Neural correlates of learning and working memory in the primate posterior parietal cortex

The posterior parietal cortex has been traditionally associated with coordinate transformations necessary for interaction with the environment and with visual-spatial attention. More recently, involvement of posterior parietal cortex in other cognitive functions such as working memory and task learning has become evident. Neurophysiological experiments in non-human primates and human imaging studies have revealed neural correlates of memory and learning at the single neuron and at the brain network level. During working memory, posterior parietal neurons continue to discharge and to represent stimuli that are no longer present. This activation resembles the responses of prefrontal neurons, although important differences have been identified in terms of the ability to resist stimulation by distracting stimuli, which is more evident in the prefrontal than the posterior parietal cortex. Posterior parietal neurons also become active during tasks that require the organization of information into larger structured elements and their activity is modulated according to learned context-dependent rules. Neural correlates of learning can be observed in the mean discharge rate and spectral power of neuronal spike trains after training to perform new task sets or rules. These findings demonstrate the importance of posterior parietal cortex in brain networks mediating working memory and learning.     

Parietal lobe contributions to episodic memory retrieval

Although the parietal lobe is not traditionally thought to support declarative memory, recent event-related fMRI studies of episodic retrieval have consistently revealed a range of memory-related influences on activation in lateral posterior parietal cortex (PPC) and precuneus extending into posterior cingulate and retrosplenial cortex. This article surveys the fMRI literature on PPC activation during remembering, a literature that complements earlier electroencephalography data. We consider these recent memory-related fMRI responses within the context of classical ideas about parietal function that emphasize space-based attention and motor intention. We conclude by proposing three hypotheses concerning how parietal cortex might contribute to memory.     

The neural regions sustaining episodic encoding and recognition of objects

In this functional MRI experiment, encoding of objects was associated with activation in left ventrolateral prefrontal/insular and right dorsolateral prefrontal and fusiform regions as well as in the left putamen. By contrast, correct recognition of previously learned objects (R judgments) produced activation in left superior frontal, bilateral inferior frontal, and right cerebellar regions, whereas correct rejection of distractor objects (N judgments) was associated with activation in bilateral prefrontal and anterior cingulate cortices, in right parietal and cerebellar regions, in the left putamen, and in the right caudate nucleus. The R minus N comparison showed activation in the left lateral prefrontal cortex and in bilateral cingulate cortices and precunei, while the N minus R comparison did not reveal any positive signal change. These results support the view that similar regions of the frontal lobe are involved in episodic encoding and retrieval processes, and that the successful episodic retrieval of newly learned objects is mainly based on a frontoparietal network.     

Age-related changes in neural mechanisms of prospective memory

The capability to remember and execute intentions in the future – termed prospective memory (PM) – may be of special significance for older adults to enable successful completion of important activities of daily living. Despite the importance of this cognitive function, mixed findings have been obtained regarding age-related decline in PM, and, currently, there is limited understanding of potential contributing mechanisms. In the current study, older (N=41) and younger adults (N=47) underwent task-functional MRI during performance of PM conditions that encouraged either spontaneous retrieval (Focal) or sustained attentional monitoring (Non-focal) to detect PM targets. Older adults exhibited a reduction in PM-related sustained activity within the anterior prefrontal cortex (aPFC) and associated dorsal frontoparietal cognitive control network, due to an increase in non-specific sustained activation in (no-PM) control blocks (i.e., an age-related compensatory shift). Transient PM-trial specific activity was observed in both age groups within a ventral parietal memory network that included the precuneus. However, within a left posterior inferior parietal node of this network, transient PM-related activity was selectively reduced in older adults during the non-focal condition. These age differences in sustained and transient brain activity statistically mediated age-related declines in PM performance, and were potentially linked via age-related changes in functional connectivity between the aPFC and precuneus. Together, they support an account consistent with the Dual Mechanisms of Control framework, in which age-related PM declines are due to neural mechanisms that support proactive cognitive control processes, such as sustained attentional monitoring, while leaving reactive control mechanisms relatively spared.     

The mid-ventrolateral prefrontal cortex and active mnemonic retrieval

The role of the mid-ventrolateral prefrontal cortex in memory retrieval is examined and compared with the role of the mid-dorsolateral prefrontal cortex in the monitoring of information in memory. It has been argued that the mid-ventrolateral prefrontal cortex (areas 47/12 and 45) is involved in the active retrieval of information from posterior cortical association areas. Active retrieval is required when stimuli in memory do not bear stable relations to each other and therefore retrieval cannot be automatically driven by strong, stable, and unambiguous stimulus or context relations. Data from functional activation studies with normal human subjects are presented that have demonstrated specific changes in activity within the mid-ventrolateral region of the frontal cortex in relation to the active retrieval of information from memory. By contrast, increases in activity in the mid-dorsolateral region of the frontal cortex occur when the performance of the tasks requires monitoring of information in memory.     

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.     

How we use rules to select actions: A review of evidence from cognitive neuroscience

Much of our behavior is guided by rules, or prescribed guides for action. In this review, I consider the current state of knowledge of how rules are learned, stored in the brain, and retrieved and used as the need arises. The focus is primarily on studies in humans, but the review is informed by relevant studies in nonhuman primates. Ventrolateral prefrontal cortex (VLPFC) has been implicated in rule learning, retrieval from long-term memory, and on-line maintenance during task preparation. Interactions between VLPFC and temporal cortex are required for rule retrieval in nonhuman primates, and brain imaging findings in humans suggest that rule knowledge is stored in the posterior middle temporal gyrus. Dorsolateral PFC appears to be more closely related to rule-based response selection than to rule retrieval. An important task for the future is to explain how PFC, basal ganglia, and temporal, parietal, and motor cortices interact to produce rule-guided behavior.     

Neural correlates of autobiographical memory retrieval in children and adults

Autobiographical memory (AM) is a critically important form of memory for life events that undergoes substantial developmental changes from childhood to adulthood. Relatively little is known regarding the functional neural correlates of AM retrieval in children as assessed with fMRI, and how they may differ from adults. We investigated this question with 14 children ages 8–11 years and 14 adults ages 19–30 years, contrasting AM retrieval with semantic memory (SM) retrieval. During scanning, participants were cued by verbal prompts to retrieve previously selected recent AMs or to verify semantic properties of words. As predicted, both groups showed AM retrieval-related increased activation in regions implicated in prior studies, including bilateral hippocampus, and prefrontal, posterior cingulate, and parietal cortices. Adults showed greater activation in the hippocampal/parahippocampal region as well as prefrontal and parietal cortex, relative to children; age-related differences were most prominent in the first 8?sec versus the second 8?sec of AM retrieval and when AM retrieval was contrasted with semantic retrieval. This study is the first to characterise similarities and differences during AM retrieval in children and adults using fMRI.     

Age-related neural activity during allocentric spatial memory

Age-related decline in allocentric (viewer-independent) spatial memory is seen across species. We employed a virtual reality analogue of the Morris Water Maze to study the effect of healthy ageing on neural activity during allocentric spatial memory using functional magnetic resonance imaging. Voxel-based morphometry was used to ascertain hippocampal volumetric integrity. A widespread neural network comprising frontal, parietal, occipital, thalamic, and cerebellar regions was activated in young and older adults, but only young adults significantly activated bilateral hippocampus and left parahippocampus, as well as right frontal pole and dorso-lateral prefrontal cortex (DLPFC) during encoding and right DLPC during retrieval. Hippocampal grey matter volume was unchanged in older adults; however, prefrontal and parahippocampal functional attenuation was accompanied by volumetric reduction. We conclude that the decline in allocentric spatial memory with age is associated with attenuated hippocampal function, as well as compromised function and structure of prefrontal and parahippocampal regions.     

心算的加工机制：来自认知神经科学的研究

心算加工分编码（表征）、运算（或提取）和反应三个阶段,这三个阶段相互影响。不同输入形式的数字表征在顶叶的不同区域完成。算术知识提取主要与左脑顶内沟有关,但当心算变得更复杂时而需要具体运算时,左脑额叶下部出现明显激活。所有与心算有关的脑区涉及大脑前额皮层和颞顶枕联合皮层的综合作用,并总体表现为左脑优势,但估算、珠心算以及某些具有特殊心算能力的人的心算还依赖视空间表征,这与右脑额顶区和楔前叶的活动有关     

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.     

重复经颅磁刺激对轻度认知障碍的干预效果

轻度认知障碍(mild cognitive impairment, MCI)是介于正常认知老化和老年痴呆的中间状态, 目前尚无有效的药物治疗方案。重复经颅磁刺激(repetitive transcranial magnetic stimulation, rTMS)可通过诱导突触可塑性的改变来改善大脑的认知功能。对rTMS干预MCI认知功能的有效性及神经机制进行分析。未来研究应优化定位手段, 延长对干预效果的随访评估, 考察不同刺激参数和刺激靶区对干预有效性的影响, 以及结合脑成像技术来探索rTMS的干预机制。     

Brain activity during the encoding, retention, and retrieval of stimulus representations

Studies of delayed nonmatching-to-sample (DNMS) performance following lesions of the monkey cortex have revealed a critical circuit of brain regions involved in forming memories and retaining and retrieving stimulus representations. Using event-related functional magnetic resonance imaging (fMRI), we measured brain activity in 10 healthy human participants during performance of a trial-unique visual DNMS task using novel barcode stimuli. The event-related design enabled the identification of activity during the different phases of the task (encoding, retention, and retrieval). Several brain regions identified by monkey studies as being important for successful DNMS performance showed selective activity during the different phases, including the mediodorsal thalamic nucleus (encoding), ventrolateral prefrontal cortex (retention), and perirhinal cortex (retrieval). Regions showing sustained activity within trials included the ventromedial and dorsal prefrontal cortices and occipital cortex. The present study shows the utility of investigating performance on tasks derived from animal models to assist in the identification of brain regions involved in human recognition memory.     

PET studies of encoding and retrieval: The HERA model

We review positron emission tomography (PET) studies whose results converge on the hemispheric encoding/retrieval asymmetry (HERA) model of the involvement of prefrontal cortical regions in the processes of human memory. The model holds that the left prefrontal cortex is differentially more involved in retrieval of information from semantic memory, and in simultaneously encoding novel aspects of the retrieved information into episodic memory, than is the right prefrontal cortex. The right prefrontal cortex, on the other hand, is differentially more involved in episodic memory retrieval than is the left prefrontal cortex. This general pattern holds for different kinds of information (e.g., verbal materials, pictures, faces) and a variety of conditions of encoding and retrieval.     

Direct comparison of episodic encoding and retrieval of words: an event-related fMRI study

Functional magnetic resonance imaging (fMRI) was used to compare directly episodic encoding and retrieval. During encoding, subjects studied visually presented words and reported via keypress whether each word represented a pleasant or unpleasant concept (intentional, deep encoding). During the retrieval phase, subjects indicated (via keypress) whether visually presented words had previously been studied. No reliable differences were found during the recognition phase for words that had been previously studied and those that had not been studied. Areas preferentially active during encoding (relative to retrieval) included left superior frontal cortex, medial frontal cortex, left superior temporal cortex, posterior cingulate, left parahippocampal gyrus, and left inferior frontal gyrus. Regions more active in retrieval than encoding included bilateral inferior parietal cortex, bilateral precuneus, right frontal polar cortex, right dorsolateral prefrontal cortex, and right inferior frontal/insular cortex.     

Direct Comparison of Episodic Encoding and Retrieval of Words: An Event-related fMRI Study

Functional magnetic resonance imaging (fMRI) was used to compare directly episodic encoding and retrieval. During encoding, subjects studied visually presented words and reported via keypress whether each word represented a pleasant or unpleasant concept (intentional, deep encoding). During the retrieval phase, subjects indicated (via keypress) whether visually presented words had previously been studied. No reliable differences were found during the recognition phase for words that had been previously studied and those that had not been studied. Areas preferentially active during encoding (relative to retrieval) included left superior frontal cortex, medial frontal cortex, left superior temporal cortex, posterior cingulate, left parahippocampal gyrus, and left inferior frontal gyrus. Regions more active in retrieval than encoding included bilateral inferior parietal cortex, bilateral precuneus, right frontal polar cortex, right dorsolateral prefrontal cortex, and right inferior frontal/insular cortex.     

Visual imagery and memory: Do retrieval strategies affect what the mind's eye sees?

A variety of visual mental imagery tasks have been shown to activate regions of visual cortex that subserve the perception of visual events. Here fMRI was used to examine whether imagery‐related visuocortical activity is modulated if imagery content is held constant but there is a change in the memory retrieval strategy used to invoke imagery. Participants were scanned while visualising common objects in two different conditions: (a) recalling recently encoded pictures and (b) based on their knowledge of concrete nouns. Results showed that retrieval‐related activations in frontal cortex were bilateral when pictures were visualised but left‐lateralised when nouns were visualised. In posterior brain regions, both imagery conditions led to activation in the same set of circumscribed areas in left temporal‐parietal cortex, including a region of the left fusiform gyrus that has previously been implicated in visual imagery. These findings suggest that the posterior network activated during imagery did not vary with strategic task‐related changes in the frontal network used to retrieve imagery content from memory.     

Common fronto-parietal activity in attention, memory, and consciousness: shared demands on integration?

Fronto-parietal activity has been frequently observed in fMRI and PET studies of attention, working memory, and episodic memory retrieval. Several recent fMRI studies have also reported fronto-parietal activity during conscious visual perception. A major goal of this review was to assess the degree of anatomical overlap among activation patterns associated with these four functions. A second goal was to shed light on the possible cognitive relationship of processes that relate to common brain activity across functions. For all reviewed functions we observed a consistent and overlapping pattern of brain activity. The overlap was most pronounced for the bilateral parietal cortex (BA 7 and BA 40; close to the intraparietal sulcus), and dorsolateral prefrontal cortex (right BA 9 and left BA 6). The common fronto-parietal activity will be discussed in terms of processes related to integration of distributed representations in the brain.     

Are errors differentiable from deceptive responses when feigning memory impairment? An fMRI study

Previous neuroimaging studies have suggested that the neural activity associated with truthful recall, with false memory, and with feigned memory impairment are different from one another. Here, we report a functional magnetic resonance imaging (fMRI) study that addressed an important but yet unanswered question: Is the neural activity associated with intentional faked responses and with errors differentiable? Using a word list learning recognition paradigm, the findings of this mixed event-related fMRI study clearly indicated that the brain activity associated with intentional faked responses was different to the activity associated with errors committed unintentionally. For intentional faked responses, significant activation was found in the ventrolateral prefrontal cortex, the posterior cingulate region, and the precuneus. However, no significant activation was observed for unintentional errors. The results suggest that deception, in terms of feigning memory impairment, is not only more cognitively demanding than making unintentional errors but also utilizes different cognitive processes.     

The neurodevelopmental basis of math anxiety

Math anxiety is a negative emotional reaction to situations involving mathematical problem solving. Math anxiety has a detrimental impact on an individual's long-term professional success, but its neurodevelopmental origins are unknown. In a functional MRI study on 7- to 9-year-old children, we showed that math anxiety was associated with hyperactivity in right amygdala regions that are important for processing negative emotions. In addition, we found that math anxiety was associated with reduced activity in posterior parietal and dorsolateral prefrontal cortex regions involved in mathematical reasoning. Multivariate classification analysis revealed distinct multivoxel activity patterns, which were independent of overall activation levels in the right amygdala. Furthermore, effective connectivity between the amygdala and ventromedial prefrontal cortex regions that regulate negative emotions was elevated in children with math anxiety. These effects were specific to math anxiety and unrelated to general anxiety, intelligence, working memory, or reading ability. Our study identified the neural correlates of math anxiety for the first time, and our findings have significant implications for its early identification and treatment.