Although storage in working memory (WM) can be tracked via measurements of ongoing neural activity, past work has shown that observers can maintain access to that information despite temporary interruptions of those neural patterns. This observation has been regarded as evidence for a neurally silent form of WM storage. Alternatively, however, unattended information could be retrieved from episodic long-term memory (eLTM) rather than being maintained in WM during the activity-silent period. Here, we tested between these possibilities by examining whether WM performance showed evidence of proactive interference (PI)—a hallmark of retrieval from eLTM—following such interruptions. Participants remembered the colors (Experiments 1–3) or locations (Experiment 4) of serially presented objects. We found PI for set sizes larger than 4, but not for smaller set sizes, suggesting that eLTM may have supported performance when WM capacity was exceeded. Critically, performance with small set sizes remained resistant to PI, even following prolonged interruptions by a challenging distractor task. Thus, we found evidence for PI-resistant memories that were maintained across likely interruptions of storage-related neural activity, an empirical pattern that implies activity-silent storage in WM.

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 before 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 toward a taxonomy of the distinct attentional processes that constrain our memories.

Multiple neural signals have been found to track the number of items stored in working memory (WM). These signals include oscillatory activity in the alpha band and slow-wave components in human EEG, both of which vary with storage loads and predict individual differences in WM capacity. However, recent evidence suggests that these two signals play distinct roles in spatial attention and item-based storage in WM. Here, we examine the hypothesis that sustained negative voltage deflections over parieto-occipital electrodes reflect the number of individuated items in WM, whereas oscillatory activity in the alpha frequency band (8–12 Hz) within the same electrodes tracks the attended positions in the visual display. We measured EEG activity while participants stored the orientation of visual elements that were either grouped by collinearity or not. This grouping manipulation altered the number of individuated items perceived while holding constant the number of locations occupied by visual stimuli. The negative slow wave tracked the number of items stored and was reduced in amplitude in the grouped condition. By contrast, oscillatory activity in the alpha frequency band tracked the number of positions occupied by the memoranda and was unaffected by perceptual grouping. Perceptual grouping, then, reduced the number of individuated representations stored in WM as reflected by the negative slow wave, whereas the location of each element was actively maintained as indicated by alpha power. These findings contribute to the emerging idea that distinct classes of EEG signals work in concert to successfully maintain on-line representations in WM.

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.

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.

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.

Current theories assume a functional role for covert attention in the maintenance of spatial information in working memory. Consistent with this view, both the locus of attention and positions stored in working memory can be decoded based on the topography of oscillatory alpha-band (8–12 Hz) activity on the scalp. Thus far, however, alpha modulation has been studied in isolation for covert attention and working memory tasks. Here, we applied an inverted spatial encoding model in combination with EEG to study the temporal dynamics of spatially specific alpha activity during a task that required observers to visually select a target location while maintaining another independently varying location in working memory. During the memory delay period, alpha-based spatial tuning functions shifted from the position stored in working memory to the covertly attended position and back again after the attention task was completed. The findings provide further evidence for a common oscillatory mechanism in both the selection and the maintenance of relevant spatial visual information and demonstrate the dynamic trade-off in prioritization between two spatial tasks.

A great deal of prior research has examined the relation between estimates of working memory and cognitive abilities. Yet, the neural mechanisms that account for these relations are still not very well understood. The current study explored whether individual differences in working memory delay activity would be a significant predictor of cognitive abilities. A large number of participants performed multiple measures of capacity, attention control, long-term memory, working memory span, and fluid intelligence, and latent variable analyses were used to examine the data. During two working memory change detection tasks, we acquired EEG data and examined the contralateral delay activity. The results demonstrated that the contralateral delay activity was significantly related to cognitive abilities, and importantly these relations were because of individual differences in both capacity and attention control. These results suggest that individual differences in working memory delay activity predict individual differences in a broad range of cognitive abilities, and this is because of both differences in the number of items that can be maintained and the ability to control access to working memory.

By the request of the authors, the following two research articles will be retracted from the Journal of Cognitive Neuroscience : 1. Anderson, D. E., Ester, E. F., Klee, D., Vogel, E. K., & Awh, E. (2014). Electrophysiological evidence for failures of item individuation in crowded visual displays. Journal of Cognitive Neuroscience , 26(10), 2298– 2309. https://dx.doi.org/10.1162/jocn_a_00649 . 2. Anderson, D. E., Bell, T. A., & Awh, E. (2012). Polymorphisms in the 5-HTTLPR gene mediate storage capacity of visual working memory. Journal of Cognitive Neuroscience , 24(5), 1069–1076. https://dx.doi. org/10.1162/jocn_a_00207 . On August 1, 2015, the Office of Research Integrity (ORI) announced a settlement agreement with David E. Anderson, the Respondent ( http://ori.hhs.gov/content/ case-summary-anderson-david ). On the basis of the Respondent’s admission and an analysis by the University of Oregon, ORI concluded that the Respondent had engaged in research misconduct by falsifying and/or fabricating data in four publications. Those publications were retracted immediately after the release of the ORI findings. Since that time, additional problems have been discovered with Article 1 above. Data points shown in Figure 8 were removed without justification and in contradiction to the analytic approach described in the methods and results. In light of this discovery and of the previous ORI findings, authors Bell and Awh no longer have confidence in the integrity of the data in Article 2. For these reasons, all authors on both articles (including the Respondent) have agreed to the retraction of Articles 1 and 2 above.

Recent studies suggest that the temporary storage of visual detail in working memory is mediated by sensory recruitment or sustained patterns of stimulus-specific activation within feature-selective regions of visual cortex. According to a strong version of this hypothesis, the relative “quality” of these patterns should determine the clarity of an individual's memory. Here, we provide a direct test of this claim. We used fMRI and a forward encoding model to characterize population-level orientation-selective responses in visual cortex while human participants held an oriented grating in memory. This analysis, which enables a precise quantitative description of multivoxel, population-level activity measured during working memory storage, revealed graded response profiles whose amplitudes were greatest for the remembered orientation and fell monotonically as the angular distance from this orientation increased. Moreover, interparticipant differences in the dispersion—but not the amplitude—of these response profiles were strongly correlated with performance on a concurrent memory recall task. These findings provide important new evidence linking the precision of sustained population-level responses in visual cortex and memory acuity.

By the request of the authors, the following two research articles will be retracted from the Journal of Cognitive Neuroscience : 1. Anderson, D. E., Ester, E. F., Klee, D., Vogel, E. K., & Awh, E. (2014). Electrophysiological evidence for failures of item individuation in crowded visual displays. Journal of Cognitive Neuroscience , 26(10), 2298– 2309. https://dx.doi.org/10.1162/jocn_a_00649 . 2. Anderson, D. E., Bell, T. A., & Awh, E. (2012). Polymorphisms in the 5-HTTLPR gene mediate storage capacity of visual working memory. Journal of Cognitive Neuroscience , 24(5), 1069–1076. https://dx.doi. org/10.1162/jocn_a_00207 . On August 1, 2015, the Office of Research Integrity (ORI) announced a settlement agreement with David E. Anderson, the Respondent ( http://ori.hhs.gov/content/ case-summary-anderson-david ). On the basis of the Respondent’s admission and an analysis by the University of Oregon, ORI concluded that the Respondent had engaged in research misconduct by falsifying and/or fabricating data in four publications. Those publications were retracted immediately after the release of the ORI findings. Since that time, additional problems have been discovered with Article 1 above. Data points shown in Figure 8 were removed without justification and in contradiction to the analytic approach described in the methods and results. In light of this discovery and of the previous ORI findings, authors Bell and Awh no longer have confidence in the integrity of the data in Article 2. For these reasons, all authors on both articles (including the Respondent) have agreed to the retraction of Articles 1 and 2 above.

We investigated the hypothesis that the covert focusing of spatial attention mediates the on-line maintenance of location information in spatial working memory. During the delay period of a spatial working-memory task, behaviorally irrelevant probe stimuli were flashed at both memorized and nonmemorized locations. Multichannel recordings of event-related potentials (ERPs) were used to assess visual processing of the probes at the different locations. Consistent with the hypothesis of attention-based rehearsal, early ERP components were enlarged in response to probes that appeared at memorized locations. These visual modulations were similar in latency and topography to those observed after explicit manipulations of spatial selective attention in a parallel experimental condition that employed an identical stimulus display.

We report an experiment that assesses the effect of variations in memory load on brain activations that mediate verbal working memory. The paradigm that forms the basis of this experiment is the “ n -back” task in which subjects must decide for each letter in a series whether it matches the one presented n items back in the series. This task is of interest because it recruits processes involved in both the storage and manipulation of information in working memory. Variations in task difficulty were accomplished by varying the value of n . As n increased, subjects showed poorer behavioral performance as well as monotonically increasing magnitudes of brain activation in a large number of sites that together have been identified with verbal working-memory processes. By contrast, there was no reliable increase in activation in sites that are unrelated to working memory. These results validate the use of parametric manipulation of task variables in neuroimaging research, and they converge with the subtraction paradigm used most often in neuroimaging. In addition, the data support a model of working memory that includes both storage and executive processes that recruit a network of brain areas, all of which are involved in task performance.

We used positron emission tomography (PET) to answer the following question: Is working memory a unitary storage system, or does it instead include different storage buffers for different kinds of information? In Experiment 1, PET measures were taken while subjects engaged in either a spatial-memory task (retain the position of three dots for 3 sec) or an object-memory task (retain the identity of two objects for 3 sec). The results manifested a striking double dissociation, as the spatial task activated only right-hemisphere regions, whereas the object task activated primarily left-hemisphere regions. The spatial (right-hemisphere) regions included occipital, parietal, and prefrontal areas, while the object (left-hemisphere) regions included inferotemporal and parietal areas. Experiment 2 was similar to Experiment 1 except that the stimuli and trial events were identical for the spatial and object tasks; whether spatial or object memory was required was manipulated by instructions. The PET results once more showed a double dissociation, as the spatial task activated primarily right-hemisphere regions (again including occipital, parietal and prefrontal areas), whereas the object task activated only left-hemisphere regions (again including inferotemporal and parietal areas). Experiment 3 was a strictly behavioral study, which produced another double dissociation. It used the same tasks as Experiment 2, and showed that a variation in spatial similarity affected performance in the spatial but not the object task, whereas a variation in shape similarity affected performance in the object but not the spatial task. Taken together, the results of the three experiments clearly imply that different working-memory buffers are used for storing spatial and object information.