Biology subjects

Awh, E.

Publications and source records attributed to Awh, E..

At least 19 recordsLinked to original sources

bioRxiv2025

Distributed and drifting signals for working memory load in human cortex

Increasing working memory (WM) load incurs behavioral costs, and whether the neural constraints on behavioral costs are localized (i.e., emanating from the intraparietal sulcus) or distributed across cortex remains an active area of debate. In a pre-registered fMRI experiment, 12 humans (12 scanner-hours each) performed a visual WM task with varying memory load (0-4 items). We replicated a localized, load-dependent increase in univariate BOLD activity in parietal cortex. However, we also observed both systematic increases and decreases in univariate activity with load across the visual hierarchy. Importantly, multivariate activation patterns encoded WM load regardless of the direction of the univariate effect, arguing against a restricted locus of load signals in parietal cortex. Finally, we observed representational drift in activity patterns encoding memory load across scanning sessions. Our results suggest a distributed code for memory load that may be continually refined over time to support more efficient information storage.

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bioRxiv2025

Contributions from Long-Term Memory Explain Superior Visual Working Memory Performance with Meaningful Objects

Visual working memory (WM) capacity has been claimed to be larger for meaningful objects than for simple features, possibly because richer semantic representations enhance the distinctiveness of stored items. However, prior demonstrations typically compared trial-unique meaningful objects with a small set of repeated simple features. This design confounds meaningfulness with proactive interference (PI), such that PI is minimal for trial-unique objects but substantial for repeated features. As a result, superior performance for meaningful objects may reflect contributions from episodic long-term memory (LTM) rather than expanded WM capacity. To test this, Experiment 1 measured WM for repeated colors, repeated meaningful objects, and trial-unique meaningful objects. The advantage for objects over colors was replicated in the trial-unique condition, but eliminated for repeated objects that equated PI across stimulus types. Hierarchical Bayesian dual-process modeling revealed that the trial-unique advantage reflected stronger familiarity signals, whereas recollection remained stable across stimulus types. Experiment 2 assessed WM storage directly using contralateral delay activity (CDA), an electrophysiological marker of the number of items stored. Although trial-unique objects again yielded behavioral advantages, CDA activity across increasing set sizes revealed a common slope and plateau for trial-unique meaningful objects and repeated colors. The CDA difference between stimulus types was additive and did not vary with set size, providing no evidence for an increased number of stored items. These findings demonstrate that previously reported advantages for meaningful objects primarily reflect reduced PI and enhanced LTM familiarity. When PI is equated, WM storage limits for simple and meaningful stimuli are equivalent. Significance StatementWorking memory provides the mental workspace that underlies reasoning, learning, and everyday decision making, yet its capacity is sharply limited. Previous studies suggested that meaningful, real-world objects are remembered better, raising the possibility that knowledge can expand this known capacity limit. However, many designs confound working memory with long-term familiarity. Here, equating proactive interference removed the behavioral advantage for meaningful items. A neural marker of active storage showed additive differences between stimulus types that did not vary with load, indicating no increase in the number of stored items. These findings identify interference, rather than expanded storage, as the source of the reported advantage and offer practical guidance for future experimental design and theories of memory limits.

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bioRxiv2025

Mechanisms of Output Gating for visual working memory

Working memory (WM) tasks often require comparing remembered items to test displays, but little is known about how people selectively remove irrelevant information at test. Across three experiments, we used contralateral delay activity (CDA) to track WM load and examine selective removal. In Experiment 1, CDA amplitudes increased with set size even when only one item was probed, suggesting minimal removal based on spatial location. Experiment 2 ruled out spatial grouping by presenting items sequentially in the same location, yet more items were retained for larger set size. In Experiment 3, however, when items belonged to distinct mnemonic categories, CDA amplitudes at test were reduced, consistent with selective removal based on category relevance. Additionally, P3 old-new effects showed that decision speed and strength were influenced by the number of items maintained. Together, these results suggest that people selectively remove WM contents based on categorical relevance, not spatial cues, enabling more efficient memory-based decisions.

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bioRxiv2025

Population-Level Activity Dissociates Preparatory Overt from Covert Attention

The neural signatures of preparing overt eye movements and directing covert spatial attention overlap as they recruit the same brain areas. Yet, these neural signatures are dissociable at the single-cell level: Specific cells within visuo-oculomotor areas are exclusively involved in motor preparation or covert attention. Nevertheless, it has been proposed that many cells in visuo-oculomotor areas are involved in both motor preparation and covert attention, and consequently their neural signatures should functionally overlap to a large degree. Here, we directly tested this proposal: we combined human (both sexes) EEG with sensitive decoding techniques to investigate whether the neural signatures of preparatory overt and covert attention are dissociable across large-scale neuronal populations. We found that neural decoding reliably discerned whether overt or covert attention was shifted well before saccade initiation. Further, inverted encoding modeling revealed earlier and sharper spatially-tuned activity in preparatory overt than in covert attention. We then asked whether preparatory overt attention achieved sharper spatially-tuned activity by using more-of-the-same covert attention, or by recruiting additional spatially selective neural processing. Cross-decoding results demonstrated that preparatory overt attention recruited at least one additional, frontal process. This additional spatially selective process emerged early and likely reflects motor preparation or predictive remapping. To summarize, we found that the neural signatures of overt and covert attention overlap, yet diverge rapidly, in part because overt attention employs an additional spatially selective neural process. Extending beyond a dissociation on the single-cell level, our findings demonstrate that population-level neural activity dissociates preparatory overt from covert attention. Significance statementThe world provides much more visual input than the brain can process simultaneously. Spatial visual attention allows for the selective processing of only the most important parts of this input. Spatial attention shifts either overtly (with an eye movement) or covertly (without an accompanying eye movement). The neural signatures underlying these types of spatial attention have long thought to overlap to a large degree. This strong overlap was thought to only break down at the level of single neurons. In our EEG study, we instead demonstrate that large populations of neurons dissociate overt from covert attention. Our results show that the neural signatures of overt and covert attention differ on a much broader scale than previously thought.

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bioRxiv2025

Differential representations of spatial location by aperiodic and alpha oscillatory activity in working memory

Decades of research have shown working memory (WM) relies on sustained pre-frontal cortical activity and visual extrastriate activity, particularly in the alpha (8-12 Hz) frequency range. This alpha activity tracks the spatial location of WM items, even when spatial position is task-irrelevant and there is no stimulus currently being presented. Traditional analyses of putative oscillations using bandpass filters, however, conflate oscillations with non-oscillatory aperiodic activity. Here, we reanalyzed seven different human electroencephalography (EEG) visual WM datasets to test the hypothesis that aperiodic activity-which is thought to reflect the relative contributions of excitatory and inhibitory drive-plays a distinct role in visual WM from true alpha oscillations. To do this, we developed a novel, time-resolved spectral parameterization approach to disentangle oscillations from aperiodic activity during WM encoding and maintenance. Across all seven tasks, totaling 112 participants, we captured the representation of spatial location from total alpha power using an inverted encoding model (IEM), replicating traditional analyses. We then trained separate IEMs to estimate the strength of spatial location representation from aperiodic-adjusted alpha (reflecting just the oscillatory component) and aperiodic activity, and find that IEM performance improves for aperiodic-adjusted alpha compared to total alpha power that blends the two signals. We also discover a novel role for aperiodic activity, where IEM performance trained on aperiodic activity is highest during stimulus presentation, but not during the WM maintenance period. Our results emphasize the importance of controlling for aperiodic activity when studying neural oscillations while uncovering a novel functional role for aperiodic activity in the encoding of visual WM information. Significance statementWorking memory is a crucial component of cognition, yet its neural mechanisms are not fully understood. Research shows that alpha activity - presumed to reflect neural oscillations - tracks the location of items we hold in memory. However, these analyses assume that all alpha power is oscillatory, even though oscillations are mixed with non-oscillatory, aperiodic activity that may be physiologically and functionally distinct. Here, we use a novel analytical approach for separating alpha oscillations and aperiodic activity dynamically across time. Our results reveal distinct roles for each in human visual working memory: aperiodic activity encodes the spatial location of information whereas alpha oscillations maintain the location of that information.

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bioRxiv2022

Overlapping Neural Representations for Dynamic Visual Imagery and Stationary Storage in Spatial Working Memory

Representations in working memory need to be flexibly transformed to adapt to our dynamic environment and variable task demands. Recent work has demonstrated that activity in the alpha frequency band enables precise decoding of visual information during both perception and sustained storage in working memory. Extant work, however, has focused exclusively on the representation of static visual images. Here we used EEG recordings to examine whether alpha-band activity supports the dynamic transformation of representations in spatial working memory using an imagery task that required the active shifting of a stored position to a new position. In line with recent findings, a common format of alpha-band activity precisely tracked both the initial position stored in working memory as well as the transformed position, with the latter emerging approximately 800-1200 ms following an auditory cue to "rotate" to a new position. Moreover, the time course of this transformation of alpha activity predicted between-subject differences in manual reaction time to indicate the new position (Experiment 1), as well as within-subject variations in saccade latency in a speeded version of the task (Experiment 2). Finally, cross-training analyses revealed robust generalization of alpha-band reconstruction of working memory contents before and after mental transformation. These findings demonstrate that alpha activity tracks dynamic transformations of representations in spatial working memory, and that the format of this activity is preserved across the initial and transformed memory representations. These findings highlight a new approach for measuring voluntary shifts in online memory representations and show common representational formats during dynamic mental imagery and the maintenance of static representations in working memory.

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bioRxiv2021

Shared long-term and short-term memory representational formats in occipital and parietal cortex

Current theories propose that the short-term retention of information in working memory (WM) and the recall of information from long-term memory (LTM) are supported by overlapping neural mechanisms in occipital and parietal cortex. Both are thought to rely on reinstating patterns of sensory activity evoked by the perception of the remembered item. However, the extent of the shared representations between WM and LTM are unclear, and it is unknown how WM and LTM representations may differ across cortical regions. We designed a spatial memory task that allowed us to directly compare the representations of remembered spatial information in WM and LTM. Critically, we carefully matched the precision of behavioral responses in these tasks. We used fMRI and multivariate pattern analyses to examine representations in (1) retinotopic cortex and (2) lateral parietal cortex (LPC) regions previously implicated in LTM. We show that visual memories were represented in a sensory-like code in both tasks across retinotopic regions in occipital and parietal cortex. LPC regions also encoded remembered locations in both WM and LTM, but in a format that differed from the sensory-evoked activity. These results suggest a striking correspondence in the format of WM and LTM representations across occipital and parietal cortex. On the other hand, we show that activity patterns in nearly all parietal regions, but not occipital regions, contained information that could discriminate between WM trials and LTM trials. Our data provide new evidence for theories of memory systems and the representation of mnemonic content.

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bioRxiv2021

Pupillometry signatures of sustained attention and working memory

There exists an intricate relationship between attention and working memory. Recent work has further established that attention and working memory fluctuate synchronously, by tightly interleaving sustained attention and working memory tasks. This work has raised many open questions about physiological signatures underlying these behavioral fluctuations. Across two experiments, we explore pupil dynamics using real-time triggering in conjunction with an interleaved sustained attention and working memory task. In Experiment 1, we use behavioral real-time triggering and replicate recent findings from our lab (deBettencourt et al., 2019) that sustained attention fluctuates concurrently with the number of items maintained in working memory. Furthermore, highly attentive moments, detected via behavior, also exhibited larger pupil sizes. In Experiment 2, we develop a novel real-time pupil triggering technique to track pupil size fluctuations in real time and trigger working memory probes. We reveal that this pupil triggering procedure elicits differences in sustained attention, as indexed by response time. These experiments reflect methodological advances in real-time triggering and further characterize an important biomarker of sustained attention.

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bioRxiv2020

Sustained attention and spatial attention distinctly influence long-term memory encoding

Our attention is critically important for what we remember. Prior measures of the relationship between attention and memory, however, have largely treated "attention" as a monolith. Here, across three experiments, we provide evidence for two dissociable aspects of attention that influence encoding into long-term memory. Using spatial cues together with a sensitive continuous report procedure, we find that long-term memory response error is affected by both trial-by-trial fluctuations of sustained attention and prioritization via covert spatial attention. Furthermore, using multivariate analyses of EEG, we track both sustained attention and spatial attention prior to stimulus onset. Intriguingly, even during moments of low sustained attention, there is no decline in the representation of the spatially attended location, showing that these two aspects of attention have robust but independent effects on long term memory encoding. Finally, sustained and spatial attention predicted distinct variance in long-term memory performance across individuals. That is, the relationship between attention and long-term memory suggests a composite model, wherein distinct attentional subcomponents influence encoding into long-term memory. These results point towards a taxonomy of the distinct attentional processes that constrain our memories.

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bioRxiv2020

Controlling the flow of interruption 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 sub-component 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 sub-component processes of attentional capture, we utilized a series of EEG components that track WM maintenance (contralateral delay activity), suppression (distractor positivity), item individuation (N2pc), and spatial attention (lateralized alpha power). We show that relevant interrupters trigger both spatial and item-based capture, which means that they undermine WM maintenance more. Irrelevant interrupters, however, only trigger spatial capture from which ongoing WM representations can recover more easily. This fractionation of attentional capture into distinct sub-component processes provides a framework by which the fate of ongoing WM processes after interruption can be explained.

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bioRxiv2020

Spatial attention enhances the spatial selectivity of population codes during the first wave of visual processing

Covert spatial attention has a variety of effects on the responses of individual neurons. However, relatively little is known about the net effect of these changes on sensory population codes, even though perception ultimately depends on population activity. Here, we measured the electroencephalogram (EEG) in human observers (male and female), and isolated stimulus-evoked activity that was phase-locked to the onset of attended and ignored visual stimuli. Using an encoding model, we reconstructed spatially selective population tuning functions from the pattern of stimulus-evoked activity across the scalp. Our EEG-based approach allowed us to measure very early visually evoked responses occurring ~100 ms after stimulus onset. In Experiment 1, we found that covert attention increased the amplitude of spatially tuned population responses at this early stage of sensory processing. In Experiment 2, we parametrically varied stimulus contrast to test how this effect scaled with stimulus contrast. We found that the effect of attention on the amplitude of spatially tuned responses increased with stimulus contrast, and was well-described by an increase in response gain (i.e., a multiplicative scaling of the population response). Together, our results show that attention increases the gain of spatial population codes during the first wave of visual processing. Significance StatementWe know relatively little about how attention improves population codes, even though perception is thought to critically depend on population activity. In this study, we used an encoding-model approach to test how attention modulates the spatial tuning of stimulus-evoked population responses measured with EEG. We found that attention multiplicatively scales the amplitude of spatially tuned population responses. Furthermore, this effect was present within 100 ms of stimulus onset. Thus, our results show that attention improves spatial population codes by increasing their gain at this early stage of processing.

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bioRxiv2020

Salient distractors induce a gradient of attentional suppression to cope with spatial and nonspatial filtering costs

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, 2015) and neural signals such as the Pd that track the position and number of distractors in the visual field (Feldmann-Wustefeld 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 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 theta 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.

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bioRxiv2020

Predicting cognitive abilities across individuals using sparse EEG connectivity

Human brains share a broadly similar functional organization with consequential individual variation. This duality in brain function has primarily been observed when using techniques that consider the spatial organization of the brain, such as MRI. Here, we ask whether these common and unique signals of cognition are also present in temporally sensitive, but spatially insensitive, neural signals. To address this question, we compiled EEG data from individuals performing multiple working memory tasks at two different data-collection sites (ns = 171 and 165). Results revealed that EEG connectivity patterns were stable within individuals and unique across individuals. Furthermore, models based on these connectivity patterns generalized across datasets to predict participants working memory capacity and general fluid intelligence. Thus, EEG connectivity provides a signature of working memory and fluid intelligence in humans and a new framework for characterizing individual differences in cognitive abilities.

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bioRxiv2020

Multivariate analysis reveals a generalizable human electrophysiological signature of working memory load

Working memory (WM) is an online memory system that is critical for holding information in a rapidly accessible state during ongoing cognitive processing. Thus, there is strong value in methods that provide a temporally-resolved index of WM load. While univariate EEG signals have been identified that vary with WM load, recent advances in multivariate analytic approaches suggest that there may be rich sources of information that do not generate reliable univariate signatures. Here, using data from 4 published studies (n = 286 and >250,000 trials), we demonstrate that multivariate analysis of EEG voltage topography provides a sensitive index of the number of items stored in WM that generalizes to novel human observers. Moreover, multivariate load detection ("mvLoad") can provide robust information at the single-trial level, exceeding the sensitivity of extant univariate approaches. We show that this method tracks WM load in a manner that is (1) independent of the spatial position of the memoranda, (2) precise enough to differentiate item-by-item increments in the number of stored items, (3) generalizable across distinct tasks and stimulus displays and (4) correlated with individual differences in WM behavior. Thus, this approach provides a powerful complement to univariate analytic approaches, enabling temporally-resolved tracking of online memory storage in humans.

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bioRxiv2019

Covert spatial attention speeds target individuation

Covert spatial attention has long been thought to speed visual processing. Psychophysics studies have shown that target information accrues faster at attended locations than at unattended locations. However, with behavioral evidence alone, it is difficult to determine whether attention speeds visual processing of the target, or subsequent post-perceptual stages of processing (e.g. decision making and response selection). Moreover, while many studies have shown that that attention can boost the amplitude of visually-evoked neural responses, no effect has been observed on the latency of those neural responses. Here, we offer new evidence that may reconcile the neural and behavioral findings. Our study focused on the N2pc, an EEG marker of visual selection that has been linked with object individuation - the formation of an object representation that is distinct from the background and from other objects. In two experiments, we manipulated whether or not covert attention was precisely deployed to the location of an impending search target. We found that the target-evoked N2pc onset approximately 20 ms earlier when the target location was cued than when it was not cued. Thus, although attention may not speed the earliest stages of sensory processing, attention does speed the critical transition between raw sensory encoding and the formation of individuated object representations. Significance StatementCovert spatial attention improves processing at attended locations. Past behavioral studies have shown that information about visual targets accrues faster at attended than at unattended locations. However, it has remained unclear whether attention speeds perceptual analysis or subsequent post-perceptual stages of processing. Here we present robust evidence that attention speeds the N2pc, an electrophysiological signal that indexes the formation of individuated object representations. Our findings show that attention speeds a relatively early stage of perceptual processing, while also elucidating the specific perceptual process that is speeded.

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bioRxiv2019

Multivariate analysis of EEG activity indexes contingent and non-contingent attentional capture

It is well known that salient yet irrelevant singleton can capture attention, even when this is inconsistent with the current goals of the observer (Theeuwes, 1992; 2010). Others however have claimed that capture is critically contingent on the goals of the observer: Capture is strongly modulated (or even eliminated) when the irrelevant singleton does not match the target-defining properties (Folk, Remington, & Johnston, 1992). There has been a long-standing debate on whether attentional capture can be explained by goal-driven and/or stimulus-driven accounts. Here, we shed further light on this phenomenon by using EEG activity (raw EEG and alpha power) to provide a time-resolved index of attentional orienting. Participants searched for a target defined by a pre-specified color. The search display was preceded by a singleton cue that either matched the color of the upcoming target (contingent cues), or that appeared in an irrelevant color (non-contingent cues). Multivariate analysis of raw EEG and alpha power revealed preferential tuning to the location of both contingent and non-contingent cues, with a stronger bias towards contingent than non-contingent cues. The time course of these effects, however, depended on the neural signal. Raw EEG data revealed attentional orienting towards the cue early on in the trial (>156 ms), while alpha power revealed sustained spatial selection in the cued locations at a later moment in the trial (>250 ms). Moreover, while raw EEG showed stronger capture by contingent cues during this early time window, the advantage for contingent cues arose during a later time window in alpha band activity. Thus, our findings suggest that raw EEG activity and alpha-band power tap into distinct neural processes that index movements of covert spatial attention. Both signals provide clear neural evidence that both contingent and non-contingent cues can capture attention, and that this process is robustly shaped by the target-defining properties in the current block of trials.

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bioRxiv2019

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. In this series of experiments, we investigated the impact of task-irrelevant interruptions on neural representations of working memory. We recorded electroencephalogram (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 (CDA) and lateralized alpha power (8-12hz). Following interruption, we found that CDA 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. Additionally, we found that task expectancy modulated the timing and magnitude of how these two neural signals responded to task-irrelevant interruptions, suggesting that the brains response to task-irrelevant interruption is shaped by task context. The distinct time courses of and influence of task context on these two neural signatures of working memory have many interesting theoretical implications about how information is actively maintained in working memory.\n\nSignificance statementWorking memory plays a central role in intelligent behaviors because it actively maintains relevant information that is easily accessible and manipulatable. In everyday life, we are often interrupted while performing such complex cognitive tasks. Therefore, understanding how working memory responds to and overcomes momentary task-irrelevant interruptions is critical for us to understand how complex cognition works. Here, we unveil how two distinct neural signatures of working memory respond to task-irrelevant interruptions by recording electroencephalogram activity in humans. Our findings raise long-standing theoretical questions about how different neural and cognitive processes contribute to the maintenance of information in working memory.

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bioRxiv2018

Item-specific delay activity demonstrates concurrent storage of multiple items in working memory

AbstractA longstanding view holds that information is maintained in working memory (WM) via persistent neural activity that encodes the content of WM. Recent work, however, has challenged the view that all items stored in WM are actively maintained. Instead, \"activity-silent\" models propose that items can be maintained in WM without the need for persistent neural activity, raising the possibility that only a subset of items - perhaps just a single item - may be actively represented at a given time. While past studies have successfully decoded multiple items stored in WM, these studies cannot rule out an active switching account in which only a single item is actively represented at a time. Here, we directly tested whether multiple representations can be held concurrently in an active state. We tracked spatial representations in WM using alpha-band (8-12 Hz) activity, which encodes spatial positions held in WM. Human observers (male and female) remembered one or two positions over a short delay while we recorded EEG. Using a spatial encoding model, we reconstructed stimulus-specific working memory representations (channel tuning functions, CTFs) from the scalp distribution of alphaband power. Consistent with past work, we found the selectivity of spatial CTFs was lower when two items were stored than when one item was stored. Critically, data-driven simulations revealed that the selectivity of spatial representations in the two-item condition could not be explained by models restricting storage to a single item at a time. Thus, our findings provide robust evidence for the concurrent storage of multiple items in visual working memory.\n\nAuthor SummaryWorking memory (WM) is a mental workspace where we temporarily hold information \"online\" in pursuit of our current goals. However, recent activity-silent models of WM have challenged the view that all items are held in an \"online\" state, instead proposing that only a subset of representations in WM - perhaps just one item - are represented by persistent activity at a time. To directly test a single-item model of persistent activity, we used a spatial encoding model to read out the strength of two representations from alpha-band (8-12 Hz) power in the human EEG signal. We provide direct evidence that both locations were maintained concurrently, ruling out the possibility that declines in stimulus-specific activity are due to storing one of two items in an activity-silent state.

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