Working memory retrieval differences between medial temporal lobe epilepsy patients and controls: A three memory layer approach

Multi-store models of working memory (WM) have given way to more dynamic approaches that conceive WM as an activated subset of long-term memory (LTM). The resulting framework considers that memory representations are governed by a hierarchy of accessibility. The activated part of LTM holds representations in a heightened state of activation, some of which can reach a state of immediate accessibility according to task demands. Recent neuroimaging studies have studied the neural basis of retrieval information with different states of accessibility. It was found that the medial temporal lobe (MTL) was involved in retrieving information within immediate access store and outside this privileged zone. In the current study we further explored the contribution of MTL to WM retrieval by analyzing the consequences of MTL damage to this process considering the state of accessibility of memory representations. The performance of a group of epilepsy patients with left hippocampal sclerosis in a 12-item recognition task was compared with that of a healthy control group. We adopted an embedded model of WM that distinguishes three components: the activated LTM, the region of direct access, and a single-item focus of attention. Groups did not differ when retrieving information from single-item focus, but patients were less accurate retrieving information outside focal attention, either items from LTM or items expected to be in the WM range. Analyses focused on items held in the direct access buffer showed that consequences of MTL damage were modulated by the level of accessibility of memory representations, producing a reduced capacity.     

Neuroimaging studies of working memory:

We performed meta-analyses on 60 neuroimaging (PET and fMRI) studies of working memory (WM), considering three types of storage material (spatial, verbal, and object), three types of executive function (continuous updating of WM, memory for temporal order, and manipulation of information in WM), and interactions between material and executive function. Analyses of material type showed the expected dorsal-ventral dissociation between spatial and nonspatial storage in the posterior cortex, but not in the frontal cortex. Some support was found for left frontal dominance in verbal WM, but only for tasks with low executive demand. Executive demand increased right lateralization in the frontal cortex for spatial WM. Tasks requiring executive processing generally produce more dorsal frontal activations than do storage-only tasks, but not all executive processes show this pattern. Brodmann’s areas (BAs) 6, 8, and 9, in the superior frontal cortex, respond most when WM must be continuously updated and when memory for temporal order must be maintained. Right BAs 10 and 47, in the ventral frontal cortex, respond more frequently with demand for manipulation (including dual-task requirements or mental operations). BA 7, in the posterior parietal cortex, is involved in all types of executive function. Finally, we consider a potential fourth executive function: selective attention to features of a stimulus to be stored in WM, which leads to increased probability of activating the medial prefrontal cortex (BA 32) in storage tasks.     

Working memory, perceptual priming, and the perception of hierarchical forms: Opposite effects of priming and working memory without memory refreshing

Previous research has shown that stimuli held in working memory (WM) can influence spatial attention. Using Navon stimuli, we explored whether and how items in WM affect the perception of visual targets at local and global levels in compound letters. Participants looked for a target letter presented at a local or global level while holding a regular block letter as a memory item. An effect of holding the target’s identity in WM was found. When memory items and targets were the same, performance was better than in a neutral condition when the memory item did not appear in the hierarchical letter (a benefit from valid cuing). When the memory item matched the distractor in the hierarchical stimulus, performance was worse than in the neutral baseline (a cost on invalid trials). These effects were greatest when the WM cue matched the global level of the hierarchical stimulus, suggesting that WM biases attention to the global level of form. Interestingly, in a no-memory priming condition, target perception was faster in the invalid condition than in the neutral baseline, reversing the effect in the WM condition. A further control experiment ruled out the effects of WM being due to participants’ refreshing their memory from the hierarchical stimulus display. The data show that information in WM biases the selection of hierarchical forms, whereas priming does not. Priming alters the perceptual processing of repeated stimuli without biasing attention.     

Dynamic states in working memory modulate guidance of visual attention: Evidence from an n-back paradigm

Items held in working memory (WM) can automatically bias attention when they reappear in visual displays. Recent evidence, however, suggests that WM biases of attention may be reduced under certain conditions, for example with increasing memory load. We employed a dual task paradigm to investigate how WM biases are affected by dynamic updating of memory contents. 1-back and 2-back versions of a memory task with colour stimuli were interrupted at intervals by an unrelated visual search task. Reappearance in the search display of the item that was currently active in WM guided attention, while suppressed or inactive items did not. We conclude that the rapid updating of memory contents facilitates the shifting of memory representations into different activity states on a moment-to-moment basis. The finding is consistent with models that propose that only one item can be “active” in WM at any one time to guide attention.     

What happens to an individual visual working memory representation when it is interrupted?

This study tested the hypothesis that even the simplest cognitive tasks require the storage of information in working memory (WM), distorting any information that was previously stored in WM. Experiment 1 tested this hypothesis by requiring observers to perform a simple letter discrimination task while they were holding a single orientation in WM. We predicted that performing the task on the interposed letter stimulus would cause the orientation memory to become less precise and more categorical compared to when the letter was absent or when it was present but could be ignored. This prediction was confirmed. Experiment 2 tested the modality specificity of this effect by replacing the visual letter discrimination task with an auditory pitch discrimination task. Unlike the interposed visual stimulus, the interposed auditory stimulus produced little or no disruption of WM, consistent with the use of modality‐specific representations. Thus, performing a simple visual discrimination task, but not a simple auditory discrimination task, distorts information about a single feature being maintained in visual WM. We suggest that the interposed task eliminates information stored within the focus of attention, leaving behind a WM representation outside the focus of attention that is relatively imprecise and categorical.     

Serial attention within working memory

It is proposed that people are limited to attending to just one “object” in working memory (WM) at any one time. Consequently, many cognitive tasks, and much of everyday thought, necessitate switches between WM items. The research to be presented measured the time involved in switching attention between objects in WM and sought to elaborate the processes underlying such switches. Two experiments required subjects to maintain two running counts; the order in which the counts were updated necessitated frequent switches between them. Even after intensive practice, a time cost was incurred when subjects updated the two counts in succession, relative to updating the same count twice. This time cost was interpreted as being due to a distinct switching mechanism that controls an internal focus of attention large enough for just one object (count) at a time. This internal focus of attention is a subset of WM (Cowan, 1988). Alternative visual and conceptual repetition-priming and memory retrieval explanations for the cost involved in switching between items in WM are addressed.     

Working memory load can both improve and impair selective attention: Evidence from the Navon paradigm

Selective attention to relevant targets has been shown to depend on the availability of working memory (WM). Under conditions of high WM load, processing of irrelevant distractors is enhanced. Here we showed that this detrimental effect of WM load on selective attention efficiency is reversed when the task requires global- rather than local-level processing. Participants were asked to attend to either the local or the global level of a hierarchical Navon stimulus while keeping either a low or a high load in WM. In line with previous findings, during attention to the local level, distractors at the global level produced more interference under high than under low WM load. By contrast, loading WM had the opposite effect of improving selective attention during attention to the global level. The findings demonstrate that the impact of WM load on selective attention is not invariant, but rather is dependent on the level of the to-be-attended information.     

Distractor interference stays constant despite variation in working memory load

Previous studies have shown that working memory (WM) plays an important role in selective attention, so that high WM load leads to inefficient distractor inhibition, in comparison with low WM load. In the present study, we examined the effect of WM on distractor processing while the extent of attentional focus was held constant. Our results show that WM load affected distractor processing only when it was positively correlated with the extent of attentional focus. When the latter was held constant, the effect of WM became negligible. Furthermore, when low WM load was paired with a wide attentional focus and high WM load was matched with a narrow attentional focus, greater distractor processing was found when the WM load was low than when it was high. These results suggest that efficient distractor inhibition may require only minimal WM resources and that the effect of WM on distractor processing is more complex than has previously been assumed.     

Interference with prior knowledge while writing from sources: Effects of working memory capacity

Writing from sources requires the processing of both external (documentary) and internal (long-term memory; LTM) sources of information, which can sometimes interfere with each other. We hypothesised that writers with high working memory (WM) capacity would resolve interference faster than those with low-capacity WM. However, results showed that writing pause duration was shorter for low-capacity writers. This result suggests that when WM resources are limited, access to domain knowledge in LTM is restricted, thereby avoiding interference between the two sources. Results are discussed within the broader context of text composition from sources and individual differences.     

Exploring the forgetting mechanisms in working memory: Evidence from a reasoning span test

In working memory (WM) span tests participants have to maintain to-be-remembered information while processing other, potentially distracting, information. Previous studies have shown that WM span scores are greater when span lists start with a long processing task and end with a short processing task than when these processing tasks are presented in the reverse order (e.g., Towse, Hitch, & Hutton, 2000). In Experiment 1, we obtained a similar stimulus order effect in a reasoning span test, using reasoning sentences that were equated for length in terms of the number of constituent words, but which differed in processing complexity; span scores were greater when lists began with a complex sentence and ended with a simple sentence than when this stimulus order was reversed. In Experiment 2, the stimulus order effect was not found when processing duration was held constant while sentence complexity was varied using a computer-paced moving window presentation paradigm. These results suggest that duration-based constraints can affect degree of forgetting independently of the load generated during processing phases in WM span performance and therefore imply that time-related forgetting can occur in WM span tests, particularly when the difficulty of the processing component blocks active maintenance of to-be-remembered material.     

Short-term memory and working memory as indices of children's cognitive skills

In the current literature, empirical and conceptual distinctions have been drawn between a more or less passive short-term memory (STM) system and a more dynamic working memory (WM) system. Distinct tasks have been developed to measure their capacity and research has generally shown that, for adults, WM, and not STM, is a reliable predictor of general cognitive ability. However, the locus of the differences between the tasks has received little attention. We present data from children concerning measures of matrices reasoning ability, reading, and numerical skill along with forward and backward order serial recall of WM, STM, and STM with articulatory suppression tasks. As indices of children's cognitive skills, STM and WM are shown to be rather similar in terms of memory per se. Neither the opportunity for rehearsal nor task complexity provides satisfactory explanations for differences between memory tests.     

Short-term memory and working memory as indices of children's cognitive skills

In the current literature, empirical and conceptual distinctions have been drawn between a more or less passive short-term memory (STM) system and a more dynamic working memory (WM) system. Distinct tasks have been developed to measure their capacity and research has generally shown that, for adults, WM, and not STM, is a reliable predictor of general cognitive ability. However, the locus of the differences between the tasks has received little attention. We present data from children concerning measures of matrices reasoning ability, reading, and numerical skill along with forward and backward order serial recall of WM, STM, and STM with articulatory suppression tasks. As indices of children's cognitive skills, STM and WM are shown to be rather similar in terms of memory per se. Neither the opportunity for rehearsal nor task complexity provides satisfactory explanations for differences between memory tests.     

Making working memory work: The effects of extended practice on focus capacity and the processes of updating,forward access,and random access

We investigated the effects of 10 h of practice on variations of the N-Back task to investigate the processes underlying possible expansion of the focus of attention within working memory. Using subtractive logic, we showed that random access (i.e., Sternberg-like search) yielded a modest effect (a 50% increase in speed) whereas the processes of forward access (i.e., retrieval in order, as in a standard N-Back task) and updating (i.e., changing the contents of working memory) were executed about 5 times faster after extended practice. We additionally found that extended practice increased working memory capacity as measured by the size of the focus of attention for the forward-access task, but not for variations where probing was in random order. This suggests that working memory capacity may depend on the type of search process engaged, and that certain working-memory-related cognitive processes are more amenable to practice than others.     

Top-down modulation: bridging selective attention and working memory

Selective attention, the ability to focus our cognitive resources on information relevant to our goals, influences working memory (WM) performance. Indeed, attention and working memory are increasingly viewed as overlapping constructs. Here, we review recent evidence from human neurophysiological studies demonstrating that top-down modulation serves as a common neural mechanism underlying these two cognitive operations. The core features include activity modulation in stimulus-selective sensory cortices with concurrent engagement of prefrontal and parietal control regions that function as sources of top-down signals. Notably, top-down modulation is engaged during both stimulus-present and stimulus-absent stages of WM tasks; that is, expectation of an ensuing stimulus to be remembered, selection and encoding of stimuli, maintenance of relevant information in mind and memory retrieval.     

Better working memory for non-social targets in infant siblings of children with Autism Spectrum Disorder

We compared working memory (WM) for the location of social versus non-social targets in infant siblings of children with Autism Spectrum Disorders (sibs-ASD, n  =   25) and of typically developing children (sibs-TD, n  =   30) at 6.5 and 9 months of age. There was a significant interaction of risk group and target type on WM, in which the sibs-ASD had better WM for non-social targets as compared with controls. There was no group by stimulus interaction on two non-memory measures. The results suggest that the increased competency of sibs-ASD in WM (creating, updating and using transient representations) for non-social stimuli distinguishes them from sibs-TD by 9 months of age. This early emerging strength is discussed as a developmental pathway that may have implications for social attention and learning in children at risk for ASD.     

Two dissociable updating processes in working memory

The authors show that the updating of working memory (WM) representations is carried out by the cooperative act of 2 dissociable reaction time (RT) components: a global updating process that provides stability by shielding WM contents against interference and a local process that provides flexibility. Participants kept track of 1-3 items (digits or Gibson figures). In each trial, the items either were similar to those in the previous trial or were different in any or all of the items. Experiments 1 and 2 established the existence of 2 independent RT components representing the 2 updating processes. Global updating cost was sensitive to total number of items in WM (set size), regardless of the number of items that actually were modified. Local updating cost was sensitive to the number of modified items, regardless of the set size. Experiment 3 showed that participants had to dismantle the representation formed by previous global updating in order to carry out new updating.     

Aging and working memory inside and outside the focus of attention: dissociations of availability and accessibility

Two experiments used the N-Back task to test for age differences in working memory inside and outside the focus of attention. Manipulations of the difficulty of item-context binding (Experiment 1) and of stimulus feature binding (Experiment 2) were used to create conditions that varied in their demand on working memory, with the expectation that greater demand might increase age differences in focus-switching costs and the search rate outside the focus of attention. Results showed, however, that although age differences were evident in measures of overall speed and accuracy, and the manipulations significantly affected response times and accuracy in the expected direction, the experimental manipulations had no impact on age differences. Findings instead pointed to age-related reductions in accuracy but not speed of focus-switching and search outside the focus of attention. Thus, age-related deficits appear to involve the availability of representations in working memory, but not their accessibility.     

The influence of working memory load on the Simon effect

The contribution of spatial and verbal working memory (WM) to the processing of stimulus location in the Simon task was examined in two experiments. Subjects performed the Simon task in single- and dual-task conditions. In the dual-task conditions, individuals first encoded the locations of four boxes/pseudocharacters or the identity of seven characters in memory, then responded to the color of a Simon stimulus, and finally received a test for the WM set. The Simon effect was insensitive to spatial WM load and was abolished by verbal WM load. These data reveal a role of verbal WM in the processing of spatial location in the Simon task. In addition, the data are inconsistent with dual-route models of the Simon effect and may support response discrimination accounts of the mechanisms underlying the processing of spatial location in the Simon task.     

Selective attention to elements in working memory

Three experiments with an arithmetic working memory task examine the object switch effect first reported by Garavan (1998), which was interpreted as evidence for a focus of attention within working memory. Experiments 1a and 1b showed object switch costs with a task that requires selective access to items in working memory, but did not involve counting, and did not require updating of working memory contents, thus ruling out two alternative explanations of Garavan's results. Experiment 2 showed object switch costs with a task that required no selective access to working memory contents, but involved updating, thus providing evidence for a second component to the overall object switch costs. Further analyses revealed that the object switch cost increased with memory set size; that there were (smaller) switch costs when the switch was to an item of the same type; that repeating an arithmetic operation does not have the same effect as repeating the object it is applied to; and that object switching is not mediated by backward inhibition of the previously focused object.     

Analogous mechanisms of selection and updating in declarative and procedural working memory: Experiments and a computational model

The article investigates the mechanisms of selecting and updating representations in declarative and procedural working memory (WM). Declarative WM holds the objects of thought available, whereas procedural WM holds representations of what to do with these objects. Both systems consist of three embedded components: activated long-term memory, a central capacity-limited component for building structures through temporary bindings, and a single-element focus of attention. Five experiments test the hypothesis of analogous mechanisms in declarative and procedural WM, investigating repetition effects across trials for individual representations (objects and responses) and for sets (memory sets and task sets), as well as set-congruency effects. Evidence for analogous processes was obtained from three phenomena: (1) Costs of task switching and of list switching are reduced with longer preparation interval. (2) The effects of task congruency and of list congruency are undiminished with longer preparation interval. (3) Response repetition interacts with task repetition in procedural WM; here we show an analogous interaction of list repetition with item repetition in declarative WM. All three patterns were reproduced by a connectionist model implementing the assumed selection and updating mechanisms. The model consists of two modules, an item-selection module selecting individual items from a memory set, or responses from a task set, and a set-selection module for selecting memory sets or task sets. The model codes the matrix of binding weights in the item-selection module as a pattern of activation in the set-selection module, thereby providing a mechanism for building chunks in LTM, and for unpacking them as structures into working memory.