Fundamentally different representations of color and motion revealed by individual differences in perceptual scaling.

The coordinate frames for color and motion are often defined by three dimensions (e.g., responses from the three types of human cone photoreceptors for color and the three dimensions of space for motion). Does this common dimensionality lead to similar perceptual representations? Here we show that the organizational principles for the representation of hue and motion direction are instead profoundly different. We compared observers' judgments of hue and motion direction using functionally equivalent stimulus metrics, behavioral tasks, and computational analyses, and used the pattern of individual differences to decode the underlying representational structure for these features. Hue judgments were assessed using a standard "hue-scaling" task (i.e., judging the proportion of red/green and blue/yellow in each hue). Motion judgments were measured using a "motion-scaling" task (i.e., judging the proportion of left/right and up/down motion in moving dots). Analyses of the interobserver variability in hue scaling revealed multiple independent factors limited to different local regions of color space. This is inconsistent with the influences across a broad range of hues predicted by conventional color-opponent models. In contrast, variations in motion scaling were characterized by more global factors plausibly related to variation in the relative weightings of the cardinal spatial axes. These results suggest that although the coordinate frames for specifying color and motion share a common dimensional structure, the perceptual coding principles for hue and motion direction are distinct. These differences might reflect a distinction between the computational strategies required for the visual analysis of spatial vs. nonspatial attributes of the world.

Automatic, Early Color-Specific Neural Responses to Object Color Knowledge.

Some familiar objects are associated with specific colors, e.g., rubber ducks with yellow. Whether and at what stage neural responses occur to these color associations remain open questions. We recorded frequency-tagged electroencephalogram (EEG) responses to periodic presentations of yellow-associated objects, shown among sequences of non-periodic blue-, red-, and green-associated objects. Both color and grayscale versions of the objects elicited yellow-specific responses, indicating an automatic activation of color knowledge from object shape. Follow-up experiments replicated these effects with green-specific responses, and demonstrated modulated responses for incongruent color/object associations. Importantly, the onset of color-specific responses was as early to grayscale as actually colored stimuli (before 100 ms), the latter additionally eliciting a conventional later response (approximately 140-230 ms) to actual stimulus color. This suggests that the neural representation of familiar objects includes both diagnostic shape and color properties, such that shape can elicit associated color-specific responses before actual color-specific responses occur.

Gaining the system: limits to compensating color deficiencies through post-receptoral gain changes.

Color percepts of anomalous trichromats are often more similar to normal trichromats than predicted from their receptor spectral sensitivities, suggesting that post-receptoral mechanisms can compensate for chromatic losses. The basis for these adjustments and the extent to which they could discount the deficiency are poorly understood. We modeled the patterns of compensation that might result from increasing the gains in post-receptoral neurons to offset their weakened inputs. Individual neurons and the population responses jointly encode luminance and chromatic signals. As a result, they cannot independently adjust for a change in the chromatic inputs, predicting only partial recovery of the chromatic responses and increased responses to achromatic contrast. These analyses constrain the potential sites and mechanisms of compensation for a color loss and characterize the utility and limits of neural gain changes for calibrating color vision.

Using equiluminance settings to estimate the cardinal chromatic directions for individuals.

Color information is processed by the retina and lateral geniculate along principal dimensions known as the cardinal directions of color space. Normal differences in spectral sensitivity can impact the stimulus directions that isolate these axes for individual observers and can arise from variation in lens and macular pigment density, photopigment opsins, photoreceptor optical density, and relative cone numbers. Some of these factors that influence the chromatic cardinal axes also impact luminance sensitivity. We modeled and empirically tested how well tilts on the individual's equiluminant plane are correlated with rotations in the directions of their cardinal chromatic axes. Our results show that, especially for the SvsLM axis, the chromatic axes can be partially predicted by luminance settings, providing a potential procedure for efficiently characterizing the cardinal chromatic axes for observers.

Adapting to an enhanced color gamut - implications for color vision and color deficiencies.

One strategy for aiding color deficiencies is to use three narrow passbands to filter the light spectrum to increase the saturation of colors. This filtering is analogous to the narrow emission bands used in wide gamut lighting or displays. We examined how perception adapts to the greater color gamut area produced by such devices, testing color-normal observers and simulated environments. Narrowband spectra increased chromatic contrasts but also increased contrast adaptation, partially offsetting the perceived contrast enhancements. Such adaptation adjustments are important for understanding the perceptual consequences of exposure to naturally or artificially enhanced color gamut areas for both color-deficient and color-normal observers.

Temporal dynamics of face adaptation.

The appearance of a face can be strongly affected by adaptation to faces seen previously. A number of studies have examined the time course of these aftereffects, but the integration time over which adaptation pools signals to control the adaptation state remains uncertain. Here we examined the effects of temporal frequency on face gender aftereffects induced by a pair of faces alternating between the two genders to assess when the aftereffects were pooled over successive faces versus driven by the last face seen. In the first experiment, we found that temporal frequencies between 0.25 and 2.00 Hz all failed to produce an aftereffect, suggesting a fairly long integration time. In the second experiment, we therefore probed slower alternation rates of 0.03 to 0.25 Hz. A rate of 0.0625 Hz (i.e., 8 seconds per face) was required to generate significant aftereffects from the last presented face and was consistent with an average time constant of 15 to 20 seconds for an exponentially decaying integration window. This integration time is substantially longer than found previously for analogous effects for alternating colors, and thus points to a potentially slower mechanism of adaptation for faces compared with chromatic adaptation.

All-or-none face categorization in the human brain.

Visual categorization is integral for our interaction with the natural environment. In this process, similar selective responses are produced to a class of variable visual inputs. Whether categorization is supported by partial (graded) or absolute (all-or-none) neural responses in high-level human brain regions is largely unknown. We address this issue with a novel frequency-sweep paradigm probing the evolution of face categorization responses between the minimal and optimal stimulus presentation times. In a first experiment, natural images of variable non-face objects were progressively swept from 120 to 3 â€‹Hz (8.33-333 â€‹ms duration) in rapid serial visual presentation sequences. Widely variable face exemplars appeared every 1 â€‹s, enabling an implicit frequency-tagged face-categorization electroencephalographic (EEG) response at 1 â€‹Hz. Face-categorization activity emerged with stimulus durations as brief as 17 â€‹ms (17-83 â€‹ms across individual participants) but was significant with 33 â€‹ms durations at the group level. The face categorization response amplitude increased until 83 â€‹ms stimulus duration (12 â€‹Hz), implying graded categorization responses. In a second EEG experiment, faces appeared non-periodically throughout such sequences at fixed presentation rates, while participants explicitly categorized faces. A strong correlation between response amplitude and behavioral accuracy across frequency rates suggested that dilution from missed categorizations, rather than a decreased response to each face stimulus, accounted for the graded categorization responses as found in Experiment 1. This was supported by (1) the absence of neural responses to faces that participants failed to categorize explicitly in Experiment 2 and (2) equivalent amplitudes and spatio-temporal signatures of neural responses to behaviorally categorized faces across presentation rates. Overall, these observations provide original evidence that high-level visual categorization of faces, starting at about 100 â€‹ms following stimulus onset in the human brain, is variable across observers tested under tight temporal constraints, but occurs in an all-or-none fashion.

The Verriest Lecture: Adventures in blue and yellow.

Conventional models of color vision assume that blue and yellow (along with red and green) are the fundamental building blocks of color appearance, yet how these hues are represented in the brain and whether and why they might be special are questions that remain shrouded in mystery. Many studies have explored the visual encoding of color categories, from the statistics of the environment to neural processing to perceptual experience. Blue and yellow are tied to salient features of the natural color world, and these features have likely shaped several important aspects of color vision. However, it remains less certain that these dimensions are encoded as primary or "unique" in the visual representation of color. There are also striking differences between blue and yellow percepts that may reflect high-level inferences about the world, specifically about the colors of light and surfaces. Moreover, while the stimuli labeled as blue or yellow or other basic categories show a remarkable degree of constancy within the observer, they all vary independently of one another across observers. This pattern of variation again suggests that blue and yellow and red and green are not a primary or unitary dimension of color appearance, and instead suggests a representation in which different hues reflect qualitatively different categories rather than quantitative differences within an underlying low-dimensional "color space."

Color variance and achromatic settings.

The average color in a scene is a potentially important cue to the illuminant and thus for color constancy, but it remains unknown how well and in what ways observers can estimate the mean chromaticity. We examined this by measuring the variability in "achromatic" settings for stimuli composed of different distributions of colors with varying contrast ranges along the luminance, SvsLM, and LvsM cardinal axes. Observers adjusted the mean chromaticity of the palette to set the average to gray. Variability in the settings increased as chromatic contrast or (to a lesser extent) luminance contrast increased. Signals along the cardinal axes are relatively independent in many detection and discrimination tasks, but showed strong interference in the white estimates. This "cross-masking" and the effects of chromatic variance in general may occur because observers cannot explicitly perceive or represent the mean of a set of qualitatively different hues (e.g., that red and green hues average to gray), and thus may infer the mean only indirectly (e.g., from the relative saturation of different hues).

Predicting color matches from luminance matches.

Color vision and spectral sensitivity vary among individuals with normal color vision; thus, for many applications, it is important to measure and correct for an observer's sensitivity. Full correction would require measuring color and luminance matches and is rarely implemented. However, luminance matches (equiluminance settings) are routinely measured and simple to conduct. We modeled how well an observer's color matches could be approximated by measuring only luminance sensitivity, since both depend on a common set of factors. We show that lens and macular pigment density and $L/M$L/M cone ratios alter equiluminance settings in different ways and can therefore be estimated from the settings. In turn, the density variations can account for a large proportion of the normal variation in color matching. Thus, luminance matches may provide a simple method to at least partially predict an observer's color matches without requiring more complex tasks or equipment.

Comparison of two methods of hue scaling.

Hue-scaling functions are designed to characterize color appearance by assessing the relative strength of the red versus green and blue versus yellow opponent sensations comprising different hues. However, these judgments can be non-intuitive and may pose difficulties for measurement and analysis. We explored an alternative scaling method based on positioning a dial to represent the relative similarity or distance of each hue from the labeled positions for the opponent categories. The hue-scaling and hue-similarity rating methods were compared for 28 observers. Settings on both tasks were comparable though the similarity ratings showed less inter-observer variability and weaker categorical bias, suggesting that these categorical biases may reflect properties of the task rather than the percepts. Alternatively, properties that are concordant for the two paradigms provide evidence for characteristics that do reflect color appearance. Individual differences on both tasks suggest that color appearance depends on multiple, narrowly tuned color processes, which are inconsistent with conventional color-opponent theory.

Neural correlates of perceptual color inferences as revealed by #thedress.

Color constancy involves disambiguating the spectral characteristics of lights and surfaces, for example to distinguish red in white light from white in red light. Solving this problem appears especially challenging for bluish tints, which may be attributed more often to shading, and this bias may underlie the individual differences in whether people described the widely publicized image of #thedress as blue-black or white-gold. To probe these higher-level color inferences, we examined neural correlates of the blue-bias, using frequency-tagging and high-density electroencephalography to monitor responses to 3-Hz alternations between different color versions of #thedress. Specifically, we compared relative neural responses to the original "blue" dress image alternated with the complementary "yellow" image (formed by inverting the chromatic contrast of each pixel). This image pair produced a large modulation of the electroencephalography amplitude at the alternation frequency, consistent with a perceived contrast difference between the blue and yellow images. Furthermore, decoding topographical differences in the blue-yellow asymmetries over occipitoparietal channels predicted blue-black and white-gold observers with over 80% accuracy. The blue-yellow asymmetry was stronger than for a "red" versus "green" pair matched for the same component differences in L versus M or S versus LM chromatic contrast as the blue-yellow pair and thus cannot be accounted for by asymmetries within either precortical cardinal mechanism. Instead, the results may point to neural correlates of a higher-level perceptual representation of surface colors.

Directional Visual Motion Is Represented in the Auditory and Association Cortices of Early Deaf Individuals.

Individuals who are deaf since early life may show enhanced performance at some visual tasks, including discrimination of directional motion. The neural substrates of such behavioral enhancements remain difficult to identify in humans, although neural plasticity has been shown for early deaf people in the auditory and association cortices, including the primary auditory cortex (PAC) and STS region, respectively. Here, we investigated whether neural responses in auditory and association cortices of early deaf individuals are reorganized to be sensitive to directional visual motion. To capture direction-selective responses, we recorded fMRI responses frequency-tagged to the 0.1-Hz presentation of central directional (100% coherent random dot) motion persisting for 2 sec contrasted with nondirectional (0% coherent) motion for 8 sec. We found direction-selective responses in the STS region in both deaf and hearing participants, but the extent of activation in the right STS region was 5.5 times larger for deaf participants. Minimal but significant direction-selective responses were also found in the PAC of deaf participants, both at the group level and in five of six individuals. In response to stimuli presented separately in the right and left visual fields, the relative activation across the right and left hemispheres was similar in both the PAC and STS region of deaf participants. Notably, the enhanced right-hemisphere activation could support the right visual field advantage reported previously in behavioral studies. Taken together, these results show that the reorganized auditory cortices of early deaf individuals are sensitive to directional motion. Speculatively, these results suggest that auditory and association regions can be remapped to support enhanced visual performance.

Dynamics of contrast adaptation in central and peripheral vision.

Adaptation aftereffects are generally stronger for peripheral than for foveal viewing. We examined whether there are also differences in the dynamics of visual adaptation in central and peripheral vision. We tracked the time course of contrast adaptation to binocularly presented Gabor patterns in both the central visual field (within 5°) and in the periphery (beyond 10° eccentricity) using a yes/no detection task to monitor contrast thresholds. Consistent with previous studies, sensitivity losses were stronger in the periphery than in the center when adapting to equivalent high contrast (90% contrast) patterns. The time course of the threshold changes was fitted with separate exponential functions to estimate the time constants during the adapt and post-adapt phases. When adapting to equivalent high contrast, adaptation effects built up and decayed more slowly in the periphery compared with central adaptation. Surprisingly, the aftereffect in the periphery did not decay completely to the baseline within the monitored post-adapt period (400 s), and instead asymptoted to a higher level than for central adaptation. Even when contrast was reduced to one-third (30% contrast) of the central contrast, peripheral adaptation remained stronger and decayed more slowly. This slower dynamic was also confirmed at suprathreshold test contrasts by tracking tilt-aftereffects with a 2AFC orientation discrimination task. Our results indicate that the dynamics of contrast adaptation differ between central and peripheral vision, with the periphery adapting not only more strongly but also more slowly, and provide another example of potential qualitative processing differences between central and peripheral vision.

Color Vision 2018: Introduction by the feature editors.

This feature issue of the Journal of the Optical Society of America A (JOSA A) reflects the basic and applied research interests of members of the color vision community. Most of the articles stem from presentations at the 24th Biennial Symposium of the International Colour Vision Society (ICVS).

Adjusting to a sudden "aging" of the lens.

Color perception is known to remain largely stable across the lifespan despite the pronounced changes in sensitivity from factors such as the progressive brunescence of the lens. However, the mechanisms and timescales controlling these compensatory adjustments are still poorly understood. In a series of experiments, we tracked adaptation in observers after introducing a sudden change in lens density by having observers wear glasses with yellow filters that approximated the average spectral transmittance of a 70-year-old lens. Individuals were young adults and wore the glasses for 5 days for 8 h per day while engaged in their normal activities. Achromatic settings were measured on a CRT before and after each daily exposure with the lenses on and off, and were preceded by 5 min of dark adaptation to control for short-term chromatic adaptation. During each day, there was a large shift in the white settings consistent with a partial compensation for the added lens density. However, there was little to no evidence of an afterimage at the end of each daily session, and participants' perceptual nulls were roughly aligned with the nulls for short-term chromatic adaptation, suggesting a rapid renormalization when the lenses were removed. The long-term drift was also extinguished by brief exposure to a white adapting field. The results point to distinct timescales and potentially distinct mechanisms compensating for changes in the chromatic sensitivity of the observer.

A simple principled approach for modeling and understanding uniform color metrics.

An important goal in characterizing human color vision is to order color percepts in a way that captures their similarities and differences. This has resulted in the continuing evolution of "uniform color spaces," in which the distances within the space represent the perceptual differences between the stimuli. While these metrics are now very successful in predicting how color percepts are scaled, they do so in largely empirical, ad hoc ways, with limited reference to actual mechanisms of color vision. In this article our aim is to instead begin with general and plausible assumptions about color coding, and then develop a model of color appearance that explicitly incorporates them. We show that many of the features of empirically defined color order systems (those of Munsell, Pantone, NCS, and others) as well as many of the basic phenomena of color perception, emerge naturally from fairly simple principles of color information encoding in the visual system and how it can be optimized for the spectral characteristics of the environment.

Color vision: introduction by the feature editors.

This feature issue of the Journal of the Optical Society of America A (JOSA A) reflects the basic and applied research interests of members of the color vision community. Most of the articles stem from presentations at the 23rd Biennial Symposium of the International Colour Vision Society (ICVS).

Dynamics of color contrast adaptation.

Many forms of color adaptation have been found to reflect both short- and long-term adjustments. We explored the buildup and decay of adaptation to chromatic contrast (temporal modulations of color) for which the dynamics are unknown. A matching task was used to track the perceived contrast of chromatic pulses of varying physical contrast during and after adapting for 1 h to a high contrast modulation repeated over five successive days. The adaptation was characterized by rapid response changes that remained stable in both time course and form across sessions. There was no consistent evidence for long-term plasticity over the time scales we tested.

Filling in, filling out, or filtering out: processes stabilizing color appearance near the center of gaze.

Spectral sensitivity varies markedly across the center of gaze, in part because of the rapid decline in the density of macular pigment outside the fovea. Yet despite these retinal inhomogeneities, the color appearance of large uniform fields remains very uniform. We explored some of the processes contributing to these stable color percepts by measuring the effects of field size and eccentricity on saturated purples, whose spectra should show the largest biases with macular pigment screening. Small purple fields at 0° and 8° eccentricities differ in appearance but by much less than predicted by the macular screening or by compensation for the average effects of this screening at the two loci. This shows that the compensation is already nearly complete because of local adjustments that filter out the sensitivity variation and confirms that this filtering includes adjustments beyond average gain changes in the cones. In large fields, the appearance is dominated by the local peripheral color. This bias persists when the field edge is fixated or when abrupt edges are removed in Gaussian spots, suggesting that the spreading is not strongly dependent on luminance edges.