Surround suppression and facilitation in the fovea: very long-range spatial interactions in contrast perception.

Surround modulation of perceived contrast has been almost exclusively studied in short-range conditions, i.e., in situations where a tiny gap, at most, separates center from surround. Existing long-range studies suggest that suppression extends to 12-cycle distance, whereas facilitation of perceived contrast is suggested to arise from visual field regions enclosing the center. In V1 neurons, however, long-range surround modulation involves both suppression and facilitation. Thus, we investigated short- and long-range surround modulation by measuring the perceived contrast of a center in the presence of a surround either near (0.3 cycles, 0.1 degree) or far (19.8 cycles, 6.6 degrees) from the center. This study demonstrates that in addition to the well-known suppression, surround modulation involves remarkably long-range facilitation of perceived contrast. At low center contrasts, the long-range facilitation was stronger than the long-range suppression, whereas at high center contrast we found mainly long-range suppression. Because the current models of perceived contrast could not account for our data, we considered our results in the context of models developed for surround modulation in V1 neurons. However, neither mechanistic nor phenomenological models proved satisfactory. Moreover, with the current knowledge, it seems that straightforward pooling of V1 neurons' responses cannot account for surround modulation of perceived contrast.

Area summation in human visual system: psychophysics, fMRI, and modeling.

Contextual modulation is a fundamental feature of sensory processing, both on perceptual and on single-neuron level. When the diameter of a visual stimulus is increased, the firing rate of a cell typically first increases (summation field) and then decreases (surround field). Such an area summation function draws a comprehensive profile of the receptive field structure of a neuron, including areas outside the classical receptive field. We investigated area summation in human vision with psychophysics and functional magnetic resonance imaging (fMRI). The stimuli were drifting sine wave gratings similar to those used in previous macaque single-cell area summation studies [corrected]. A model was developed to facilitate comparison of area summation in fMRI to area summation in psychophysics and single cells. The model consisted of units with an antagonistic receptive field structure found in single cells in the primary visual cortex. The receptive field centers of the model neurons were distributed in the region of the visual field covered by a single voxel. The measured area summation functions were qualitatively similar to earlier single-cell data. The model with parameters derived from psychophysics captured the spatial structure of the summation field in the primary visual cortex as measured with fMRI. The model also generalized to a novel situation in which the neural population was displaced from the stimulus center. The current study shows that contextual modulation arises from similar spatially antagonistic and overlapping excitatory and inhibitory mechanisms, both in single cells and in human vision.

Low-level features determine brightness in White's and Benary's illusions.

We masked White's and Benary's brightness illusions and simultaneous contrast with narrowband visual noise and measured detection thresholds and brightness. The noise was either isotropic or orientation filtered. A narrow spatial frequency tuning was found for detection and brightness for every stimulus. A narrow orientation tuning was also found: the strength of the illusions decreased (White and Benary) or increased (White) depending on the orientation of the mask. The critical borders were always of the same contrast polarity. The results suggest that the brightness in figure-ground scenes is determined by mechanisms integrating incremental and decremental borders in early visual cortices.

Time course of suppression by surround gratings: highly contrast-dependent, but consistently fast.

Timing is critical for the effectiveness of a modulating surround signal. In this study, the optimal timing of a suppressing surround signal was measured psychophysically in human subjects. The perceived contrast of a fixated 1-deg circular patch of vertical sinusoidal grating (the target: 4 cpd, Michelson contrast 0.2) was measured as a function of the onset asynchrony between the target and an annular "surround" grating with the same orientation and spatial frequency. The contrast and area of the surround stimulus were varied parametrically. The suppressive signal peaked at earlier times the higher the surround contrast (0.1-0.4), following a function consistent with the contrast-dependence of retinal response dynamics. Increasing the area of the surround grating also moved peak suppression to earlier times. At ca. 2 deg annulus outer diameter the time to peak of the suppressive signal was shortest, although its amplitude grew with annulus area even beyond that. When both the contrast and the area of the centre and surround gratings were equal, suppression was maximal if the surround stimulus was presented ca. 5 ms before the target. Such a short delay of suppression is consistent with a neural implementation based on feedforward-feedback connections, but not with horizontal connections.

Brightness processing in the visual cortex.

Single cell recordings have shown that some cells in the primary visual cortex (V1) signal surface brightness. However, fMRI experiments have found brightness related activation only in the higher cortical areas. In a psychophysical setup, we were able to dissociate the reduction of brightness caused by Gabor flankers, similar to the receptive fields in V1, from the edge induced brightness change. The former resemble the single cell recording results and the latter the fMRI results.

Spatial frequency difference between textures interferes with brightness perception.

Abrupt changes in luminance trigger and restrict brightness filling-in. If brightness was actively filled-in and mediated by cells signaling both luminance borders and surface brightness, then brightness spreading could also get disrupted by changes in texture. We measured psychophysically the brightness of a uniform luminance disk, which was segmented into two parts by different textures. The brightness of the central part of the disk was substantially reduced, and the reduction depended on spatial frequency, but not on the orientation difference between the textures. The results show that texture borders are able to block brightness filling-in. The bandwidth of brightness spreading was estimated to be approximately 1.5 octaves. This suggests that brightness information spreads only between neurons of similar spatial frequency characteristics.

Collinear context (and learning) change the profile of the perceptual filter.

The effect of collinear context on the filter mediating the detection of a Gabor stimulus was investigated by using the classification image method. Classification images were estimated for a 1.5 cpd horizontal Gabor target and the same target flanked by two collinear Gabors horizontally 1.7 degrees displaced from the target. The target was masked by a low-contrast white-noise mask. Obtained classification images were fitted by Gabor functions. The results show that collinear flankers increase the length of the classification image profiles along the collinear axis. At the same time, modest facilitory effects were observed in most subjects. The specificity and the amount of context-induced elongation in the classification images makes it hard to be explained by uncertainty reduction alone. In previous studies, collinear facilitation has been reported to abolish due to perceptual learning. We report a possibly related phenomenon: classification image data was re-analyzed in two parts consisting of the early and the late trials. In the latter trials, differences between the classification images in flankers and no-flankers condition are no longer significant.

Spatial frequency tuning of brightness polarity identification.

Recent studies have shown that cells in the primary visual cortex can, in addition to borders, also encode surface brightness. Whether the brightness is encoded by a large extraclassical receptive field or by a filling-in type mechanism activated by the luminance border is not known. These explanations imply different spatial frequency tunings for the underlying mechanism. In a psychophysical masking paradigm we measured spatial frequency tuning functions for identification of both luminance polarity (bright/dark) and luminance border orientation of oval and circular luminance patches with variable diameters (0.2-10 deg). For both tasks we obtained nearly overlapping narrow (1.5 octave) bandpass masking tuning functions centered at 1.5-5.0 c/deg. Stimulus size and shape had only minimal effect on the tuning functions. The results favor the idea of brightness filling-in and suggest that the cells activated by the luminance border modulate the activity of the cells signaling surface brightness. Further, the brightness processing mechanism is spatial frequency selective.

Contextual effects in fine spatial discriminations.

The context in which a pattern is viewed can greatly affect its apparent contrast, a phenomenon commonly attributed to pooled contrast gain control processes. A low-contrast surround may slightly enhance apparent contrast, whereas increasing the contrast of the surround leads to a monotonic decline in contrast appearance. We ask here how the presence of a patterned surround affects the ability to perform fine, suprathreshold orientation, contrast, and spatial frequency discriminations as a function of surround contrast and phase. Our results revealed an unexpected dip in performance when center and surround were in phase and similar in contrast. These results suggest that additional processes, perhaps those involved in scene segregation, play a role in contextual effects on discrimination.

Separation of edge detection and brightness perception.

When a low spatial frequency noise mask is superimposed onto a luminance staircase, the perceived brightness pattern is dramatically altered although the edges remain visible. We measured contrast thresholds for the edges and for the illusory scalloping (Chevreul-illusion), as a function of noise center spatial frequency. The masking tuning functions overlapped, but peaked at different spatial frequencies and contrast levels. The results suggest that perceived brightness is triggered only by the low spatial frequency components of the edges--the high spatial frequency components are not able to produce a brightness pattern.

Spatial integration in Glass patterns.

The extraction of a global orientation structure presumably has a different neural mechanism from that of the analysis of its local features. We investigated spatial integration within these two mechanisms using stimulus patterns composed of dot pairs (dipoles). The stimuli targeted local feature detection, contained no global configuration, but rather contained randomly oriented dipoles of a fixed length (the distance between the dots in a pair). For the detection of a global orientation structure, local dipole orientations were arranged in a concentric Glass pattern. Thresholds as a function of a stimulus area were determined by measuring the minimum proportion of dipoles among random-dot noise (signal-to-noise ratio) required for the detection of dipoles (features), as well as for the detection of an orientation structure. Thresholds for feature detection were significantly higher than those for the detection of the global structure--regardless of the stimulus size. Spatial integration, however, did not differ between the two tasks: the exponents of the power functions fitted to data for six observers were -0.48 +/- 0.07 for random dipole orientations and -0.62 +/- 0.1 for Glass patterns.