Min-Suk Kang

Nonhuman primate event-related potentials indexing covert shifts of attention

A half-centurys worth of research has established the existence of numerous event-related potential components measuring different cognitive operations in humans including the selection of stimuli by covert attention mechanisms. Surprisingly, it is unknown whether nonhuman primates exhibit homologous electrophysiological signatures of selective visual processing while viewing complex scenes. We used an electrophysiological technique with macaque monkeys analogous to procedures for recording scalp event-related potentials from humans and found that monkeys exhibit short-latency visual components sensitive to sensory processing demands and lateralizations related to shifting of covert attention similar to the human N2pc component. These findings begin to bridge the gap between the disparate literatures by using electrophysiological measurements to study the deployment of visual attention in the brains of humans and nonhuman primates.

Semantic Analysis Does Not Occur in the Absence of Awareness Induced by Interocular Suppression

It has been intensely debated whether visual stimuli are processed to the point of semantic analysis in the absence of awareness. In the present study, we measured the extent to which the meaning of a stimulus was registered using the N400 component of human event-related potentials (ERPs), a highly sensitive index of the semantic mismatch between a stimulus and the context in which it is presented. Observers judged the semantic relatedness of a context and target word while ERPs were recorded under continuous flash suppression (Experiments 1 and 2) and binocular rivalry (Experiment 3). Finally, we parametrically manipulated the visibility of the target word by increasing the contrast between the target word and the suppressive stimulus presented to the other eye (Experiment 4). We found that the amplitude of the N400 was attenuated with increasing suppression depth and was absent whenever the observers could not discriminate the meaning of suppressed words. We discuss these findings in the context of single-process models of consciousness, which can account for a large body of empirical evidence obtained from visual masking, attentional manipulations, and, now, interocular suppression paradigms.

The benefit of forgetting.

Recent research using change-detection tasks has shown that a directed-forgetting cue, indicating that a subset of the information stored in memory can be forgotten, significantly benefits the other information stored in visual working memory. How do these directed-forgetting cues aid the memory representations that are retained? We addressed this question in the present study by using a recall paradigm to measure the nature of the retained memory representations. Our results demonstrated that a directed-forgetting cue leads to higher-fidelity representations of the remaining items and a lower probability of dropping these representations from memory. Next, we showed that this is made possible by the to-be-forgotten item being expelled from visual working memory following the cue, allowing maintenance mechanisms to be focused on only the items that remain in visual working memory. Thus, the present findings show that cues to forget benefit the remaining information in visual working memory by fundamentally improving their quality relative to conditions in which just as many items are encoded but no cue is provided.

What causes alternations in dominance during binocular rivalry

Several mechanisms have been proposed to account for perceptual alterations during binocular rivalry, including neural adaptation and neural noise. However, the importance of neural adaptation for producing perceptual alterations has been challenged in several articles (Y.-J. Kim, Grabowecky, & Suzuki, 2006; Moreno-Bote, Rinzel, & Rubin, 2007). We devised an “online” adaptation procedure to reexamine the role of adaptation in binocular rivalry. Periods of adaptation inserted into rivalry observation periods parametrically alter the dynamics of rivalry such that increased adaptation duration decreases dominance duration, which cannot be accounted for by neural noise. Analysis of the average dominance durations and their variance (coefficient of variation) provides evidence for an increasingly important role of noise in rivalry alternations as a given dominance period continues in time, consistent with recent computational models.

The Effect of Visual Search Efficiency on Response Preparation Neurophysiological Evidence for Discrete Flow

Most models assume that response time (RT) comprises the time required for successive processing stages, but they disagree about whether information is transmitted continuously or discretely between stages. We tested these alternative hypotheses by measuring when movement-related activity began in the frontal eye field (FEF) of macaque monkeys performing visual search. Previous work showed that RT was longer when visual neurons in FEF took longer to select the target, a finding consistent with prolonged perceptual processing during less efficient search. We now report that the buildup of saccadic movement-related activity in FEF is delayed in inefficient visual search. Variability in the delay of movement-related activity accounted for the difference in RT between search conditions and for the variability of RT within conditions. These findings provide neurophysiological support for the hypothesis that information is transmitted discretely between perceptual and response stages of processing during visual search.

Visual working memory contaminates perception

Indirect evidence suggests that the contents of visual working memory may be maintained within sensory areas early in the visual hierarchy. We tested this possibility using a well-studied motion repulsion phenomenon in which perception of one direction of motion is distorted when another direction of motion is viewed simultaneously. We found that observers misperceived the actual direction of motion of a single motion stimulus if, while viewing that stimulus, they were holding a different motion direction in visual working memory. Control experiments showed that none of a variety of alternative explanations could account for this repulsion effect induced by working memory. Our findings provide compelling evidence that visual working memory representations directly interact with the same neural mechanisms as those involved in processing basic sensory events.

Size matters: a study of binocular rivalry dynamics.

W.J.M. Levelt systematized the influence of stimulus strength on binocular rivalry dynamics in several formal propositions. His counterintuitive 2nd proposition states that mean dominance duration of one eyes stimulus depends not on the strength of that stimulus but, instead, on the strength of the stimulus viewed by the other eye. Some studies have reported results consistent with this proposition but others have found violations of the proposition. This paper examines the dynamics of binocular rivalry by changing the size of rival stimuli and the tracking instructions during rivalry tracking periods in which the contrasts of the two rival stimuli are varied independently. Levelts 2nd proposition was validated when those stimuli were large, but it was violated when the rival stimuli were small, suggesting that the dynamics of binocular rivalry are spatiotemporal in nature. A simple energy model with coupling among neighboring areas of rivalry can account for these findings. Other dynamics depending on the size of rival stimuli are discussed.

Event-related potentials elicited by errors during the stop-signal task. II: Human effector specific error responses

Although previous research with human and nonhuman primates has examined the neural correlates of performance monitoring, discrepancies in methodology have limited our ability to make cross-species generalizations. One major obstacle arises from the use of different behavioral responses and tasks across different primate species. Specifically, it is unknown whether performance-monitoring mechanisms rely on different neural circuitry in tasks requiring oculomotor vs. skeletomotor responses. Here, we show that the human error-related negativity (ERN) elicited by a saccadic eye-movement response relative to a manual response differs in several critical ways. The human saccadic ERN exhibits a prolonged duration, a broader frontomedial voltage distribution, and different neural source estimates than the manual ERN in exactly the same stop-signal task. The human saccadic error positivity (Pe) exhibited a frontomedial voltage distribution with estimated electrical sources in supplementary motor area and rostral anterior cingulate cortex for saccadic responses, whereas the manual Pe showed a posterior scalp distribution and potential origins in the superior parietal lobule. These findings constrain models of the cognitive mechanisms indexed by the ERN/Pe complex. Moreover, by paralleling work with nonhuman primates performing the same saccadic stop-signal task (Godlove et al. 2011), we demonstrate a cross-species homology of error event-related potentials (ERPs) and lay the groundwork for definitively localizing the neural sources of performance-monitoring ERPs.

An Integrated Framework of Spatiotemporal Dynamics of Binocular Rivalry

Fluctuations in perceptual dominance during binocular rivalry exhibit several hallmark characteristics. First, dominance switches are not periodic but, instead, stochastic: perception changes unpredictably. Second, despite being stochastic, average durations of rivalry dominance vary dependent on the strength of the rival stimuli: variations in contrast, luminance, or spatial frequency produce predictable changes in average dominance durations and, hence, in alternation rate. Third, perceptual switches originate locally and spread globally over time, sometimes as traveling waves of dominance: rivalry transitions are spatiotemporal events. This essay (1) reviews recent advances in our understanding of the bases of these three hallmark characteristics of binocular rivalry dynamics and (2) provides an integrated framework to account for those dynamics using cooperative and competitive spatial interactions among local neural circuits distributed over the visual fields retinotopic map. We close with speculations about how that framework might incorporate top-down influences on rivalry dynamics.

Distinct neural mechanisms for spatially lateralized and spatially global visual working memory representations

Visual working memory (VWM) allows humans to actively maintain a limited amount of information. Whereas previous electrophysiological studies have found that lateralized event-related potentials (ERPs) track the maintenance of information in VWM, recent imaging experiments have shown that spatially global representations can be read out using the activity across the visual cortex. The goal of the present study was to determine whether both lateralized and spatially global electrophysiological signatures coexist. We first show that it is possible to simultaneously measure lateralized ERPs that track the number of items held in VWM from one visual hemfield and parietooccipital α (8-12 Hz) power over both hemispheres indexing spatially global VWM representations. Next, we replicated our findings and went on to show that this bilateral parietooccipital α power as well as the contralaterally biased ERP correlate of VWM carries a signal that can be used to decode the identity of the representations stored in VWM. Our findings not only unify observations across electrophysiology and imaging techniques but also suggest that ERPs and α-band oscillations index different neural mechanisms that map on to lateralized and spatially global representations, respectively.