Achromatic and chromatic perceived contrast are reduced in the visual periphery.

The loss of contrast sensitivity with eccentricity is well documented, and is steeper for higher spatial frequencies, and for L/M cone-opponent stimuli compared to achromatic or S-cone-opponent. Here, we ask how perceived contrast depends on eccentricity when stimuli are presented at suprathreshold contrasts, and test two opposing predictions. Contrast constancy predicts no loss in perceived contrast across the visual field regardless of changes in detection threshold - appearance depends only on physical contrast. Conversely, perceived contrast may be scaled in the same way as detection threshold, reflecting the proportional increase in stimulus contrast above threshold. We measured perceived contrast for L/M cone-opponent, S-cone opponent, and Ach stimuli up to 18 degrees of eccentricity using a 2AFC contrast matching method between fovea and periphery. We tested a range of reference contrasts from low (close to detection threshold) to high suprathreshold contrasts and we relate suprathreshold perceived contrast to measured detection thresholds. We find evidence for a hybrid model in which apparent contrast is reduced with eccentricity for stimuli in the low and mid contrast range, with contrast constancy only attained at the highest contrasts. When equated for similar sensitivity losses, we find no difference between chromatic and Ach contrast responses.

Magnetoencephalography contrast adaptation reflects perceptual adaptation.

Contrast adaptation is a fundamental visual process that has been extensively investigated and used to infer the selectivity of visual cortex. We recently reported an apparent disconnect between the effects of contrast adaptation on perception and functional magnetic resonance imaging BOLD response adaptation, in which adaptation between chromatic and achromatic stimuli measured psychophysically showed greater selectivity than adaptation measured using BOLD signals. Here we used magnetoencephalography (MEG) recordings of neural responses to the same chromatic and achromatic adaptation conditions to characterize the neural effects of contrast adaptation and to determine whether BOLD adaptation or MEG better reflect the measured perceptual effects. Participants viewed achromatic, L-M isolating, or S-cone isolating radial sinusoids before adaptation and after adaptation to each of the three contrast directions. We measured adaptation-related changes in the neural response to a range of stimulus contrast amplitudes using two measures of the MEG response: the overall response amplitude, and a novel time-resolved measure of the contrast response function, derived from a classification analysis combined with multidimensional scaling. Within-stimulus adaptation effects on the contrast response functions in each case showed a pattern of contrast-gain or a combination of contrast-gain and response-gain effects. Cross-stimulus adaptation conditions showed that adaptation effects were highly stimulus selective across early, ventral, and dorsal visual cortical areas, consistent with the perceptual effects.

Linking perceived to physical contrast: Comparing results from discrimination and difference-scaling experiments.

Psychophysical approaches that allow us to estimate how perceived stimulus intensity is linked to physical intensity are import tools for studying nonlinear transformations of visual signals within different visual pathways. Here, we investigated how stimulus contrast is encoded in achromatic and chromatic pathways using simple grating stimuli. We compared two experimental approaches to this question: contrast discrimination (increment detection thresholds measured on contrast pedestals) and the maximum likelihood difference scaling (MLDS) approach introduced by Maloney and Yang (2003). The results of both experiments are expressed using simple models that include a transducer function mapping physical contrast to an internal signal the observer uses in making judgments, and an estimate of the variability of this representation (internal "noise"). We found that the transducers derived from both experiments have a similar form, but occupy different ranges of physical contrast in different stimulus conditions, reflecting difference in contrast sensitivity. This is consistent with past discrimination results, and in the difference-scaling case provides new evidence supporting the idea that suprathreshold chromatic and achromatic contrast are processed similarly, once differences in contrast sensitivity are taken into account. Model estimates of internal noise were higher in the difference-scaling experiment than the discrimination experiment, a finding we attribute to a difference in task complexity. Finally, we fit an alternative version of the MLDS model in which internal noise increased with response level. This alternative was no better at predicting holdout data in a cross-validation analysis than the original constant-variance model.

Shifting eye balance using monocularly directed attention in normal vision.

In binocular vision, even without conscious awareness of eye of origin, attention can be selectively biased toward one eye by presenting a visual stimulus uniquely to that eye. Monocularly directed visual cues can bias perceptual dominance, as shown by studies using discrete measures of percept changes in continuous-flash suppression. Here, we use binocular rivalry to determine whether eye-based visual cues can modulate eye balance using continuous percept reporting. Using a dual-task versus single-task paradigm, we investigated whether the attentional load of these cues differentially modulates eye balance. Furthermore, both color-based and motion-based cue stimuli, non-overlaid and peripheral to the rivalry grating stimuli, were used to determine whether shifts in eye balance were stimulus specific. Aligned to cue stimulus onset, time series of percept reports were constructed and averaged across trials and participants. Specifically, for the monocular attention conditions, we found a significant shift in eye balance toward the cued eye and a significant difference in the time taken to switch from the dominating percept, regardless of whether the attention stimuli is color based or motion based. Although we did not find a significant main effect of attentional load, we found a significant interaction effect between the attentionally cued eye and attentional load on the shift in eye balance, indicating an influence of monocular attention on the shift in eye balance.

Attention selectively enhances stimulus information for surround over foveal stimulus representations in occipital cortex.

By attending to part of a visual scene, we can prioritize processing of the most relevant visual information and so use our limited resources effectively. Previous functional magnetic resonance imaging (fMRI) work has shown that attention can increase overall blood-oxygen-level-dependent (BOLD) signal responsiveness but also enhances the stimulus information in terms of classifier performance. Here, we investigate how these effects vary across the visual field. We compare attention-enhanced fMRI-BOLD amplitude responses and classifier accuracy in fovea and surrounding stimulus regions using a set of four simple stimuli subdivided into a foveal region (1.4° diameter) and a surround region (15° diameter). We found dissociations between the effects of attention on average response and in enhancing stimulus information. In early visual cortex, we found that attention increased the amplitude of responses to both foveal and surround parts of the stimuli and increased classifier performance only for the surround stimulus. Conversely, ventral visual areas showed less change in average response but greater changes in decoding. Unlike for early visual cortex, in the ventral visual cortex attention produced similar changes in decoding for center and surround stimuli.

fMRI representational similarity analysis reveals graded preferences for chromatic and achromatic stimulus contrast across human visual cortex.

Human visual cortex is partitioned into different functional areas that, from lower to higher, become increasingly selective and responsive to complex feature dimensions. Here we use a Representational Similarity Analysis (RSA) of fMRI-BOLD signals to make quantitative comparisons across LGN and multiple visual areas of the low-level stimulus information encoded in the patterns of voxel responses. Our stimulus set was picked to target the four functionally distinct subcortical channels that input visual cortex from the LGN: two achromatic sinewave stimuli that favor the responses of the high-temporal magnocellular and high-spatial parvocellular pathways, respectively, and two chromatic stimuli isolating the L/M-cone opponent and S-cone opponent pathways, respectively. Each stimulus type had three spatial extents to sample both foveal and para-central visual field. With the RSA, we compare quantitatively the response specializations for individual stimuli and combinations of stimuli in each area and how these change across visual cortex. First, our results replicate the known response preferences for motion/flicker in the dorsal visual areas. In addition, we identify two distinct gradients along the ventral visual stream. In the early visual areas (V1-V3), the strongest differential representation is for the achromatic high spatial frequency stimuli, suitable for form vision, and a very weak differentiation of chromatic versus achromatic contrast. Emerging in ventral occipital areas (V4, VO1 and VO2), however, is an increasingly strong separation of the responses to chromatic versus achromatic contrast and a decline in the high spatial frequency representation. These gradients provide new insight into how visual information is transformed across the visual cortex.

Selective Colour Vision Deficits in Multiple Sclerosis at Different Temporal Stages.

Multiple sclerosis (MS) without optic neuritis causes color-vision deficit but the evidence for selective color deficits in parvocellular-Red/Green (PC-RG) and koniocellular-Blue/Yellow (KC-BY) pathways is inconclusive. We investigated selective color-vision deficits at different MS stages. Thirty-one MS and twenty normal participants were tested for achromatic, red-green and blue-yellow sinewave-gratings (0.5 and 2 cycles-per-degree (cpd)) contrast orientation discrimination threshold. Red-green mean threshold at 0.5cpd in established-MS and blue-yellow mean threshold in all MS participants were abnormal. These findings show blue-yellow versus red-green color test is useful in differentiating MS chronicity, which helps to better understand the mechanism of colour-vision involvement in MS.

Enhanced luminance sensitivity on color and luminance pedestals: Threshold measurements and a model of parvocellular luminance processing.

Psychophysical interactions between chromatic and achromatic stimuli may inform our understanding of the cortical processing of signals of parvocellular origin, which carry both luminance and color information. We measured observers' sensitivity in discriminating the luminance of circular patch stimuli with a range of baseline ("pedestal") luminance and chromaticity. Pedestal stimuli were defined along vectors in cone-contrast space in a plane spanned by the red-green cone-opponent (L-M) and achromatic (L + M + S) axes. For a range of pedestal directions and intensities within this plane, we measured thresholds for discriminating increments from decrements along the achromatic axis. Low-contrast pedestals lowered luminance thresholds for every pedestal type. Thresholds began to increase with higher pedestal contrasts, forming a "dipper"-shaped function. Dipper functions varied systematically with pedestal chromaticity: Compared to the achromatic case, chromatic pedestals were effective at lower contrast. We suggest that the enhancement of luminance sensitivity caused by both achromatic and chromatic pedestals stems from a single mechanism, which is maximally sensitive to chromatic stimuli. We fit our data with a computational model of such a mechanism, in which luminance is computed from the rectified output of cone-opponent mechanisms similar to parvocellular neurons.

Color contrast adaptation: fMRI fails to predict behavioral adaptation.

fMRI-adaptation is a valuable tool for inferring the selectivity of neural responses. Here we use it in human color vision to test the selectivity of responses to S-cone opponent (blue-yellow), L/M-cone opponent (red-green), and achromatic (Ach) contrast across nine regions of interest in visual cortex. We measure psychophysical adaptation, using comparable stimuli to the fMRI-adaptation, and find significant selective adaptation for all three stimulus types, implying separable visual responses to each. For fMRI-adaptation, we find robust adaptation but, surprisingly, much less selectivity due to high levels of cross-stimulus adaptation in all conditions. For all BY and Ach test/adaptor pairs, selectivity is absent across all ROIs. For RG/Ach stimulus pairs, this paradigm has previously shown selectivity for RG in ventral areas and for Ach in dorsal areas. For chromatic stimulus pairs (RG/BY), we find a trend for selectivity in ventral areas. In conclusion, we find an overall lack of correspondence between BOLD and behavioral adaptation suggesting they reflect different aspects of the underlying neural processes. For example, raised cross-stimulus adaptation in fMRI may reflect adaptation of the broadly-tuned normalization pool. Finally, we also identify a longer-timescale adaptation (1h) in both BOLD and behavioral data. This is greater for chromatic than achromatic contrast. The longer-timescale BOLD effect was more evident in the higher ventral areas than in V1, consistent with increasing windows of temporal integration for higher-order areas.

Reevaluating hMT+ and hV4 functional specialization for motion and static contrast using fMRI-guided repetitive transcranial magnetic stimulation.

Although visual areas hMT+ and hV4 are considered to have segregated functions for the processing of motion and form within dorsal and ventral streams, respectively, more recent evidence favors some functional overlap. Here we use fMRI-guided online repetitive transcranial magnetic stimulation (rTMS) to test two associated hypotheses: that area hV4 is causally involved in the perception of motion and hMT+ in the perception of static form. We use variations of a common global stimulus to test two dynamic motion-based tasks and two static form-based tasks in ipsilateral and contralateral visual fields. We find that rTMS to both hMT+ and hV4 significantly impairs direction discrimination and causes a perceptual slowing of motion, implicating hV4 in motion perception. Stimulation of hMT+ impairs motion in both visual fields, implying that disruption to one hMT+ disrupts the other with both needed for optimal performance. For the second hypothesis, we find the novel result that hV4 stimulation markedly reduces perceived contrast of a static stimulus. hMT+ stimulation also produces an effect, implicating it in static contrast perception. Our findings are the first to show that rTMS of hV4 can produce a large perceptual effect and, taken together, suggest a less rigid functional segregation between hMT+ and hV4 than previously thought.

Evidence for chromatic edge detectors in human vision using classification images.

Edge detection plays an important role in human vision, and although it is clear that there are luminance edge detectors, it is not known whether there are chromatic edge detectors as well. We showed observers a horizontal edge blurred by a Gaussian filter (with widths of σ = 0.1125, 0.225, or 0.45°) embedded in blurred Brown noise. Observers had to choose which of two stimuli contained the edge. Brown noise was used in preference to white noise to reveal localized edge detectors. Edges and noise were defined by either luminance or chromatic contrast (isoluminant L/M and S-cone opponent). Classification image analysis was applied to observer responses. In this analysis, the random components of the stimulus are correlated with observer responses to reveal a template that shows how observers weighted different parts of the stimulus to arrive at their decision. We found classification images for both luminance and isoluminant chromatic stimuli that had shapes very similar to derivatives of Gaussian filters. The widths of these classification images tracked the widths of the edges, but the chromatic edge classification images were wider than the luminance ones. These results are consistent with edge detection filters sensitive to luminance contrast and isoluminant chromatic contrast.

Chromatic and achromatic monocular deprivation produce separable changes of eye dominance in adults.

Temporarily depriving one eye of its input, in whole or in part, results in a transient shift in eye dominance in human adults, with the patched eye becoming stronger and the unpatched eye weaker. However, little is known about the role of colour contrast in these behavioural changes. Here, we first show that the changes in eye dominance and contrast sensitivity induced by monocular eye patching affect colour and achromatic contrast sensitivity equally. We next use dichoptic movies, customized and filtered to stimulate the two eyes differentially. We show that a strong imbalance in achromatic contrast between the eyes, with no colour content, also produces similar, unselective shifts in eye dominance for both colour and achromatic contrast sensitivity. Interestingly, if this achromatic imbalance is paired with similar colour contrast in both eyes, the shift in eye dominance is selective, affecting achromatic but not chromatic contrast sensitivity and revealing a dissociation in eye dominance for colour and achromatic image content. On the other hand, a strong imbalance in chromatic contrast between the eyes, with no achromatic content, produces small, unselective changes in eye dominance, but if paired with similar achromatic contrast in both eyes, no changes occur. We conclude that perceptual changes in eye dominance are strongly driven by interocular imbalances in achromatic contrast, with colour contrast having a significant counter balancing effect. In the short term, eyes can have different dominances for achromatic and chromatic contrast, suggesting separate pathways at the site of these neuroplastic changes.

A Normative Data Set for the Clinical Assessment of Achromatic and Chromatic Contrast Sensitivity Using a qCSF Approach.

Purpose: The measurement of achromatic sensitivity has been an important tool for monitoring subtle changes in vision as the result of disease or response to therapy. In this study, we aimed to provide a normative data set for achromatic and chromatic contrast sensitivity functions within a common cone contrast space using an abbreviated measurement approach suitable for clinical practice. In addition, we aimed to provide comparisons of achromatic and chromatic binocular summation across spatial frequency. Methods: We estimated monocular cone contrast sensitivity functions (CCSFs) using a quick Contrast Sensitivity Function (qCSF) approach for achromatic as well as isoluminant, L/M cone opponent, and S cone opponent stimuli in a healthy population of 51 subjects. We determined the binocular CCSFs for achromatic and chromatic vision to evaluate the degree of binocular summation across spatial frequency for these three different mechanisms in a subset of 20 subjects. Results: Each data set shows consistent contrast sensitivity across the population. They highlight the extremely high cone contrast sensitivity of L/M cone opponency compared with the S-cone and achromatic responses. We also find that the two chromatic sensitivities are correlated across the healthy population. In addition, binocular summation for all mechanisms depends strongly on stimulus spatial frequency. Conclusions: This study, using an approach well suited to the clinic, is the first to provide a comparative normative data set for the chromatic and achromatic contrast sensitivity functions, yielding quantitative comparisons of achromatic, L/M cone opponent, and S cone opponent chromatic sensitivities as a function of spatial frequency.

The Whole is Other Than the Sum: Perceived Contrast Summation Within Color and Luminance Plaids.

The apparent contrast of a plaid is a reflection of the neural relationship between the responses to its two orthogonal component gratings. To investigate the perceived contrast summation of the responses to component gratings in plaids, we compared the apparent contrasts of monocular plaids to a component grating presented alone across chromaticity and spatial frequency. Observers performed a contrast-matching task for red-green color and luminance stimuli at low- and medium-spatial frequencies. Using the measured points of subjective equality between plaids and gratings, we evaluate perceived contrast summation across conditions, which may vary between 1 (no summation) and 2 (full summation). We show that achromatic plaids have higher perceived contrast summation than chromatic plaids. The greatest difference occurs at the medium-spatial frequency, with summation highest for achromatic plaids (1.87) and lowest for chromatic plaids (1.49), while at low-spatial frequencies, there is a smaller summation difference between achromatic (1.72) and chromatic (1.65) plaids. These results are consistent with recent theories of distinct cross-orientation suppression and summation mechanisms in color and luminance vision. Two control experiments for binocular versus monocular viewing, and the overall size of the stimulus patches did not reveal any differences from our main results.

Effect of overlaid luminance contrast on perceived color contrast: Shadows enhance, borders suppress.

Natural scenes contain both color and luminance variations at different sizes and orientations that are sometimes spatially overlaid and sometimes not. Here, we explore visual interactions between overlaid color and luminance contrast that are both suprathreshold and highly visible. We used a color-luminance plaid in which the perception of the color contrast and luminance contrast components were measured separately using a method of constant stimuli, to reveal how overlaid cross-oriented luminance contrast affects perceived color contrast, and how color contrast affects perceived luminance contrast. Binocular, monocular, and dichoptic viewing conditions were used for different spatial frequencies (0.375-1.5 cpd, 2 Hz) and base contrasts. We find that overlaid, cross-oriented luminance contrast enhances perceived color contrast by an average of 32% (monocularly and binocularly) across a wide range of luminance contrasts, but interocularly suppresses color contrast. For the reverse condition, we found no effect of color contrast on perceived luminance contrast. If, however, the cross-oriented arrangement is changed to co-oriented, specifically with the color and luminance borders aligned and in-phase, the color enhancement disappears and becomes mild suppression. Likewise, if the phase of the co-aligned components is varied, color enhancement returns once the color and luminance borders are misaligned and out of phase. Thus the relative position of the color and luminance borders is a crucial factor in determining the type of interaction, with color suppression occurring when the luminance and color borders coincide, as when demarcating an object boundary, and color enhancement when they do not coincide, as occurs in shadows and shading.

Orientation tuning of binocular summation: a comparison of colour to achromatic contrast.

A key function of the primary visual cortex is to combine the input from the two eyes into a unified binocular percept. At low, near threshold, contrasts a process of summation occurs if the visual inputs from the two eyes are similar. Here we measure the orientation tuning of binocular summation for chromatic and equivalent achromatic contrast. We derive estimates of orientation tuning by measuring binocular summation as a function of the orientation difference between two sinusoidal gratings presented dichoptically to different eyes. We then use a model to estimate the orientation bandwidth of the neural detectors underlying the binocular combination. We find that orientation bandwidths are similar for chromatic and achromatic stimuli at both low (0.375 c/deg) and mid (1.5 c/deg) spatial frequencies, with an overall average of 29 ± 3 degs (HWHH, s.e.m). This effect occurs despite the overall greater binocular summation found for the low spatial frequency chromatic stimuli. These results suggest that similar, oriented processes underlie both chromatic and achromatic binocular contrast combination. The non-oriented detection process found in colour vision at low spatial frequencies under monocular viewing is not evident at the binocular combination stage.

Color responses and their adaptation in human superior colliculus and lateral geniculate nucleus.

We use an fMRI adaptation paradigm to explore the selectivity of human responses in the lateral geniculate nucleus (LGN) and superior colliculus (SC) to red-green color and achromatic contrast. We measured responses to red-green (RG) and achromatic (ACH) high contrast sinewave counter-phasing rings with and without adaptation, within a block design. The signal for the RG test stimulus was reduced following both RG and ACH adaptation, whereas the signal for the ACH test was unaffected by either adaptor. These results provide compelling evidence that the human LGN and SC have significant capacity for color adaptation. Since in the LGN red-green responses are mediated by P cells, these findings are in contrast to earlier neurophysiological data from non-human primates that have shown weak or no contrast adaptation in the P pathway. Cross-adaptation of the red-green color response by achromatic contrast suggests unselective response adaptation and points to a dual role for P cells in responding to both color and achromatic contrast. We further show that subcortical adaptation is not restricted to the geniculostriate system, but is also present in the superior colliculus (SC), an oculomotor region that until recently, has been thought to be color-blind. Our data show that the human SC not only responds to red-green color contrast, but like the LGN, shows reliable but unselective adaptation.

The dynamics of cross-orientation masking at monocular and interocular sites.

We investigated the temporal properties of monocular and dichoptic cross-orientation masking (XOM) mediating suppressive or facilitatory cross-channel interactions between the neural detectors for the test and orthogonal mask stimuli. We measured the evolution of masking as a function of the duration of the test and mask stimuli to determine its time constant, and determined its dependence on stimulus onset asynchrony (SOA), for three contrast combinations: color-only (red-green color test and mask), luminance-only (luminance test and mask) and color-luminance (color test and luminance mask). Results show that the temporal properties of monocular and dichoptic masking differ markedly from each other and across contrast type. For the color-only condition, the dichoptic suppressive interaction is significantly longer than for the monocular one and both are largely independent of SOA. For the luminance-only condition, the suppressive interactions in both presentations are faster than for color, have similar time constants, but have different dependencies on SOA. For the color-luminance condition under the monocular conditions, cross-orientation facilitation (XOF) occurs with the luminance mask speeding up the processing of the color test with greatest XOF when the luminance mask precedes the color test by around 22 ms. No significant effects are observed for the dichoptic condition. Effects are invariant across spatial frequency. These strongly differential dynamic effects suggest that there is separate encoding of color contrast, luminance contrast, and their combination at the relatively early within-eye stage of processing, which is distinct from the dichoptic site.

The selectivity of responses to red-green colour and achromatic contrast in the human visual cortex: an fMRI adaptation study.

There is controversy as to how responses to colour in the human brain are organized within the visual pathways. A key issue is whether there are modular pathways that respond selectively to colour or whether there are common neural substrates for both colour and achromatic (Ach) contrast. We used functional magnetic resonance imaging (fMRI) adaptation to investigate the responses of early and extrastriate visual areas to colour and Ach contrast. High-contrast red-green (RG) and Ach sinewave rings (0.5 cycles/degree, 2 Hz) were used as both adapting stimuli and test stimuli in a block design. We found robust adaptation to RG or Ach contrast in all visual areas. Cross-adaptation between RG and Ach contrast occurred in all areas indicating the presence of integrated, colour and Ach responses. Notably, we revealed contrasting trends for the two test stimuli. For the RG test, unselective processing (robust adaptation to both RG and Ach contrast) was most evident in the early visual areas (V1 and V2), but selective responses, revealed as greater adaptation between the same stimuli than cross-adaptation between different stimuli, emerged in the ventral cortex, in V4 and VO in particular. For the Ach test, unselective responses were again most evident in early visual areas but Ach selectivity emerged in the dorsal cortex (V3a and hMT+). Our findings support a strong presence of integrated mechanisms for colour and Ach contrast across the visual hierarchy, with a progression towards selective processing in extrastriate visual areas.

The role of human brain area hMT+ in the perception of global motion investigated with repetitive transcranial magnetic stimulation (rTMS).

BACKGROUND: Psychophysical evidence suggests that the perception of the motion and color of moving stimuli are determined separately in the human brain. Here we aim to determine the role of visual cortical areas hMT+ and V1/V2 in each task by measuring the effect of rTMS of each area using an off-line continuous theta-burst stimulation (cTBS) protocol. METHODS: In the motion task, the direction of moving dots was identified using a global motion stimulus that avoids tracking, and in the detection task for the same stimulus, the presence of the dots was detected regardless of motion. Performance was measured using forced-choice methods in 8 subjects, both before and at 4 time-intervals in the 1-hour after brain stimulation. All experiments were done using achromatic and isoluminant, red-green chromatic stimuli. RESULTS: Performance on global motion for both achromatic and chromatic stimuli was significantly impaired following cTBS of visual area hMT+, with a maximum effect occurring 11 min after stimulation. In comparison, there was no effect of cTBS on the motion task for areas V1/V2 or the vertex (control). cTBS did not affect the detection task in either area. CONCLUSIONS: Our experiments validate the use of cTBS as an advantageous off-line rTMS protocol for studying visual areas. The results indicate a causal link between neural activity in area hMT+ and perception of motion of isoluminant chromatic stimuli. We conclude that area hMT+ is part of a common pathway processing the global motion of chromatic and achromatic stimuli, but is not involved in their detection.