of human CNGA3, is a hydrophobic residue forming part of the carboxy-terminal leucine zipper (CLZ) domain, which is functionally important in the mediation of intracellular interactions. To our knowledge, this is the first report of a Japanese complete achromat with CNGA3 mutations, and of any patient with a missense mutation within the CLZ domain. The outcome suggests low frequency (7%, 1/14) of CNGA3 mutations in Japanese patients.

Inherited tritan color vision deficiency is caused by defects in the function of the short-wavelength-sensitive (S) cones. This heterozygous group of disorders has an autosomal dominant pattern of inheritance. Amino acid variations of the S cone opsin are rare and all that have been identified thus far are associated with inherited tritan color vision defects. Here we report the identification of a 30-year-old male who made errors on standard color vision tests consistent with the presence of a mild tritan color vision deficiency. We tested the hypothesis that his color vision impairment was due to a mutation in the S cone photopigment gene. He was found to be heterozygous for a mutation that caused the amino acid proline to be substituted in place of a highly conserved leucine at amino acid position 56 in the S cone opsin. This mutation was absent in 564 S cone photopigment genes from 282 subjects who did not make tritan errors. Thus, we conclude that this mutation disrupts the normal function of S cones.

In normal trichromats, the long- (L) and middle-wavelength-sensitive (M) pigment genes are arranged in a head-to-tandem array on the X chromosome. Two amino acids at positions 277 and 285, encoded by exon 5 of the L and M genes, respectively, are essential for the spectral difference between L and M pigments whose spectral peaks are at approximately 560 and 530 nm. Intragenic or intergenic unequal crossing-over commonly occurs between the highly homologous L and M genes, resulting in red-green color vision deficiencies. The dichromacy is usually associated with a single L gene for deuteranopia or a single 5′ L-M 3′ hybrid gene with M-gene exon 5 for protanopia. We clinically diagnosed a total of 88 male dichromats using a Nagel model I anomaloscope, which included one unclassified subject in addition to 31 protanopes and 56 deuteranopes. The objective of this study was to characterize the phenotype of the subject and to determine the genotype of his X-linked pigment genes. The subject accepted not only any red-green mixture but also an extended yellow-scale range at each matching point (i.e. 20 to 32 scale units at the green primary and 3.5 to 6 scale units at the red primary). The slopes of regression lines were in the range of −0.34 to −0.23, while the mean slopes for the protanopes and deuteranopes were −0.38 and −0.01, respectively. Spectral sensitivity tests showed that the subject's curve was shifted between the protanope and deuteranope curves. Molecular analysis revealed a novel form of a single pigment gene with a unique arrangement of exon 5 (Y277 from the L gene and A285 from the M gene). The predicted λmax (541 to 546 nm) of the unique pigment was closer to the M than to the L pigment. Our outcome suggests that intragenic unequal crossing-over may have occurred between amino acid positions 279 and 283.

Shift in the wavelength of peak sensitivity of the cone photo pigments is a major cause of inter-individual variations in the Rayleigh match. Normal color observers performed multiple Rayleigh matches (i.e., a series of seven Rayleigh-type color matches using various sets of test and primary lights) in order to derive estimates of the M- and L-photo pigment wavelengths of maximum absorbance. We predicted the ratio of incident radiances Pr/Pg for a range of plausible values in terms of wavelength of the peak sensitivity of the M-cone and L-cone. An algorithm allowed us to adjust the values of the peak wavelength of the cone photo pigments in order to minimize the sum of squared differences between predicted and real results of matches. To create candidate M-cone and L-cone photo pigments that best predict the set of equation values, we used the low density spectral absorbance curves of the M-cone and L-cone photo pigments respectively, as tabulated by Stockman and Sharpe (2000), and shifted each template along a logarithmic wavelength scale. For all observers, the system of seven Rayleigh-like equations converges to a unique solution. M- and L-cone photo pigments are estimated separately. The distribution of the wavelength of maximum sensitivity of the photo pigments includes a cluster. Only a few women's results lie outside the cluster. The choice of the template has a considerable influence on the convergence of the algorithm.

In mammals, cone photoreceptor subtypes are thought to establish topographies that reflect the species-relevant properties of the visual environment. Middle- to long-wavelength-sensitive (M) cones are the dominant population and in most species they form an area centralis at the visual axis. Short-wavelength-sensitive (S) cone topographies do not always match this pattern. We here correlate the interrelationship of S and M cone topographies in representatives of several mammalian orders with different visual ecology, including man, cheetah, cat, Eurasian lynx, African lion, wild hog, roe deer, and red deer. Retinas were labeled with opsin antisera and S and M cone distributions as well as S/M cone ratios were mapped. We find that species inhabiting open environments show M cone horizontal streaks (cheetah, pig, deer). Species living in structured habitats (tiger, lynx, red deer) have increased S cone densities along the retinal margin. In species with active vision (cheetah, bear, tiger, man), S cone distributions are more likely to follow the centripetal M cone gradients. Small species show a ventral bias of peak S cone density which either matches the peak of M cone density in a temporal area centralis (diurnal sciurid rodents, tree shrews) or not (cat, manul, roe deer).

Thus, in addition to habitat structure, physical size and specific lifestyle patterns (e.g. food acquisition) appear to underlie the independent variations of M and S cone topographies.

The objective of the present work was to determine the interaction of cone inputs in the response of horizontal cells using heterochromatic flicker photometry (HFP). Intracellular electrophysiological recordings were made in horizontal cells of isolated retinae of carp maintained in physiological solution, with the receptor side up. Sharp glass microelectrodes filled with 3 M KCl solution with resistances between 100 and 120 MΩ were used. Stimuli comprised six cycles of two 6-Hz sinusoidal light waves in counterphase adjusted for the same number of quanta: a green light (550 nm) from a monochromator with a Xenon lamp and an LED red light (628 nm). The stimulation program consisted of 10 steps with the 550-nm wave at constant amplitude, while the 628-nm wave varied in increments of 10% up to 100%, followed by another 10 steps with the 628-nm wave at constant amplitude while the 550-nm wave varied in increments of 10% up to 100%. We recorded responses from four different horizontal cell classes: H1 (monophasic, broadband, n = 37), H2 (biphasic, red-green color-opponent, n = 13), and H3 (biphasic, blue-yellow color-opponent, n = 2) cone horizontal cells; and RH (monophasic, broadband, n = 3) rod horizontal cells. H1 and RH horizontal cells showed a similar cancellation point at a heterochromatic mixture consistent with mixed inputs from 630- and 550-nm cones. No cancellation point was found for the H2 cell class. Fish H1 cells add cone inputs and signal “luminance” in light levels appropriate for cone stimulation. The same occurs with RH cells, which also signal “luminance,” but in light levels appropriate for rod work. For both cell classes there is an HFP cancellation point occurring at a combination of 628-nm and 550-nm lights in opposing phase that leads to the cancellation of the cell's response. No cancellation was found for H2 and H3 cells, which are the chromatically opponent horizontal cells in lower vertebrates.

We recently developed a new technique to measure cone inputs to visual neurons and used this technique to seek short-wavelength-sensitive (S) cone inputs to parasol, magnocellular (MC) and midget, parvocellular (PC) ganglion cells. Here, we compare our physiological measurements of S-cone weights to those predicted by a random wiring model that assumes cells' receptive fields receive input from mixed cone types. The random wiring model predicts the average weights of S-cone input to be similar to the total percentage of S-cones but with considerable scatter, and the S-cone input polarity to be consistent with that of PC cells' surround and of MC cells' center. This is not consistent with our physiological measurements. We suggest that the ganglion cells' receptive fields may have a mechanism to avoid S-cone inputs, as is the case in the H1 horizontal cells. Previous reports of S-cone inputs, in particular substantial input to MC cells, are likely to reflect variation in prereceptoral filtering and/or the failure to correct for variation in macular pigment.