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.

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.

Color perception and compensation in color deficiencies assessed with hue scaling.

Anomalous trichromats have three classes of cone receptors but with smaller separation in the spectral sensitivities of their longer-wave (L or M) cones compared to normal trichromats. As a result, the differences in the responses of the longer-wave cones are smaller, resulting in a weaker input to opponent mechanisms that compare the LvsM responses. Despite this, previous studies have found that their color percepts are more similar to normal trichromats than the smaller LvsM differences predict, suggesting that post-receptoral processes might amplify their responses to compensate for the weaker opponent inputs. We evaluated the degree and form of compensation using a hue-scaling task, in which the appearance of different hues is described by the perceived proportions of red-green or blue-yellow primary colors. The scaling functions were modeled to estimate the relative salience of the red-green to blue-yellow components. The red-green amplitudes of the 10 anomalous observers were 1.5 times weaker than for a group of 26 normal controls. However, their relative sensitivity at threshold for detecting LvsM chromatic contrast was on average 6 times higher, consistent with a 4-fold gain in the suprathreshold hue-scaling responses. Within-observer variability in the settings was similar for the two groups, suggesting that the suprathreshold gain did not similarly amplify the noise, at least for the dimension of hue. While the compensation was pronounced it was nevertheless partial, and anomalous observers differed systematically from the controls in the shapes of the hue-scaling functions and the corresponding loci of their color categories. Factor analyses further revealed different patterns of individual differences between the groups. We discuss the implications of these results for understanding both the processes of compensation for a color deficiency and the limits of these processes.

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.

Individual differences and their implications for color perception.

Individual differences are a conspicuous feature of color vision and arise from many sources, in both the observer and the world. These differences have important practical implications for comparing and correcting perception and performance, and important theoretical implications for understanding the design principles underlying color coding. Color percepts within and between individuals often vary less than the variations in spectral sensitivity might predict. This stability is achieved by a variety of processes that compensate perception for the sensitivity limits of the eye and brain. Yet judgments of color between individuals can also vary widely, and in ways that are not readily explained by differences in sensitivity or the environment. These differences are uncorrelated across different color categories, and could reflect how these categories are learned or represented.

Variations in normal color vision. VII. Relationships between color naming and hue scaling.

A longstanding and unresolved question is how observers construct a discrete set of color categories to partition and label the continuous variations in light spectra, and how these categories might reflect the neural representation of color. We explored the properties of color naming and its relationship to color appearance by analyzing individual differences in color-naming and hue-scaling patterns, using factor analysis of individual differences to identify separate and shared processes underlying hue naming (labeling) and hue scaling (color appearance). Observers labeled the hues of 36 stimuli spanning different angles in cone-opponent space, using a set of eight terms corresponding to primary (red, green, blue, yellow) or binary (orange, purple, blue-green, yellow-green) hues. The boundaries defining different terms varied mostly independently, reflecting the influence of at least seven to eight factors. This finding is inconsistent with conventional color-opponent models in which all colors derive from the relative responses of underlying red-green and blue-yellow dimensions. Instead, color categories may reflect qualitatively distinct attributes that are free to vary with the specific spectral stimuli they label. Inter-observer differences in color naming were large and systematic, and we examined whether these differences were associated with differences in color appearance by comparing the hue naming to color percepts assessed by hue scaling measured in the same observers (from Emery et al., 2017). Variability in both tasks again depended on multiple (7 or 8) factors, with some Varimax-rotated factors specific to hue naming or hue scaling, but others common to corresponding stimuli for both judgments. The latter suggests that at least some of the differences in how individuals name or categorize color are related to differences in how the stimuli are perceived.

Variations in normal color vision. VI. Factors underlying individual differences in hue scaling and their implications for models of color appearance.

Observers with normal color vision vary widely in their judgments of color appearance, such as the specific spectral stimuli they perceive as pure or unique hues. We examined the basis of these individual differences by using factor analysis to examine the variations in hue-scaling functions from both new and previously published data. Observers reported the perceived proportion of red, green, blue or yellow in chromatic stimuli sampling angles at fixed intervals within the LM and S cone-opponent plane. These proportions were converted to hue angles in a perceptual-opponent space defined by red vs. green and blue vs. yellow axes. Factors were then extracted from the correlation matrix using PCA and Varimax rotation. These analyses revealed that inter-observer differences depend on seven or more narrowly-tuned factors. Moreover, although the task required observers to decompose the stimuli into four primary colors, there was no evidence for factors corresponding to these four primaries, or for opponent relationships between primaries. Perceptions of "redness" in orange, red, and purple, for instance, involved separate factors rather than one shared process for red. This pattern was compared to factor analyses of Monte Carlo simulations of the individual differences in scaling predicted by variations in standard opponent mechanisms, such as their spectral tuning or relative sensitivity. The observed factor pattern is inconsistent with these models and thus with conventional accounts of color appearance based on the Hering primaries. Instead, our analysis points to a perceptual representation of color in terms of multiple mechanisms or decision rules that each influence the perception of only a relatively narrow range of hues, potentially consistent with a population code for color suggested by cortical physiology.