Luminance mechanisms mediate the motion of red-green isoluminant gratings: the role of "temporal chromatic aberration"

In this paper we use a dynamic noise-masking paradigm to explore the nature of the mechanisms mediating the motion perception of drifting isoluminant red-green gratings. We compare contrast thresholds for the detection and direction discrimination of drifting gratings (1.5 cpd), over a range of temporal frequencies (0.5-9 Hz) in the presence of variable luminance or chromatic noise. In the first experiment, we used dynamic luminance noise to show that direction thresholds for red-green grating motion are masked by luminance noise over the entire temporal range tested, whereas detection thresholds are unaffected. This result indicates that the motion of nominally isoluminant red-green gratings is mediated by luminance signals. We suggest that stimulus-based luminance artifacts are not responsible for this effect because there is no masking of the detection thresholds. Instead we propose that chromatic motion thresholds for red-green isoluminant gratings are mediated by dynamic luminance artifacts that have an internal, physiological origin. We have termed these "temporal chromatic aberration". In the second experiment, we used dynamic chromatic noise masking to test for a chromatic contribution to red-green grating motion. We were unable to find conclusive evidence for a contribution of chromatic mechanisms to the chromatic grating motion, although a contribution at very high chromatic contrasts cannot be ruled out. Our results add to a growing body of evidence indicating the presence of dynamic, internal luminance artifacts in the motion of chromatic stimuli and we show that these occur even at very low temporal rates. Our results are compatible with our previous work indicating the absence of a chromatic mechanism for first order (quasi-linear) apparent motion [Vision Res. 40 (2000) 1993]. We conclude that previous conclusions based on the motion of chromatic red-green gratings should be reassessed to determine the contribution of dynamic luminance artifacts.     

S-cone contributions to linear and non-linear motion processing

We investigated the characteristics of mechanisms mediating motion discrimination of S-cone isolating stimuli and found a double dissociation between the effects of luminance noise, which masks linear but not non-linear motion, and chromatic noise, which masks non-linear but not linear motion. We conclude that S-cones contribute to motion via two different pathways: a non-linear motion mechanism via a chromatic pathway and a linear motion mechanism via a luminance pathway. Additionally, motion discrimination and detection thresholds for drifting, S-cone isolating Gabors are unaffected by luminance noise, indicating that grating motion is mediated via chromatic mechanisms and based on higher-order motion processing.     

The contribution of color to global motion processing

This study investigates the contribution of color vision to global motion. We present evidence demonstrating that performance on a global motion task attains similar levels for both types of chromatic (L/M-cone opponent and S-cone opponent) and luminance stimuli at suprathreshold contrasts. We show, however, that the motion thresholds for isoluminant chromatic stimuli are luminance based, on the grounds that they are masked by luminance noise but robust to chromatic noise. Detection thresholds, on the other hand, are chromatic in origin (masked by chromatic but not luminance noise), indicating that there is no luminance artifact in the stimulus. We suggest that for color vision at isoluminance the global motion task is based on the integration of many local, luminance-based signals.     

Global motion processing in human color vision: a deficit for second-order stimuli

The investigation of the mechanism of global motion in color vision has been limited because the processing of the first-order chromatic RDK elements, based on low-level linear motion detectors, is impaired. Here we return to this problem by using second-order elements in a global motion stimulus. Second-order RDK elements were circular contrast-modulated (CM) envelopes of a low-pass filtered noise carrier. The stimuli were achromatic or isolated L/M- or S-cone opponent mechanisms. We measured simultaneously detection and motion direction identification thresholds at 100% motion coherence and at different RDK speeds with a 2-AFC paradigm. We found that direction identification thresholds were higher than detection thresholds for both chromatic and achromatic stimuli. The gap between these thresholds was greater for the chromatic than the achromatic stimuli and motion direction thresholds for the chromatic RDK were very high or impossible to obtain. We also measured global motion performance (RDK speed of 4 deg/s) by varying the coherence of limited lifetime RDK stimuli. Global motion thresholds could only be obtained for achromatic stimuli and not for chromatic ones. Within the limits of the present stimulus conditions, we found no global motion processing of second-order chromatic stimuli.     

The detection of motion in chromatic stimuli: first-order and second-order spatial structure

This study provides evidence for the existence of a low-level chromatic motion mechanism and further elucidates the conditions under which its operation becomes measurable in an experimental stimulus. Observers discriminated the direction of motion of amplitude modulated (AM) gratings that were defined by luminance or chromatic variation and masked with spatiotemporally broadband luminance or chromatic noise. The size and retinal location of the stimuli were varied and the effects of broadband noise and grating masks were both compared with the cohort of stimuli. Some significant disparities in the published literature were well explained by the results. In conclusion, evidence for a chromatically sensitive motion mechanism that evades the, detrimental effects of a luminance mask was found only at the fovea and only when the stimulus was small and centrally placed.     

Theoretical predictions and experimental findings of visual deficits in glaucoma

Purpose: To investigate the motion and colour processing in normal subjects and in patients with primary open angle glaucoma, and to develop theoretical models in order to explain present findings and to understand the processes involved. Methods: Movement processing was investigated using an optical motion projection system, capable of generating continuous motion of a single dot target with controlled speed, displacement and motion direction. Displacement and direction discrimination thresholds (i.e. target displacement at a constant speed needed for 75% probability of detection of motion or correct discrimination of its direction) were measured for both foveal and peripheral stimulus presentations and for a range of speeds. Colour vision was tested using a novel method which employs a random luminance modulation technique to mask out achromatic contrast changes and allowing detection of a spatially structured object based on the processing of chromatic signals only (Barbur et al. 1993). Results: Changes in the parameters of the motion detection model (e.g., delay time and receptor regularity) that was developed to predict the experimental data in normal subjects also predicts an increase in displacement threshold and poorer sensitivity for discrimination of motion direction in glaucoma patients. The results obtained show that patients may report stimulus motion in 80% of presentations whilst performing at chance when required to discriminate the direction of motion. Chromatic discrimination ellipses measured for foveal stimulus presentations in glaucoma patients were often similar to those measured in normal subjects. In the periphery, however, much larger chromatic displacement thresholds were measured in all directions. Conclusions: The technique used to measure the chromatic discrimination ellipses is sensitive to glaucomatous nerve damage and the results suggest a larger threshold elevation in the periphery than at the fovea. The motion model developed to explain performance in normal vision also predicts the experimental findings in glaucoma patients and may be used to predict degraded motion discrimination in patient groups with similar pathologies.     

The perception of speed based on L-M and S-(L+M) cone opponent processing

We have measured perceived speed and speed discrimination thresholds for stimuli that selectively activate the L-M, S-(L+M) cone opponent and L+M (luminance) post-receptoral pathways. For low speeds and low contrasts speed discrimination thresholds for L-M and S-(L+M) are similar but are higher and have a greater dependency upon contrast than those for luminance motion. These differences between chromatic and luminance speed perception can be eliminated when stimuli are equated with respect to their individual motion detection thresholds (MDTs). For fast moving gratings speed perception based upon L-M, S-(L+M) and L+M signals is similar in terms of threshold performance and contrast dependency. These results are consistent with the view that there are separate mechanisms for the analysis of chromatic and luminance motion, the relative contributions of which may change as a function of stimulus contrast and speed. The similarity in performance for S-(L+M) and L+M chromatic stimuli across a range of stimulus parameters suggests that signals derived from the two cone opponent pathways can be used equally well. Our results argue against the idea that speed perception is compromised when it is based upon information derived from the S-(L+M) cone opponent pathway.     

Velocity tuned mechanisms in human motion processing

We determined two-dimensional motion discrimination contours in the spatio-temporal frequency plane to characterize the mechanisms underlying velocity perception. In particular, we wanted to determine whether there exist mechanisms tuned specifically to velocity, rather than separable mechanisms tuned to spatial and temporal frequency. A 4-AFC paradigm was used to determine spatio-temporal frequency discrimination thresholds for moving sinewave gratings defined by luminance contrast. Three of the grating patches used were defined by the same spatial and temporal frequency (standard), the other (test) differed by a fixed proportional change in spatial and temporal frequency. Subjects had to indicate which grating differed most from the others and the thresholds determined for varying proportions of change in spatial and temporal frequency were used to trace out complete threshold contours in the plane spanned by these attributes. Some of the contours, primarily at speeds above 1 deg/s, were noticeably oriented along lines of constant velocity. To further isolate these mechanisms, spatio-temporal noise was added to the standard stimuli either along a line of constant velocity or in the direction orthogonal to it. When spatio-temporal noise of constant velocity was added to the standard stimuli, threshold contours became elongated only along the direction of the noise. The same amount of noise in the orthogonal direction produced an overall increase in thresholds without changing the shape of the contour, presenting clear evidence for velocity tuned mechanisms. In further experiments we discovered that velocity tuned mechanisms interact with separable mechanisms to produce optimal discriminability. Analogous experiments with isoluminant stimuli failed to exhibit evidence for velocity tuning, supporting the notion that the human color vision system is impaired in its coding of stimulus speed, despite excellent sensitivity to direction of motion.     

The contribution of color to motion in normal and color-deficient observers

By opposing drifting luminance and color gratings, we have measured the "equivalent luminance contrast" of color, the contribution that color makes to motion. We found that this equivalent contrast was highest (greater than 10%) for low spatial and temporal frequencies and was higher for red/green than for blue/yellow stimuli. Equivalent luminance contrast was about 4% for a green/purple stimulus that fell along the tritan confusion line, indicating a modest input to the motion pathway from the short wavelength-sensitive cones (B-cones). Contrast thresholds for the discrimination of the direction of motion showed that the contribution of color to motion was about the same (within a factor of 2) as that for luminance in terms of multiples of threshold contrast. These responses to moving, chromatic gratings could be mediated by any of several factors that can create a residual response in a luminance pathway: temporal phase lag between the responses to the colors of the stimuli, second harmonic distortion in the response and variability in equiluminance points across units. Each of these factors was evaluated experimentally and their combined effect could account for only a small portion of the contribution of color to motion. As a result, we attribute the perception of the motion of equiluminous stimuli to an opponent-color input to directionally selective cortical units. Chromatic stimuli had little or no equivalent contrast for color-deficient observers, whether the stimulus was red/green, which they discriminate less well than normals, or blue/yellow, which they discriminate almost as well as normals. The equivalent contrast measure provided an excellent basis for classifying normal, protan and deutan observers.     

Orientation selectivity in luminance and color vision assessed using 2-d band-pass filtered spatial noise

We evaluated orientation discrimination in color and luminance vision using an external noise paradigm. Stimuli were spatiotemporal patches of 2D orientation noise isolating the achromatic, red-green and blue-yellow mechanisms, and matched in multiples of contrast detection threshold. We found a monotonic increase of orientation discrimination thresholds with the stimuli orientation bandwidths that is similar for both color and luminance contrasts. This dependence was fitted with two suitable models. A variance summation model suggests that internal orientation noise is significantly greater for the chromatic than for the achromatic mechanisms, while the efficiencies are similar. A gain control model of orientation tuning suggests that both chromatic and achromatic mechanisms are characterized by broadly tuned orientation detectors and that the relative chromatic deficit in orientation discrimination may only result from a slightly broader orientation tuning for the chromatic mechanisms. The moderate deficiency in chromatic orientation discrimination may account for the small differences found in shape perception between color and luminance vision.     

Spatial frequency discrimination: a comparison of achromatic and chromatic conditions

In this study, we have compared foveal SF discriminations for luminance and color-defined stimuli using two different tasks (criteria): in criterion-A, the discrimination is based on spatial (size of the stimuli) and/or spatial frequency; in criterion-B, it is based on apparent motion (contraction/expansion). We used high contrast (75%) spatially localized D6 stimuli and cosine gratings (0.25-9.5 cpd). The SF discrimination was measured by the method of constant stimuli with a two-interval forced-choice procedure. Data show that: (i) for criterion-A, the discrimination thresholds for color stimuli were lower than that for luminance stimuli at low SFs, but similar or higher at higher SFs; for criterion-B, the thresholds to chromatic stimuli were higher than that to achromatic stimuli for all SFs; (ii) SF discrimination was best at inter-stimulus-interval (ISI) of about 200 ms for color stimuli and at ISI of 0 ms for luminance stimuli; (iii) SF discrimination got better with stimulus duration and reached to plateau at 200 ms (or more) for color stimuli and at 67 ms (or more) for luminance stimuli; (iv) SF discrimination threshold (mean Delta(f)=0.19 octaves) is about one-tenth of the full bandwidth (mean=1.96 octaves) of SF tuned mechanisms and is in hyperacuity range; both (discrimination and hyperacuity) can be explained by the relative activities within a population of tuned mechanisms. We conclude that color and luminance SF discrimination thresholds have a different SF dependence. While color appears to perform better than luminance vision at low SFs, this effect is lost or even reversed at high SFs. Data imply that color and form interact, but color and motion are largely segregated (i.e. they weakly interact).     

Determining the visibility of noise in motion signals.

A procedure is described for generating stimuli to study the detection of noise components in motion signals. By using random dots with intensities distributed according to a Gaussian probability function, a temporally and spatially continuous mixture of signal and noise components can be realized in random dot kinematograms. These stimuli were used in a noise detection task, a signal detection task and a direction discrimination task. Signal-to-noise ratio ('coherence') thresholds for the signal detection and direction discrimination tasks were consistent with previous research. Noise can be detected at levels of approximately 0.5-2.5%, depending on the size of the motion stimulus. We argue that the noise in the motion stimulus becomes detectable when it exceeds the noise intrinsic to the various stages of motion processing. Therefore,the method provides a simple procedure for obtaining measures of equivalent input noise and can be used for estimating internal noise levels of motion processing mechanisms.     

A single mechanism for both luminance and chromatic grating vernier tasks: evidence from temporal summation

Vernier thresholds are determined by luminance rather than chromatic contrast when both are present in vernier targets. The role of luminance and chromatic mechanisms in vernier performance under equiluminant conditions remains uncertain. Temporal summation functions for vernier thresholds with luminance and red-green equiluminant gratings were compared to those for detection thresholds with similar stimuli. Vernier thresholds showed similar temporal summation for luminance and chromatic gratings, which is consistent with a single mechanism underlying vernier performance in the two conditions. However, detection thresholds showed a shorter temporal summation duration for luminance gratings than for chromatic gratings, which suggests that two different mechanisms underlie detection thresholds. Analysis of physiological data supports the hypothesis that the frequency-doubled response of ganglion cells in the magnocellular pathway can provide accurate spatiotemporal information for vernier performance at equiluminance.     

The temporal properties of the response of macaque ganglion cells and central mechanisms of flicker detection

This analysis assesses sensitivity of primate ganglion cells to sinusoidal modulation as a function of temporal frequency, based on the structure of their impulse trains; sensitivity to luminance and chromatic modulation was compared to human psychophysical sensitivity to similar stimuli. Each stimulus cycle was Fourier analyzed, and response amplitudes subjected to neurometric analysis; this assumes a detector with duration inversely proportional to frequency, that is, the stimulus epoch analyzed is a single cycle rather than a fixed duration, and provides an upper bound for a detection by an observer who bases judgments on a single cell. Signal-to-noise ratio for a given Fourier amplitude rapidly decreased with temporal frequency. This is a consequence of the statistics of impulse trains making up the response; at higher temporal frequencies, there are fewer impulses per cycle. Performance of this "single-cell" observer was then compared with that of modeled central detection mechanisms of fixed duration. For chromatic modulation, a filter/detector with a time constant of approximately 40 ms operating upon the parvocellular (PC) pathway provided a match to psychophysical results, whereas for luminance modulation, a filter/detection mechanism operating upon the magnocellular (MC) pathway with a time constant of approximately 5-10 ms provided a suitable match. The effects of summation and nonlinear interactions between cell inputs to detection are also considered in terms of enhanced sensitivity and "sharpness" of thresholds, that is, the steepness of the neurometric function. For both luminance (MC cells) and chromatic modulation (PC cells), restricted convergence (<20 cells) appears adequate to provide sharp thresholds and sensitivity comparable to psychophysical performance.     

Chromatic discrimination of natural objects

Studies of chromatic discrimination are typically based on homogeneously colored patches. Surfaces of natural objects, however, cannot be characterized by a single color. Instead, they have a chromatic texture, that is, a distribution of different chromaticities. Here we study chromatic discrimination for natural images and synthetic stimuli with a distribution of different chromaticities under various states of adaptation. Discrimination was measured at the adaptation point, where the mean chromaticity of the test stimuli was the same as the chromaticity of the adapting background, and away from the adaptation point. At the adaptation point, discrimination for natural objects resulted in threshold contours that were selectively elongated in a direction of color space matching the chromatic variation of the colors within the natural object. Similar effects occurred for synthetic stimuli. Away from the adaptation point, discrimination thresholds increased and threshold ellipses were elongated along the contrast axis connecting adapting color and test color. Away from the adaptation point, no significant differences between the different stimulus classes were found. The effect of the chromatic texture on discrimination seemed to be masked by the overall increase in discrimination thresholds. Our results show that discrimination of chromatic textures, either synthetic or natural, differs from that of simple uniform patches when the chromatic variation is centered at the adaptation point.     

The spatial tuning of color and luminance peripheral vision measured with notch filtered noise masking.

We have measured the spatial bandwidths of the bandpass red-green chromatic and luminance mechanisms at four locations in the nasal visual field (0, 10, 20 and 30 degrees) using a method of notch filtered noise masking which effectively removes the artifact of off-frequency looking for our stimuli. Detection thresholds were measured for luminance or isoluminant red-green Gaussian enveloped test gratings of 0.5 cpd embedded in 1/f noise. Firstly, thresholds were obtained as a function of increasing noise spectral density and were fitted using a standard noise masking model. These results support the existence across the visual field of independent, red-green chromatic and luminance mechanisms with similar sampling efficiencies. Secondly, we measured thresholds in notch filtered noise as a function of notch width and derived the spatial bandwidth of the detection mechanism. We find both color and luminance mechanisms have similar bandwidths which remain virtually constant across eccentricity. These results indicate strong overall similarities between the early processing of color and luminance vision, and lend support to the role of color as an 'intrinsic image' in spatial vision. The results are discussed in the light of the anchored channel and shifting channel models of peripheral contrast sensitivity and pattern detection.     

Failure of direction identification for briefly presented second-order motion stimuli: evidence for weak direction selectivity of the mechanisms encoding motion

We sought to investigate why the direction of second-order motion, unlike first-order motion, cannot be identified when the stimulus exposure duration is brief (<200 ms). In a series of experiments observers identified both the orientation (vertical or horizontal) and the direction (left, right, down or up) of a drifting sinusoidal modulation (0.93 c/ degrees ) in either the luminance (first order) or the contrast (second order) of a two-dimensional noise carrier. All motion stimuli were equated for visibility, and the duration was varied using the method of constant stimuli. Performance was measured for second-order motion over a range of drift temporal frequencies (0.63-5.04 Hz) and for first-order motion stimuli composed of two, opposite drifting modulations in luminance of unequal modulation depth. Orientation-identification performance was nearly 100% correct for both first-order and second-order motion stimuli, even at the briefest stimulus duration tested (26.49 ms). Direction identification for first-order motion was also typically good with brief presentations, but was poor for second-order motion when the exposure duration was < approximately 200 ms. Importantly increasing either the drift temporal frequency of second-order motion or the bidirectional nature of the first-order motion patterns produced comparable levels of performance for the two varieties of motion (i.e. the minimum duration required for reliable direction identification could be equated). As orientation-identification performance for the first-order and second-order motion stimuli was comparably good and minimally affected by duration, the marked differences on the direction-identification task must be specific to mechanisms that encode drift direction, rather than spatial structure. We propose that second-order motion detectors are much less selective for stimulus direction than first-order motion sensors, and thus are more susceptible to the deleterious effects of limiting stimulus duration (which introduces spurious motion in the opposite direction, particularly at low drift rates). Alternative explanations based on the delayed propagation of second-order motion signals or the temporal characteristics of the underlying motion mechanisms are not supported by our findings.     

The influence of spatial and temporal noise on the detection of first-order and second-order orientation and motion direction

Thresholds for identifying the direction of second-order motion (contrast-modulated dynamic noise) are consistently higher than those for identifying spatial orientation, unlike first-order gratings for which the two thresholds are typically the same. Two explanations of this phenomenon have been proposed: either first-order and second-order patterns are encoded by separate mechanisms with different properties, or dynamic noise selectively impairs ("masks") sensitivity to second-order motion direction but not orientation. The former predicts the two thresholds should remain distinct for second-order patterns, irrespective of the temporal structure (static vs. dynamic) of the noise carrier. The latter predicts direction thresholds should be higher than orientation thresholds, for both second-order and first-order motion patterns, when dynamic (but not static) noise is present. To resolve this issue we measured direction and orientation thresholds for first-order (luminance) and second-order (contrast or polarity) modulations of static or dynamic noise. Results were decisive: The two thresholds were invariably the same for first-order stimuli but markedly different (direction thresholds approximately 50% higher) for second-order stimuli, regardless of the temporal properties (static or dynamic) and the overall contrast of the noise, or the drift temporal frequency of the envelope. This suggests that first-order and second-order motion are encoded separately and that the mechanisms encoding second-order stimuli cannot determine direction at the absolute threshold for spatial form.     

The chromatic input to global motion perception

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.     

Visual Ageing: Unspecific Decline of the Responses to Luminance and Colour

We have investigated whether ageing affects selectively the responses to equiluminant patterns of pure colour contrast. In two groups of subjects (mean ages 29 and 72 yr) contrast thresholds were measured psychophysically for the detection and for the discrimination of the direction of motion of drifting gratings. The gratings were modulated either in pure luminance contrast (and uniform colour), or pure chromatic contrast (red-green equiluminant gratings). In subjects of the same age groups, visual evoked potentials (VEP) were recorded in response to gratings with either pure luminance contrast or pure colour contrast sinusoidally reversed in contrast at various temporal frequencies. It was shown that psychophysical contrast sensitivity for equiluminant patterns deteriorates significantly with age, and VEP latency increases. However, these effects of ageing on the responses to patterns of pure colour contrast are substantially the same as those observed in the same subjects for stimuli with pure luminance contrast. The results suggest that ageing causes a small and unspecific decline of the response of the visual system to luminance and colour contrast. Copyright © 1996 Elsevier Science Ltd.