Luminance Pathway Research Articles

Long-range suppressive interactions between S-cone and luminance channels

Surround suppression (SS) refers to a reduction in the effective stimulus contrast in one visual location produced by a stimulus presented in an adjacent location. This type of suppression is tuned for orientation and spatial frequency and is thought to be a cortical process. In this paper we used psychophysical measurements to determine whether S-cone-driven signals are affected by surround suppression and, if so, whether S-cone and achromatic signals interact at spatially-remote locations. Our results revealed three important aspects of surround suppression. Firstly, we show that S-cone probes are suppressed by simultaneous S-cone contrast surrounds and that this suppression has the characteristics of a cortical mechanism. Secondly, we show that when probes and surrounds are presented simultaneously, there are no suppressive interactions between S-cone and luminance stimuli. Finally, we demonstrate that this apparent independence is an artifact of signal timing: when the S-cone components of the stimuli precede the luminance components by approximately 40 ms, we find a significant interaction between the two pathways. The amplitude of this interaction depends critically upon the relative onset times of the two components. These results indicate that some component of surround suppression depends on neural computations that occur after the S- and luminance pathways are combined in striate cortex. In addition, the strong dependence of the magnitude of surround suppression on temporal ordering suggests that much of the effect is driven by transient signals.

Two distinct cone-opponent processes in the L+M luminance pathway

We measured phase shifts between Long-wavelength cone (L-cone) and Middle-wavelength cone (M-cone) signals as well as sensitivity in the luminance pathway either following a cone-silent substitution of colored background or on a steady colored background. In background substitution, the phase shifts between L- and M-cone signals varied only slightly depending on the substituted color, whereas marked elevation of the threshold following the substitution of colored background was found. In contrast, the phase shifts, as well as threshold, varied largely, depending on the background color in the steady background. These facts suggest that suppression by the cone-opponent process for background color substitution is different from the one for a steady colored background.

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.

Paradoxical shifts in human color sensitivity caused by constructive and destructive interference between signals from the same cone class

Paradoxical shifts in human color (spectral) sensitivity occur on deep-red (658 nm) background fields. As the radiance of the deep-red background is increased from low to moderate levels, the spectral sensitivity for detecting 15-Hz flicker shifts toward shorter wavelengths, although by more than is predicted by selective chromatic adaptation (e.g., Eisner & MacLeod, 1981; Stromeyer et al., 1987; Stockman et al., 1993). Remarkably, though, at higher background radiances, the spectral sensitivity then shifts precipitously back towards longer wavelengths. Here, we show that both effects are due in large part to destructive and constructive interference between signals generated by the same cone type. Contrary to the conventional model of the human visual system, the M- and L-cone types contribute not just the customary fast signals to the achromatic or luminance pathway, but also slower signals of the same or opposite sign. The predominant signs of the slow M- and L-cone signals change with background radiance, but always remain spectrally opposed (M-L or L-M). Consequently, when the slow and fast signals from one cone type destructively interfere, as they do near 15 Hz, those from the other cone type constructively interfere, causing the paradoxical shifts in spectral sensitivity. The shift in spectral sensitivity towards longer wavelengths is accentuated at higher temporal frequencies by a suppression of fast M-cone signals by deep-red fields.

Pupil Response to Color Signals in Cone-Contrast Space

Purpose: It is widely accepted that pupil responses to visual stimuli are determined by the ambient illuminance, and recently it has been shown that changes in stimulus color also contributes to a pupillary control mechanism. However, the role of pupillary responses to chromatic stimuli is not clear. The aim of this study was to investigate how color and luminance signals contribute to the pupillary control mechanism. Methods: We measured pupillary iso-response contours in M-and L-cone contrast space. The iso-response contours in cone-contrast space have been determined to examine what mechanisms contribute to the pupillary pathway. The shapes of the iso-response contour change when different mechanisms determine the response. Results: It was shown that for all subjects, the pupillary iso-response contours form an ellipse with positive slope in cone-contrast space, indicating that the sensitivities to the chromatic stimuli are higher than those for the luminance stimuli. The pupil responds maximally to a grating that has a stronger L-cone modulation than the red-green isoluminant grating. Conclusions: The sensitivity of the chromatic pathway, in terms of pupillary response, is three times larger than that of the luminance pathway, a property that might have utility in clinical applications.

Which Visual Pathways Cause Fixation-Related Inhibition?

Visual stimuli can both inhibit and activate motor mechanisms. In one well-known example, the latency of saccadic eye movements is prolonged in the presence of a fixation stimulus, relative to the case in which the fixation stimulus disappears before the target appears. This automatic sensory-motor effect, known as the gap effect or fixation-offset effect, has been associated with inhibitory connections within the superior colliculus (SC). Visual information is provided to the SC and other oculomotor areas, such as the frontal eye fields (FEF), mainly by the magnocellular geniculostriate pathway, and also by the retinotectal pathway. We tested whether signals in these pathways are necessary to create fixation-related inhibition, by using stimuli invisible to them. We found that such stimuli, visible only to short-wave-sensitive cones (S cones), do produce fixation-related inhibition (including when warning effects were equated). We also demonstrate that this fixation-related inhibition cannot be explained by residual activation of luminance pathways and must be caused by a route separate from that of luminance fixation signals. Thus there are at least two routes that cause fixation-related inhibition, and direct sensory input to the SC or FEF by the magnocellular or retinotectal pathways is not required. We discuss the implications that there may be both cortical and collicular mechanisms.

Long-wavelength adaptation reveals slow, spectrally opponent inputs to the human luminance pathway

In addition to its expected fast, additive L- and M-cone inputs (L + M), the luminance pathway has slow, spectrally opponent inputs. We have previously shown that on long-wavelength fields, the dominant slow signals change from L-M at moderate intensity levels to M-L signals at high. Here, we focus on the transition between them, which we find is marked by substantial changes in temporal phase delay, and by large and unexpected shifts in flicker spectral sensitivity. At moderate temporal frequencies, counter to the selective adaptation caused by the field, spectral sensitivity changes from being M-cone-like to more L-cone-like. These changes can be accounted for by a change in the relative strengths of the slow spectrally opponent cone signals from L-M exceeding M-L below the transition to M-L exceeding L-M above it, and by the resulting changes in constructive and destructive interference between the dominant signal components. We speculate that the transition is caused by the deep-red field becoming equivalent, postreceptorally, to a green field at high bleaching levels. These results further challenge the dogma that there are separable psychophysical channels for the transmission and processing of color and luminance information. Although its output generates an achromatic percept, the luminance channel has spectrally opponent inputs.

Spectrally opponent inputs to the human luminance pathway: slow +L and −M cone inputs revealed by low to moderate long‐wavelength adaptation

The luminance pathway has slow (s), spectrally opponent cone inputs in addition to the expected fast (f), non-opponent inputs. The nature of these inputs to luminance flicker perception was further explored psychophysically by measuring middle- (M-) and long-wavelength-sensitive (L-) cone modulation sensitivities, M- and L-cone phase delays, and flicker spectral sensitivities under three conditions of low to moderate long-wavelength adaptation. Under these conditions we find that the luminance channel has fast M- and L-cone input signals (+fM and +fL), and slow, spectrally opponent cone input signals (+sL and -sM). The slow signals found under these conditions are therefore of the opposite polarity to those (+sM and -sL) found under more intense long-wavelength adaptation. At these less intense levels, fast and slow M-cone signals of opposite polarity (-sM and +fM) cancel at low frequencies, but then constructively interfere at intermediate frequencies (ca 12.5-22.5 Hz, depending on adapting level) because of the delay between them. In contrast, fast and slow L-cone signals of the same polarity (+sL and +fL) sum at low frequencies, but then destructively interfere at intermediate frequencies. Importantly, the spectrally opponent signals (+sL and -sM) contribute to flicker nulls without producing visible colour variation. Although its output generates an achromatic percept, the luminance channel has slow spectrally opponent as well as fast non-opponent inputs.

Spectrally opponent inputs to the human luminance pathway: slow +M and −L cone inputs revealed by intense long‐wavelength adaptation

The nature of the inputs to achromatic luminance flicker perception was explored psychophysically by measuring middle- (M-) and long-wavelength-sensitive (L-) cone modulation sensitivities, M- and L-cone phase delays, and spectral sensitivities as a function of temporal frequency. Under intense long-wavelength adaptation, the existence of multiple luminance inputs was revealed by substantial frequency-dependent changes in all three types of measure. Fast (f) and slow (s) M-cone input signals of the same polarity (+sM and +fM) sum at low frequencies, but then destructively interfere near 16 Hz because of the delay between them. In contrast, fast and slow L-cone input signals of opposite polarity (-sL and +fL) cancel at low frequencies, but then constructively interfere near 16 Hz. Although these slow, spectrally opponent luminance inputs (+sM and -sL) would usually be characterized as chromatic, and the fast, non-opponent inputs (+fM and +fL) as achromatic, both contribute to flicker photometric nulls without producing visible colour variation. Although its output produces an achromatic percept, the luminance channel has slow, spectrally opponent inputs in addition to the expected non-opponent ones. Consequently, it is not possible in general to silence this channel with pairs of 'equiluminant' alternating stimuli, since stimuli equated for the non-opponent luminance mechanism (+fM and +fL) may still generate spectrally opponent signals (+sM and +sL).

Accommodation responses to stimuli in cone contrast space

The aim was to identify the cone contributions and pathways for reflex accommodation. Twelve illumination conditions were used to test specified locations in cone-contrast space. Accommodation was monitored continuously in a Badal optometer while the grating stimulus (2.2 c/d sine-wave; 0.27 modulation) moved sinusoidally (0.195 Hz) towards and away from the eye from a mean position of 2.00 D (±1.00 D). Mean accommodation level and dynamic gain and phase at 0.195 Hz were calculated. Mean accommodation level varied significantly when the long- and middle-wavelength cone contrast ratio was altered in both the luminance and chromatic quadrants of cone-contrast space. This experiment indicates that L- and M-cones contribute to luminance and chromatic signals that produce the accommodation response, most likely through magno-cellular and parvo-cellular pathways, respectively. The L:M cone weighting to the luminance pathway that mediates accommodation is 1.63:1. The amplitude and direction of the response depends on changes in chromatic contrast and luminance contrast signals that result from longitudinal chromatic aberration and defocus of the image.

Thin edges and the induced contrast asynchrony

The Induced Contrast Asynchrony is a visual effect that demonstrates a perceptual separation between luminance and contrast (Shapiro et al, in press). The basic version of the effect consists of two physically identical rectangular patches whose luminance levels are modulated at 2 Hz (.75×1.5 deg, 40 + 15*sin(tw) cd/m2); one patch is surrounded by a dark field (6×6 deg, 20 cd/m2) and the other by a light field (6×6 deg, 60 cd/m2). This produces a configuration in which the luminance of the two patches modulates in phase and the contrast modulates in antiphase. Observers perceive primarily the contrast signal (i.e., the patches modulate in antiphase), but at low frequencies, observers can also attend to the luminance signal. This year we present the effects of the edge widths on the appearance of the asynchronous contrast signal. We measure psychometric functions for the perceived contrast signal versus the width of the introduced lines over a range of patch sizes (width .15 to 1.2 deg). We show that 1) the contrast asynchrony can be produced when the surrounding field is of minimal width, indicating that the contrast signal arises primarily at the patch/surround edge; 2) placing a thin (.05 deg) white or black line adjacent to one side of the modulating rectangular patch suppresses perception of the asynchronous contrast signal; 3) a variety of stimulus configurations (e.g., circular patch with a two-tone surround or gradient bar adjacent to a modulating patch) can produce new versions of the contrast asynchrony; and 4) the combination of thin edges and single-patch modulation can produce a novel form of contrast-defined apparent position shifts. Most of the results can be explained by a model with separate contrast (i.e., rectified difference) and luminance pathways. A sophisticated spatiotemporal analysis is required to account for apparent changes in spatial position. The results offer new insights into static lightness illusions such as White's effect. R15-EY12946

An oscillatory correlation model of visual motion analysis.

We describe and evaluate a model of motion perception based on the integration of information from two parallel pathways: a motion pathway and a luminance pathway. The motion pathway has two stages. The first stage measures and pools local motion across the input animation sequence and assigns reliability indices to these pooled measurements. The second stage groups locations on the basis of these measurements. In the luminance pathway, the input scene is segmented into regions on the basis of similarities in luminance. In a subsequent integration stage, motion and luminance segments are combined to obtain the final estimates of object motion. The neural network architecture we employ is based on LEGION (locally excitatory globally inhibitory oscillator networks), a scheme for feature binding and region labeling based on oscillatory correlation. Many aspects of the model are implemented at the neural network level, whereas others are implemented at a more abstract level. We apply this model to the computation of moving, uniformly illuminated, two-dimensional surfaces that are either opaque or transparent. Model performance replicates a number of distinctive features of human motion perception.

Long-term maturation of visual pathways.

Previous research in adults has demonstrated the utility of the visual evoked potential (VEP) to measure the integrity of the chromatic and achromatic visual pathways. The VEP has also been shown to be a valuable indicator of maturation of these pathways in infants up to 1 year of age. The present manuscript reports changes in the visual pathways from 2 years to adulthood as measured by the spatio-chromatic VEP. The responses to achromatic reversal stimuli designed to preferentially activate the low spatial-frequency achromatic (luminance) pathways appear adult-like by 1 year of age. The responses to low spatial-frequency isoluminant onset stimuli designed to preferentially activate the chromatic pathway do not appear as they do in the adult until after 12-13 years of age. The shapes of the chromatic VEP waveforms shift from a positive-negative complex to a negative-positive complex. These changes can be modeled by a decrease in the latency of a large negative component between the ages of 1 year and adulthood. The results suggest that for low spatial-frequency stimuli, there are long-term changes in the development of the chromatic pathways that are not observed in the low spatial-frequency achromatic pathways. The changes in the chromatic VEP waveforms with age may be a physiological correlate of reported behavioral changes.

Luminance processing in apparent motion, Vernier offset and stereoscopic depth

We obtained (apparently) linear responses to luminance from three special displays of apparent motion, Vernier offset and stereoscopic depth. In our motion stimulus a dark and a light bar exchanged luminances repetitively on a grey surround. Motion was attributed to the bar that differed more from the surround, that is, on a dark surround the light bar appeared to jump, and on a light surround the dark bar appeared to jump. The apparent motion disappeared when the luminance of the surround lay halfway between that of the bars — on a linear, not a logarithmic scale. Similar results were obtained for special Vernier offset and stereo stimuli. These results cannot be explained if all luminances are processed within the same luminance pathway and that pathway transforms input luminance using non-linear compression. However, the apparent linearity of our results could arise from opposite and equal non-linearities cancelling out within separate ON- and OFF-spatial luminance pathways. A second set of experiments presented one bar separately into each eye on different surrounds (dichoptic presentation of competing apparent motion signals) or manipulated the display spatially so that different surrounds were associated with different bars (binocular presentation of competing Vernier targets). Results showed that apparent motion and Vernier signals of equal Weber contrast (normalisation of linear difference to surround luminance) evoked equal-motion and equal Vernier offset strengths. Given that motion and Vernier strength followed Weber’s law, we infer that the ON- and OFF-pathways transform luminance non-linearly. Our third experiment presents an example of a brightness bisection task in which we were able to influence the bisection steps, to follow either a linear or non-linear series. The benefits of parsing the visual scene so that visual information is processed within two opposite luminance pathways is discussed.

Selective cone suppression by the L-M- and M-L-cone-opponent mechanisms in the luminance pathway.

We investigated how transient changes of background color influence the L- and M- (long- and middle-wavelength-sensitive-) cone signals in the luminance pathway. Motion identification thresholds were measured for a drifting sinusoidal grating (1 cycle/deg) modulated along different vector directions in L- and M-cone contrast space. The color of a central 4-deg-diameter region was briefly altered (500 ms) by incrementing or decrementing either L- or M-cone excitation. Incrementing L-cone and decrementing M-cone excitation produced a field that appeared reddish relative to the yellow surround. Likewise, incrementing M-cone and decrementing L-cone produced a field that appeared greenish. Motion identification thresholds were obtained on the yellow field following the brief color transitions. The results show that the threshold for the L-cone direction was selectively elevated by the background substitution of incrementing L-cone and decrementing M-cone excitation (shift toward reddish color). The same substitution, however, did not affect the threshold in the M-cone direction. Similarly, the threshold for the M-cone direction was selectively elevated by the background substitution of incrementing M-cone, decrementing L-cone excitation (shift toward greenish) without affecting the threshold in the L-cone direction. Experiments using the motion quadrature paradigm confirmed that these effects occur within the luminance mechanisms. These results indicate that the activation of L-on plus M-off signals suppresses the L-cone signal and that the activation of L-off plus M-on signals suppresses the M-cone signals in the luminance pathway. We propose a retinal model based on the experimental results.

The effect of photon noise on the detection of white flashes.

Thresholds for detecting brief, white, foveal test flashes drop abruptly within 0.2 sec of the offset of a white adapting field. The magnitude of the abrupt drop is proportional to the square root of field intensity (square root of I) correct for bleaching and dark light. Thresholds are then stable out to 1.6 sec for 200 msec tests, or recover only slightly for 20 msec tests. These results exclude some simple deterministic models in which Weber-like gain controls in the luminance pathway are assumed to recover exponentially in the dark, but can be explained parsimoniously if turning off the field abolishes photon-driven noise, improving the S/N ratio while leaving visual responsivity virtually unaltered. This theory was first put forward by Krauskopf and Reeves [(1980) Vision Research, 20, 193-196] for S-cone thresholds; it implies that the Weber law for increment thresholds is not due to a single gain control, but rather expresses the product of two distinct square root of I factors, adjustment of responsivity and photon-driven noise. Removal of the noise, not recovery of gain, permits thresholds to fall in early dark adaptation.

The peripheral flicker effect: desensitization of the luminance pathway by static and modulated light.

Previous studies have shown that luminance flicker, presented peripheral to a foveal test target, increases thresholds for target detection: the peripheral flicker (PF) effect. These studies have also shown that thresholds are elevated more for luminance targets, relative to chromatic targets. In the present study we examined the specificity of the PF effect on the luminance mechanism and assessed the contribution of modulated stray-light to the test field, as well as longer range spatial interactions. We found that the presence of a foveal luminance pedestal, as well as PF, caused a notch to appear in the spectral sensitivity function around 570 nm. This result confirms the hypothesis that the PF effect decreases the sensitivity of the luminance pathway. To assess the contribution of stray-light to the PF effect, we modulated a luminance pedestal without the presence of PF in order to simulate the stray-light effect in isolation. A decrease in sensitivity for wavelengths around 570 nm occurred with modulated stray-light, suggesting that modulated stray-light contributes substantially to this effect. We then minimized the modulated stray-light by phase-reversing a checkerboard pattern in the periphery. A significant, though smaller, threshold elevation to mid-spectrum stimuli was obtained, suggesting that long range spatial effects are also active in the PF effect. We conclude that the PF effect causes a desensitization of foveal luminance pathways via local and more long range spatial interactions. Our results are consistent with previous data which suggest that the PF effect is due to selective adaptation of cells in the magnocellular pathway (M-cells). Our data imply that local network adaptation may be a property of the magnocellular pathway.

Colour tuning in human visual cortex measured with functional magnetic resonance imaging.

The primate retina contains three classes of cones, the L, M and S cones, which respond preferentially to long-, middle- and short-wavelength visible light, respectively. Colour appearance results from neural processing of these cone signals within the retina and the brain. Perceptual experiments have identified three types of neural pathways that represent colour: a red-green pathway that signals differences between L- and M-cone responses; a blue-yellow pathway that signals differences between S-cone responses and a sum of L- and M-cone responses; and a luminance pathway that signals a sum of L- and M-cone responses. It might be expected that there are neurons in the primary visual cortex with response properties that resemble these three perceptual pathways, but attempts to find them have led to inconsistent results. We have therefore used functional magnetic resonance imaging (fMRI) to examine responses in the human brain to a large number of colours. In visual cortical areas V1 and V2, the strongest response is to red-green stimuli, and much of this activity is from neurons receiving opposing inputs from L and M cones. A strong response is also seen with blue-yellow stimuli, and this response declines rapidly as the temporal frequency of the stimulus is increased. These responses resemble psychophysical measurements, suggesting that colour signals relevant for perception are encoded in a large population of neurons in areas V1 and V2.

Faster than the eye can see: blue cones respond to rapid flicker

Flickering lights that are detected by the blue cones of the human visual system fuse to yield a steady sensation at much lower rates of flicker than do lights that are detected by the red or green cones. Yet, although blue-cone-detected lights flickering at 30-40 Hz appear to be steady, they are still able to interact with red- or green-cone-detected flickering lights to produce clearly detectable beats in the form of an amplitude modulation of the red- or green-cone flicker. Thus the blue cones produce a viable high-frequency flicker signal, as do the red and green cones, but one that is normally lost before it reaches sensation. The temporal-frequency response for the blue-cone beat interaction is similar in shape to the temporal-frequency response for directly detected red- or green-cone flicker. When measured through the same pathway (which we identify as the luminance pathway, since it is able to transmit high-frequency flicker), the response of the blue cones seems to be as fast as that of the other cones.

輪郭線形状の両眼立体視形成への影響

We investigated cue interactions between binocular stereopsis and contour shapes for depth perception. We used random-dot stereograms of corrugated surfaces, where the upper and lower sides were shaped like sinusoidal waves to provide pictorial depth cues. The depth information of contour shapes was either coherent or conflictive to that of binocular disparity. Observation of the conflict stimulus showed that two different depths are never seen simultaneously. When the observing duration was longer than several tenths of a second, a stable surface provided by binocular stereopsis was observed. However, measuring the time required for perceiving stereoscopic depth revealed the influence of the contour shape on time. Coherent depth from the contour shape speeded up the processing of stereoscopic depth in comparison with the conflictive depth cue (experiment 1). Since the effect of pictorial cues on the stereoscopic depth was found even when the contour was defined only by texture differences (experiment 2), it seens that depth based on contour shape is not restricted to the luminance pathway.