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.     

How verbal memory loads consume attention

According to a traditional assumption about working memory, participants retain a series of verbal items for immediate recall using covert verbal rehearsal, without much need for attention. We reassessed this assumption by imposing a speeded, nonverbal choice reaction time (CRT) task following the presentation of each digit in a list to be recalled. When the memory load surpassed a few items, performance on the speeded CRT task became increasingly impaired. This CRT task impairment depended only on attention-related components of working memory; it was not alleviated by the presence of an auditory memory trace that automatically helped the recall of items at the ends of spoken lists. We suggest that attention-demanding refreshing of verbal stimuli occurs along with any covert rehearsal.     

Working memory selection and competition between target and distractor representations

A working memory selection task combining directed-forgetting and memory-scanning paradigms was used to test the hypothesis that difficulty in selecting target from distractor items in working memory depends on the competition between target and distractor representations, as is commonly assumed in perceptual selection research. Participants memorized two trigrams and were then cued to select one as the new memory set and forget the other. They later saw a test letter and made judgment as to whether the letter was in the selected trigram, i.e., the new memory set. Selection difficulty was examined by manipulating the type of selection cues and the time the cue could be utilized. While re-presenting the targets in the cue display facilitated selection, re-presenting distractors did not impede selection. The results suggest that working memory selection may depend more on the activation from representations of the target than of the distractor.     

Neural dynamics of attention switching and temporal-order information in short-term memory

Reeves and Sperling (1986) have developed an experimental paradigm and a model to explain how attention switching influences the storage of temporal-order information in short-term memory (STM), or working memory. The present article suggests that attention switching influences initial storage of items in STM, but that competitive interactions among the STM representations of stored items control the further evolution of temporal-order information as new items are processed. The laws governing these competitive interactions, called the long-term memory (LTM) invariance principle and the STM normalization rule, were originally derived from postulates that ensure that STM is updated in a way that enables temporally stable list learning in LTM to occur. Despite these adaptive constraints, and often because of them, temporal-order information is not always stored veridically. Both feedforward and feedback STM processes, with different invariant properties, are identified in the storage of temporal-order information.     

Behavioral and neurophysiological correlates of episodic coding,proactive interference,and list length effects in a running span verbal working memory task

Updating refers to (1) discarding items from, (2) repositioning items in, and (3) adding items to a running working memory span. Our behavioral and fMRI experiments varied three factors: trial length, proactive interference (PI), and group integrity. Group integrity reflected whether the grouping of items at the encoding stage was violated at discarding. Behavioral results were consistent with the idea that updating processes have a relatively short refractory period and may not fatigue, and they revealed that episodic information about group context is encoded automatically in working memory stimulus representations. The fMRI results did not show evidence that updating requirements in a task recruit executive control processes other than those supporting performance on nonupdating trials. They did reveal an item-accumulation effect, in which signal increased monotonically with the number of items presented during the trial, despite the insensitivity of behavioral measures to this factor. Behavioral and fMRI correlates of PI extended previous results and rejected an alternative explanation of PI effects in working memory.     

The number and quality of representations in working memory

Flexible-resource theories characterize working memory as a flexible resource that can store either a large number of low-quality representations or a small number of high-quality representations. In contrast, limited-item theories propose that the number of items that can be stored in working memory is strictly limited and cannot be increased by decreasing the quality of the representations. We tested these fundamentally different conceptualizations of working memory capacity by determining whether observers could trade quality for quantity in working memory when given incentives to do so. We found no evidence that observers could increase the number of representations by decreasing their quality in working memory, but observers could make such a trade-off at earlier processing stages. Our results show that the capacity limit of working memory is best characterized as a limit on the number of items that can be stored and not as a limit on a finely divisible resource that simultaneously determines the number and quality of the representations.     

SKOOL versus ZOOL: effects of orthographic and phonological long-term memory on nonword immediate serial recall

In a now-classic study Besner and Davelaar (1982) reported an advantage of pseudohomophone (PSH) over nonword recall in a visual immediate serial recall (ISR) task, which remained under articulatory suppression (AS), and interpreted the findings as indicating PSH items obtain support from stored phonological long-term memory (LTM) representations even when phonological rehearsal is disrupted. However, one key question relating to this PSH effect remains: could the results have been contaminated by a potential confound of orthographic familiarity (i.e., PSH items often look like the word they sound like)? As a result, the present study examined the impact of orthography on PSH ISR. Our findings indicate that PSH accuracy was consistently higher for items that had an orthographic similarity to the parent word, and this effect did not interact with concurrent task. We therefore argue that PSH items in ISR obtain independent support from both orthographic and phonological LTM representations. The present study demonstrates the critical impact of orthographic LTM representations on visual nonword ISR, and we suggest that this may be a fruitful avenue for further research.     

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.     

Working memory, long-term memory, and medial temporal lobe function

Early studies of memory-impaired patients with medial temporal lobe (MTL) damage led to the view that the hippocampus and related MTL structures are involved in the formation of long-term memory and that immediate memory and working memory are independent of these structures. This traditional idea has recently been revisited. Impaired performance in patients with MTL lesions on tasks with short retention intervals, or no retention interval, and neuroimaging findings with similar tasks have been interpreted to mean that the MTL is sometimes needed for working memory and possibly even for visual perception itself. We present a reappraisal of this interpretation. Our main conclusion is that, if the material to be learned exceeds working memory capacity, if the material is difficult to rehearse, or if attention is diverted, performance depends on long-term memory even when the retention interval is brief. This fundamental notion is better captured by the terms subspan memory and supraspan memory than by the terms short-term memory and long-term memory. We propose methods for determining when performance on short-delay tasks must depend on long-term (supraspan) memory and suggest that MTL lesions impair performance only when immediate memory and working memory are insufficient to support performance. In neuroimaging studies, MTL activity during encoding is influenced by the memory load and correlates positively with long-term retention of the material that was presented. The most parsimonious and consistent interpretation of all the data is that subspan memoranda are supported by immediate memory and working memory and are independent of the MTL.     

Declarative working memory: A bio-inspired cognitive architecture proposal

Memory is considered one of the most important functions since it allows us to code, store and retrieve knowledge. These qualities make it an indispensable function for a virtual creature. In general, memory can be classified based on the durability of the stored data in working memory and long-term memory. Working memory refers to the capacity to maintain temporarily a limited amount of information in mind, which can then be used to support various abilities, including learning, reasoning, planning and decision-making. Unlike short-term memory, working memory is not only a storage site, but it is also a framework of interacting processes that involve the temporary storage and manipulation of information in the service of performing complex cognitive activities. Declarative memory is a type of long-term memory related with the storage of facts and events. This research focuses on the development of a cognitive architecture for the type of working memory that maintains and manipulates declarative information. The construction of the model was grounded in theoretical evidence taken from cognitive sciences such as neuroscience and psychology, which gave us the components and their processes. The model was evaluated through a case study that covers the encoding, storing, and retrieval stages. Our hypothesis is that a virtual creature endowed with our working memory model will provide faster access to the information needed for the ongoing task. Therefore, it improves the planning and decision-making processes.     

Contribution of lexico-semantic processes to verbal short-term memory tasks: a PET activation study

Recent studies have demonstrated the intervention of long-term memory processes in verbal STM tasks and several cognitive models have been proposed to explain these effects. A PET study was performed in order to determine whether supplementary cerebral areas are involved when subjects have to execute short-term memory tasks for items having representations in long-term memory (in comparison to items without such representations: words vs non-words). Results indicate that verbal STM for words specifically involves the left middle temporal gyrus (BA 21) and temporo-parietal junction (BA 39). These areas can be associated with lexical and semantic processes. These results are in agreement with cognitive models that postulate the simultaneous influence of lexical and semantic long-term representations on verbal STM processes and/or a lexico-semantic buffer.     

Dissociation of short- and long-term face memory: evidence from long-term recency effects in temporal lobe epilepsy

We tested whether memory deficits in temporal lobe epilepsy (TLE) are better described by a single- or dual-store memory model. To this aim, we analyzed the influence of TLE and proactive interference (PI) on immediate and 24-h long-term recency effects during face recognition in 16 healthy participants and 18 right and 21 left non-surgical TLE patients. PI in healthy participants or TLE erased the long-term recency effect, but left the immediate recency effect unaffected. Although the immediate recency effect was still visible in right TLE patients, the number of detected recency items during immediate recognition was decreased in right TLE compared to left TLE. Right TLE was also related to decreased detection of pre-recency items during delayed recognition compared to left TLE, and decreased detection of pre-recency items during immediate recognition under PI. The results show that the temporal lobes are necessary for the long-term recency effect, but not for the immediate recency effect, and thus speak for a dissociation of short- and long-term memory for faces. Right TLE is related to more severe long-term memory deficits than left TLE and is also related to additional short-term memory deficits for faces.     

Searching while loaded: Visual working memory does not interfere with hybrid search efficiency but hybrid search uses working memory capacity

In “hybrid search” tasks, such as finding items on a grocery list, one must search the scene for targets while also searching the list in memory. How is the representation of a visual item compared with the representations of items in the memory set? Predominant theories would propose a role for visual working memory (VWM) either as the site of the comparison or as a conduit between visual and memory systems. In seven experiments, we loaded VWM in different ways and found little or no effect on hybrid search performance. However, the presence of a hybrid search task did reduce the measured capacity of VWM by a constant amount regardless of the size of the memory or visual sets. These data are broadly consistent with an account in which VWM must dedicate a fixed amount of its capacity to passing visual representations to long-term memory for comparison to the items in the memory set. The data cast doubt on models in which the search template resides in VWM or where memory set item representations are moved from LTM through VWM to earlier areas for comparison to visual items.     

Interactions between visual working memory representations

We investigated whether the representations of different objects are maintained independently in working memory or interact with each other. Observers were shown two sequentially presented orientations and required to reproduce each orientation after a delay. The sequential presentation minimized perceptual interactions so that we could isolate interactions between memory representations per se. We found that similar orientations were repelled from each other whereas dissimilar orientations were attracted to each other. In addition, when one of the items was given greater attentional priority by means of a cue, the representation of the high-priority item was not influenced very much by the orientation of the low-priority item, but the representation of the low-priority item was strongly influenced by the orientation of the high-priority item. This indicates that attention modulates the interactions between working memory representations. In addition, errors in the reported orientations of the two objects were positively correlated under some conditions, suggesting that representations of distinct objects may become grouped together in memory. Together, these results demonstrate that working-memory representations are not independent but instead interact with each other in a manner that depends on attentional priority.     

Adaptive memory: enhanced source memory for animate entities

ABSTRACT

The animacy effect refers to enhanced memory for animate over inanimate items. In two studies, we examined whether this memory advantage generalises to source memory. A multinomial processing tree model was used to disentangle item recognition, source memory, and guessing processes. In Study 1, animate and inanimate words were presented at different spatial locations on the screen. Animacy was associated with enhanced source memory for the spatial locations of the items. In Study 2, pseudowords were associated with animate and inanimate properties. Replicating previous results, the pseudowords were better remembered when they were associated with animate properties than when they were associated with inanimate properties. What is more, participants had enhanced source memory for the association between the pseudowords and the animate properties. The results strengthen the idea that animate items are associated with richer mnemonic representations than inanimate items.     

Ageing and feature binding in visual working memory: The role of presentation time

A large body of research has clearly demonstrated that healthy ageing is accompanied by an associative memory deficit. Older adults exhibit disproportionately poor performance on memory tasks requiring the retention of associations between items (e.g., pairs of unrelated words). In contrast to this robust deficit, older adults’ ability to form and temporarily hold bound representations of an object's surface features, such as colour and shape, appears to be relatively well preserved. However, the findings of one set of experiments suggest that older adults may struggle to form temporary bound representations in visual working memory when given more time to study objects. However, these findings were based on between-participant comparisons across experimental paradigms. The present study directly assesses the role of presentation time in the ability of younger and older adults to bind shape and colour in visual working memory using a within-participant design. We report new evidence that giving older adults longer to study memory objects does not differentially affect their immediate memory for feature combinations relative to individual features. This is in line with a growing body of research suggesting that there is no age-related impairment in immediate memory for colour-shape binding.     

The effects of golf expertise and presentation modality on memory for golf and everyday items

The present study assessed whether golf expertise, presentation modality, and domain relevance affected memory for golf-related and everyday items. Forty-eight experienced golfers and 48 non-golfers were compared in their memory for golf-related ("putt to the hole") and everyday ("turn on the lamp") items. To-be-remembered items were presented verbally, visually, or were enacted. Enacted information was recalled best, followed by visually presented information. Combined effects of modality and golf expertise on recall of golf items were demonstrated on immediate but not on delayed recall. The findings suggest that recall of domain-relevant information is optimal when one has relevant background knowledge, and under conditions of visual encoding. The data support research on facilitation of domain-relevant knowledge on recall [Weber, N., & Brewer, N. (2003). Expert memory: The interaction of stimulus structure, attention, and expertise. Applied Cognitive Psychology, 17, 295-308]. Interpersonal body representation may have played a role in recall processes among experts [Thomas, R., Press, C., & Haggard, P. (2006). Shared representations in body perception. Acta Psychologica, 121, 317-330].     

Working memory load differentially affects tip-of-the-tongue states and feeling-of-knowing judgments

Tip-of-the-tongue states (TOTs) are judgments of the likelihood of imminent retrieval for items currently not recalled, whereas feeling-of-knowing judgments (FOKs) are predictions of successful recognition for items not recalled. The assumption has been that similar metacognitive processes dictate these similar judgments. In Experiment 1, TOTs and FOKs were compared for general information questions. Participants remembered four digits (working memory load) during target retrieval for half of the questions, and there was no memory load for the other questions. Working memory did not affect recall but decreased the number of TOTs and increased FOKs. In Experiment 2, participants maintained six digits during retrieval. TOTs decreased in the working memory condition, but FOKs remained constant. Experiment 3 replicated the results of Experiment 2 while asking for FOKs for recall. In each of the first three experiments, positive metacognitive judgments also affected working memory performance, supporting the idea that working memory and metamemory use similar monitoring processes. In Experiment 4, visual working memory did not affect TOTs or FOKs. The data support a view that TOTs and FOKs are separable metacognitive entities.     

The effects of word co-occurrence on short-term memory: associative links in long-term memory affect short-term memory performance

In immediate serial recall tasks, high-frequency words are recalled better than low-frequency words. This has been attributed to high-frequency words' being better represented and providing more effective support to a redintegration process at retrieval (C. Hulme et al., 1997). In studies of free recall, there is evidence that frequency of word co-occurrence, rather than word frequency per se, may explain the recall advantage enjoyed by high-frequency words (J. Deese, 1960). The authors present evidence that preexposing pairs of low-frequency words, so as to create associative links between them, has substantial beneficial effects on immediate serial recall performance. These benefits, which are not attributable to simple familiarization with the words per se, do not occur for high-frequency words. These findings indicate that associative links between items in long-term memory have important effects on short-term memory performance and suggest that the effects of word frequency in short-term memory tasks are related to differences in interitem associations in long-term memory.     

Cognitive control of familiarity: Directed forgetting reduces proactive interference in working memory

Proactive interference (PI) occurs when previously learned information interferes with new learning. In a working memory task, PI induces longer response times and more errors to recent negative probes than to new probes, presumably because the recent probe’s familiarity invites a “yes” response. Warnings, longer intertrial intervals, and the increased contextual salience of the probes can reduce but not eliminate PI, suggesting that cognitive control over PI is limited. Here we tested whether control exerted in the form of intentional forgetting performed during working memory can reduce the magnitude of PI. In two experiments, participants performed a working memory task with directed-forgetting instructions and the occasional presentation of recent probes. Surprise long-term memory testing indicated better memory for to-be-remembered than for to-be-forgotten items, documenting the classic directed-forgetting effect. Critically, in working memory, PI was virtually eliminated for recent probes from prior to-be-forgotten lists, as compared to recent probes from prior to-be-remembered lists. Thus cognitive control, when executed via directed forgetting, can reduce the adverse and otherwise persistent interference from familiarity, an effect that we attribute to attenuated memory representations of the to-be-forgotten items.