Abstract There is growing evidence for the relationship between working memory and academic attainment. The aim of the current study was to investigate whether working memory is simply a proxy for IQ or whether there is a unique contribution to learning outcomes. The findings indicate that children's working memory skills at 5 years of age were the best predictor of literacy and numeracy 6 years later. IQ, in contrast, accounted for a smaller portion of unique variance to these learning outcomes. The results demonstrate that working memory is not a proxy for IQ but rather represents a dissociable cognitive skill with unique links to academic attainment. Critically, we find that working memory at the start of formal education is a more powerful predictor of subsequent academic success than IQ. This result has important implications for education, particularly with respect to intervention. 2009 Elsevier Inc. All rights reserved.

Abstract Over a century ago, Freud proposed that unwanted memories can be excluded from awareness, a process called repression. It is unknown, however, how repression occurs in the brain. We used functional magnetic resonance imaging to identify the neural systems involved in keeping unwanted memories out of awareness. Controlling unwanted memories was associated with increased dorsolateral prefrontal activation, reduced hippocampal activation, and impaired retention of those memories. Both prefrontal cortical and right hippocampal activations predicted the magnitude of forgetting. These results confirm the existence of an active forgetting process and establish a neurobiological model for guiding inquiry into motivated forgetting.

Transcranial direct current stimulation (tDCS), applied to the left dorsolateral prefrontal cortex (DLPFC) has been found to improve working memory (WM) performance in both healthy and clinical participants. However, whether this effect can be enhanced by cognitive activity undertaken during tDCS has not yet been explored.ObjectiveThis study aimed to explore whether tDCS applied to the left DLPFC during the persistent performance of one WM task would improve performance on a subsequent WM task, to a greater extent than either tDCS or cognitive activity alone.MethodsTen healthy participants took part in three counterbalanced conditions. The conditions involved 10 minutes of either anodal tDCS while completing an n-back task, anodal tDCS while at rest, or sham tDCS while completing an n-back task. The n-back that was used in this study was a computer-based letter WM task that involved 5 minutes of two-back, followed by 5 minutes of three-back. Digit span forward and backward was administered immediately before and after each treatment, and performance change (pre- to posttreatment) calculated and compared across conditions. The digit span tasks involved a series of numbers being read to the participant, and the participant was required to repeat them back, either in the same order (Digits forward) or in the reverse order (Digits backward).ResultstDCS applied during completion of the n-back task was found to result in greater improvement in performance on digit span forward, compared with tDCS applied while at rest and sham tDCS during the n-back task. This finding was not evident with digit span backward.ConclusionsThese results indicate that there may be potential for the use of adjunctive cognitive remediation techniques to enhance the effects of tDCS. However, further research needs to be undertaken in this area to replicate and extend this finding.

Cognitive performance, including performance on working memory (WM) tasks declines with age. Changes in brain activations are one presumed contributor to WM decline in the healthy aging population. In particular, neuroimaging studies show that when older adults perform WM tasks there tends to be greater bilateral frontal activity than in younger adults. We hypothesized that stimulating the prefrontal cortex in healthy older adults would improve WM performance. To test this hypothesis we employed transcranial direct current stimulation (tDCS), a neurostimulation technique in which small amounts of electrical current are applied to the scalp with the intent of modulating the activity in underlying neurons. Across three testing sessions we applied sham stimulation or anodal tDCS to the left (F3) or right (F4) prefrontal cortex to healthy older adults as they performed trials of verbal and visual 2-back WM tasks. Surprisingly, tDCS was uniformly beneficial across site and WM task, but only in older adults with more education. In the less educated group, tDCS provided no benefit to verbal or visual WM performance. We interpret these findings as evidence for differential frontal recruitment as a function of strategy when older adults perform WM tasks.

This article reviews studies demonstrating enhancement with transcranial direct current stimulation (tDCS) of attention, learning, and memory processes in healthy adults. Given that these are fundamental cognitive functions, they may also mediate stimulation effects on other higher-order processes such as decision-making and problem solving. Although tDCS research is still young, there have been a variety of methods used and cognitive processes tested. While these different methods have resulted in seemingly contradictory results among studies, many consistent and noteworthy effects of tDCS on attention, learning, and memory have been reported. The literature suggests that although tDCS as typically applied may not be as useful for localization of function in the brain as some other methods of brain stimulation, tDCS may be particularly well-suited for practical applications involving the enhancement of attention, learning, and memory, in both healthy subjects and in clinical populations.

Abstract Working memory is the process of maintaining an active representation of information so that it is available for use. In monkeys, a prefrontal cortical region important for spatial working memory lies in and around the principal sulcus, but in humans the location, and even the existence, of a region for spatial working memory is in dispute. By using functional magnetic resonance imaging in humans, an area in the superior frontal sulcus was identified that is specialized for spatial working memory. This area is located more superiorly and posteriorly in the human than in the monkey brain, which may explain why it was not recognized previously.

Commentators expressed a wide variety of views on whether there is a basic capacity limit of 3 to 5 chunks and, among those who believe in it, about why it occurs. In this response, I conclude that the capacity limit is real and that the concept is strengthened by additional evidence offered by a number of commentators. I consider various arguments why the limit occurs and try to organize these arguments into a conceptual framework or of storage capacity limits meant to be useful in future research to settle the issue. I suggest that principles of memory representation determine what parts of the representation will be most prominent but that limits of attention (or of a memory store that includes only items that have been most recently attended) determine the 3- to 5-chunk capacity limit.

Abstract Interactions between prefrontal cortex (PFC) and stimulus-specific visual cortical association areas are hypothesized to mediate visual working memory in behaving monkeys. To clarify the roles for homologous regions in humans, event-related fMRI was used to assess neural activity in PFC and fusiform face area (FFA) of subjects performing a delay-recognition task for faces. In both PFC and FFA, activity increased parametrically with memory load during encoding and maintenance of face stimuli, despite quantitative differences in the magnitude of activation. Moreover, timing differences in PFC and FFA activation during memory encoding and retrieval implied a context dependence in the flow of neural information. These results support existing neurophysiological models of visual working memory developed in the nonhuman primate.

Previous research has demonstrated that the maintenance of visual information in working memory is associated with a sustained posterior contralateral negativity. Here we show that this component is also elicited during the spatially selective access to visual working memory. Participants memorized a bilateral visual search array that contained two potential targets on the left and right side. The task-relevant side was signalled by post-cues that were presented either 150 ms after array offset or after a longer interval (700 1000 ms). Enhanced negativities at posterior electrodes contralateral to the cued side of a target were elicited in response to both early and late post-cues, suggesting that they reflect not only memory maintenance, but also processes involved in the access to stored visual working memory representations. Results provide new electrophysiological evidence for the retinotopic organization of visual working memory.

Abstract Previous studies have claimed that weak transcranial direct current stimulation (tDCS) induces persisting excitability changes in the human motor cortex that can be more pronounced than cortical modulation induced by transcranial magnetic stimulation, but there are no studies that have evaluated the effects of tDCS on working memory. Our aim was to determine whether anodal transcranial direct current stimulation, which enhances brain cortical excitability and activity, would modify performance in a sequential-letter working memory task when administered to the dorsolateral prefrontal cortex (DLPFC). Fifteen subjects underwent a three-back working memory task based on letters. This task was performed during sham and anodal stimulation applied over the left DLPFC. Moreover seven of these subjects performed the same task, but with inverse polarity (cathodal stimulation of the left DLPFC) and anodal stimulation of the primary motor cortex (M1). Our results indicate that only anodal stimulation of the left prefrontal cortex, but not cathodal stimulation of left DLPFC or anodal stimulation of M1, increases the accuracy of the task performance when compared to sham stimulation of the same area. This accuracy enhancement during active stimulation cannot be accounted for by slowed responses, as response times were not changed by stimulation. Our results indicate that left prefrontal anodal stimulation leads to an enhancement of working memory performance. Furthermore, this effect depends on the stimulation polarity and is specific to the site of stimulation. This result may be helpful to develop future interventions aiming at clinical benefits.

A key motivation for understanding capacity in working memory (WM) is its relationship with fluid intelligence. Recent evidence has suggested a two-factor model that distinguishes between the number of representations that can be maintained in WM and the resolution of those representations. To determine how these factors relate to fluid intelligence, we conducted an exploratory factor analysis on multiple number-limited and resolution-limited measures of WM ability. The results strongly supported the two-factor model, with fully orthogonal factors accounting for performance in the number-limited and resolution-limited conditions. Furthermore, the reliable relationship between WM capacity and fluid intelligence was exclusively supported by the number factor ( r =.66), whereas the resolution factor made no reliable contribution ( r = .05). Thus, the relationship between WM capacity and standard measures of fluid intelligence is mediated by the number of representations that can be simultaneously maintained in WM, rather than by the precision of those representations.

Several studies have shown that transcranial direct current stimulation (tDCS) is able to enhance performances on verbal and visual working memory (WM) tasks. Available evidence points to the right dorsolateral prefrontal cortex (DLPFC) as a critical area in visual WM, but to date direct comparisons of the effects obtained by stimulating the left versus the right DLPFC in the same subject are lacking. Our aim was to determine whether tDCS over the right DLPFC can differently affect performance as compared with left DLPFC stimulation. Ten healthy subjects performed a memory-guided visuospatial task in three conditions: baseline, during anodal stimulation applied over the right and during anodal stimulation applied over the left DLPFC. All the subjects also underwent a sham stimulation as control. Our results show that only active stimulation over the right DLPFC is able to increase performance when compared to the other conditions. Our findings confirm the crucial role played by the right DLPFC in spatial WM tasks.

Single-unit recordings in macaque extrastriate cortex have shown that attentional selection of nonspatial features can operate in a location-independent manner. Here, we investigated analogous neural correlates at the neural population level in human observers by using simultaneous event-related potential (ERP) and event-related magnetic field (ERMF) recordings. The goals were to determine (1) whether task-relevant features are selected before attention is allocated to the location of the target, and (2) whether this selection reflects the locations of the relevant features. A visual search task was used in which the spatial distribution of nontarget items with attended feature values was varied independently of the location of the target. The presence of task-relevant features in a given location led to a change in ERP/ERMF activity beginning approximately 140 msec after stimulus onset, with a neural origin in the ventral occipito-temporal cortex. This effect was independent of the location of the actual target. This effect was followed by lateralized activity reflecting the allocation of attention to the location of the target (the well known N2pc component), which began at approximately 170 msec poststimulus. Current source localization indicated that the allocation of attention to the location of the target originated in more anterior regions of occipito-temporal cortex anterior than the feature-related effects. These findings suggest that target detection in visual search begins with the detection of task-relevant features, which then allows spatial attention to be allocated to the location of a likely target, which in turn allows the target to be positively identified.

In vivo effects of transcranial direct current stimulation (tDCS) have attracted much attention nowadays as this area of research spreads to both the motor and cognitive domains. The common assumption is that the anode electrode causes an enhancement of cortical excitability during stimulation, which then lasts for a few minutes thereafter, while the cathode electrode generates the opposite effect, i.e., anodal-excitation and cathodal-inhibition effects (AeCi). Yet, this dual-polarity effect has not been observed in all tDCS studies. Here, we conducted a meta-analytical review aimed to investigate the homogeneity/heterogeneity of the effect sizes of the AeCi dichotomy in both motor and cognitive functions. The AeCi effect was found to occur quite commonly with motor investigations and rarely in cognitive studies. When the anode electrode is applied over a non-motor area, in most cases, it will cause an excitation as measured by a relevant cognitive or perceptual task; however, the cathode electrode rarely causes an inhibition. We found homogeneity in motor studies and heterogeneity in cognitive studies with the electrode polarity serving as a moderator that can explain the source of heterogeneity in cognitive studies. The lack of inhibitory cathodal effects might reflect compensation processes as cognitive functions are typically supported by rich brain networks. Further insights as to the polarity and domain interaction are offered, including subdivision to different classes of cognitive functions according to their likelihood of being affected by stimulation.

We used electrophysiological methods to track the deployment of visual spatial attention while observers were engaged in concurrent central attentional processing, using a variant of the attentional blink paradigm. Two visual targets (T 1 , T 2 ) were presented at a stimulus onset asynchrony of either 200 ms or 800 ms. T 1 was a white digit among white letters presented on a dark background using rapid serial visual presentation at fixation. T 2 was another digit that was presented to the left or right of fixation simultaneously with a distractor digit in the opposite visual field, each followed by a pattern mask. In each T 2 display, one digit was red and one was green. Half of the subjects reported the red digit and ignored the green one, whereas the other half reported the green digit and ignored the red one. T 1 and T 2 were reported in one block of trials, and only T 2 in another block (order counterbalanced across subjects). Accuracy of report of T 2 was lower at short SOA than at long SOA when both T 1 and T 2 were reported, but was similar across SOA when only T 2 was reported. The electrophysiological results focused on the N2pc component, which was used as an index of the locus of spatial attention. N2pc was reduced in amplitude when subjects reported T 1 , and particularly so at the short SOA. The results suggest that attention to T 1 interfered with the deployment of visual spatial attention to T 2 .

The nature of parietal contributions to working memory (WM) remain poorly understood but of considerable interest. We previously reported that posterior parietal damage selectively impaired WM probed by recognition (Berryhill and Olson, 2008a). Recent studies provided support using a neuromodulatory technique, transcranial direct current stimulation (tDCS) applied to the right parietal cortex (P4). These studies confirmed parietal involvement in WM because parietal tDCS altered WM performance: anodal current tDCS improved performance in a change detection task, and cathodal current tDCS impaired performance on a sequential presentation task. Here, we tested whether these complementary results were due to different degrees of parietal involvement as a function of WM task demands, WM task difficulty, and/or participants WM capacity. In Experiment 1, we applied cathodal and anodal tDCS to the right parietal cortex and tested participants on both previously used WM tasks. We observed an interaction between tDCS (anodal, cathodal), WM task difficulty, and participants WM capacity. When the WM task was difficult, parietal stimulation (anodal or cathodal) improved WM performance selectively in participants with high WM capacity. In the low WM capacity group, parietal stimulation (anodal or cathodal) impaired WM performance. These nearly equal and opposite effects were only observed when the WM task was challenging, as in the change detection task. Experiment 2 probed the interplay of WM task difficulty and WM capacity in a parametric manner by varying set size in the WM change detection task. Here, the effect of parietal stimulation (anodal or cathodal) on the high WM capacity group followed a linear function as WM task difficulty increased with set size. The low WM capacity participants were largely unaffected by tDCS. These findings provide evidence that parietal involvement in WM performance depends on both WM capacity and WM task demands. We discuss these findings in terms of alternative WM strategies employed by low and high WM capacity individuals. We speculate that low WM capacity individuals do not recruit the posterior parietal lobe for WM tasks as efficiently as high WM capacity individuals. Consequently, tDCS provides greater benefit to individuals with high WM capacity.

Event-related potentials (ERPs) were recorded as 12 subjects performed a delayed matching to sample task. We presented two bilateral abstract shapes and cued spatially which had to be memorized for a subsequent matching task: left, right or both. During memorization a posterior slow negative ERP wave developed over the hemisphere contralateral to the memorized shape. This effect was similar in high and low memory load trials while the memory figures were visible (for 1000 ms). As the figures disappeared (for 1500 ms), the effect persisted only in the low memory load conditions. We suggest that the contralateral negativity reflects a visual short-term memory process and that capacity limitation in the high memory load condition causes this process to collapse.

J Neurophysiol. 1971 May;34(3):337-47.

61Attention control and attention scope during visual WM were examined by tDCS.61Anodal tDCS on the right PPC specifically enlarged attention scope in visual WM.61Anodal tDCS on the right PFC specifically enhanced attention control in visual WM.61Bilateral presentation introduced an additional demand on attention control.

The identification of targets in visual search arrays may be improved by suppressing competing information from the surrounding distractor items. The present study provided evidence that this hypothetical filtering process has a neural correlate, the "N2pc" component of the event-related potential waveform. The N2pc was observed when a target item was surrounded by competing distractor items but was absent when the array could be rejected as a nontarget on the basis of simple feature information. In addition, the N2pc was eliminated when filtering was discouraged by removing the distractor items, making the distractors relevant, or making all items within an array identical. Combined with previous topographic analyses, these results suggest that attentional filtering occurs in occipital cortex under the control of feedback from higher cortical regions after a preliminary feature-based analysis of the stimulus array.

Event-related brain potentials (ERPs) were recorded from normal young adults during visual search tasks in which the stimulus arrays contained either eight identical items (homogeneous arrays) or seven identical items and one deviant item (pop-out arrays). Four experiments were conducted in which different classes of stimulus arrays were designated targets and the remaining stimulus arrays were designated nontargets. In Experiments 1 and 2, both target and nontarget pop-out stimuli elicited an enhanced anterior N2 wave and a contralaterally larger posterior P1 wave, but Experiments 3 and 4 demonstrated that these components do not reflect fully automatic pop-out detection processes. In all four experiments, target pop-outs elicited enlarged anterior P2, posterior N2, occipital P3, and parietal P3 waves. The target-elicited posterior N2 wave contained a contralateral subcomponent (N2pc) that exhibited a focus over occipital cortex in maps of current source density. The overall pattern of results was consistent with guided search models in which preattentive stimulus information is used to guide attention to task-relevant stimuli.

Abstract We used lateralized Event-Related Potential (ERP) measures - the N2pc and CDA/SPCN components - to assess the role of grouping by target similarity during enumeration. Participants saw a variable number (0, 1, 2 or 3) of same- or differently-colored targets presented among homogeneous distracters, and performed an enumeration task. Results showed that the N2pc, but not the CDA, was larger for multiple targets of identical color relative to targets of different colors. The findings are interpreted in terms of the effects of grouping on early versus late stages of multiple object processing. Within this framework, they reveal that grouping has an effect on early individuation mechanisms, while later processing mechanisms are less prone to such an influence.

Background: Recent studies revealed that anodal transcranial direct current stimulation (tDCS) to the left dorsolateral prefrontal cortex (DLPFC) may improve verbal working memory (WM) performance in humans. In the present study, we evaluated executive attention, which is the core of WM capacity, considered to be significantly involved in tasks that require active maintenance of memory representations in interference-rich conditions, and is highly dependent on DLPFC function.Objectives: We investigated verbal WM accuracy using a WM task that is highly sensitive to executive attention function. We were interested in how verbal WM accuracy may be affected by WM load, unilateral DLPFC stimulation, and gender, as previous studies showed gender-dependent brain activation during verbal WM tasks.Methods: We utilized a modified verbal n-Back task hypothesized to increase demands on executive attention. We examined "online" WM performance while participants received transcranial direct current stimulation (tDCS), and implicit learning performance in a post-stimulation WM task.Results: Significant lateralized "online" stimulation effects were found only in the highest WM load condition revealing that males benefit from left DLPFC stimulation, while females benefit from right DLPFC stimulation. High WM load performance in the left DLPFC stimulation was significantly related to post-stimulation recall performance.Conclusions: Our findings support the idea that lateralized stimulation effects in high verbal WM load may be gender-dependent. Further, our post-stimulation results support the idea that increased left hemisphere activity may be important for encoding verbal information into episodic memory as well as for facilitating retrieval of context-specific targets from semantic memory. (C) 2013 Elsevier Inc. All rights reserved.

Abstract 1 In this paper we demonstrate in the intact human the possibility of a non-invasive modulation of motor cortex excitability by the application of weak direct current through the scalp. 2 Excitability changes of up to 40 %, revealed by transcranial magnetic stimulation, were accomplished and lasted for several minutes after the end of current stimulation. 3 Excitation could be achieved selectively by anodal stimulation, and inhibition by cathodal stimulation. 4 By varying the current intensity and duration, the strength and duration of the after-effects could be controlled. 5 The effects were probably induced by modification of membrane polarisation. Functional alterations related to post-tetanic potentiation, short-term potentiation and processes similar to postexcitatory central inhibition are the likely candidates for the excitability changes after the end of stimulation. Transcranial electrical stimulation using weak current may thus be a promising tool to modulate cerebral excitability in a non-invasive, painless, reversible, selective and focal way.

Abstract Transcranial direct current stimulation (tDCS) of the human motor cortex results in polarity-specific shifts of cortical excitability during and after stimulation. Anodal tDCS enhances and cathodal stimulation reduces excitability. Animal experiments have demonstrated that the effect of anodal tDCS is caused by neuronal depolarisation, while cathodal tDCS hyperpolarises cortical neurones. However, not much is known about the ion channels and receptors involved in these effects. Thus, the impact of the sodium channel blocker carbamazepine, the calcium channel blocker flunarizine and the NMDA receptor antagonist dextromethorphane on tDCS-elicited motor cortical excitability changes of healthy human subjects were tested. tDCS-protocols inducing excitability alterations (1) only during tDCS and (2) eliciting long-lasting after-effects were applied after drug administration. Carbamazepine selectively eliminated the excitability enhancement induced by anodal stimulation during and after tDCS. Flunarizine resulted in similar changes. Antagonising NMDA receptors did not alter current-generated excitability changes during a short stimulation, which elicits no after-effects, but prevented the induction of long-lasting after-effects independent of their direction. These results suggest that, like in other animals, cortical excitability shifts induced during tDCS in humans also depend on membrane polarisation, thus modulating the conductance of sodium and calcium channels. Moreover, they suggest that the after-effects may be NMDA receptor dependent. Since NMDA receptors are involved in neuroplastic changes, the results suggest a possible application of tDCS in the modulation or induction of these processes in a clinical setting. The selective elimination of tDCS-driven excitability enhancements by carbamazepine proposes a role for this drug in focussing the effects of cathodal tDCS, which may have important future clinical applications.

Clin Neurophysiol. 2003 Nov;114(11):2220-2; author reply 2222-3. Letter

The time-dependent effect of transcranial direct current stimulation (tDCS) on working memory was investigated by applying anodal stimulation over the left prefrontal cortex. This single-blind, sham-controlled crossover study recruited 15 healthy participants. A three-back verbal working-memory task was performed before, during, and 30 min after 1 mA anodal or sham tDCS. Anodal tDCS, compared with sham stimulation, significantly improved working-memory performance. Accuracy of response was significantly increased after 20 min of tDCS application, and was further enhanced after 30 min of stimulation. This effect was maintained for 30 min after the completion of stimulation. These results suggest that tDCS at 1 mA enhances working memory in a time-dependent manner for at least 30 min in healthy participants.

Abstract Direct current (DC) is very effective in modulating spontaneous neuronal firing. The history of electrophysiology starts with the discovery of the biological effects of DC and as early as two centuries ago scalp DC was used to treat mental disorder. Psychophysiological investigations suggested a possible effect of scalp DC in humans. More recently several studies assessed, with motor potentials evoked by transcranial brain stimulation, the motor-cortical excitability changes induced by scalp DC. Even weak DCs pass through the scalp and influence human brain activity. DCs delivered at relatively strong intensities (1 mA) and for long periods (10 min or so), not only influence (either increase or decrease) brain excitability during their application in normal subjects, but induce persistent changes in excitability after their offset that, at least in the motor cortex, can last for almost 1 h. Scalp DC might represent a non-invasive simple and valuable potential treatment for psychiatric and neurologic diseases with changes in brain excitability or focally abnormal (increased or decreased) function.

Previous functional neuroimaging studies have shown that maintenance of centrally presented objects in visual short-term memory (VSTM) leads to bilateral increases of BOLD activations in IPS/IOS cortex, while prior electrophysiological work suggests that maintaining stimuli encoded from a single hemifield leads to a sustained posterior contralateral negativity (SPCN) in electrophysiology and magnetoencephalography. These two findings have never been investigated using the same physiological measures. We recorded the BOLD response using fMRI, magnetoencephalography (MEG), and electrophysiology (EEG), while subjects encoded visual stimuli from a single hemifield of a balanced display. The EEG showed an SPCN. However, no SPCN-like activation was observed in the BOLD signals. The BOLD response in parietal cortex remained bilateral, even after unilateral encoding of the stimuli, but MEG showed both bilateral and contralateral activations, each likely reflecting a sub portion of the neuronal populations participating in the maintenance of information in VSTM. Contrary to the assumption that BOLD, EEG, and MEG responses that were each linked to the maintenance of information in VSTM are markers of the same neuronal processes, our findings suggest that each technique reveals a somewhat distinct but overlapping neural signature of the mechanisms supporting visual short-term memory.

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.

Abstract Three experiments used position emission tomography (PET) to study the neural basis of human working memory. These studies ask whether different neural circuits underly verbal and spatial memory. In Experiment 1, subjects had to retain for 3 sec. either the names of four letters (verbal memory) or the positions of three dots (spatial memory). The PET results manifested a clear cut double dissociation, as the verbal task activated primarily left-hemisphere regions whereas the spatial task activated only right-hemisphere regions. In Experiment 2, the identical sequence of letters was presented in all conditions, and what varied was whether subjects had to remember the names of the letters (verbal memory) or their positions in the display (spatial memory). In the verbal task, activation was concentrated more in the left than the right hemisphere; in the spatial task, there was substantial activation in both hemispheres, though in key regions, there was more activation in the right than the left hemisphere. Experiment 3 studied only verbal memory, and showed that a continuous memory task activated the same regions as the discrete verbal task used in Experiment 1. Taken together, these results indicate that verbal and spatial working memory are implemented by different neural structures.

Executive working memory operations are related to prefrontal regions in the healthy brain. Moreover, neuroimaging data provide evidence for a functional dissociation of ventrolateral and dorsolateral prefrontal cortex. Most authors either suggest a modality-specific or a function-specific prefrontal cortex organization. In the present study we particularly aimed at the identification of different prefrontal cerebral areas that are involved in executive inhibitory processes during spatial working memory encoding. In an fMRI study (functional magnetic resonance imaging) we examined the neural correlates of spatial working memory processing by varying the amount of executive demands of the task. Twenty healthy volunteers performed the Corsi Block-Tapping test (CBT) during fMRI. The CBT requires the storage and reproduction of spatial target sequences. In a second condition, we presented an adapted version of the Block-Suppression-Test (BST). The BST is based on the original CBT but additionally requires the active suppression of visual distraction within the target sequences. In comparison to the CBT performance, particularly the left dorsolateral prefrontal cortex (BA 9) showed more activity during the BST condition. Our results show that the left dorsolateral prefrontal cortex plays a crucial role for executive controlled inhibition of spatial distraction. Furthermore, our findings are in line with the processing model of a functional dorsolateral-ventrolateral prefrontal cortex organization.

The limits of visual (VSTM) have been well documented, and recent neuroscientific studies suggest that VSTM performance is associated with activity in the posterior parietal cortex. Here we show that artificially elevating parietal activity via positively charged electric current through the skull can rapidly and effortlessly improve people's VSTM performance. This artificial improvement, however, comes with an interesting twist: it interacts with people's natural VSTM capability such that low performers who tend to remember less information benefitted from the stimulation, whereas high performers did not. This behavioral dichotomy is explained by event-related potentials around the parietal regions: low performers showed increased waveforms in N2pc and contralateral delay activity (), which implies improvement in attention deployment and access in the current paradigm, respectively. Interestingly, these components are found during the presentation of the test array instead of the retention interval, from the parietal sites ipsilateral to the target location, thus suggesting that transcranial direct current stimulation (tDCS) was mainly improving one's ability to suppress no-change distractors located on the irrelevant side of the display during the comparison stage. The high performers, however, did not benefit from tDCS as they showed equally large waveforms in N2pc and , or SPCN (sustained parietal contralateral negativity), before and after the stimulation such that electrical stimulation could not help any further, which also accurately accounts for our behavioral observations. Together, these results suggest that there is indeed a fixed upper limit in VSTM, but the low performers can benefit from neurostimulation to reach that maximum via enhanced comparison processes, and such behavioral improvement can be directly quantified and visualized by the magnitude of its associated electrophysiological waveforms.

Abstract Neuroscience research has identified the involvement of the dorsolateral prefrontal cortex (DLPFC) in cognitive control. Questions remain, however, about its lateralization correlates during Stroop task performance, an experimental cornerstone on which a large amount of cognitive control research is based. After reviewing the literature, we find that three Stroop variants have been used in an attempt to uncover different aspects of cognitive control related to DLPFC involvement. In sum, rapid and sequential up-regulation of the attentional set seems to be related to the left DLPFC. These attentional adjustments are based on participants' expectancies regarding the conflicting nature of the upcoming trial, and not on the conflict itself. In contrast, the right DLPFC is associated with an overall up-regulation of the attentional set when attentional conflict is experienced.

The capacity of visual short-term memory is highly limited, maintaining only three to four objects simultaneously. This extreme limitation necessitates efficient mechanisms to select only the most relevant objects from the immediate environment to be represented in memory and to restrict irrelevant items from consuming capacity. Here we report a neurophysiological measure of this memory selection mechanism in humans that gauges an individual's efficiency at excluding irrelevant items from being stored in memory. By examining the moment-by-moment contents of visual memory, we observe that selection efficiency varies substantially across individuals and is strongly predicted by the particular memory capacity of each person. Specifically, high capacity individuals are much more efficient at representing only the relevant items than are low capacity individuals, who inefficiently encode and maintain information about the irrelevant items present in the display. These results provide evidence that under many circumstances low capacity individuals may actually store more information in memory than high capacity individuals. Indeed, this ancillary allocation of memory capacity to irrelevant objects may be a primary source of putative differences in overall storage capacity.

Neuroimaging studies in humans have shown that different working memory (WM) tasks recruit a common bilateral fronto-parietal cortical network. Animal studies as well as neuroimaging studies in humans have suggested that this network, in particular the prefrontal cortex, is preferentially recruited when material from different domains (e.g. spatial information or verbal/object information) has to be memorized. Early imaging studies have suggested qualitative dissociations in the prefrontal cortex for spatial and object/verbal WM, either in a left-right or a ventral-dorsal dimension. However, results from different studies are inconsistent. Moreover, recent fMRI studies have failed to find evidence for domain dependent dissociations of WM-related activity in prefrontal cortex. Here we present evidence from two independent fMRI studies using physically identical stimuli in a verbal and spatial WM task showing that domain dominance for WM does indeed exist, although only in the form of quantitative differences in activation and not in the form of a dissociation with different prefrontal regions showing mutually exclusive activation in different domains. Our results support a mixed dimension model of domain dominance for WM within the prefrontal cortex, with left ventral prefrontal cortex (PFC) supporting preferentially verbal WM and right dorsal PFC supporting preferentially spatial WM. The concept of domain dominance is discussed in the light of recent theories of prefrontal cortex function.

Because of the precise temporal resolution of electrophysiological recordings, the event-related potential (ERP) technique has proven particularly valuable for testing theories of perception and attention. Here, I provide a brief tutorial on the ERP technique for consumers of such research and those considering the use of human electrophysiology in their own work. My discussion begins with the basics regarding what brain activity ERPs measure and why they are well suited to reveal critical aspects of perceptual processing, attentional selection, and cognition, which are unobservable with behavioral methods alone. I then review a number of important methodological issues and often-forgotten facts that should be considered when evaluating or planning ERP experiments.

Symbolic visual cues indicating the location of an upcoming target are believed to invoke endogenous shifts of attention to cued locations. In the present study, we investigated how visual attention is shifted during such cuing paradigms by recording event-related potentials (ERPs). We focused on a component known to index lateralized shifts of perceptual attention during visual search tasks, known as the N2pc component. The ERP data show that attention was shifted to a cued location in anticipation of a target shape when the location is marked by a placeholder object (Experiments 1 and 2). However, when the possible locations were not marked by placeholder objects, we found no evidence for an anticipatory shift of attention to the cued location (Experiment 3). These findings indicate that the perceptual attention mechanism indexed by the N2pc is deployed to objects and not simply locations in space devoid of object structure.

This study examined whether objects are attended in serial or in parallel during a demanding visual search task. A component of the event-related potential waveform, the N2pc wave, was used as a continuous measure of the allocation of attention to possible targets in the search arrays. Experiment 1 demonstrated that the relative allocation of attention shifts rapidly, favoring one item and then another. In Experiment 2, a paradigm was used that made it possible to track the absolute allocation of attention to individual items. This experiment showed that attention was allocated to one object for 100-150 ms before attention began to be allocated to the next object. These findings support models of attention that posit serial processing in demanding visual search tasks.

Spatial working memory (SWM) is the ability to temporarily store and manipulate spatial information. It has a limited capacity and is quite vulnerable to interference. Dorsolateral prefrontal cortex (DLPFC) has been shown to be a part of the SWM network but its specific functional role still remains unknown. Here we applied transcranial direct current stimulation (tDCS), a noninvasive brain stimulation technique that provides polarity-specific stimulation over the targeted region, to investigate the specific role of the right DLPFC in resolving interference in SWM. A forward- and backward-recall computerized Corsi Block Tapping task (CBT), both with and without a concurrent motor interference task (the modified Luria manual sequencing task) was used to measure SWM capacity and reaction time. The results showed that motor interference impeded accuracy and prolonged reaction time in forward and backward recall for SWM. Anodal tDCS over right DLPFC yielded the tendency to shorten participants’ reaction time in the conditions with interference (forward with interference, and backward with interference). Most importantly, anodal tDCS significantly improved participants’ SWM span when cognitive demand was the highest (the “backward-recall with motor interference” condition). These results suggest that (1) the right DLPFC plays a crucial role in dealing with the cross-domain motor interference for spatial working memory and (2) the anodal tDCS over right DLPFC improved SWM capacity particularly when task difficulty demands more complex mental manipulations that could be due to the facilitatory effect of anodal tDCS which enhanced the DLPFC function within central executive system at the top-down attentional level.