Nancy B. Carlisle

Attentional Templates in Visual Working Memory

Most theories of attention propose that we maintain attentional templates in visual working memory to control what information is selected. In the present study, we directly tested this proposal by measuring the contralateral-delay activity (CDA) of human event-related potentials during visual search tasks in which the target is cued on each trial. Here we show that the CDA can be used to measure the maintenance of attentional templates in visual working memory while processing complex visual scenes. In addition, this method allowed us to directly observe the shift from working memory to long-term memory representations controlling attention as learning occurred and experience accrued searching for the same target object. Our findings provide definitive support for several critical proposals made in theories of attention, learning, and automaticity.

Where do we store the memory representations that guide attention

During the last decade one of the most contentious and heavily studied topics in the attention literature has been the role that working memory representations play in controlling perceptual selection. The hypothesis has been advanced that to have attention select a certain perceptual input from the environment, we only need to represent that item in working memory. Here we summarize the work indicating that the relationship between what representations are maintained in working memory and what perceptual inputs are selected is not so simple. First, it appears that attentional selection is also determined by high-level task goals that mediate the relationship between working memory storage and attentional selection. Second, much of the recent work from our laboratory has focused on the role of long-term memory in controlling attentional selection. We review recent evidence supporting the proposal that working memory representations are critical during the initial configuration of attentional control settings, but that after those settings are established long-term memory representations play an important role in controlling which perceptual inputs are selected by mechanisms of attention.

Automatic and strategic effects in the guidance of attention by working memory representations.

Theories of visual attention suggest that working memory representations automatically guide attention toward memory-matching objects. Some empirical tests of this prediction have produced results consistent with working memory automatically guiding attention. However, others have shown that individuals can strategically control whether working memory representations guide visual attention. Previous studies have not independently measured automatic and strategic contributions to the interactions between working memory and attention. In this study, we used a classic manipulation of the probability of valid, neutral, and invalid cues to tease apart the nature of such interactions. This framework utilizes measures of reaction time (RT) to quantify the costs and benefits of attending to memory-matching items and infer the relative magnitudes of automatic and strategic effects. We found both costs and benefits even when the memory-matching item was no more likely to be the target than other items, indicating an automatic component of attentional guidance. However, the costs and benefits essentially doubled as the probability of a trial with a valid cue increased from 20% to 80%, demonstrating a potent strategic effect. We also show that the instructions given to participants led to a significant change in guidance distinct from the actual probability of events during the experiment. Together, these findings demonstrate that the influence of working memory representations on attention is driven by both automatic and strategic interactions.

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.

Templates for Rejection: Configuring Attention to Ignore Task-Irrelevant Features

Theories of attention are compatible with the idea that we can bias attention to avoid selecting objects that have known nontarget features. Although this may underlie several existing phenomena, the explicit guidance of attention away from known nontargets has yet to be demonstrated. Here we show that observers can use feature cues (i.e., color) to bias attention away from nontarget items during visual search. These negative cues were used to quickly instantiate a template for rejection that reliably facilitated search across the cue-to-search stimulus onset asynchronies (SOAs), although negative cues were not as potent as cues that guide attention toward target features. Furthermore, by varying the search set size we show a template for rejection is increasingly effective in facilitating search as scene complexity increases. Our findings demonstrate that knowing what not to look for can be used to configure attention to avoid certain features, complimenting what is known about setting attention to select certain target features.

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.

Reconciling conflicting electrophysiological findings on the guidance of attention by working memory

Maintaining a representation in working memory has been proposed to be sufficient for the execution of top-down attentional control. Two recent electrophysiological studies that recorded event-related potentials (ERPs) during similar paradigms have tested this proposal, but have reported contradictory findings. The goal of the present study was to reconcile these previous reports. To this end, we used the stimuli from one study (Kumar, Soto, & Humphreys, 2009) combined with the task manipulations from the other (Carlisle & Woodman, 2011b). We found that when an item matching a working memory representation was presented in a visual search array, we could use ERPs to quantify the size of the covert attention effect. When the working memory matches were consistently task-irrelevant, we observed a weak attentional bias to these items. However, when the same item indicated the location of the search target, we found that the covert attention effect was approximately four times larger. This shows that simply maintaining a representation in working memory is not equivalent to having a top-down attentional set for that item. Our findings indicate that high-level goals mediate the relationship between the contents of working memory and perceptual attention.

The effects of self-focus on attentional biases in social anxiety:An ERP study.

Cognitive theories of social anxiety disorder suggest that biased attention plays a key role in maintaining symptoms. These biases include self-focus and attention to socially threatening stimuli in the environment. The goal of this study was to utilize ERPs that are elicited by a change detection task to examine biases in selective attention (i.e., N2pc) and working memory maintenance (i.e., contralateral delay activity; CDA). Additionally, the effect of self-focus was examined using false heart rate feedback. In support of the manipulation, self-focus cues resulted in greater self-reported self-consciousness and task interference, enhanced anterior P2 amplitude and reduced SPN amplitude. Moreover, P2 amplitude for self-focus cues was correlated with reduced task performance for socially anxious subjects only. The difference in P2 amplitude between self-focus and standard cues was correlated with social anxiety independent of depression. As hypothesized, socially anxious participants (n = 20) showed early selection and maintenance of disgust faces relative to neutral faces as indicated by the N2pc and CDA components. Nonanxious controls (n = 22) did not show these biases. During self-focus cues, controls showed marginal evidence of biased selection for disgust faces, whereas socially anxious subjects showed no bias in this condition. Controls showed an ipsilateral delay activity after being cued to attend to one hemifield. Overall, this study supports early and persistent attentional bias for social threat in socially anxious individuals. Furthermore, self-focus may disrupt these biases. These findings and supplementary data are discussed in light of cognitive models of social anxiety disorder, recent empirical findings, and treatment.

The control of single-color and multiple-color visual search by attentional templates in working memory and in long-term memory

The question whether target selection in visual search can be effectively controlled by simultaneous attentional templates for multiple features is still under dispute. We investigated whether multiple-color attentional guidance is possible when target colors remain constant and can thus be represented in long-term memory but not when they change frequently and have to be held in working memory. Participants searched for one, two, or three possible target colors that were specified by cue displays at the start of each trial. In constant-color blocks, the same colors remained task-relevant throughout. In variable-color blocks, target colors changed between trials. The contralateral delay activity (CDA) to cue displays increased in amplitude as a function of color memory load in variable-color blocks, which indicates that cued target colors were held in working memory. In constant-color blocks, the CDA was much smaller, suggesting that color representations were primarily stored in long-term memory. N2pc components to targets were measured as a marker of attentional target selection. Target N2pcs were attenuated and delayed during multiple-color search, demonstrating less efficient attentional deployment to color-defined target objects relative to single-color search. Importantly, these costs were the same in constant-color and variable-color blocks. These results demonstrate that attentional guidance by multiple-feature as compared with single-feature templates is less efficient both when target features remain constant and can be represented in long-term memory and when they change across trials and therefore have to be maintained in working memory.

Visual working memory gives up attentional control early in learning: Ruling out interhemispheric cancellation

Current research suggests that we can watch visual working memory surrender the control of attention early in the process of learning to search for a specific object. This inference is based on the observation that the contralateral delay activity (CDA) rapidly decreases in amplitude across trials when subjects search for the same target object. Here, we tested the alternative explanation that the role of visual working memory does not actually decline across learning, but instead lateralized representations accumulate in both hemispheres across trials and wash out the lateralized CDA. We show that the decline in CDA amplitude occurred even when the target objects were consistently lateralized to a single visual hemifield. Our findings demonstrate that reductions in the amplitude of the CDA during learning are not simply due to the dilution of the CDA from interhemispheric cancellation.