Items In Working Memory Research Articles

Persistent activity during working memory maintenance predicts long-term memory formation in the human hippocampus.

Working memory (WM) and long-term memory (LTM) are often viewed as separate cognitive systems. Little is known about how these systems interact when forming memories. We recorded single neurons in the human medial temporal lobe while patients maintained novel items in WM and completed a subsequent recognition memory test for the same items. In the hippocampus, but not in the amygdala, the level of WM content-selective persistent activity during WM maintenance was predictive of whether the item was later recognized with high confidence or forgotten. By contrast, visually evoked activity in the same cells was not predictive of LTM formation. During LTM retrieval, memory-selective neurons responded more strongly to familiar stimuli for which persistent activity was high while they were maintained in WM. Our study suggests that hippocampal persistent activity of the same cells supports both WM maintenance and LTM encoding, thereby revealing a common single-neuron component of these two memory systems.

Latent memory traces for prospective items in visual working memory

Latent memory traces for prospective items in visual working memory

Persistent activity during working memory maintenance predicts long-term memory formation in the human hippocampus.

Working Memory (WM) and Long-Term Memory (LTM) are often viewed as separate cognitive systems. Little is known about how these systems interact when forming memories. We recorded single neurons in the human medial temporal lobe while patients maintained novel items in WM and a subsequent recognition memory test for the same items. In the hippocampus but not the amygdala, the level of WM content-selective persist activity during WM maintenance was predictive of whether the item was later recognized with high confidence or forgotten. In contrast, visually evoked activity in the same cells was not predictive of LTM formation. During LTM retrieval, memory-selective neurons responded more strongly to familiar stimuli for which persistent activity was high while they were maintained in WM. Our study suggests that hippocampal persistent activity of the same cell supports both WM maintenance and LTM encoding, thereby revealing a common single-neuron component of these two memory systems.

Ocular working memory signals are flexible to behavioral priority and subjective imagery strength.

The pupillary light response was long considered a brainstem reflex, outside of cognitive influence. However, newer findings indicate that pupil dilation (and eye movements) can reflect content held "in mind" with working memory (WM). These findings may reshape understanding of ocular and WM mechanisms, but it is unclear whether the signals are artifactual or functional to WM. Here, we ask whether peripheral and oculomotor WM signals are sensitive to the task-relevance or "attentional state" of WM content. During eye-tracking, human participants saw both dark and bright WM stimuli, then were retroactively cued to the item that would most likely be tested. Critically, we manipulated the attentional priority among items by varying the cue reliability across blocks. We confirmed previous findings that remembering darker items is associated with larger pupils (vs. brighter), and that gaze is biased toward cued item locations. Moreover, we discovered that pupil and eye movement responses were influenced differently by WM item relevance. Feature-specific pupillary effects emerged only for highly prioritized WM items but were eliminated when cues were less reliable, and pupil effects also increased with self-reported visual imagery strength. Conversely, gaze position consistently veered toward the cued item location, regardless of cue reliability. However, biased microsaccades occurred at a higher frequency when cues were more reliable, though only during a limited post-cue time window. Therefore, peripheral sensorimotor processing is sensitive to the task-relevance or functional state of internal WM content, but pupillary and eye movement WM signals show distinct profiles. These results highlight a potential role for early visual processing in maintaining multiple WM content dimensions.NEW & NOTEWORTHY Here, we found that working memory (WM)-driven ocular inflections-feature-specific pupillary and saccadic biases-were muted for memory items that were less behaviorally relevant. This work illustrates that functionally informative goal signals may extend as early as the sensorimotor periphery, that pupil size may be under more fine-grained control than originally thought, and that ocular signals carry multiple dimensions of cognitively relevant information.

Steady-state Visual Evoked Potentials Reveal Dynamic (Re)allocation of Spatial Attention during Maintenance and Utilization of Visual Working Memory.

Visual working memory (VWM) allows storing goal-relevant information to guide future behavior. Prior work suggests that VWM is spatially organized and relies on spatial attention directed toward locations at which memory items were encoded, even if location is task-irrelevant. Importantly, attention often needs to be dynamically redistributed between locations, for example, in preparation for an upcoming probe. Very little is known about how attentional resources are distributed between multiple locations during a VWM task and even less about the dynamic changes governing such attentional shifts over time. This is largely due to the inability to use behavioral outcomes to reveal fast dynamic changes within trials. We here demonstrated that EEG steady-state visual evoked potentials (SSVEPs) successfully track the dynamic allocation of spatial attention during a VWM task. Participants were presented with to-be-memorized gratings and distractors at two distinct locations, tagged with flickering discs. This allowed us to dynamically track attention allocated to memory and distractor items via their coupling with space by quantifying the amplitude and coherence of SSVEP responses in the EEG signal to flickering stimuli at the former memory and distractor locations. SSVEP responses did not differ between memory and distractor locations during early maintenance. However, shortly before probe comparison, we observed a decrease in SSVEP coherence over distractor locations indicative of a reallocation of spatial attentional resources. RTs were shorter when preceded by stronger decreases in SSVEP coherence at distractor locations, likely reflecting attentional shifts from the distractor to the probe or memory location. We demonstrate that SSVEPs can inform about dynamic processes in VWM, even if location does not have to be reported by participants. This finding not only supports the notion of a spatially organized VWM but also reveals that SSVEPs betray a dynamic prioritization process of working memory items and locations over time that is directly predictive of memory performance.

Time course of attentional guidance by visual and verbal working memory representations

ABSTRACT Visual and verbal representations in working memory guide visual attention to a memory-matching item. It is well documented that visual cues have an advantage in target detection over verbal cues; however, little research has explored the differences in visual search processes and between attentional guidance by visual and verbal working memory. To address this issue, here we investigated the time course of attentional guidance through visual and verbal representations by comparing the inhibition of return effects. Previous research shows that both types of working memory items produce early facilitation effects and later inhibition effects, but there are lesser facilitation effects for verbal than for visual working memory guidance of attention. Results showed that in all conditions, attention was guided by memory representations, but no inhibition of return was observed. Thus, the robustness of previous findings on the time course of attentional guidance by working memory is low.

Behavioral fluctuation reflecting theta-rhythmic activation of sequential working memory

Sequential working memory, the ability to actively maintain sequential information, is essential for human cognition. The neural representation of each item in sequential working memory is thought to be activated rhythmically within the theta (3–7 Hz) range of human electrophysiology. In the current study, we predicted that if neural representations of sequential working memory items were truly activated rhythmically, periodic fluctuations in behavior would be evident. That is, the ease and speed of recalling each memory item would oscillate depending on the interval between memory encoding and recall, affected by the rhythmic neural representation. We conducted detailed analyses of reaction times for retrieving sequential and non-sequential information in eight experiments (total n = 125). The results revealed that reaction times for recalling sequential information showed fluctuation in the theta range as a function of the interval between memory encoding and recall, which was significantly stronger than that observed when the task did not require participants to remember the sequential order. Taken together, the current findings revealed that participants’ behavior exhibited theta-rhythmic fluctuation when recalling sequential information in a relatively large sample, supporting theta phase-dependent coding of sequential working memory.

Are latent working memory items retrieved from long-term memory?

Switching one's focus of attention between to-be-remembered items in working memory (WM) is critical for cognition, but the mechanisms by which this is accomplished are unclear. A long-term memory (LTM) account suggests that switching attention away from an item, and passively retaining and reactivating such "latent" items back into the focus of attention involves episodic LTM retrieval processes, even for delays of only a few seconds. We tested this hypothesis using a two-item, double-retrocue WM task that requires participants to switch attention away from and reactivate items followed by subsequent LTM tests for reactivated items from the initial WM task (vs. continuously retained or untested control items). We compared performance on these tests between older adults (a population with LTM deficits) and young adults with either full (Experiment 1) or divided (Experiment 2) attention during the WM delay periods. The effects of reactivating latent items, as well as ageing and divided attention, had significant effects on WM performance, but did not interact with or systematically affect subsequent LTM for reactivated versus control items on item-, location-, or associative-recognition memory judgements made with either high or low confidence. Experiment 3 confirmed that these effects did not depend on whether or not young participants were warned about the subsequent LTM tests before performing the WM task. These dissociations between WM and LTM are inconsistent with the LTM account of latent WM; they are more consistent with the dynamic processing model of WM (Current Directions in Psychological Science).

The reactivation of task rules triggers the reactivation of task-relevant items

Working memory (WM) describes the temporary storage of task-relevant items and procedural rules to guide action. Despite its central importance for goal-directed behavior, the interplay between WM and long-term memory (LTM) remains poorly understood. Recent studies have shown that repeated use of the same task-relevant item in WM results in a hand-off of the storage of that item to LTM, and switching to a new item reactivates WM. To further elucidate the rules governing WM-LTM interactions, we here planned to probe whether a change in task rules, independent of a switch in task-relevant items, would also lead to WM reactivation of maintained items. To this end, we used scalp-recorded electroencephalogram (EEG) data, specifically the contralateral delay activity (CDA), to track WM item storage while manipulating repetitions and changes in task rules and task-relevant items across trials in a visual WM task. We tested two rival hypotheses: If changes in task rules result in a reactivation of the target item representation, then the CDA should increase when a task change is cued even when the same target has been repeated across trials. However, if the reactivation of a task-relevant item only depends on the mnemonic availability of the item itself instead of the task it is used for, then only the changes in task-relevant items should reactivate the representations. Accordingly, the CDA amplitude should decrease for repeated task-relevant items independently of a task change. We found a larger CDA on task-switch compared to task-repeat trials, suggesting that the reactivation of task rules triggers the reactivation of task-relevant items in WM. By demonstrating that WM reactivation of LTM is interdependent for task rules and task-relevant items, this study informs our understanding of visual WM and its interplay with LTM. Preregistered Stage 1 protocolhttps://osf.io/zp9e8 (date of in-principle acceptance: 19/12/2021).

"Leader-Follower" Dynamic Perturbation Manipulates Multi-Item Working Memory in Humans.

Manipulating working memory (WM) is a central yet challenging notion. Previous studies suggest that WM items with varied memory strengths reactivate at different latencies, supporting a time-based mechanism. Motivated by this view, here we developed a purely bottom-up "Leader-Follower" behavioral approach to manipulate WM in humans. Specifically, task-irrelevant flickering color disks that are bound to each of the memorized items are presented during the delay period, and the ongoing luminance sequences of the color disks follow a Leader-Follower relationship, that is, a hundreds of milliseconds temporal lag. We show that this dynamic behavioral approach leads to better memory performance for the item associated with the temporally advanced luminance sequence (Leader) than the item with the temporally lagged luminance sequence (Follower), yet with limited effectiveness. Together, our findings constitute evidence for the essential role of temporal dynamics in WM operation and offer a promising, noninvasive WM manipulation approach.

The ordinal distance effect in working memory: does it exist in the absence of confounds?

In most daily-life situations, briefly remembering actions or words is not sufficient to reach a goal. You often have to remember them in a specific order. One behavioural observation of the processing of ordinal information in working memory is the ordinal distance effect. It refers to the facilitation in the ordinal processing of items that are at distant positions in comparison to items close to each other in working memory. So far, the ordinal distance effect has always been investigated with a simultaneous presentation of the items of the memory sequences. Such a presentation created a confound: items distant by their ordinal distance were also distant by their physical distance (i.e., the visuospatial distance between their positions on the screen). In the present study, we investigated whether the ordinal distance effect can be observed in the absence of a physical confound using a sequential presentation of the items of the memory sequence. Our findings revealed a combination of reversed and standard distance effects, unchanged by physical characteristics. The presence of a reversed distance effect suggests that a serial scanning strategy confers an advantage for adjacent items. Different strategies apply to the ordinal judgment of adjacent versus more distant items in verbal working memory. Interestingly, when ruling out the confound of primacy and recency effects, the standard distance effect disappeared while the reversed distance effect remained. Ultimately, our findings question the existence of the ordinal distance effect as a separate effect from other working memory confounds.

Alternating capture of attention by multiple visual working memory representations

Can we look for multiple objects simultaneously? Previous studies have proposed that the representation of an item in visual working memory (VWM) can bias the deployment of attention to memory-matching items in visual search. However, it is still controversial whether multiple VWM item representations are able to capture attention. In the present study, we adopted an eye-tracking technique to reveal this issue. In Experiment 1, we replicated Chen and Du’s Experiment 2 and adopted an eye-tracking technique to determine whether multiple VWM item representations are able to bias attention. In Chen and Du’s paradigm, the memory test was always followed by the search test, and the participants might intentionally prepare for the search task, which can affect the results of the research. Thus, in Experiment 2, we prevented participants from predicting the temporal occurrence of the visual search task by randomly conducting a search test or a memory test after memoranda. The findings of the present study suggested that only one working memory item at a time influences attention and multiple working memory items may affect attention through alternation.Protocol registrationThe stage 1 protocol for this Registered Report was accepted in principle on September 5, 2022. The protocol, as accepted by the journal, can be found at: https://doi.org/10.17605/OSF.IO/BE529.

Can items in visual working memory be shielded from visual interference while in use?

Can items in visual working memory be shielded from visual interference while in use?

Differently prioritized working memory items are differently protected from perceptual interference.

Differently prioritized working memory items are differently protected from perceptual interference.

Spatial Configuration Effects on the Dissociation between Active and Latent States in Visual Working Memory.

Visual working memory includes both active, recallable items and latent items not directly available for recall. During the online manipulation of active working memory, latent items gain robust retention. According to the dissociation account, active and passive memories exist in independent states, both of which take up their own cognitive resources. However, it is unclear whether dissociation is a universal phenomenon during memory maintenance. Given that memory information is retained as a binding of identity and location, the spatial proximity of memory items might shield the dissociation property. To test this, we adopted a retro-cue memory task where the cued and uncued items were separated in the active and latent states. In Experiment 1, the memory items were presented at a relatively large spatial distance. The results supported the dissociation account for well-separated items. However, Experiment 2 rejected the dissociation for closer-in items, possibly because items in visual working memory were spatially labeled. These findings suggest that while memory maintenance generally conforms to the dissociation account, the spatial configuration of memory items can affect the dissociation property between the active and latent neural states in visual working memory.

Changes in behavioral priority influence the accessibility of working memory content

Evolving behavioral goals require the existence of selection mechanisms that prioritize task-relevant working memory (WM) content for action. Selecting an item stored in WM is known to blunt and/or reverse information loss in stimulus-specific representations of that item reconstructed from human brain activity, but extant studies have focused on all-or-none circumstances that allow or disallow an agent to select one of several items stored in WM. Conversely, behavioral studies suggest that humans can flexibly assign different levels of priority to different items stored in WM, but how doing so influences neural representations of WM content is unclear. One possibility is that assigning different levels of priority to items in WM influences the quality of those representations, resulting in more robust neural representations of high- vs. low-priority WM content. A second – and non-exclusive – possibility is that asymmetries in behavioral priority influence how rapidly neural representations of high- vs. low-priority WM content can be selected and reported. We tested these possibilities in two experiments by decoding high- and low-priority WM content from EEG recordings obtained while human volunteers performed a retrospectively cued WM task. Probabilistic changes in the behavioral relevance of a remembered item had no effect on our ability to decode it from EEG signals; instead, these changes influenced the latency at which above-chance decoding performance was reached. Thus, our results indicate that probabilistic changes in the behavioral relevance of WM content influence the ease with which memories can be selected independently of their strength.

Spectrotemporal content of human auditory working memory represented in functional connectivity patterns

Recent research suggests that working memory (WM), the mental sketchpad underlying thinking and communication, is maintained by multiple regions throughout the brain. Whether parts of a stable WM representation could be distributed across these brain regions is, however, an open question. We addressed this question by examining the content-specificity of connectivity-pattern matrices between subparts of cortical regions-of-interest (ROI). These connectivity patterns were calculated from functional MRI obtained during a ripple-sound auditory WM task. Statistical significance was assessed by comparing the decoding results to a null distribution derived from a permutation test considering all comparable two- to four-ROI connectivity patterns. Maintained WM items could be decoded from connectivity patterns across ROIs in frontal, parietal, and superior temporal cortices. All functional connectivity patterns that were specific to maintained sound content extended from early auditory to frontoparietal cortices. Our results demonstrate that WM maintenance is supported by content-specific patterns of functional connectivity across different levels of cortical hierarchy.

Working memory control dynamics follow principles of spatial computing

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.

Working Memory Precision and Associative Binding in Mild Cognitive Impairment

ABSTRACT To better understand working memory (WM) deficits in Mild Cognitive Impairment (MCI), we examined information precision and associative binding in WM in 21 participants with MCI, compared to 16 healthy controls, using an item-location delayed reproduction task. WM, along with other executive functions (i.e. Trail Making Task (TMT) and Stroop task), were measured before and after a 2-h nap. The napping manipulation was intended as an exploratory element to this study exploring potential impacts of napping on executive functions. Compared to healthy participants, participants with MCI exhibited inferior performance not only in identifying encoded WM items but also on item-location associative binding and location precision even when only one item was involved. We also found changes on TMT and Stroop tasks in MCI, reflecting inferior attention and inhibitory control. Post-napping performance improved in most of these WM and other executive measures, both in MCI and their healthy peers. Our study shows that associative binding and WM precision can reliably differentiate MCIs from their healthy peers. Additionally, most measures showed no differential effect of group pre- and post-napping. These findings may contribute to better understanding cognitive deficits in MCI therefore improving the diagnosis of MCI.

10 Minutes Frontal 40 Hz tACS-Effects on Working Memory Tested by Luck-Vogel Task.

Working memory is a cognitive process that involves short-term active maintenance, flexible updating, and processing of goal- or task-relevant information. All frequency bands are involved in working memory. The activities of the theta and gamma frequency bands in the frontoparietal network are highly involved in working memory processes; theta oscillations play a role in the temporal organization of working memory items, and gamma oscillations influence the maintenance of information in working memory. Transcranial alternating current stimulation (tACS) results in frequency-specific modulation of endogenous oscillations and has shown promising results in cognitive neuroscience. The electrophysiological and behavioral changes induced by the modulation of endogenous gamma frequency in the prefrontal cortex using tACS have not been extensively studied in the context of working memory. Therefore, we aimed to investigate the effects of frontal gamma-tACS on working memory outcomes. We hypothesized that a 10-min gamma tACS administered over the frontal cortex would significantly improve working memory outcomes. Young healthy participants performed Luck-Vogel cognitive behavioral tasks with simultaneous pre- and post-intervention EEG recording (Sham versus 40 Hz tACS). Data from forty-one participants: sham (15 participants) and tACS (26 participants), were used for the statistical and behavioral analysis. The relative changes in behavioral outcomes and EEG due to the intervention were analyzed. The results show that tACS caused an increase in the power spectral density in the high beta and low gamma EEG bands and a decrease in left-right coherence. On the other hand, tACS had no significant effect on success rates and response times. Conclusion: 10 min of frontal 40 Hz tACS was not sufficient to produce detectable behavioral effects on working memory, whereas electrophysiological changes were evident. The limitations of the current stimulation protocol and future directions are discussed in detail in the following sections.