Working memory training: improving intelligence--changing brain activity

The main objectives of the study were: to investigate whether training on working memory (WM) could improve fluid intelligence, and to investigate the effects WM training had on neuroelectric (electroencephalography - EEG) and hemodynamic (near-infrared spectroscopy - NIRS) patterns of brain activity. In a parallel group experimental design, respondents of the working memory group after 30 h of training significantly increased performance on all tests of fluid intelligence. By contrast, respondents of the active control group (participating in a 30-h communication training course) showed no improvements in performance. The influence of WM training on patterns of neuroelectric brain activity was most pronounced in the theta and alpha bands. Theta and lower-1 alpha band synchronization was accompanied by increased lower-2 and upper alpha desynchronization. The hemodynamic patterns of brain activity after the training changed from higher right hemispheric activation to a balanced activity of both frontal areas. The neuroelectric as well as hemodynamic patterns of brain activity suggest that the training influenced WM maintenance functions as well as processes directed by the central executive. The changes in upper alpha band desynchronization could further indicate that processes related to long term memory were also influenced.     

Dorsolateral prefrontal cortex mediates working memory processes in motor skill learning

Neuroimaging studies have shown that the dorsolateral prefrontal cortex (DLPFC) is recruited during motor skill learning, which suggests the involvement of the DLPFC in working memory (WM) processes, such as selection and integration of motor representations temporarily stored in WM. However, direct evidence linking activation of the DLPFC to WM storage and manipulation during motor skill learning in real-time is rare. In this study, we conducted two experiments to investigate the causal role of DLPFC activity in WM storage and manipulation during motor skill learning under low and high WM-demand conditions. Participants received continuous theta burst stimulation (cTBS) and sham stimulation (crossover design) over the left DLPFC (experiment 1) or right DLPFC (experiment 2). Before and after stimulation, participants in both experiments performed a sequential finger-tapping (SFT) task containing repeated sequence (low-WM demand) and non-repeated sequence (high-WM demand) conditions which are used to study WM processes. The number of correct sequences (NoCS) and reproduction error rate were analyzed. Learning gains in NoCS improved significantly with the practice for both sequence types in the presence of either stimulation type. Compared to sham stimulation, cTBS over the left DLPFC resulted in significantly reduced learning gains in NoCS for non-repeated sequences. These results suggest that the left DLPFC contributes to WM manipulation during motor skill learning.     

Neuroimaging studies of working memory:

We performed meta-analyses on 60 neuroimaging (PET and fMRI) studies of working memory (WM), considering three types of storage material (spatial, verbal, and object), three types of executive function (continuous updating of WM, memory for temporal order, and manipulation of information in WM), and interactions between material and executive function. Analyses of material type showed the expected dorsal-ventral dissociation between spatial and nonspatial storage in the posterior cortex, but not in the frontal cortex. Some support was found for left frontal dominance in verbal WM, but only for tasks with low executive demand. Executive demand increased right lateralization in the frontal cortex for spatial WM. Tasks requiring executive processing generally produce more dorsal frontal activations than do storage-only tasks, but not all executive processes show this pattern. Brodmann’s areas (BAs) 6, 8, and 9, in the superior frontal cortex, respond most when WM must be continuously updated and when memory for temporal order must be maintained. Right BAs 10 and 47, in the ventral frontal cortex, respond more frequently with demand for manipulation (including dual-task requirements or mental operations). BA 7, in the posterior parietal cortex, is involved in all types of executive function. Finally, we consider a potential fourth executive function: selective attention to features of a stimulus to be stored in WM, which leads to increased probability of activating the medial prefrontal cortex (BA 32) in storage tasks.     

Increased arithmetic complexity is associated with domain-general but not domain-specific magnitude processing in children: A simultaneous fNIRS-EEG study

The investigation of the neural underpinnings of increased arithmetic complexity in children is essential for developing educational and therapeutic approaches and might provide novel measures to assess the effects of interventions. Although a few studies in adults and children have revealed the activation of bilateral brain regions during more complex calculations, little is known about children. We investigated 24 children undergoing one-digit and two-digit multiplication tasks while simultaneously recording functional near-infrared spectroscopy (fNIRS) and electroencephalography (EEG) data. FNIRS data indicated that one-digit multiplication was associated with brain activity in the left superior parietal lobule (SPL) and intraparietal sulcus (IPS) extending to the left motor area, and two-digit multiplication was associated with activity in bilateral SPL, IPS, middle frontal gyrus (MFG), left inferior parietal lobule (IPL), and motor areas. Oscillatory EEG data indicated theta increase and alpha decrease in parieto-occipital sites for both one-digit and two-digit multiplication. The contrast of two-digit versus one-digit multiplication yielded greater activity in right MFG and greater theta increase in frontocentral sites. Activation in frontal areas and theta band data jointly indicate additional domain-general cognitive control and working memory demands for heightened arithmetic complexity in children. The similarity in parietal activation between conditions suggests that children rely on domain-specific magnitude processing not only for two-digit but—in contrast to adults—also for one-digit multiplication problem solving. We conclude that in children, increased arithmetic complexity tested in an ecologically valid setting is associated with domain-general processes but not with alteration of domain-specific magnitude processing.     

工作记忆的神经振荡调控：基于神经振荡夹带现象

工作记忆的神经振荡机制研究是当前记忆领域的研究热点之一。那么, 神经振荡仅仅是工作记忆过程的伴随现象, 还是直接参与并调控了工作记忆的加工过程?已有研究发现, 大脑内部的神经振荡活动在外界节律性刺激的驱动下, 逐步与外界刺激节律相位同步化, 这一现象被称为“神经振荡夹带”。重复经颅磁刺激(repetitive Transcranial Magnetic Stimulation, rTMS)和经颅交流电刺激(transcranial Alternating Current Stimulation, tACS)干预研究基于此现象, 对大脑局部脑区施加节律性磁、电刺激, 进而调控工作记忆过程中特定频段的神经振荡活动、跨频段的神经振荡耦合或跨脑区的神经振荡相位同步, 为神经振荡参与工作记忆加工提供较为直接的因果证据。未来研究需考虑从脑网络的角度出发, 调控多个脑区之间的神经振荡活动, 进一步考察神经振荡对工作记忆的影响。此外, 还需注意探索和优化rTMS/tACS调控工作记忆的刺激方案, 并辅以客观的脑电记录, 提高该类研究的有效性和可重复性, 最终达到提高工作记忆能力的目的。     

Attention training improves aberrant neural dynamics during working memory processing in veterans with PTSD

Posttraumatic stress disorder (PTSD) is associated with executive functioning deficits, including disruptions in working memory (WM). Recent studies suggest that attention training reduces PTSD symptomatology, but the underlying neural mechanisms are unknown. We used high-density magnetoencephalography (MEG) to evaluate whether attention training modulates brain regions serving WM processing in PTSD. Fourteen veterans with PTSD completed a WM task during a 306-sensor MEG recording before and after 8 sessions of attention training treatment. A matched comparison sample of 12 combat-exposed veterans without PTSD completed the same WM task during a single MEG session. To identify the spatiotemporal dynamics, each group’s data were transformed into the time-frequency domain, and significant oscillatory brain responses were imaged using a beamforming approach. All participants exhibited activity in left hemispheric language areas consistent with a verbal WM task. Additionally, veterans with PTSD and combat-exposed healthy controls each exhibited oscillatory responses in right hemispheric homologue regions (e.g., right Broca’s area); however, these responses were in opposite directions. Group differences in oscillatory activity emerged in the theta band (4–8 Hz) during encoding and in the alpha band (9–12 Hz) during maintenance and were significant in right prefrontal and right supramarginal and inferior parietal regions. Importantly, following attention training, these significant group differences were reduced or eliminated. This study provides initial evidence that attention training improves aberrant neural activity in brain networks serving WM processing.     

An EEG coherence test of the frontal dorsal versus ventral hypothesis in N-back working memory

We examined Goldman-Rakic's conceptualization of working memory (WM) involving modality-specific reverbatory circuits using electrophysiological measures. An n-back WM task, with either spatial information or object information to be remembered, was administered with EEG coherence used to provide a functional measure of corticocortical communication. Support was not found for modality-specific WM systems in this task. Instead, results suggest that there is a circuit that connects dorsal frontal with dorsal parietal areas which is activated during all levels of the WM vigilance task, regardless of the type of stimuli involved. Data also suggest that left frontal areas are sensitive to the level of difficulty of the WM task (0,1,2, and 3-back), as well as familiarity of task demands. Overall, our results support a model that conceptualizes a unitary WM system rather than a system composed of separate WM circuits for different modalities.     

An fMRI study of phonological and spatial working memory using identical stimuli

The aim of the present fMRI study was to localize brain areas that were uniquely activated for phonological versus spatial working memory. Previous studies have reported inconsistent results, most likely because of methodological heterogeneity varying both stimuli and instructions in the same study. Here, identical consonant-vowel-consonant non-words were visually presented to the subjects in a 2-back paradigm under two different instructions; the subjects either had to memorize the non-words per se or their location. The results give evidence for a hemispheric organization of working memory, with dominance for processing of phonological information in the left hemisphere and frontal cortex, and spatial information in the right hemisphere and parietal cortex. The results also reflect a certain overlap between the neuronal network for working memory and processing of verbal and spatial material. These findings are discussed with regard to processing specificity and the extent that activated areas also may reflect perceptual processes.     

双特征空间客体捆绑关系的存储机制——来自ERP的证据

利用事件相关电位技术(ERPs),采用延迟匹配任务的实验范式,测查了16名正常被试完成位置客体、方向客体和位置—方向捆绑客体的工作记忆诱发的皮层慢电位。实验发现:在第1个客体呈现后的300ms到600ms之间,在顶叶皮层(CP5、P7、P3),位置客体、方向客体比捆绑客体诱发了一个更负的成分; 在右额(F4、FC6、FC2、F8)、右后颞(TP10)、左颞(T7)、左后颞(TP9)等脑区,捆绑客体比单一特征客体诱发出了一个更正的成分; 在800至1400ms的慢波成分上,位置和方向捆绑客体在F4、FC6和F8三个记录点诱发的波形比单一特征客体的波形波幅更正。右侧前额叶参与了客体整合表征的存储,该结果支持了情景缓冲器的假设。     

Age-Related Changes in Neural Activation During Working Memory Performance

Changes in frontal lobe functions are a typical part of aging of the brain. There are age-related declines in working memory performance, a skill requiring frontal lobe activation. This study examined neural activation, using [15 O] water positron emission tomography (PET) methodology, during performance on two verbal working memory tasks in younger and older participants. The results demonstrated the typical areas of activation associated with working memory performance (e.g., dorsolateral prefrontal cortex and inferior parietal cortex) in both groups. However, the younger participants utilized the right dorsolateral prefrontal cortex and anterior cingulate gyrus significantly more than the older participants. In turn, the older participants used the left dorsolateral prefrontal cortex significantly more than the younger participants and maintained material-specific lateralization in their pattern of activation. These findings are consistent with a previous report of different age-related patterns of frontal activation during working memory.     

Neural substrates of successful working memory and long-term memory formation in a relational spatial memory task

Working memory (WM) tasks may involve brain activation actually implicated in long-term memory (LTM). In order to disentangle these two memory systems, we employed a combined WM/LTM task, using a spatial relational (object-location) memory paradigm and analyzed which brain areas were associated with successful performance for either task using fMRI. Critically, we corrected for the performance on the respective memory task when analyzing subsequent memory effects. The WM task consisted of a delayed-match-to-sample task assessed in an MRI scanner. Each trial consisted of an indoor or outdoor scene in which the exact configuration of four objects had to be remembered. After a short delay (7–13 s), the scene was presented from a different angle and spatial recognition for two objects was tested. After scanning, participants received an unexpected subsequent recognition memory (LTM) task, where the two previously unprobed objects were tested. Brain activity during encoding, delay phase and probe phase was analyzed based on WM and LTM performance. Results showed that successful WM performance, when corrected for LTM performance, was associated with greater activation in the inferior frontal gyrus and left fusiform gyrus during the early stage of the maintenance phase. A correct decision during the WM probe was accompanied by greater activation in a wide network, including bilateral hippocampus, right superior parietal gyrus and bilateral insula. No voxels exhibited supra-threshold activity during the encoding phase, and we did not find any differential activity for correct versus incorrect trials in the WM task when comparing LTM correct versus LTM incorrect trials.     

Working memory maintenance contributes to long-term memory formation: Evidence from slow event-related brain potentials

Behavioral research has led to conflicting views regarding the relationship between working memory (WM) maintenance and long-term memory (LTM) formation. We used slow event-related brain potentials to investigate the degree to which neural activity during WM maintenance is associated with successful LTM formation. Participants performed a WM task with objects and letter strings, followed by a surprise LTM test. Slow potentials were found to be more negative over the parietal and occipital cortex for objects and over the left frontal cortex for letter strings during WM maintenance. Within each category, they were enhanced for items that were subsequently successfully remembered. These effects were topographically distinct, with maximum effects at those electrodes that showed the maximum negativity during WM maintenance in general. Together, these results are strongly consistent with the ideas that WM maintenance contributes to LTM formation and that this may occur through strengthening of stimulus-specific cortical memory traces.     

Regional and inter-regional theta oscillation during episodic novelty processing

Recent event-related potential (ERP) and functional magnetic resonance imaging (fMRI) studies suggest that novelty processing may be involved in processes that recognize the meaning of a novel sound, during which widespread cortical regions including the right prefrontal cortex are engaged. However, it remains unclear how those cortical regions are functionally integrated during novelty processing. Because theta oscillation has been assumed to have a crucial role in memory operations, we examined local and inter-regional neural synchrony of theta band activity during novelty processing. Fifteen right-handed healthy university students participated in this study. Subjects performed an auditory novelty oddball task that consisted of the random sequence of three types of stimuli such as a target (1000 Hz pure tone), novel (familiar environmental sounds such as dog bark, buzz, car crashing sound and so on), and standard sounds (950 Hz pure tone). Event-related spectra perturbation (ERSP) and the phase-locking value (PLV) were measured from human scalp EEG during task. Non-parametric statistical tests were applied to test for significant differences between stimulus novelty and stimulus targets in ERSP and PLV. The novelty P3 showed significant higher amplitude and shorter latency compared with target P3 in frontocentral regions. Overall, theta activity was significantly higher in the novel stimuli compared with the target stimuli. Specifically, the difference in theta power between novel and target stimuli was most significant in the right frontal region. This right frontal theta activity was accompanied by phase synchronization with the left temporal region. Our results imply that theta phase synchronization between right frontal and left temporal regions underlie the retrieval of memory traces for unexpected but familiar sounds from long term memory in addition to working memory retrieval or novelty encoding.     

信息保持、短时存贮与执行控制的脑模型

工作记忆的脑机制是认知神经科学研究的重要问题。研究者在分析大量脑成像研究数据的基础上,创建了许多解释工作记忆神经基础的模型。Smith和Jonides发展了工作记忆成分结构模型和贮存与执行加工分离模型。Postle和D’Esposito建构了工作记忆相对表征混合模型,D’Esposito, Postle和Rypma阐述了工作记忆加工阶段动态模型。Fletcher和Henson提出工作记忆额叶分区整合理论。该文通过对上述模型或理论的评价,总结了工作记忆脑成像研究中存在的问题,展望了该研究领域的发展趋势。     

Dissociation of the neural systems for working memory maintenance of verbal and nonspatial visual information

Working memory for names and faces was investigated to ascertain whether verbal and nonspatial visual information is maintained in working memory by separate neural systems. The subjects performed a delayed match-to-sample task for famous or unfamous faces and names and a sensorimotor control task. Several occipital, temporal, parietal, and prefrontal areas were activated during all memory delays, in comparison with the control delays. Greater delay activity for unfamous faces than for names was obtained in the right fusiform gyrus, right inferior frontal gyrus (IFG), right IFG/ precentral gyrus, and right medial superior frontal gyrus, whereas greater delay activity for unfamous names than for faces was observed in the precuneus, left insula/postcentral gyrus, and left IFG/ precentral gyrus. There was no significant difference in the prefrontal activity in the comparison between famous faces and names. Greater delay activity for famous names than for faces was obtained in visual association and parietal areas. The results indicate that there is a functional dissociation based on information type within the neural system that is responsible for working memory maintenance of verbal and nonspatial visual information.     

视觉空间工作记忆组间差异的神经机制

为考察视觉空间工作记忆（working memory, WM）维持和操作的组间差异及其神经机制,本研究记录了高、低WM组完成延迟再认（维持）任务和心理旋转（操作）任务时的行为和事件相关电位数据。结果发现,在操作任务中,高WM组比低WM组的反应时显著更短;高WM组的中前额叶慢波显著更正、双侧后顶区慢波显著更负,并且两者的波幅显著负相关。在维持任务中,两组被试的反应时无显著差异;高WM组的中前额叶慢波显著更正。结果表明,高WM组的执行注意能力可能更强,能通过有效调节和分配加工资源来表征视觉信息。     

EEG theta phase coupling during executive control of visual working memory investigated in individuals with schizophrenia and in healthy controls

In healthy humans, it has been shown that executive functions are associated with increased frontal-midline EEG theta activity and theta phase coupling between frontal and posterior brain regions. In individuals with schizophrenia, central executive functions are supposed to be heavily impaired. Given that theta phase coupling is causally involved in central executive functions, one would expect that patients with an executive function deficit should display abnormal EEG theta synchronization. We therefore investigated executive functioning in 21 healthy controls and 21 individuals with schizophrenia while they performed a visuospatial delayed match to sample task. The task required either high executive demands (manipulation of content in working memory [WM]) or low executive demands (retention of WM content). In addition, WM load (one vs. three items) was varied. Results indicated higher frontal theta activity for manipulation processes than for retention processes in patients with schizophrenia, as compared with healthy controls, independently of WM load. Furthermore, individuals with schizophrenia revealed a reduction in theta phase coupling during early stages of the delay period for retention, as well as for manipulation processes at high-WM loads. Deviations in theta phase coupling in individuals with schizophrenia were mainly characterized by aberrant fronto-posterior connections, but also by attenuated posterior connections during manipulation of high-WM load. To conclude, fronto-parietal theta coupling seems to be substantially involved in executive control, whereas frontal theta activity seems to reflect general task demands, such as deployment of attentional resources during WM.     

Neural mechanisms of interference control underlie the relationship between fluid intelligence and working memory span

Fluid intelligence (gF) and working memory (WM) span predict success in demanding cognitive situations. Recent studies show that much of the variance in gF and WM span is shared, suggesting common neural mechanisms. This study provides a direct investigation of the degree to which shared variance in gF and WM span can be explained by neural mechanisms of interference control. The authors measured performance and functional magnetic resonance imaging activity in 102 participants during the n-back WM task, focusing on the selective activation effects associated with high-interference lure trials. Brain activity on these trials was correlated with gF, WM span, and task performance in core brain regions linked to WM and executive control, including bilateral dorsolateral prefrontal cortex (middle frontal gyrus; BA9) and parietal cortex (inferior parietal cortex; BA 40/7). Interference-related performance and interference-related activity accounted for a significant proportion of the shared variance in gF and WM span. Path analyses indicate that interference control activity may affect gF through a common set of processes that also influence WM span. These results suggest that individual differences in interference-control mechanisms are important for understanding the relationship between gF and WM span.     

The effect of memory load on cortical activity in the spatial working memory circuit

Accumulating evidence from electrophysiology and neuroimaging studies suggests that spatial working memory is subserved by a network of frontal and parietal regions. In the present study, we parametrically varied the memory set size (one to four spatial locations) of a delayed-response task and applied time-resolved fMRI to study the influence of memory load upon the spatial working memory circuit. Our behavioral results showed that performance deteriorates (lower accuracy and longer reaction time) as memory load increases. Memory load influenced cortical activity during the cue, delay, and response phases of the delayed-response task. Although delay-related activity in many regions increased with increasing memory load, it also was significantly reduced in the middle frontal gyrus and frontal eye fields and leveled off in the parietal areas when memory load increased further. Delayrelated activity in the left posterior parietal cortex was also lower during the error trials, in comparison with the correct trials. Our findings indicate that the delay period activity in the spatial working memory circuit is load sensitive and that the attenuation of this signal is the neural manifestation of performance limitation in the face of excessive memory load.     

Positive affect improves working memory: Implications for controlled cognitive processing

This study examined the effects of positive affect on working memory (WM) and short-term memory (STM). Given that WM involves both storage and controlled processing and that STM primarily involves storage processing, we hypothesised that if positive affect facilitates controlled processing, it should improve WM more than STM. The results demonstrated that positive affect, compared with neutral affect, significantly enhanced WM, as measured by the operation span task. The influence of positive affect on STM, however, was weaker. These results suggest that positive affect enhances WM, a task that involves controlled processing, not just storage processing. Additional analyses of recall and processing times and accuracy further suggest that improved WM under positive affect is not attributable to motivational differences, but results instead from improved controlled cognitive processing.