bioRxiv Science⌕ Search

Biology subjects

Adam, K. C. S.

Publications and source records attributed to Adam, K. C. S..

8 recordsLinked to original sources

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.

neuroscience↗

Exploring neural mechanisms underlying error-related impairments in active working memory suggests an adaptive shielding of contents during cognitive control

Goal-directed behavior relies on cognitive flexibility - the ability to rapidly adapt ongoing thoughts and behaviors while preserving task-relevant information. The performance monitoring system optimizes such behavior by detecting and evaluating errors, while the working memory (WM) system maintains relevant information and protects it from interference. We investigated how these two systems interact. In prior work (Wessel et al., 2022), we found that motor errors impaired active WM maintenance (Error-Related Impairment of Active working Memory; ERIAM). Here, we aimed to identify the source of ERIAM by tracking a neurophysiological marker of visual WM maintenance - the contralateral delay activity (CDA) - throughout the error-making process. Forty-two human participants maintained visual information in WM while performing a motoric task during the delay period. Consistent with prior results, a significant ERIAM effect occurred: motor errors impaired WM performance. Critically, CDA amplitudes did not differ between motor correct and error trials before the flanker task, ruling out a general performance deficit. The CDA was also unaffected immediately after flankers, ruling out a perceptual interference explanation. Significant CDA differences only emerged after motor errors, supporting a genuinely error-related origin of the ERIAM effect. Contrary to prediction, however, CDA was more disrupted after correct responses than errors, and greater disruptions predicted a smaller ERIAM effect. These findings suggest that participants might store WM in multiple states to reduce interference from errors and that the CDA dynamics reflect these adaptive shielding strategies. These findings provide new insights into the source of error-related interference in active WM.

neuroscience↗

Dissociating Contributions of Theta and Alpha Oscillations from Aperiodic Neural Activity in Human Visual Working Memory

While visual working memory (WM) is strongly associated with reductions in occipitoparietal alpha (8-12 Hz) power, the role of frontal midline theta (4-7 Hz) power is less clear, with both increases and decreases widely reported. Here, we test the hypothesis that this theta paradox can be explained by non-oscillatory, aperiodic neural activity dynamics. Because traditional time-frequency analyses of electroencephalography (EEG) data conflate oscillations and aperiodic activity, event-related changes in aperiodic activity can manifest as task-related changes in apparent oscillations, even when none are present. Reanalyzing EEG data from two visual WM experiments (n = 74, of either sex), and leveraging spectral parameterization, we found systematic changes in aperiodic activity with WM load, and we replicated classic alpha, but not theta, oscillatory effects after controlling for aperiodic changes. Aperiodic activity decreased during WM retention, and further flattened over the occipitoparietal cortex with an increase in WM load. After controlling for these dynamics, aperiodic-adjusted alpha power decreased with increasing WM load. In contrast, aperiodic-adjusted theta power appeared to increase during WM retention, but because aperiodic activity reduces more, it falsely appears as though theta "oscillatory" power (e.g., total band power) is reduced. Furthermore, only a minority of participants (31/74) had a detectable degree of theta oscillations. These results offer a potential resolution to the theta paradox where studies show contrasting power changes. Additionally, we have identified novel aperiodic dynamics during human visual WM. Significance statementWorking Memory (WM) is our ability to hold information in mind without it being present in our external environment. Years of research focused on oscillatory brain dynamics to discover the mechanisms of WM. Here, we specifically look at oscillatory and non-oscillatory, aperiodic activity as measured with scalp EEG to test their significance in supporting WM. We challenge earlier findings regarding theta oscillations with our analysis approach, while replicating alpha oscillation findings. Furthermore, aperiodic activity is found to be involved in WM, over frontal regions in a task-general manner, and over anterior regions this activity is reduced with an increase in the number of remembered items. Thus, we have identified novel aperiodic dynamics during human visual WM.

neuroscience↗

History-driven modulations of population codes in early visual cortex during visual search

To find important objects, we must focus on our goals, ignore distractions, and take our changing environment into account. This is formalized in models of visual search whereby goal-driven, stimulus-driven and history-driven factors are integrated into a priority map that guides attention. Stimulus history robustly influences where attention is allocated even when the physical stimulus is the same: when a salient distractor is repeated over time, it captures attention less effectively. A key open question is how we come to ignore salient distractors when they are repeated. Goal-driven accounts propose that we use an active, expectation-driven mechanism to attenuate the distractor signal (e.g., predictive coding), whereas stimulus-driven accounts propose that the distractor signal is attenuated due to passive changes to neural activity and inter-item competition (e.g., adaptation). To test these competing accounts, we measured item-specific fMRI responses in human visual cortex during a visual search task where trial history was manipulated (colors unpredictably switched or were repeated). Consistent with a stimulus-driven account of history-based distractor suppression, we found that repeated singleton distractors were suppressed starting in V1, and distractor suppression did not increase in later visual areas. In contrast, we observed signatures of goal-driven target enhancement that were absent in V1, increased across visual areas, and were not modulated by stimulus history. Our data suggest that stimulus history does not alter goal-driven expectations, but rather modulates canonically stimulus-driven sensory responses to contribute to a temporally-integrated representation of priority. Significance StatementVisual search refers to our ability to find what we are looking for in a cluttered visual world (e.g., finding your keys). To perform visual search, we must integrate information about our goals (e.g., find the red key-chain), the environment (e.g., salient items capture your attention), and changes to the environment (i.e., stimulus history). Although stimulus history impacts behavior, the neural mechanisms that mediate history-driven effects remain debated. Here, we leveraged fMRI and multivariate analysis techniques to measure history-driven changes to the neural representation of items during visual search. We found that stimulus history influenced the representation of a salient pop-out distractor starting in V1, suggesting that stimulus history operates via modulations in early sensory processing rather than goal-driven expectations.

neuroscience↗

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.

neuroscience↗

Δ9-Tetrahydrocannabinol (THC) impairs visual working memory performance

With the increasing prevalence of legal cannabis use and availability, there is an urgent need to identify cognitive impairments related to its use. It is widely believed that cannabis, or its main psychoactive component {Delta}9-tetrahydrocannabinol (THC), impairs working memory, i.e., the ability to temporarily hold information in mind. However, our review of the literature yielded surprisingly little empirical support for an effect of THC or cannabis on working memory. We thus conducted a study with 3 main goals: (1) quantify the effect of THC on visual working memory in a well-powered sample (2) test the potential role of cognitive effects (mind wandering and metacognition) in disrupting working memory, and (3) demonstrate how insufficient sample size and task duration reduce the likelihood of detecting a drug effect. We conducted two double-blind, randomized crossover experiments in which healthy adults (N=23, 23) performed a reliable and validated visual working memory task (the "Discrete Whole-Report task", 90 trials) after administration of THC (7.5 and/or 15 mg oral) or placebo. We also assessed self-reported mind wandering (Exp 1) and metacognitive accuracy about ongoing task performance (Exp 2). THC impaired working memory performance (d = .65), increased mind wandering (Exp 1), and decreased metacognitive accuracy about task performance (Exp 2). Thus, our findings indicate that THC does impair visual working memory, and that this impairment may be related to both increased mind-wandering and decreased monitoring of task performance. Finally, we used a down-sampling procedure to illustrate the effects of task length and sample size on power to detect the acute effect of THC on working memory.

pharmacology and toxicology↗

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.

neuroscience↗

Dissecting the neural focus of attention reveals distinct processes for spatial attention and object-based storage in visual working memory

Complex cognition relies on both online representations in working memory (WM) said to reside in the focus of attention, and passive offline representations of related information. Here, we dissect the focus of attention by showing that distinct neural signals index the online storage of objects and sustained spatial attention. We recorded EEG activity during two tasks that employed identical stimulus displays while the relative demands for object storage and spatial attention varied. We found distinct delay-period signatures for an attention task (which only required spatial attention) and WM task (which invoked both spatial attention and object storage). Although both tasks required active maintenance of spatial information, only the WM task elicited robust contralateral delay activity that was sensitive to mnemonic load. Thus, we argue that the focus of attention is maintained via a collaboration between distinct processes for covert spatial orienting and object-based storage.

neuroscience↗