Seeing unique hues

Hue can be described by four separate sensations of red (R), green (G), yellow (Y), and blue (B). These are combined in the spectrally opponent RG and YB mechanisms, whose null points correspond to the unique sensations of Y, G, B. Participants used a form of magnitude estimation to describe color appearance of light flashes that were systematically varied in size, luminance, duration, purity, and retinal eccentricity. Wavelengths of the unique hues were derived from the hue and saturation scaling functions. Only unique Y remained invariant across all the viewing conditions. The shifts in unique hues with test conditions place strong restrictions on models of how the RG and YB mechanisms are assembled. Despite polymorphism of L and M cones and variation of their ratios across participants and across the retina, the frequency distribution of unique Y was very narrow, implying some reweighting of cone inputs to individuals' RG mechanisms.     

The relationship between peripherally matched invariant hues and unique hues: a cone-contrast approach

A characteristic shift in hue and saturation occurs when colored targets are viewed peripherally compared with centrally. Four hues, one in each of the red, blue, green, and yellow regions of color space, remain unchanged when presented in the peripheral field. Apart from green, these peripherally invariant hues correspond almost exactly in color space with the unique hues. We explore this puzzling observation using asymmetric color-matching and color-naming experiments and computing cone contrasts for peripheral and central stimuli. We find that the difference between cone contrasts for the peripheral and central stimuli reaches a maximum at the chromatic axis corresponding to peripherally invariant green. We speculate that the effect is linked to a weakened signal from M-cones and probably associated with a reduced number of M-cones in peripheral retina.     

Perceived shifts in saturation and hue of chromatic stimuli in the near peripheral retina

Using an asymmetric color matching technique, we measured the perceived changes that occur in the saturation and hue of colored stimuli at different eccentricities within the central 25 degrees of the human retina in nine color-normal subjects. A cone-opponent-based vector model was used to compute the activity of the L-M and S-(L+M) channels. The results show that a large proportion of the shifts in saturation and hue that occur with increasing retinal eccentricity are mirrored by decreased activity of the L-M channel. In comparison, the contribution of the S cone-opponent system undergoes relatively little change within the central 20 degrees . In addition, we also found that changes in saturation and hue are different from each other in terms of their variation across color space and their variation with stimulus size. Our findings suggest that perceived shifts in saturation and hue are mediated largely via the reduction in activation of the L-M cone-opponent channel but that saturation and hue might be subject to different retinal and/or cortical influences that contribute to their differing size dependencies in the peripheral retina.     

Color appearance in the entire visual field: color zone map based on the unique hue component

To provide the fundamental data for a color zone map, the color appearances of nearly unique hue stimuli presented over the entire visual field were qualitatively and quantitatively evaluated by hue and saturation judgments, blackness evaluation, and categorical color naming. The hue of red and green stimuli shifts toward a unique yellow, while that of yellow and blue does not change with an increase in eccentricity. The saturation of all the stimuli falls with an increase in the eccentricity in all directions. On the basis of the unique hue component, color zone maps for red, dark yellow, yellow, green, and blue stimuli are drawn. All the color zone maps extend over a wider region in the temporal and lower directions than in the nasal and upper directions of the visual field. The results are compared with the color zones of previous studies. The relationship between the color zones and the color categorization, as well as the underlying mechanisms of reduced saturation and hue shift, is discussed.     

Variations in normal color vision. IV. Binary hues and hue scaling

We used hue cancellation and focal naming to compare individual differences in stimuli selected for unique hues (e.g., pure blue or green) and binary hues (e.g., blue-green). Standard models assume that binary hues depend on the component responses of red-green and blue-yellow processes. However, variance was comparable for unique and binary hues, and settings across categories showed little correlation. Thus, the choices for the binary mixtures are poorly predicted by the unique hue settings. Hue scaling was used to compare individual differences both within and between categories. Ratings for distant stimuli were again independent, while neighboring stimuli covaried and revealed clusters near the poles of the LvsM and SvsLM cardinal axes. While individual differences were large, mean focal choices for red, blue-green, yellow-green, and (to a lesser extent) purple fall near the cardinal axes, such that the cardinal axes roughly delineate the boundaries for blue vs. green and yellow vs. green categories. This suggests a weak tie between the cone-opponent axes and the structure of color appearance.     

Variations in normal color vision. II. Unique hues

We examined individual differences in the color appearance of nonspectral lights and asked how they might be related to individual differences in sensitivity to chromatic stimuli. Observers set unique hues for moderately saturated equiluminant stimuli by varying their hue angle within a plane defined by the LvsM and SvsLM cone-opponent axes that are thought to characterize early postreceptoral color coding. Unique red settings were close to the +L pole of the LvsM axis, while green, blue, and yellow settings clustered along directions intermediate to the LvsM and SvsLM axes and thus corresponded to particular ratios of LvsM to SvsLM activity. Interobserver differences in the unique hues were substantial. However, no relationship was found between hue settings and relative sensitivity to the LvsM and SvsLM axes. Moreover, interobserver variations in different unique hues were uncorrelated and were thus inconsistent with a common underlying factor such as relative sensitivity or changes in the spectral sensitivities of the cones. Thus for the moderately saturated lights we tested, the unique hues appear largely unconstrained by normal individual differences in the cone-opponent axes. In turn, this suggests that the perceived hue for these stimuli does not depend on fixed (common) physiological weightings of the cone-opponent axes or on fixed (common) color signals in the environment.     

Real-world stimuli show perceived hue shifts in the peripheral visual field

Certain hues undergo shifts in their appearance when they are viewed by the peripheral retina. This has often been shown on a 3-primary color CRT monitor. To investigate the possible role of metamerism, we replicated our peripheral color matching experiments using Munsell paper stimuli viewed under real and simulated daylight (using a 3-primary projection system). Using stimuli of constant value and chroma (7/4), observers adjusted the hue of a 3 deg target presented 18 deg nasally, until it matched a 1 deg target presented 1 deg nasally. The magnitude and pattern of measured hue shifts were similar to those measured using CRT stimuli. We conclude that the perceived hue shifts that have previously been reported in the peripheral retina are independent of the nature of the stimulus and of the illuminant.     

How well are color components of samples of the Natural Color System estimated?

The aim of this study was to determine how accurately color-normal subjects that have received basic information about, but do not have practical experience with, the Natural Color System (NCS) can estimate the Heringian components of a representative selection of samples. Twenty-five color-normal subjects, taking part in two trials with at least a 24 h gap between assessments, selected four samples representing individual unique hues (uHs) from a set of 40 highly chromatic NCS samples on a rotatable tray. The samples selected for assessment of components were displayed to the subjects who estimated the hue components of 16 high-chroma samples, hue and white/black components of 16 tonal color samples, and three achromatic samples with different blackness values. Variability in selection of samples representing uHs as well as the relationship between the subjects' estimates of unique hue components and the defined values of the system was obtained. It was found that hues alone are easier to correctly estimate than hues together with white and black and that the components of colors of higher chroma are easier to estimate accurately than those of lower chroma. It was also found that, for R and G, the mean uH choices of subjects differed very little from the NCS's R and G, whereas selections for yellow and blue deviated, the former by 1.22 hue steps (slightly greener than G90Y), and the latter by 1.36 hue steps (represented approximately by R85B). This may impact the accuracy of color models that employ NCS unique hues.     

Bezold-Brücke effect in normal trichromats and protanopes

Luminance-dependent change in color appearance--the Bezold-Brücke effect--was investigated in protanopes and related to that in normal trichromats. Spectral lights were presented at six luminance levels covering mesopic, low, and high photopic vision-across three log steps from 0.76 to 760 Td. To judge color appearance, a variant of the color-naming method was used with four primary basic color terms and a "White" response. This modification enabled us to examine apparent saturation changes along with the Bezold-Brücke hue shift. Color-naming frequency functions were acquired across ten presentations of each stimulus. Since protanopes name colors idiosyncratically, changes in color appearance cannot be quantified directly from the color-naming functions. To circumvent the difficulty, these functions were transformed into color similarity measures for analysis with multidimensional scaling purported to reconstruct individual color spaces. In these, luminance-dependent shifts in color appearance were represented by means of geometric displacements. We found that for normal trichromats, shifts measured in this way agreed with those derived in our study directly, and with the hue shifts reported in earlier studies. For protanopes, contrary to some models of dichromatic vision, changes in color appearance are significant and indicate superimposed shifts in hue and saturation. The results obtained for normal trichromats, especially for protanopes, imply that nonlinearity in the yellow-blue opponent system is insufficient to explain the Bezold-Brücke effect, given the nature of the saturation shift and the demonstrated divergence between unique hues and invariant hues.     

A tritan Waldo would be easier to detect in the periphery than a red/green one: evidence from visual search

In a color naming task from 0° to 55° eccentricity, we found that red/green performance (n=10 subjects) declines around 40° eccentricity, 5° earlier than does tritan performance (main effect of color, p=0.009; eccentricity, p<0.001; interaction, p=0.005). In a feature visual search task (e.g., red target dot among green distractor dots; twelve 2.5° diameter dots; 0, 20, and 45° eccentricity; 12 subjects), performance was significantly more impaired for red/green than for tritan stimuli, especially in the periphery (main effect of color, p=0.007; eccentricity, p<0.001; interaction, p=0.003). This effect occurred even following a rod bleach. Our results are consistent with influences from both the retina (especially random rather than selective peripheral cone input to midget ganglion cells for red/green perception, and selective cone input to small bistratified cells for tritan perception) and the cortex (differential cortical magnification across the two chromatic axes).     

Rod hue biases produced on CRT displays

Studies of rod hue biases using monochromatic stimuli have shown that rod stimulation can shift the balance of hues at mesopic light levels. We found that the CRT display produced all three previously identified rod hue biases, which shifted the loci of all four unique hues at low mesopic light levels. Rod hue biases occurred at 2.6 cd/m(2) for some observers but not at 26 cd/m(2). At optimal light levels below 0.5 cd/m(2), rod hue biases varied among observers but generally (1) enhanced green versus red at unique yellow and sometimes at unique blue, (2) enhanced blue versus yellow at both unique green and unique red, and (3) enhanced red versus green at unique blue. Rod hue biases persisted for some observers even for smaller foveal stimuli.     

Direction in the color plane as a factor in chromatic flicker and chromatic motion

A percept of motion results when a chromatic grating, formed from a spatial alternation between two isoluminant hues, drifts across the visual field. With hue pairs chosen to be equally subjectively dissimilar, the motion is greater for alternation along some directions in color space (orange/blue) than others (green/purple), suggesting a specific interaction between the (L-M) and S(0) chromatic opponent channels. This phenomenon was explored systematically by choosing 24 pairs of hues across the color circle and using the method of paired comparisons to scale their movement-inducing contrast. The flicker-inducing contrast observed from rapid alternation between the pairs was measured in the same way. Both phenomena consistently drew upon both chromatic channels, though in different proportions, as if chromatic and temporal variation information are multiplexed along motion-processing pathways. Border-distinctness data were also collected to isolate the (L-M) channel.     

Effect of stimulus intensity on the sizes of chromatic perceptive fields

The effects of intensity on chromatic perceptive field size were investigated along the horizontal meridian at 10 degrees temporal eccentricity by manipulating stimulus intensity from 0.3 to 3.3 log trolands. Following light adaptation, observers described the hue and saturation of monochromatic stimuli (440-660 nm, in 10 nm steps) for a series of test sizes (0.098-3 degrees) presented along the time period associated with the cone plateau of the dark-adaptation function. Perceptive field sizes of the four elemental hues (red, green, yellow, and blue) and the saturation component were estimated by three observers at each intensity level for each wavelength. In general, perceptive field sizes of blue and red are the smallest, and yellow and green are the largest. Furthermore, perceptive field sizes of all four hues decrease with increasing stimulus intensity, though the absolute change is largest for green and yellow. The decrease in size with increase in intensity cannot be completely explained in terms of saturation or rod signals and is likely, then, attributable to a cone-based mechanism.     

Variations in normal color vision. III. Unique hues in Indian and United States observers

Basic color categories are thought to share a common pattern across linguistic groups, yet the focal colors defining those categories can vary substantially within any single group. We asked whether focal colors can also differ systematically across different groups of individuals living in potentially different color environments, by measuring focal and unique hues for observers in India and the United States. Differences between groups were generally small relative to the within-group variations, consistent with a strong common basis for color naming across diverse contexts. However, for most hues the average settings differed significantly across subpopulations. These differences persisted across testing conditions and thus probably reflect longer-term contextual influences on color appearance judgments. They suggest that while color categories may be qualitatively similar, precisely how the hue spectrum is parsed may differ quantitatively across different populations of observers. Both the between-group and the within-group differences are inconsistent with the differences predicted by common peripheral sources of variation in color vision (e.g., in lens or macular pigment) and may reflect an influence of environmental or cultural differences in focal color choices.     

The perceptual balance of color

The cone contrasts carrying different dimensions of color vision vary greatly in magnitude, yet the perceived contrast of color and luminance in the world appears similar. We examined how this perceptual balance is adjusted by adaptation to the contrast in images. Observers set the level of L vs. M and S vs. LM contrast in 1/f noise images to match the perceived strength of a fixed level of luminance contrast. The perceptual balance of color in the images was roughly consistent with the range of contrast characteristic of natural images. Relative perceived contrast could be strongly biased by brief prior exposure to images with lower or higher levels of chromatic contrast. Similar adaptation effects were found for luminance contrast in images of natural scenes. For both, observers reliably chose the contrast balance that appeared correct, and these choices were rapidly recalibrated by adaptation. This recalibration of the norm for contrast could reflect both changes in sensitivity and shifts in criterion. Our results are consistent with the possibility that color mechanisms adjust the range of their responses to match the range of signals in the environment, and that contrast adaptation plays an important role in these adjustments.     

Aftereffect of contrast adaptation to a chromatic notched-noise stimulus

One of the most challenging topics in the study of human color vision is the investigation of the number of hue-selective channels that are necessary for the representation of color appearance at the post-opponent level and the bandwidth of their sensitivity. The present study aims to elucidate this issue by using a chromatic version of the notch-filtered noise (herein, notched-noise) stimulus for contrast adaptation. After adaptation to this stimulus, some color-sensitive mechanisms that selectively respond to missing hues in the notched-noise stimulus may remain sensitive, while the other mechanisms may be desensitized. The shifts in the color appearance of a gray test field after the adaptation to such a notched noise were measured using the method of adjustment. The results showed systematic shifts in the hue and saturation. They showed neither point nor line symmetric profiles with respect to the achromatic point in an isoluminant plane. The fittings of the results, obtained by using a tiny numerical model for assessing the hue-selective mechanisms, suggested that there are at least two narrowly tuned and at least three broadly tuned mechanisms. The narrowly tuned mechanisms are the most sensitive along the blue and yellow directions. The present study confirmed the variation of multiple channels at the post-opponent level and suggested that this variation may be responsible for the processing of color appearance.     

Hue discrimination, unique hues and naming

The hue discrimination curve (HDC) that characterizes performances over the entire hue circle was determined by using sinusoidally modulated spectral power distributions of 1.5 c/300 nm with fixed amplitude and twelve reference phases. To investigate relationship between hue discrimination and appearance, observers further performed a free color naming and unique hue tasks. The HDC consistently displayed two minima and two maxima; discrimination is optimal at the yellow/orange and blue/magenta boundaries and pessimal in green and in the extra-spectral magenta colors. A linear model based on Müller zone theory correctly predicts a periodical profile but with a phase-opponency (minima/maxima at 180° apart) which is inconsistent with the empirical HDC's profile.     

Evaluation of single-pigment shift model of anomalous trichromacy.

The spectral sensitivity of the visual photopigments, the interobserver variability in color judgments, and the spectral locus of unique yellow provide three major problems for accounts of X-chromosomal-linked anomalous trichromacy. According to the single-pigment hypothesis, the primary defect in anomalous trichromacy is a wavelength shift in the peak sensitivity of one of the three visual photopigments. We show that this shift results in reduction of the anomalous trichromat's r-g opponent chromatic channel. The distribution of response variability in Rayleigh equation match widths due to factors other than the spectral characteristics of the photopigments is similar in normal and anomalous trichromats. When normal and anomalous trichromats make hue estimations of sets of stimuli designed to contain similar chromatic information, their judgments show similar variability. Calculation of the r-g opponent chromatic channel can provide correct predictions of the spectral loci for unique yellow for anomalous trichromats.     

Perception of forbidden colors in retinally stabilized equiluminant images: an indication of softwired cortical color opponency?

In color theory and perceptual practice, two color naming combinations are forbidden-reddish greens and bluish yellows-however, when multicolored images are stabilized on the retina, their borders fade and filling-in mechanisms can create forbidden colors. The sole report of such events found that only some observers saw forbidden colors, while others saw illusory multicolored patterns. We found that when colors were equiluminant, subjects saw reddish greens, bluish yellows, or a multistable spatial color exchange (an entirely novel perceptual phenomena); when the colors were nonequiluminant, subjects saw spurious pattern formation. To make sense of color opponency violations, we created a soft-wired model of cortical color opponency (based on winner-take-all competition) whose opponency can be disabled.     

Senescent changes in parafoveal color appearance: saturation as a function of stimulus area.

The chromatic content (saturation) of monochromatic stimuli (480, 505, 577, and 650 nm) was scaled as a function of field size at three different retinal locations by 58 observers ranging from 18 to 83 yr of age. The different retinal locations (6 deg nasal, 2.5 deg inferior and 6 deg temporal eccentricity) were chosen according to anatomical studies demonstrating different degrees of senescent losses of cones or ganglion cells. Nine field sizes were tested, ranging from 0.0096 to 0.96 deg in diameter. The subjects used a percentage scale to judge the saturation of the flashed stimulus presentations (2 s on, 5 s off). The data analysis demonstrated that older observers require larger field sizes than younger observers to perceive hue as well as larger field sizes to reach the same level of scaled saturation. The spatial dependency of color appearance for younger and older observers was not correlated with senescent losses in retinal cells reported for the different retinal locations. The data were modeled by using an impulse-response function (i.e., Naka-Rushton equation) so that perceptive fields could be compared to electrophysiological measures of receptive fields or dendritic fields of retinal and cortical cells.