Visual localization was studied by flashing small stimuli on a green background and requiring observers to press keys to indicate whether the stimulus appeared to the left or right of fixation. The results suggest that, for small (0.25 deg) briefly flashed (17 ms) stimuli at an eccentric location (10 deg), color contrast is not useable and localization presumably must rely on the magnocellular pathway. When stimulus size and duration were increased at 10-deg eccentricity, isochromatic stimuli could be localized at less than 10% luminance contrast (again suggesting use of the magnocellular high sensitivity luminance-contrast system), but isoluminant color-contrast stimuli could also be localized (suggesting use of the color-contrast sensitive parvocellular system). Thus, the results indicate that, dependent on stimulus conditions, both magnocellular and parvocellular pathways were utilized by normal observers in this localization task.

We recorded from single cells in area V2 of cynomolgus monkeys using standard acute recording techniques. After measuring each cell's spatial and temporal properties, we performed several tests of its chromatic properties using sine-wave gratings modulated around a mean gray background. Most cells behaved like neurons in area V1 and their responses were adequately described by a model that assumes a linear combination of cone signals. Unlike in V1, we found a subpopulation of cells whose activity was increased or inhibited by stimuli within a narrow range of color combinations. No particular color directions were preferentially represented. V2 cells showing color specificity, including cells showing narrow chromatic tuning, were present in any of the stripe compartments, as defined by cytochrome-oxidase (CO) staining. An addition of chromatic contrast facilitated the responses of most neurons to gratings with various luminance contrasts. Neurons in all three CO compartments gave significant responses to isoluminant gratings. Receptive-field properties of cells were generally similar for luminance and chromatically defined stimuli. We found only a small number of cells with a clearly identifiable double-opponent receptive-field organization.

We investigated the representation of color in cortical area V2 of macaque monkeys, and the association of color with other stimulus attributes. We measured the selectivity of individual V2 neurons for color, motion, and form. Most neurons in V2 were orientation selective, about half of them were selective for color, and a minority of cells (about 20%) were selective for size or direction. We correlated these physiological measurements with the anatomical location of the cells with respect to the cytochrome oxidase (CO) compartments of area V2. There was a tendency for color-selective cells to be found more frequently in the thin stripes, but color-selective cells also occurred frequently in thick stripes and inter-stripes. We found no difference in the degree of color selectivity between the different CO compartments. Furthermore, there was no negative correlation between color selectivity and selectivity for other stimulus attributes. We found many cells capable of encoding information along more than one stimulus dimension, regardless of their location with respect to the CO compartments. We suggest that area V2 plays an important role in integrating information about color, motion, and form. By this integration of stimulus attributes a cue-invariant representation of the visual world might be achieved.

Previous studies report that background luminance flicker, which is asynchronous with signal averaging, reduces the amplitude and increases the latency of the pattern-onset visual evoked potential (VEP). This effect has been attributed to saturation of the magnocellular (m-) pathway by the flicker stimulus. In the current study, we evaluate this hypothesis and further characterize this effect. We found that flicker had similar effects on the pattern-onset and pattern-reversal VEP, suggesting that the reversal and onset responses have similar generators. Chromatic flicker decreased latency of the chromatic VEP whereas luminance flicker increased peak latency to luminance targets. This result indicates that luminance flicker saturates a rapidly conducting m-pathway whereas chromatic flicker saturates a more slowly conducting parvocellular (p-) pathway. Finally, evoked potentials to chromatic and luminance stimuli were recorded from 34 electrodes over the scalp in the presence of static and asynchronously modulated backgrounds. An equivalent dipole model was used to assess occipital, parietal, and temporal lobe components of the surface response topography. Results showed that chromatic flicker reduced activity to a greater extent in the ventral visual pathway whereas luminance flicker reduced activity to a greater extent in the dorsal visual pathway to parietal lobe. We conclude that the VEP to isoluminant color and luminance stimuli contains both m- and p-pathway components. Asynchronous flicker can be used to selectively reduce the contribution of these pathways to the surface recorded VEP. Our results provide evidence of parallel pathways in the human visual system, with a dorsal luminance channel projecting predominantly to the posterior parietal lobe and a ventral color channel projecting predominantly to inferior temporal lobe.

Saccadic reaction times (SRTs) of three human subjects were analyzed. The gap paradigm was used (i.e. fixation point offset precedes target onset) to obtain high proportions of express saccades (i.e. saccades of extremely short reaction times) in the SRT distributions. In one set of experiments, the luminance of the (red) saccade target was varied from brighter to darker than the (green) background including an isoluminance condition. Express saccades were obtained in response to pure color contrast stimuli with about the same frequency and reaction time as to stimuli with both color and luminance contrast. In a second experiment, the luminance contrast of a white target on a white background was lowered below 10%. Again the number of express saccades was not reduced. Thus, in contrast to other perceptual phenomena the visual neural mechanisms underlying the generation of express saccades are not affected by isoluminance nor low contrast luminance.

We have studied the responses of MT neurons to moving gratings, spatially modulated in luminance and chromaticity. Most MT neurons responded briskly and with high contrast sensitivity to targets whose luminance was modulated, with or without added chromatic contrast. When luminance modulation was removed and only chromatic stimulation was used, the responses of all MT neurons were attenuated. Most were completely unresponsive to stimulation with targets whose modulation fell within a “null” plane in color space; these null planes varied from neuron to neuron, but all lay close to the plane of constant photometric luminance. For about a third of the neurons, there was no color direction in which responses were completely abolished; almost all of these neurons had a definite minimum response for chromatic modulation near the isoluminant plane. MT neurons that responded to isoluminant targets did so inconsistently and with poor contrast sensitivity, so that only intensely modulated targets were effective. Whereas the best thresholds of MT neurons for luminance targets are close to behavioral contrast threshold, the thresholds for isoluminant targets lie considerably above behavioral contrast threshold. Therefore, although some MT neurons do give responses to isoluminant targets, they are unlikely to be the source of the chromatic motion signals revealed behaviorally.

Simple reaction times (RTs) were measured to brief temporally blurred (total onset 570 ms) Gaussian isoluminant chromatic patches (s.d. 0.5°) whose chromaticities lay along the cardinal chromatic axes (0°, 90°, 180°, and 270° in MBDKL color space). Bipolar adapting stimuli were employed (0° versus 180° or 90° versus 270°). These were larger Gaussian blobs (s.d. 1°), modulating sinusoidally between the two hues at 1 Hz. Throughout, the background was illuminant “C” (x = 0.31, y = 0.316, L = 12.5). In a single run, a series of 64 or 32 stimuli were presented without adaptation, followed by 64 or 32 stimuli each of which was preceded by 3 s of adaptation, either along the same or the orthogonal chromatic axis. Finally, 192 or 128 RTs were recorded to measure the time course of recovery from adaptation. Both adapting and test stimuli were presented at fixed supra-threshold contrasts. The effect of adaptation was seen as a lengthening of the RT, which occurred in the first few seconds of the adaptation period. After cessation of adaptation, there was a similarly rapid shortening of RT, although full recovery took 60–90 s. Adaptation gain functions suggested that the S-(L + M) system was less prone to adaptation than L-M.

There is common consensus now that color-defined motion can be perceived by the human visual system. For global motion integration tasks based on isoluminant random dot kinematograms conflicting evidence exists, whether observers can (Ruppertsberg et al., 2003) or cannot (Bilodeau & Faubert, 1999) extract a common motion direction for stimuli modulated along the isoluminant red-green axis. Here we report conditions, in which S-cones contribute to chromatic global motion processing. When the display included extra-foveal regions, the individual elements were large (∼0.3°) and the displacement was large (∼1°), stimuli modulated along the yellowish-violet axis proved to be effective in a global motion task. The color contrast thresholds for detection for both color axes were well below the contrasts required for global motion integration, and therefore the discrimination-to-detection ratio was >1. We conclude that there is significant S-cone input to chromatic global motion processing and the extraction of global motion is not mediated by the same mechanism as simple detection. Whether the koniocellular or the magnocellular pathway is involved in transmitting S-cone signals is a topic of current debate (Chatterjee & Callaway, 2002).

The Zoellner illusion is a geometric distortion occurring when nonorthogonal inducing lines appear to tilt veridically parallel bars. The retinal pathways contributing to such illusions are unknown. The goal of this experiment was to investigate the retinal origin of the illusion. This was accomplished by determining the contrast gain for illusion thresholds. The magnocellular (MC-) and parvocellular (PC-) pathways exhibit different contrast gains, and this difference can be used psychophysically to identify the pathway. The stimulus pattern was four vertical bars with a series of inducing lines. The bars were always 5% higher in contrast than the inducing bars. The pattern was presented on a larger pedestal. Two paradigms were used. In the pulsed-pedestal paradigm, the observer adapted to the background and the pedestal and pattern were presented together as a brief pulse. In the steady-pedestal paradigm, the observer adapted to the continuously presented pedestal and the pattern appeared as a brief pulse. The contrast between the pedestal and the pattern was varied to obtain thresholds for two criteria: perceiving the directions of the inner inducing lines, and perceiving the distortion of the bars. The results for both criteria were similar in shape, but displaced in sensitivity. Detection of the directions of the inner inducing lines was 0.16–0.29 log unit more sensitive than perception of the illusion. The data for the pulsed-pedestal paradigm depended on the contrast between the pedestal and the pattern and produced a shallow V-shape. These results were associated with mediation in the PC-pathway. The data for the steady-pedestal paradigm depended on the pedestal luminance in a linear relation and showed similar sensitivity to the data for the pulsed-pedestal paradigm. Perception of the illusion required 10–15% Weber contrast.

For over 30 years there has been a controversy over whether color-defined motion can be perceived by the human visual system. Some results suggest that there is no chromatic motion mechanism at all, whereas others do find evidence for a purely chromatic motion mechanism. Here we examine the chromatic input to global motion processing for a range of color directions in the photopic luminance range. We measure contrast thresholds for global motion identification and simple detection using sparse random-dot kinematograms. The results show a discrepancy between the two chromatic axes: whereas it is possible for observers to perform the global motion task for stimuli modulated along the red–green axis, we could not assess the contrast threshold required for stimuli modulated along the yellowish-violet axis. The contrast required for detection for both axes, however, are well below the contrasts required for global motion identification. We conclude that there is a significant red–green input to global motion processing providing further evidence for the involvement of the parvocellular pathway. The lack of S-cone input to global motion processing suggests that the koniocellular pathway mediates the detection but not the processing of complex motion for our parameter range.

The color vision of Old World primates and humans uses two cone-opponent systems; one differences the outputs of L and M cones forming a red–green (RG) system, and the other differences S cones with a combination of L and M cones forming a blue–yellow (BY) system. In this paper, we show that in human vision these two systems have a differential distribution across the visual field. Cone contrast sensitivities for sine-wave grating stimuli (smoothly enveloped in space and time) were measured for the two color systems (RG & BY) and the achromatic (Ach) system at a range of eccentricities in the nasal field (0–25 deg). We spatially scaled our stimuli independently for each system (RG, BY, & Ach) in order to activate that system optimally at each eccentricity. This controlled for any differential variations in spatial scale with eccentricity and provided a comparison between the three systems under equivalent conditions. We find that while red–green cone opponency has a steep decline away from the fovea, the loss in blue–yellow cone opponency is more gradual, showing a similar loss to that found for achromatic vision. Thus only red–green opponency, and not blue–yellow opponency, can be considered a foveal specialization of primate vision with an overrepresentation at the fovea. In addition, statistical calculations of the level of chance cone opponency in the two systems indicate that selective S cone connections to postreceptoral neurons are essential to maintain peripheral blue–yellow sensitivity in human vision. In the red–green system, an assumption of cone selectivity is not required to account for losses in peripheral sensitivity. Overall, these results provide behavioral evidence for functionally distinct neuro-architectural origins of the two color systems in human vision, supporting recent physiological results in primates.

We report VEP studies which delineate interactions between chromatic and luminance contrast signals. We examined responses to sinusoidal luminance gratings undergoing 4-Hz square-wave contrast reversal, upon which standing gratings with various admixtures of luminance and chromatic contrast were alternately superimposed and withdrawn. The presence of the standing grating induced a VEP component at the fundamental frequency of the contrast-reversal grating. This VEP component appeared without any appreciable lag, and did not vary in amplitude over the 4 s during which the standing grating was present. The observed fundamental response differed from the fundamental component that would be expected from the known interaction between the luminance component of the standing grating with the modulated grating (Bodis-Wollner et al., 1972; Bobak et al., 1988), in three ways: (1) The fundamental response was not nulled for standing gratings that were isoluminant or near-isoluminant. (2) The chromatic dependence of the fundamental response implied an S-cone input to the interaction. (3) No single mechanism (driven by a linear combination of cone signals) could account quantitatively for the size of this response, particularly when the standing grating strongly modulated two cones in phase.