Controlling the Flow of Distracting Information in Working Memory.

Visual working memory (WM) must maintain relevant information, despite the constant influx of both relevant and irrelevant information. Attentional control mechanisms help determine which of this new information gets access to our capacity-limited WM system. Previous work has treated attentional control as a monolithic process-either distractors capture attention or they are suppressed. Here, we provide evidence that attentional capture may instead be broken down into at least two distinct subcomponent processes: (1) Spatial capture, which refers to when spatial attention shifts towards the location of irrelevant stimuli and (2) item-based capture, which refers to when item-based WM representations of irrelevant stimuli are formed. To dissociate these two subcomponent processes of attentional capture, we utilized a series of electroencephalography components that track WM maintenance (contralateral delay activity), suppression (distractor positivity), item individuation (N2pc), and spatial attention (lateralized alpha power). We show that new, relevant information (i.e., a task-relevant distractor) triggers both spatial and item-based capture. Irrelevant distractors, however, only trigger spatial capture from which ongoing WM representations can recover more easily. This fractionation of attentional capture into distinct subcomponent processes provides a refined framework for understanding how distracting stimuli affect attention and WM.

Spatially Guided Distractor Suppression during Visual Search.

Past work has demonstrated that active suppression of salient distractors is a critical part of visual selection. Evidence for goal-driven suppression includes below-baseline visual encoding at the position of salient distractors (Gaspelin and Luck, 2018) and neural signals such as the distractor positivity (Pd) that track how many distractors are presented in a given hemifield (Feldmann-Wüstefeld and Vogel, 2019). One basic question regarding distractor suppression is whether it is inherently spatial or nonspatial in character. Indeed, past work has shown that distractors evoke both spatial (Theeuwes, 1992) and nonspatial forms of interference (Folk and Remington, 1998), motivating a direct examination of whether space is integral to goal-driven distractor suppression. Here, we use behavioral and EEG data from adult humans (male and female) to provide clear evidence for a spatial gradient of suppression surrounding salient singleton distractors. Replicating past work, both reaction time and neural indices of target selection improved monotonically as the distance between target and distractor increased. Importantly, these target selection effects were paralleled by a monotonic decline in the amplitude of the Pd, an electrophysiological index of distractor suppression. Moreover, multivariate analyses revealed spatially selective activity in the θ-band that tracked the position of the target and, critically, revealed suppressed activity at spatial channels centered on distractor positions. Thus, goal-driven selection of relevant over irrelevant information benefits from a spatial gradient of suppression surrounding salient distractors.SIGNIFICANCE STATEMENT Past work has shown that distractor suppression is an important part of goal-driven attentional selection, but has not yet revealed whether suppression is spatially directed. Using behavioral data, event-related potentials (ERPs) of the EEG signal [N2pc and distractor positivity (Pd) component], as well as a multivariate model of EEG data [channel tuning functions (CTF)], we show that suppression-related neural activity increases monotonically as the distance between targets and distractors decreases, and that spatially-selective activity in the θ-band reveals depressed activity in spatial channels that index distractor positions. Thus, we provide robust evidence for spatially-guided distractor suppression, a result that has important implications for models of goal-driven attentional control.

Neural measures of working memory in a bilateral change detection task.

The change detection task is a widely used paradigm to examine visual working memory processes. Participants memorize a set of items and then, try to detect changes in the set after a retention period. The negative slow wave (NSW) and contralateral delay activity (CDA) are event-related potentials in the EEG signal that are commonly used in change detection tasks to track working memory load, as both increase with the number of items maintained in working memory (set size). While the CDA was argued to more purely reflect the memory-specific neural activity than the NSW, it also requires a lateralized design and attention shifts prior to memoranda onset, imposing more restrictions on the task than the NSW. The present study proposes a novel change detection task in which both CDA and NSW can be measured at the same time. Memory items were presented bilaterally, but their distribution in the left and right hemifield varied, inducing a target imbalance or "net load." NSW increased with set size, whereas CDA increased with net load. In addition, a multivariate linear classifier was able to decode the set size and net load from the EEG signal. CDA, NSW, and decoding accuracy predicted an individual's working memory capacity. In line with the notion of a bilateral advantage in working memory, accuracy, and CDA data suggest that participants tended to encode items relatively balanced. In sum, this novel change detection task offers a basis to make use of converging neural measures of working memory in a comprehensive paradigm.

Perturbing Neural Representations of Working Memory with Task-irrelevant Interruption.

Working memory maintains information so that it can be used in complex cognitive tasks. A key challenge for this system is to maintain relevant information in the face of task-irrelevant perturbations. Across two experiments, we investigated the impact of task-irrelevant interruptions on neural representations of working memory. We recorded EEG activity in humans while they performed a working memory task. On a subset of trials, we interrupted participants with salient but task-irrelevant objects. To track the impact of these task-irrelevant interruptions on neural representations of working memory, we measured two well-characterized, temporally sensitive EEG markers that reflect active, prioritized working memory representations: the contralateral delay activity and lateralized alpha power (8-12 Hz). After interruption, we found that contralateral delay activity amplitude momentarily sustained but was gone by the end of the trial. Lateralized alpha power was immediately influenced by the interrupters but recovered by the end of the trial. This suggests that dissociable neural processes contribute to the maintenance of working memory information and that brief irrelevant onsets disrupt two distinct online aspects of working memory. In addition, we found that task expectancy modulated the timing and magnitude of how these two neural signals responded to task-irrelevant interruptions, suggesting that the brain's response to task-irrelevant interruption is shaped by task context.

Alpha-band Activity Tracks the Zoom Lens of Attention.

Voluntary control over spatial attention has been likened to the operation of a zoom lens, such that processing quality declines as the size of the attended region increases, with a gradient of performance that peaks at the center of the selected area. Although concurrent changes in activity in visual regions suggest that zoom lens adjustments influence perceptual stages of processing, extant work has not distinguished between changes in the spatial selectivity of attention-driven neural activity and baseline shift of activity that can increase mean levels of activity without changes in selectivity. Here, we distinguished between these alternatives by measuring EEG activity in humans to track preparatory changes in alpha activity that indexed the precise topography of attention across the possible target positions. We observed increased spatial selectivity in alpha activity when observers voluntarily directed attention toward a narrower region of space, a pattern that was mirrored in target discrimination accuracy. Thus, alpha activity tracks both the centroid and spatial extent of covert spatial attention before the onset of the target display, lending support to the hypothesis that narrowing the zoom lens of attention shapes the initial encoding of sensory information.

Failed Suppression of Salient Stimuli Precedes Behavioral Errors.

Our visual system is constantly confronted with more information than it can process. To deal with the limited capacity, attention allows us to enhance relevant information and suppress irrelevant information. Particularly, the suppression of salient irrelevant stimuli has shown to be important as it prevents attention to be captured and thus attentional resources to be wasted. This study aimed at directly connecting failures to suppress distraction with a neural marker of suppression, the distractor positivity (Pd). We measured participants' EEG signal while they performed a visual search task in which they had to report a digit inside a shape target while ignoring distractors, one of which could be a salient color singleton. Reports of target digits served as a behavioral index of enhancement, and reports of color distractor digits served as a behavioral index of failed suppression, each measured against reports of neutral distractor digits serving as a baseline. Participants reported the target identity more often than any distractor identity. The singleton identity was reported least often, suggesting suppression of the singleton below baseline. Suppression of salient stimuli was absent in the beginning and then increased throughout the experiment. When the singleton identity was reported, the Pd was observed in a later time window, suggesting that behavioral errors were preceded by failed suppression. Our results provide evidence for the signal suppression hypothesis that states salient items have to be actively suppressed to avoid attentional capture. Our results also provide direct evidence that the Pd is reflecting such active suppression.

Statistical regularities induce spatial as well as feature-specific suppression.

[Correction Notice: An Erratum for this article was reported in Vol 45(10) of Journal of Experimental Psychology: Human Perception and Performance (see record 2019-57445-001). In the article, Figure 1 was an older version and has been updated. All versions of this article have been corrected.] We are constantly extracting regularities from the visual environment to optimize attentional orienting. Here we examine the phenomenon that recurrent presentation of distractors in a specific location leads to its attentional suppression. Specifically, we address the question whether suppression is specific to the spatial regularities of distractors or also extends to visual features bearing statistical regularities. To that end, we used a visual search task with two high-probability locations, each showing one of two distractor types more often than the other. At these high-probability locations, target processing was impaired and attentional capture by either distractor was reduced, consistent with feature-unspecific spatial suppression. However, suppression was more facilitated when the distractor feature was presented at the high-probability location that matched its features, suggesting feature-specific suppression. Interestingly, feature-unspecific spatial suppression only spread between locations when distractors varied within a feature dimension (e.g., red and green) but not when they varied across feature dimensions (e.g., red and square). Our findings thus demonstrate a joint influence of implicitly learned spatial and feature regularities on attention and reveal how the visual system can benefit from complex statistical regularities. (PsycINFO Database Record (c) 2019 APA, all rights reserved).

Neural Evidence for the Contribution of Active Suppression During Working Memory Filtering.

In order to efficiently process incoming visual information, selective attention acts as a filter that enhances relevant and suppresses irrelevant information. In this study, we used an event-related potential (ERP) approach with systematic lateralization to investigate enhancement and suppression during encoding of information into visual working memory (WM) separately. We used a change detection task in which observers had to memorize some items while ignoring other items. We found that the to-be-ignored items elicited a PD component in the ERP, suggesting that irrelevant information is actively suppressed from WM. The PD amplitude increased with distractor load and decreased with the ability to group distractors according to Gestalt principles. This suggests that the PD can be used as an indicator of how efficiently items can be suppressed from entering WM. Furthermore, while lateral memory-targets elicited a "traditional" CDA (starting ~300 ms), lateral memory-distractors elicited a sustained positivity contralateral to memory-distractors (CDAp, starting ~400 ms). In sum the results suggest that inhibition of irrelevant information is an important factor for efficient WM and is reflected in spontaneous (PD) and sustained suppression (CDAp).

Contralateral Delay Activity Indexes Working Memory Storage, Not the Current Focus of Spatial Attention.

Contralateral delay activity (CDA) has long been argued to track the number of items stored in visual working memory (WM). Recently, however, Berggren and Eimer [Berggren, N., & Eimer, M. Does contralateral delay activity reflect working memory storage or the current focus of spatial attention within visual working memory? Journal of Cognitive Neuroscience, 28, 2003-2020, 2016] proposed the alternative hypothesis that the CDA tracks the current focus of spatial attention instead of WM storage. This hypothesis was based on the finding that, when two successive arrays of memoranda were placed in opposite hemifields, CDA amplitude was primarily determined by the position and number of items in the second display, not the total memory load across both displays. Here, we considered the alternative interpretation that participants dropped the first array from WM when they encoded the second array because the format of the probe display was spatially incompatible with the initial sample display. In this case, even if the CDA indexes active storage rather than spatial attention, CDA activity would be determined by the second array. We tested this idea by directly manipulating the spatial compatibility of sample and probe displays. With spatially incompatible displays, we replicated Berggren and Eimer's findings. However, with spatially compatible displays, we found clear evidence that CDA activity tracked the full storage load across both arrays, in line with a WM storage account of CDA activity. We propose that expectations of display compatibility influenced whether participants viewed the arrays as parts of a single extended event or two independent episodes. Thus, these findings raise interesting new questions about how event boundaries may shape the interplay between passive and active representations of task-relevant information.

Reduced visual attention in heterogeneous textures is reflected in occipital alpha and theta band activity.

Increasing context heterogeneity has been found to reduce attention deployment towards an embedded target item. Heterogeneity in visual search tasks is typically induced by segmenting the background into several perceptual groups. In the present study, however, context heterogeneity was induced by varying whole-field heterogeneity, i.e., the degree of distractor variability within the entire context. This allowed us to (i) more gradually vary context heterogeneity, and (ii) investigate attention deployment on a whole-field scale. Results showed that both search performance and amplitude of the N2pc (lateralized ERP; posterior contralateral negativity in the N2 range) monotonically decreased with increasing context heterogeneity, which confirmed that there was less efficient attention deployment for more heterogeneous contexts. The amplitude of the bilateral N2 exhibited a U-shaped function, suggesting global perception for the lowest and highest levels of heterogeneity, but local processing for intermediate heterogeneity levels. Independent component analyses revealed an occipital ERP-contributing effective source cluster that may reflect stimulus representations on a saliency map. With increasing heterogeneity, these sources exhibited more theta band activity for distractors and less theta band activity for targets. Alpha band activity of a second component cluster varied with heterogeneity level, and low-theta/delta activity of a third source cluster distinguished target presence versus absence. In sum, our results suggest that independent brain sources contributed to the differential processing of heterogeneous versus homogeneous contexts.

Selection history alters attentional filter settings persistently and beyond top-down control.

Visual selective attention is known to be guided by stimulus-based (bottom-up) and goal-oriented (top-down) control mechanisms. Recent work has pointed out that selection history (i.e., the bias to prioritize items that have been previously attended) can result in a learning experience that also has a substantial impact on subsequent attention guidance. The present study examined to what extent goal-oriented top-down control mechanisms interact with an observer's individual selection history in guiding attention. Selection history was manipulated in a categorization task in a between-subjects design, where participants learned that either color or shape was the response-relevant dimension. The impact of this experience was assessed in a compound visual search task with an additional color distractor. Top-down preparation for each search trial was enabled by a pretrial task cue (Experiment 1) or a fixed, predictable trial sequence (Experiment 2). Reaction times and ERPs served as indicators of attention deployment. Results showed that attention was captured by the color distractor when participants had learned that color predicted the correct response in the categorization learning task, suggesting that a bias for predictive stimulus features had developed. The possibility to prepare for the search task reduced the bias, but could not entirely overrule this selection history effect. In Experiment 3, both tasks were performed in separate sessions, and the bias still persisted. These results indicate that selection history considerably shapes selective attention and continues to do so persistently even when the task allowed for high top-down control.

Intertrial priming due to distractor repetition is eliminated in homogeneous contexts.

Targets are found more easily in a visual search task when their feature is repeatedly presented, an effect known as intertrial priming. Recent findings suggest that priming of distractors can also improve search performance by facilitated suppression of repeated distractor features. The efficacy of intertrial priming for targets can be potentiated by the expectancy of a specific target feature; systematic repetition shows larger intertrial priming than random repetition. For distractors, the underlying mechanism is less clear. We used the systematic lateralization approach to disentangle target- and distractor-related processing with subcomponents of the N2pc. We found no modulation of the NT component, which reflects prioritization of target processing. The ND component, which reflects attentional capture by irrelevant stimuli, however, showed intertrial priming: ND monotonically decreased with repetition of a distractor color, but only if a specific distractor feature was expected, and if the context induced a search that was vulnerable to attentional capture. These observations suggest that distractor priming only improves visual search if volitional control is relatively high. The results also suggest that intertrial priming for distractors is due to decreased attentional capture by repeatedly presented distractors, whereas target processing remains unaffected.

Rewarded visual items capture attention only in heterogeneous contexts.

Reward is known to affect visual search performance. Rewarding targets can increase search performance, whereas rewarding distractors can decrease search performance. We used subcomponents of the N2pc in the event-related EEG, the NT (target negativity) and ND /PD (distractor negativity/positivity), in a visual search task to disentangle target and distractor processing related to reward. The visual search task comprised homogeneous and heterogeneous contexts in which a target and a colored distractor were embedded. After each correct trial, participants were given a monetary reward that depended on the color of the distractor. We found longer response times for displays with high-reward distractors compared to displays with low-reward distractors, indicating reward-induced interference, however, only for heterogeneous contexts. The NT component, indicative of attention deployment to the target, showed that target selection was impaired by high-reward distractors, regardless of the context homogeneity. Processing of distractors was not affected by reward in homogeneous contexts. In heterogeneous contexts, however, high-reward distractors were more likely to capture attention (ND ) and required more effort to be suppressed (PD ) than low-reward distractors. In sum the results showed that, despite the fact that target selection is impaired by high-reward distractors in both homogeneous and heterogeneous background contexts, high-reward distractors capture attention only in scenarios that foster attentional capture.

Target discrimination delays attentional benefit for grouped contexts: An ERP study.

Searching a target was shown to be facilitated when embedded in a homogeneous compared to a heterogeneous context. This study investigated extended attention deployment after visual search for a target that observers had to discriminate in contexts of varying heterogeneity. Results showed more efficient attention deployment towards targets in homogeneous contexts as evident in higher accuracy and larger N2pc amplitudes than in random contexts. RTs to subsequently presented probes were shorter and a larger posterior positivity was observed in the ERP when probes were presented at the target location compared to other locations. This on-target advantage was larger for homogeneous contexts at ISIs of 90ms, but not for shorter (30ms) or longer (150ms) ISIs. These results show that visual search tasks induce an accelerated attention deployment after homogeneous contexts, however with a delay after target discrimination compared to less demanding target detection.

You see what you have learned. Evidence for an interrelation of associative learning and visual selective attention.

Besides visual salience and observers' current intention, prior learning experience may influence deployment of visual attention. Associative learning models postulate that observers pay more attention to stimuli previously experienced as reliable predictors of specific outcomes. To investigate the impact of learning experience on deployment of attention, we combined an associative learning task with a visual search task and measured event-related potentials of the EEG as neural markers of attention deployment. In the learning task, participants categorized stimuli varying in color/shape with only one dimension being predictive of category membership. In the search task, participants searched a shape target while disregarding irrelevant color distractors. Behavioral results showed that color distractors impaired performance to a greater degree when color rather than shape was predictive in the learning task. Neurophysiological results show that the amplified distraction was due to differential attention deployment (N2pc). Experiment 2 showed that when color was predictive for learning, color distractors captured more attention in the search task (ND component) and more suppression of color distractor was required (PD component). The present results thus demonstrate that priority in visual attention is biased toward predictive stimuli, which allows learning experience to shape selection. We also show that learning experience can overrule strong top-down control (blocked tasks, Experiment 3) and that learning experience has a longer-term effect on attention deployment (tasks on two successive days, Experiment 4).

Action Planning Mediates Guidance of Visual Attention from Working Memory.

Visual search is impaired when a salient task-irrelevant stimulus is presented together with the target. Recent research has shown that this attentional capture effect is enhanced when the salient stimulus matches working memory (WM) content, arguing in favor of attention guidance from WM. Visual attention was also shown to be closely coupled with action planning. Preparing a movement renders action-relevant perceptual dimensions more salient and thus increases search efficiency for stimuli sharing that dimension. The present study aimed at revealing common underlying mechanisms for selective attention, WM, and action planning. Participants both prepared a specific movement (grasping or pointing) and memorized a color hue. Before the movement was executed towards an object of the memorized color, a visual search task (additional singleton) was performed. Results showed that distraction from target was more pronounced when the additional singleton had a memorized color. This WM-guided attention deployment was more pronounced when participants prepared a grasping movement. We argue that preparing a grasping movement mediates attention guidance from WM content by enhancing representations of memory content that matches the distractor shape (i.e., circles), thus encouraging attentional capture by circle distractors of the memorized color. We conclude that templates for visual search, action planning, and WM compete for resources and thus cause interferences.

Stimulus homogeneity enhances implicit learning: evidence from contextual cueing.

Visual search for a target object is faster if the target is embedded in a repeatedly presented invariant configuration of distractors ('contextual cueing'). It has also been shown that the homogeneity of a context affects the efficiency of visual search: targets receive prioritized processing when presented in a homogeneous context compared to a heterogeneous context, presumably due to grouping processes at early stages of visual processing. The present study investigated in three Experiments whether context homogeneity also affects contextual cueing. In Experiment 1, context homogeneity varied on three levels of the task-relevant dimension (orientation) and contextual cueing was most pronounced for context configurations with high orientation homogeneity. When context homogeneity varied on three levels of the task-irrelevant dimension (color) and orientation homogeneity was fixed, no modulation of contextual cueing was observed: high orientation homogeneity led to large contextual cueing effects (Experiment 2) and low orientation homogeneity led to low contextual cueing effects (Experiment 3), irrespective of color homogeneity. Enhanced contextual cueing for homogeneous context configurations suggest that grouping processes do not only affect visual search but also implicit learning. We conclude that memory representation of context configurations are more easily acquired when context configurations can be processed as larger, grouped perceptual units. However, this form of implicit perceptual learning is only improved by stimulus homogeneity when stimulus homogeneity facilitates grouping processes on a dimension that is currently relevant in the task.

Textures shape the attentional focus: evidence from exogenous and endogenous cueing.

The spatial cueing paradigm (Posner Quarterly Journal of Experimental Psychology 32:3-25, 1980) has often been used to investigate the time course of the deployment of visual attention in space. In a series of eight experiments we investigated whether spatial cues would not only enhance processing of stimuli presented at cued locations, but also enhance processing of the entire texture in which the stimuli were presented. Results showed highest accuracy for responses to stimuli presented at cued locations, a replication of the traditional cueing effect (Posner 1980). Additionally, stimuli presented at uncued locations were responded to with higher accuracy when they were presented inside the same texture as the cued location, as compared with stimuli presented outside the texture with the cued location. To investigate this texture advantage for both automatic and voluntary attention deployment, exogenous and endogenous cues were used. The texture advantage was observed for short interstimulus intervals (ISIs) of 50 and 100 ms for exogenous cues and for a longer ISI of 200 ms for endogenous cues. These findings indicate that the arrangement of task-irrelevant visual stimuli also can have a large impact on the cueing effect. This suggests that visual spatial attention spreads texture-wise across the visual field. Control experiments revealed that the homogeneity within texture elements contributes most to the effect but that the texture advantage is a function of both orientation contrast at the texture border and homogeneity within texture elements.

Context heterogeneity has a sustained impact on attention deployment: behavioral and electrophysiological evidence.

In visual search, similar nearby stimuli can be grouped and thus enhance processing of an embedded target. The aim of the present study was to examine the time course of attention deployment after a brief presentation of stimulus arrays of different heterogeneity. Targets in less heterogeneous, grouped contexts yielded higher accuracy and larger N2pc amplitudes than targets in more heterogeneous, random contexts, indicating more efficient selection in the former. Subsequently presented probes yielded shorter reaction times and a larger posterior positivity when presented at the target location. This advantage was more pronounced after grouped compared to random contexts at the shorter compared to the longer interstimulus interval. The results show that less heterogeneous contexts that allow for grouping not only enhance processing of stimuli within that context, but have a sustained effect on visual attention.

Context homogeneity facilitates both distractor inhibition and target enhancement.

Homogeneous contexts were shown to result in prioritized processing of embedded targets compared to heterogeneous contexts (Duncan & Humphreys, 1989). The present experiment used behavioral and ERP measures to examine whether context homogeneity affects both enhancing relevant information and inhibiting irrelevant in contexts of varying homogeneity. Targets and distractors were presented laterally or on the vertical midline which allowed disentangling target- and distractor-related activity in the lateralized ERP (Hickey, diLollo, & McDonald, 2009). In homogeneous contexts, targets elicited an NT component from 150 ms on and a PD component from 200 ms on, showing early attention deployment at target locations and active suppression of distractors. In heterogeneous contexts, an NT component was also found from 150 ms on and PD was found from 250 ms on, suggesting delayed suppression of the distractor. Before 250 ms, distractors in heterogeneous contexts elicited a contralateral negativity, indicating attentional capture of the distractor prior to active suppression. In sum the present results suggest that top-down control of attention is more pronounced in homogeneous than in heterogeneous contexts.