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Visual working memory for image statistics.

To define the role of statistical features of images in visual working memory, we compared the ability of subjects (N=6) to identify changes in arrays of black and white checks when these changes altered some aspect of their statistical structure, versus when these changes did not. Alteration of luminance statistics or local higher-order statistics improved performance, but alteration of the degree of bilateral symmetry did not. The dependence of performance on the degree of statistical change indicated that statistical information was represented in a graded, rather than categorical, fashion.

Adult↗

Functional topography of a distributed neural system for spatial and nonspatial information maintenance in working memory.

We investigated the degree to which the distributed and overlapping patterns of activity for working memory (WM) maintenance of objects and spatial locations are functionally dissociable. Previous studies of the neural system responsible for maintenance of different types of information in WM have reported seemingly contradictory results concerning the degree to which spatial and nonspatial information maintenance leads to distinct patterns of activation in prefrontal cortex. These inconsistent results may be partly attributable to the fact that different types of objects were used for the "object WM task" across studies. In the current study, we directly compared the patterns of response during WM tasks for face identity, house identity, and spatial location using functional magnetic resonance imaging (fMRI). Furthermore, independence of the neural resources available for spatial and object WM was tested behaviorally using a dual-task paradigm. Together, these results suggest that the mechanisms for the maintenance of house identity information are distributed and overlapping with those that maintain spatial location information, while the mechanisms for maintenance of face identity information are relatively more independent. There is, however, a consistent functional topography that results in superior prefrontal cortex producing the greatest response during spatial WM tasks, and middle and inferior prefrontal cortices producing their greatest responses during object WM tasks, independent of the object type. These results argue for a dorsal-ventral functional organization for spatial and nonspatial information. However, objects may contain both spatial and nonspatial information and, thus, have a distributed but not equipotent representation across both dorsal and ventral prefrontal cortex.

Adolescent↗

Working memory and retrieval: a resource-dependent inhibition model.

Four experiments examined individual differences in working memory (WM) capacity and how those differences affect performance on retrieval from both primary and secondary memory. The results showed that WM differences appear only in retrieval from primary memory and then only under conditions that lead to interference or response competition within the task. This suggests that WM capacity is important to retrieval that is based on controlled effortful search but not search that is based on automatic activation. A view is presented suggesting that individual differences in attentional resources lead to differences in the ability to inhibit or suppress irrelevant information. The paradigm also allowed more general comparisons between the processes involved in retrieval from primary and secondary memory. As expected, it was found that retrieval from primary memory was a function of set size. However, for sets larger than 2 items, retrieval from secondary memory was independent of set size.

Analysis of Variance↗

The phonological store of working memory: is it phonological and is it a store?

The phonological store construct of the working memory model is critically evaluated. Three experiments test the prediction that the effect of irrelevant sound and the effect of phonological similarity each survive the action of articulatory suppression but only when presentation of to-be-remembered lists is auditory, not visual. No evidence was found to support the interaction predicted among irrelevant speech, modality, and articulatory suppression. Although evidence for an interaction among modality, phonological similarity, and articulatory suppression was found, its presence could be diminished by a suffix, which is an acoustic, not a phonological factor. Coupled with other evidence--from the irrelevant sound effect and errors in natural speech--the action attributed to the phonological store seems better described in terms of a combination of auditory-perceptual and output planning mechanisms.

Humans↗

Changes in working memory after traumatic brain injury in children.

The impact of traumatic brain injury (TBI) on working memory (WM) was studied in 144 children (79 with mild, 23 with moderate, and 42 with severe injuries) who underwent magnetic resonance imaging (MRI) at 3 months and were tested at baseline and at 3, 6, 12, and 24 months postinjury. An n-back WM task for letter identity was administered with memory load ranging from 1- to 3-back and a 0-back condition. A TBI Severity x Quadratic Tune interaction showed that net percentage correct (correct detections of targets minus false alarms) was significantly lower in severe than in mild TBI groups. The Left Frontal Lesions x Age interaction approached significance. Mechanisms mediating late decline in WM and the effects of left frontal lesions are discussed.

Adolescent↗

Spiders crawl easily through the bottleneck: visual working memory for negative stimuli.

The special status of spiders in the attentional bottleneck and visual working memory (VWM) was studied. 23 spider-fearfuls (SF) and 23 non-anxious controls (NACs) participated in a serial VWM-task. Each trial showed a 4 x 4 matrix of images and 5 of these were subsequently cued for 150 ms each. Afterwards, one of the 16 displayed images was hidden and probed. The spider image was included in the string of 5 cued images, among the 11 uncued items, or not at all. For both groups, memory was better for cued spiders than for other cued items. SFs also showed improved memory for uncued spiders. The relevance of the results for theories of attention and cognitive models of phobias are discussed.

Adolescent↗

Age differences in working memory tasks: the role of secondary memory and the central executive system.

Two experiments are reported in which age differences in working memory performance are explored. In the first study, young and older adults held 2, 3, 4, or 5 unrelated words in mind while verifying a single or complex sentence. An age-related decrement was found in subsequent serial recall of the words, and this decrement was larger with longer word lists. Experiment 2 confirmed the interaction between age and list length, using list lengths of 4, 6, and 8 words and a free-recall procedure. There was no interaction between age and divided attention in either experiment. Surprisingly, sentence complexity had a greater detrimental effect on recall in the younger group. The results are discussed in terms of articulatory rehearsal being augmented by using secondary memory in the case of younger subjects.

Adolescent↗

Interference with rehearsal in spatial working memory in the absence of eye movements.

We have previously argued that rehearsal in spatial working memory is interfered with by spatial attention shifts rather than simply by movements to locations in space (Smyth & Scholey, 1994). It is possible, however, that the stimuli intended to induce attention shifts in our experiments also induced eye movements and interfered either with an overt eye movement rehearsal strategy or with a covert one. In the first experiment reported here, subjects fixated while they maintained a sequence of spatial items in memory before recalling them in order. Fixation did not affect recall, but auditory spatial stimuli presented during the interval did decrease performance, and it was further decreased if the stimuli were categorized as corning from the right or the left. A second experiment investigated the effects of auditory spatial stimuli to which no response was ever required and found that these did not interfere with performance, indicating that it is the spatial salience of targets that leads to interference. This interference from spatial input in the absence of any overt movement of the eyes or limbs is interpreted in terms of shifts of spatial attention or spatial monitoring, which Morris (1989) has suggested affects spatial encoding and which our findings suggest also affects reactivation in rehearsal.

Adult↗

The effects of electromagnetic field emitted by GSM phones on working memory.

The influence of pulsed radiofrequency (RF) electromagnetic fields of digital GSM mobile phones on working memory in healthy subjects were studied. Memory load was varied from 0 to 3 items in an n-back task. Each subject was tested twice within a single session, with and without the RF exposure (902MHz, 217Hz). The RF field speeded up response times when the memory load was three items but no effects of RF were observed with lower loads. The results suggest that RF fields have a measurable effect on human cognitive performance and encourage further studies on the interactions of RF fields with brain function.

Adolescent↗

Computerized training of working memory in children with ADHD--a randomized, controlled trial.

OBJECTIVE: Deficits in executive functioning, including working memory (WM) deficits, have been suggested to be important in attention-deficit/hyperactivity disorder (ADHD). During 2002 to 2003, the authors conducted a multicenter, randomized, controlled, double-blind trial to investigate the effect of improving WM by computerized, systematic practice of WM tasks. METHOD: Included in the trial were 53 children with ADHD (9 girls; 15 of 53 inattentive subtype), aged 7 to 12 years, without stimulant medication. The compliance criterion (>20 days of training) was met by 44 subjects, 42 of whom were also evaluated at follow-up 3 months later. Participants were randomly assigned to use either the treatment computer program for training WM or a comparison program. The main outcome measure was the span-board task, a visuospatial WM task that was not part of the training program. RESULTS: For the span-board task, there was a significant treatment effect both post-intervention and at follow-up. In addition, there were significant effects for secondary outcome tasks measuring verbal WM, response inhibition, and complex reasoning. Parent ratings showed significant reduction in symptoms of inattention and hyperactivity/impulsivity, both post-intervention and at follow-up. CONCLUSIONS: This study shows that WM can be improved by training in children with ADHD. This training also improved response inhibition and reasoning and resulted in a reduction of the parent-rated inattentive symptoms of ADHD.

Attention Deficit Disorder with Hyperactivity↗

Event-related potential correlates of impaired visuospatial working memory in schizophrenia.

Previous studies have reported that patients with schizophrenia demonstrate impaired performance during working memory (WM) tasks. The current study aimed to determine whether WM impairments in schizophrenia are accompanied by reduced slow wave (SW) activity during on-line maintenance of mnemonic information. Event-related potentials were obtained from patients with schizophrenia and well controls as they performed a visuospatial delayed response task. On 50% of trials, a distractor stimulus was introduced during the delay. Compared with controls, patients with schizophrenia produced less SW memory negativity, particularly over the right hemisphere, together with reduced frontal enhancement of SW memory negativity in response to distraction. The results indicate that patients with schizophrenia generate less maintenance phase neuronal activity during WM performance, especially under conditions of distraction.

Adult↗

Prefrontal task-related activity representing visual cue location or saccade direction in spatial working memory tasks.

To examine what kind of information task-related activity encodes during spatial working memory processes, we analyzed single-neuron activity in the prefrontal cortex while two monkeys performed two different oculomotor delayed-response (ODR) tasks. In the standard ODR task, monkeys were required to make a saccade to the cue location after a 3-s delay, whereas in the rotatory ODR (R-ODR) task, they were required to make a saccade 90 degrees clockwise from the cue location after the 3-s delay. By comparing the same task-related activities in these two tasks, we could determine whether such activities encoded the location of the visual cue or the direction of the saccade. One hundred twenty one neurons exhibited task-related activity in relation to at least one task event in both tasks. Among them, 41 neurons exhibited directional cue-period activity, most of which encoded the location of the visual cue. Among 56 neurons with directional delay-period activity, 86% encoded the location of the visual cue, whereas 13% encoded the direction of the saccade. Among 57 neurons with directional response-period activity, 58% encoded the direction of the saccade, whereas 35% encoded the location of the visual cue. Most neurons whose response-period activity encoded the location of the visual cue also exhibited directional delay-period activity that encoded the location of the visual cue as well. The best directions of these two activities were identical, and most of these response-period activities were postsaccadic. Therefore this postsaccadic activity can be considered a signal to terminate unnecessary delay-period activity. Population histograms encoding the location of the visual cue showed tonic sustained activation during the delay period. However, population histograms encoding the direction of the saccade showed a gradual increase in activation during the delay period. These results indicate that the transformation from visual input to motor output occurs in the dorsolateral prefrontal cortex. The analysis using population histograms suggests that this transformation occurs gradually during the delay period.

Animals↗

Motion information is spatially localized in a visual working-memory task.

We asked if the information about stimulus motion used in a visual working-memory task is localized in space. Monkeys compared the directions of two moving random-dot stimuli, sample and test, separated by a temporal delay and reported whether the stimuli moved in the same or in different directions. By presenting the two comparison stimuli in separate locations in the visual field, we determined whether information about stimulus direction was spatially localized during the storage and retrieval/comparison components of the task. Two psychophysical measures of direction discrimination provided nearly identical estimates of the critical spatial separation between sample and test stimuli that lead to a loss in threshold. Direction range thresholds measured with dot stimuli consisting of a range of local directional vectors were affected by spatial separation when a random-motion mask was introduced during the delay into the location of the upcoming test. The selective masking at the test location suggests that the information about the remembered direction was localized and available at that location. Direction difference thresholds, measured with coherently moving random dots, were also affected by separation between the two comparison stimuli. The separation at which performance was affected in both tasks increased with retinal eccentricity in parallel with the increase in receptive-field size in neurons in cortical area MT. The loss with transfer of visual information between different spatial locations suggests a contribution of cortical areas with localized receptive fields to the performance of the memory task. The similarity in the spatial scale of the storage mechanism derived psychophysically and the receptive field size of neurons in area MT suggest that MT neurons are central to this task.

Animals↗

The influence of working-memory demand and subject performance on prefrontal cortical activity.

Brain imaging and behavioral studies of working memory (WM) converge to suggest that the ventrolateral prefrontal cortex (PFC) mediates a capacity-limited storage buffer and that the dorsolateral PFC mediates memory organization processes that support supracapacity memory storage. Previous research from our laboratory has shown that the extent to which such memory organization processes are required depends on both task factors (i.e., memory load) and subject factors (i.e., response speed). Task factors exert their effects mainly during WM encoding while subject factors exert their effects mainly during WM retrieval. In this study, we sought to test the generalizability of these phenomena under more difficult memory-demand conditions than have been used previously. During scanning, subjects performed a WM task in which they were required to maintain between 1 and 8 letters over a brief delay. Neural activity was measured during encoding, maintenance, and retrieval task periods using event-related functional magnetic resonance imaging. With increasing memory load, there were reaction time increases and accuracy rate decreases, ventrolateral PFC activation decreases during encoding, and dorsolateral PFC activation increases during maintenance and retrieval. These results suggest that the ventrolateral PFC mediates WM storage and that the dorsolateral PFC mediates strategic memory organization processes that facilitate supracapacity WM storage. Additionally, high-performing subjects showed overall less activation than low-performing subjects, but activation increases with increasing memory load in the lateral PFC during maintenance and retrieval. Low-performing subjects showed overall more activation than high-performing subjects, but minimal activation increases in the dorsolateral PFC with increasing memory load. These results suggest that individual differences in both neural efficiency and cognitive strategy underlie individual differences in the quality of subjects' WM performance.

Adult↗

Cognitive processes in children's reading and attention: the role of working memory, divided attention, and response inhibition.

Children experiencing attention difficulties have documented cognitive deficits in working memory (WM), response inhibition and dual tasks. Recent evidence suggests however that these same cognitive processes are also closely associated with reading acquisition. This paper therefore explores whether these variables predicted attention difficulties or reading among 123 children with and without significant attention problems sampled from the school population. Children were screened using current WM and attention task measures. Three factors explained variance in WM and attention tasks. Response inhibition tasks loaded mainly with central executive measures, but a dual processing task loaded with the visual-spatial WM measures. Phonological loop measures loaded independently of attention measures. After controls for age, IQ and attention-group membership, phonological loop and 'central processing' measures both predicted reading ability. A 'visual memory/dual-task' factor predicted attention group membership after controls for age, IQ and reading ability. Results thus suggest that some of the processes previously assumed to be predictive of attention problems may reflect processes involved in reading acquisition. Visual memory and dual-task functioning are, however, purer indices of cognitive difficulty in children experiencing attention problems.

Attention↗

Serial attention within working memory.

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

Adult↗

Visual working memory for simple and complex visual stimuli.

Does the magical number four characterize our visual working memory (VWM) capacity for all kinds of objects, or is the capacity of VWM inversely related to the perceptual complexity of those objects? To find out how perceptual complexity affects VWM, we used a change detection task to measure VWM capacity for six types of stimuli of different complexity: colors, letters, polygons, squiggles, cubes, and faces. We found that the estimated capacity decreased for more complex stimuli, suggesting that perceptual complexity was an important factor in determining VWM capacity. However, the considerable correlation between perceptual complexity and VWM capacity declined significantly if subjects were allowed to view the sample memory display longer. We conclude that when encoding limitations are minimized, perceptual complexity affects, but does not determine, VWM capacity.

Adolescent↗

Involvement of the mammillary bodies in spatial working memory revealed by cytochrome oxidase activity.

In view of the inconclusive findings relating the nuclei of the mammillary bodies (MB) with spatial memory, we evaluated the oxidative metabolic activity of the medial and lateral nuclei of the mammillary bodies (MB) after training young rats (30 days) of both sexes in the Morris water maze. Different groups were trained in spatial working (WM) or reference memory (RM) tasks, respectively. The corresponding naïve groups swam for the same amount of time as the trained groups but without the escape platform. Control groups were added that had not been manipulated in any way. No sex-related differences were detected in the working memory task although males exhibited better reference memory than females. Cytochrome oxidase (CO) activity, an endogenous metabolic marker for neuronal activity, was measured in all the groups. CO activity increased significantly in both MB nuclei of male and female rats only in the spatial working memory group. In addition, high CO activity in the lateral nucleus of the MB was linearly correlated with lower escape latencies in both sexes after training in the working memory task. No CO activity changes were found in the basolateral amygdala (BL) in any of the experimental groups. This nucleus was used as a control brain region because of its participation in emotional behavior. The results suggest a specific role of the MB nuclei in spatial working memory in both sexes.

Animals↗