Nonspatial Working Memory Tasks Research Articles

Interaction between visual working memory and upright postural control in young adults: an event-related potential study based on the n-back paradigm.

As a part of the overall information-processing system of the brain, postural control is related to the cognitive processes of working memory. Previous studies have suggested that cognitive tasks and postural control processes can compete for resources in common brain areas, although there is an "inverted U" relationship between arousal level and behavioral control - the arousal level of individuals changes when performing cognitive tasks. However, the exact neural connections between the two are unclear. This may be related to the nature of cognitive tasks. Some studies believe that posture occupies not only spatial information processing resources but also visual non-spatial information processing resources. Other studies believe that posture control only occupies spatial information processing resources in the central system, but does not occupy non-spatial information processing resources. Previous studies used different cognitive task materials and reached different conclusions. In this study, we used the same visuospatial and non-spatial materials, the n-back visual working memory paradigm, the event-related potential technique to investigate the effects of visuospatial and non-spatial working memory tasks on adolescents' postural control under different cognitive loads. The results of this study showed that in both visuospatial and non-spatial conditions, the N1 effect of the parieto-occipital lobe was larger during upright posture than in the sitting position (160-180 ms), the P300 effect of the central parieto-occipital region (280-460 ms) was induced by working memory in different postures, and the P300 wave amplitude was higher in the sitting position than in the upright position. We demonstrated that upright postural control enhances early selective attention but interferes with central memory encoding, thus confirming that postural control and visuospatial and non-spatial working memory share brain regions and compete with each other.

Stimulus-driven attention and cognitive control during encoding: An event related brain potentials study

Stimulus-driven attention drawn to relevant items can improve working memory (WM) whether attentional capture is driven by salient, low level features or by contingent salience from shared features with targets. In the current work, we examined the time course of enhanced attention to contingently salient information in a non-spatial WM task using event related brain potentials (ERPs). In line with previous work, we predicted that the encoding of contingently salient stimuli would be associated with an enhancement of cognitive control processes rather than low-level salience detection. The results of this study supported this hypothesis, evidenced by a posterior P3 component of greater amplitude for contingently salient stimuli relative to stimuli of a control color, which is thought to reflect enhanced attention to information that matches a target held in WM. However, P3 amplitude during encoding was unrelated to subsequent memory accuracy. As an exploratory follow up on these results, we conducted a regression analysis including beliefs about ability to focus attention as a moderator, which interacted with P3 amplitude to predict WM recall of salient letters. Furthermore, source localization analyses implicated a significant contribution of regions in the salience network to the detection of target stimuli, but only frontal control regions showed a greater response to salient than control letters. Thus, the results of this experiment suggest that participants enhance cognitive control during the encoding of contingently salient stimuli, but that the relationship between this neural process during encoding and subsequent benefits to WM recall might depend on individual differences in attentional focus.

Different neural activities support auditory working memory in musicians and bilinguals.

Musical training and bilingualism benefit executive functioning and working memory (WM)-however, the brain networks supporting this advantage are not well specified. Here, we used functional magnetic resonance imaging and the n-back task to assess WM for spatial (sound location) and nonspatial (sound category) auditory information in musician monolingual (musicians), nonmusician bilinguals (bilinguals), and nonmusician monolinguals (controls). Musicians outperformed bilinguals and controls on the nonspatial WM task. Overall, spatial and nonspatial WM were associated with greater activity in dorsal and ventral brain regions, respectively. Increasing WM load yielded similar recruitment of the anterior-posterior attention network in all three groups. In both tasks and both levels of difficulty, musicians showed lower brain activity than controls in superior prefrontal frontal gyrus and dorsolateral prefrontal cortex (DLPFC) bilaterally, a finding that may reflect improved and more efficient use of neural resources. Bilinguals showed enhanced activity in language-related areas (i.e., left DLPFC and left supramarginal gyrus) relative to musicians and controls, which could be associated with the need to suppress interference associated with competing semantic activations from multiple languages. These findings indicate that the auditory WM advantage in musicians and bilinguals is mediated by different neural networks specific to each life experience.

Alpha-Band Activity Reveals Spontaneous Representations of Spatial Position in Visual Working Memory

An emerging view suggests that spatial position is an integral component of working memory (WM), such that non-spatial features are bound to locations regardless of whether space is relevant [1, 2]. For instance, past work has shown that stimulus position is spontaneously remembered when non-spatial features are stored. Item recognition is enhanced when memoranda appear at the same location where they were encoded [3-5], and accessing non-spatial information elicits shifts of spatial attention to the original position of the stimulus [6, 7]. However, these findings do not establish that a persistent, active representation of stimulus position is maintained in WM because similar effects have also been documented following storage in long-term memory [8, 9]. Here we show that the spatial position of the memorandum is actively coded by persistent neural activity during a non-spatial WM task. We used a spatial encoding model in conjunction with electroencephalogram (EEG) measurements of oscillatory alpha-band (8-12Hz) activity to track active representations of spatial position. The position of the stimulus varied trial to trial but was wholly irrelevant to the tasks. We nevertheless observed active neural representations of the original stimulus position that persisted throughout the retention interval. Further experiments established that these spatial representations are dependent on the volitional storage of non-spatial features rather than being a lingering effect of sensory energy or initial encoding demands. These findings provide strong evidence that online spatial representations are spontaneously maintained in WM-regardless of task relevance-during the storage of non-spatial features.

CAMK2A polymorphisms predict working memory performance in humans

Working memory is impaired in a number of psychiatric conditions.1, 2 An important role of alpha Ca2+/calmodulin dependent protein kinase II in learning and memory has been shown in rodent models. However, these findings have not been confirmed in humans yet. Here, we report genetic associations between the human CAMK2A gene and the performance in a spatial and a non-spatial working memory (non-SWM) task in two independent human samples.

Object Working Memory Performance Depends on Microstructure of the Frontal-Occipital Fasciculus

Re-entrant circuits involving communication between the frontal cortex and other brain areas have been hypothesized to be necessary for maintaining the sustained patterns of neural activity that represent information in working memory, but evidence has so far been indirect. If working memory maintenance indeed depends on such temporally precise and robust long-distance communication, then performance on a delayed recognition task should be highly dependent on the microstructural integrity of white-matter tracts connecting sensory areas with prefrontal cortex. This study explored the effect of variations in white-matter microstructure on working memory performance in two separate groups of participants: patients with multiple sclerosis and age- and sex-matched healthy adults. Functional magnetic resonance imaging was performed to reveal cortical regions involved in spatial and object working memory, which, in turn, were used to define specific frontal to extrastriate white-matter tracts of interest via diffusion tensor tractography. After factoring out variance due to age and the microstructure of a control tract (the corticospinal tract), the number of errors produced in the object working memory task was specifically related to the microstructure of the inferior frontal-occipital fasciculus. This result held for both groups, independently, providing a within-study replication with two different types of white-matter structural variability: multiple sclerosis-related damage and normal variation. The results demonstrate the importance of interactions between specific regions of the prefrontal cortex and sensory cortices for a nonspatial working memory task that preferentially activates those regions.

Changes in cognitive functions of students in the transitional period from elementary school to junior high school

Background: When students proceed to junior high school from elementary school, rapid changes in the environment occur, which may cause various behavioral and emotional problems. However, the changes in cognitive functions during this transitional period have rarely been studied. Methods: In 158 elementary school students from 4th- to 6th-grades and 159 junior high school students from 7th- to 9th-grades, we assessed various cognitive functions, including motor processing, spatial construction ability, semantic fluency, immediate memory, delayed memory, spatial and non-spatial working memory, and selective, alternative, and divided attention. Results: Our findings showed that performance on spatial and non-spatial working memory, alternative attention, divided attention, and semantic fluency tasks improved from elementary to junior high school. In particular, performance on alternative and divided attention tasks improved during the transitional period from elementary to junior high school. Conclusion: Our finding suggests that development of alternative and divided attention is of crucial importance in the transitional period from elementary to junior high school.

Overlapping functional anatomy for working memory and visual search

Recent behavioural findings using dual-task paradigms demonstrate the importance of both spatial and non-spatial working memory processes in inefficient visual search (Anderson et al. in Exp Psychol 55:301–312, 2008). Here, using functional magnetic resonance imaging (fMRI), we sought to determine whether brain areas recruited during visual search are also involved in working memory. Using visually matched spatial and non-spatial working memory tasks, we confirmed previous behavioural findings that show significant dual-task interference effects occur when inefficient visual search is performed concurrently with either working memory task. Furthermore, we find considerable overlap in the cortical network activated by inefficient search and both working memory tasks. Our findings suggest that the interference effects observed behaviourally may have arisen from competition for cortical processes subserved by these overlapping regions. Drawing on previous findings (Anderson et al. in Exp Brain Res 180:289–302, 2007), we propose that the most likely anatomical locus for these interference effects is the inferior and middle frontal cortex of the right hemisphere. These areas are associated with attentional selection from memory as well as manipulation of information in memory, and we propose that the visual search and working memory tasks used here compete for common processing resources underlying these mechanisms.

Transient and selective overexpression of D 2 receptors in the striatum causes persistent deficits in conditional associative learning

Cognitive deficits in schizophrenia are thought to derive from a hypofunction of the prefrontal cortex (PFC), but the origin of the hypofunction is unclear. To explore the nature of this deficit, we genetically modified mice to model the increase in striatal dopamine D(2) receptors (D(2)Rs) observed in patients with schizophrenia. Previously, we reported deficits in spatial working memory tasks in these mice, congruent with the working memory deficits observed in schizophrenia. However, patients with schizophrenia suffer from deficits in many executive functions, including associative learning, planning, problem solving, and nonspatial working memory. We therefore developed operant tasks to assay two executive functions, conditional associative learning (CAL) and nonspatial working memory. Striatal D(2)R-overexpressing mice show a deficit in CAL because of perseverative behavior, caused by interference from the previous trial. D(2)R up-regulation during development was sufficient to cause this deficit, because switching off the transgene in adulthood did not rescue the phenotype. We validated prefrontal dependency of CAL by using neurotoxic lesions. Lesions of the medial PFC including the anterior cingulate, infralimbic, and prelimbic cortices impair CAL because of increased interference from previously rewarded trials, exactly as observed in D(2)R transgenic mice. In contrast, lesions restricted to the infralimbic and prelimbic cortices have no effect on CAL but impair performance in the nonspatial working memory task. These assays not only give us insight into how excess striatal D(2)Rs affect cognition but also provide tools for studying cognitive endophenotypes in mice.

A role for spatial and nonspatial working memory processes in visual search.

Searching a cluttered visual scene for a specific item of interest can take several seconds to perform if the target item is difficult to discriminate from surrounding items. Whether working memory processes are utilized to guide the path of attentional selection during such searches remains under debate. Previous studies have found evidence to support a role for spatial working memory in inefficient search, but the role of nonspatial working memory remains unclear. Here, we directly compared the role of spatial and nonspatial working memory for both an efficient and inefficient search task. In Experiment 1, we used a dual-task paradigm to investigate the effect of performing visual search within the retention interval of a spatial working memory task. Importantly, by incorporating two working memory loads (low and high) we were able to make comparisons between dual-task conditions, rather than between dual-task and single-task conditions. This design allows any interference effects observed to be attributed to changes in memory load, rather than to nonspecific effects related to "dual-task" performance. We found that the efficiency of the inefficient search task declined as spatial memory load increased, but that the efficient search task remained efficient. These results suggest that spatial memory plays an important role in inefficient but not efficient search. In Experiment 2, participants performed the same visual search tasks within the retention interval of visually matched spatial and verbal working memory tasks. Critically, we found comparable dual-task interference between inefficient search and both the spatial and nonspatial working memory tasks, indicating that inefficient search recruits working memory processes common to both domains.

Domain‐related differentiation of working memory in the Japanese macaque (Macaca fuscata) frontal cortex: a positron emission tomography study

The lateral prefrontal cortex (LPFC) is important for working memory (WM) task performance. Neuropsychological and neurophysiological studies in monkeys suggest that the lateral prefrontal cortex is functionally segregated based on the working memory domain (spatial vs. non-spatial). However, this is not supported by most human neuroimaging studies, and the discrepancy might be due to differences in methods and/or species (monkey neuropsychology/physiology vs. human neuroimaging). We used positron emission topography to examine the functional segregation of the lateral prefrontal cortex of Japanese macaques (Macaca fuscata) that showed near 100% accuracy on spatial and non-spatial working memory tasks. Compared with activity during the non-working memory control tasks, the dorsolateral prefrontal cortex (DLPFC) was more active during the non-spatial, but not during the spatial, working memory task, although a muscimol microinjection into the dorsolateral prefrontal cortex significantly impaired the performance of both working memory tasks. A direct comparison of the brain activity between the two working memory tasks revealed no differences within the lateral prefrontal cortex, whereas the premotor area was more active during the spatial working memory task. Comparing the delay-specific activity, which did not include task-associated stimulus/response-related activity, revealed more spatial working memory-related activity in the posterior parietal and premotor areas, and more non-spatial working memory-related activity in the dorsolateral prefrontal cortex and hippocampus. These results suggest that working memory in the monkey brain is segregated based on domain, not within the lateral prefrontal cortex but rather between the posterior parietal-premotor areas and the dorsolateral prefrontal-hippocampus areas.

Combined D1/D2 receptor stimulation under conditions of dopamine depletion impairs spatial working memory performance in humans

The mesocortical dopamine system is regarded as an important modulator of working memory. While it has been established that stimulation of the D1/D2 receptor in primates can improve spatial working memory performance, findings in humans are less consistent. Recent studies in humans suggest that global depletion of dopamine via tyrosine/phenylalanine depletion may impair spatial working memory performance, although these results are also inconsistent, and it has been suggested that task differences may partly underlie the inconsistent findings. This study had two aims: (1) to investigate the effects of acute tyrosine depletion (TPD) on a number of working memory tasks and (2) to examine whether stimulation of D1/D2 receptors under conditions of TPD can attenuate or "reverse" TPD-induced working memory impairments. Eighteen healthy male participants performed a spatial working memory delayed-recognition task, non-spatial working memory task and spatial n-back task on three separate occasions, after TPD, TPD and pergolide (D1/D2 agonist), and placebo. TPD did not impair working memory performance on any of the tasks administered. However, stimulation of D1/D2 receptors under TPD conditions caused a subtle impairment in spatial working memory performance. The finding that D1/D2 stimulation under TPD conditions impairs working memory highlights the complexity of functional effects of augmenting dopaminergic transmission within a dopamine-depleted state. The lack of TPD-related effects on a range of working memory tasks questions the reliability of TPD as a modulator of dopamine function and working memory performance in humans.

Altered resting state networks in mild cognitive impairment and mild Alzheimer's disease: An fMRI study

Activity and reactivity of the default mode network in the brain was studied using functional magnetic resonance imaging (fMRI) in 28 nondemented individuals with mild cognitive impairment (MCI), 18 patients with mild Alzheimer's disease (AD), and 41 healthy elderly controls (HC). The default mode network was interrogated by means of decreases in brain activity, termed deactivations, during a visual encoding task and during a nonspatial working memory task. Deactivation was found in the default mode network involving the anterior frontal, precuneus, and posterior cingulate cortex. MCI patients showed less deactivation than HC, but more than AD. The most pronounced differences between MCI, HC, and AD occurred in the very early phase of deactivation, reflecting the reactivity and adaptation of the network. The default mode network response in the anterior frontal cortex significantly distinguished MCI from both HC (in the medial frontal) and AD (in the anterior cingulate cortex). The response in the precuneus could only distinguish between patients and HC, not between MCI and AD. These findings may be consistent with the notion that MCI is a transitional state between healthy aging and dementia and with the proposed early changes in MCI in the posterior cingulate cortex and precuneus. These findings suggest that altered activity in the default mode network may act as an early marker for AD pathology.

GluR-A-dependent synaptic plasticity is required for the temporal encoding of nonspatial information.

Four related experiments studied operant performance of mice on differential reinforcement of low rates of responding (DRL) paradigms. Experiment 1 showed that excitotoxic hippocampal lesions impaired performance of a 10-s DRL schedule (DRL-10). Experiments 2 and 3 showed that GluR-A AMPA receptor subunit knockout mice, which are deficient in CA3-CA1 long-term potentiation (LTP), were markedly impaired at 15 s (DRL-15), but less impaired at DRL-10. Experiment 4 compared DRL-15 performance in mice from the 2 strains from which the GluR-A colony was derived and showed that they did not differ. The results show that GluR-A-containing AMPA receptors are required for normal performance on hippocampus-dependent, nonspatial working memory tasks, consistent with a role for GluR-A in the temporal encoding (what happened when) of nonspatial information.

The role of memory representation in the vigilance decrement

Working memory load is critically important for the overall level of performance on vigilance tasks. However, its role in a key aspect of vigilance-sensitivity decrement over time-is unclear. We used a dual-task procedure in which either a spatial or a nonspatial working memory task was performed simultaneously with a spatial vigilance task for 20 min. Sensitivity in the vigilance task declined over time when the concurrent task involved spatial working memory. In contrast, there was no sensitivity decrement with a nonspatial working memory task. The results provide the first evidence of a specific role for working memory representation in vigilance decrement. The findings are also consistent with a multiple resource theory in which separate resources for memory representation and cognitive control operations are differentially susceptible to depletion over time, depending on the demands of the task at hand.

The role of spatial working memory in visual search efficiency.

Many theories have proposed that visual working memory plays an important role in visual search. In contrast, by showing that a nonspatial working memory load did not interfere with search efficiency, Woodman, Vogel, and Luck (2001) recently proposed that the role of working memory in visual search is insignificant. However, the visual search process may interfere with spatial working memory. In the present study, a visual search task was performed concurrently with either a spatial working memory task (Experiment 1) or a nonspatial working memory task (Experiment 2). We found that the visual search process interfered with a spatial working memory load, but not with a nonspatial working memory load. These results suggest that there is a distinction between spatial and nonspatial working memory in terms of interactions with visual search tasks. These results imply that the visual search process and spatial working memory storage require the same limited-capacity mechanisms.

Differences between auditory evoked responses recorded during spatial and nonspatial working memory tasks

Results from several recent studies suggest that neuronal processing of sound content and its spatial location may be dissociated. The use of modern neuroimaging techniques has allowed for the determination that different brain structures may be specifically activated during working memory processing of pitch and location of sound. The time course of these task-related differences, however, remains uncertain. In the present study, we performed simultaneous whole-head electroencephalogram and magnetoencephalogram recordings, using a new behavioral paradigm, to investigate the dynamics of differences between “what” and “where” evoked responses in the auditory system as a function of memory load. In the location task the latency of the N1m was shorter and its generator was situated more inferiorly than in the pitch task. Working memory processing of the tonal frequency enhanced the amplitude of the N2 component, as well as the negative-going deflection at a latency around 400 ms. A memory-load-dependent task-related difference was found in the positive slow wave which was higher during the location than pitch task at the low load. Late slow waves were affected by memory load but not type of task. These results suggest that separate neuronal networks are involved in the attribute-specific analysis of auditory stimuli and their encoding into working memory, whereas the maintenance of auditory information is accomplished by a common, nonspecific neuronal network.

Dissociable effects of dopaminergic therapy on spatial versus non-spatial working memory in Parkinson’s disease

There is now evidence for definite and early cognitive deficits in Parkinson’s disease (PD), involving, in particular, executive functions and working memory. However, the distinction between visuo-spatial and non-spatial working memory deficits and the impact of dopamine on these deficits are still open to debate. The aim of this study was therefore to investigate cognitive and motor performance in PD patients in two conditional associative learning tasks requiring either spatial or non-spatial visual working memory. The subject had to point to visual targets according to the visual characteristics of memorised visual cues (colour, position and form). To assess the effect of l-dopa therapy, PD patients were studied over two consecutive days: one ON/OFF group of nine PD patients with treatment (ON condition) on the first day and without treatment (OFF condition) on the second day; and another OFF/ON group of nine PD patients tested on reverse. The PD groups were compared to a control group of nine age-matched healthy subjects. Our main data demonstrate that: (1) in PD patients with OFF treatment, the response time of manual pointing is increased mainly in the non-spatial working memory task; and (2) in PD patients with ON treatment, either the response time is normal (on the first day) or is increased in both visuo-spatial and non-spatial tasks. We suggest that this dissociation between spatial versus non-spatial working memory deficits in non-medicated PD might be related to compensatory mechanisms that occur following fronto-striatal dysfunction.

Rat spatial memory tasks adapted for humans: characterization in subjects with intact brain and subjects with selective medial temporal lobe thermal lesions.

In the present paper we describe five tests, 3 of which were designed to be similar to tasks used with rodents. Results obtained from control subjects, patients with selective thermo-coagulation lesions to the medial temporal lobe and results from non-human primates and rodents are discussed. The tests involve memory for spatial locations acquired by moving around in a room, memory for objects subjects interacted with, or memory for objects and their locations. Two of the spatial memory tasks were designed specifically as analogs of the Morris water task and the 8-arm radial-maze tasks used with rats. The Morris water task was modeled by hiding a sensor under the carpet of a room (Invisible Sensor Task). Subjects had to learn its location by using an array of visual cues available in the room. A path integration task was developed in order to study the non-visual acquisition of a cognitive representation of the spatial location of objects. In the non-visual spatial memory task, we blindfolded subjects and led them to a room where they had to find 3 objects and remember their locations. We designed an object location task by placing 4 objects in a room that subjects observed for later recall of their locations. A recognition task, and a novelty detection task were given subsequent to the recall task. An 8-arm radial-maze was recreated by placing stands at equal distance from each other around the room, and asking subjects to visit each stand once, from a central point. A non-spatial working memory task was designed to be the non-spatial equivalent of the radial maze. Search paths recorded on the first trial of the Invisible Sensor Task, when subjects search for the target by trial and error are reported. An analysis of the search paths revealed that patients with lesions to the right or left hippocampus or parahippocampal cortex employed the same type of search strategies as normal controls did, showing similarities and differences to the search behavior recorded in rats. Interestingly, patients with lesions that included the right parahippocampal cortex were impaired relative to patients with lesions to the right hippocampus that spared the parahippocampal cortex, when recall of the sensor was tested after a 30 min delay (Bohbot et al. 1998). No differences were obtained between control subjects and patients with selective thermal lesions to the medial temporal lobe, when tested on the radial-maze, the non-spatial analogue to the radial-maze and the path integration tasks. Differences in methodological procedures, learning strategies and lesion location could account for some of the discrepant results between humans and non-human species. Patients with lesions to the right hippocampus, irrespective of whether the right parahippocampal cortex was spared or damaged, had difficulties remembering the particular configuration and identity of objects in the novelty detection of the object location task. This supports the role of the human right hippocampus for spatial memory, in this case, involving memory for the location of elements in the room; learning known to require the hippocampus in the rat.

Distinct patterns of brain oscillations underlie two basic parameters of human maze learning.

We examine how oscillations in the intracranial electroencephalogram (iEEG) relate to human maze learning. Theta- band activity (4-12 Hz in rodents; 4-8 Hz in humans) plays a significant role in memory function in rodents and in humans. Recording intracranially in humans, we have reported task-related, theta-band rhythmic activity in the raw trace during virtual maze learning and during a nonspatial working memory task. Here we analyze oscillations during virtual maze learning across a much broader range of frequencies and analyze their relationship to two task variables relevant to learning. We describe a new algorithm for detecting oscillatory episodes that takes advantage of the high signal-to-noise ratio and high temporal resolution of the iEEG. Accounting for the background power spectrum of the iEEG, the algorithm allows us to directly compare levels of oscillatory activity across frequencies within the 2- to 45-Hz band. We report that while episodes of oscillatory activity are found at various frequencies, most of the rhythmic activity during virtual maze learning occurs within the theta band. Theta oscillations are more prevalent when the task is made more difficult (manipulation of maze length). However, these oscillations do not tend to covary significantly with decision time, a good index of encoding and retrieval operations. In contrast, lower- and higher-frequency oscillations do covary with this variable. These results suggest that while human cortically recorded theta might play a role in encoding, the overall levels of theta oscillations tell us little about the immediate demands on encoding or retrieval. Finally, different patterns of oscillations may reflect distinct underlying aspects of memory function.