Single-pulse Transcranial Magnetic Stimulation Affects Working-memory Performance via Posterior Beta-band Oscillations.

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.

Representing Context and Priority in Working Memory.

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.

Neural Signatures of Competition between Voluntary and Involuntary Influences over the Focus of Attention in Visual Working Memory.

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.

Investigating the Roles of the Visual and Parietal Cortex in Representing Content versus Context in Visual Working Memory.

Content-to-context binding is crucial for working memory performance. Using a dual-serial retrocueing (DSR) task on oriented gratings, Yu et al. (2020) found that content (orientation) of both prioritized and unprioritized memory items (PMI; UMI) was represented simultaneously in visual cortex, while their context (location) was represented in intraparietal sulcus (IPS), with a priority-based remapping of the representation of content and context of the UMI in each region, respectively. This registered report acquired fMRI of 24 healthy adults while they performed a DSR task with location as the to-be-reported content and orientation as the task-relevant context. We contrasted three accounts: domain-dependent, the engagement of visual and parietal regions depends on the feature domain (orientation vs location); functional, the engagement of these regions depends on their function (content vs context); and hybrid-a combination of the domain-dependent account and the additional stipulation that IPS encodes context regardless of domain. Delay-period activity in early visual cortex conformed most closely with functional predictions: robust priority-sensitive representation of stimulus location (content), but no evidence for the active representation of stimulus orientation (context). Delay-period activity in IPS, in contrast, conformed most closely to predictions of the hybrid account: active representation of content (location) and of prioritized context (orientation). Exploratory analyses further supported the hybrid account of IPS, revealing univariate sensitivity to variation in both content and context load, the latter in a manner that predicted individual differences in behavior. The representation of visual information in working memory is highly dependent on behavioral context.

Strategic control of location and ordinal context in visual working memory.

Working memory (WM) requires encoding stimulus identity and context (e.g. where or when stimuli were encountered). To explore the neural bases of the strategic control of context binding in WM, we acquired fMRI while subjects performed delayed recognition of 3 orientation patches presented serially and at different locations. The recognition probe was an orientation patch with a superimposed digit, and pretrial instructions directed subjects to respond according to its location ("location-relevant"), to the ordinal position corresponding to its digit ("order-relevant"), or to just its orientation (relative to all three samples; "context-irrelevant"). Delay period signal in PPC was greater for context-relevant than for "context-irrelevant" trials, and multivariate decoding revealed strong sensitivity to context binding requirements (relevant vs. "irrelevant") and to context domain ("location-" vs. "order-relevant") in both occipital cortex and PPC. At recognition, multivariate inverted encoding modeling revealed markedly different patterns in these 2 regions, suggesting different context-processing functions. In occipital cortex, an active representation of the location of each of the 3 samples was reinstated regardless of the trial type. The pattern in PPC, by contrast, suggested a trial type-dependent filtering of sample information. These results indicate that PPC exerts strategic control over the representation of stimulus context in visual WM.

Context Binding in Visual Working Memory Is Reflected in Bilateral Event-Related Potentials, But Not in Contralateral Delay Activity.

Successful retrieval of a specific item from visual working memory (VWM) depends on the binding of that item to its unique context. Recent functional magnetic resonance imaging studies of VWM manipulating memory set homogeneity have identified an important role for the intraparietal sulcus in context binding, independent of any role in representing stimulus identity. The current study explored whether the contralateral delay activity (CDA), which is an event-related potential (ERP) component derived from posterior electrodes that tracks the amount of information held in VWM, might also be sensitive to context-binding demands. In experiment 1, human participants performed lateralized delayed recognition with memory sets containing one, three, or five items that were drawn from the same category (orientations: "homogeneous") or from different categories (orientation, color, and luminance: "heterogeneous"). Because the location and identity of the memory probe indicated the item to be retrieved, homogeneous trials placed higher context-binding demands. VWM capacity was higher in heterogeneous trials. ERPs contralateral (contra) and ipsilateral (ipsi) to the remembered stimuli were higher for homogeneous trials, but these differences were removed in the contra - ipsi subtraction that produced the CDA. In experiment 2, human participants performed lateralized delayed recall with memory sets of one or three items (homogeneous or heterogeneous). Behavior was superior for three-item heterogeneous trials than for homogeneous trials, with modeling revealing context-binding errors in the latter. Bilateral ERPs and CDA results replicated experiment 1. These results support that the CDA tracks the number of object files engaged by VWM and establish that it is not sensitive to context-binding demands.

Flexible top-down control in the interaction between working memory and perception.

Successful goal-directed behavior often requires continuous sensory processing while simultaneously maintaining task-related information in working memory (WM). Although WM and perception are known to interact, little is known about how their interactions are controlled. Here, we tested the hypothesis that WM perception interactions engage two distinct modes of control - proactive and reactive - in a manner similar to classic conflict-adaptation tasks (e.g. Stroop, flanker, and Simon). Participants performed a delayed recall-of-orientation WM task, plus a standalone visual discrimination-of-orientation task the occurred during the delay period, and with the congruity in orientation between the tasks manipulated. Proactive control was seen in the sensitivity of task performance to the previous trial's congruity (i.e. a Gratton effect). Reactive control was observed in a repulsive serial-dependence produced by incongruent discriminanda. Quantitatively, these effects were explained by parameters from a reinforcement learning-based model that tracks trial-to-trial fluctuations in control demand: reactive control by a phasic control prediction error (control PE), and proactive control by a tonic level of predicted conflict updated each trial by the control PE. Thus, WM-perception interactions may be controlled by the same mechanisms that govern conflict in other domains of cognition, such as response selection.

The Influence of Active Removal from Working Memory on Serial Dependence.

Flexible control of the contents of working memory (WM) includes removing no-longer-relevant information. Although simply withdrawing attention offers a "passive" mechanism, empirical findings suggest that it is also possible to actively remove information from WM. In this Registered Report we tested evidence that the bias (serial dependence) that an item exerts on the subsequent trial will be opposite in sign-attraction vs. repulsion - depending on whether it was passively or actively removed, respectively. A repulsive bias would be consistent with a specific mechanism for active removal: a rapid adaptation-like modification of perceptual circuitry. In a preliminary study, trials of two types were administered in pairs, multi-item WM followed by 1-item delayed recall, and we evaluated serial dependence of the latter on items from the former. In the first trial of each pair, two memoranda were presented, then one was designated irrelevant, then a third memorandum was presented. The critical manipulation was whether the third item was presented at the same location as the now "irrelevant memory item" (IMI). Overlap between the two should prompt the active removal of the IMI, whereas nonoverlap might prompt just the withdrawal of attention. Whereas the IMI exerted the expected attractive bias on 1-item recall in the no-overlap condition, we found an (unexpected) repulsive bias in the overlap condition. Because repulsive biases have been attributed to the adaptation-like modification of perceptual circuitry, replication of this result in this Registered Report would provide independent evidence for this mechanism for active removal from WM. Interpretation of the Stage 2 results are complicated by the fact that the approved Registered Report, carried out online, generated data that failed to meet a basic sanity check, and were therefore uninterpretable. Consequently, a follow-up lab-based experiment using procedures similar to the Registered Report generated results consistent with the hypothesis of principal theoretical interest: The IMI in the overlap condition exerted a repulsive bias on the subsequent trial.

Priority-based transformations of stimulus representation in visual working memory.

How does the brain prioritize among the contents of working memory (WM) to appropriately guide behavior? Previous work, employing inverted encoding modeling (IEM) of electroencephalography (EEG) and functional magnetic resonance imaging (fMRI) datasets, has shown that unprioritized memory items (UMI) are actively represented in the brain, but in a "flipped", or opposite, format compared to prioritized memory items (PMI). To acquire independent evidence for such a priority-based representational transformation, and to explore underlying mechanisms, we trained recurrent neural networks (RNNs) with a long short-term memory (LSTM) architecture to perform a 2-back WM task. Visualization of LSTM hidden layer activity using Principal Component Analysis (PCA) confirmed that stimulus representations undergo a representational transformation-consistent with a flip-while transitioning from the functional status of UMI to PMI. Demixed (d)PCA of the same data identified two representational trajectories, one each within a UMI subspace and a PMI subspace, both undergoing a reversal of stimulus coding axes. dPCA of data from an EEG dataset also provided evidence for priority-based transformations of the representational code, albeit with some differences. This type of transformation could allow for retention of unprioritized information in WM while preventing it from interfering with concurrent behavior. The results from this initial exploration suggest that the algorithmic details of how this transformation is carried out by RNNs, versus by the human brain, may differ.

Spontaneous alpha-band amplitude predicts subjective visibility but not discrimination accuracy during high-level perception.

Near-threshold perception is a paradigm case of awareness diverging from reality - the perception of an unchanging stimulus can vacillate from undetected to clearly perceived. The amplitude of low-frequency brain oscillations - particularly in the alpha-band (8-13 Hz) - has emerged as a reliable predictor of trial-to-trial variability in perceptual decisions based on simple, low-level stimuli. Here, we addressed the question of how spontaneous oscillatory amplitude impacts subjective and objective aspects of perception using high-level visual stimuli. Human observers completed a near-threshold face/house discrimination task with subjective visibility ratings while electroencephalograms (EEG) were recorded. Using single-trial multiple regression analysis, we found that spontaneous fluctuations in prestimulus alpha-band amplitude were negatively related to visibility judgments but did not predict trial-by-trial accuracy. These results extend previous findings that indicate that strong prestimulus alpha diminishes subjective perception without affecting the accuracy or sensitivity (d') of perceptual decisions into the domain of high-level perception.

Spectral Distribution Dynamics across Different Attentional Priority States.

Anticipatory covert spatial attention improves performance on tests of visual detection and discrimination, and shifts are accompanied by decreases and increases of α band power at electroencephalography (EEG) electrodes corresponding to the attended and unattended location, respectively. Although the increase at the unattended location is often interpreted as an active mechanism (e.g., inhibiting processing at the unattended location), most experiments cannot rule out the alternative possibility that it is a secondary consequence of selection elsewhere. To adjudicate between these accounts, we designed a Posner-style visual cueing task in which male and female human participants made orientation judgments of targets appearing at one of four locations: up, down, right, or left. Critically, trials were blocked such that within a block the locations along one meridian alternated in status between attended and unattended, and targets never appeared at the other two, making them irrelevant. Analyses of the concurrently measured EEG signal were conducted on "traditional" narrowband α (8-14 Hz), as well as on two components resulting from the decomposition of this signal: "periodic" α; and the slope of the aperiodic 1/f-like component. Although data from right-left blocks replicated the familiar pattern of lateralized asymmetry in narrowband α power, with neither α signal could we find evidence for any difference in the time course at unattended versus irrelevant locations, an outcome consistent with the secondary-consequence interpretation of attention-related dynamics in the α band. Additionally, 1/f slope was shallower at attended and unattended locations, relative to irrelevant, suggesting a tonic adjustment of physiological state.SIGNIFICANCE STATEMENT Visual spatial attention, the prioritization of one location in the visual field, is critical for guiding behavior in cluttered environments. Although influential theories posit an important role for α band oscillations in the inhibition of processing at unattended locations, we used a novel procedure to find evidence for an alternative interpretation: selection of one location may simply result in a return to physiological baseline at all others. In addition to determining one way that attention does not work (important for future progress in this field), we also discovered novel evidence for one way that it does work: by modifying the tonic physiological state (indexed by an aperiodic component of the electroencephalography (EEG)] at locations where spatial selection is likely to occur.

The Neural Codes Underlying Internally Generated Representations in Visual 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.

Understanding occipital and parietal contributions to visual working memory: Commentary on Xu (2020).

In her commentary, Xu (2020) admonishes the reader that "To have a full understanding of the cognitive mechanisms underlying VWM [visual working memory], both behavioral and neural evidence needs to be taken into account. This is a must, and not a choice, for any study that attempts to capture the nature of VWM" (p. 11). Although we don't disagree with this statement, our overall assessment of this commentary is that it, itself, fails to satisfy several "musts" and, consequently, does not pose a serious challenge for the sensory recruitment framework for understanding visual working memory. These "musts" include accurately characterizing the framework being critiqued, not favoring verbal models and intuition at the expense of formal quantitative models, and providing even-handed interpretation of the work of others. We'll conclude with a summary of how the sensory recruitment framework can be incorporated into a broader working model of visual working memory.

Spatial specificity of feature-based interaction between working memory and visual processing.

Visual working memory interacts with ongoing visual processing in a stimulus-specific manner, potentially through a common neural substrate supporting visual perception and working memory maintenance. The spatial specificity of this effect, however, remains unknown. The current study tested whether features in working memory influence perception in a spatially specific or global fashion. Across four experiments, subjects performed perceptual discrimination tasks on orientation or on contrast while concurrently holding an orientation in working memory. Experiments 1 and 2 revealed that memory content boosted the perceived contrast of the discrimination stimulus when the two matched in orientation, but only when the locations of the memorandum and the discriminandum also matched. In turn, feature-based influence on memory precision was also greater when locations matched. Experiments 3 and 4 demonstrated that the spatial specificity of this interaction was influenced by task demands. When encoding of location was discouraged in Experiment 3, memory interacted with perception in a global fashion, whereas when location was task-relevant in otherwise very similar Experiment 4, the feature-based enhancement was again modulated by location. These results suggest that context-binding demand is an important determinant of the spatial specificity of memory-perception interaction and highlight the flexible configurability of working memory representations. (PsycInfo Database Record (c) 2021 APA, all rights reserved).

Attention Biases Competition for Visual Representation via Dissociable Influences from Frontal and Parietal Cortex.

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.

Gender (Im)balance in Citation Practices in Cognitive Neuroscience.

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.

The Role of Location-Context Binding in Nonspatial Visual Working Memory.

Successful retrieval of an item from visual working memory (VWM) often requires an associated representation of the trial-unique context in which that item was presented. In experiment 1, fMRI of 16 male and female humans replicated a previous dissociation of the effects of manipulating memory load in comparison to the effects of manipulating context binding, by comparing VWM for one oriented line versus for three lines individuated by their location versus for three "heterogeneous" items drawn from different categories (orientation, color, and luminance): delay-period fMRI signal in frontal cortex and intraparietal sulcus (IPS) was sensitive to stimulus homogeneity rather than to memory load per se. Additionally, inspection of behavioral performance revealed a broad range of individual differences in the probability of responses to nontargets (also known as "swap errors"), and a post hoc comparison of high swap-error versus low swap-error groups generated several intriguing results: at recall, high swap-error subjects were seen to represent both the orientation and the location of the probed item less strongly, and with less differentiation from nonprobed items, and delay-period signal in IPS predicted behavioral and neural correlates of context binding at recall. In experiment 2, which was a preregistered replication, the 27 male and female humans were grouped into low and high swap-error groups by median split, and the results were broadly consistent with experiment 1. These results present a neural correlate of swap errors, and suggest that delay-period activity of the IPS may be more important for the operation of context binding than for representation per se of stimulus identity.