A common rule for integration and suppression of luminance contrast across eyes, space, time, and pattern

Visual perception begins by dissecting the retinal image into millions of small patches for local analyses by local receptive fields. However, image structures extend well beyond these receptive fields and so further processes must be involved in sewing the image fragments back together to derive representations of higher order (more global) structures. To investigate the integration process, we also need to understand the opposite process of suppression. To investigate both processes together, we measured triplets of dipper functions for targets and pedestals involving interdigitated stimulus pairs (A, B). Previous work has shown that summation and suppression operate over the full contrast range for the domains of ocularity and space. Here, we extend that work to include orientation and time domains. Temporal stimuli were 15-Hz counter-phase sine-wave gratings, where A and B were the positive and negative phases of the oscillation, respectively. For orientation, we used orthogonally oriented contrast patches (A, B) whose sum was an isotropic difference of Gaussians. Results from all four domains could be understood within a common framework in which summation operates separately within the numerator and denominator of a contrast gain control equation. This simple arrangement of summation and counter-suppression achieves integration of various stimulus attributes without distorting the underlying contrast code.

Induced gamma activity in primary visual cortex is related to luminance and not color contrast: an MEG study

Gamma activity in the visual cortex has been reported in numerous EEG studies of coherent and illusory figures. A dominant theme of many such findings has been that temporal synchronization in the gamma band in response to these identifiable percepts is related to perceptual binding of the common features of the stimulus. In two recent studies using magnetoencephalography (MEG) and the beamformer analysis technique, we have shown that the magnitude of induced gamma activity in visual cortex is dependent upon independent stimulus features such as spatial frequency and contrast. In particular, we showed that induced gamma activity is maximal in response to gratings of 3 cycles per degree (3 cpd) of high luminance contrast. In this work, we set out to examine stimulus contrast further by using isoluminant red/green gratings that possess color but not luminance contrast using the same cohort of subjects. We found no induced gamma activity in V1 or visual cortex in response to the isoluminant gratings in these subjects who had previously shown strong induced gamma activity in V1 for luminance contrast gratings.

Binocular contrast vision at and above threshold

A fundamental problem for any visual system with binocular overlap is the combination of information from the two eyes. Electrophysiology shows that binocular integration of luminance contrast occurs early in visual cortex, but a specific systems architecture has not been established for human vision. Here, we address this by performing binocular summation and monocular, binocular, and dichoptic masking experiments for horizontal 1 cycle per degree test and masking gratings. These data reject three previously published proposals, each of which predict too little binocular summation and insufficient dichoptic facilitation. However, a simple development of one of the rejected models (the twin summation model) and a completely new model (the two-stage model) provide very good fits to the data. Two features common to both models are gently accelerating (almost linear) contrast transduction prior to binocular summation and suppressive ocular interactions that contribute to contrast gain control. With all model parameters fixed, both models correctly predict (1) systematic variation in psychometric slopes, (2) dichoptic contrast matching, and (3) high levels of binocular summation for various levels of binocular pedestal contrast. A review of evidence from elsewhere leads us to favor the two-stage model. © 2006 ARVO.

The interaction of luminance and texture amplitude in surface depth perception

Previous studies have suggested separate channels for detection of first-order luminance modulations (LM) and second-order modulations of the local amplitude (AM) of a texture. Mixtures of LM and AM with different phase relationships appear very different: in-phase compounds (LM + AM) look like 3-D corrugated surfaces, while out-of-phase compounds (LM - AM) appear flat and/or transparent. This difference may arise because the in-phase compounds are consistent with multiplicative shading, while the out-of-phase compounds are not. We investigated the role of these modulation components in surface depth perception. We used a textured background with thin bars formed by local changes in luminance and/or texture amplitude. These stimuli appear as embossed surfaces with wide and narrow regions. Keeping the AM modulation depth fixed at a suprathreshold level, we determined the amount of luminance contrast required for observers to correctly indicate the width (narrow or wide) of 'raised' regions in the display. Performance (compared to the LM-only case) was facilitated by the presence of AM, but, unexpectedly, performance for LM - AM was as good as for LM + AM. Thus, these results suggest that there is an interaction between first-order and second-order mechanisms during depth perception based on shading cues, but the phase dependence is not yet understood.

The interaction of luminance and texture amplitude in depth perception

Previous studies have suggested separate channels for the detection of first-order luminance (LM) and second-order modulations of the local amplitude (AM) of a texture (Schofield and Georgeson, 1999 Vision Research 39 2697 - 2716; Georgeson and Schofield, 2002 Spatial Vision 16 59). It has also been shown that LM and AM mixtures with different phase relationships are easily separated in identification tasks, and (informally) appear very different with the in-phase compound (LM + AM), producing the most realistic depth percept. We investigated the role of these LM and AM components in depth perception. Stimuli consisted of a noise texture background with thin bars formed as local increments or decrements in luminance and/or noise amplitude. These stimuli appear as embossed surfaces with wide and narrow regions. When luminance and amplitude changes have the same sign and magnitude (LM + AM) the overall modulation is consistent with multiplicative shading, but this is not so when the two modulations have opposite sign (LM - AM). Keeping the AM modulation depth fixed at a suprathreshold level, we determined the amount of luminance contrast required for observers to correctly indicate the width (narrow or wide) of raised regions in the display. Performance (compared to the LM-only case) was facilitated by the presence of AM, but, unexpectedly, performance for LM - AM was even better than for LM + AM. Further tests suggested that this improvement in performance is not due to an increase in the detectability of luminance in the compound stimuli. Thus, contrary to previous findings, these results suggest the possibility of interaction between first-order and second-order mechanisms in depth perception.

Motion sharpening and contrast:Gain control precedes compressive non-linearity?

Blurred edges appear sharper in motion than when they are stationary. We (Vision Research 38 (1998) 2108) have previously shown how such distortions in perceived edge blur may be accounted for by a model which assumes that luminance contrast is encoded by a local contrast transducer whose response becomes progressively more compressive as speed increases. If the form of the transducer is fixed (independent of contrast) for a given speed, then a strong prediction of the model is that motion sharpening should increase with increasing contrast. We measured the sharpening of periodic patterns over a large range of contrasts, blur widths and speeds. The results indicate that whilst sharpening increases with speed it is practically invariant with contrast. The contrast invariance of motion sharpening is not explained by an early, static compressive non-linearity alone. However, several alternative explanations are also inconsistent with these results. We show that if a dynamic contrast gain control precedes the static non-linear transducer then motion sharpening, its speed dependence, and its invariance with contrast, can be predicted with reasonable accuracy. © 2003 Elsevier Science Ltd. All rights reserved.

A model for motion sharpening: contrast gain control precedes compressive nonlinearity

Blurred edges appear sharper in motion than when they are stationary. We have previously shown how such distortions in perceived edge blur may be explained by a model which assumes that luminance contrast is encoded by a local contrast transducer whose response becomes progressively more compressive as speed increases. To test this model further, we measured the sharpening of drifting, periodic patterns over a large range of contrasts, blur widths, and speeds Human Vision. The results indicate that, while sharpening increased with speed, it was practically invariant with contrast. This contrast invariance cannot be explained by a fixed compressive nonlinearity since that predicts almost no sharpening at low contrasts.We show by computational modelling of spatiotemporal responses that, if a dynamic contrast gain control precedes the static nonlinear transducer, then motion sharpening, its speed dependence, and its invariance with contrast can be predicted with reasonable accuracy.

Grating and plaid masks indicate linear summation in a contrast gain pool

In human vision, the response to luminance contrast at each small region in the image is controlled by a more global process where suppressive signals are pooled over spatial frequency and orientation bands. But what rules govern summation among stimulus components within the suppressive pool? We addressed this question by extending a pedestal plus pattern mask paradigm to use a stimulus with up to three mask components: a vertical 1 c/deg pedestal, plus pattern masks made from either a grating (orientation = -45°) or a plaid (orientation = ±45°), with component spatial frequency of 3 c/deg. The overall contrast of both types of pattern mask was fixed at 20% (i.e., plaid component contrasts were 10%). We found that both of these masks transformed conventional dipper functions (threshold vs. pedestal contrast with no pattern mask) in exactly the same way: The dipper region was raised and shifted to the right, but the dipper handles superimposed. This equivalence of the two pattern masks indicates that contrast summation between the plaid components was perfectly linear prior to the masking stage. Furthermore, the pattern masks did not drive the detecting mechanism above its detection threshold because they did not abolish facilitation by the pedestal (Foley, 1994). Therefore, the pattern masking could not be attributed to within-channel masking, suggesting that linear summation of contrast signals takes place within a suppressive contrast gain pool. We present a quantitative model of the effects and discuss the implications for neurophysiological models of the process. © 2004 ARVO.

Contrast summation across eyes and space is revealed along the entire dipper function by a "Swiss cheese" stimulus.

Previous contrast discrimination experiments have shown that luminance contrast is summed across ocular (T. S. Meese, M. A. Georgeson, & D. H. Baker, 2006) and spatial (T. S. Meese & R. J. Summers, 2007) dimensions at threshold and above. However, is this process sufficiently general to operate across the conjunction of eyes and space? Here we used a "Swiss cheese" stimulus where the blurred "holes" in sine-wave carriers were of equal area to the blurred target ("cheese") regions. The locations of the target regions in the monocular image pairs were interdigitated across eyes such that their binocular sum was a uniform grating. When pedestal contrasts were above threshold, the monocular neural images contained strong evidence that the high-contrast regions in the two eyes did not overlap. Nevertheless, sensitivity to dual contrast increments (i.e., to contrast increments in different locations in the two eyes) was a factor of ∼1.7 greater than to single increments (i.e., increments in a single eye), comparable with conventional binocular summation. This provides evidence for a contiguous area summation process that operates at all contrasts and is influenced little, if at all, by eye of origin. A three-stage model of contrast gain control fitted the results and possessed the properties of ocularity invariance and area invariance owing to its cascade of normalization stages. The implications for a population code for pattern size are discussed.

Spatially extensive summation of contrast-energy is revealed by contrast detection of micro-pattern textures

Vision must analyze the retinal image over both small and large areas to represent fine-scale spatial details and extensive textures. The long-range neuronal convergence that this implies might lead us to expect that contrast sensitivity should improve markedly with the contrast area of the image. But this is at odds with the orthodox view that contrast sensitivity is determined merely by probability summation over local independent detectors. To address this puzzle, I aimed to assess the summation of luminance contrast without the confounding influence of area-dependent internal noise. I measured contrast detection thresholds for novel Battenberg stimuli that had identical overall dimensions (to clamp the aggregation of noise) but were constructed from either dense or sparse arrays of micro-patterns. The results unveiled a three-stage visual hierarchy of contrast summation involving (i) spatial filtering, (ii) long-range summation of coherent textures, and (iii) pooling across orthogonal textures. Linear summation over local energy detectors was spatially extensive (as much as 16 cycles) at Stage 2, but the resulting model is also consistent with earlier classical results of contrast summation (J. G. Robson & N. Graham, 1981), where co-aggregation of internal noise has obscured these long-range interactions.

Grid-texture mechanisms in human vision:contrast detection of regular sparse micro-patterns requires specialist templates

Previous work has shown that human vision performs spatial integration of luminance contrast energy, where signals are squared and summed (with internal noise) over area at detection threshold. We tested that model here in an experiment using arrays of micro-pattern textures that varied in overall stimulus area and sparseness of their target elements, where the contrast of each element was normalised for sensitivity across the visual field. We found a power-law improvement in performance with stimulus area, and a decrease in sensitivity with sparseness. While the contrast integrator model performed well when target elements constituted 50–100% of the target area (replicating previous results), observers outperformed the model when texture elements were sparser than this. This result required the inclusion of further templates in our model, selective for grids of various regular texture densities. By assuming a MAX operation across these noisy mechanisms the model also accounted for the increase in the slope of the psychometric function that occurred as texture density decreased. Thus, for the first time, mechanisms that are selective for texture density have been revealed at contrast detection threshold. We suggest that these mechanisms have a role to play in the perception of visual textures.

Contrast and lustre:a model that accounts for eleven different forms of contrast discrimination in binocular vision

Our goal here is a more complete understanding of how information about luminance contrast is encoded and used by the binocular visual system. In two-interval forced-choice experiments we assessed observers' ability to discriminate changes in contrast that could be an increase or decrease of contrast in one or both eyes, or an increase in one eye coupled with a decrease in the other (termed IncDec). The base or pedestal contrasts were either in-phase or out-of-phase in the two eyes. The opposed changes in the IncDec condition did not cancel each other out, implying that along with binocular summation, information is also available from mechanisms that do not sum the two eyes' inputs. These might be monocular mechanisms. With a binocular pedestal, monocular increments of contrast were much easier to see than monocular decrements. These findings suggest that there are separate binocular (B) and monocular (L,R) channels, but only the largest of the three responses, max(L,B,R), is available to perception and decision. Results from contrast discrimination and contrast matching tasks were described very accurately by this model. Stimuli, data, and model responses can all be visualized in a common binocular contrast space, allowing a more direct comparison between models and data. Some results with out-of-phase pedestals were not accounted for by the max model of contrast coding, but were well explained by an extended model in which gratings of opposite polarity create the sensation of lustre. Observers can discriminate changes in lustre alongside changes in contrast.

Discriminating depth in stereo surfaces: dipper effect found, and lost

Measurement of detection and discrimination thresholds yields information about visual signal processing. For luminance contrast, we are 2 - 3 times more sensitive to a small increase in the contrast of a weak 'pedestal' grating, than when the pedestal is absent. This is the 'dipper effect' - a reliable improvement whose interpretation remains controversial. Analogies between luminance and depth (disparity) processing have attracted interest in the existence of a 'disparity dipper' - are thresholds for disparity, or disparity modulation (corrugated surfaces), facilitated by the presence of a weak pedestal? Lunn and Morgan (1997 Journal of the Optical Society of America A 14 360 - 371) found no dipper for disparity-modulated gratings, but technical limitations (8-bit greyscale) might have prevented the necessary measurement of very small disparity thresholds. We used a true 14-bit greyscale to render small disparities accurately, and measured 2AFC discrimination thresholds for disparity modulation (0.6 cycle deg-1) of a random texture at various pedestal levels. Which interval contained greater modulation of depth? In the first experiment, a clear dipper was found. Thresholds were about 2X1 lower with weak pedestals than without. But here the phase of modulation (0° or 180°) was randomised from trial to trial. In a noisy signal-detection framework, this creates uncertainty that is reduced by the pedestal, thus improving performance. When the uncertainty was eliminated by keeping phase constant within sessions, the dipper effect disappeared, confirming Lunn and Morgan's result. The absence of a dipper, coupled with shallow psychometric slopes, suggests that the visual response to small disparities is essentially linear, with no threshold-like nonlinearity.

The role of the cortical occipital spike in photosensitive epilepsy

Chapters one to three are an introduction to photosensitive epilepsy, electroencephalography (EEG) and the magnocellular and parvocellular visual pathways. Photoparoxysmal response (PPR) are strongly associated with photosensitive epilepsy. Chapters four to nine investigated whether occipital spikes were associated with PPR and hence with photosensitive epilepsy. The chapters investigated whether the response types showed similar dependence on stimulus characteristics using EEG. Chapters four and five found that occipital spikes and PPR showed different dependence on colour and luminance contrast. The differences were consistent with the magnocellular pathway mediating occipital spikes and the pavocellular pathway mediating PPR. The study in chapter eight found that monocular occlusion had a significantly greater effect on PPR than on occipital spikes, which is further evidence against an association between the two types of response. Chapters six and seven showed that occipital spikes and PPR had similar optimum spatial and temporal frequencies. Chapter nine showed that both response types could be generated via stimulation of the periphery of the retina. However, these three chapters are not strong evidence of an association, as the results do not contradict the theory that the responses are generated via different pathways. The magnocellular and pavocellular pathways have similar optimum temporal and spatial frequencies and both are present in the periphery. In chapter ten, magnetoencephalography was used to estimate the source of activity underlying the components of the VEP and occipital spike. Changes in the amplitude and latency in the components of the normal VEP are associated with epilepsy. However, the source underlying the occipital spikes was not related to that underlying the components of the VEP so this is also removed as a source of evidence for an association between occipital spikes and photosensitive epilepsy.

Nonlinearities in the binocular combination of luminance and contrast

We studied the rules by which visual responses to luminous targets are combined across the two eyes. Previous work has found very different forms of binocular combination for targets defined by increments and by decrements of luminance, with decrement data implying a severe nonlinearity before binocular combination. We ask whether this difference is due to the luminance of the target, the luminance of the background, or the sign of the luminance excursion. We estimated the pre-binocular nonlinearity (power exponent) by fitting a computational model to ocular equibrightness matches. The severity of the nonlinearity had a monotonic dependence on the signed difference between target and background luminance. For dual targets, in which there was both a luminance increment and a luminance decrement (e.g. contrast), perception was governed largely by the decrement. The asymmetry in the nonlinearities derived from the subjective matching data made a clear prediction for visual performance: there should be more binocular summation for detecting luminance increments than for detecting luminance decrements. This prediction was confirmed by the results of a subsequent experiment. We discuss the relation between these results and luminance nonlinearities such as a logarithmic transform, as well as the involvement of contemporary model architectures of binocular vision.