A single pulse of TMS (spTMS) during the delay period of a double serial retrocuing working-memory task can briefly rescue decodability of an unprioritized memory item (UMI). This physiological phenomenon, which is paralleled in behavior by involuntary retrieval of the UMI, is carried by the beta frequency band, implicating beta-band dynamics in priority coding in working memory. We decomposed EEG data from 12 participants performing double serial retrocuing with concurrent delivery of spTMS using Spatially distributed PhAse Coupling Extraction. This procedure decomposes the scalp-level signal into a set of discrete coupled oscillators, each with a component strength that can vary over time. The decomposition revealed a diversity of low-frequency components, a subset of them strengthening with the onset of the task, and the majority declining in strength across the trial, as well as within each delay period. Results with spTMS revealed no evidence that it works by activating previously “silent” sources; instead, it had the effect of modulating ongoing activity, specifically by exaggerating the within-delay decrease in strength of posterior beta components. Furthermore, the magnitude of the effect of spTMS on the loading strength of a posterior beta component correlated with the disruptive effect of spTMS on performance, a pattern also seen when analyses were restricted to trials with “UMI-lure” memory probes. Rather than reflecting the “activation” of a putatively “activity silent” UMI, these results implicate beta-band dynamics in a mechanism that distinguishes prioritized from unprioritized, and suggest that the effect of spTMS is to disrupt this code.

The ability to prioritize among contents in working memory (WM) is critical for successful control of thought and behavior. Recent work has demonstrated that prioritization in WM can be implemented by representing different states of priority in different representational formats. Here, we explored the mechanisms underlying WM prioritization by simulating the double serial retrocuing task with recurrent neural networks. Visualization of stimulus representational dynamics using principal component analysis revealed that the network represented trial context (order of presentation) and priority via different mechanisms. Ordinal context, a stable property lasting the duration of the trial, was accomplished by segregating representations into orthogonal subspaces. Priority, which changed multiple times during a trial, was accomplished by separating representations into different strata within each subspace. We assessed the generality of these mechanisms by applying dimensionality reduction and multiclass decoding to fMRI and EEG data sets and found that priority and context are represented differently along the dorsal visual stream and that behavioral performance is sensitive to trial-by-trial variability of priority coding, but not context coding.

Adaptive behavior relies on the selection and prioritization of relevant sensory inputs from the external environment as well as from among internal sensory representations held in working memory. Recent behavioral evidence suggests that the classic distinction between voluntary (goal-driven) and involuntary (stimulus-driven) influences over attentional allocation also applies to the selection of internal representations held in working memory. In the current EEG study, we set out to investigate the neural dynamics associated with the competition between voluntary and involuntary control over the focus of attention in visual working memory. We show that when voluntary and involuntary factors compete for the internal focus of attention, prioritization of the appropriate item is delayed—as reflected both in delayed gaze biases that track internal selection and in delayed neural beta (15–25 Hz) dynamics that track the planning for the upcoming memory-guided manual action. We further show how this competition is paralleled—possibly resolved—by an increase in frontal midline theta (4–8 Hz) activity that, moreover, predicts the speed of ensuing memory-guided behavior. Finally, because theta increased following retrocues that effectively reduced working-memory load, our data unveil how frontal theta activity during internal attentional focusing tracks demands on cognitive control over and above working-memory load. Together, these data yield new insight into the neural dynamics that govern the focus of attention in visual working memory, and disentangle the contributions of frontal midline theta activity to the processes of control versus retention in working memory.

Humans can construct rich subjective experience even when no information is available in the external world. Here, we investigated the neural representation of purely internally generated stimulus-like information during visual working memory. Participants performed delayed recall of oriented gratings embedded in noise with varying contrast during fMRI scanning. Their trialwise behavioral responses provided an estimate of their mental representation of the to-be-reported orientation. We used multivariate inverted encoding models to reconstruct the neural representations of orientation in reference to the response. We found that response orientation could be successfully reconstructed from activity in early visual cortex, even on 0% contrast trials when no orientation information was actually presented, suggesting the existence of a purely internally generated neural code in early visual cortex. In addition, cross-generalization and multidimensional scaling analyses demonstrated that information derived from internal sources was represented differently from typical working memory representations, which receive influences from both external and internal sources. Similar results were also observed in intraparietal sulcus, with slightly different cross-generalization patterns. These results suggest a potential mechanism for how externally driven and internally generated information is maintained in working memory.

What mechanisms underlie the prioritization of neural representations of visually perceived information to guide behavior? We assessed the dynamics whereby attention biases competition for representation of visual stimuli by enhancing representations of relevant information and suppressing the irrelevant. Multivariate pattern analysis (MVPA) classifiers were trained to discriminate patterns of fMRI activity associated with each of three stimuli, within several predefined ROIs. Participants performed a change-detection task wherein two of three presented items flashed at 1 Hz, one to each side of central fixation. Both flashing stimuli would unpredictably change state, but participants covertly counted the number of changes only for the cued item. In the ventral occipito-temporal ROI, MVPA evidence (a proxy for representational fidelity) was dynamically enhanced for attended stimuli and suppressed for unattended stimuli, consistent with a mechanism of biased competition between stimulus representations. Frontal and parietal ROIs displayed a qualitatively distinct, more “source-like” profile, wherein MVPA evidence for only the attended stimulus could be observed above baseline levels. To assess how attentional modulation of ventral occipito-temporal representations might relate to signals originating in the frontal and/or parietal ROIs, we analyzed informational connectivity (IC), which indexes time-varying covariation between regional levels of MVPA evidence. Parietal-posterior IC was elevated during the task, but did not differ for cued versus uncued items. Frontal-posterior IC, in contrast, was sensitive to an item's priority status. Thus, although regions of frontal and parietal cortex act as sources of top–down attentional control, their precise functions likely differ.

In the field of neuroscience, despite the fact that the proportion of peer-reviewed publications authored by women has increased in recent decades, the proportion of citations of women-led publications has not seen a commensurate increase: In five broad-scope journals, citations of papers first- and/or last-authored by women have been shown to be fewer than would be expected if gender was not a factor in citation decisions [Dworkin, J. D., Linn, K. A., Teich, E. G., Zurn, P., Shinohara, R. T., & Bassett, D. S. The extent and drivers of gender imbalance in neuroscience reference lists. Nature Neuroscience , 23 , 918–926, 2020]. Given the important implications that such underrepresentation may have on the careers of women researchers, it is important to determine whether this same trend is true in subdisciplines of the field, where interventions might be more targeted. Here, we report the results of an extension of the analyses carried out by Dworkin et al. (2020) to citation patterns in the Journal of Cognitive Neuroscience . The results indicate that the underrepresentation of women-led publications in reference sections is also characteristic of papers published in Journal of Cognitive Neuroscience over the past decade. Furthermore, this pattern of citation imbalances is present regardless of author gender, implicating systemic factors. These results contribute to the growing body of evidence that intentional action is needed to address inequities in the way that we carry out and communicate our science.

Although humans can hold multiple items in mind simultaneously, the contents of working memory (WM) can be selectively prioritized to guide future behavior. We explored whether the “same-object” benefits in visual processing may also be observed in visual WM. fMRI data were collected while participants performed a multistep serial retrocuing task in which they first viewed two 2-D objects (coherently moving colored dots). During retention, an initial relevance cue then indicated whether only the first or only the second object (“object-relevant”), or only the color of both objects or only their direction of motion would be relevant for the remainder of the trial (“feature-relevant”). On “object-relevant” trials, the ensuing priority cues selected either one of the features (“color” or “direction”) bound to the relevance-cued object, whereas on “feature-relevant” trials, the priority cues selected one of the two relevance-cued features. Using multivariate inverted encoding models, we found a same-object benefit on object-relevant trials in occipitotemporal regions: On feature-relevant trials, the first priority cue triggered a strengthening of the neural representation of the cued feature and a concomitant weakening to baseline of the uncued feature, whereas on object-relevant trials, the cued item remained active but did not increase in strength and the uncued item weakened but remained significantly elevated throughout the delay period. Although the stimulus-specific representation in frontoparietal regions was weak and uneven, these regions closely tracked the higher order information of which stimulus category was relevant for behavior throughout the trial, suggesting an important role in controlling the prioritization of information in visual WM.

Numerous studies have demonstrated that visual STM (VSTM) and attention are tightly linked processes that share a number of neuroanatomical substrates. Here, we used repetitive TMS (rTMS) along with simultaneous EEG to examine the causal relationship between intraparietal sulcus functioning and performance on tasks of attention and VSTM. Participants performed two tasks in which they were required to attend to or remember colored items over a brief interval, with 10-Hz rTMS applied on some of the trials. Although no overall behavioral changes were observed across either task, rTMS did affect individual performance on both the attention and VSTM tasks in a manner that was predicted by individual differences in baseline performance. Furthermore, rTMS also affected ongoing oscillations in the alpha and beta bands, and these changes were related to the observed change in behavioral performance. The results reveal a causal relationship between intraparietal sulcus activity and tasks measuring both visual attention and VSTM.

Many aspects of perception and cognition are supported by activity in neural populations that are tuned to different stimulus features (e.g., orientation, spatial location, color). Goal-directed behavior, such as sustained attention, requires a mechanism for the selective prioritization of contextually appropriate representations. A candidate mechanism of sustained spatial attention is neural activity in the alpha band (8–13 Hz), whose power in the human EEG covaries with the focus of covert attention. Here, we applied an inverted encoding model to assess whether spatially selective neural responses could be recovered from the topography of alpha-band oscillations during spatial attention. Participants were cued to covertly attend to one of six spatial locations arranged concentrically around fixation while EEG was recorded. A linear classifier applied to EEG data during sustained attention demonstrated successful classification of the attended location from the topography of alpha power, although not from other frequency bands. We next sought to reconstruct the focus of spatial attention over time by applying inverted encoding models to the topography of alpha power and phase. Alpha power, but not phase, allowed for robust reconstructions of the specific attended location beginning around 450 msec postcue, an onset earlier than previous reports. These results demonstrate that posterior alpha-band oscillations can be used to track activity in feature-selective neural populations with high temporal precision during the deployment of covert spatial attention.

For decades it has been assumed that sustained, elevated neural activity—the so-called active trace—is the neural correlate of the short-term retention of information. However, a recent fMRI study has suggested that this activity may be more related to attention than to retention. Specifically, a multivariate pattern analysis failed to find evidence that information that was outside the focus of attention, but nonetheless in STM, was retained in an active state. Here, we replicate and extend this finding by querying the neural signatures of attended versus unattended information within STM with electroencephalograpy (EEG), a method sensitive to oscillatory neural activity to which the previous fMRI study was insensitive. We demonstrate that in the delay-period EEG activity, there is information only about memory items that are also in the focus of attention. Information about items outside the focus of attention is not detectable. This result converges with the fMRI findings to suggest that, contrary to conventional wisdom, an active memory trace may be unnecessary for the short-term retention of information.

The concept of the “mnemonic scotoma,” a spatially circumscribed region of working memory impairment produced by unilateral lesions of the PFC, is central to the view that PFC is critical for the short-term retention of information. Presented here, however, are previously unpublished data that offer an alternative, nonmnemonic interpretation of this pattern of deficit. In their study, Wajima and Sawaguchi [Wajima, K., & Sawaguchi, T. The role of GABAergic inhibiton in suppressing perseverative responses in the monkey prefrontal cortex. Neuroscience Research, 50(Suppl. 1), P3–P317, 2004] applied the GABA A antagonist bicuculline methiodide unilaterally to the PFC of two monkeys while they performed an oculomotor delayed-response task. Consistent with previous studies, errors for the initial memory-guided saccade were markedly higher when the cued location fell into the region of the visual field affected by the infusion. These erroneous saccades tended to select an alternative target location (out of a possible 16) that had not been cued on that trial. By extending the analysis window, however, it was observed that the second, “corrective” saccade often acquired the location that had been cued on that trial. Further analysis of the erroneous initial saccades indicated that they tended to be directed to a location that had been relevant on the previous trial. Thus, the deficit was not one of “forgetting” the cued location. Rather, it was one of selecting between currently and previously relevant locations. These findings suggest a need for a reconsideration of the concept of the mnemonic scotoma, which in turn invites a reconsideration of functional interpretations of sustained neuronal activity in PFC.

It is widely assumed that the short-term retention of information is accomplished via maintenance of an active neural trace. However, we demonstrate that memory can be preserved across a brief delay despite the apparent loss of sustained representations. Delay period activity may, in fact, reflect the focus of attention, rather than STM. We unconfounded attention and memory by causing external and internal shifts of attention away from items that were being actively retained. Multivariate pattern analysis of fMRI indicated that only items within the focus of attention elicited an active neural trace. Activity corresponding to representations of items outside the focus quickly dropped to baseline. Nevertheless, this information was remembered after a brief delay. Our data also show that refocusing attention toward a previously unattended memory item can reactivate its neural signature. The loss of sustained activity has long been thought to indicate a disruption of STM, but our results suggest that, even for small memory loads not exceeding the capacity limits of STM, the active maintenance of a stimulus representation may not be necessary for its short-term retention.

Verbal working memory (VWM), the ability to maintain and manipulate representations of speech sounds over short periods, is held by some influential models to be independent from the systems responsible for language production and comprehension [e.g., Baddeley, A. D. Working memory, thought, and action . New York, NY: Oxford University Press, 2007]. We explore the alternative hypothesis that maintenance in VWM is subserved by temporary activation of the language production system [Acheson, D. J., & MacDonald, M. C. Verbal working memory and language production: Common approaches to the serial ordering of verbal information. Psychological Bulletin, 135, 50–68, 2009b]. Specifically, we hypothesized that for stimuli lacking a semantic representation (e.g., nonwords such as mun ), maintenance in VWM can be achieved by cycling information back and forth between the stages of phonological encoding and articulatory planning. First, fMRI was used to identify regions associated with two different stages of language production planning: the posterior superior temporal gyrus (pSTG) for phonological encoding (critical for VWM of nonwords) and the middle temporal gyrus (MTG) for lexical–semantic retrieval (not critical for VWM of nonwords). Next, in the same subjects, these regions were targeted with repetitive transcranial magnetic stimulation (rTMS) during language production and VWM task performance. Results showed that rTMS to the pSTG, but not the MTG, increased error rates on paced reading (a language production task) and on delayed serial recall of nonwords (a test of VWM). Performance on a lexical–semantic retrieval task (picture naming), in contrast, was significantly sensitive to rTMS of the MTG. Because rTMS was guided by language production-related activity, these results provide the first causal evidence that maintenance in VWM directly depends on the long-term representations and processes used in speech production.

Left inferior frontal gyrus (IFG) is a critical neural substrate for the resolution of proactive interference (PI) in working memory. We hypothesized that left IFG achieves this by controlling the influence of familiarity- versus recollection-based information about memory probes. Consistent with this idea, we observed evidence for an early (200 msec)-peaking signal corresponding to memory probe familiarity and a late (500 msec)-resolving signal corresponding to full accrual of trial-related contextual (“recollection-based”) information. Next, we applied brief trains of repetitive transcranial magnetic stimulation (rTMS) time locked to these mnemonic signals, to left IFG and to a control region. Only early rTMS of left IFG produced a modulation of the false alarm rate for high-PI probes. Additionally, the magnitude of this effect was predicted by individual differences in susceptibility to PI. These results suggest that left IFG-based control may bias the influence of familiarity- and recollection-based signals on recognition decisions.

Understanding the contributions of the prefrontal cortex (PFC) to working memory is central to understanding the neural bases of high-level cognition. One question that remains controversial is whether the same areas of the dorsolateral PFC (dlPFC) that participate in the manipulation of information in working memory also contribute to its short-term retention (STR). We evaluated this question by first identifying, with functional magnetic resonance imaging (fMRI), brain areas involved in manipulation. Next, these areas were targeted with repetitive transcranial magnetic stimulation (rTMS) while subjects performed tasks requiring only the STR or the STR plus manipulation of information in working memory. fMRI indicated that manipulation-related activity was independent of retention-related activity in both the PFC and superior parietal lobule (SPL). rTMS, however, yielded a different pattern of results. Although rTMS of the dlPFC selectively disrupted manipulation, rTMS of the SPL disrupted manipulation and STR to the same extent. rTMS of the postcentral gyrus (a control region) had no effect on performance. The implications of these results are twofold. In the PFC, they are consistent with the view that this region contributes more importantly to the control of information in working memory than to its STR. In the SPL, they illustrate the importance of supplementing the fundamentally correlational data from neuroimaging with a disruptive method, which affords stronger inference about structure-function relations.

The prefrontal cortex (PFC) contributes to working memory functions via executive control processes that do not entail the storage, per se, of mnemonic representations. One of these control processes may be a sensory gating mechanism that facilitates retention of representations in working memory by down-regulating the gain of the sensory processing of intervening irrelevant stimuli. This idea was tested by scanning healthy young adults with functional magnetic resonance imaging while they performed a delayed face-recognition task. The 2 × 2 factorial design varied the factors of Memory (present, absent) and Distraction (present, absent). During memory-present trials, target and probe stimuli were individual gray-scale male faces. Memory-absent trials were identical, except that they employed the same recurring female faces (denoting a “no memory” trial). Distraction-present trials featured rapid serial visual presentation of bespectacled male faces during the two middle seconds of the delay. The first step of the analyses identified dorsolateral PFC (dlPFC) and inferior occipitotemporal cortex (IOTC) voxels exhibiting delay-period activity in memory-present/distraction-absent trials, that is, the “unfilled” delay. Within these voxels, distraction-evoked activity in the dlPFC was markedly higher during trials that required the concurrent short-term retention of information than on those that did not, whereas the opposite effect was seen in the IOTC. These results are consistent with the view that processes related to sensory gating account for a portion of the delay-period activity that is routinely observed in the dlPFC.