Abstract: Representation of the visual and spatial properties of our environment is a pivotal requirement of everyday cognition. We can mentally represent the visual form of objects. We can extract information from several of the senses as to the location of objects in relation to ourselves and to other objects nearby. For some of those objects we can reach out and manipulate them. We can also imagine ourselves manipulating objects in advance of doing so, or even when it would be impossible to do so physically. The problem posed to science is how these cognitive operations are accomplished, and proffered accounts lie in two essentially parallel research endeavours, working memory and imagery. Working memory is thought to pervade everyday cognition, to provide on-line processing and temporary storage, and to update, moment to moment, our representation of the current state of our environment and our interactions with that environment. There is now a strong case for the claims of working memory in the area of phonological and articulatory functions, all of which appear to contribute to everyday activities such as counting, arithmetic, vocabulary acquisition, and some aspects of reading and language comprehension. The claims for visual and spatial working memory functions are less convincing. Most notable has been the assumption that visual and spatial working memory are intimately involved in the generation, retention and manipulations of visual images. There has until recently been little hard evidence to justify that assumption, and the research on visual and spatial working memory has focused on a relatively restricted range of imagery tasks and phenomena. In a more or less independent development, the literature on visual imagery has now amassed a voluminous corpus of data and theory about a wide range of imagery phenomena. Despite this, few books on imagery refer to the concept of working memory in any detail, or specify the nature of the working memory system that might be involved in mental imagery. This essay follows a line of reconciliation and positive critiquing in exploring the possible overlap between mental imagery and working memory. Theoretical development in the book draws on data from both cognitive psychology and cognitive neuropsychology. The aim is to stimulate debate, to address directly a number of assumptions that hitherto have been implicit, and to assess the contribution of the concept of working memory to our understanding of these intriguing core aspects of human cognition.

Abstract: Working memory is the retention of a small amount of information in a readily accessible form. It facilitates planning, comprehension, reasoning, and problem-solving. I examine the historical roots and conceptual development of the concept and the theoretical and practical implications of current debates about working memory mechanisms. Then I explore the nature of cognitive developmental improvements in working memory, the role of working memory in learning, and some potential implications of working memory and its development for the education of children and adults. The use of working memory is quite ubiquitous in human thought, but the best way to improve education using what we know about working memory is still controversial. I hope to provide some directions for research and educational practice.

Abstract: Experimental evidence suggests that the maintenance of an item in working memory is achieved through persistent activity in selective neural assemblies of the cortex. To understand the mechanisms underlying this phenomenon, it is essential to investigate how persistent activity is affected by external inputs or neuromodulation. We have addressed these questions using a recurrent network model of object working memory. Recurrence is dominated by inhibition, although persistent activity is generated through recurrent excitation in small subsets of excitatory neurons. Our main findings are as follows. (1) Because of the strong feedback inhibition, persistent activity shows an inverted U shape as a function of increased external drive to the network. (2) A transient external excitation can switch off a network from a selective persistent state to its spontaneous state. (3) The maintenance of the sample stimulus in working memory is not affected by intervening stimuli (distractors) during the delay period, provided the stimulation intensity is not large. On the other hand, if stimulation intensity is large enough, distractors disrupt sample-related persistent activity, and the network is able to maintain a memory only of the last shown stimulus. (4) A concerted modulation of GABA A and NMDA conductances leads to a decrease of spontaneous activity but an increase of persistent activity; the enhanced signal-to-noise ratio is shown to increase the resistance of the network to distractors. (5) Two mechanisms are identified that produce an inverted U shaped dependence of persistent activity on modulation. The present study therefore points to several mechanisms that enhance the signal-to-noise ratio in working memory states. These mechanisms could be implemented in the prefrontal cortex by dopaminergic projections from the midbrain.

Abstract: Maneuvering safely through the environment is central to survival of almost all species. The ability to do this depends on learning and remembering locations. This capacity is encoded in the brain by two systems: one using cues outside the organism (distal cues), allocentric navigation, and one using self-movement, internal cues and nearby proximal cues, egocentric navigation. Allocentric navigation involves the hippocampus, entorhinal cortex, and surrounding structures; in humans this system encodes allocentric, semantic, and episodic memory. This form of memory is assessed in laboratory animals in many ways, but the dominant form of assessment is the Morris water maze (MWM). Egocentric navigation involves the dorsal striatum and connected structures; in humans this system encodes routes and integrated paths and, when overlearned, becomes procedural memory. In this article, several allocentric assessment methods for rodents are reviewed and compared with the MWM. MWM advantages (little training required, no food deprivation, ease of testing, rapid and reliable learning, insensitivity to differences in body weight and appetite, absence of nonperformers, control methods for proximal cue learning, and performance effects) and disadvantages (concern about stress, perhaps not as sensitive for working memory) are discussed. Evidence-based design improvements and testing methods are reviewed for both rats and mice. Experimental factors that apply generally to spatial navigation and to MWM specifically are considered. It is concluded that, on balance, the MWM has more advantages than disadvantages and compares favorably with other allocentric navigation tasks.

Abstract: This meta-analysis examined the effects of process-based executive-function and working memory training (49 articles, 61 independent samples) in older adults (> 60 years). The interventions resulted in significant effects on performance on the trained task and near-transfer tasks; significant results were obtained for the net pretest-to-posttest gain relative to active and passive control groups and for the net effect at posttest relative to active and passive control groups. Far-transfer effects were smaller than near-transfer effects but were significant for the net pretest-to-posttest gain relative to passive control groups and for the net gain at posttest relative to both active and passive control groups. We detected marginally significant differences in training-induced improvements between working memory and executive-function training, but no differences between the training-induced improvements observed in older adults and younger adults, between the benefits associated with adaptive and nonadaptive training, or between the effects in active and passive control conditions. Gains did not vary with total training time.

Abstract: Prefrontal persistent activity during the delay of spatial working memory tasks is thought to maintain spatial location in memory. A 'bump attractor' computational model can account for this physiology and its relationship to behavior. However, direct experimental evidence linking parameters of prefrontal firing to the memory report in individual trials is lacking, and, to date, no demonstration exists that bump attractor dynamics underlies spatial working memory. We analyzed monkey data and found model-derived predictive relationships between the variability of prefrontal activity in the delay and the fine details of recalled spatial location, as evident in trial-to-trial imprecise oculomotor responses. Our results support a diffusing bump representation for spatial working memory instantiated in persistent prefrontal activity. These findings reinforce persistent activity as a basis for spatial working memory, provide evidence for a continuous prefrontal representation of memorized space and offer experimental support for bump attractor dynamics mediating cognitive tasks in the cortex.

Abstract: A growing interest in cerebellar function and its involvement in higher cognition have prompted much research in recent years. Cerebellar presence in a wide range of cognitive functions examined within an increasing body of neuroimaging literature has been observed. We applied a meta-analytic approach, which employed the activation likelihood estimate method, to consolidate results of cerebellar involvement accumulated in different cognitive tasks of interest and systematically identified similarities among the studies. The current analysis included 88 neuroimaging studies demonstrating cerebellar activations in higher cognitive domains involving emotion, executive function, language, music, timing and working memory. While largely consistent with a prior meta-analysis by Stoodley and Schmahmann ([2009]: Neuroimage 44:489-501), our results extended their findings to include music and timing domains to provide further insights into cerebellar involvement and elucidate its role in higher cognition. In addition, we conducted inter- and intradomain comparisons for the cognitive domains of emotion, language, and working memory. We also considered task differences within the domain of verbal working memory by conducting a comparison of the Sternberg with the n-back task, as well as an analysis of the differential components within the Sternberg task. Results showed a consistent cerebellar presence in the timing domain, providing evidence for a role in time keeping. Unique clusters identified within the domain further refine the topographic organization of the cerebellum.

Abstract: Although working memory (WM) figures centrally in many theories of second language (L2) proficiency development and processing, some have argued that the importance of WM is overstated (e.g., Juffs, Transactions of the Philological Society, 102, 199–225, 2004). Despite many studies over the past two decades, the literature lacks a quantitative synthesis of the extant results. In this article, we report a meta-analysis of data from 79 samples involving 3,707 participants providing 748 effect sizes. The results indicate that WM is positively associated with both L2 processing and proficiency outcomes, with an estimated population effect size (ρ) of .255. In additional analyses, we assessed whether the WM–criterion relationship was modulated by potential covariates identified in the literature search (i.e., participant characteristics, WM measure features, criterion measure factors, and publication status). The results of the covariate analyses indicated larger effect sizes for the executive control (vs. storage) component of WM, and for verbal (vs. nonverbal) measures of WM. Minimal publication bias was detected, suggesting that WM has a robust, positive relationship with L2 outcomes. We discuss the implications of these results for models of WM and theories of L2 processing and L2 proficiency development.

Abstract: Video gaming is a highly pervasive activity, providing a multitude of complex cognitive and motor demands. Gaming can be seen as an intense training of several skills. Associated cerebral structural plasticity induced has not been investigated so far. Comparing a control with a video gaming training group that was trained for 2 months for at least 30 min per day with a platformer game, we found significant gray matter (GM) increase in right hippocampal formation (HC), right dorsolateral prefrontal cortex (DLPFC) and bilateral cerebellum in the training group. The HC increase correlated with changes from egocentric to allocentric navigation strategy. GM increases in HC and DLPFC correlated with participants' desire for video gaming, evidence suggesting a predictive role of desire in volume change. Video game training augments GM in brain areas crucial for spatial navigation, strategic planning, working memory and motor performance going along with evidence for behavioral changes of navigation strategy. The presented video game training could therefore be used to counteract known risk factors for mental disease such as smaller hippocampus and prefrontal cortex volume in, for example, post-traumatic stress disorder, schizophrenia and neurodegenerative disease.

Abstract: Executive function is a collection of cognitive processes essential for higher order mental function. Processes involved in executive function include, but are not limited to, working memory, attention, cognitive flexibility, and impulse control. These complex behaviors are largely mediated by prefrontal cortical function but are modulated by dopaminergic, noradrenergic, serotonergic, and cholinergic input. The ability of these neurotransmitter systems to modulate executive function allows for adaptation in cognitive behavior in response to changes in the environment. Because of the important role these neurotransmitter systems play in regulating executive function, changes in these systems can also have a grave impact on executive function. In addition, polymorphisms in genes associated with these neurotransmitters are associated with phenotypic differences in executive function. Understanding how these naturally occurring polymorphisms contribute to different executive function phenotypes will advance basic knowledge of cognition and potentially further understanding and treatment of mental illness that involve changes in executive function. In this review, we will examine the influence of dopamine, norepinephrine, serotonin, and acetylcholine on the following measures of executive function: attention, cognitive flexibility, and impulse control. We will also review the effects of polymorphisms in genes associated with these neurotransmitter systems on these measures of executive function.

Abstract: The aim of this review is to illustrate the role of working memory and executive functions for scholastic achievement as an introduction to the question of whether and how working memory and executive control training may improve academic abilities. The review of current research showed limited but converging evidence for positive effects of process-based complex working-memory training on academic abilities, particularly in the domain of reading. These benefits occurred in children suffering from cognitive and academic deficits as well as in healthy students. Transfer of training to mathematical abilities seemed to be very limited and to depend on the training regime and the characteristics of the study sample. A core issue in training research is whether high- or low-achieving children benefit more from cognitive training. Individual differences in terms of training-related benefits suggested that process-based working memory and executive control training often induced compensation effects with larger benefits in low performing individuals. Finally, we discuss the effects of process-based training in relation to other types of interventions aimed at improving academic achievement.

Abstract: In the research reported here, we tested the hypothesis that sustained engagement in learning new skills that activated working memory, episodic memory, and reasoning over a period of 3 months would enhance cognitive function in older adults. In three conditions with high cognitive demands, participants learned to quilt, learned digital photography, or engaged in both activities for an average of 16.51 hr a week for 3 months. Results at posttest indicated that episodic memory was enhanced in these productive-engagement conditions relative to receptive-engagement conditions, in which participants either engaged in nonintellectual activities with a social group or performed low-demand cognitive tasks with no social contact. The findings suggest that sustained engagement in cognitively demanding, novel activities enhances memory function in older adulthood, but, somewhat surprisingly, we found limited cognitive benefits of sustained engagement in social activities.

Abstract: Can cognitive abilities such as reasoning be improved through working memory training? This question is still highly controversial, with prior studies providing contradictory findings. The lack of theory-driven, systematic approaches and (occasionally serious) methodological shortcomings complicates this debate even more. This review suggests two general mechanisms mediating transfer effects that are (or are not) observed after working memory training: enhanced working memory capacity, enabling people to hold more items in working memory than before training, or enhanced efficiency using the working memory capacity available (e.g., using chunking strategies to remember more items correctly). We then highlight multiple factors that could influence these mechanisms of transfer and thus the success of training interventions. These factors include (1) the nature of the training regime (i.e., intensity, duration, and adaptivity of the training tasks) and, with it, the magnitude of improvements during training, and (2) individual differences in age, cognitive abilities, biological factors, and motivational and personality factors. Finally, we summarize the findings revealed by existing training studies for each of these factors, and thereby present a roadmap for accumulating further empirical evidence regarding the efficacy of working memory training in a systematic way.

Abstract: To find objects of interest in a cluttered and continually changing visual environment, humans must often ignore salient stimuli that are not currently relevant to the task at hand. Recent neuroimaging results indicate that the ability to prevent salience-driven distraction depends on the current level of attentional control activity in frontal cortex, but the specific mechanism by which this control activity prevents salience-driven distraction is still poorly understood. Here, we asked whether salience-driven distraction is prevented by suppressing salient distractors or by preferentially up-weighting the relevant visual dimension. We found that salient distractors were suppressed even when they resided in the same feature dimension as the target (that is, when dimensional weighting was not a viable selection strategy). Our neurophysiological measure of suppression--the PD component of the event-related potential--was associated with variations in the amount of time it took to perform the search task: distractors triggered the PD on fast-response trials, but on slow-response trials they triggered activity associated with working memory representation instead. These results demonstrate that during search salience-driven distraction is mitigated by a suppressive mechanism that reduces the salience of potentially distracting visual objects.

Abstract: Errors in short-term memory increase with the quantity of information stored, limiting the complexity of cognition and behavior. In visual memory, attempts to account for errors in terms of allocation of a limited pool of working memory resources have met with some success, but the biological basis for this cognitive architecture is unclear. An alternative perspective attributes recall errors to noise in tuned populations of neurons that encode stimulus features in spiking activity. I show that errors associated with decreasing signal strength in probabilistically spiking neurons reproduce the pattern of failures in human recall under increasing memory load. In particular, deviations from the normal distribution that are characteristic of working memory errors and have been attributed previously to guesses or variability in precision are shown to arise as a natural consequence of decoding populations of tuned neurons. Observers possess fine control over memory representations and prioritize accurate storage of behaviorally relevant information, at a cost to lower priority stimuli. I show that changing the input drive to neurons encoding a prioritized stimulus biases population activity in a manner that reproduces this empirical tradeoff in memory precision. In a task in which predictive cues indicate stimuli most probable for test, human observers use the cues in an optimal manner to maximize performance, within the constraints imposed by neural noise.

Abstract: Executive function (EF) has come to be an umbrella term used for a diversity of hypothesized cognitive processes, including planning, working memory, attention, inhibition, self-monitoring, self-regulation, and initiation carried out by prefrontal areas of the frontal lobes.

Abstract: The modality effect refers to a cognitive load learning effect that occurs when a mixed-mode (partly visual and partly auditory) presentation of information is more effective than a single-mode (either visual or auditory alone) presentation of the same information. For learning to occur, novel material must be organised and incorporated into long-term memory via a limited working memory. For instruction to be effective, it has to be designed in ways in which the limitations of working memory are overcome. As distraction and interference impose an additional memory load, their impact on the limited working memory system has to be taken into consideration in a multimedia context where the different formats of words and pictures allow for many possible ways of presenting information. The instructional predictions that flow from the experimental work on the modality effect are straightforward. From a practical perspective, the modality effect provides guidelines for effective instruction.

Abstract: Excitation–inhibition balance (E/I balance) is a fundamental property of cortical microcircuitry. Disruption of E/I balance in prefrontal cortex is hypothesized to underlie cognitive deficits observed in neuropsychiatric illnesses such as schizophrenia. To elucidate the link between these phenomena, we incorporated synaptic disinhibition, via N-methyl-D-aspartate receptor perturbation on interneurons, into a network model of spatial working memory (WM). At the neural level, disinhibition broadens the tuning of WM-related, stimulus-selective persistent activity patterns. The model predicts that this change at the neural level leads to 2 primary behavioral deficits: 1) increased behavioral variability that degrades the precision of stored information and 2) decreased ability to filter out distractors during WM maintenance. We specifically tested the main model prediction, broadened WM representation under disinhibition, using behavioral data from human subjects performing a spatial WM task combined with ketamine infusion, a pharmacological model of schizophrenia hypothesized to induce disinhibition. Ketamine increased errors in a pattern predicted by the model. Finally, as proof-of-principle, we demonstrate that WM deteriorations in the model can be ameliorated by compensations that restore E/I balance. Our findings identify specific ways by which cortical disinhibition affects WM, suggesting new experimental designs for probing the brain mechanisms of WM deficits in schizophrenia.