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Working memory for sign language: a window into the architecture of the working memory system.

Traditionally, working memory has been divided into two major domains: verbal and visuo-spatial. The verbal domain of working memory can be characterized either by its relationship to language or by its grounding in auditory processing. Because of this ambiguity, languages that are not auditory and vocal (i.e., signed languages) pose a challenge to this conception of working memory. We describe several experiments with deaf users of American Sign Language (ASL) that explore the extent to which the architecture of working memory is determined by the constraints of auditory and visual processing and the extent to which it is determined by the characteristics of language. Various working memory effects were investigated: phonological similarity, word length, and articulatory suppression. The pattern of evidence strongly supports the existence of a sign-based 'rehearsal loop' mechanism parallel to the speech-based rehearsal loop. However, we also discuss evidence pointing to differences between the speech loop and the sign loop from forward and backward digit span tasks with deaf and hearing subjects. Despite their similarities based on linguistic properties, the speech loop and the sign loop appear to diverge due to the differing processing demands of audition and vision. Overall, the results suggest that the architecture of working memory is shaped both by the properties of language structure and by the constraints imposed by sensorimotor modality.

Journal Article↗

Separable neuronal circuitries for manipulable and non-manipulable objects in working memory.

Previous work using single-cell recordings in monkeys and neuro-imaging studies in humans has shown that perceiving an object or imaging the action associated with the object recruits the same brain regions in the ventral premotor cortex as performing an action with the object. We used functional magnetic resonance imaging (fMRI) for examining whether similar brain regions are also activated while maintaining information about manipulable objects in working memory. Holding information about manipulable objects in working memory activated the left ventral premotor cortex and the left inferior frontal gyrus (Broca's area). Conversely, non-manipulable objects to be held in working memory co-activated Broca's area and the left angular gyrus. When contrasted directly, manipulable relative to non-manipulable objects activated the left ventral premotor cortex and the anterior intraparietal sulcus, a circuitry that is assumed to mediate the transformation of movement-relevant object properties into hand actions. These results indicate that visual working memory for manipulable objects is based on motor programmes associated with their use. Similar to speech motor programmes in verbal memory tasks, hand motor programmes may allow the maintenance of objects in working memory over short intervals.

Adult↗

Phonological deterioration in adults with an acquired severe hearing impairment: a deterioration in long-term memory or working memory?

The purpose of this study was to examine the phonological processing skills in individuals with an acquired severe hearing impairment. The subjects were tested on cognitive tasks that vary in their demands on phonological processing. The severely hearing impaired individuals performed at a significantly lower level on the word-word rhyme judgements task, but performed on a par with the control group on the picture-word rhyme judgements task and the lexical decision-making task. The results indicate that the phonological processing skills in individuals who have acquired a severe hearing loss in adult life deteriorate. The results are discussed with respect to theoretical and clinical implications.

Adult↗

Episodic memory meets working memory in the frontal lobe: functional neuroimaging studies of encoding and retrieval.

Recent functional-neuroimaging studies have provided a wealth of new information suggesting that regions of the prefrontal cortex play a role in episodic memory encoding and retrieval. This review seeks to evaluate the results of these studies in the context of one general model that has proposed that the left prefrontal cortex is preferentially involved in episodic memory encoding, whereas the right prefrontal cortex is preferentially involved in episodic memory retrieval, irrespective of the type (e.g., modality) of information being remembered. The origins of this framework are considered in some detail and then all relevant functional-neuroimaging studies are critically reviewed. The results of this review fail to provide support for the functional-asymmetry model, suggesting instead that episodic memory encoding and retrieval may actually involve similar regions of the lateral prefrontal cortex when all factors relating to the type of stimulus material (i.e., modality), are appropriately controlled.

Brain Mapping↗

Distinct capacity limits for attention and working memory: Evidence from attentive tracking and visual working memory paradigms.

A hallmark of both visual attention and working memory is their severe capacity limit: People can attentively track only about four objects in a multiple object tracking (MOT) task and can hold only up to four objects in visual working memory (VWM). It has been proposed that attention underlies the capacity limit of VWM. We tested this hypothesis by determining the effect of varying the load of a MOT task performed during the retention interval of a VWM task and comparing the resulting dual-task costs with those observed when a VWM task was performed concurrently with another VWM task or with a verbal working memory task. Instead of supporting the view that the capacity limit of VWM is solely attention based, the results indicate that VWM capacity is set by the interaction of visuospatial attentional, central amodal, and local task-specific sources of processing.

Attention↗

Roles of working memory capacity and long-term working memory skill in complex task performance.

In the present research, we examined the relative roles of domain-general and domain-specific individual difference characteristics in complex cognitive task performance. Specifically, we examined the impact both of working memory (WM) capacity and of acquired skills used to encode presented information in an accessible form in long-term working memory (LTWM) on performance in a complex aviation task environment. Measures of WM capacity and LTWM skill served as performance predictors. A criterion measure of task performance was related to the predictor measures. The results indicated that an increase in LTWM skill decreases the role of WM capacity as the determinant of complex task performance, although both measures are important performance predictors. We discuss how the two distinct WM constructs coexist and interact to support complex task performance.

Cognition↗

Memory processes of flight situation awareness: interactive roles of working memory capacity, long-term working memory, and expertise.

This research examined the role of working memory (WM) capacity and long-term working memory (LT-WM) in flight situation awareness (SA). We developed spatial and verbal measures of WM capacity and LT-WM skill and then determined the ability of these measures to predict pilot performance on SA tasks. Although both spatial measures of WM capacity and LT-WM skills were important predictors of SA performance, their importance varied as a function of pilot expertise. Spatial WM capacity was most predictive of SA performance for novices, whereas spatial LT-WM skill based on configurations of control flight elements (attitude and power) was most predictive for experts. Furthermore, evidence for an interactive role of WM and LT-WM mechanisms was indicated. Actual or potential applications of this research include cognitive analysis of pilot expertise and aviation training.

Acceleration↗

Methylphenidate restores visual memory, but not working memory function in attention deficit-hyperkinetic disorder.

RATIONALE: Dysfunction of executive neuropsychological performance, mediated by the prefrontal cortex, has been the central focus of recent attention deficit/ hyperkinetic disorder (AD-HKD) research. The role of other potential neuropsychological "risk factors", such as recognition memory, remains understudied. Further, the impact of methylphenidate (MPH) on key neuropsychological processes in AD-HKD remains poorly understood. OBJECTIVES: To compare the performance of boys with AD-HKD on a spatial working memory (SWM) task and on two non-working memory tasks [a simultaneous and delayed matching-to-sample task (DMtS) and a pattern-recognition task] with that of healthy boys, and to investigate the impact of acute and chronic MPH on performance of these tasks. METHODS: Baseline performance of 75 stimulant-naive boys with AD-HKD was compared with that of 70 healthy boys. The AD-HKD boys were then re-tested following the administration of acute and chronic challenges with MPH (0.3 mg/kg and 0.6 mg/kg) under randomised double-blind placebo controlled conditions. RESULTS: Compared with healthy boys, the AD-HKD boys demonstrated performance deficits on all neuropsychological tasks. A single dose of MPH restored performance on the DMtS task but had no impact on the SWM or pattern-recognition tasks. Chronic MPH administration did not alter performance on the SWM task but did improve performance on both the pattern-recognition and DMtS tasks. However, the acute restorative effect of MPH on DMtS diminished with repeated administration. CONCLUSIONS: Our results suggest that current conceptualisations of the neuropsychological basis of AD-HKD and the proposed therapeutic mechanisms of MPH require broadening.

Adolescent↗

Brain areas involved in spatial working memory.

Spatial working memory entails the ability to keep spatial information active in working memory over a short period of time. To study the areas of the brain that are involved in spatial working memory, a group of stroke patients was tested with a spatial search task. Patients and healthy controls were asked to search through a number of boxes shown at different locations on a touch-sensitive computer screen in order to find a target object. In subsequent trials, new target objects were hidden in boxes that were previously empty. Within-search errors were made if a participant returned to an already searched box; between-search errors occurred if a participant returned to a box that was already known to contain a target item. The use of a strategy to remember the locations of the target objects was calculated as well. Damage to the right posterior parietal and right dorsolateral prefrontal cortex impaired the ability to keep spatial information 'on-line', as was indicated by performance on the Corsi Block-Tapping task and the within-search errors. Moreover, patients with damage to the right posterior parietal cortex, the right dorsolateral prefrontal cortex and the hippocampal formation bilaterally made more between-search errors, indicating the importance of these areas in maintaining spatial information in working memory over an extended time period.

Brain↗

Treatment with magnesium improves reference memory but not working memory while reducing GFAP expression following traumatic brain injury.

PURPOSE: Previous studies have shown that administration of MgCl2 in animal models of brain injury significantly improves functional recovery: however, few studies have examined cognitive recovery. The present study evaluated the effect of MgCl2 pharmacotherapy on recovery of function following medial frontal cortex contusion injury. METHODS: Groups of rats were assigned to either MgCl2 (1.0 mmol/kg) or saline treatment conditions and prepared with contusion injuries or shams. Drug treatment was administered 15 min and 24 hr following injury. Rats were examined on tests of sensorimotor performance (bilateral tactile adhesive removal) and cognitive ability (reference and working memory). RESULTS: Administration of MgCl2 following injury significantly reduced the behavioral impairments observed on the bilateral tactile removal test. The acquisition of reference memory was also significantly improved compared to saline-treated rats; however, treatment did not improve working memory performance. Lesion analysis revealed that administration of MgCl2 did not significantly reduce lesion size compared to saline-treatment. Examination of glial fibrillary acidic protein (GFAP) expression showed that MgCl2 did significantly reduce the number of GFAP+ cells. CONCLUSION: These results indicate that MgCl2 administration significantly improved behavioral outcome following injury in a task dependent manner and reduced GFAP expression.

Animals↗

Working memory and inferences: evidence from eye fixations during reading.

Eye fixations during reading were monitored to examine the relationship between individual differences in working memory capacity-as assessed by the reading span task-and inferences about predictable events. Context sentences predicting likely events, or non-predicting control sentences, were presented. They were followed by continuation sentences in which a target word represented an event to be inferred (inferential word) or an unlikely event (non-predictable word). A main effect of reading span showed that high working memory capacity was related to shorter gaze durations across sentence regions. More specific findings involved an interaction between context, target, and reading span on late processing measures and regions. Thus, for high- but not for low-span readers, the predicting condition, relative to the control condition, facilitated reanalysis of the continuation sentence that represented the inference concept. This effect was revealed by a reduction in regression-path reading time in the last region of the sentence, involving less time reading that region and fewer regressions from it. These results indicate that working memory facilitates elaborative inferences during reading, but that this occurs at late text-integration processes, rather than at early lexical-access processes.

Cognition↗

Word length and phonological similarity effects in simple, complex, and delayed serial recall tasks: implications for working memory.

Some current models of working memory argue that a passive short-term store is not involved in more dynamic working memory tasks. Other models argue that standard short-term memory and working memory tasks rely on common storage facilities. We examine these issues by exploring two signature effects of passive short-term storage in simple span, complex span, and Brown-Peterson tasks. The finding that all three tasks show word length and phonological similarity effects suggests that common processes or storage mechanisms are involved in all tasks. The implications for models of working memory are discussed.

Humans↗

Prefrontal cortex and working memory processes.

Working memory is a mechanism for short-term active maintenance of information as well as for processing maintained information. The dorsolateral prefrontal cortex has been known to participate in working memory. The analysis of task-related dorsolateral prefrontal cortex activity while monkeys performed a variety of working memory tasks revealed that delay-period activity is a neural correlate of a mechanism for temporary active maintenance of information, because this activity persisted throughout the delay period, showed selectivity to a particular visual feature, and was related to correct behavioral performances. Information processing can be considered as a change of the information represented by a population of neural activities during the progress of the trial. Using population vectors calculated by a population of task-related dorsolateral prefrontal cortex activities, we demonstrated the temporal change of information represented by a population of dorsolateral prefrontal cortex activities during performances of spatial working memory tasks. Cross-correlation analysis using spike firings of simultaneously isolated pairs of neurons reveals widespread functional interactions among neighboring neurons, especially neurons having delay-period activity, and their dynamic modulation depending on the context of the trial. Functional interactions among neurons and their dynamic modulation could be a mechanism of information processing in the dorsolateral prefrontal cortex.

Action Potentials↗

Temporal dynamics of brain activation during a working memory task.

Working memory is responsible for the short-term storage and online manipulation of information necessary for higher cognitive functions, such as language, planning and problem-solving. Traditionally, working memory has been divided into two types of processes: executive control (governing the encoding manipulation and retrieval of information in working memory) and active maintenance (keeping information available 'online'). It has also been proposed that these two types of processes may be subserved by distinct cortical structures, with the prefrontal cortex housing the executive control processes, and more posterior regions housing the content-specific buffers (for example verbal versus visuospatial) responsible for active maintenance. However, studies in non-human primates suggest that dorsolateral regions of the prefrontal cortex may also be involved in active maintenance. We have used functional magnetic resonance imaging to examine brain activation in human subjects during performance of a working memory task. We used the temporal resolution of this technique to examine the dynamics of regional activation, and to show that prefrontal cortex along with parietal cortex appears to play a role in active maintenance.

Adult↗

The primate working memory networks.

Working memory has long been associated with the prefrontal cortex, since damage to this brain area can critically impair the ability to maintain and update mnemonic information. Anatomical and physiological evidence suggests, however, that the prefrontal cortex is part of a broader network of interconnected brain areas involved in working memory. These include the parietal and temporal association areas of the cerebral cortex, cingulate and limbic areas, and subcortical structures such as the mediodorsal thalamus and the basal ganglia. Neurophysiological studies in primates confirm the involvement of areas beyond the frontal lobe and illustrate that working memory involves parallel, distributed neuronal networks. In this article, we review the current understanding of the anatomical organization of networks mediating working memory and the neural correlates of memory manifested in each of their nodes. The neural mechanisms of memory maintenance and the integrative role of the prefrontal cortex are also discussed.

Animals↗

Working memory in spelling: evidence from backward typing.

Theories of spelling (Margolin, 1984; Nolan & Caramazza, 1983) propose a working memory system for storing order and identity information of letters during the spelling process. Capacity limitations related to the use of such a graphemic buffer were explored. Participants had to type words backwards. Longer pauses between key presses were assumed to signal points at which graphemic buffer contents were refreshed. Five- and six-letter words were divided by a major pause into chunks of two and three letters, partly coinciding with syllables. Articulatory suppression had no effect on performance. Increasing the length of the stimuli to seven to eight letters resulted in major pauses occurring at syllable boundaries, and performance becoming vulnerable to articulatory suppression but not foot tapping. Forward typing resulted in a similar pause pattern. The results suggest that chunks of approximately three letters can be handled at any one time. For short words the task seems to rely on non-phonological modes of coding, whereas longer words appear to require the use of a phonological code, possibly for keeping track of progress through the word.

Humans↗

Organization of working memory within the human prefrontal cortex: a PET study of self-ordered object working memory.

The prefrontal cortex plays a critical role in working memory, the active maintenance of information for brief periods of time for guiding future motor and cognitive processes. Two competing models have emerged to account for the growing human and non-human primate literature examining the functional neuroanatomy of working memory. One theory holds that the lateral frontal cortex plays a domain-specific role in working memory with the dorsolateral and ventrolateral cortical regions supporting working memory for spatial and non-spatial material, respectively. Alternatively, the lateral frontal cortex may play a process-specific role with the more dorsal regions becoming recruited whenever active manipulation or monitoring of information in working memory becomes necessary. Many working memory tasks do not allow for direct tests of these competing models. The present study used a novel self-ordered working memory task and positron emission tomography to identify whether dorsal or ventral lateral cortical areas are recruited during a working memory task that required extensive monitoring of non-spatial information held within working memory. We observed increased blood flow in the right dorsolateral, but not ventrolateral, prefrontal cortex. Increases in blood flow in the dorsolateral region correlated strongly with task performance. Thus, the results support the process-specific hypothesis.

Adult↗

Self-initiated encoding facilitates object working memory in schizophrenia: implications for the etiology of working memory deficit.

BACKGROUND: Working memory (WM) deficit is present in a majority of patients with schizophrenia but it is unclear which components of WM are impaired. Past studies suggest that encoding may be compromised. One important determinant of encoding is the deployment of selective attention to the target stimulus. In addition, attention and encoding are modulated by motivational factors. In this study, we investigated the effects of self-initiated encoding (i.e., voluntary attention) on WM. METHODS: 19 patients with schizophrenia and 19 matched control subjects participated in visual WM and control tasks. Encoding was manipulated by asking subjects to select from two face targets and memorize 1) one of the two identical faces (Non-preference condition), 2) one that is marked (Non-choice condition), and 3) one they prefer (Preference condition). WM accuracy for both location (spatial) and identity (object) was measured. RESULTS: Overall, patients with schizophrenia were less accurate and slower than the control subjects but the deficit was greater for object WM. However, patients were more accurate in object WM when they selected a preferred face as their target during encoding (preference condition) compared with the other two conditions. This effect was not significant for spatial WM. CONCLUSIONS: These results suggest that voluntary, self-initiated attention may facilitate object encoding especially if the selection of the target involves affective choice, and that attention may play different roles in encoding 'what' versus 'where' in WM. Since encoding affects all forms of memory, these results may have a more general implication for memory.

Adult↗