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Right parietal contributions to verbal working memory: spatial or executive?

The left inferior parietal cortex has been claimed to be the site of the verbal short-term store, yet imaging studies report activity of a homologous right-hemisphere region in verbal working memory tasks as well. In spite of its prevalent activity, right parietal contributions to verbal working memory are poorly understood. To clarify its role in verbal working memory performance, we tested a patient with a lesion in the right parietal lobe on verbal and spatial versions of the N-back task. The patient was impaired in all the spatial conditions regardless of load (0-, 1-, and 2-back), whereas in the verbal N-back he was impaired only in the conditions with a memory demand (1- and 2-back). Given that we had presented stimuli at multiple locations in the verbal N-back, however, it remained possible that the lesion impaired spatial representation rather than verbal working memory per se. With central stimulus presentation, his performance dramatically improved indicating that his difficulty with the N-back task was largely due to his poor visuospatial abilities.

Aged↗

Case studies in working memory: a case for single cases?

Patterns of cognitive deficit in single neuropsychological cases are common sources of evidence for theories of normal cognition. In particular, the working memory model has benefited from data obtained from a number of contrasting patients, in some cases resulting in modifications of the working memory model. In this paper, patterns of data from short-term memory patients and anarthric patients are compared with patterns of data from normal subjects. The patterns of patient data that were unlike those patterns typically found for groups of normal subjects, could be incorporated within a modified version of the articulatory loop component of the working memory model. However a small number of individual normal subjects also did not show the pattern that is reported on the basis of average performance of groups of normal subjects. This causes some difficulty in interpreting those data from such 'aberrant normal' patterns, and those data from single patients with functional cognitive deficits. The implications of these findings for the interpretation of neuropsychological data are discussed in the context of the working memory model, but with the intention of making a general point pertaining to the development of functional models of cognition. It is argued that single case studies should continue to provide a useful source of evidence, providing that care is exercised in considering the implications of such data for models of normal cognition.

Adult↗

The role of the dorsal raphe-serotonergic system and cholinergic receptors in the modulation of working memory.

The present study investigated the role of the dorsal raphe-serotonergic system and its interaction with muscarinic or nicotinic receptors in the modulation of working memory and motor activity by assessing the effects of serotonin lesion with pCA and cholinergic receptor blockade on the performance of rats in a working memory (delayed non-matching to position, DNMTP) task. The pCA lesion did not impair the choice accuracy or motor activity of rats in the DNMTP-task. The lower dose of scopolamine (0.075 mg/kg) impaired percent correct responses already at the shortest delay which is not indicative of a working memory impairment per se. Scopolamine also disrupted motor activity markedly. The effects of scopolamine 0.075 mg/kg on the choice accuracy were aggravated by pCA treatment. Furthermore, the effects of N-methylscopolamine (0.150 mg/kg) were comparable with scopolamine. The higher dose of mecamylamine (3.0 mg/kg) also interfered with motor activity and it decreased the choice accuracy. The performance disruption induced by mecamylamine was not as severe as that seen with scopolamine. Mecamylamine did not reveal any interaction with the serotonergic lesion. Hexamethonium slightly decreased the percent correct responses, while not interfering with motor activity of rats. The present results suggest that: (i) lesion of serotonergic fibers with pCA does not significantly impair the choice accuracy or interfere with motor activity of rats; (ii) the blockade of cholinergic receptors does not impair working memory per se, but disrupts motor activity, and (iii) pCA lesion of serotonergic fibers aggravates the non-mnemonic choice accuracy impairment induced by central muscarinic blockade, while not interacting with the cholinolytics in modulation of motor activity.

Animals↗

Effect of working memory on evaluation-related frontocentral negativity.

Recent results suggest that a negative ERP potential emerges not only on error trials, but is also found after correct responses. There is broad evidence that this component is generated in the anterior cingulate cortex (ACC). The present study aimed to explore the influence of working memory demands on the evaluation process probably reflected by the negative potential. To this purpose, a modified continuous performance task (CPT) was used, with variations of the delay between cues and imperative stimuli. Data were analyzed using conventional averaging techniques as well as source localization with LORETA (low resolution brain electromagnetic tomography). Results suggest a significant effect of working memory delay on the amplitude of the post-response negative ERP component. Its source was located in the anterior cingulate cortex, with the exact location being dependent upon working memory demands. The results support the notion of a general response evaluation system reflected by a post-response negative component. The findings of a working memory dependent modification of this potential suggest a functional link between the medial frontal cortex and the lateral frontal regions primarily involved in working memory processing.

Adult↗

A new one-trial test for neurobiological studies of memory in rats. III. Spatial vs. non-spatial working memory.

Rats were submitted to object and spatial recognition tests (both based on the same paradigm) and to the radial-arm maze. The results are as follows: (1) rats could discriminate between a new and a familiar object when the retention delay was 1 min, 15 min or 60 min but not 24 h. The relationship between the level of discrimination and intertrial delays is quadratic with a maximum for 15 min. (2) Exposure to distractive stimuli during the retention delay may impair object recognition. (3) Rats discriminated between a new and a familiar space. (4) There is no correlation between the three tests which argues for a multiple form of working memory, especially a spatial and a non-spatial one. (5) Medial septal lesion did not impair object and spatial recognition memory, but the level of discrimination in the spatial recognition test was significantly reduced compared to that of control.

Aging↗

Working memory, metacognitive uncertainty, and belief bias in syllogistic reasoning.

Studies of syllogistic reasoning have shown that the size of the belief bias effect varies with manipulations of logical validity and problem form. This paper presents a mental models-based account, which explains these findings in terms of variations in the working-memory demands of different problem types. We propose that belief bias may reflect the use of a heuristic that is applied when a threshold of uncertainty in one's processing--attributable to working-memory overload--is exceeded during reasoning. Three experiments are reported, which tested predictions deriving from this account. In Experiment 1, conclusions of neutral believability were presented for evaluation, and a predicted dissociation was observed in confidence ratings for responses to valid and invalid arguments, with participants being more confident in the former. In Experiment 2, an attempt to manipulate working-memory loads indirectly by varying syllogistic figure failed to produce predicted effects upon the size of the belief bias effect. It is argued that the employment of a conclusion evaluation methodology minimized the effect of the figural manipulation in this experiment. In Experiment 3, participants' articulatory and spatial recall capacities were calibrated as a direct test of working-memory involvement in belief bias. Predicted differences in the pattern of belief bias observed between high and low spatial recall groups supported the view that limited working memory plays a key role in belief bias.

Concept Formation↗

Neural mechanisms of spatial working memory: contributions of the dorsolateral prefrontal cortex and the thalamic mediodorsal nucleus.

The dorsolateral prefrontal cortex (DLPFC) has been known to play an important role in working memory. Neurophysiological studies have revealed that delay period activity observed in the DLPFC is a neural correlate of the temporary storage mechanism for information and that this activity represents either retrospective or prospective information, although the majority represents retrospective information. However, the DLPFC is not the only brain area related to working memory. The analysis of neural activity in the thalamic mediodorsal (MD) nucleus reveals that the MD also participates in working memory. Although similar task-related activities were observed in the MD, the directional bias of these activities and the proportion of presaccadic activity are different between the MD and the DLPFC. These results indicate that, although the MD participates in working memory, the way it participates in this process is different between these two areas, in that the MD participates more in motor control aspects than the DLPFC does.

Animals↗

Functional neuroanatomical double dissociation of mnemonic and executive control processes contributing to working memory performance.

We used event-related functional MRI to investigate the neural bases of two categories of mental processes believed to contribute to performance of an alphabetization working memory task: memory storage and memory manipulation. Our delayed-response tasks required memory for the identity and position-in-the-display of items in two- or five-letter memory sets (to identify load-sensitive regions) or memory for the identity and relative position-in-the-alphabet of items in five-letter memory sets (to identify manipulation-sensitive regions). Results revealed voxels in the left perisylvian cortex of five of five subjects showing load sensitivity (as contrasted with alphabetization-sensitive voxels in this region in only one subject) and voxels of dorsolateral prefrontal cortex in all subjects showing alphabetization sensitivity (as contrasted with load-sensitive voxels in this region in two subjects). This double dissociation was reliable at the group level. These data are consistent with the hypothesis that the nonmnemonic executive control processes that can contribute to working memory function are primarily prefrontal cortex-mediated whereas mnemonic processes necessary for working memory storage are primarily posteriorly mediated. More broadly, they support the view that working memory is a faculty that arises from the coordinated interaction of computationally and neuroanatomically dissociable processes.

Cerebral Cortex↗

Response shifting and inhibition, but not working memory, are impaired after long-term heavy alcohol consumption.

Chronic alcohol abuse leads to cognitive deficits. The authors investigated whether a systematic increase of interference in a 2-back working memory paradigm would lead to cognitive deficits in alcoholic participants and compared their performance in such a task with that in an alternate-response task. Twenty-four nonamnesic and nondemented alcohol abuse (AA) patients and 12 patients with Korsakoff's syndrome (KS) were compared with a control group. AA patients were impaired in the alternate-response task but not in working memory interference resolution. KS patients performed worse than the AA patients and the controls in both tasks. The neurotoxic side effects of alcohol therefore lead to a specific deficit in alternating between response rules but not in working memory, independently of whether the working memory task involves interference resolution or not.

Adult↗

Does the capacity of working memory change with age?

Young and elderly adults were compared for recall performance on simple digit and word spans (traditional tests of primary memory), versus a "loaded" auditory word span test designed to emphasize working memory capacity. Although digit spans were identical for the two age groups, there were small but significant age differences in word span, and even larger differences in working memory performance. An analysis of correlations between span measures and verbal ability scores supported the position that working memory loading represents a special problem for the elderly.

Adolescent↗

The roles of sequencing and verbal working memory in sentence comprehension deficits in Parkinson's disease.

Studies of sentence comprehension deficits in Parkinson's disease (PD) patients suggest that language processing involves circuits connecting subcortical and cortical regions. Anatomically segregated neural circuits appear to support different cognitive and motor functions. To investigate which functions are implicated in PD comprehension deficits, we tested comprehension, verbal working memory span, and cognitive set-switching in a non-linguistic task in 41 PD patients; we also obtained speech measurements reflecting motor sequencing processes that may be involved in articulatory rehearsal within working memory. Comprehension of sentences with center-embedded or final relative clauses was impaired when they could not be understood from lexical semantic content alone. Overall comprehension error rates correlated strongly with impaired set-switching and significantly with reduced working memory span and speech motor sequencing deficits. Correlations with comprehension of different sentence structures indicate that these impairments do not represent a single deficit; rather, PD comprehension deficits appear to arise from several independent mechanisms. Deficits in cognitive set-switching or underlying inhibitory processes may compromise the ability to process relative clauses. Deficits in verbal working memory appear to impair comprehension of long-distance dependencies. Speech sequencing correlated with neither set-switching nor verbal working memory span, consistent with their being supported by independent, segregated cortico-subcortical circuits.

Aged↗

"What"-Then-Where" in visual working memory: an event-related fMRI study.

Behavioral studies indicate that spatial and object working memory are computed by dissociable subsystems. We investigated the neural bases of this dissociation with a whole-brain fMRI design and analysis technique that permitted direct assessment of delay-period activity, uncontaminated by other components of the trial. The task employed a "what"-then-"where" design, with an object and a spatial delay period incorporated in each trial; within-trial order of delay conditions was balanced across each scan. Our experiment failed to find evidence, at the single-subject level and at the group level, for anatomical segregation of spatial and object working memory function in the frontal cortex. Delay-period activity in the caudate nucleus revealed a sensitivity to position in the trial in the spatial, but not the object, condition. In posterior regions, spatial delay-period activity was associated with preferential recruitment of extrastriate areas falling within Brodmann's area 19 and, less reliably, the superior parietal lobule. Object-specific delay-period activity was found predominantly in ventral regions of the posterior cortex and demonstrated more topographic variability across subjects than did spatial working memory activity.

Adult↗

Effects of domain knowledge, working memory capacity, and age on cognitive performance: an investigation of the knowledge-is-power hypothesis.

Domain knowledge facilitates performance in many cognitive tasks. However, very little is known about the interplay between domain knowledge and factors that are believed to reflect general, and relatively stable, characteristics of the individual. The primary goal of this study was to investigate the interplay between domain knowledge and one such factor: working memory capacity. Adults from wide ranges of working memory capacity, age, and knowledge about the game of baseball listened to, and then answered questions about, simulated radio broadcasts of baseball games. There was a strong facilitative effect of preexisting knowledge of baseball on memory performance, particularly for information judged to be directly relevant to the baseball games. However, there was a positive effect of working memory capacity on memory performance as well, and there was no indication that domain knowledge attenuated this effect. That is, working memory capacity contributed to memory performance even at high levels of domain knowledge. Similarly, there was no evidence that domain knowledge attenuated age-related differences (favoring young adults) in memory performance. We discuss implications of the results for understanding proficiency in cognitive domains from an individual-differences perspective.

Adolescent↗

High-resolution EEG mapping of cortical activation related to working memory: effects of task difficulty, type of processing, and practice.

Changes in cortical activity during working memory tasks were examined with electroencephalograms (EEGs) sampled from 115 channels and spatially sharpened with magnetic resonance imaging (MRI)-based finite element deblurring. Eight subjects performed tasks requiring comparison of each stimulus to a preceding one on verbal or spatial attributes. A frontal midline theta rhythm increased in magnitude with increased memory load. Dipole models localized this signal to the region of the anterior cingulate cortex. A slow (low-frequency), parietocentral, alpha signal decreased with increased working memory load. These signals were insensitive to the type of stimulus attribute being processed. A faster (higher-frequency), occipitoparietal, alpha signal was relatively attenuated in the spatial version of the task, especially over the posterior right hemisphere. Theta and alpha signals increased, and overt performance improved, after practice on the tasks. Increases in theta with both increased task difficulty and with practice suggests that focusing attention required more effort after an extended test session. Decreased alpha in the difficult tasks indicates that this signal is inversely related to the amount of cortical resources allocated to task performance. Practice-related increases in alpha suggest that fewer cortical resources are required after skill development. These results serve: (i) to dissociate the effects of task difficulty and practice; (ii) to differentiate the involvement of posterior cortex in spatial versus verbal tasks; (iii) to localize frontal midline theta to the anteromedial cortex; and (iv) to demonstrate the feasibility of using anatomical MRIs to remove the blurring effect of the skull and scalp from the ongoing EEG. The results are discussed with respect to those obtained in a prior study of transient evoked potentials during working memory.

Adult↗

Backward inhibition in a task of switching attention within verbal working memory.

Three experiments were conducted to examine the backward inhibition effect in attention switching within verbal working memory. Experiment one showed significant backward inhibition effect in a "tri-count task". Experiment two suggested that the effect was not due to a perceptual inhibition on the previously presented figure. Experiment three excluded the sequential expectancy explanation for this inhibition effect. Our results suggest that attention switching between working memory items is accompanied by inhibition of the previously attended working memory item. The findings are discussed in respect to the account of the executive function.

Adolescent↗

A biophysical model of multiple-item working memory: a computational and neuroimaging study.

Biophysically based computational models have successfully accounted for the persistent neural activity underlying the maintenance of single items of information in working memory. The aim of the present study was to extend previous models in order to retain multiple items, in agreement with the observed human storage capacity. This was done by implementing cellular mechanisms known to occur during the childhood development of working memory, such as an increased synaptic strength and improved contrast and specificity of the neural response. Our computational study shows that these mechanisms are sufficient to create a neural network which can store information about multiple items through sustained neural activity. Furthermore, by using functional magnetic resonance imaging, we found that the information-activity curve predicted by the model corresponds to that in the human posterior parietal cortex during performance of working memory tasks, which is consistent with previous studies of brain activity related to working memory capacity in humans.

Adult↗

Dopaminergic basis for deficits in working memory but not attentional set-shifting in Parkinson's disease.

Although Parkinson's disease is a common neurodegenerative disorder characterised by its motoric symptoms, there is an increasing recognition of accompanying impairments in cognition that have a profound impact on the quality of life of these patients. These deficits predominantly affect executive function and impairments of working memory have been frequently reported. However, the underlying neurochemical and pathological basis for these deficits are not well understood. In this study, 20 patients were tested 'on' and 'off' levodopa (L-dopa) medication on a task that allowed different aspects of working memory function such as maintenance, retrieval and manipulation to be tested within the same general paradigm as well as on an unrelated test of attentional set-shifting, which is known to be sensitive to deficits in early Parkinson's disease. Compared to healthy volunteers, PD patients were impaired at manipulation more than maintenance or retrieval of information within working memory. The patients were also impaired at the attentional set-shifting task. However, whereas L-dopa ameliorated the working memory deficit in manipulation (improving both accuracy and cognitive response time), it had no effect on the attentional set-shifting impairment. These results confirm that working memory deficits in PD are both psychologically specific and related to dopamine depletion. It is anticipated that greater understanding of these mechanisms will lead to future therapeutic improvements.

Aged↗

Locating and fractionating working memory using functional neuroimaging: storage, maintenance, and executive functions.

Working memory encompasses the short-term store of information and operations on that information. We review functional neuroimaging studies that have attempted to determine cortical areas involved in working memory functions. Current research suggests distinct systems for verbal information, visual objects, and spatial locations. Passive storage buffers appear to be located in posterior brain areas, whereas active maintenance of the information involves ventrolateral prefrontal areas. More complex, executive operations appear to recruit dorsolateral prefrontal cortex and anterior cingulate cortex. The possible involvement of reciprocal circuits including frontal cortex, basal ganglia, and thalamus is also discussed.

Brain↗