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Multiple neural mechanisms underlying gating to working memory have been proposed with divergent results obtained in human and animal studies. Previous findings from non-human primates suggest prefrontal beta frequency bursts as a correlate of transient inhibition during selective encoding. Human studies instead suggest a similar role for sensory alpha power fluctuations. To cast light on these discrepancies we employed a sequential working memory task with distractors for human participants. In particular, we examined their whole-brain electrophysiological activity in both alpha and beta bands with the same single-trial burst analysis earlier performed on non-human primates. Our results reconcile earlier findings by demonstrating that both alpha and beta bursts in humans correlate with the filtering and control of memory items, but with region and task-specific differences between the two rhythms. Occipital beta burst patterns were selectively modulated during the transition from sensory processing to memory retention whereas prefrontal and parietal beta bursts tracked sequence order and were proactively upregulated prior to upcoming target encoding. Occipital alpha bursts instead increased during the actual presentation of unwanted sensory stimuli. Source reconstruction additionally suggested the involvement of striatal and thalamic alpha and beta. Thus, specific whole-brain burst patterns correlate with different aspects of working memory control., (© 2024. The Author(s).)

Working memory (WM) allows us to remember and selectively control a limited set of items. Neural evidence suggests it is achieved by interactions between bursts of beta and gamma oscillations. However, it is not clear how oscillations, reflecting coherent activity of millions of neurons, can selectively control individual WM items. Here we propose the novel concept of spatial computing where beta and gamma interactions cause item-specific activity to flow spatially across the network during a task. This way, control-related information such as item order is stored in the spatial activity independent of the detailed recurrent connectivity supporting the item-specific activity itself. The spatial flow is in turn reflected in low-dimensional activity shared by many neurons. We verify these predictions by analyzing local field potentials and neuronal spiking. We hypothesize that spatial computing can facilitate generalization and zero-shot learning by utilizing spatial component as an additional information encoding dimension., (© 2023. The Author(s).)

Working memory has long been thought to arise from sustained spiking/attractor dynamics. However, recent work has suggested that short-term synaptic plasticity (STSP) may help maintain attractor states over gaps in time with little or no spiking. To determine if STSP endows additional functional advantages, we trained artificial recurrent neural networks (RNNs) with and without STSP to perform an object working memory task. We found that RNNs with and without STSP were able to maintain memories despite distractors presented in the middle of the memory delay. However, RNNs with STSP showed activity that was similar to that seen in the cortex of a non-human primate (NHP) performing the same task. By contrast, RNNs without STSP showed activity that was less brain-like. Further, RNNs with STSP were more robust to network degradation than RNNs without STSP. These results show that STSP can not only help maintain working memories, it also makes neural networks more robust and brain-like., Competing Interests: The authors have no competing interests., (Copyright: © 2022 Kozachkov et al. This is an open access article distributed under the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.)

Working memories have long been thought to be maintained by persistent spiking. However, mounting evidence from multiple-electrode recording (and single-trial analyses) shows that the underlying spiking is better characterized by intermittent bursts of activity. A counterargument suggested this intermittent activity is at odds with observations that spike-time variability reduces during task performance. However, this counterargument rests on assumptions, such as randomness in the timing of the bursts, which may not be correct. Thus, we analyzed spiking and LFPs from monkeys' prefrontal cortex (PFC) to determine if task-related reductions in variability can co-exist with intermittent spiking. We found that it does because both spiking and associated gamma bursts were task-modulated, not random. In fact, the task-related reduction in spike variability could largely be explained by a related reduction in gamma burst variability. Our results provide further support for the intermittent activity models of working memory as well as novel mechanistic insights into how spike variability is reduced during cognitive tasks., (© 2022. The Author(s).)

Neural oscillations are evident across cortex but their spatial structure is not well- explored. Are oscillations stationary or do they form "traveling waves", i.e., spatially organized patterns whose peaks and troughs move sequentially across cortex? Here, we show that oscillations in the prefrontal cortex (PFC) organized as traveling waves in the theta (4-8Hz), alpha (8-12Hz) and beta (12-30Hz) bands. Some traveling waves were planar but most rotated. The waves were modulated during performance of a working memory task. During baseline conditions, waves flowed bidirectionally along a specific axis of orientation. Waves in different frequency bands could travel in different directions. During task performance, there was an increase in waves in one direction over the other, especially in the beta band., Competing Interests: The authors have declared that no competing interests exist.

Visual working memory (WM) storage is largely independent between the left and right visual hemifields/cerebral hemispheres, yet somehow WM feels seamless. We studied how WM is integrated across hemifields by recording neural activity bilaterally from lateral prefrontal cortex. An instructed saccade during the WM delay shifted the remembered location from one hemifield to the other. Before the shift, spike rates and oscillatory power showed clear signatures of memory laterality. After the shift, the lateralization inverted, consistent with transfer of the memory trace from one hemisphere to the other. Transferred traces initially used different neural ensembles from feedforward-induced ones, but they converged at the end of the delay. Around the time of transfer, synchrony between the two prefrontal hemispheres peaked in theta and beta frequencies, with a directionality consistent with memory trace transfer. This illustrates how dynamics between the two cortical hemispheres can stitch together WM traces across visual hemifields., Competing Interests: Declaration of interests The authors declare no competing interests., (Copyright © 2021 The Author(s). Published by Elsevier Inc. All rights reserved.)

Beta oscillations (∼13 to 30 Hz) have been observed during many perceptual, cognitive, and motor processes in a plethora of brain recording studies. Although the function of beta oscillations (hereafter "beta" for short) is unlikely to be explained by any single monolithic description, we here discuss several convergent findings. In prefrontal cortex (PFC), increased beta appears at the end of a trial when working memory information needs to be erased. A similar "clear-out" function might apply during the stopping of action and the stopping of long-term memory retrieval (stopping thoughts), where increased prefrontal beta is also observed. A different apparent role for beta in PFC occurs during the delay period of working memory tasks: it might serve to maintain the current contents and/or to prevent interference from distraction. We confront the challenge of relating these observations to the large literature on beta recorded from sensorimotor cortex. Potentially, the clear-out of working memory in PFC has its counterpart in the postmovement clear-out of the motor plan in sensorimotor cortex. However, recent studies support alternative interpretations. In addition, we flag emerging research on different frequencies of beta and the relationship between beta and single-neuron spiking. We also discuss where beta might be generated: basal ganglia, cortex, or both. We end by considering the clinical implications for adaptive deep-brain stimulation., (Copyright © 2019 the authors.)

Working memory is the fundamental function by which we break free from reflexive input-output reactions to gain control over our own thoughts. It has two types of mechanisms: online maintenance of information and its volitional or executive control. Classic models proposed persistent spiking for maintenance but have not explicitly addressed executive control. We review recent theoretical and empirical studies that suggest updates and additions to the classic model. Synaptic weight changes between sparse bursts of spiking strengthen working memory maintenance. Executive control acts via interplay between network oscillations in gamma (30-100 Hz) in superficial cortical layers (layers 2 and 3) and alpha and beta (10-30 Hz) in deep cortical layers (layers 5 and 6). Deep-layer alpha and beta are associated with top-down information and inhibition. It regulates the flow of bottom-up sensory information associated with superficial layer gamma. We propose that interactions between different rhythms in distinct cortical layers underlie working memory maintenance and its volitional control., (Copyright © 2018 Elsevier Inc. All rights reserved.)

Persistent spiking has been thought to underlie working memory (WM). However, virtually all of the evidence for this comes from studies that averaged spiking across time and across trials, which masks the details. On single trials, activity often occurs in sparse transient bursts. This has important computational and functional advantages. In addition, examination of more complex tasks reveals neural coding in WM is dynamic over the course of a trial. All this suggests that spiking is important for WM, but that its role is more complex than simply persistent spiking. Dual Perspectives Companion Paper: Persistent Spiking Activity Underlies Working Memory, by Christos Constantinidis, Shintaro Funahashi, Daeyeol Lee, John D. Murray, Xue-Lian Qi, Min Wang, and Amy F.T. Arnsten., (Copyright © 2018 the authors 0270-6474/18/387013-07$15.00/0.)

All of the cerebral cortex has some degree of laminar organization. These different layers are composed of neurons with distinct connectivity patterns, embryonic origins, and molecular profiles. There are little data on the laminar specificity of cognitive functions in the frontal cortex, however. We recorded neuronal spiking/local field potentials (LFPs) using laminar probes in the frontal cortex (PMd, 8A, 8B, SMA/ACC, DLPFC, and VLPFC) of monkeys performing working memory (WM) tasks. LFP power in the gamma band (50-250 Hz) was strongest in superficial layers, and LFP power in the alpha/beta band (4-22 Hz) was strongest in deep layers. Memory delay activity, including spiking and stimulus-specific gamma bursting, was predominately in superficial layers. LFPs from superficial and deep layers were synchronized in the alpha/beta bands. This was primarily unidirectional, with alpha/beta bands in deep layers driving superficial layer activity. The phase of deep layer alpha/beta modulated superficial gamma bursting associated with WM encoding. Thus, alpha/beta rhythms in deep layers may regulate the superficial layer gamma bands and hence maintenance of the contents of WM., Competing Interests: The authors declare no conflict of interest., (Copyright © 2018 the Author(s). Published by PNAS.)

Working memory (WM) activity is not as stationary or sustained as previously thought. There are brief bursts of gamma (~50-120 Hz) and beta (~20-35 Hz) oscillations, the former linked to stimulus information in spiking. We examined these dynamics in relation to readout and control mechanisms of WM. Monkeys held sequences of two objects in WM to match to subsequent sequences. Changes in beta and gamma bursting suggested their distinct roles. In anticipation of having to use an object for the match decision, there was an increase in gamma and spiking information about that object and reduced beta bursting. This readout signal was only seen before relevant test objects, and was related to premotor activity. When the objects were no longer needed, beta increased and gamma decreased together with object spiking information. Deviations from these dynamics predicted behavioral errors. Thus, beta could regulate gamma and the information in WM.

Working memory is thought to result from sustained neuron spiking. However, computational models suggest complex dynamics with discrete oscillatory bursts. We analyzed local field potential (LFP) and spiking from the prefrontal cortex (PFC) of monkeys performing a working memory task. There were brief bursts of narrow-band gamma oscillations (45-100 Hz), varied in time and frequency, accompanying encoding and re-activation of sensory information. They appeared at a minority of recording sites associated with spiking reflecting the to-be-remembered items. Beta oscillations (20-35 Hz) also occurred in brief, variable bursts but reflected a default state interrupted by encoding and decoding. Only activity of neurons reflecting encoding/decoding correlated with changes in gamma burst rate. Thus, gamma bursts could gate access to, and prevent sensory interference with, working memory. This supports the hypothesis that working memory is manifested by discrete oscillatory dynamics and spiking, not sustained activity., (Copyright © 2016 Elsevier Inc. All rights reserved.)

Changes in oscillatory brain activity are strongly correlated with performance in cognitive tasks and modulations in specific frequency bands are associated with working memory tasks. Mesoscale network models allow the study of oscillations as an emergent feature of neuronal activity. Here we extend a previously developed attractor network model, shown to faithfully reproduce single-cell activity during retention and memory recall, with synaptic augmentation. This enables the network to function as a multi-item working memory by cyclic reactivation of up to six items. The reactivation happens at theta frequency, consistently with recent experimental findings, with increasing theta power for each additional item loaded in the network's memory. Furthermore, each memory reactivation is associated with gamma oscillations. Thus, single-cell spike trains as well as gamma oscillations in local groups are nested in the theta cycle. The network also exhibits an idling rhythm in the alpha/beta band associated with a noncoding global attractor. Put together, the resulting effect is increasing theta and gamma power and decreasing alpha/beta power with growing working memory load, rendering the network mechanisms involved a plausible explanation for this often reported behavior.