Cortical visual neurons in the cat and monkey are inhibited by stimuli surrounding their receptive fields (surround suppression) or presented within their receptive fields (cross-orientation or overlay suppression). We show that human contrast sensitivity is similarly affected by two distinct suppression mechanisms. In agreement with the animal studies, human surround suppression is tightly tuned to the orientation and spatial frequency of the test, unlike overlay suppression. Using a double-masking paradigm, we also show that in humans, overlay suppression precedes surround suppression in the processing sequence. Surprisingly, we find that, unlike overlay suppression, surround suppression is only strong in the periphery (Ͼ1°eccentricity). This result argues for a new functional distinction between foveal and peripheral operations.

Crowding (mutual scrambling of nearby peripheral stimuli) has several known asymmetries. We explored these and other asymmetries systematically across the visual field. Crowding strength for 16 target (Gabor) positions in the visual field (8 directions × 2 eccentricities) were determined by positioning a plaid mask made of two transparently overlaid Gabors either inward, outward, clockwise, or counter-clockwise around the target. Overall, we found a surprisingly large individual variation in crowding strength appearing as idiosyncratic hotspots across the visual field. No correlations were found between the idiosyncratic variations of crowding and visual acuity either across the visual field or across subjects. When averaged across observers the results replicated most of the previously reported asymmetries of crowding. No new types of asymmetries were observed, but we found that the inward-outward asymmetry of crowding is present only along the horizontal meridian. Most surprisingly, we discovered that this asymmetry increases two-fold, if the observer is forced to attend to both left and right visual fields. This indicates that besides other factors attention allocation has a strong effect on the crowding asymmetry.

Crowding and surround suppression share many similarities, which suggests the possibility of a common mechanism. Despite decades of research, there has been little effort to compare the two phenomena in a consistent fashion. A recent study by D. M. Levi, S. Hariharan, and S. A. Klein (2002) argues that the two are unrelated because crowding effects can be much stronger than suppression effects. Here we report experiments in which the same Gabor target was used both for orientation identification (crowding) and contrast detection (suppression) tasks. In agreement with early crowding studies (e.g., H. Bouma, 1973) we found, that an outward mask is much more effective than an inward mask for the orientation identification task. Notably, no such anisotropy was observed for the contrast detection task, commonly used to measure surround suppression. The anisotropic masking, which defines crowding, is observed only at fine scales (roughly within an octave of the acuity limit), whereas surround suppression is observed at all scales. Our results demonstrate that surround suppression and crowding are indeed two distinct phenomena. We used this characteristic anisotropy to show that a popular crowding paradigm in which target contrast is varied to measure crowding is confounding it with surround suppression. Surround suppression apparently dominates at low contrasts, which would explain some of the reported similarities between the two phenomena.

Contrast sensitivity is known to be strongly influenced by the target surround, yet the role of the surround interaction in visual processing remains unclear. Previously, we have shown that the surround strongly suppresses contrast sensitivity in the periphery when the surround spatial frequency and orientation match those of the target (Petrov, Carandini, & McKee, 2005). Here, we explore how various spatial characteristics of the iso-oriented and frequency-matched surround, such as surround phase and spatial layout, affect suppression. We manipulated surround geometry (annulus ring, half annulus, and bow tie) and its separation from the target (both laterally and in depth) and varied the position of the half-annulus and bow-tie surrounds with respect to Gabor target's orientation and with respect to its location in the visual field (i.e., radial vs. tangential surrounds). We also compared monoptic, dichoptic, and binocular surround suppression. Except for a significant radial-tangential anisotropy, only the area of the surround and the lateral separation between the surround and target had a significant effect on the magnitude of suppression. We showed that, although suppression amplitude remains constant with stimulus eccentricity, the lateral extent of suppression scales in proportion to the eccentricity. The most surprising finding was that the extent of surround suppression does not scale with stimulus size or spatial frequency. We suggest that the properties of surround suppression are best explained by a mechanism that selects salient targets for subsequent saccades.

It has long been known that an outward mask is much more disruptive than an inward mask in crowding (H. Bouma, 1973). We show that the locus of attention strongly affects this inward-outward anisotropy, removing it in some conditions and reversing it in others. In a 2AFC paradigm, subjects identified whether a high-contrast Gabor target of a given orientation was presented left or right of fixation. When a fixed eccentricity (8°) was used, the outward plaid mask produced much stronger crowding than the inward mask. When 7°, 8°, and 9° eccentricities were interleaved within the same run, diffusing attention, the inward and outward masks produced the same amount of crowding for all three eccentricities. When target identification was contingent on a foveal cue, biasing attention inward, the inward mask produced stronger crowding. Finally, a new contrast-detection paradigm was used to demonstrate that attention is generally mislocalized outward of the target, which may explain the commonly observed anisotropy in crowding. Our results suggest that spatial attention is intimately involved in the mechanism of crowding.

When flanked by collinear Gabor patches, detection thresholds for a target Gabor patch improve by up to a factor of 2. This result has been interpreted as evidence for collinear facilitation. However, facilitation has been observed only for targets near detection threshold, where observers seem uncertain about the location and other properties of the stimulus. So the effect of the flankers may be to reduce this uncertainty. If this is true, then other cues to target location should produce a similar improvement in thresholds. To test this hypothesis, we measured contrast detection thresholds for a Gabor target alone, and in the presence of either a faint circle surrounding the target location, or two high-energy flanking Gabor patches. We also used an adaptive procedure to measure the slope of the psychometric function to determine whether the slopes were considerably lower in the presence of location cues or flanking Gabors, as predicted by signal detection theory when uncertainty is reduced. As observed previously, the presence of collinear flankers improved detection thresholds by a factor of two. Yet, on average, the circle alone accounted for the most of the facilitation; for three of our five observers, it improved thresholds as much as the collinear flankers. Other cues that specified target location produced similar improvements in detection thresholds. The slopes of the psychometric functions were much shallower in the presence of these location cues or the collinear flankers compared to the target-alone condition. This change in the slopes indicates that the threshold improvement is largely due to a significant reduction in uncertainty.

The abundant literature on crowding offers fairly simple explanations for the phenomenon, such as position uncertainty or feature pooling, but convincing evidence to support these explanations is lacking. In part, this is because the stimuli used for crowding studies are usually letters or other complex shapes, which makes it hard to determine exactly what kind of information is lost. In our experiment, we asked observers to identify simultaneously the slants (left or right) of three horizontally aligned Gabor targets. The targets were presented at 6 degrees in the periphery, and their size and separation were chosen to incur strong crowding. The loss of information about the position or orientation of individual members of the Gabor triads does not explain our results. Instead, crowding appears to be a particular form of collective information loss. Firstly, the outmost target was crowded much less than the other targets, which rules out explanations based on simple pooling and shows that crowding has a pronounced foveal directionality. Secondly, the specific pattern of confusion shown by all the observers indicates that the only reliable information available to them was orientation contrast, that is, the number (and, to a lesser degree, the location) of sites where slant changed. Thus, crowding appears to spare only the most salient peripheral information, which supports the hypothesis that crowding is caused by limitations of attentional resolution.

We explored the time course of surround suppression and found clear evidence for two distinct mechanisms: one strong, transient, and largely monocular, the other weaker, sustained, and binocular. We measured detection thresholds for a Gabor target at 8 deg eccentricity surrounded by a large annulus of matching spatial frequency and orientation. At short stimulus durations surround suppression was very strong, but the suppression strength decreased precipitously for durations longer than ~100 msec. The strong transient component did not transfer between the eyes and occurred almost instantaneously (<1 frame delay, 12 msec) irrespective of the separation between target and surround. Both suppression components were tightly tuned to orientation, peaking at target orientation, but neither was tuned to target spatial phase. These results are in good agreement with surround suppression properties measured in macaque V1 neurons. The absence of interocular transfer, the strong orientation selectivity, and the high propagation speed incommensurate with slow horizontal connections in V1 suggest that the transient component of suppression originates between input layers and the subsequent layers in V1.