Information In Working Memory Research Articles

The entorhinal-DG/CA3 pathway in the medial temporal lobe retains visual working memory of a simple surface feature.

Classic models consider working memory (WM) and long-term memory as distinct mental faculties that are supported by different neural mechanisms. Yet, there are significant parallels in the computation that both types of memory require. For instance, the representation of precise item-specific memory requires the separation of overlapping neural representations of similar information. This computation has been referred to as pattern separation, which can be mediated by the entorhinal-DG/CA3 pathway of the medial temporal lobe (MTL) in service of long-term episodic memory. However, although recent evidence has suggested that the MTL is involved in WM, the extent to which the entorhinal-DG/CA3 pathway supports precise item-specific WM has remained elusive. Here, we combine an established orientation WM task with high-resolution fMRI to test the hypothesis that the entorhinal-DG/CA3 pathway retains visual WM of a simple surface feature. Participants were retrospectively cued to retain one of the two studied orientation gratings during a brief delay period and then tried to reproduce the cued orientation as precisely as possible. By modeling the delay-period activity to reconstruct the retained WM content, we found that the anterior-lateral entorhinal cortex (aLEC) and the hippocampal DG/CA3 subfield both contain item-specific WM information that is associated with subsequent recall fidelity. Together, these results highlight the contribution of MTL circuitry to item-specific WM representation.

Humans, fish, spiders and bees inherited working memory and attention from their last common ancestor.

All brain processes that generate behaviour, apart from reflexes, operate with information that is in an "activated" state. This activated information, which is known as working memory (WM), is generated by the effect of attentional processes on incoming information or information previously stored in short-term or long-term memory (STM or LTM). Information in WM tends to remain the focus of attention; and WM, attention and STM together enable information to be available to mental processes and the behaviours that follow on from them. WM and attention underpin all flexible mental processes, such as solving problems, making choices, preparing for opportunities or threats that could be nearby, or simply finding the way home. Neither WM nor attention are necessarily conscious, and both may have evolved long before consciousness. WM and attention, with similar properties, are possessed by humans, archerfish, and other vertebrates; jumping spiders, honey bees, and other arthropods; and members of other clades, whose last common ancestor (LCA) is believed to have lived more than 600 million years ago. It has been reported that very similar genes control the development of vertebrate and arthropod brains, and were likely inherited from their LCA. Genes that control brain development are conserved because brains generate adaptive behaviour. However, the neural processes that generate behaviour operate with the activated information in WM, so WM and attention must have existed prior to the evolution of brains. It is proposed that WM and attention are widespread amongst animal species because they are phylogenetically conserved mechanisms that are essential to all mental processing, and were inherited from the LCA of vertebrates, arthropods, and some other animal clades.

Mountains of memory in a sea of uncertainty: Sampling the external world despite useful information in visual working memory

A large part of research on visual working memory (VWM) has traditionally focused on estimating its maximum capacity. Yet, humans rarely need to load up their VWM maximally during natural behavior, since visual information often remains accessible in the external world. Recent work, using paradigms that take into account the accessibility of information in the outside world, has indeed shown that observers utilize only one or two items in VWM before sampling from the external world again. One straightforward interpretation of this finding is that, in daily behavior, much fewer items are memorized than the typically reported capacity limits. Here, we first investigate whether this lower reliance on VWM when information is externally accessible might instead reflect resampling before VWM is actually depleted. To this aim we devised an online task, in which participants copied a model (six items in a 4x4 grid; always accessible) in an adjacent empty 4x4 grid. A key aspect of our paradigm is that we (unpredictably) interrupted participants just before inspection of the model with a 2-alternative-forced-choice (2-AFC) question, probing their VWM content. Critically, we observed above-chance performance on probes appearing just before model inspection. This finding shows that the external world was resampled, despite VWM still containing relevant information. We then asked whether increasing the cost of sampling causes participants to load up more information in VWM or, alternatively, to squeeze out more information from VWM (at the cost of making more errors). To manipulate the cost of resampling, we made it more difficult (specifically, more time-consuming) to access the model. We show that with increased cost of accessing the model (which lead to fewer, but longer model inspections), participants could place more items correctly immediately after sampling, and they kept attempting to place items for longer after their first error. These findings demonstrate that participants both encoded more information in VWM and made attempts to squeeze out more information from VWM when sampling became more costly. We argue that human observers constantly evaluate how certain they are of their VWM contents, and only use that VWM content of which their certainty exceeds a context-dependent “action threshold”. This threshold, in turn, depends on the trade-off between the cost of resampling and the benefits of making an action. We argue that considering the interplay between the available VWM contents and a context-dependent action threshold, is key for reconciling the traditional VWM literature with VWM use in our day-to-day behavior.

Manipulating information in visual working memory

Manipulating information in visual working memory

Reactivating ordinal position information from auditory sequence memory in human brains.

Retaining a sequence of events in their order is a core ability of many cognitive functions, such as speech recognition, movement control, and episodic memory. Although content representations have been widely studied in working memory (WM), little is known about how ordinal position information of an auditory sequence is retained in the human brain as well as its coding characteristics. In fact, there is still a lack of an efficient approach to directly accessing the stored ordinal position code during WM retention. Here, 31 participants performed an auditory sequence WM task with their brain activities recorded using electroencephalography (EEG). We developed new triggering events that could successfully reactivate neural representations of ordinal position during the delay period. Importantly, the ordinal position reactivation is further related to recognition behavior, confirming its indexing of WM storage. Furthermore, the ordinal position code displays an intriguing "stable-dynamic" format, i.e. undergoing the same dynamic neutral trajectory in the multivariate neural space during both encoding and retention (whenever reactivated). Overall, our results provide an effective approach to accessing the behaviorally-relevant ordinal position information in auditory sequence WM and reveal its new temporal characteristics.

The proportion of working memory items recoverable from long-term memory remains fixed despite adult aging.

We explored whether long-term memory (LTM) retrieval is constrained by working memory (WM) limitations, in 80 younger and 80 older adults. Participants performed a WM task with images of unique everyday items, presented at varying set sizes. Subsequently, we tested participants' LTM for items from the WM task and examined the ratio of LTM/WM retention. While older adults' WM and LTM were generally poorer than that of younger adults, their LTM deficit was no greater than what was predicted from their WM performance. The ability to encode WM information into LTM appeared immune to age-related cognitive decline. (PsycInfo Database Record (c) 2022 APA, all rights reserved).

A ventral stream-prefrontal cortex processing cascade enables working memory gating dynamics

The representation of incoming information, goals and the flexible processing of these are required for cognitive control. Efficient mechanisms are needed to decide when it is important that novel information enters working memory (WM) and when these WM ‘gates’ have to be closed. Compared to neural foundations of maintaining information in WM, considerably less is known about what neural mechanisms underlie the representational dynamics during WM gating. Using different EEG analysis methods, we trace the path of mental representations along the human cortex during WM gate opening and closing. We show temporally nested representational dynamics during WM gate opening and closing depending on multiple independent neural activity profiles. These activity profiles are attributable to a ventral stream-prefrontal cortex processing cascade. The representational dynamics start in the ventral stream during WM gate opening and WM gate closing before prefrontal cortical regions are modulated. A regional specific activity profile is shown within the prefrontal cortex depending on whether WM gates are opened or closed, matching overarching concepts of prefrontal cortex functions. The study closes an essential conceptual gap detailing the neural dynamics underlying how mental representations drive the WM gate to open or close to enable WM functions such as updating and maintenance.

Daily fluctuations in children’s working memory accuracy and precision: Variability at multiple time scales and links to daily sleep behavior and fluid intelligence

Children often experience better and worse days when performing cognitive tasks. Whether such fluctuations are systematic and how they are linked to fluctuations at faster time scales within days is less clear. To fill these gaps, we probed N = 108 fifth graders on WM tasks twice daily in morning and afternoon sessions, and also assessed nightly sleep behavior, over a period of four weeks using ambulatory assessment. Children systematically fluctuated in their recall of visuospatial and numerical information in WM across multiple time scales. These fluctuations showed consistencies but also discrepancies among each other. Especially, fast variability of memory precision across moments was related to load and fluid intelligence. Daily WM accuracy was positively coupled to sleep quality, but only in a subset of children with larger daily WM fluctuations. We propose that short-term WM fluctuations and their couplings to other time-varying constructs could help to explain long-term cognitive development. • Children show systematic intra-individual working memory (WM) fluctuations. • WM load particularly affected the rapid item-to-item variability of recall precision. • Lower momentary recall variability was linked to higher levels in fluid intelligence. • Nightly sleep quality predicted higher daily WM accuracy, but only in a subset of children. • These dynamics emphasize a nuanced distinction of faster and slower WM variability.

The semantic relatedness effect in serial recall: Deconfounding encoding and recall order

The ability to store information in verbal working memory (WM) closely interacts with our linguistic knowledge. For instance, we can hold semantically related words (e.g., “cat, dog, bird”) better in our WM than unrelated ones (e.g., “desk, pillow, mouse”). This study investigates boundary conditions of the beneficial effect of semantic relatedness of words on immediate memory for lists. Independently varying the encoding and recall order for lists of related words, we unraveled several mechanistic explanations of the semantic relatedness effect. We first tested a semantic cueing mechanism, according to which the recall of an item facilitates the recall of other semantically related items. We next disentangled an interactive activation versus feature overlap account of semantic relatedness. Whereas the former predicts that the semantic relatedness effect emerges from the temporal co-activation of related items, the latter predicts that it emerges from the superposition of semantic features bound to similar contexts. Our results demonstrate that semantic relatedness affects WM performance at the encoding stage of WM processing, which rules out semantic cueing as a plausible mechanism. Further, the temporal order in which words are presented was the most important determinant of the semantic relatedness effect, in agreement with the interactive activation account. This study supports a model in which semantic relatedness supports WM through interactive activation occurring in semantic long-term memory.

Perceptual comparisons modulate memory biases induced by new visual inputs.

It is well-established that stimulus-specific information in visual working memory (VWM) can be systematically biased by new perceptual inputs. These memory biases are commonly attributed to interference that arises when perceptual inputs are physically similar to VWM contents. However, recent work has suggested that explicitly comparing the similarity between VWM contents and new perceptual inputs modulates the size of memory biases above and beyond stimulus-driven effects. Here, we sought to directly investigate this modulation hypothesis by comparing the size of memory biases following explicit comparisons to those induced when new perceptual inputs are ignored (Experiment 1) or maintained in VWM alongside target information (Experiment 2). We found that VWM reports showed larger attraction biases following explicit perceptual comparisons than when new perceptual inputs were ignored or maintained in VWM. An analysis of participants' perceptual comparisons revealed that memory biases were amplified after perceptual inputs were endorsed as similar-but not dissimilar-to one's VWM representation. These patterns were found to persist even after accounting for variability in the physical similarity between the target and perceptual stimuli across trials, as well as the baseline memory precision between the distinct task demands. Together, these findings illustrate a causal role of perceptual comparisons in modulating naturally-occurring memory biases.

The impact of working memory on divergent thinking flexibility

The current study focuses on the relationship between working memory (WM) and flexibility, which is a dimension of divergent thinking. The research consisted of two experiments in which the participants’ task was to categorize given objects into as many categories as possible. We assumed that manipulation of the visual or relational complexity of a set of figures would respectively affect WM’s ability to maintain and process goal-relevant information. Additionally, the dual-task paradigm was used to observe the relocation of attention resources in the task because WM capacity is limited by attention resources. We hypothesized that both attentional control and the ability to maintain/process goal-relevant information in WM would affect the level of flexibility. The results show that an increased load on WM storage and on processing mechanisms leads to a decline in flexibility scores, which becomes especially apparent when the secondary task, which makes WM capacity more vulnerable, is applied. We conclude that the flexibility of divergent thinking is equally positively associated with the efficiency of WM mechanisms, i.e., storage and processing of current information, and the ability to use controlled attention.

Unintended emotions in the laboratory: Emotions incidentally induced by a standard visual working memory task relate to task performance.

The ability to temporarily hold information in visual working memory (VWM) is among the most crucial and most extensively examined human cognitive functions. Here, we empirically confirm previous speculations (a) that a standard VWM task arouses emotions in participants and (b) that these task-induced emotions are related to VWM performance. In a first qualitative study (N = 19), by adapting a qualitative method of inquiry, the think-aloud technique, we found that the task induced different positive and negative emotions, such as joy and anger, which varied on the inter- as well as on the intraindividual level. The emotional experiences seemed to be tied to the implicit achievement requirement of the VWM task (getting it right vs. wrong). Encouraged by these findings, two quantitative studies (N = 45, and N = 44, respectively) revealed that VWM performance was positively linked to joy and pride, and negatively linked to anger, frustration, and boredom on the inter- and on the intraindividual level. Notably, these emotions were also affected by an experimental manipulation of task difficulty (set size 4 vs. 8). Further, the findings from Study 3 were replicated in a fourth high-powered online study (N = 110). This research is the first to demonstrate that a task designed to measure VWM in itself triggers emotions, specifically achievement emotions, which, in turn, are linked with VWM performance. Our findings suggest that these task-induced emotions should be considered as potential confounding variables in future research on VWM and in cognitive research in general. (PsycInfo Database Record (c) 2022 APA, all rights reserved).

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.

Gaze behaviour: A window into distinct cognitive processes revealed by the Tower of London test

The analysis of gaze behaviour during complex tasks provides a promising non-invasive method to examine how specific eye movement patterns relate to various aspects of cognition and action. Notably, the association between aspects of gaze behaviour and subsequent goal-directed action during high-level visuospatial problem solving remains elusive. Therefore, the current study comprehensively examined gaze behaviour using traditional and entropy-based gaze analyses in healthy adults (N = 27) while they performed the Freiburg version of the Tower of London task. Results demonstrated that both gaze analyses provided crucial temporal and spatial information related to planning, solution elaboration and execution. Specifically, gaze biases toward task-relevant areas (i.e., the work space) and an increase in gaze complexity (i.e., gaze transition entropy) during optimal performance reflected changes in cognitive demands as task difficulty increased. A comparison between optimal and non-optimal performance revealed sub-optimal gaze patterns that occurred in the early stages of planning, which were taken to reflect poor information extraction from the task environment and impaired maintenance of information in visuospatial working memory. Gaze behaviour during movement execution indicated an increased need to extract and process information from the goal space. Consequently, movement execution time increased in order to reverse erroneous movements and re-sequence the problem solution. Taken together, the traditional and entropy-based gaze analyses applied in the present study provide a promising approach to identify eye movement patterns that support neurocognitive performance on tasks relying on visuospatial planning and problem solving.

Individual differences in executive attention and inhibitory control are related to spatial memory biases in adults.

Executive attention is involved in working memory; however, the role of executive attention in the maintenance of information in spatial working memory is debated. This study examined whether inhibitory control was related to spatial working memory biases in adults in a simple spatial memory task where participants had to remember one location on an otherwise blank computer screen. On some trials, a distractor was presented during the maintenance period. Eighty-four participants completed the spatial working memory task and a battery of cognitive control measures. When a distractor was presented during the maintenance period of the spatial memory task, performance on two of the cognitive control measures, a measure of overall attention and a measure of inhibitory control was related to memory errors. When a distractor was not presented during the spatial memory task, memory errors were not related to performance on the cognitive control tasks. Overall, these effects demonstrated that attention is related to maintaining locations in spatial working memory in adults, and inhibitory control may also be related such that those with more efficient inhibitory control were less influenced by distractors presented during the maintenance period.

Sequential versus simultaneous presentation of memoranda in verbal working memory: (How) does it matter?

To-be-memorized information in verbal working memory (WM) can be presented sequentially, like in oral language, and simultaneously, like in written language. Few studies have addressed the importance and implications for verbal WM processing of these two presentation modes. While sequential presentation may favor discrete, temporal encoding processes, simultaneous presentation may favor spatial encoding processes. We compared immediate serial recall tasks for sequential versus simultaneous word list presentation with a specific focus on serial position curves of recall performance, transposition gradients, and the nature of serial order errors. First, we observed higher recall performance in the simultaneous compared to the sequential conditions, with a particularly large effect at end-of-list items. Moreover, results showed more transposition errors between non-adjacent items for the sequential condition, as well as more omission errors especially for start-of-list items. This observation can be explained in terms of differences in refreshing opportunities for start-of-list items during encoding between conditions. This study shows that the presentation mode of sequential material can have a significant impact on verbal WM performance, with an advantage for simultaneous encoding of sequence information.

Recruitment of a long-term memory supporting neural network during repeated maintenance of a multi-item abstract visual image in working memory

Humans can flexibly transfer information between different memory systems. Information in visual working memory (VWM) can for instance be stored in long-term memory (LTM). Conversely, information can be retrieved from LTM and temporarily held in WM when needed. It has previously been suggested that a neural transition from parietal- to midfrontal activity during repeated visual search reflects transfer of information from WM to LTM. Whether this neural transition indeed reflects consolidation and is also observed when memorizing a rich visual scene (rather than responding to a single target), is not known. To investigate this, we employed an EEG paradigm, in which abstract six-item colour-arrays were repeatedly memorized and explicitly visualized, or merely attended to. Importantly, we tested the functional significance of a potential neural shift for longer-term consolidation in a subsequent recognition task. Our results show a gradually enhanced- and sustained modulation of the midfrontal P170 component and a decline in parietal CDA, during repeated WM maintenance. Improved recollection/visualization of memoranda upon WM-cueing, was associated with contralateral parietal- and right temporal activity. Importantly, only colour-arrays previously held in WM, induced a greater midfrontal P170-response, together with left temporal- and late centro-parietal activity, upon re-exposure. These findings provide evidence for recruitment of an LTM-supporting neural network which facilitates visual WM maintenance.

Classification images reveal uniform decay of facial feature information in visual working memory

Classification images reveal uniform decay of facial feature information in visual working memory

Shared Representational Formats for Information Maintained in Working Memory and Information Retrieved from Long-Term Memory.

Current theories propose that the short-term retention of information in working memory (WM) and the recall of information from long-term memory (LTM) are supported by overlapping neural mechanisms in occipital and parietal cortex. However, the extent of the shared representations between WM and LTM is unclear. We designed a spatial memory task that allowed us to directly compare the representations of remembered spatial information in WM and LTM with carefully matched behavioral response precision between tasks. Using multivariate pattern analyses on functional magnetic resonance imaging data, we show that visual memories were represented in a sensory-like code in both memory tasks across retinotopic regions in occipital and parietal cortex. Regions in lateral parietal cortex also encoded remembered locations in both tasks, but in a format that differed from sensory-evoked activity. These results suggest a striking correspondence in the format of representations maintained in WM and retrieved from LTM across occipital and parietal cortex. On the other hand, we also show that activity patterns in nearly all parietal regions, but not occipital regions, contained information that could discriminate between WM and LTM trials. Our data provide new evidence for theories of memory systems and the representation of mnemonic content.

The Neural Representation of Ordinal Information: Domain-Specific or Domain-General?

Ordinal processing allows for the representation of the sequential relations between stimuli and is a fundamental aspect of different cognitive domains such as verbal working memory (WM), language and numerical cognition. Several studies suggest common ordinal coding mechanisms across these different domains but direct between-domain comparisons of ordinal coding are rare and have led to contradictory evidence. This fMRI study examined the commonality of ordinal representations across the WM, the number, and the letter domains by using a multivoxel pattern analysis approach and by focusing on triplet stimuli associated with robust ordinal distance effects. Neural patterns in fronto-parietal cortices distinguished ordinal distance in all domains. Critically, between-task predictions of ordinal distance in fronto-parietal cortices were robust between serial order WM, alphabetical order judgment but not when involving the numerical order judgment tasks. Moreover, frontal ROIs further supported between-task prediction of distance for the luminance judgment control task, the serial order WM, and the alphabetical tasks. These results suggest that common neural substrates characterize processing of ordinal information in WM and alphabetical but not numerical domains. This commonality, particularly in frontal cortices, may however reflect attentional control processes involved in judging ordinal distances rather than the intervention of domain-general ordinal codes.