Absence of linear subthreshold summation between red-green and luminance mechanisms over a wide range of spatio-temporal conditions

We have tested the independence of red-green chromatic and luminance mechanisms at detection threshold using a method of subthreshold summation. Stimuli were isoluminant red-green gratings and yellow-black luminance gratings that uniquely activate the red-green color and luminance mechanisms, respectively. Stimuli were Gaussian enveloped 0.25, 1 or 4 cpd sinewave gratings, counter-phase flickered at 0, 5 or 9 Hz. The threshold detection of red-green color contrast was measured in the presence of a subthreshold amount of luminance contrast, and vice versa. The results allow a model of linear summation between the color and luminance mechanisms to be rejected, but are well fitted by a model, assuming that these mechanisms are independent but combine to determine detection by probability summation, with a high summation index (median value = 4). We conclude that there are independent red-green chromatic mechanism and luminance detection mechanisms over this range of spatio-temporal conditions.     

Evidence for separate pathways for color and luminance detection mechanisms

We measure threshold versus contrast (TvC) functions for chromatic (red-green) and luminance sine-wave-grating stimuli for (1) the detection of luminance in the presence of color contrast and (2) the detection of color in the presence of luminance contrast. We find that, although these crossed TvC functions both display a dipperlike shape, their facilitation differs from that found for standard uncrossed dipper functions (luminance on luminance or color on color contrast). Their facilitation disappears (cross condition 1) or is reduced (cross condition 2) by randomized presentation of the phase of the test and the mask, and the remaining facilitation (cross condition 2) displays no spatial tuning. We argue that these crossed facilitatory interactions cannot be explained by detection mechanisms with common inputs from color and luminance contrast (a nonindependence of transduction), and we present evidence that instead they reflect the use of local cues in the stimuli. We also measure the luminance-luminance TvC function in the presence of a fixed suprathreshold color contrast. The results demonstrate that, even when the color contrast produces a masking of the luminance thresholds, luminance-luminance facilitation still occurs. Thus the opposing effects of masking and facilitation can occur simultaneously. Furthermore, while luminance-luminance facilitation occurs independently of color contrast, masking can be produced by either contrast. This suggests that masking and facilitation have different underlying origins. Similar results are found for the color detection thresholds in the presence of a luminance pedestal. We conclude that there are separate pathways for the detection of color and luminance contrast, each with no input from the other contrast. We suggest that the cross masking reflects divisive interactions between these pathways that is restricted to high contrasts.     

Stereoacuity and colour contrast

We have measured the contrast dependence of stereoacuity using both horizontally and vertically oriented, isoluminant (red-green) and isochromatic (yellow-black), 0.5 c/deg Gabor patches. For comparison, contrasts were computed in multiples of detection threshold, where detection threshold was defined as the contrast required for the stimulus to be simultaneously detectable in each eye. Disparity thresholds (1/stereoacuity) for vertical chromatic Gabors were higher than those for vertical luminance Gabors by a factor of between 4 and 9 depending on contrast, and declined less steeply with contrast. Disparity thresholds for horizontal chromatic Gabors were very high (130-210 min arc) compared with horizontal luminance Gabors (by a factor of between 9 and 17) and were only measurable at contrasts above 10 times simultaneous monocular detection threshold. These results support the view that chromatic stereoscopic processing is less precise than luminance stereoscopic processing, and that there is a special deficit in the processing of disparity with horizontally oriented chromatic stimuli. The implications of these results for the role of colour vision in stereopsis are discussed.     

Two motion systems with common and separate pathways for color and luminance

We present psychological experiments that reveal two motion systems, a specific and an unspecific one. The specific system prevails at medium to high temporal frequencies. It comprises at least two separate motion pathways that are selective for color and for luminance and that do not interact until after the motion signal is extracted separately in each. By contrast, the unspecific system prevails at low temporal frequencies and it combines color and luminance signals at an earlier stage, before motion extraction. The successful implementation of an efficient and accurate technique for assessing equiluminance corroborates further the main findings. These results offer a general framework for understanding the nature of interactions between color and luminance signals in motion perception and suggest that previously proposed dichotomies in motion processing may be encompassed by the specific/unspecific dichotomy proposed here.     

Differential effect of two calcium channel blockers--nifedipine and diltiazem--on atherogenesis in hypercholesterolemic hamster

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.     

Visual evoked potential assessment of the effects of glaucoma on visual subsystems

The purpose of this study is to test the hypothesis that glaucoma leads to selective deficits in parallel pathways or channels. Sweep VEPs were obtained to isolated-check stimuli that were modulated sinusoidally in either isoluminant chromatic contrast or in positive and negative luminance contrast. Response functions were obtained from 14 control subjects, 15 patients with open-angle glaucoma, and seven glaucoma suspects. For all three groups of subjects we found characteristic differences between the VEP response functions to isoluminant chromatic contrast stimuli and to luminance contrast stimuli. The isoluminant chromatic stimulus conditions appeared to favor activity of the P-pathway, whereas the luminance contrast stimuli at low depths of modulation favored M-pathway activity. VEP responses for patients with OAG were significantly reduced for chromatic contrast and luminance contrast conditions, whereas VEP responses for glaucoma suspects were significantly reduced only for the 15-Hz positive luminance contrast condition. Our results suggest that both M- and P-pathways are affected by glaucoma.     

Separate colour-opponent mechanisms underlie the detection and discrimination of moving chromatic targets

Current opinion holds that human colour vision is mediated primarily via a colour-opponent pathway that carries information about both wavelength and luminance contrast (type I). However, some authors argue that chromatic sensitivity may be limited by a different geniculostriate pathway, which carries information about wavelength alone (type II). We provide psychophysical evidence that both pathways may contribute to the perception of moving, chromatic targets in humans, depending on the nature of the visual discrimination. In experiment 1, we show that adaptation to drifting, red-green stimuli causes reductions in contrast sensitivity for both the detection and direction discrimination of moving chromatic targets. Importantly, the effects of adaptation are not directionally specific. In experiment 2, we show that adaptation to luminance gratings results in reduced sensitivity for the direction discrimination, but not the detection of moving chromatic targets. We suggest that sensitivity for the direction discrimination of chromatic targets is limited by a colour-opponent pathway that also conveys luminance-contrast information, whereas the detection of such targets is limited by a pathway with access to colour information alone. The properties of these pathways are consistent with the known properties of type-I and type-II neurons of the primate parvocellular lateral geniculate nucleus and their cortical projections. These findings may explain the known differences between detection and direction discrimination thresholds for chromatic targets moving at low to moderate velocities.     

Red-green and achromatic temporal filters: a ratio model predicts contrast-dependent speed perception

We simultaneously measured detection and identification performance by using isoluminant red-green (RG) and achromatic flickering stimuli and fitted these data with a modified line-element model that does not make high-threshold assumptions. The modeling shows that detection and identification data are adequately described by postulating only two underlying temporal filters each for RG and achromatic vision, even when more than two threshold classifications are evident. We use a spatial frequency of 1.5 cycles per degree (c/deg) and compare the derived temporal impulse response functions with those obtained previously with the use of 0.25 c/deg stimuli under otherwise identical conditions [J. Opt. Soc. Am. A 13, 1969 (1996)]. We find that at 1.5 c/deg the luminance impulse response functions peak later and integrate out to longer times compared with those measured at 0.25 c/deg. For RG stimuli, although their relative overall sensitivities change, the impulse response functions are similar across spatial frequency, indicating a constancy of chromatic temporal properties across spatial scales. In a second experiment, we measured RG and achromatic flicker discrimination over a wide range of suprathreshold contrasts. These data suggest a common nonlinear contrast response function operating after initial temporal filtering. Using a ratio model of speed perception in which both RG and achromatic filters are combined at a common motion site, we can predict (1) the perceived slowing of RG stimuli compared with the perceived drift of achromatic drifting stimuli, (2) the contrast dependency of speed perception for RG and achromatic drifting stimuli, and (3) how this dependency changes with base speed. Thus we conclude that there is no need to postulate separate mechanisms for fast and slow motion [Nature (London) 367, 268 (1994)], since a unified ratio model can explain both RG and achromatic contrast-speed dependency.     

Development of contrast sensitivity and temporal-frequency selectivity in primate lateral geniculate nucleus

We studied the development of spatial contrast-sensitivity and temporal-frequency selectivity for neurons in the monkey lateral geniculate nucleus. During postnatal week 1, the spatial properties of P-cells and M-cells are hardly distinguishable, with low contrast-sensitivity, sluggish responses, and poor spatial resolution. The acuity of P-cells improves progressively until at least 8 months, but there is no obvious increase in their maximum contrast-sensitivity with age. The contrast sensitivity of M-cells is already clearly higher than that of P-cells by 2 months, and at 8 months of age this characteristic difference between M- and P-cells approaches the adult pattern. There is a major increase in responsiveness during the first 2 postnatal months, especially for M-cells, the peak firing rate of which rises fivefold, on average, between birth and 2 months. Many P-cells in the neonatal and 2-month-old animals did not give statistically reliable responses to achromatic gratings, even at the highest contrasts: this unresponsiveness of P-cells might result from low gain and/or chromatic opponency. The upper limit of temporal resolution in the neonate is low--about one-third of that in the adult. Among M-cells, the improvement in temporal resolution, like that in contrast sensitivity, is rapid over the first 2 months, followed by a slower change approaching the adult value by 8 months of age. The development of contrast sensitivity, responsiveness and temporal tuning are little affected, if at all, by binocular deprivation of pattern vision from birth for even a prolonged period.     

Perifollicular clear space under skirt-like epithelial structure of human small vellus hair follicle

Peak latencies of pattern electroretinogram (PERG) were compared between glaucomatous eyes and non-glaucomatous eyes. Contrast threshold for motion perception (CTMP) was also measured with a new device in addition to routine static contrast sensitivity, static visual field and visual acuity. In the present recording, configurations for PERG, i.e. low reversal rate, low mean luminance and presence of background illumination, the PERG peak and trough (P1 and N2, respectively) latencies, were significantly prolonged in the glaucomatous eyes. Although both of the two PERG latencies were strongly correlated with the CTMP, only the P1 peak latency was strongly correlated with the perimetric indices. These results suggest that the P1 and N2 latencies reflect different aspects of signal processing in the retina, especially for moving targets.     

Reaction time to motion onset of luminance and chromatic gratings is determined by perceived speed

We measured reaction times for detecting motion onset for sinusoidal gratings whose contrast was modulated in either luminance or chromaticity, for various drift rates and contrasts. In general, reaction times to chromatic gratings were slower than to luminance gratings of matched cone contrast, but the difference in response depended critically on both contrast and speed. At high image speeds there was virtually no difference, whereas at low speeds, the difference was pronounced, especially at low contrasts. At high image speeds there was little dependence of reaction times on contrast (for either luminance or colour), whereas at low speeds the dependence was greater, particularly for chromatic stimuli. This pattern of results is reminiscent of those found for apparent speed of drifting luminance and chromatic gratings. We verified the effects of contrast on perceived speed, and went on to show that the effects of contrast on reaction times are totally predictable by the perceived speed of the stimuli, as if it were perceived rather than physical speed that determined reaction times. Our results support that idea of separate systems for fast and slow motion (with separate channels for luminance and colour at slower speeds), and further suggest that apparent speed and reaction times may be determined at a similar stage of motion analysis.     

On the binocular summation of chromatic contrast

The binocular summation of chromatic contrast was investigated under a variety of stimulus conditions. Binocular and monocular contrast detection thresholds were measured using 0.5 cpd Gabor patches. It was found that, using stimuli which contained combinations of chromatic and luminance contrast, binocular detection could take place independently in luminance-contrast- and chromatic-contrast-sensitive mechanisms. It was also found that, with chromatic stimuli, levels of binocular summation were above those expected from probability summation between the eyes, and thus showed evidence for binocular neural summation within chromatic detection mechanisms. The implications of these results for (a) the binocularity of chromatic detection mechanisms, and (b) the suggested link between stereopsis and binocular neural summation, are discussed.     

Scholarly mission. Fostering scholarship in research, theory, and practice

PURPOSE: Color vision testing in young children typically is precluded by the motor and cognitive skills required by standard tests; yet this information can be useful for diagnosis and counseling in many conditions. The purpose of this study is to evaluate a visual evoked potential (VEP) method for assessing red-green color vision anomalies in pediatric patients. METHOD: The relative chromatic luminance (C = R/R + G) of a rapidly reversing red-green checkerboard was varied across a wide range within a short viewing period (10 sec). Swept-parameter VEP methods were used to measure the cortical response to the range of C presented. RESULTS: Individuals with normal color vision exhibit a VEP response that exceeds noise levels across all values of C, often with an amplitude minima near the photopic equiluminant point (C = 0.5). Results from children with established protan and deutan color vision anomalies show loss of VEP amplitude and phase at values of C consistent with the respective color defect. A patient with achromatopsia showed a generalized depression of VEP response across all values of C tested. CONCLUSION: Color sweep VEP techniques appear promising for the clinical assessment of color status in pediatric patients.     

Comparison of red-green, blue-yellow and achromatic losses in glaucoma

Achromatic losses in glaucoma would be expected to be greater than, or equal to, red-green chromatic losses if the following assumptions are made: (1) the function of the remaining axons is either unchanged or non-selectively reduced; (2) red-green chromatic information is signaled by the midget ganglion cell system; and (3) the function of the magnocellular system is reduced at least as much as that of the midget ganglion cells. This prediction was tested by measuring red-green (along with blue-yellow) mixture thresholds for 1 deg, 0.2 sec test spots presented on a color monitor on a white background of 50 cd/m2. Ellipses were fitted to plots of green contrast as a function of red contrast (or yellow as a function of blue), and major and minor axes of these ellipses were taken as measures of chromatic and achromatic thresholds, respectively. The study population consisted of 29 eyes in 29 patients with early glaucoma; control data were derived from a data bank of 83 normal eyes. Red-green losses were significantly (P < 0.05) greater than achromatic losses in 6 out of the 11 eyes which showed significant losses of either chromatic or achromatic sensitivity (or both). It is concluded that, for these eyes, at least one of the above three assumptions is incorrect.     

Interaction of motion and color in the visual pathways

We measured perceived velocity as a function of contrast for luminance and isoluminant sinusoidal gratings, luminance and isoluminant plaids, and second-order, amplitude-modulated, drift-balanced stimuli. For all types of stimuli perceived velocity was contrast-invariant for fast moving patterns at or above 4 deg/sec. For slowly moving stimuli the log of perceived velocity was a linear function of the log of the contrast. The slope of this perceived velocity-vs-contrast line (velocity gain) was relatively shallow for luminance gratings and luminance plaids, but was steep for isoluminant gratings and isoluminant plaids, as well as for drift-balanced stimuli. Independent variation of spatial and temporal frequency showed that these variables, and not velocity alone, determine the velocity gain. Overall, the results indicate that slow moving stimuli defined by chromaticity or by second-order statistics are processed in a different manner from luminance defined stimuli. We propose that there are a number of independent mechanisms processing motion targets and it is the interplay of these mechanisms that is responsible for the final percept.     

Influence of stimulus size and contrast on the temporal parameters of saccadic eye movements: implications for road traffic

Saccadic eye movements are required for the recognition of peripheral objects in road traffic. Their latencies largely determine reaction time in emergency situations. The aim of this study was to investigate the influence of stimulus and surround parameters on the temporal characteristics of saccadic eye movements under conditions of object size, contrast and luminance corresponding to nighttime traffic. Square stimuli of 1 degree or 5 degrees size were presented under an eccentricity of 5 degrees and 15 degrees. The luminance of the surround was 4 x 10(-4) cd/m2 and 1 cd/m2. More than approx. 2,000 saccades of 7 normal subjects were registered and evaluated with respect to the latency, maximal velocity, and frequency of secondary saccades. At high stimulus contrast, latency approaches a minimum of approx. 200 ms. Latency increases with decreasing contrast up to maximal values of more than 600 ms. Transforming the contrast values into a relative decibel scale shows that this increase in latency occurs at significantly higher relative contrast values under scotopic as compared with photopic conditions. Our results demonstrate that an overall latency of 200-300 ms is not adequate for the assessment of accidents, especially under the stimulus conditions of nighttime traffic.     

Detection of chromatic and luminance contrast modulation by the visual system

The data presented in this paper examine the ability of observers to detect a modulation in the contrast of chromatic and luminance gratings as a function of the carrier contrast, duration, and spatial frequency. The nature of the signal underlying this ability is investigated by examining both the paradigm used to make the measurement and the effect of grating masks on performance in the tasks. The results show that observers' ability to discriminate amplitude modulation from an unmodulated carrier is dependent on carrier contrast but only up to approximately 5-8 times carrier-detection threshold. Discrimination is, however, independent of spatial frequency [10-1 cycles per degree (cpd) component-frequency range], carrier color, and, most surprisingly, stimulus duration (1000-30 ms). This set of experiments compliments data from previous papers and assimilates many of the conclusions drawn from this previous data. There is absolutely no evidence for the existence of a distortion product mediating performance under any of the current conditions, and the data seriously question whether the visual system might use such a signal even if it does exist under more extreme conditions than those used here. The evidence suggests that the visual system detects variations in both chromatic and luminance contrast by means of a mechanism operating locally upon the spatial structure of the carrier.     

Motion coherence across different chromatic axes

It has been reported that equiluminant plaid patterns constructed from component gratings modulated along different axes of a cardinal colour space fail to create a coherent impression of two-dimensional motion [Krauskopf and Farell (1990). Nature, 348, 328-331]. In this paper we assess whether this lack of interaction between cardinal axes is a general finding or is instead dependent upon specific stimulus parameters. Type I and Type II plaids were made from sinusoidal components (1 cpd) each modulated along axes in a cardinal colour space and presented at equivalent perceived contrasts. The spatial angular difference between the two components was varied from 5 to 90 deg whilst keeping the Intersection of Constraints (I.O.C.) solution of the pattern constant. Observers were required to indicate the perceived direction of motion of the pattern in a single interval direction-identification task. We find that: (i) When plaids were made from components modulated along the same cardinal axis, coherent "pattern" motion was perceived at all angular differences. As the angular difference between the components decreased in a Type II plaid, the perceived direction of motion moved closer to the I.O.C. solution and away from that predicted by the vector sum. (ii) A plaid made from components modulated along red-green and blue-yellow cardinal axes (cross-cardinal axis) did not cohere at high angular differences (> 30 deg) but had a perceived direction of the fastest moving component. At lower angular differences, however, pattern motion was detected and approached the I.O.C. solution in much the same way as a same-cardinal axis Type II plaid. (iii) A plaid made from a luminance grating and a cardinal chromatic grating (red-green or blue-yellow) failed to cohere under all conditions, demonstrating that there is no interaction between luminance and chromatic cardinal axes. These results indicate that there are conditions under which red-green and blue-yellow cardinal components interact for the purposes of motion detection.     

Luminance and spatial attention effects on early visual processing

Event-related potentials (ERPs) were recorded from healthy subjects in response to unilaterally flashed high and low luminance bar stimuli presented randomly to left and right field locations. Their task was to covertly and selectively attend to either the left or right stimulus locations (separate blocks) in order to detect infrequent shorter target bars of either luminance. Independent of attention, higher stimulus luminance resulted in higher ERP amplitudes for the posterior N95 (80-110 ms), occipital P1 (110-140 ms), and parietal N1 (130-180 ms). Brighter stimuli also resulted in shorter peak latency for the occipital N1 component (135-220 ms); this effect was not observed for the N1 components over parietal, central or frontal regions. Significant attention-related amplitude modulations were obtained for the occipital P1, occipital, parietal and central N1, the occipital and parietal P2, and the parietal N2 components; these components were larger to stimuli at the attended location. In contrast to the relatively short latencies of both spatial attention and luminance effects, the first interaction between luminance and spatial attention effects was observed for the P3 component to the target stimuli (350-750 ms). This suggests that interactions of spatial attention and stimulus luminance previously reported for reaction time measures may not reflect the earliest stages of sensory/perceptual processing. Differences in the way in which luminance and attention affected the occipital P1, occipital N1 and parietal N1 components suggest dissociations among these ERPs in the mechanisms of visual and attentional processing they reflect.(ABSTRACT TRUNCATED AT 250 WORDS)