Neural Patterns in Parietal Cortex and Hippocampus Distinguish Retrieval of Start versus End Positions in Working Memory.

Coding serial order of information is a fundamental ability of our cognitive system, and still, little is known about its neural substrate. This study examined the neural substrates involved in the retrieval of information that is serially stored in verbal working memory task using a sensitive multivariate analysis approach. We compared neural activity for memorized items stemming from the beginning versus the end of a memory list assessing the degree of neural pattern discordance between order positions (beginning vs. end). The present results confirmed and refined the role of the intraparietal sulcus in the processing of serial order information in working memory. An important finding is that the hippocampus showed sensitivity to serial order information. Our results indicate that the representation of serial order information relies on a broader set of neural areas and highlight the role of the intraparietal sulcus and the hippocampus, in addition to the supramarginal gyrus and the SMA. The contribution of different neural regions might reflect the involvement of distinct levels of serial order coding (i.e., spatial, attentional, temporal) that support the representation of serial order information.

Spatial Attention in Serial Order Working Memory: An EEG Study.

Theoretical models explaining serial order processing link order information to specified position markers. However, the precise characteristics of position marking have remained largely elusive. Recent studies have shown that space is involved in marking serial position of items in verbal working memory (WM). Furthermore, it has been suggested, but not proven, that accessing these items involves horizontal shifts of spatial attention. We used continuous electroencephalography recordings to show that memory search in serial order verbal WM involves spatial attention processes that share the same electrophysiological signatures as those operating on the visuospatial WM and external space. Accessing an item from a sequence in verbal WM induced posterior "early directing attention negativity" and "anterior directing attention negativity" contralateral to the position of the item in mental space (i.e., begin items on the left; end items on the right). In the frequency domain, we observed posterior alpha suppression contralateral to the position of the item. Our results provide clear evidence for the involvement of spatial attention in retrieving serial information from verbal WM. Implications for WM models are discussed.

From Counting to Retrieving: Neural Networks Underlying Alphabet Arithmetic Learning.

This fMRI study aimed at unraveling the neural basis of learning alphabet arithmetic facts, as a proxy of the transition from slow and effortful procedural counting-based processing to fast and effortless processing as it occurs in learning addition arithmetic facts. Neural changes were tracked while participants solved alphabet arithmetic problems in a verification task (e.g., F + 4 = J). Problems were repeated across four learning blocks. Two neural networks with opposed learning-related changes were identified. Activity in a network consisting of basal ganglia and parieto-frontal areas decreased with learning, which is in line with a reduction of the involvement of procedure-based processing. Conversely, activity in a network involving the left angular gyrus and, to a lesser extent, the hippocampus gradually increases with learning, evidencing the gradual involvement of retrieval-based processing. Connectivity analyses gave insight in the functional relationship between the two networks. Despite the opposing learning-related trajectories, it was found that both networks become more integrated. Taking alphabet arithmetic as a proxy for learning arithmetic, the present results have implications for current theories of learning arithmetic facts and can give direction to future developments.

The neural basis of metacognitive monitoring during arithmetic in the developing brain.

In contrast to a substantial body of research on the neural basis of cognitive performance in several academic domains, less is known about how the brain generates metacognitive (MC) awareness of such performance. The existing work on the neurobiological underpinnings of metacognition has almost exclusively been done in adults and has largely focused on lower level cognitive processing domains, such as perceptual decision-making. Extending this body of evidence, we investigated MC monitoring by asking children to solve arithmetic problems, an educationally relevant higher-order process, while providing concurrent MC reports during fMRI acquisition. Results are reported on 50 primary school children aged 9-10 years old. The current study is the first to demonstrate that brain activity during MC monitoring, relative to the control task, increased in the left inferior frontal gyrus in children. This brain activity further correlated with children's arithmetic development over a 3-year time period. These data are in line with the frequently suggested, yet never empirically tested, hypothesis that activity in the prefrontal cortex during arithmetic is related to the higher-order process of MC monitoring.

Disentangling Neural Sources of Problem Size and Interference Effects in Multiplication.

Multiplication is thought to be primarily solved via direct retrieval from memory. Two of the main factors known to influence the retrieval of multiplication facts are problem size and interference. Because these factors are often intertwined, we sought to investigate the unique influences of problem size and interference on both performance and neural responses during multiplication fact retrieval in healthy adults. Behavioral results showed that both problem size and interference explained separate unique portions of RT variance, but with significantly stronger contribution from problem size, which contrasts with previous work in children. Whole-brain fMRI results relying on a paradigm that isolated multiplication fact retrieval from response selection showed highly overlapping brain areas parametrically modulated by both problem size and interference in a large network of frontal, parietal, and subcortical brain areas. Subsequent analysis within these regions revealed problem size to be the stronger and more consistent "unique" modulating factor in overlapping regions as well as those that appeared to respond only to problem size or interference at the whole-brain level, thus underscoring the need to look beyond anatomical overlap using arbitrary thresholds. Additional unique contributions of interference (beyond problem size) were identified in right angular gyrus and subcortical regions associated with procedural processing. Together, our results suggest that problem size, relative to interference, tends to be the more dominant factor in driving behavioral and neural responses during multiplication fact retrieval in adults. Nevertheless, unique contributions of both factors demonstrate the importance of considering the overlapping and unique contributions of each in explaining the cognitive and neural bases of mental multiplication.

Category specific recall in acute stroke: a case with letter speech.

Category selective recall in spontaneous speech after stroke has been reported only rarely. We recently described three cases demonstrating transient number speech in the acute stage of left hemispheric stroke and hypothesized a link with multilingualism and mathematical proficiency. In this report, we describe a similar case with a transient episode of utterances of randomly selected letters. Like in the three previous cases, this episode was preceded by a brief stage of mutism and ultimately evolved to Wernicke's aphasia over a period of days. This phenomenon is reviewed with reference to linguistic models and neuroanatomic and neurophysiological correlates.

More than number sense: The additional role of executive functions and metacognition in arithmetic.

Arithmetic is a major building block for children's development of more complex mathematical abilities. Knowing which cognitive factors underlie individual differences in arithmetic is key to gaining further insight into children's mathematical development. The current study investigated the role of executive functions and metacognition (domain-general cognitive factors) as well as symbolic numerical magnitude processing (domain-specific cognitive factor) in arithmetic, enabling the investigation of their unique contribution in addition to each other. Participants were 127 typically developing second graders (7- and 8-year-olds). Our within-participant design took into account different components of executive functions (i.e., inhibition, shifting, and updating), different aspects of metacognitive skills (i.e., task-specific and general metacognition), and different levels of experience in arithmetic, namely addition (where second graders had extensive experience) and multiplication (where second graders were still in the learning phase). This study reveals that both updating and metacognitive monitoring are important unique predictors of arithmetic in addition to each other and to symbolic numerical magnitude processing. Our results point to a strong and unique role of task-specific metacognitive monitoring skills. These individual differences in noticing one's own errors might help one to learn from his or her mistakes.

Physical activity to improve cognition in older adults: can physical activity programs enriched with cognitive challenges enhance the effects? A systematic review and meta-analysis.

BACKGROUND: Aging-related cognitive decline and cognitive impairment greatly impacts older adults' daily life. The worldwide ageing of the population and associated wave of dementia urgently calls for prevention strategies to reduce the risk of cognitive decline. Physical activity (PA) is known to improve cognitive function at older age through processes of neuroplasticity. Yet, emerging studies suggest that larger cognitive gains may be induced when PA interventions are combined with cognitive activity (CA). This meta-analysis evaluates these potential synergistic effects by comparing cognitive effects following combined PA + CA interventions to PA interventions (PA only), CA interventions (CA only) and control groups. METHODS: Pubmed, Embase, PsycInfo, CINAHL and Sportdiscus were searched for English peer-reviewed papers until April 2018. Data were extracted on cognition and factors potentially influencing the cognitive effects: mode of PA + CA combination (sequential or simultaneous), session frequency and duration, intervention length and study quality. Differences between older adults with and without mild cognitive impairments were also explored. RESULTS: Forty-one studies were included. Relative to the control group, combined PA + CA intervention showed significantly larger gains in cognition (g = 0.316; 95% CI 0.188-0.443; p < .001). Studies that compared combined PA + CA with PA only, showed small but significantly greater cognitive improvement in favor of combined interventions (g = 0.160; 95% CI 0.041-0.279; p = .008). No significant difference was found between combined PA + CA and CA only interventions. Furthermore, cognitive effects tended to be more pronounced for studies using simultaneous designs (g = 0.385; 95%CI 0.214-0.555; p < .001) versus sequential designs (g = 0.114; 95%CI -0.102- 0.331, p = .301). Effects were not moderated by session frequency, session duration, intervention length or study quality. Also, no differences in effects were found between older adults with and without mild cognitive impairments. CONCLUSION: Findings of the current meta-analysis suggest that PA programs for older adults could integrate challenging cognitive exercises to improve cognitive health. Combined PA + CA programs should be promoted as a modality for preventing as well as treating cognitive decline in older adults. Sufficient cognitive challenge seems more important to obtain cognitive effects than high doses of intervention sessions.

Common and distinct brain regions in both parietal and frontal cortex support symbolic and nonsymbolic number processing in humans: A functional neuroimaging meta-analysis.

In recent years, there has been substantial growth in neuroimaging studies investigating neural correlates of symbolic (e.g. Arabic numerals) and non-symbolic (e.g. dot arrays) number processing. At present it remains contested whether number is represented abstractly, or if number representations in the brain are format-dependent. In order to quantitatively evaluate the available neuroimaging evidence, we used activation likelihood estimation (ALE) to conduct quantitative meta-analyses of the results reported in 57 neuroimaging papers. Consistent with the existence of an abstract representation of number in the brain, conjunction analyses revealed overlapping activation for symbolic and nonsymbolic numbers in frontal and parietal lobes. Consistent with the notion of format-dependent activation, contrast analyses demonstrated anatomically distinct fronto-parietal activation for symbolic and non-symbolic processing. Therefore, symbolic and non-symbolic numbers are subserved by format-dependent and abstract neural systems. Moreover, the present results suggest that regions across the parietal cortex, not just the intraparietal sulcus, are engaged in both symbolic and non-symbolic number processing, challenging the notion that the intraparietal sulcus is the key region for number processing. Additionally, our analyses indicate that regions in the frontal cortex subserve magnitude representations rather than non-numerical cognitive processes associated with number tasks, thereby highlighting the importance of considering both frontal and parietal regions as important for number processing.

Brain networks under attack: robustness properties and the impact of lesions.

A growing number of studies approach the brain as a complex network, the so-called 'connectome'. Adopting this framework, we examine what types or extent of damage the brain can withstand-referred to as network 'robustness'-and conversely, which kind of distortions can be expected after brain lesions. To this end, we review computational lesion studies and empirical studies investigating network alterations in brain tumour, stroke and traumatic brain injury patients. Common to these three types of focal injury is that there is no unequivocal relationship between the anatomical lesion site and its topological characteristics within the brain network. Furthermore, large-scale network effects of these focal lesions are compared to those of a widely studied multifocal neurodegenerative disorder, Alzheimer's disease, in which central parts of the connectome are preferentially affected. Results indicate that human brain networks are remarkably resilient to different types of lesions, compared to other types of complex networks such as random or scale-free networks. However, lesion effects have been found to depend critically on the topological position of the lesion. In particular, damage to network hub regions-and especially those connecting different subnetworks-was found to cause the largest disturbances in network organization. Regardless of lesion location, evidence from empirical and computational lesion studies shows that lesions cause significant alterations in global network topology. The direction of these changes though remains to be elucidated. Encouragingly, both empirical and modelling studies have indicated that after focal damage, the connectome carries the potential to recover at least to some extent, with normalization of graph metrics being related to improved behavioural and cognitive functioning. To conclude, we highlight possible clinical implications of these findings, point out several methodological limitations that pertain to the study of brain diseases adopting a network approach, and provide suggestions for future research.

How serially organized working memory information interacts with timing.

The temporary storage of serial order information in working memory (WM) has been demonstrated to be crucial to higher order cognition. The previous studies have shown that the maintenance of serial order can be a consequence of the construction of position markers to which to-be-remembered information will be bound. However, the nature of these position markers remains unclear. In this study, we demonstrate the crucial involvement of time in the construction of these markers by establishing a bidirectional relationship. First, results of the first experiment show that the initial items in WM result in faster responding after shorter time presentations, while we observe the opposite for items stored further in WM. Second, in the next experiment, we observe an effect of temporal cueing on WM retrieval; longer time cues facilitate responding to later WM items compared with items stored at the beginning of WM. These findings are discussed in the context of position marker theories, reviewing the functional involvement of time in the construction of these markers and its association with space.

Unsigned value prediction-error modulates the motor system in absence of choice.

Human actions are driven by the pursuit of goals, especially when achieving these goals entails a reward. Accordingly, recent work showed that anticipating a reward in a motor task influences the motor system, boosting motor excitability and increasing overall readiness. Attaining a reward typically requires some mental or physical effort. Recent neuroimaging evidence suggested that both reward expectation and effort requirements are encoded by a partially overlapping brain network. Moreover, reward and effort information are combined in an integrative value signal. However, whether and how mental effort is integrated with reward at the motor level during task preparation remains unclear. To address these issues, we implemented a mental effort task where reward expectation and effort requirements were manipulated. During task preparation, TMS was delivered on the motor cortex and motor-evoked potentials (MEPs) were recorded on the right hand muscles to probe motor excitability. The results showed an interaction of effort and reward in modulating the motor system, reflecting an unsigned value prediction-error signal. Crucially, this was observed in the motor system in absence of a value-based decision or value-driven action selection. This suggests a high-level cognitive factor such as unsigned value prediction-error can modulate the motor system. Interestingly, effort-related motor excitability was also modulated by individual differences in tendency to engage in (and enjoy) mental effort, as measured by the Need for Cognition questionnaire, underlining a role of subjective effort experience in value-driven preparation for action.

Anticipatory processes in brain state switching - evidence from a novel cued-switching task implicating default mode and salience networks.

The default mode network (DMN) is the core brain system supporting internally oriented cognition. The ability to attenuate the DMN when switching to externally oriented processing is a prerequisite for effective performance and adaptive self-regulation. Right anterior insula (rAI), a core hub of the salience network (SN), has been proposed to control the switching from DMN to task-relevant brain networks. Little is currently known about the extent of anticipatory processes subserved by DMN and SN during switching. We investigated anticipatory DMN and SN modulation using a novel cued-switching task of between-state (rest-to-task/task-to-rest) and within-state (task-to-task) transitions. Twenty healthy adults performed the task implemented in an event-related functional magnetic resonance imaging (fMRI) design. Increases in activity were observed in the DMN regions in response to cues signalling upcoming rest. DMN attenuation was observed for rest-to-task switch cues. Obversely, DMN was up-regulated by task-to-rest cues. The strongest rAI response was observed to rest-to-task switch cues. Task-to-task switch cues elicited smaller rAI activation, whereas no significant rAI activation occurred for task-to-rest switches. Our data provide the first evidence that DMN modulation occurs rapidly and can be elicited by short duration cues signalling rest- and task-related state switches. The role of rAI appears to be limited to certain switch types - those implicating transition from a resting state and to tasks involving active cognitive engagement.

Attention-based rehearsal: Eye movements reveal how visuospatial information is maintained in working memory.

The human eye scans visual information through scan paths, series of fixations. Analogous to these scan paths during the process of actual "seeing," we investigated whether similar scan paths are also observed while subjects are "rehearsing" stimuli in visuospatial working memory. Participants performed a continuous recall task in which they rehearsed the precise location and color of three serially presented discs during a retention interval, and later reproduced either the precise location or the color of a single probed item. In two experiments, we varied the direction along which the items were presented and investigated whether scan paths during rehearsal followed the pattern of stimulus presentation during encoding (left-to-right in Experiment 1; left-to-right/right-to-left in Experiment 2). In both experiments, we confirmed that the eyes follow similar scan paths during encoding and rehearsal. Specifically, we observed that during rehearsal participants refixated the memorized locations they saw during encoding. Most interestingly, the precision with which these locations were refixated was associated with smaller recall errors. Assuming that eye position reflects the focus of attention, our findings suggest a functional contribution of spatial attention shifts to working memory and are in line with the hypothesis that maintenance of information in visuospatial working memory is supported by attention-based rehearsal. (PsycInfo Database Record (c) 2024 APA, all rights reserved).

From long-term to short-term: Distinct neural networks underlying semantic knowledge and its recruitment in working memory.

Although numerous studies suggest that working memory (WM) and semantic long-term knowledge interact, the nature and underlying neural mechanisms of this intervention remain poorly understood. Using functional magnetic resonance imaging (fMRI), this study investigated the extent to which neural markers of semantic knowledge in long-term memory (LTM) are activated during the WM maintenance stage in 32 young adults. First, the multivariate neural patterns associated with four semantic categories were determined via an implicit semantic activation task. Next, the participants maintained words - the names of the four semantic categories implicitly activated in the first task - in a verbal WM task. Multi-voxel pattern analyses showed reliable neural decoding of the four semantic categories in the implicit semantic activation and the verbal WM tasks. Critically, however, no between-task classification of semantic categories was observed. Searchlight analyses showed that for the WM task, semantic category information could be decoded in anterior temporal areas associated with abstract semantic category knowledge. In the implicit semantic activation task, semantic category information was decoded in superior temporal, occipital and frontal cortices associated with domain-specific semantic feature representations. These results indicate that item-level semantic activation during verbal WM involves shallow rather than deep semantic information.

Picture novelty attenuates semantic interference and modulates concomitant neural activity in the anterior cingulate cortex and the locus coeruleus.

Goal-directed behavior requires the ability to focus on information that is relevant to a given task and to ignore information that might interfere with it. In the Stroop task, for example, the influence of an irrelevant word needs to be overcome, which is believed to be difficult because it arises in a fast and automatic fashion, which effectively renders it very salient. Here we address the question of whether this can be counteracted by increasing the saliency of the task-relevant input, for example by modulating its relative novelty, which increases saliency in a fairly implicit and controlled fashion. To test the influence of novelty on interference processing, we employed a picture-word interference task in the fMRI scanner, in which we manipulated the novelty of the task-relevant picture. We found that picture novelty indeed reduced typical behavioral interference from incongruent words. Moreover, familiar incongruent trials were associated with activity increases in the anterior cingulate cortex (ACC), a prime conflict-processing region, as well as in the noradrenergic locus coeruleus (LC), which entertains connections both to and from the ACC. The lack of analogous activations in novel incongruent trials suggests that the reduction of behavioral interference was not related to enhanced conflict-resolution processes, but rather to the automatic prioritization of novel pictures which appears to avert the influence of irrelevant words at the front end. Interestingly, activity in the ACC and LC was slightly stronger in novel congruent trials compared to incongruent ones, which may reflect increased relevance of novel stimuli when encoded in a congruent context. In summary, the present data demonstrate that stimulus novelty clearly reduces semantic interference, and highlights a complex interaction of interference and novelty processing on the neural level, including an involvement of the noradrenergic system in the processing of cognitively and perceptually salient events.

Spatial attention interacts with serial-order retrieval from verbal working memory.

The ability to maintain the serial order of events is recognized as a major function of working memory. Although general models of working memory postulate a close link between working memory and attention, such a link has so far not been proposed specifically for serial-order working memory. The present study provided the first empirical demonstration of a direct link between serial order in verbal working memory and spatial selective attention. We show that the retrieval of later items of a sequence stored in working memory-compared with that of earlier items-produces covert attentional shifts toward the right. This observation suggests the conceptually surprising notion that serial-order working memory, even for nonspatially defined verbal items, draws on spatial attention.

Attention supports verbal short-term memory via competition between dorsal and ventral attention networks.

Interactions between the neural correlates of short-term memory (STM) and attention have been actively studied in the visual STM domain but much less in the verbal STM domain. Here we show that the same attention mechanisms that have been shown to shape the neural networks of visual STM also shape those of verbal STM. Based on previous research in visual STM, we contrasted the involvement of a dorsal attention network centered on the intraparietal sulcus supporting task-related attention and a ventral attention network centered on the temporoparietal junction supporting stimulus-related attention. We observed that, with increasing STM load, the dorsal attention network was activated while the ventral attention network was deactivated, especially during early maintenance. Importantly, activation in the ventral attention network increased in response to task-irrelevant stimuli briefly presented during the maintenance phase of the STM trials but only during low-load STM conditions, which were associated with the lowest levels of activity in the dorsal attention network during encoding and early maintenance. By demonstrating a trade-off between task-related and stimulus-related attention networks during verbal STM, this study highlights the dynamics of attentional processes involved in verbal STM.

Brain correlates of subjective freedom of choice.

The subjective feeling of free choice is an important feature of human experience. Experimental tasks have typically studied free choice by contrasting free and instructed selection of response alternatives. These tasks have been criticised, and it remains unclear how they relate to the subjective feeling of freely choosing. We replicated previous findings of the fMRI correlates of free choice, defined objectively. We introduced a novel task in which participants could experience and report a graded sense of free choice. BOLD responses for conditions subjectively experienced as free identified a postcentral area distinct from the areas typically considered to be involved in free action. Thus, the brain correlates of subjective feeling of free action were not directly related to any established brain correlates of objectively-defined free action. Our results call into question traditional assumptions about the relation between subjective experience of choosing and activity in the brain's so-called voluntary motor areas.

Errors and conflict at the task level and the response level.

In the last decade, research on error and conflict processing has become one of the most influential research areas in the domain of cognitive control. There is now converging evidence that a specific part of the posterior frontomedian cortex (pFMC), the rostral cingulate zone (RCZ), is crucially involved in the processing of errors and conflict. However, error-related research has focused primarily on a specific error type, namely, response errors. The aim of the present study was to investigate whether errors on the task level rely on the same neural and functional mechanisms. Here we report a dissociation of both error types in the pFMC: whereas response errors activate the RCZ, task errors activate the dorsal frontomedian cortex. Although this last region shows an overlap in activation for task and response errors on the group level, a closer inspection of the single-subject data is more in accordance with a functional anatomical dissociation. When investigating brain areas related to conflict on the task and response levels, a clear dissociation was perceived between areas associated with response conflict and with task conflict. Overall, our data support a dissociation between response and task levels of processing in the pFMC. In addition, we provide additional evidence for a dissociation between conflict and errors both at the response level and at the task level.