Erich W. Graf

High-Level Face Adaptation Without Awareness

When a visual stimulus is suppressed from awareness, processing of the suppressed image is necessarily reduced. Although adaptation to simple image properties such as orientation still occurs, adaptation to high-level properties such as face identity is eliminated. Here we show that emotional facial expression continues to be processed even under complete suppression, as indexed by substantial facial expression aftereffects.

Natural images dominate in binocular rivalry

Ecological approaches to perception have demonstrated that information encoding by the visual system is informed by the natural environment, both in terms of simple image attributes like luminance and contrast, and more complex relationships corresponding to Gestalt principles of perceptual organization. Here, we ask if this optimization biases perception of visual inputs that are perceptually bistable. Using the binocular rivalry paradigm, we designed stimuli that varied in either their spatiotemporal amplitude spectra or their phase spectra. We found that noise stimuli with “natural” amplitude spectra (i.e., amplitude content proportional to 1/f, where f is spatial or temporal frequency) dominate over those with any other systematic spectral slope, along both spatial and temporal dimensions. This could not be explained by perceived contrast measurements, and occurred even though all stimuli had equal energy. Calculating the effective contrast following attenuation by a model contrast sensitivity function suggested that the strong contrast dependency of rivalry provides the mechanism by which binocular vision is optimized for viewing natural images. We also compared rivalry between natural and phase-scrambled images and found a strong preference for natural phase spectra that could not be accounted for by observer biases in a control task. We propose that this phase specificity relates to contour information, and arises either from the activity of V1 complex cells, or from later visual areas, consistent with recent neuroimaging and single-cell work. Our findings demonstrate that human vision integrates information across space, time, and phase to select the input most likely to hold behavioral relevance.

The spatial scale of perceptual memory in ambiguous figure perception.

Ambiguous visual stimuli highlight the constructive nature of vision: perception alternates between two plausible interpretations of unchanging input. However, when a previously viewed ambiguous stimulus reappears, its earlier perception almost entirely determines the new interpretation; memory disambiguates the input. Here, we investigate the spatial properties of this perceptual memory, taking into account strong anisotropies in percept preference across the visual field. Countering previous findings, we show that perceptual memory is not confined to the location in which it was instilled. Rather, it spreads to noncontiguous regions of the visual field, falling off at larger distances. Furthermore, this spread of perceptual memory takes place in a frame of reference that is tied to the surface of the retina. These results place the neural locus of perceptual memory in retinotopically organized sensory cortical areas, with implications for the wider function of perceptual memory in facilitating stable vision in natural, dynamic environments.

On the relation between dichoptic masking and binocular rivalry

When our two eyes view incompatible images, the brain invokes suppressive processes to inhibit one image, and favor the other. Two phenomena are typically observed: dichoptic masking (reduced sensitivity to one image) for brief presentations, and binocular rivalry (alternation between the two images), over longer exposures. However, it is not clear if these two phenomena arise from a common suppressive process. We investigated this by measuring both threshold elevation in simultaneous dichoptic masking and mean percept durations in rivalry, whilst varying relative stimulus orientation. Masking and rivalry showed significant correlations, such that strong masking was associated with long dominance durations. A second experiment suggested that individual differences across both measures are also correlated. These findings are consistent with varying the magnitude of interocular suppression in computational models of both rivalry and masking, and imply the existence of a common suppressive process. Since dichoptic masking has been localised to the monocular neurons of V1, this is a plausible first stage of binocular rivalry.

Plasticity of convergence-dependent variations of cyclovergence with vertical gaze

Binocular alignment of foveal images is facilitated by cross-couplings of vergence eye movements with distance and direction of gaze. These couplings reduce horizontal, vertical and cyclodisparities at the fovea without using feedback from retinal image disparity. Horizontal vergence is coupled with accommodation. Vertical vergence that aligns tertiary targets in asymmetric convergence is thought to be coupled with convergence and horizontal gaze. Cyclovergence aligns the horizontal retinal meridians during gaze elevation in symmetrical convergence and is coupled with convergence and vertical gaze. The latter vergence-dependent changes of cyclovergence have been described in terms of the orientation of Listings plane and have been referred to as the binocular extension of Listings law. Can these couplings be modified? Plasticity has been demonstrated previously for two of the three dimensions of vergence (horizontal and vertical). The current study demonstrates that convergence-dependent changes of the orientation of Listings plane can be adapted to either exaggerate or to reduce the cyclovergence that normally facilitates alignment of the horizontal meridians of the retinas with one another during gaze elevation in symmetrical convergence. The adaptability of cyclovergence demonstrates a neural mechanism that, in conjunction with the passive forces determined by biomechanical properties of the orbit, could play an active role in implementing Listings extended law and provide a means for calibrating binocular eye alignment in three dimensions.

The fate of task-irrelevant visual motion: Perceptual load versus feature-based attention

We tested contrasting predictions derived from perceptual load theory and from recent feature-based selection accounts. Observers viewed moving, colored stimuli and performed low or high load tasks associated with one stimulus feature, either color or motion. The resultant motion aftereffect (MAE) was used to evaluate attentional allocation. We found that task-irrelevant visual features received less attention than co-localized task-relevant features of the same objects. Moreover, when color and motion features were co-localized yet perceived to belong to two distinct surfaces, feature-based selection was further increased at the expense of object-based co-selection. Load theory predicts that the MAE for task-irrelevant motion would be reduced with a higher load color task. However, this was not seen for co-localized features; perceptual load only modulated the MAE for task-irrelevant motion when this was spatially separated from the attended color location. Our results suggest that perceptual load effects are mediated by spatial selection and do not generalize to the feature domain. Feature-based selection operates to suppress processing of task-irrelevant, co-localized features, irrespective of perceptual load.

Changes in cyclotorsion and vertical eye alignment during prolonged monocular occlusion

When binocular vision is prevented with monocular occlusion, the two eyes assume a position of rest related to the combination of underlying tonic innervation of the oculomotor system, cross-coupled accommodative-vergence input and vergence responses to perceptual cues for spatial location relative to the head. When the latter two are controlled, the covered eye has been shown in the majority of subjects to turn outward (exophoria) and upward (hyperphoria) after prolonged monocular occlusion. The present study investigates the change in torsional eye alignment and its relation to vertical eye alignment after eight hours of monocular occlusion. The results revealed an excyclophoria during occlusion in four out of five subjects. The patched eye also became elevated in two subjects and depressed in two others. Thus, during prolonged monocular occlusion, the relative directions of cyclophoria and vertical phoria appear to be independent. In addition, there were non-concomitant changes in vertical phoria with horizontal gaze, toward a state where the adducted eye was elevated relative to the abducted eye. Simulations with Orbit(TM) suggest that these non-concomitant changes in vertical phoria with a concomitant excyclophoria may be based upon orbital mechanics. Excyclophoria appears to be the baseline state of binocular alignment.

Equivalence of physical and perceived speed in binocular rivalry.

The relative dominance of gratings engaged in binocular rivalry can be influenced by their surroundings. One striking example occurs when surrounding motion is congruent with one but not the other grating (C. L. Paffen, S. F. te Pas, R. Kanai, M. J. van der Smagt, & F. A. Verstraten, 2004). However, such center-surround stimulus configurations can also modulate perceived speed, via a directionally tuned process (H. P. Norman, J. F. Norman, J. T. Todd, & D. T. Lindsey, 1996). We recorded rivalry for Gabor patches embedded in a drifting noise texture. Gratings whose directions opposed the background motion tended to dominate more, and vice versa, consistent with previous findings. Observers then matched the speed of a drifting noise-embedded Gabor to that of a Gabor surrounded by mean luminance. Surround motion produced substantial changes in perceived speed, by at least a factor of two for all observers. We then asked whether perceived speed could account for the contextual effects on dominance. We measured the effects of speed on rivalry dominance by changing the physical speeds of rivaling gratings, as determined by the matching data. We found the same pattern of dominance as for the context experiment, indicating that perceived and true speed influence rivalry in the same manner. We propose a Bayesian interpretation of the perceived speed illusion.

Spatial and temporal tuning of motion in depth.

We used the Pulfrich effect to investigate perception of motion in depth. Independent manipulation of spatial and temporal frequency content in stereoscopic motion stimuli revealed the tuning characteristics of motion-in-depth perception. Sensitivity to interocular phase difference between sinusoidally oscillating sine-wave gratings was measured in four observers who judged direction of motion in depth. Discrimination thresholds in terms of interocular phase difference were determined to investigate spatial and temporal tuning characteristics of a system that is based on interocular phase difference, interocular delay, binocular disparity and velocity difference. Temporal frequency tuning of interocular phase difference thresholds was band pass and relatively dependent on spatial frequency variation. These results together with evidence from two control experiments support the idea that sensitivity to direction of motion in depth is limited by a stereo-motion system that monitors binocular horizontal disparity and motion rather than interocular phase difference, interocular delay, or interocular velocity difference.

Motion-aftereffect-induced blindness

Motion-induced blindness (MIB) describes the occasional disappearance of salient visual objects in the presence of moving features (Y. S. Bonneh, A. Cooperman, & D. Sagi, 2001). Here we test whether motion adaptation and the ensuing motion aftereffect (MAE) are sufficient to trigger disappearance of salient targets. In three experiments, observers adapted to either rotating or static stimuli. Immediately afterwards, a static test pattern was presented consisting of a mask with texture elements and three superimposed target dots in a triangular arrangement. Observers reported dot disappearance and reappearance. The results clearly show that illusory motion in a static test pattern, following motion adaptation, promotes the disappearance of target dots. Furthermore, disappearance is modulated by the depth relationship between test pattern and targets, increasing for targets placed stereoscopically behind the test pattern. We conclude that MIB is influenced by perceived relative motion between depth-segregated features.