Using ambiguous plaid stimuli to investigate the influence of immediate prior experience on perception.

In a series of three experiments, we used an ambiguous plaid motion stimulus to explore the behavioral and electrophysiological effects of prior stimulus exposures and perceptual states on current awareness. The results showed that prior exposure to a stimulus biased toward one percept led to subsequent suppression of that percept. In contrast, in the absence of stimulus bias, prior perceptual experience can have a facilitative influence. The suppressive effects caused by the prior stimulus were found to transfer to an ambiguous plaid test stimulus rotated 180º relative to the adaptation stimulus, but were abolished if (1) the ambiguous test stimulus was only rotated 90º relative to the adaptation stimulus or (2) the adaptation stimulus was heavily biased toward the component grating percept. Event-related potential recordings were consistent with the involvement of visual cortical areas and suggested that the influence of recent stimulus exposure may involve recruitment of additional brain processes beyond those responsible for initial stimulus encoding. In contrast, the effects of prior and current perceptual experience appeared to depend on similar brain processes. Although the data presented here focus on vision, the work is discussed within the context of data from a parallel series of experiments in audition.

Situation-based social anxiety enhances the neural processing of faces: evidence from an intergroup context.

Social anxiety is the intense fear of negative evaluation by others, and it emerges uniquely from a social situation. Given its social origin, we asked whether an anxiety-inducing social situation could enhance the processing of faces linked to the situational threat. While past research has focused on how individual differences in social anxiety relate to face processing, we tested the effect of manipulated social anxiety in the context of anxiety about appearing racially prejudiced in front of a peer. Visual processing of faces was indexed by the N170 component of the event-related potential. Participants viewed faces of Black and White males, along with nonfaces, either in private or while being monitored by the experimenter for signs of prejudice in a 'public' condition. Results revealed a difference in the N170 response to Black and Whites faces that emerged only in the public condition and only among participants high in dispositional social anxiety. These results provide new evidence that anxiety arising from the social situation modulates the earliest stages of face processing in a way that is specific to a social threat, and they shed new light on how anxiety effects on perception may contribute to the regulation of intergroup responses.

Art reaches within: aesthetic experience, the self and the default mode network.

In a task of rating images of artworks in an fMRI scanner, regions in the medial prefrontal cortex that are known to be part of the default mode network (DMN) were positively activated on the highest-rated trials. This is surprising given the DMN's original characterization as the set of brain regions that show greater fMRI activity during rest periods than during performance of tasks requiring focus on external stimuli. But further research showed that DMN regions could be positively activated also in structured tasks, if those tasks involved self-referential thought or self-relevant information. How may our findings be understood in this context? Although our task had no explicit self-referential aspect and the stimuli had no a priori self-relevance to the observers, the experimental design we employed emphasized the personal aspects of aesthetic experience. Observers were told that we were interested in their individual tastes, and asked to base their ratings on how much each artwork "moved" them. Moreover, we used little-known artworks that covered a wide range of styles, which led to high individual variability: each artwork was rated highly by some observers and poorly by others. This means that rating-specific neural responses cannot be attributed to the features of any particular artworks, but rather to the aesthetic experience itself. The DMN activity therefore suggests that certain artworks, albeit unfamiliar, may be so well-matched to an individual's unique makeup that they obtain access to the neural substrates concerned with the self-access which other external stimuli normally do not get. This mediates a sense of being "moved," or "touched from within." This account is consistent with the modern notion that individuals' taste in art is linked with their sense of identity, and suggests that DMN activity may serve to signal "self-relevance" in a broader sense than has been thought so far.

Slow cortical dynamics and the accumulation of information over long timescales.

Making sense of the world requires us to process information over multiple timescales. We sought to identify brain regions that accumulate information over short and long timescales and to characterize the distinguishing features of their dynamics. We recorded electrocorticographic (ECoG) signals from individuals watching intact and scrambled movies. Within sensory regions, fluctuations of high-frequency (64-200 Hz) power reliably tracked instantaneous low-level properties of the intact and scrambled movies. Within higher order regions, the power fluctuations were more reliable for the intact movie than the scrambled movie, indicating that these regions accumulate information over relatively long time periods (several seconds or longer). Slow (<0.1 Hz) fluctuations of high-frequency power with time courses locked to the movies were observed throughout the cortex. Slow fluctuations were relatively larger in regions that accumulated information over longer time periods, suggesting a connection between slow neuronal population dynamics and temporally extended information processing.

The brain on art: intense aesthetic experience activates the default mode network.

Aesthetic responses to visual art comprise multiple types of experiences, from sensation and perception to emotion and self-reflection. Moreover, aesthetic experience is highly individual, with observers varying significantly in their responses to the same artwork. Combining fMRI and behavioral analysis of individual differences in aesthetic response, we identify two distinct patterns of neural activity exhibited by different sub-networks. Activity increased linearly with observers' ratings (4-level scale) in sensory (occipito-temporal) regions. Activity in the striatum (STR) also varied linearly with ratings, with below-baseline activations for low-rated artworks. In contrast, a network of frontal regions showed a step-like increase only for the most moving artworks ("4" ratings) and non-differential activity for all others. This included several regions belonging to the "default mode network" (DMN) previously associated with self-referential mentation. Our results suggest that aesthetic experience involves the integration of sensory and emotional reactions in a manner linked with their personal relevance.

Feeling without seeing? Engagement of ventral, but not dorsal, amygdala during unaware exposure to emotional faces.

The ability to selectively perceive items in the environment may be modulated by the emotional content of those items. The neural mechanism that underlies the privileged processing of emotionally salient content is poorly understood. Here, using fMRI, we investigated this issue via a binocular rivalry procedure when face stimuli depicting fearful or neutral expressions competed for awareness with a house. Results revealed an interesting dissociation in the amygdala during rivalry condition: Whereas its dorsal component exhibited dominant activation to aware fearful faces, a ventral component was more active during the suppression of fearful faces. Moreover, during rivalry, the dorsal and ventral components of the amygdala were coupled with segregated cortical activations in the brainstem and medial PFC, respectively. In summary, this study points to a differential involvement of two clusters within the amygdala and their connected networks in naturally occurring perceptual biases of emotional content in faces.

Seeing race: N170 responses to race and their relation to automatic racial attitudes and controlled processing.

We examined the relation between neural activity reflecting early face perception processes and automatic and controlled responses to race. Participants completed a sequential evaluative priming task, in which two-tone images of Black faces, White faces, and cars appeared as primes, followed by target words categorized as pleasant or unpleasant, while encephalography was recorded. Half of these participants were alerted that the task assessed racial prejudice and could reveal their personal bias ("alerted" condition). To assess face perception processes, the N170 component of the ERP was examined. For all participants, stronger automatic pro-White bias was associated with larger N170 amplitudes to Black than White faces. For participants in the alerted condition only, larger N170 amplitudes to Black versus White faces were also associated with less controlled processing on the word categorization task. These findings suggest that preexisting racial attitudes affect early face processing and that situational factors moderate the link between early face processing and behavior.

Uncovering camouflage: amygdala activation predicts long-term memory of induced perceptual insight.

What brain mechanisms underlie learning of new knowledge from single events? We studied encoding in long-term memory of a unique type of one-shot experience, induced perceptual insight. While undergoing an fMRI brain scan, participants viewed degraded images of real-world pictures where the underlying objects were hard to recognize ("camouflage"), followed by brief exposures to the original images ("solution"), which led to induced insight ("Aha!"). A week later, the participants' memory was tested; a solution image was classified as "remembered" if detailed perceptual knowledge was elicited from the camouflage image alone. During encoding, subsequently remembered images were associated with higher activity in midlevel visual cortex and medial frontal cortex, but most pronouncedly, in the amygdala, whose activity could be used to predict which solutions will remain in long-term memory. Our findings extend the known roles of amygdala in memory to include promotion of long-term memory of the sudden reorganization of internal representations.

Alternation rate in perceptual bistability is maximal at and symmetric around equi-dominance.

When an ambiguous stimulus is viewed for a prolonged time, perception alternates between the different possible interpretations of the stimulus. The alternations seem haphazard, but closer inspection of their dynamics reveals systematic properties in many bistable phenomena. Parametric manipulations result in gradual changes in the fraction of time a given interpretation dominates perception, often over the entire possible range of zero to one. The mean dominance durations of the competing interpretations can also vary over wide ranges (from less than a second to dozens of seconds or more), but finding systematic relations in how they vary has proven difficult. Following the pioneering work of W. J. M. Levelt (1968) in binocular rivalry, previous studies have sought to formulate a relation in terms of the effect of physical parameters of the stimulus, such as image contrast in binocular rivalry. However, the link between external parameters and "stimulus strength" is not as obvious for other bistable phenomena. Here we show that systematic relations readily emerge when the mean dominance durations are examined instead as a function of "percept strength," as measured by the fraction of dominance time, and provide theoretical rationale for this observation. For three different bistable phenomena, plotting the mean dominance durations of the two percepts against the fraction of dominance time resulted in complementary curves with near-perfect symmetry around equi-dominance (the point where each percept dominates half the time). As a consequence, the alternation rate reaches a maximum at equi-dominance. We next show that the observed behavior arises naturally in simple double-well energy models and in neural competition models with cross-inhibition and input normalization. Finally, we discuss the possibility that bistable perceptual switches reflect a perceptual "exploratory" strategy, akin to foraging behavior, which leads naturally to maximal alternation rate at equi-dominance if perceptual switches come with a cost.

Beauty and the beholder: highly individual taste for abstract, but not real-world images.

How individual are visual preferences? For real-world scenes, there is high agreement in observer's preference ratings. This could be driven by visual attributes of the images but also by non-visual associations, since those are common to most individuals. To investigate this, we developed a set of novel abstract, visually diverse images. At the individual observer level both abstract and real-world images yielded robust and consistent visual preferences, and yet abstract images yielded much lower across observer agreement in preferences than did real-world images. This suggests that visual preferences are typically driven by the semantic content of stimuli, and that shared semantic interpretations then lead to shared preferences. Further experiments showed that highly individual preferences can nevertheless emerge also for real-world scenes, in contexts which de-emphasize their semantic associations.

Balance between noise and adaptation in competition models of perceptual bistability.

Perceptual bistability occurs when a physical stimulus gives rise to two distinct interpretations that alternate irregularly. Noise and adaptation processes are two possible mechanisms for switching in neuronal competition models that describe the alternating behaviors. Either of these processes, if strong enough, could alone cause the alternations in dominance. We examined their relative role in producing alternations by studying models where by smoothly varying the parameters, one can change the rhythmogenesis mechanism from being adaptation-driven to noise-driven. In consideration of the experimental constraints on the statistics of the alternations (mean and shape of the dominance duration distribution and correlations between successive durations) we ask whether we can rule out one of the mechanisms. We conclude that in order to comply with the observed mean of the dominance durations and their coefficient of variation, the models must operate within a balance between the noise and adaptation strength-both mechanisms are involved in producing alternations, in such a way that the system operates near the boundary between being adaptation-driven and noise-driven.

A hierarchy of temporal receptive windows in human cortex.

Real-world events unfold at different time scales and, therefore, cognitive and neuronal processes must likewise occur at different time scales. We present a novel procedure that identifies brain regions responsive to sensory information accumulated over different time scales. We measured functional magnetic resonance imaging activity while observers viewed silent films presented forward, backward, or piecewise-scrambled in time. Early visual areas (e.g., primary visual cortex and the motion-sensitive area MT+) exhibited high response reliability regardless of disruptions in temporal structure. In contrast, the reliability of responses in several higher brain areas, including the superior temporal sulcus (STS), precuneus, posterior lateral sulcus (LS), temporal parietal junction (TPJ), and frontal eye field (FEF), was affected by information accumulated over longer time scales. These regions showed highly reproducible responses for repeated forward, but not for backward or piecewise-scrambled presentations. Moreover, these regions exhibited marked differences in temporal characteristics, with LS, TPJ, and FEF responses depending on information accumulated over longer durations (approximately 36 s) than STS and precuneus (approximately 12 s). We conclude that, similar to the known cortical hierarchy of spatial receptive fields, there is a hierarchy of progressively longer temporal receptive windows in the human brain.

Bi-stable depth ordering of superimposed moving gratings.

Ambiguous stimuli with two distinct interpretations give rise to perceptual alternations between them. During prolonged viewing of transparently moving gratings, observers report periods of perceiving one grating in front of the other, alternating with periods of the reverse depth ordering. We measured the percepts' dominance times to study the effect of depth cues (wavelength, duty cycle, and speed) on the perceived depth ordering. The grating with shorter wavelength, lower duty cycle, or higher speed was perceived as being behind the other for a fraction of time larger than one half. The fraction of time spent perceiving each grating as behind changed gradually as a function of the parameters. The fraction of dominance depended on the ratio between the gratings' wavelengths, not on their absolute sizes. The wavelength ratio had a stronger effect on perceived depth than that of duty cycle or speed and could override stereoscopic disparity cues. Similar results were obtained with superimposed moving surfaces of random dots. The findings are interpreted in terms of their relation to statistical properties of natural surfaces and provide evidence that the fraction of dominance of each percept represents the likelihood that it corresponds to the true interpretation of the underlying scene.

Mechanisms for Frequency Control in Neuronal Competition Models.

We investigate analytically a firing rate model for a two-population network based on mutual inhibition and slow negative feedback in the form of spike frequency adaptation. Both neuronal populations receive external constant input whose strength determines the system's dynamical state-a steady state of identical activity levels or periodic oscillations or a winner-take-all state of bistability. We prove that oscillations appear in the system through supercritical Hopf bifurcations and that they are antiphase. The period of oscillations depends on the input strength in a nonmonotonic fashion, and we show that the increasing branch of the period versus input curve corresponds to a release mechanism and the decreasing branch to an escape mechanism. In the limiting case of infinitely slow feedback we characterize the conditions for release, escape, and occurrence of the winner-take-all behavior. Some extensions of the model are also discussed.

Noise-induced alternations in an attractor network model of perceptual bistability.

When a stimulus supports two distinct interpretations, perception alternates in an irregular manner between them. What causes the bistable perceptual switches remains an open question. Most existing models assume that switches arise from a slow fatiguing process, such as adaptation or synaptic depression. We develop a new, attractor-based framework in which alternations are induced by noise and are absent without it. Our model goes beyond previous energy-based conceptualizations of perceptual bistability by constructing a neurally plausible attractor model that is implemented in both firing rate mean-field and spiking cell-based networks. The model accounts for known properties of bistable perceptual phenomena, most notably the increase in alternation rate with stimulation strength observed in binocular rivalry. Furthermore, it makes a novel prediction about the effect of changing stimulus strength on the activity levels of the dominant and suppressed neural populations, a prediction that could be tested with functional MRI or electrophysiological recordings. The neural architecture derived from the energy-based model readily generalizes to several competing populations, providing a natural extension for multistability phenomena.

Brain areas selective for both observed and executed movements.

When observing a particular movement a subset of movement-selective visual and visuomotor neurons are active in the observer's brain, forming a representation of the observed movement. Similarly, when executing a movement a subset of movement-selective motor and visuomotor neurons are active, forming a representation of the executed movement. In this study we used an fMRI-adaptation protocol to assess cortical response selectivity to observed and executed movements simultaneously. Subjects freely played the rock-paper-scissors game against a videotaped opponent, sometimes repeatedly observing or executing the same movement on subsequent trials. Numerous brain areas exhibited adaptation (repetition suppression) during either repeated observations or repeated executions of the same movement. A subset of areas exhibited an overlap of both effects, containing neurons with selective responses for both executed and observed movements. We describe the function of these movement representation areas in the context of the human mirror system, which is expected to respond selectively to both observed and executed movements.

Dynamical characteristics common to neuronal competition models.

Models implementing neuronal competition by reciprocally inhibitory populations are widely used to characterize bistable phenomena such as binocular rivalry. We find common dynamical behavior in several models of this general type, which differ in their architecture in the form of their gain functions, and in how they implement the slow process that underlies alternating dominance. We focus on examining the effect of the input strength on the rate (and existence) of oscillations. In spite of their differences, all considered models possess similar qualitative features, some of which we report here for the first time. Experimentally, dominance durations have been reported to decrease monotonically with increasing stimulus strength (such as Levelt's "Proposition IV"). The models predict this behavior; however, they also predict that at a lower range of input strength dominance durations increase with increasing stimulus strength. The nonmonotonic dependency of duration on stimulus strength is common to both deterministic and stochastic models. We conclude that additional experimental tests of Levelt's Proposition IV are needed to reconcile models and perception.

Cluster-based analysis of FMRI data.

We propose a method for the statistical analysis of fMRI data that tests cluster units rather than voxel units for activation. The advantages of this analysis over previous ones are both conceptual and statistical. Recognizing that the fundamental units of interest are the spatially contiguous clusters of voxels that are activated together, we set out to approximate these cluster units from the data by a clustering algorithm especially tailored for fMRI data. Testing the cluster units has a two-fold statistical advantage over testing each voxel separately: the signal to noise ratio within the unit tested is higher, and the number of hypotheses tests compared is smaller. We suggest controlling FDR on clusters, i.e., the proportion of clusters rejected erroneously out of all clusters rejected and explain the meaning of controlling this error rate. We introduce the powerful adaptive procedure to control the FDR on clusters. We apply our cluster-based analysis (CBA) to both an event-related and a block design fMRI vision experiment and demonstrate its increased power over voxel-by-voxel analysis in these examples as well as in simulations.

Rapid detection of salient regions: evidence from apparent motion.

Most studies that have used Kanizsa-type illusory figures to investigate perceptual completion have treated the crisp bounding illusory contours (ICs) and the enclosed region as nondissociable stimulus attributes. However, there is evidence that enclosed "salient regions" (SRs; Stanley & Rubin, 2003) are detected even in cases when bounding ICs are not perceptually completed. Here we used apparent motion (AM) to test whether SRs are detected in the absence of crisp bounding ICs. Kanizsa-type stimuli were modified in ways that eliminated the bounding ICs, but the clear impression of an enclosed region remained. SR stimuli were embedded in an array of like inducers. On successive frames, the inducers in the array rotated in a way that resulted in translation of the enclosed region. Four speeds of translation were tested. Observers performed a two-alternative forced-choice task on the direction of translation. Perceptually completed SRs produced robust AM whether they were bound by crisp ICs or not-observer performance was as good and, in certain cases, even better for SRs with no bounding ICs. We interpret these findings within a theoretical framework that makes a distinction between region-based and contour-based segmentation processes that operate in concert to achieve segmentation of the visual scene.

Differences in real and illusory shape perception revealed by backward masking.

Illusory contours (ICs) are thought to be a result of processes involved in the perceptual recovery of occluded surfaces. Here, we investigate the relationship between real and illusory contour perception using a shape discrimination task and backward masking paradigm. ICs can mask other ICs when times between mask onset and stimulus onset, or SOAs, are very long ( approximately 300 ms), but real contours (RCs) are not similarly effective. Masking is absent for RC masks at perceptually salient contrasts, as well as for those with contrast lowered to match the perceived brightness of the illusory surface. We also find that RCs are not masked at long SOAs, either by ICs or by other RCs. Finally, the masking seen between ICs can occur for different sizes of target and mask. The cross-size masking would not be expected if the masking were at a level sensitive to retinal contour location. The late masking therefore may be related to a higher level of processing of shape categories and surfaces, the level at which shapes defined by ICs and RCs are differentially represented.