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.

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.

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.

Binocular vs. monocular hue perception.

Hue perception has been shown to differ for the same stimulus when presented to the temporal and the nasal areas of the retina. The present study investigated perceptual differences in stimuli viewed binocularly or monocularly in the peripheral retina to determine how hue information combines across the two retinas for a stimulus falling on the temporal retina of one eye and the nasal retina of the other. A hue-scaling procedure was utilized to ascertain hue perception for three color- and binocular-normal observers viewing monochromatic stimuli (450-670nm, 20nm steps) ranging in size from 1.0° to 3.7°. Peripherally-presented binocular stimuli fell upon the nasal retina of one eye and the temporal retina of the other. Hue-scaling results indicated that peripheral binocular hue and saturation perceptions for smaller stimuli were more similar to those of stimuli falling on the temporal retina in the monocular condition. Hue-scaling data were also used to determine perceptive field sizes for the four elemental hues. Binocular perceptive field sizes were more similar to those obtained for stimuli falling on the temporal retina in the monocular conditions. Eye dominance did not appear to have an effect on hue perception. The results seem to indicate that visual information from the temporal retina is weighted more heavily when information from the two eyes is combined cortically. This finding may relate to differences in V1 cortical activation for stimuli presented to the nasal retina versus the temporal retina.

Small field tritanopia in the peripheral retina.

If stimuli are made sufficiently small, color-normal individuals report a loss in hue perception, in particular a decrease in the perception of green, in both the fovea and peripheral retina. This effect is referred to as small field tritanopia. It is not clear, however, how rod input may alter the dynamics of small field tritanopia in the peripheral retina. This paper looks at peripheral hue-naming data obtained for small stimuli at mesopic and photopic retinal illuminances under conditions that minimize (bleach) and maximize (no bleach) rod contribution. The data show that attenuation in the perception of green occurs with larger stimuli in the no-bleach condition than in the bleach condition. As retinal illuminance increases, the stimulus size that elicits small field tritanopia decreases, but the stimulus size is still larger under the no-bleach condition. Small field tritanopia in both the bleach and no-bleach conditions may be related to short-wavelength-sensitive (S) cone activity and its potential role in the mediation of the perception of green. The differences in stimulus size for small field tritanopia may be explained by rod input into the magnocellular and koniocellular pathways, which compromises the strength of the chromatic signals and creates a differential loss in the perception of green as compared to the other elemental hues.

Supersaturation in the peripheral retina.

Foveal and peripheral hue-scaling data were obtained for a 1° foveal stimulus and a 3° stimulus presented at 10° retinal eccentricity under both bleach (reducing rod input) and no-bleach (permitting rod input) conditions. Uniform appearance diagrams (UADs) were generated from the data. Peripheral stimuli appeared more saturated than foveal stimuli (i.e., supersaturated), especially in the green-yellow region of the UADs. This effect was particularly pronounced for the peripheral bleach condition. The range of wavelengths perceived as green-yellow in the peripheral retina was expanded as compared to the fovea, while the range of wavelengths experienced as blue-green was compressed. This indicates that there are shifts in the unique hue loci with retinal location. While several factors can be ruled out as potential causes for these perceptual differences, the underlying mechanism of this supersaturation effect in the peripheral retina is unknown.

Color appearance at ±10° along the vertical and horizontal meridians.

Hue-scaling data were collected from three observers using the "4+1" color-naming procedure for circular (0.25°-5°), monochromatic (440-660 nm) stimuli. Stimuli were presented at ±10° along the vertical and horizontal meridians under conditions chosen to include both rod and cone signals (no bleach) and to minimize rod contribution (bleach). All color-naming data were analyzed and compared using uniform appearance diagrams. Smaller stimuli appear more desaturated under both bleach conditions. This effect is particularly detrimental for the perception of green and is influenced by retinal location and exacerbated with rod input. As stimulus size increases and perceptive field sizes are filled for all four elemental hues, the differences in hue perception among the four peripheral locations and the two bleach conditions are attenuated. Results are consistent with predictions based on known differences in the underlying retinal mosaic among the four locations.

Middle- and long-wavelength discrimination declines with rod photopigment regeneration.

Hue-discrimination functions were derived from hue-naming data (480-620 nm, 20 nm steps) obtained in 4 min intervals from 4 min to 28 min postbleach at 10° temporal retinal eccentricity. Hue-naming data were also obtained in the fovea. Hue-discrimination functions derived at the 4, 8, and 12 min intervals were very similar to those derived in the fovea. As time postbleach exceeded 12 min and rod sensitivity increased, the shape of the hue-discrimination functions changed. Most notably, the minimum between 560-580 nm disappeared and the just noticeable differences (JNDs) for the longer wavelength stimuli increased. The long-wavelength suppression in hue discrimination may be due to rod input in the magnocellular pathway interacting and affecting the long-wavelength sensitivity of the parvocellular pathway.

Unique hue loci differ with methodology.

BACKGROUND: Studies investigating the effect of rods on unique hue loci in the peripheral retina generally obtain measures at two time points associated with the dark adaptation function - the cone plateau and the rod plateau. In comparison, this study used a color-naming procedure to identify the loci of unique green and unique yellow as a function of time associated with the entire dark adaptation function. The unique hue loci derived by this procedure were then compared to those obtained directly with a staircase procedure. METHOD: Hue-scaling functions were obtained for monochromatic stimuli for four observers using the '4 + 1' procedure. Data were collected every 4 min following extinction of a bleaching stimulus. These hue-scaling functions were then converted to uniform appearance diagrams (UADs) to derive unique green and unique yellow loci. Unique green and unique yellow loci were also obtained from the same observers via a staircase procedure at 4-9 min post-bleach (minimal rod input) and after 28 min dark adaptation (maximal rod input). Measurements were made in the peripheral retina at 10° temporal retinal eccentricity and at the fovea. RESULTS: Unique green loci derived from UADs are at longer wavelengths compared to those measured directly with the staircase procedure. In addition, unique green loci derived from UADs show a progressive shift to longer wavelengths as time post-bleach increases; whereas, unique green loci obtained from the staircase procedure differ little between the rod-bleach and no-bleach conditions. Unique yellow loci are similar across both experimental procedures. CONCLUSION: Unique green loci derived from UADs are not the same as those measured with traditional psychophysical procedures. These differences may be due to the different response criteria used by observers in the color-naming and staircase procedures. The unique green loci obtained from UADs indicate that rod signals shift unique green loci to longer wavelengths as time post-bleach increases. More direct methods need to be employed to determine if this rod effect is valid.

Scotopic spatiotemporal sensitivity differences between young and old adults.

BACKGROUND: Our lab has previously demonstrated losses in contrast sensitivity to low spatial frequencies under scotopic conditions with older adults. It is not clear, however, whether the temporal frequency of a stimulus alters the relation between age and the spatial contrast sensitivity function (sCSF) under scotopic conditions. METHODS: A maximum-likelihood, two-alternative, temporal forced-choice QUEST procedure was used to measure threshold to spatially and temporally modulated stimuli in both young (mean = 26 years) and old (mean = 75 years) adults. RESULTS: In general, the shapes of the spatial and temporal CSFs were low-pass for both young and old observers; contrast sensitivity decreased at approximately the same rate with increasing spatial frequency and temporal frequency for both age groups, although the overall sensitivity of the old group was lower than that of the young group. The high-frequency resolution limit was lower for the old group compared to the young group. CONCLUSIONS: The differences in contrast sensitivity between the young and old groups suggest a uniform loss in sensitivity of the channels mediating spatial and temporal vision. Because of this loss, the spatial and temporal window of visibility for the older adults is compromised relative to the younger adults.

Chromatic perceptive field sizes measured at 10 degrees eccentricity along the horizontal and vertical meridians.

The different hemifields in the retina are known to vary in photoreceptor density as well as in the number of photoreceptors converging onto one ganglion cell. The effect of these differences among the retinal hemifields at 10 degrees retinal eccentricity was investigated using a color-naming procedure to derive perceptive field sizes for the hue terms of blue, green, yellow, and red. Color-naming data were obtained under two conditions: (1) after a bleach condition, chosen to minimize rod contribution, and (2) after 30 min dark adaptation, chosen to maximize rod contribution. Perceptive field sizes measured in the bleach condition were consistent with degree of neural convergence of cones to ganglion cells across the retina rather than differences in cone density. Rod densities relative to cone densities correlated with the size of perceptive fields in the no-bleach condition, i.e., the greater the rod:cone ratio, the larger the perceptive field.

Effect of stimulus intensity on the sizes of chromatic perceptive fields.

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

Chromatic perceptive field sizes change with retinal illuminance.

The effect of retinal illuminance (0.3-3.3 log td) on chromatic perceptive field size was investigated at 10 degrees eccentricity along the horizontal meridian of the temporal retina. Using the 4+1 color-naming procedure, observers described the hue and saturation of a series of monochromatic stimuli (440-660 nm, in 10-nm steps) of various test sizes (.098-5 degrees) after 30-min dark adaptation. Perceptive field sizes of the four elemental hues and the saturation component were estimated for each wavelength at each retinal illuminance. Results indicate that perceptive field sizes for blue, green, yellow, and saturation all decrease with increasing retinal illuminance; the perceptive field size for red is the smallest and invariant with intensity. The influence of rods on perceptive field size may account for some of the results; other factors are also considered.

A new look at the Bezold-Brücke hue shift in the peripheral retina.

Experiments were conducted with a bipartite field to better understand the Bezold-Brücke hue shift in the peripheral retina. The first experiment measured hue shift in the fovea and at 1 degrees and 8 degrees along the horizontal meridian of the nasal retina for nominal test wavelengths of 430, 450, 490, 520 and 610 nm. Peripheral measurements were obtained under two adaptation conditions: after 30 min dark adaptation and following a rod-bleach. Results indicated that foveal hue shifts differed from those obtained after a rod-bleach. Data from the rod-bleach and no-bleach conditions in the periphery were similar, indicating that rods could not account for the differences between the foveal data and the rod-bleach peripheral data. Hue shifts obtained for the 520 nm test stimulus, and to a smaller extent other test wavelengths, at 8 degrees nasal retinal eccentricity revealed that the wavelength of the matching stimulus depended upon the lateral position of the matching and test fields, and this effect was greater in the no-bleach condition than the rod-bleach condition. Several factors were investigated in experiments 2 and 3 to explain the results with the 520 nm test field. It appears that differential rod density under the two half fields and the compression of photoreceptors by the optic disk may partially, but not fully, account for the 520 nm effect.

Ricco's Areas for S- and L-Cone Mechanisms Across the Retina.

The purposes of this study were to measure areas of complete spatial summation (i.e., Ricco's area) for S- and L-cone mechanisms and to evaluate whether the sizes of Ricco's area could be explained in terms of either the densities of photoreceptors or ganglion cells. Increment thresholds were measured at the fovea and at 1.5°, 4°, 8°, and 20° in the superior retina using a temporal two-alternative forced-choice procedure. Test stimuli ranging from -0.36 to 4.61 log area (min(2)) were presented on concentric 12.3° adapting and auxiliary fields, which isolated either an S- or L-cone mechanism on the plateau of the respective threshold vs. intensity function. The data indicate that from 0-20° retinal eccentricity, the size of Ricco's area is larger for the S-cone mechanism than the L-cone mechanism, increases monotonically for the L-cone mechanism, and, for both cone mechanisms, increases between 8-20° retinal eccentricity. This latter finding suggests that ganglion cell density rather than cone density defines the size of Ricco's area in the parafoveal and peripheral retina.