Updating working memory in aircraft noise and speech noise causes different fMRI activations

The present study used fMRI/BOLD neuroimaging to investigate how visual‐verbal working memory is updated when exposed to three different background‐noise conditions: speech noise, aircraft noise and silence. The number‐updating task that was used can distinguish between “substitution processes,” which involve adding new items to the working memory representation and suppressing old items, and “exclusion processes,” which involve rejecting new items and maintaining an intact memory set. The current findings supported the findings of a previous study by showing that substitution activated the dorsolateral prefrontal cortex, the posterior medial frontal cortex and the parietal lobes, whereas exclusion activated the anterior medial frontal cortex. Moreover, the prefrontal cortex was activated more by substitution processes when exposed to background speech than when exposed to aircraft noise. These results indicate that (a) the prefrontal cortex plays a special role when task‐irrelevant materials should be denied access to working memory and (b) that, when compensating for different types of noise, either different cognitive mechanisms are involved or those cognitive mechanisms that are involved are involved to different degrees.     

Interactions between frontal cortex and basal ganglia in working memory: a computational model

The frontal cortex and the basal ganglia interact via a relatively well understood and elaborate system of interconnections. In the context of motor function, these interconnections can be understood as disinhibiting, or “releasing the brakes,” on frontal motor action plans: The basal ganglia detect appropriate contexts for performing motor actions and enable the frontal cortex to execute such actions at the appropriate time. We build on this idea in the domain of working memory through the use of computational neural network models of this circuit. In our model, the frontal cortex exhibits robust active maintenance, whereas the basal ganglia contribute a selective, dynamic gating function that enables frontal memory representations to be rapidly updated in a task-relevant manner. We apply the model to a novel version of the continuous performance task that requires subroutine-like selective working memory updating and compare and contrast our model with other existing models and theories of frontal-cortex-basal-ganglia interactions.     

Do young and older adults rely on different processes in source memory tasks? A neuropsychological study

Source memory has consistently been associated with prefrontal function in both normal and clinical populations. Nevertheless, the exact contribution of this brain region to source memory remains uncertain, and evidence suggests that processes used by young and older adults may differ. The authors explored the extent to which scores on composite measures of neuropsychological tests of frontal and medial temporal function differentially predicted the performance of young and older adults on source memory tasks. Results indicated that a frontal composite measure, consistently associated with source memory performance in older adults, was unrelated to source memory in young adults, although it was sensitive to a demanding working memory task. The memory composite score, however, predicted performance in the young group. In addition, item and source memory were correlated in young but not older people. Findings are discussed in terms of age-related differences in working memory and executive functions, and differential binding processes necessary for item and source memory. The requirement to integrate item and source information at encoding appears to place greater demands on executive or working memory processes in older adults than in younger adults.     

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.     

Updating of context in working memory: An event-related potential study

Flexible control of behavior depends on the representation, maintenance, and updating of context information in working memory, which is thought to rely on the prefrontal cortex (PFC). However, in contrast to maintenance, the dynamics of context activation and updating have not been well studied. To identify neural signals associated with context updating, we compared event-related potentials associated with cues that did or did not provide task-relevant context information. The earliest effect of context was detected 200 msec following cue onset and had a scalp topography consistent with a generator in the PFC. Subsequent effects of context were detected at 400-700 msec following cue onset (P3b), with a broad scalp distribution spanning posterior areas, and during the final 300 msec preceding the target, with a probable generator in the medial frontal cortex. We propose that the effect of context on P2 is consistent with the onset of context updating in the PFC. Subsequent components may be indicative of activation of task-relevant posterior regions and context maintenance.     

Which factors influence number updating in working memory? The effects of size distance and suppression

Updating information in working memory is a critical process which makes possible to have available, at every moment, the information most relevant for mind operations. However, the specific mechanisms underlying the updating process have rarely been analysed. This paper examines the importance of two of the mechanisms implicated in a numerical updating task: item comparison and item substitution. The item comparison mechanism was studied by manipulating the size distance between items. The item substitution mechanism was investigated by increasing/decreasing the number of updates within trials. Furthermore, in order to examine the effects of time constraints, presentation rate was manipulated. Over three experiments, the results obtained highlighted that updating performance is mainly influenced by suppression request, even when the presentation rate is self‐paced. However, errors depend on the distance between items. The implications of the results for the understanding of updating are discussed.     

Encoding location and serial order in auditory working memory: evidence for separable processes

In this study, we investigated the interactions between temporal and spatial information in auditory working memory. In two experiments, participants were presented with sequences of sounds originating from different locations in space and were then asked to recall either their position or their serial order. In Experiment 1, attention during encoding was manipulated by contrasting 'pure' blocks (i.e., location-only or serial-order-only trials) to 'mixed' blocks (i.e., different percentages of spatial and serial-order trials). In Experiment 2, 'pure' blocks were contrasted to blocks in which spatial and serial-order trials were intermixed with a third task requiring a semantic categorization of sounds. Results from both experiments showed that, whereas serial-order recall is linearly affected by the simultaneous encoding of a concurrent feature, the recall of position is mostly unaffected by concurrent feature encoding. Contrastingly, overall performance level was lower for spatial recall than serial recall. We concluded that serial order and location of items appear to be independently encoded in auditory working memory. Serial order is easier to recall, but strongly affected by the processing of concurrent item dimensions, while item location is more difficult to recall, but relatively automatic, as shown by its strong resistance to interfering dimensions in encoding.     

Evidence for neocortical involvement in reference memory

Rats were trained on an eight-arm radial maze task using a procedure that provides for an assessment of both working and reference memory. Following training, rats received parietal cortex, medial prefrontal cortex, visual cortex, or nucleus basalis magnocellularis lesions. Rats with visual cortex lesions showed no change in performance on either working or reference memory. Rats with parietal cortex lesions displayed a temporary deficit in reference, but no deficit on working memory. Animals with medial prefrontal cortex lesions showed a temporary deficit on both working and reference memory. Rats with extensive lateral frontal and parietal cortex depletion of acetylcholinesterase following nucleus basalis magnocellularis lesions had a marked disruption only of reference but not of working memory. It is concluded that neocortex and possibly the cholinergic projections to neocortex play an important role in mediating reference memory.     

Cognitive neuroscience of episodic memory encoding

This paper presents a cognitive neuroscientific perspective on how human episodic memories are formed. Convergent evidence from multiple brain imaging studies using positron emission tomography (PET) and functional magnetic resonance imaging (fMRI) suggests a role for frontal cortex in episodic memory encoding. Activity levels within frontal cortex can predict episodic memory encoding across a wide range of behavioral manipulations known to influence memory performance, such as those present during levels of processing and divided attention manipulations. Activity levels within specific frontal and medial temporal regions can even predict, on an item by item basis, whether an episodic memory is likely to form. Furthermore, separate frontal regions appear to participate in supplying code-specific information, including distinct regions which process semantic attributes of verbal information as well as right-lateralized regions which process nonverbal information. We hypothesize that activity within these multiple frontal regions provides a functional influence (input) to medical temporal regions that bind the information together into a lasting episodic memory trace.     

Large-scale neural network for sentence processing

Our model of sentence comprehension includes at least grammatical processes important for structure-building, and executive resources such as working memory that support these grammatical processes. We hypothesized that a core network of brain regions supports grammatical processes, and that additional brain regions are activated depending on the working memory demands associated with processing a particular grammatical feature. We used functional magnetic resonance imaging (fMRI) to test this hypothesis by comparing cortical activation patterns during coherence judgments of sentences with three different syntactic features. We found activation of the ventral portion of left inferior frontal cortex during judgments of violations of each grammatical feature. Increased recruitment of the dorsal portion of left inferior frontal cortex was seen during judgments of violations of specific grammatical features that appear to involve a more prominent working memory component. Left posterolateral temporal cortex and anterior cingulate were also implicated in judging some of the grammatical features. Our observations are consistent with a large-scale neural network for sentence processing that includes a core set of regions for detecting and repairing several different kinds of grammatical features, and additional regions that appear to participate depending on the working memory demands associated with processing a particular grammatical feature.     

Strategy-dependent changes in memory: Effects on behavior and brain activity

In the present study, an implicit strategy manipulation was used to explore the contribution of memory strategy to brain activation and behavioral performance. Participants were biased to use either a short-term (maintenance-focused) or long-term (retrieval-focused) memory strategy within a single memory task through manipulation of task context. In comparing directly matched trials across the different task contexts, we observed clear changes in both behavioral performance and brain activity across a network of regions located primarily within lateral and medial frontal cortex. These effects of the memory strategy manipulation suggest that when a retrieval-focused strategy is induced, mnemonic processes are preferentially engaged during the encoding period. In contrast, when a maintenance-focused strategy is induced, mnemonic processes are preferentially engaged during the delay and response periods. Taken together, the results imply that covert cognitive strategies play an important role in modulating brain activation and behavior during memory tasks.     

An fMRI study of phonological and spatial working memory using identical stimuli

The aim of the present fMRI study was to localize brain areas that were uniquely activated for phonological versus spatial working memory. Previous studies have reported inconsistent results, most likely because of methodological heterogeneity varying both stimuli and instructions in the same study. Here, identical consonant-vowel-consonant non-words were visually presented to the subjects in a 2-back paradigm under two different instructions; the subjects either had to memorize the non-words per se or their location. The results give evidence for a hemispheric organization of working memory, with dominance for processing of phonological information in the left hemisphere and frontal cortex, and spatial information in the right hemisphere and parietal cortex. The results also reflect a certain overlap between the neuronal network for working memory and processing of verbal and spatial material. These findings are discussed with regard to processing specificity and the extent that activated areas also may reflect perceptual processes.     

Effects of frontal lobe damage on interference effects in working memory

Working memory is hypothesized to comprise a collection of distinct components or processes, each of which may have a unique neural substrate. Recent neuroimaging studies have isolated a region of the left inferior frontal gyrus that appears to be related specifically to one such component: resolving interference from previous items in working memory. In the present study, we examined working memory in patients with unilateral frontal lobe lesions by using a modified version of an item recognition task in which interference from previous trials was manipulated. In particular, we focused on patient R.C., whose lesion uniquely impinged on the region identified in the neuroimaging studies of interference effects. We measured baseline working memory performance and interference effects in R.C. and other frontal patients and in age-matched control subjects and young control subjects. Comparisons of each of these groups supported the following conclusions. Normal aging is associated with changes to both working memory and interference effects. Patients with frontal damage exhibited further declines in working memory but normal interference effects, with the exception of R.C., who exhibited a pronounced interference effect on both response time and accuracy. We propose that the left inferior frontal gyrus subserves a general, nonmnemonic function of selecting relevant information in the face of competing alternatives and that this function may be required by some working memory tasks.     

Encoding processes and sex-role preferences

Seven- and 10-year-old children were tested on memory and sex-role preference tasks. The memory task was the Wickens release from proactive inhibition paradigm in which short-term recall of words is tested on successive trials. On Trials 1–4, words were selected from one of two categories, either words with masculine or feminine connotations. On Trial 5, words were drawn from the second category. Sex-role preference was assessed by asking the child to select his favorite pictures from an array that included masculine and feminine items. Recall by boys at both ages increased following a shift between words with masculine or feminine connotations, suggesting that this dimension of a word's meaning was encoded in memory. Recall by girls who selected a feminine item as their favorite on the sex-role preference task increased following a category shift; recall by girls who chose a masculine item did not increase. These results are discussed in relation to previous research on the attributes of encoding in children's memory.     

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.     

Updating information in verbal working memory and executive functioning

50 older adults (M age=77.9 yr., SD=7.3; 35 women and 15 men) were tested using the updating working-memory task. They were also given the neuropsychological Wisconsin Card Sorting Test, assumed to evaluate executive functioning and the frontal cortex. A factor analysis with age, education, and verbal ability partialled out was computed on the updating task outcomes and resulted in a two-factor solution, indicating that this task requires two independent processes, interpreted as reflecting a storage component and an updating component. Partial correlations with age, education, and verbal ability partialled out indicated that Wisconsin Card Sorting Test measures were significantly associated with the factor supposed to reflect the updating process. Such results appeared consistent with the model of working memory with a central executive system involved in the updating process and related to the executive-frontal functioning, and a phonological loop system involved in the storage of verbal information and not linked to executive-frontal functions.     

The focus of attention in working memory: Evidence from a word updating task

Three experiments examined the cognitive costs of item switching within working memory with a novel word updating task, thereby extending previous research to the field of linguistic stimuli and linguistic-graphemic updating operations. In Experiments 1 and 2 costs for switching between working memory items were evident on the word level, and they increased with an increasing word set size (Experiment 2). In contrast, a surprisingly similar switch effect on the level of letters was not affected by word set size (Experiment 2). Experiment 3 showed that this effect is not simply based on the need for re-orienting visual spatial attention. To account for the overall picture of results, a recursive model of attentional foci is proposed. Moreover, individual working memory span appears to be associated with the accuracy of item switching, but not with its speed.     

The focus of attention in working memory: evidence from a word updating task

Three experiments examined the cognitive costs of item switching within working memory with a novel word updating task, thereby extending previous research to the field of linguistic stimuli and linguistic-graphemic updating operations. In Experiments 1 and 2 costs for switching between working memory items were evident on the word level, and they increased with an increasing word set size (Experiment 2). In contrast, a surprisingly similar switch effect on the level of letters was not affected by word set size (Experiment 2). Experiment 3 showed that this effect is not simply based on the need for re-orienting visual spatial attention. To account for the overall picture of results, a recursive model of attentional foci is proposed. Moreover, individual working memory span appears to be associated with the accuracy of item switching, but not with its speed.     

Content-context binding in verbal working memory updating: on-line and off-line effects

The role of content–context binding in working memory updating has received only marginal interest, despite its undoubted relevance for the updating process. In the classical updating paradigm, the main focus has been on the process of discarding information when new information is presented. Its efficacy has been determined by measurement of the accuracy of recalled, updated information. In the current study we measured the working memory updating process directly, employing a dynamic memory task composed of alternating sequences of learning, active maintenance and updating phases. Binding was manipulated by changing the type of updating (total or partial) and the strength of the perceptual connection between items. The on-line updating process and the off-line efficacy were separately analysed. Both on-line and off-line measures indicated that partial updating conditions, where the operation of updating content–context binding was required, were more demanding than both total updating conditions and memory and maintenance phases. Our results suggest that working memory updating can be identified not only as a process of substitution of information, but also as inhibition of no longer relevant information and, above all, as a binding updating.     

Segmentation in the perception and memory of events

People make sense of continuous streams of observed behavior in part by segmenting them into events. Event segmentation seems to be an ongoing component of everyday perception. Events are segmented simultaneously at multiple timescales, and are grouped hierarchically. Activity in brain regions including the posterior temporal and parietal cortex and lateral frontal cortex increases transiently at event boundaries. The parsing of ongoing activity into events is related to the updating of working memory, to the contents of long-term memory, and to the learning of new procedures. Event segmentation might arise as a side effect of an adaptive mechanism that integrates information over the recent past to improve predictions about the near future.