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Ambiguity is implicit in neural representations of the physical world. Previous work has examined how the visual system resolves ambiguous neural signals that represent various features, such as the percept resulting from rivalrous chromaticities or forms. Relatively little is known, however, about the contribution of unambiguous neural representations to perceptual resolution of ambiguous ones. This is addressed here by measuring perceptual resolution of ambiguity by grouping, which is operationalized as the tendency for multiple similar ambiguous representations to be seen as identical to each other. Multiple chromatically ambiguous representations were created using interocular switch rivalry and presented together with a nearby but separate unambiguous (non-rivalrous) chromaticity. The magnitude of grouping the chromatic regions was compared when ambiguous regions were seen alone versus with unambiguous regions seen simultaneously. Contrary to prevailing theory that the resolution of the ambiguous percepts would follow the unambiguous ones, the ambiguous chromatic regions consistently appeared identical to each other, but their appearance was not found to be attracted to the unambiguous color percept. This supports the proposition that the ambiguity itself in a neural representation is a linking feature contributing to perceptual disambiguation.

Color is a perceptual construct that arises from neural processing in hierarchically organized cortical visual areas. Previous research, however, often failed to distinguish between neural responses driven by stimulus chromaticity versus perceptual color experience. An unsolved question is whether the neural responses at each stage of cortical processing represent a physical stimulus or a color we see. The present study dissociated the perceptual domain of color experience from the physical domain of chromatic stimulation at each stage of cortical processing by using a switch rivalry paradigm that caused the color percept to vary over time without changing the retinal stimulation. Using functional MRI (fMRI) and a model-based encoding approach, we found that neural representations in higher visual areas, such as V4 and VO1, corresponded to the perceived color, whereas responses in early visual areas V1 and V2 were modulated by the chromatic light stimulus rather than color perception. Our findings support a transition in the ascending human ventral visual pathway, from a representation of the chromatic stimulus at the retina in early visual areas to responses that correspond to perceptually experienced colors in higher visual areas.

A long-standing question in the field of vision research is whether scalp-recorded EEG activity contains sufficient information to identify stimulus chromaticity. Recent multivariate work suggests that it is possible to decode which chromaticity an observer is viewing from the multielectrode pattern of EEG activity. There is debate, however, about whether the claimed effects of stimulus chromaticity on visual evoked potentials (VEPs) are instead caused by unequal stimulus luminances, which are achromatic differences. Here, we tested whether stimulus chromaticity could be decoded when potential confounds with luminance were minimized by (1) equating chromatic stimuli in luminance using heterochromatic flicker photometry for each observer and (2) independently varying the chromaticity and luminance of target stimuli, enabling us to test whether the pattern for a given chromaticity generalized across wide variations in luminance. We also tested whether luminance variations can be decoded from the topography of voltage across the scalp. In Experiment 1, we presented two chromaticities (appearing red and green) at three luminance levels during separate trials. In Experiment 2, we presented four chromaticities (appearing red, orange, yellow, and green) at two luminance levels. Using a pattern classifier and the multielectrode pattern of EEG activity, we were able to accurately decode the chromaticity and luminance level of each stimulus. Furthermore, we were able to decode stimulus chromaticity when we trained the classifier on chromaticities presented at one luminance level and tested at a different luminance level. Thus, EEG topography contains robust information regarding stimulus chromaticity, despite large variations in stimulus luminance.

A coherent percept of our visual world is important for functioning. Ambiguities, however, are implicit in visual neural representations and must be resolved for stable perception of objects and scenes. Grouping processes can link multiple neurally ambiguous fragments across the visual field. Experiments here determined how multiple visual features of each fragment contribute to perceptual resolution of ambiguity by grouping. Chromatic interocular-switch rivalry, a technique for presenting competing dichoptic images, was used to induce ambiguous neural representations for equiluminant chromatic discs and gratings. Two dichoptic stimuli were presented simultaneously to measure the amount of time they both appeared the same in at least one feature domain. The two stimuli were grouped when they appeared to share ambiguous features such as color, orientation, and spatial frequency more often than chance. Experiments here tested whether unshared and unambiguous features impeded grouping of the ambiguous components. Overall, the results show that grouping can be driven by neural ambiguity that is common for fragments across the visual field, even when the fragments also have other unshared, unambiguous features.

In a classic study, Kovacs et al. [Proc. Natl. Acad. Sci. USA93, 15508 (1996)PNASA60027-842410.1073/pnas.93.26.15508] used an array of many disks presented dichoptically with half of the disks in one eye "red" and the other half "green;" disk chromaticities in the fellow eye were reversed, resulting in binocular color rivalry for every disk, thus creating color ambiguity. Surprisingly, the binocularly fused percept sometimes was all disks of the same color (red or green), which showed that perceptual resolution of the many ambiguous neural representations did not rely completely on monocular dominance or on independent resolution for each disk. The present study replicates and expands on the original with the aim to isolate binocularly driven neural mechanisms of perceptual resolution without contamination from monocular dominance. Observers viewed a color-rivalrous array with 16 disks presented either steadily to each eye, as in Kovacs et al., or with chromatic interocular-switch rivalry (CISR), which swaps the two images between the eyes every 133 ms. The total proportion of viewing time when the 16 disks were perceived to be all red or all green was measured. For three observers, the disks all appeared the same color more often with CISR than with steady rivalrous presentation, suggesting that monocular dominance interferes with grouped perceptual resolution of ambiguous stimuli in the Kovacs paradigm. This conclusion was supported by an additional condition using CISR, but with every disk the same color in one eye at each instant (e.g., all "red" disks in one eye and all "green" in the other). This condition was never significantly different from the original CISR condition, as expected if CISR reveals only binocularly mediated perceptual resolution of the disks' color, irrespective of monocular neural representations. In conclusion, chromatically tuned binocularly driven neurons account for perceptual resolution of CISR.

Very-long-term (VLT) chromatic adaptation results from exposure to an altered chromatic environment for days or weeks. Color shifts from VLT adaptation are observed hours or days after leaving the altered environment. Short-term chromatic adaptation, on the other hand, results from exposure for a few minutes or less, with color shifts measured within seconds or a few minutes after the adapting light is extinguished; recovery to the pre-adapted state is complete in less than an hour. Here, both types of adaptation were combined. All adaptation was to reddish-appearing long-wavelength light. Shifts in unique yellow were measured following adaptation. Previous studies demonstrate shifts in unique yellow due to VLT chromatic adaptation, but shifts from short-term chromatic adaptation to comparable adapting light can be far greater than from VLT adaptation. The question considered here is whether the color shifts from VLT adaptation are cumulative with large shifts from short-term adaptation or, alternatively, does simultaneous short-term adaptation eliminate color shifts caused by VLT adaptation. The results show the color shifts from VLT and short-term adaptation together are cumulative, which indicates that both short-term and very-long-term chromatic adaptation affect color perception during natural viewing.

This research had two goals. First, a new method of very-long-term chromatic adaptation was compared to an older method of long-wavelength ambient illumination. In the new method, the observer viewed for 1 h per day for 12 or 14 days a CRT screen composed of oriented lines that appeared red. One observer also replicated a previous procedure (Neitz et al., 2002) in which she was exposed to long-wavelength room illumination for 4 h per day for 14 days. For both methods, equilibrium yellow was measured each day about 20 h after the end of the adaptation period. Both methods of very-long-term chromatic adaptation gave similar results. Second, shifts in equilibrium yellow were measured over a 30:1 range of light levels to determine if changes in color percepts were explained solely by a gain change in cone sensitivities (von Kries coefficient law). The magnitude of shift of equilibrium yellow depended on the level of the test light, which was not consistent with a gain theory of very-long-term chromatic adaptation.

Under dichoptic viewing conditions, rivalrous gratings that differ in both color and form can give the percept of the color from one eye in part of the form in the other eye. This study examined the afterimage following such misbinding of color to form. The first experiment established that afterimages of the misbound percept were seen. Two possible mechanisms for the misbound afterimage are (1) persisting retinal representations that are rivalrous and subsequently resolved to give misbinding, as during rivalrous viewing, and (2) a persisting response from a central neural representation of the misbound percept with the form from one eye and color from the other eye. The results support afterimage formation from a central representation of the misbound percept, not from resolution of rivalrous monocular representations.

The appearance of an object or surface depends strongly on the light from other objects and surfaces in view. This review focuses on color in complex scenes, which have regions of different colors in view simultaneously and/or successively, as in natural viewing. Two fundamental properties distinguish the chromatic representation evoked by a complex scene from the representation for an isolated patch of light. First, in complex scenes, the color of an object is not fully determined by the light from that object reaching the eye. Second, the chromatic representation of a complex scene contributes not only to hue, saturation, and brightness, but also to other percepts such as shape, texture, and object segmentation. These two properties are cornerstones of this review, which examines color perception with context that varies over space or time, including color constancy, and chromatic contributions to such percepts as orientation, contour, depth, and motion.

The color appearance of a physically steady central region can appear to vary over time if a surrounding chromatic light varies in time. The induced temporal variation, however, is strongly attenuated at surround temporal frequencies above ∼3 Hz. At these higher temporal frequencies, the central region appears steady (De Valois et al., 1986). The posited explanation is a cortical low-pass temporal filter. Here, we investigate whether higher temporal-frequency surrounds induce color shifts in the steady appearance of the central test. Surrounds modulated in time along the l or s chromatic direction of MacLeod-Boynton color space were symmetric around equal-energy white (EEW). The temporal frequency of the surround was varied. If observers perceived the central test to be temporally modulating between two points in time, they set two separate matches to the extreme points of this modulation. If the central test appeared steady in time, then color matches were made to this steady appearance. Corroborating previous reports, measurements showed that surround temporal frequencies below ∼3 Hz induced temporal modulation. At higher temporal frequencies, however, the surround induced steady color shifts, compared to a steady surround at its time average (EEW). The measurements imply that a nonlinear neural process affects chromatic induction from time-varying context.

Temporally varying chromatic-inducing light was used to infer receptive-field organization. Time-varying shifts in color appearance within a test field were induced by a surrounding chromatic pattern; the shifts were then nulled by adding a time-varying stimulus to the test area so the observer perceived a steady test. This method measured chromatic induction without requiring an observer to judge the color appearance of the test. The induced color shifts were consistent with a +s/−s spatially antagonistic neural receptive field, which also accounts for color shifts induced by static chromatic patterns (Monnier & Shevell, 2003, Monnier & Shevell, 2004). The response of this type of receptive-field, which is found only in the visual cortex, increases with S-cone stimulation at its center and decreases with S-cone stimulation within its surround. The measurements also showed a negligible influence of temporal inducing frequency in the range 0.5–4 Hz.

Chromatic assimilation is the shift in color appearance toward nearby light. Assimilation was measured using nearby light with time-varying chromaticity. This light induced time-varying assimilation within the test area. Assimilation was quantified by the amplitude of temporally varying test-area light – in counter-phase to the induced assimilation – required to null the assimilation. Unlike previous studies of assimilation, observers here judged only the steadiness of the test area, not its color. The inducing light was varied in luminance, temporal frequency and chromaticity. The measured assimilation could not be explained by only optical factors affecting receptoral quantal absorption. This implies a neural process contributes to assimilation. The nulling measurements showed also that assimilation was not induced independently within the L/M- and S-cone pathways. � 2005 Elsevier Ltd. All rights reserved.

Chromatic assimilation is the shift in color appearance of a test field toward the appearance of nearby light. Possible explanations of chromatic assimilation include wavelength independent spread light, wavelength-dependent chromatic aberration and neural summation. This study evaluated these explanations by measuring chromatic assimilation from a concentric-ring pattern into an equal-energy-white background, as a function of the inducing rings width, separation, chromaticity and luminance. The measurements showed, in the s direction, that assimilation was observed with different inducing-ring widths and separations when the inducing luminance was lower or higher than the test luminance. In general, the thinner the inducing rings and the smaller their separation, the stronger the assimilation in s. In the l direction, either assimilation or contrast was observed, depending on the ring width, separation and luminance. Overall, the measured assimilation could not be accounted for by the joint contributions from wavelength-independent spread light and wavelength-dependent chromatic aberration. Spatial averaging of neural signals explained the assimilation in s reasonably well, but there were clear deviations from neural spatial averaging for the l direction. � 2004 Elsevier Ltd. All rights reserved.

Patterned backgrounds that selectively stimulate the S-cones cause conspicuous color shifts. These shifts are accounted for by an S-cone antagonistic (+S/-S) center-surround receptive field [Monnier, P., & Shevell, S. K. (2004). Chromatic induction from S-cone patterns. Vision Research, 44, 849-856]. The present study tested two additional implications of the S-cone receptive field for color shifts: (1) proportionality of the shifts with respect to S-cone contrast within the inducing pattern and (2) bandpass selectivity of the shifts with respect to the spatial frequency of the inducing pattern. Measurements showed that the magnitude of the color shift was linear with S-cone contrast and that the largest color shift was observed with inducing patterns at an intermediate spatial frequency. These results further support an S-cone spatially antagonistic receptive field as the neural substrate mediating the large color shifts from S-cone patterns.

Chromatic assimilation is a shift toward the color of nearby light. Several studies conclude that a neural process contributes to assimilation but the neural locus remains in question. Some studies posit a peripheral process, such as retinal receptive-field organization, while others claim the neural mechanism follows depth perception, figure/ground segregation, or perceptual grouping. The experiments here tested whether assimilation depends on a neural process that follows stereoscopic depth perception. By introducing binocular disparity, the test field judged in color was made to appear in a different depth plane than the light that induced assimilation. The chromaticity and spatial frequency of the inducing light, and the chromaticity of the test light, were varied. Chromatic assimilation was found with all inducing-light sizes and chromaticities, but the magnitude of assimilation did not depend on the perceived relative depth planes of the test and inducing fields. We found no evidence to support the view that chromatic assimilation depends on a neural process that follows binocular combination of the two eyes' signals.

Intense scrutiny has been focused on whether chromatic stimuli contribute to motion perception. The present study considers a related but different question: how does motion affect chromatic detection? Detection thresholds were measured for a disk that underwent a brief (13.3 ms) chromatic change in the L/(L+M) chromatic direction. The disk's presentation sequence and speed (0–16 deg/s) were manipulated. In the coherent presentation sequence, the disk moved smoothly along a circular path centered on the fixation point. In the random presentation sequence, the disk appeared randomly at positions along the circular path. In both types of sequences, the disk underwent a brief chromatic change midway through the temporal presentation sequence. Threshold was elevated in the coherent condition compared to the random condition, and threshold decreased with an increase in speed. The threshold elevation observed in the coherent presentation sequence can be accounted for by temporal integration. The decrease in threshold with an increase in speed can be accounted for by spatial integration. The results, therefore, can be explained by spatiotemporal integration, without invoking a neural mechanism specialized for motion.

Binocular disparity is an aspect of natural viewing. This research investigates whether disparity affects surface color perception. Achromatic settings were obtained and compared for two stereograms of a scene with specular reflections, one stereogram with binocular disparity and one without it (cyclopean view). Binocular disparity was found to improve color constancy. Next, the geometry of specular highlights, which is distorted without binocular disparity, was specifically examined. Measurements compared color constancy with specular reflections that were either normal (with stereo disparity) or distorted (cyclopean view of the specularities). No significant change in constancy was found due to the geometrical distortion of specular highlights that occurs without stereo disparity, suggesting that constancy depends on other features of the percept affected by disparity. The results are discussed in terms of illuminant estimation in surface color perception.

Chromatic induction is the change in appearance of one light caused by a second, nearby light. We measured chromatic induction in a central test viewed within an inducing field that was varied in only short-wavelength-sensitive (S) cone stimulation. The observer matched the appearance of the central test by adjusting the chromaticity of a haploscopically presented comparison field, seen by the other eye on a dark background. When the central test weakly stimulated S cones, the S-cone level in the surround caused little change in the color appearance of the test. When the central test substantially stimulated S cones, on the other hand, the appearance of the center showed S-cone contrast: raising the level of S in the surround reduced the level of S set to match the central test. Further, a surround that weakly stimulated S cones raised the matching S-cone level above that required without a surround (dark-adapted condition). These results cannot be explained by S-cone sensitivity loss or by a two-process model of adaptation. A cortical mechanism is proposed to mediate S-cone antagonism.

Interocular-switch rivalry (also known as stimulus rivalry) is a kind of binocular rivalry in which two rivalrous images are swapped between the eyes several times a second. The result is stable periods of one image and then the other, with stable intervals that span many eye swaps (Logothetis, Leopold, & Sheinberg, 1996). Previous work used this close kin of binocular rivalry with rivalrous forms. Experiments here test whether chromatic interocular-switch rivalry, in which the swapped stimuli differ in only chromaticity, results in slow alternation between two colors. Swapping equiluminant rivalrous chromaticities at 3.75 Hz resulted in slow perceptual color alternation, with one or the other color often continuously visible for two seconds or longer (during which there were 15+ eye swaps). A well-known theory for sustained percepts from interocular-switch rivalry with form is inhibitory competition between binocular neurons driven by monocular neurons with matched orientation tuning in each eye; such binocular neurons would produce a stable response when a given orientation is swapped between the eyes. A similar model can account for the percepts here from chromatic interocular-switch rivalry and is underpinned by the neurophysiological finding that color-preferring binocular neurons are driven by monocular neurons from each eye with well-matched chromatic selectivity (Peirce, Solomon, Forte, & Lennie, 2008). In contrast to chromatic interocular-switch rivalry, luminance interocular-switch rivalry with swapped stimuli that differ in only luminance did not result in slowly alternating percepts of different brightnesses.

When seen in isolation, a light that varies in chromaticity over time is perceived to oscillate in color. Perception of that same time-varying light may be altered by a surrounding light that is also temporally varying in chromaticity. The neural mechanisms that mediate these contextual interactions are the focus of this article. Observers viewed a central test stimulus that varied in chromaticity over time within a larger surround that also varied in chromaticity at the same temporal frequency. Center and surround were presented either to the same eye (monocular condition) or to opposite eyes (dichoptic condition) at the same frequency (3.125, 6.25, or 9.375 Hz). Relative phase between center and surround modulation was varied. In both the monocular and dichoptic conditions, the perceived modulation depth of the central light depended on the relative phase of the surround. A simple model implementing a linear combination of center and surround modulation fit the measurements well. At the lowest temporal frequency (3.125 Hz), the surround's influence was virtually identical for monocular and dichoptic conditions, suggesting that at this frequency, the surround's influence is mediated primarily by a binocular neural mechanism. At higher frequencies, the surround's influence was greater for the monocular condition than for the dichoptic condition, and this difference increased with temporal frequency. Our findings show that two separate neural mechanisms mediate chromatic contextual interactions: one binocular and dominant at lower temporal frequencies and the other monocular and dominant at higher frequencies (6–10 Hz).

Wu et al. [Nature 429, 262 (2004)] describe a visual illusion in which color and motion are incorrectly bound: green dots moving downward and red dots moving upward are seen as green dots going up and red dots going down. The present study determined whether S cones contribute to color-motion feature-binding errors, in order to assess the neural representation of color at the level of binding. The specific experimental question is whether binding errors depend on S-cone responses from the objects perceived to have an illusory direction of motion. Alternatively, only L and M cones may determine the neural representation of color that regulates color-motion feature binding. In two experiments, the chromatic difference was manipulated between central objects, which induce color-motion binding errors, and peripheral objects, where color-motion binding errors occur. The chromaticity difference was varied along only the L/M-cone axis or only the S-cone axis. As in Wu et al. [Nature 429, 262 (2004)], color-motion binding was frequently observed in the periphery when there were no central versus peripheral chromatic differences. Further, the results showed that the frequency of color-motion binding errors in the periphery depended on the difference in S-cone excitation between center and periphery, thereby demonstrating that the neural representation of color at the level of feature binding depends on signals from not only L and M cones but also S cones.

Observers performed red–green and yellow–blue hue cancellation tasks for a 0.8-deg circular test field on a dark surround, by manipulating the excitation level of one cone class while the other two classes were held constant. The results of the red–green judgments conformed to classical opponent color theory in that both L- and S-cone excitation levels were antagonistic to M-cone signals. The yellow–blue judgments revealed a nonmonotonic nonlinearity in which the S-cone signal could act either antagonistically or synergistically with M- and L-cone signals. These results demonstrate that fixed hue sensations should not be associated with a given class of cone, even at the level of opponent neural coding.

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.

The visual system is able to recognize a scene based on a sketch made of very simple features. This ability is likely crucial for survival, when fast image recognition is necessary, and it is believed that a primal sketch is extracted very early in the visual processing. Such highly simplified representations can be sufficient for accurate object discrimination, but an open question is the role played by color in this process. Rich color information is available in natural scenes, yet artist's sketches are usually monochromatic; and, black-and-white movies provide compelling representations of real world scenes. Also, the contrast sensitivity of color is low at fine spatial scales. We approach the question from the perspective of optimal information processing by a system endowed with limited computational resources. We show that when such limitations are taken into account, the intrinsic statistical properties of natural scenes imply that the most effective strategy is to ignore fine-scale color features and devote most of the bandwidth to gray-scale information. We find confirmation of these information-based predictions from psychophysics measurements of fast-viewing discrimination of natural scenes. We conclude that the lack of colored features in our visual representation, and our overall low sensitivity to high-frequency color components, are a consequence of an adaptation process, optimizing the size and power consumption of our brain for the visual world we live in.

We compared changes in the appearance of a test region caused by introducing an inhomogeneous chromatic background to changes caused by a space-averaged equivalent uniform background. Subjects adjusted a test field presented on a CRT so that it appeared neither reddish nor greenish. Sparse “white” or “green” dots, randomly scattered throughout a “red” background field, caused a large decrease (up to 15 nm) in the dominant wavelength of the red/green equilibrium setting, compared to measurements with a uniform “red” background. A uniform background with the same space-averaged chromaticity and luminance as the complex background had an effect similar to the uniform “red” background. These results contradict theories of color constancy that rely on the “gray world” assumption, and indicate the significance for color perception of individual chromaticities within discrete, noncontiguous regions.

The appearance of a chromatic stimulus depends on more than the wavelengths composing it. The scientific literature has countless examples showing that spatial and temporal features of light influence the colors we see. Studying chromatic stimuli that vary over space, time or direction of motion has a further benefit beyond predicting color appearance: the unveiling of otherwise concealed neural processes of color vision. Spatial or temporal stimulus variation uncovers multiple mechanisms of brightness and color perception at distinct levels of the visual pathway. Spatial variation in chromaticity and luminance can change perceived three-dimensional shape, an example of chromatic signals that affect a percept other than color. Chromatic objects in motion expose the surprisingly weak link between the chromaticity of objects and their physical direction of motion, and the role of color in inducing an illusory motion direction. Space, time and motion – color’s colleagues – reveal the richness of chromatic neural processing.

According to opponent-colors theory, a reddish surround induces greenness in a central test field. Color-appearance measurements verify this with a long-wavelength reddish surround (660 nm) but not with a short-wavelength reddish surround (440 nm). Surprisingly, a short-wavelength reddish surround shifts the appearance of a test toward redness. Four possible explanations are: (1) stray light from the short-wavelength reddish surround falls in the test area; (2) receptoral sensitivity changes over whelm induced greenness from the surround; (3) a neural process of assimilation, rather than contrast, to the surrounding light; (4) short-wavelength-sensitive (S) cones do not contribute to induced redness/greenness. Chromatic cancelation experiments confirm the fourth explanation. There was no change in induced redness/greenness when quantal absorption by only S cones in the surround was varied by 30-fold (using tritanopic metamers), even though varying stimulation of S cones strongly affected the color appearance of the surround. The redness induced by a short-wavelength surround is accounted for by opponent chromatic induction mediated by only middle- and long-wavelength sensitive cones.

How does a physical stimulus determine a conscious percept? Binocular rivalry provides useful insights into this question because constant physical stimulation during rivalry causes different visual experiences. For example, presentation of vertical stripes to one eye and horizontal stripes to the other eye results in a percept that alternates between horizontal and vertical stripes. Presentation of a different color to each eye (color rivalry) produces alternating percepts of the two colors or, in some cases, a color mixture. The experiments reported here reveal a novel and instructive resolution of rivalry for stimuli that differ in both form and color: perceptual alternation between the rivalrous forms (e.g., horizontal or vertical stripes), with both eyes' colors seen simultaneously in separate parts of the currently perceived form. Thus, the colors presented to the two eyes (a) maintain their distinct neural representations despite resolution of form rivalry and (b) can bind separately to distinct parts of the perceived form.

In natural viewing, an object's background often changes over time. Temporally varying backgrounds were investigated here with a steady test field within a time-varying surrounding chromaticity. With slow surround variation (below approximately 3 Hz), the color appearance of a steady test is also perceived to fluctuate. At somewhat higher temporal frequencies, however, temporal variation of the surround is visible but the test appears steady (R. L. De Valois, M. A. Webster, K. K. De Valois, & B. Lingelbach, 1986); also above approximately 3 Hz, temporal chromatic variation along the l- or s-axis of the MacLeod-Boynton space (symmetric about equal-energy-spectrum "white") shifts the steady appearance of the test field toward redness or yellowness, respectively (A. D. D'Antona & S. K. Shevell, 2006). In the study here, color shifts were measured with temporal surround modulation at 6 Hz or greater along axes intermediate to the l and s directions. Varying the relative phase of simultaneous surround variation in l and s should not change responses within independent l and s pathways but should differentially excite neural representations that combine l and s signals (so-called higher order chromatic mechanisms). Varying the phase of l and s showed that the induced color shifts were accounted for by neural responses both from nearly independent l and s pathways and from higher order chromatic mechanisms.

An open question in color rivalry is whether alternation between two colors is caused by a difference in receptoral stimulation or a difference in the neural representation of color appearance. This question was examined with binocular rivalry between physically identical lights that differed in appearance due to chromatic induction. Perceptual alternation was measured between gratings of the same chromaticity; each one was presented within a different patterned surround that caused the gratings, one to each eye, to appear unequal in hue because of chromatic induction. The gratings were presented dichoptically with binocular disparity so the rivalrous gratings appeared in front of the surround. Perceptual alternation in hue was found for the two physically identical chromaticities. Stereoscopic depth also was perceived, corroborating binocular neural combination despite color rivalry (Treisman, 1962). The results show that color rivalry is resolved after color-appearance shifts caused by chromatic context, and that color rivalry does not require competing unequal cone excitations from the rivalrous stimuli.

The predominance of rivalrous targets is affected by surrounding context when stimuli rival in orientation, motion or color. This study investigated the influence of chromatic context on binocular color rivalry. The predominance of rivalrous chromatic targets was measured in various surrounding contexts. The first experiment showed that a chromatic surround’s influence was stronger when the surround was uniform or a grating with luminance contrast (chromatic/black grating) compared to an equiluminant grating (chromatic/white). The second experiment revealed virtually no effect of the orientation of the surrounding chromatic context, using chromatically rivalrous vertical gratings. These results are consistent with a chromatic representation of the context by a non-oriented, chromatically selective and spatially antagonistic receptive field. Neither a double-opponent receptive field nor a receptive field without spatial antagonism accounts for the influence of context on binocular color rivalry.

Is neural binding of color and form required for perception of a unified colored object? Individual cells selectively tuned to both color and orientation are proposed to moot the binding problem. This study reveals perceptual misbinding of color, thereby revealing separate neural representations of color and form followed by a subsequent binding process. Low luminance-contrast, rivalrous chromatic gratings were presented dichoptically. Each grating had alternating chromatic and gray stripes (e.g., red/gray in the left eye, green/gray in the right eye). Observers viewed the two rivalrous, 2 cpd gratings for 1 min. The duration of exclusive visibility was measured for four percepts: left-eye stimulus, right-eye stimulus, fusion of the two colors, or a two-color grating (e.g. a red/green grating). The percept of a two-color grating (misbinding) was observed with Michelson luminance contrast in the grating up to 20%. In general, for a given level of luminance contrast either misbinding (low luminance contrast) or color mixture (high luminance contrast) was observed, but not both of them. The perceived two-color gratings show that two rivalrous chromaticities are both represented neurally when color and form are combined to give a unified percept. “Resolution” of competing chromatic signals from the two eyes is not restricted to color dominance and color mixture. The transition from misbinding to color mixture by increasing luminance contrast shows that luminance edges have an important role in correct localization of color.

Modern theories of brightness induction include an influence from regions that do not share a border with the target. This study tested whether the spatial range of neural integration is the same with incremental versus decremental contrast edges in relatively remote parts of the background. Using an asymmetric matching task, observers set the brightness of a comparison ring, within its own uniform surround, to match the brightness of a test ring within a contiguous surround and a noncontiguous background. The measurements showed that the area of integration depended on the incremental versus decremental contrast polarity at the edge between the surround and background. This implies that brightness induction from an inhomogeneous background must consider the polarity of contrast edges within the whole scene.

Chromatic induction from patterned backgrounds depends on the spatial as well as the chromatic aspects of the background light. Color appearance with patterned and uniform backgrounds was compared using chromaticities distinguished by only the S cones; all backgrounds were equivalent to equal-energy white in terms of L-cone and M-cone stimulation. The measurements showed larger shifts in color appearance with a patterned chromatic background than with a uniform background at any chromaticity within the pattern. The measurements also showed that inducing light within different spatial regions could cause opposite shifts in color appearance: inducing light near a test field shifted appearance toward the inducing chromaticity (assimilation), while the same light some distance from the test shifted appearance away from the inducing chromaticity (simultaneous contrast). The shifts in color appearance were accounted for by a neural receptive field with S-cone spatial antagonism.

Chromatic induction from a surround is attenuated by chromatic contrast within a remote region outside of the surround (Shevell & Wei, 1998, Vision Research, 38, 1561-1566). The present study reports hue-cancellation measurements that show the attenuation depends on the magnitude, spatial frequency and chromaticity of remote chromatic contrast. Spatial-frequency tuning is shown by maximal attenuation of induction with remote contrast elements of the same size as the test. Experiments with various chromaticities of remote contrast show that S-cone stimulation within the remote region has a much weaker effect than L-/M-cone chromatic contrast, and does not depend on whether the S-cone stimulation is uniform or uneven across the region. Overall, the results show that remote L/M contrast affects classical chromatic induction, with its effect depending on the spatial frequency and magnitude of contrast. The influence of remote S-cone stimulation, on the other hand, is relatively weak and depends on only the S-cone spatial average, at least when S-cone stimulation by the test and its immediate surround is minimal (as in all experiments here).

Color matches between two small patches were made in a display containing ten larger regions of different chromaticities. The spatial organization of the ten regions was varied while keeping constant the immediate surround of each patch as well as the space-average chromaticity of the entire stimulus. Different spatial arrangements were designed to alter the perceptual organization inferred by the observer without changing the ensemble of chromaticities actually in view. For example, one arrangement of the ten regions was consistent with five surfaces under two distinct illuminations, with one edge within the display (an “apparent illumination edge”) dividing the stimulus into two areas, one under illuminant A and the other under illuminant C. Another spatial arrangement had the ten regions configured to induce an observer to infer ten surfaces under a single illumination. When the ten regions were arranged with an apparent illumination edge, the patch within the area of illuminant C was perceived as bluer than when the same patch and immediate surround were presented without an apparent illumination edge. The results are accounted for by positing that observers group together regions sharing the same inferred illumination, with a consequent effect on color perception: A fixed patch-within-surround shifts in hue and saturation toward the perceived illumination. We suggest that the change in color perception in a complex scene that results from a difference in real illumination may be caused by the inferred illumination at the perceptual level, not directly by the physical change in the light absorbed by photoreceptors.

Adapting to a chromatic light can alter the color appearance of other lights in view. The chromatic adapting effect is measured here with the test and adapting field perceived in the same depth plane, or perceived in different depth planes (using stereo disparity). The measurements show only a weak, though consistent, shift in the appearance of the test when adapting field and test are perceived in different depth planes, compared to when they are in the same plane. Adding complexity to the adapting stimulus, in the form of a second chromatic light surrounding the background, alters the appearance of the test but shows no dependence on the depth relations. Overall, there is only a small difference in chromatic adaptation caused by introducing a three-dimensional representation of these stimuli.

Individual differences in abnormal color vision are well known. A fundamental unresolved problem is the great variation in color vision even among those classified as having the same color-vision defect. Several physiological hypotheses have been proposed to account for this variation but little consideration has been given to how (and how much) color matching and discrimination are affected by the posited physiological mechanisms. Advances in molecular genetics have renewed interest in this problem, which is at the foundation of the relation between genotype and phenotype. We report here theoretical Rayleigh ranges (chromatic discrimination) and quantal matches for deuteranomalous trichromats with photopigments in the red/green range that vary in their separation and optical density. The results show there is relatively little loss of discrimination with pigments of normal optical density separated by as little as 2–3 nm. With pigments separated by 4 nm or less, however, optical density can strongly influence discrimination when varied independently in the two types of cone. Moderately lower (or higher) optical density in only one cone-type affects discrimination by altering the shape of the cone's relative spectral sensitivity function. The lack of correlation between Rayleigh-match midpoint and range, which is reported in the literature, may be accounted for by independent variation in pigment separation and optical density.

The perceived color of a light varies with the background on which it is seen. In the present study, patterned backgrounds composed of two different chromaticities caused larger shifts in perceived color than did a uniform background at either chromaticity within the pattern. Cortical receptive-field organization, but not optical factors or known retinal neurons, can account for the color shifts from patterned backgrounds.

Few studies have investigated the possible role of higher-level cognitive mechanisms in color constancy. Following-up on previous work with successive color constancy (Allen, Beilock, & Shevell, J. Exp. Psychol. Learn. 37, 1014, 2011), the current study examined the relation between simultaneous color constancy and working memory – the ability to maintain a desired representation while suppressing irrelevant information. Higher working-memory was associated with poorer simultaneous color constancy for a chromatically complex stimulus, but was not associated with simultaneous color constancy for a chromatically simple stimulus. Ways in which the executive attention mechanism of working memory may play a role in color constancy when observers are not required to maintain a representation of a color in memory (as was the case here) are discussed. This finding supports a role for higher-level cognitive mechanisms in color constancy, and is the first to demonstrate a relation between simultaneous color constancy and a complex cognitive ability.

The dual Rayleigh-type color match is the ratio of 621 nm light to 550 nm light that in admixture matches 586 nm light, divided by the ratio of 667 nm light to 550 nm light that in admixture matches 586 nm light. Compared to the classical Rayleigh match, the dual-match procedure minimizes variation in color matching arising from differences in lens pigmentation and photopigment optical density, and thus amplifies individual differences due to shifts in L pigment lambda max. We hypothesized that the dual matches would provide a clearer distinction between subjects with serine and subjects with alanine than would the classical Rayleigh match because individuals with serine express L pigments with a lambda max shifted toward longer wavelengths than do those with alanine. Classical Rayleigh color matches were compared with dual Rayleigh-type color matches in 14 color-normal observers whose DNA had been analyzed previously for the presence of the amino acid serine or alanine at position 180 in the L opsin. The resulting distribution of dual-match measurements for the seven subjects with serine does not overlap the distribution of measurements for the seven subjects with alanine. The classical Rayleigh-match measurements for these two groups of subjects, on the other hand, overlap substantially. More than half of the subjects' classical Rayleigh matches are within the overlapping range. The dual Rayleigh-type matches, therefore, provide an improved psychophysical technique for assessing whether an individual observer has serine or alanine at position 180.

Individual differences in color matches of normal trichromats are well documented. Recently, variants of the classical Rayleigh match have been measured to explore the cause(s) of these individual differences. Interobserver differences in the wavelength of peak sensitivity of photopigment (lambda max) are of primary interest because they are attributed to an X-chromosome-linked polymorphism. Color-matching equations, however, show the Rayleigh match cannot distinguish between interobserver differences in lambda max and interobserver differences in the optical density of photopigment. Further analysis of color-matching equations reveals that the ratio of two particular Rayleigh-type matches amplifies the effect of individual differences in the lambda max of L cones relative to the effects of optical density and pre-receptoral spectrally selective filtering. The ratio of these two color matches was measured for 17 color-normal males. The range of the results for the 17 observers is too large to be explained by only individual differences in photopigment optical density and pre-receptoral filtering. This implies there are interobserver differences in lambda max. The results are accounted for quantitatively by a small difference (3-5 nm) in the lambda max of the L-cone photopigment. The ratio of two Rayleigh-type matches is a rapid and convenient measurement for assessing the L-cone lambda max in the eye of an individual observer and therefore may be useful for classifying normal trichomats into phenotypic sub-types.

Light at the boundary of a uniform test field (contrast) has a qualitatively different effect on color perception than light in more remote noncontiguous regions (context). Basic properties of color perception with contextual short-wavelength light are assessed here with a 1 degree test field surrounded by either contiguous or noncontiguous 440 or 491 nm light (32 td). Contrasting stimuli are 3 or 5 degrees adapting fields, a thin 1 degree i.d.-2 degrees o.d. (0.5 degree wide) contiguous band, or a large 1 degree i.d.-5 degrees o.d. contiguous surround. Contextual stimuli are a remote 3 degrees i.d.-5 degrees o.d. ring or 0.5 degree wide noncontiguous bands at various distances from the edge of the 1 degree test field (2 degrees i.d.-3 degrees o.d., 3 degrees i.d.-4 degrees o.d., or 4 degrees i.d.-5 degrees o.d. bands). Contiguous surrounds have little influence on color appearance, but remote noncontiguous short-wavelength light strong affects the color of the test field, shifting it toward redness. The shift toward redness increases as a thin 440 nm band is moved farther from the test field (up to 5 degrees), unlike the effect of distance on remote middle- and long-wavelength bands. Measurements comparing the effects of 440 nm and luminance-equated 491 nm light indicate a contribution from S cones.

Is a retinal representation of an edge necessary to constrain the reach of color filling-in? If so, then color filling-in should not be constrained by illusory contours, because they do not exist at a retinal level. Alternatively, if color filling-in is constrained by contours at a perceptual level of neural representation, regardless of whether there is a retinal representation, then color filling-in should be constrained by illusory contours. To address this question, a variety of real luminance edges and illusory contours were presented under conditions designed to cause color filling-in. The results showed that illusory contours bounded the reach of color filling-in. A neural representation of a contour may first exist at a retinal level or a cortical level; in either case, the contour exists at a perceptual level and bounds color filling-in.

We measured changes in the color appearance of one light caused by another light presented in a well-separated region. Observers viewed a 1 degrees test field superimposed on a 3 degrees, 540 or 660 nm adapting field (32 or 320 td). The change in appearance due to noncontiguous light was determined by surrounding the 3 degrees adapting field with a continguous 3 degrees i.d., 5 degrees o.d. ring of either 32 or 320 td. The ring was 540, 660 nm or achromatic (tungsten-halogen "white"). The test was an admixture of 549 and 660 nm light, and varied from 6 to 1000 td. The observer adjusted the ratio of 549 to 660 nm test light so the test appeared neither reddish nor greenish. A 540 or 660 nm ring had a chromatic inducing effect on the small test that mimicked a simple surround contiguous with the test. Results with an achromatic ring were more complex: an isolated achromatic ring (no adapting field present) had virtually no effect on the color appearance of the test, but the same achromatic ring surrounding a chromatic adapting field shifted the test toward the color appearance of the adapting light (e.g. introducing a "white" ring surrounding a "green" adapting field shifted the test toward greenness). A thin pencil-width band of "white" light superimposed on a larger 5 degrees adapting field had an effect similar to a "white" 3-5 degrees ring. These results demonstrate (1) strong effects of the remote noncontiguous lights and (2) that the change in color appearance they cause is not a simple function of only the light in the noncontinguous region. The change depends on other lights in view. The visual processes revealed in these experiments are considered in terms of inferred illumination and surface reflectances of objects in natural scenes.

The perceived color of a light depends on surrounding light. When the surround varies slowly over time, a central, physically steady light is perceived to vary also. This perceived temporal variation of the central region, however, is strongly attenuated when the surround varies faster than approximately 3 Hz (R. L. De Valois, M. A. Webster, K. K. De Valois, & B. Lingelbach, 1986). The classical explanation is low-pass temporal filtering at a cortical stage that attenuates the neural representation of temporal frequencies capable of causing induced temporal variation. This theory assumes neural responses are linear, so only temporal frequencies in the stimulus are represented in the neural response. The current experiments revealed that temporal frequencies above 3 Hz are capable of inducing temporal variation. Specifically, with two temporal frequencies superimposed in the surround, the induced temporal variation in the uniform region is at the difference frequency of these two frequencies, even though this frequency is not physically present in the stimulus. The results are accounted for by a nonlinear neural process, which causes temporal variation at the difference frequency, and a following linear temporal filter.

We measured for six male observers, the psychometric functions for the detection of two simultaneously presented points of light. The test stimuli were two 1 min point sources separated by 17 min arc, and pulsed for 0.5 msec. The stimuli varied in wavelength from 500 to 620 nm. The psychometric functions were fit with a model that assumes ideal detection and the following properties: (1) Poisson-distributed quantal absorptions; (2) binomial sampling of foveal long-wavelength-sensitive (LWS) and middle-wavelength-sensitive (MWS) cones; (3) independent responses of the LWS and MWS cones; and (4) the Smith-Pokorny fundamentals for cone spectral sensitivies. Based on χ2 fits to the psychometric functions for detecting neither, one or both of the two-point stimuli presented, estimates were derived for the minimum quantal catch by a single cone for detection (C), the number of effective cones illuminated by a point stimulus at threshold (N) and the proportion of central foveal cones of the LWS type (PL). Three observers were color-normal, two were protanopes and one was a deuteranope. A second deuteranope was included in the design but his data were too unreliable for an unambigous solution. The estimated quantal requirement C was consistently near 5 (4–6), and the effective number of illuminated cones always was 1 or 2. The plausible range of PL (98% confidence interval) for the color-normal observers was 0.48–0.68 (observer YY), 0.76–0.90 (observer MW) and 0.77–0.95 (observer DF). The best fitting PL solution for these observers were 0.61, 0.82 and 0.88, respectively. These were comparable to the values obtained from flicker photometric data. The best PL value for each of the protanopes was 0.00 and for the deuteranope the best PL value was 0.98.

Patterned backgrounds can induce large shifts in color appearance, even with patterns of only 10% S-cone contrast (S. K. Shevell & P. Monnier, 2005). The present study tested whether a background pattern could induce color shifts even at a below-threshold contrast. In the first experiment, S-cone contrast threshold for discriminating a pattern from a homogenous background was measured by a 2AFC procedure. Next, a test ring was inserted within the patterned background. With the test ring present, six of eight observers reliably distinguished trials with a patterned background from trials with a homogeneous field, even though the S-cone contrast in the pattern was too low to be discriminated from a homogeneous background. This suggested that a below-threshold S-cone pattern shifted the color appearance of the test ring; that is, the appearance of the test was used to discriminate whether the background was patterned or homogeneous. This was corroborated by asymmetric color matches, which revealed a color shift caused by subthreshold S-cone contrast within the patterned background.

This study investigates color perception in a scene with two different illuminants. The two illuminants, in opposite corners, simultaneously shine on a (simulated) scene with an opaque dividing wall, which controls how much of the scene is illuminated by each source. In the first experiment, the height of the dividing wall was varied. This changed the amount of each illuminant reaching objects on the opposite side of the wall. Results showed that the degree of color constancy decreased when a region on one side of the wall had cues to both illuminants, suggesting that cues from the second illuminant are detrimental to color constancy. In a later experiment, color constancy was found to improve when the specular highlight cues from the second illuminant were altered to be consistent with the first illuminant. This corroborates the influence of specular highlights in surface color perception, and suggests that the reduced color constancy in the first experiment is due to the inconsistent, though physically correct, cues from the two illuminants.

The method of constant stimuli was used to estimate the psychometric functions for detection of one or two flashes when two light pulses were presented. The test stimulus consisted of two simultaneous 0.5 msec, 1′ pulses separated by 17′. Observers reported seeing 0, 1 or 2 flashes. A computer-controlled direct-view apparatus allowed sampling of slightly different foveal locations on each trial. The data were analyzed assuming a binomial probability for sampling of L and M cones and Poisson distributed quantal fluctuation. Under these assumptions, the measurements imply that detection requires a minimum of 5–7 quanta absorbed per cone, and that the effective number of cones illuminated by the 1′, 0.5 msec pulse is two. The estimated LM cone ratio was 1.6 for one observer and 4.0 for the other; each observer's ratio was in general agreement with the value estimated independently by heterochromatic flicker photometry.