The relationships between working memory and long-term memory

Recent studies have led to the proposal that working memory operates not as a gateway between sensory input and long-term memory but as a workspace. The core of argument is that access to acquired knowledge and prior learning occurs before information becomes available to working memory. This proposition is a way to accomodate Baddeley's multiple component working memory model and the view that considers that working memory is nothing other than temporary activations of representations and procedures in long-term memory. However, this ‘workspace’ conception of working memory raises the question of the relationships between the central executive system and long-term memory.     

Changes in Brain Network Efficiency and Working Memory Performance in Aging

Working memory is a complex psychological construct referring to the temporary storage and active processing of information. We used functional connectivity brain network metrics quantifying local and global efficiency of information transfer for predicting individual variability in working memory performance on an n-back task in both young (n = 14) and older (n = 15) adults. Individual differences in both local and global efficiency during the working memory task were significant predictors of working memory performance in addition to age (and an interaction between age and global efficiency). Decreases in local efficiency during the working memory task were associated with better working memory performance in both age cohorts. In contrast, increases in global efficiency were associated with much better working performance for young participants; however, increases in global efficiency were associated with a slight decrease in working memory performance for older participants. Individual differences in local and global efficiency during resting-state sessions were not significant predictors of working memory performance. Significant group whole-brain functional network decreases in local efficiency also were observed during the working memory task compared to rest, whereas no significant differences were observed in network global efficiency. These results are discussed in relation to recently developed models of age-related differences in working memory.     

Mental imagery and visual working memory

Visual working memory provides an essential link between past and future events. Despite recent efforts, capacity limits, their genesis and the underlying neural structures of visual working memory remain unclear. Here we show that performance in visual working memory--but not iconic visual memory--can be predicted by the strength of mental imagery as assessed with binocular rivalry in a given individual. In addition, for individuals with strong imagery, modulating the background luminance diminished performance on visual working memory and imagery tasks, but not working memory for number strings. This suggests that luminance signals were disrupting sensory-based imagery mechanisms and not a general working memory system. Individuals with poor imagery still performed above chance in the visual working memory task, but their performance was not affected by the background luminance, suggesting a dichotomy in strategies for visual working memory: individuals with strong mental imagery rely on sensory-based imagery to support mnemonic performance, while those with poor imagery rely on different strategies. These findings could help reconcile current controversy regarding the mechanism and location of visual mnemonic storage.     

Visual working memory capacity and proactive interference

Background

Visual working memory capacity is extremely limited and appears to be relatively immune to practice effects or the use of explicit strategies. The recent discovery that visual working memory tasks, like verbal working memory tasks, are subject to proactive interference, coupled with the fact that typical visual working memory tasks are particularly conducive to proactive interference, suggests that visual working memory capacity may be systematically under-estimated.

Methodology/Principal Findings

Working memory capacity was probed behaviorally in adult humans both in laboratory settings and via the Internet. Several experiments show that although the effect of proactive interference on visual working memory is significant and can last over several trials, it only changes the capacity estimate by about 15%.

Conclusions/Significance

This study further confirms the sharp limitations on visual working memory capacity, both in absolute terms and relative to verbal working memory. It is suggested that future research take these limitations into account in understanding differences across a variety of tasks between human adults, prelinguistic infants and nonlinguistic animals.     

Transfer after Working Memory Updating Training

During the past decade, working memory training has attracted much interest. However, the training outcomes have varied between studies and methodological problems have hampered the interpretation of results. The current study examined transfer after working memory updating training by employing an extensive battery of pre-post cognitive measures with a focus on near transfer. Thirty-one healthy Finnish young adults were randomized into either a working memory training group or an active control group. The working memory training group practiced with three working memory tasks, while the control group trained with three commercial computer games with a low working memory load. The participants trained thrice a week for five weeks, with one training session lasting about 45 minutes. Compared to the control group, the working memory training group showed strongest transfer to an n-back task, followed by working memory updating, which in turn was followed by active working memory capacity. Our results support the view that working memory training produces near transfer effects, and that the degree of transfer depends on the cognitive overlap between the training and transfer measures.     

Forebrain NR2B overexpression facilitating the prefrontal cortex long-term potentiation and enhancing working memory function in mice

Prefrontal cortex plays an important role in working memory, attention regulation and behavioral inhibition. Its functions are associated with NMDA receptors. However, there is little information regarding the roles of NMDA receptor NR2B subunit in prefrontal cortical synaptic plasticity and prefrontal cortex-related working memory. Whether the up-regulation of NR2B subunit influences prefrontal cortical synaptic plasticity and working memory is not yet clear. In the present study, we measured prefrontal cortical synaptic plasticity and working memory function in NR2B overexpressing transgenic mice. In vitro electrophysiological data showed that overexpression of NR2B specifically in the forebrain region resulted in enhancement of prefrontal cortical long-term potentiation (LTP) but did not alter long-term depression (LTD). The enhanced LTP was completely abolished by a NR2B subunit selective antagonist, Ro25-6981, indicating that overexpression of NR2B subunit is responsible for enhanced LTP. In addition, NR2B transgenic mice exhibited better performance in a set of working memory paradigms including delay no-match-to-place T-maze, working memory version of water maze and odor span task. Our study provides evidence that NR2B subunit of NMDA receptor in prefrontal cortex is critical for prefrontal cortex LTP and prefrontal cortex-related working memory.     

Distinct Visual Working Memory Systems for View-Dependent and View-Invariant Representation

Background

How do people sustain a visual representation of the environment? Currently, many researchers argue that a single visual working memory system sustains non-spatial object information such as colors and shapes. However, previous studies tested visual working memory for two-dimensional objects only. In consequence, the nature of visual working memory for three-dimensional (3D) object representation remains unknown.

Methodology/Principal Findings

Here, I show that when sustaining information about 3D objects, visual working memory clearly divides into two separate, specialized memory systems, rather than one system, as was previously thought. One memory system gradually accumulates sensory information, forming an increasingly precise view-dependent representation of the scene over the course of several seconds. A second memory system sustains view-invariant representations of 3D objects. The view-dependent memory system has a storage capacity of 3–4 representations and the view-invariant memory system has a storage capacity of 1–2 representations. These systems can operate independently from one another and do not compete for working memory storage resources.

Conclusions/Significance

These results provide evidence that visual working memory sustains object information in two separate, specialized memory systems. One memory system sustains view-dependent representations of the scene, akin to the view-specific representations that guide place recognition during navigation in humans, rodents and insects. The second memory system sustains view-invariant representations of 3D objects, akin to the object-based representations that underlie object cognition.     

Continuous Theta Burst Stimulation over the Left Dorsolateral Prefrontal Cortex Decreases Medium Load Working Memory Performance in Healthy Humans

The dorsolateral prefrontal cortex (DLPFC) plays a key role in working memory. Evidence indicates that transcranial magnetic stimulation (TMS) over the DLPFC can interfere with working memory performance. Here we investigated for how long continuous theta-burst stimulation (cTBS) over the DLPFC decreases working memory performance and whether the effect of cTBS on performance depends on working memory load. Forty healthy young subjects received either cTBS over the left DLPFC or sham stimulation before performing a 2-, and 3-back working memory letter task. An additional 0-back condition served as a non-memory-related control, measuring general attention. cTBS over the left DLPFC significantly impaired 2-back working memory performance for about 15 min, whereas 3-back and 0-back performances were not significantly affected. Our results indicate that the effect of left DLPFC cTBS on working memory performance lasts for roughly 15 min and depends on working memory load.     

An Exploration of the Relations between External Representations and Working Memory

It is commonly hypothesized that external representations serve as memory aids and improve task performance by means of expanding the limited capacity of working memory. However, very few studies have directly examined this memory aid hypothesis. By systematically manipulating how information is available externally versus internally in a sequential number comparison task, three experiments were designed to investigate the relation between external representations and working memory. The experimental results show that when the task requires information from both external representations and working memory, it is the interaction of information from the two sources that determines task performance. In particular, when information from the two sources does not match well, external representations hinder instead of enhance task performance. The study highlights the important role the coordination among different representations plays in distributed cognition. The general relations between external representations and working memory are discussed.     

Effective visual working memory capacity: an emergent effect from the neural dynamics in an attractor network

The study of working memory capacity is of outmost importance in cognitive psychology as working memory is at the basis of general cognitive function. Although the working memory capacity limit has been thoroughly studied, its origin still remains a matter of strong debate. Only recently has the role of visual saliency in modulating working memory storage capacity been assessed experimentally and proved to provide valuable insights into working memory function. In the computational arena, attractor networks have successfully accounted for psychophysical and neurophysiological data in numerous working memory tasks given their ability to produce a sustained elevated firing rate during a delay period. Here we investigate the mechanisms underlying working memory capacity by means of a biophysically-realistic attractor network with spiking neurons while accounting for two recent experimental observations: 1) the presence of a visually salient item reduces the number of items that can be held in working memory, and 2) visually salient items are commonly kept in memory at the cost of not keeping as many non-salient items.OUR MODEL SUGGESTS THAT WORKING MEMORY CAPACITY IS DETERMINED BY TWO FUNDAMENTAL PROCESSES: encoding of visual items into working memory and maintenance of the encoded items upon their removal from the visual display. While maintenance critically depends on the constraints that lateral inhibition imposes to the mnemonic activity, encoding is limited by the ability of the stimulated neural assemblies to reach a sufficiently high level of excitation, a process governed by the dynamics of competition and cooperation among neuronal pools. Encoding is therefore contingent upon the visual working memory task and has led us to introduce the concept of effective working memory capacity (eWMC) in contrast to the maximal upper capacity limit only reached under ideal conditions.     

Spatial cognition and memory: a reversible lesion with lidocaine into the anteromedial/posterior parietal cortex (AM/PPC) affects differently working and long-term memory on two foraging tasks

Place memory is relevant for exploration and forage behaviour. When food supply is dispersed, a win-shift has advantage over a win-stay strategy. In the Olton Octagonal Maze, the rat follows a win-shift strategy using working memory. However, in the Olton 4x4 version, the rat follows a win-stay strategy, using both working and long-term memories. It has been suggested that the neocortex is required for the resolution of tasks demanding long-term, but not for that demanding working memory alone. The role of anteromedial/posterior parietal cortex (AM/PPC) was investigated here, using a reversible lesion induced by intracerebral lidocaine infusion. Long-Evans rats were implanted with guide cannulae into the AM/PPC and trained in an Olton 4x4 maze, counting working and long-term memory errors after a delay. Then, the animals were infused with lidocaine or saline during the delay phase and tested for three days. Another series of animals, treated as before, was tested in an Olton Octagonal Maze and subjected to the same injection schedule. In the Olton 4x4 Maze, lidocaine produced a significant increase in working and long-term memory errors, compared to saline and post-lidocaine conditions. In contrast, in the Olton Octagonal Maze, lidocaine did not induce any effect on working memory errors. Thus, AM/PPC is required when both working with previous information and long-term memories are needed, but not when only working memory is required, as it happens under ethological conditions. Whenever food supply is dispersed, a win-shift strategy is preferable.     

Visuospatial Working Memory Capacity Predicts Physiological Arousal in a Narrative Task

Physiological arousal that occurs during narrative production is thought to reflect emotional processing and cognitive effort (Bar-Haim et al. in Dev Psychobiol 44:238–249, 2004). The purpose of this study was to determine whether individual differences in visuospatial working memory and/or verbal working memory capacity predict physiological arousal in a narrative task. Visuospatial working memory was a significant predictor of skin conductance level (SCL); verbal working memory was not. When visuospatial working memory interference was imposed, visuospatial working memory was no longer a significant predictor of SCL. Visuospatial interference also resulted in a significant reduction in SCL. Furthermore, listener ratings of narrative quality were contingent upon the visuospatial working memory resources of the narrator. Potential implications for educators and clinical practitioners are discussed.     

Dual N-Back Working Memory Training in Healthy Adults: A Randomized Comparison to Processing Speed Training

Enhancing cognitive ability is an attractive concept, particularly for middle-aged adults interested in maintaining cognitive functioning and preventing age-related declines. Computerized working memory training has been investigated as a safe method of cognitive enhancement in younger and older adults, although few studies have considered the potential impact of working memory training on middle-aged adults. This study investigated dual n-back working memory training in healthy adults aged 30–60. Fifty-seven adults completed measures of working memory, processing speed, and fluid intelligence before and after a 5-week web-based dual n-back or active control (processing speed) training program. Results: Repeated measures multivariate analysis of variance failed to identify improvements across the three cognitive composites, working memory, processing speed, and fluid intelligence, after training. Follow-up Bayesian analyses supported null findings for training effects for each individual composite. Findings suggest that dual n-back working memory training may not benefit working memory or fluid intelligence in healthy adults. Further investigation is necessary to clarify if other forms of working memory training may be beneficial, and what factors impact training-related benefits, should they occur, in this population.     

Stability of Whole Brain and Regional Network Topology within and between Resting and Cognitive States

Background

Graph-theory based analyses of resting state functional Magnetic Resonance Imaging (fMRI) data have been used to map the network organization of the brain. While numerous analyses of resting state brain organization exist, many questions remain unexplored. The present study examines the stability of findings based on this approach over repeated resting state and working memory state sessions within the same individuals. This allows assessment of stability of network topology within the same state for both rest and working memory, and between rest and working memory as well.

Methodology/Principal Findings

fMRI scans were performed on five participants while at rest and while performing the 2-back working memory task five times each, with task state alternating while they were in the scanner. Voxel-based whole brain network analyses were performed on the resulting data along with analyses of functional connectivity in regions associated with resting state and working memory. Network topology was fairly stable across repeated sessions of the same task, but varied significantly between rest and working memory. In the whole brain analysis, local efficiency, Eloc, differed significantly between rest and working memory. Analyses of network statistics for the precuneus and dorsolateral prefrontal cortex revealed significant differences in degree as a function of task state for both regions and in local efficiency for the precuneus. Conversely, no significant differences were observed across repeated sessions of the same state.

Conclusions/Significance

These findings suggest that network topology is fairly stable within individuals across time for the same state, but also fluid between states. Whole brain voxel-based network analyses may prove to be a valuable tool for exploring how functional connectivity changes in response to task demands.     

Cognitive Behavioral Performance of Untreated Depressed Patients with Mild Depressive Symptoms

This study evaluated the working memory performance of 18 patients experiencing their first onset of mild depression without treatment and 18 healthy matched controls. The results demonstrated that working memory impairment in patients with mild depression occurred when memorizing the position of a picture but not when memorizing the pictures themselves. There was no significant difference between the two groups in the emotional impact on the working memory, indicating that the attenuation of spatial working memory was not affected by negative emotion; however, cognitive control selectively affected spatial working memory. In addition, the accuracy of spatial working memory in the depressed patients was not significantly reduced, but the reaction time was significantly extended compared with the healthy controls. This finding indicated that there was no damage to memory encoding and function maintenance in the patients but rather only impaired memory retrieval, suggesting that the extent of damage to the working memory system and cognitive control abilities was associated with the corresponding depressive symptoms. The development of mild to severe depressive symptoms may be accompanied by spatial working memory damage from the impaired memory retrieval function extending to memory encoding and memory retention impairments. In addition, the impaired cognitive control began with an inadequate capacity to automatically process internal negative emotions and further extended to impairment of the ability to regulate and suppress external emotions. The results of the mood-congruent study showed that the memory of patients with mild symptoms of depression was associated with a mood-congruent memory effect, demonstrating that mood-congruent memory was a typical feature of depression, regardless of the severity of depression. This study provided important information for understanding the development of cognitive dysfunction.     

Developmental continuity of working memory from infancy to preschool age

The objective of the paper was to study the developmental continuity of working memory function from infancy to preschool age. At the age of 10 to 11 months 44 participants completed delayed response task (A-not-B) that measures working memory function. Between 5 and 7 years of age the same participants performed three tasks assessing working memory for temporal order in auditory and visual modalities and a control task measuring short-term visuospatial memory. The dependence of temporal-order memory at preschool age on individual level of infant working memory was found for all methods of measurement despite the differences in way of presentation and reproducing of the stimuli order. Results indicate direct continuity in the development of working memory function from infancy to preschool age.     

Impulsive Choice Predicts Poor Working Memory in Male Rats

A number of maladaptive behaviors and poor health outcomes (e.g., substance abuse, obesity) correlate with impulsive choice, which describes the tendency to prefer smaller, immediate rewards in lieu of larger, delayed rewards. Working memory deficits are often reported in those diagnosed with the same maladaptive behaviors. Human studies suggest that impulsive choice is associated with working memory ability but, to date, only one study has explored the association between working memory and impulsive choice in rats and no relation was reported. The current study reevaluated the association between working memory and impulsive choice in 19 male Long-Evans rats. Psychophysical adjusting procedures were used to quantify working memory (titrating-delay match-to-position procedure) and impulsive choice (adjusting delay procedure). Rats were partitioned into low- and high-impulsive groups based on performance in the impulsive choice task. Low-impulsive rats performed significantly better in the working memory assessment. Across all rats, impulsive choice was negatively correlated with working memory performance. These findings support the hypothesis that prefrontal cortex function, specifically, working memory, is related to impulsive choice. Future research might profitably examine the experimental variables designed to influence working memory to evaluate the effects of these variables on impulsive choice and maladaptive behaviors with which it is correlated.     

Use of an Eight-arm Radial Water Maze to Assess Working and Reference Memory Following Neonatal Brain Injury

Working and reference memory are commonly assessed using the land based radial arm maze. However, this paradigm requires pretraining, food deprivation, and may introduce scent cue confounds. The eight-arm radial water maze is designed to evaluate reference and working memory performance simultaneously by requiring subjects to use extra-maze cues to locate escape platforms and remedies the limitations observed in land based radial arm maze designs. Specifically, subjects are required to avoid the arms previously used for escape during each testing day (working memory) as well as avoid the fixed arms, which never contain escape platforms (reference memory). Re-entries into arms that have already been used for escape during a testing session (and thus the escape platform has been removed) and re-entries into reference memory arms are indicative of working memory deficits. Alternatively, first entries into reference memory arms are indicative of reference memory deficits. We used this maze to compare performance of rats with neonatal brain injury and sham controls following induction of hypoxia-ischemia and show significant deficits in both working and reference memory after eleven days of testing. This protocol could be easily modified to examine many other models of learning impairment.     

High visual working memory capacity in trait social anxiety

Working memory capacity is one of the most important cognitive functions influencing individual traits, such as attentional control, fluid intelligence, and also psychopathological traits. Previous research suggests that anxiety is associated with impaired cognitive function, and studies have shown low verbal working memory capacity in individuals with high trait anxiety. However, the relationship between trait anxiety and visual working memory capacity is still unclear. Considering that people allocate visual attention more widely to detect danger under threat, visual working memory capacity might be higher in anxious people. In the present study, we show that visual working memory capacity increases as trait social anxiety increases by using a change detection task. When the demand to inhibit distractors increased, however, high visual working memory capacity diminished in individuals with social anxiety, and instead, impaired filtering of distractors was predicted by trait social anxiety. State anxiety was not correlated with visual working memory capacity. These results indicate that socially anxious people could potentially hold a large amount of information in working memory. However, because of an impaired cognitive function, they could not inhibit goal-irrelevant distractors and their performance decreased under highly demanding conditions.     

Is the Link from Working Memory to Analogy Causal? No Analogy Improvements following Working Memory Training Gains

Analogical reasoning has been hypothesized to critically depend upon working memory through correlational data [1], but less work has tested this relationship through experimental manipulation [2]. An opportunity for examining the connection between working memory and analogical reasoning has emerged from the growing, although somewhat controversial, body of literature suggests complex working memory training can sometimes lead to working memory improvements that transfer to novel working memory tasks. This study investigated whether working memory improvements, if replicated, would increase analogical reasoning ability. We assessed participants’ performance on verbal and visual analogy tasks after a complex working memory training program incorporating verbal and spatial tasks [3], [4]. Participants’ improvements on the working memory training tasks transferred to other short-term and working memory tasks, supporting the possibility of broad effects of working memory training. However, we found no effects on analogical reasoning. We propose several possible explanations for the lack of an impact of working memory improvements on analogical reasoning.