Mapping cognition to the brain through neural interactions

Brain imaging methods, such as positron emission tomography (PET) and functional magnetic resonance imaging (fMRI), provide a unique opportunity to study the neurobiology of human memory. As these methods can measure most of the brain, it is possible to examine the operations of large-scale neural systems and their relation to cognition. Two neuroimaging studies, one concerning working memory and the other episodic memory retrieval, serve as examples of application of two analytic methods that are optimised for the quantification of neural systems, structural equation modelling, and partial least squares. Structural equation modelling was used to explore shifting prefrontal and limbic interactions from the right to the left hemisphere in a delayed match-to-sample task for faces. A feature of the functional network for short delays was strong right hemisphere interactions between hippocampus, inferior prefrontal, and anterior cingulate cortices. At longer delays, these same three areas were strongly linked, but in the left hemisphere, which was interpreted as reflecting change in task strategy from perceptual to elaborate encoding with increasing delay. The primary manipulation in the memory retrieval study was different levels of retrieval success. The partial least squares method was used to determine whether the image-wide pattern of covariances of Brodmann areas 10 and 45/47 in right prefrontal cortex (RPFC) and the left hippocampus (LGH) could be mapped on to retrieval levels. Area 10 and LGH showed an opposite pattern of functional connectivity with a large expanse of bilateral limbic cortices that was equivalent for all levels of retrieval as well as the baseline task. However, only during high retrieval was area 45/47 included in this pattern. The results suggest that activity in portions of the RPFC can reflect either memory retrieval mode or retrieval success depending on other brain regions to which it is functionally linked, and imply that regional activity must be evaluated within the neural context in which it occurs. The general hypothesis that learning and memory are emergent properties of large-scale neural network interactions is discussed, emphasising that a region can play a different role across many functions and that role is governed by its interactions with anatomically related regions.     

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.     

Bi-hemispheric engagement in the retrieval of autobiographical episodes

Functional magnetic resonance imaging (fMRI) was used to study the neural correlates of neutral, stressful, negative and positive autobiographical memories. The brain activity produced by these different kinds of episodic memory did not differ significantly, but a common pattern of activation for different kinds of autobiographical memory was revealed that included (1) largely bilateral portions of the medial and superior temporal lobes, hippocampus and parahippocampus, (2) portions of the ventral, medial, superior and dorsolateral prefrontal cortex, (3) the anterior and posterior cingulate, including the retrosplenial, cortex, (4) the parietal cortex, and (5) portions of the cerebellum. The brain regions that were mainly activated constituted an interactive network of temporal and prefrontal areas associated with structures of the extended limbic system. The main bilateral activations with left-sided preponderance probably reflected reactivation of complex semantic and episodic self-related information representations that included previously experienced contexts. In conclusion, the earlier view of a strict left versus right prefrontal laterality in the retrieval of semantic as opposed to episodic autobiographical memory, may have to be modified by considering contextual variables such as task demands and subject variables. Consequently, autobiographical memory integration should be viewed as based on distributed bi-hemispheric neural networks supporting multi-modal, emotionally coloured components of personal episodes.     

Right amygdalar and temporofrontal activation during autobiographic, but not during fictitious memory retrieval

What distinguishes the recall of real-life experiences from that of self-created, fictitious emotionally laden information? Both kinds of information belong to the episodic memory system. Autobiographic memories constitute that part of the episodic memory system that is composed of significant life episodes, primarily of the distant past. Functional imaging was used to study the neural networks engaged in retrieving autobiographic and fictitious information of closely similar content. The principally activated brain regions overlapped considerably and constituted temporal and inferior prefrontal regions plus the cerebellum. Selective activations of the right amygdala and the right ventral prefrontal cortex (at the level of the uncinate fascicle interconnnecting prefrontal and temporopolar areas) were found when subtracting fictitious from autobiographic retrieval. Furthermore, distinct foci in the left temporal lobe were engaged. These data demonstrate that autobiographic memory retrieval uses (at least in non-brain damaged individuals) a network of right hemispheric ventral prefrontal and temporopolar regions and left hemispheric lateral temporal regions. It is concluded that it is the experiential character, its special emotional infiltration and its arousal which distinguishes memory of real-life from that of fictitious episodes. Consequently, our results point to the engagement of a bi-hemispheric network in which the right temporo-prefrontal hemisphere is likely to be responsible for the affective/arousal side of information retrieval and the left-hemispheric temporal gyrus for its engram-like representation. Portions of the neural activation found during retrieval might, however, reflect re-encoding processes as well.     

Incidental retrieval of emotional contexts in post-traumatic stress disorder and depression: an fMRI study

In the present study, we used fMRI to assess patients suffering from post-traumatic stress disorder (PTSD) or depression, and trauma-exposed controls, during an episodic memory retrieval task that included non-trauma-related emotional information. In the study phase of the task neutral pictures were presented in emotional or neutral contexts. Participants were scanned during the test phase, when they were presented with old and new neutral images in a yes/no recognition memory task. fMRI results for the contrast between old and new items revealed activation in a predominantly left-sided network of cortical regions including the left middle temporal, bilateral posterior cingulate, and left prefrontal cortices. Activity common to all three groups when correctly judging pictures encoded in emotional contexts was much more limited. Relative to the control and depressed groups the PTSD group exhibited greater sensitivity to correctly recognised stimuli in the left amygdala/ventral striatum and right occipital cortex, and more specific sensitivity to items encoded in emotional contexts in the right precuneus, left superior frontal gyrus, and bilateral insula. These results are consistent with a substantially intact neural system supporting episodic retrieval in patients suffering from PTSD. Moreover, there was little indication that PTSD is associated with a marked change in the way negatively valenced information, not of personal significance, is processed.     

Adult age differences in visual word identification: functional neuroanatomy by positron emission tomography

Adult age differences in the neural systems mediating semantic (context-independent) memory were investigated using positron emission tomography (PET). Younger (20-29 years) and older (62-70 years) participants performed lexical decision (word/nonword discrimination) and nonsemantic (simple visual search) baseline tasks during PET scanning. Within the lexical decision task, display duration and presentation rate were varied across scans. The behavioral data suggested that although an age-related slowing was evident in visual feature and response processing, the retrieval of semantic/lexical information was similar for younger and older adults. For both age groups, lexical-related activation occurred in inferior prefrontal and occipitotemporal regions of the left hemisphere. Differential activation, as a function of age group, was observed in the left occipitotemporal pathway as a result of older adults' maintaining higher levels of neural activity in striate cortex (during visual search) and in inferior temporal cortex (during lexical decision). The prefrontal activation was similar for the two age groups. Thus, although this form of semantic memory retrieval does not undergo significant age-related decline, an age-related change in the associated pattern of neural activation is evident. These findings differ from previous neuroimaging studies of episodic (context-dependent) memory retrieval, which have suggested that age-related compensatory mechanisms are expressed primarily by greater activation of prefrontal regions for older adults than for younger adults.     

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 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.     

Functional asymmetry of human prefrontal cortex: encoding and retrieval of verbally and nonverbally coded information

There are several views about the organization of memory functions in the human prefrontal cortex. One view assumes a process-specific brain lateralization according to different memory subprocesses, that is, encoding and retrieval. An alternative view emphasizes content-specific lateralization of brain systems involved in memory processes. This study addresses this apparent inconsistency between process- and content-specific lateralization of brain activity by investigating the effects of verbal and nonverbal encoding on prefrontal activations during encoding and retrieval of environmental novel sounds using fMRI. An intentional memory task was applied in which subjects were required either to judge the sounds' loudness (nonverbal encoding task) or to indicate whether or not a sound can be verbally described (verbal encoding task). Retrieval processes were examined in a subsequent yes/no recognition test. In the study phase the right posterior dorsolateral prefrontal cortex (PFC) was activated in both tasks. During verbal encoding additional activation of the left dorsolateral PFC was obtained. Retrieval-related fMRI activity varied as a function of encoding task: For the nonverbal task we detected an activation focus in the right posterior dorsolateral PFC whereas an activation in the left dorsolateral PFC was observed for the verbal task. These findings indicate that the right dorsolateral PFC is engaged in encoding of auditory information irrespective of encoding task. The lateralization of PFC activity during retrieval was shown to depend on the availability of verbal codes, with left hemispheric involvement for verbally and right hemispheric activation for nonverbally coded information.     

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.     

人类工作记忆的某些神经影像研究

采用神经影像技术研究人类工作记忆的脑机理是目前一个十分活跃的研究领域。研究表明存在负责不同信息加工的工作记忆系统 ,如词语工作记忆、空间工作记忆等 ,其中词语工作记忆主要由大脑左半球参与 ,空间工作记忆主要由大脑右半球参与。前额叶在工作记忆中的作用相当复杂 ,包括对记忆信息的注意和抑制、管理、整合等功能。合理巧妙的实验设计、多种研究手段的综合应用必将使人类工作记忆的脑机理得到充分阐明。     

ERPs associated with visual duration discriminations in prefrontal and parietal cortex

An event-related potentials (ERP) study was undertaken to examine the role of prefrontal and parietal association cortices on selective attention and short-term memory functions in a duration discrimination task. Subjects performed better when discriminating the first stimulus relative to the second and not the reverse. Two contingent negative variations (CNV) were obtained for each stimulus duration at prefrontal regions, as well as two P300s at parietal regions. The CNV(S1) component recorded during the first stimulus (S1) appeared to be involved in selective attention at bilateral sites, while the P300(S1) component in the left hemisphere may be implicated in retaining it. The CNV(S2) wave, displayed during the second stimulus (S2), at bilateral sites and the right-sided P300(S2) wave seem to be implicated in working memory. The results indicate that recorded activity at prefrontal and parietal association cortices is tightly linked to task parameters and behavioral performances.     

The neural basis of autobiographical and semantic memory: new evidence from three PET studies

A novel, neuropsychologically informed paradigm (extended retrieval of events in response to a cue word) was used to investigate the neural basis of autobiographical and semantic memory. Contrasting retrieval of autobiographical memories with retrieval of semantic facts (ABM-SEM) in 24 subjects across three PET studies revealed bilateral involvement of the middle temporal gyrus (BA 21) and medial frontal cortex (BA 9/10). The opposite contrast, SEM-ABM, resulted in increased regional cerebral blood flow in left posterior temporal regions (BA 37) and left prefrontal cortex (BA 45/46). Laterality maps suggest that the bilateral pattern seen in our studies, but not often in other neuroimaging investigations, reflects the use of a task stressing retrieval of specific personal events. Further comparisons revealed that the activation in the right anterior temporal lobe during autobiographical recall was virtually identical to that seen during retrieval of information about famous people or events in contrast with retrieval of general semantic facts. These findings suggest that the retrieval of an autobiographical event requires participation from conceptual knowledge, and that this type of knowledge is bilaterally distributed in the temporal lobes.     

Developmental shifts in fMRI activations during visuospatial relational reasoning

To investigate maturational plasticity of fluid cognition systems, functional brain imaging was undertaken in healthy 8-19 year old participants while completing visuospatial relational reasoning problems similar to Raven's matrices and current elementary grade math textbooks. Analyses revealed that visuospatial relational reasoning across this developmental age range recruited activations in the superior parietal cortices most prominently, the dorsolateral prefrontal, occipital-temporal, and premotor/supplementary cortices, the basal ganglia, and insula. There were comparable activity volumes in left and right hemispheres for nearly all of these regions. Regression analyses indicated increasing activity predominantly in the superior parietal lobes with developmental age. In contrast, multiple anterior neural systems showed significantly less activity with age, including dorsolateral and ventrolateral prefrontal, paracentral, and insula cortices bilaterally, basal ganglia, and particularly large clusters in the midline anterior cingulate/medial frontal cortex, left middle cingulate/supplementary motor cortex, left insula-putamen, and left caudate. Findings suggest that neuromaturational changes associated with visuospatial relational reasoning shift from a more widespread fronto-cingulate-striatal pattern in childhood to predominant parieto-frontal activation pattern in late adolescence.     

Recognizing one's own face

We report two studies of facial self-perception using individually tailored, standardized facial photographs of a group of volunteers and their partners. A computerized morphing procedure was used to merge each target face with an unknown control face. In the first set of experiments, a discrimination task revealed a delayed response time for the more extensively morphed self-face stimuli. In a second set of experiments, functional magnetic resonance imaging (fMRI) was used to measure brain activation while subjects viewed morphed versions of either their own or their partner's face, alternating in blocks with presentation of an unknown face. When subjects viewed themselves (minus activation for viewing an unknown face), increased blood oxygenation was detected in right limbic (hippocampal formation, insula, anterior cingulate), left prefrontal cortex and superior temporal cortex. In the partner (versus unknown) experiment, only the right insula was activated. We suggest that a neural network involving the right hemisphere in conjunction with left-sided associative and executive regions underlies the process of visual self-recognition. Together, this combination produces the unique experience of self-awareness.     

rTMS for PTSD: induced merciful oblivion or elimination of abnormal hypermnesia?

Neuroimaging studies and experimental data suggest that symptoms of posttraumatic stress disorder (PTSD) are associated with dysfunctions of neural circuits linking prefrontal cortex and the limbic system that have a role in autobiographic episodic memory. High-frequency repetitive transcranial magnetic stimulation (rTMS) of the right dorsolateral prefrontal cortex (DLPFC) has been suggested to be beneficial to patients with PTSD, transiently alleviating re-experiencing as well as avoidance reactions and associated anxiety symptoms. In healthy humans, converging evidence suggests that rTMS of the right DLPFC interferes with episodic memory retrieval. Hence, we hypothesize that daily applications of rTMS in PTSD patients may reduce access to the set of autobiographical stored events, that, if re-experienced, may cause the overt PTSD symptoms.     

青老年组不同难度下心算活动的脑功能磁共振成像研究

应用功能磁共振成像技术研究不同心算难度下脑区的活动以及年龄的影响。14名志愿者（20～29岁青年和60～69岁老年被试各7名）参加了该实验。实验任务为2个难度水平的连续减法心算,分别为1000—3和1000—17。结果表明：（1）心算加工激活了额叶和顶叶的许多脑区;（2）大脑左半球是心算加工的优势半球,但随着心算难度加大,大脑一侧化程度下降,而年老加剧了这一趋势;（3）青年组进行简单心算（1000—3）时,额中回未见明显激活,而老年组进行简单心算时,该脑区被明显激活。总体上,额叶和顶叶在心算活动中起着重要作用,而任务难度和年龄对心算加工时脑活动的影响以额中回区最为明显。     

Hemispheric asymmetries in the activation and monitoring of memory errors

Previous research on the lateralization of memory errors suggests that the right hemisphere's tendency to produce more memory errors than the left hemisphere reflects hemispheric differences in semantic activation. However, all prior research that has examined the lateralization of memory errors has used self-paced recognition judgments. Because activation occurs early in memory retrieval, with more time to make a decision, other memory processes, like strategic monitoring processes, may affect memory errors. By manipulating the time subjects were given to make memory decisions, this study separated the influence of automatic memory processes (activation) from strategic memory processes (monitoring) on the production of false memories. The results indicated that when retrieval was fast, the right hemisphere produced more memory errors than the left hemisphere. However, when retrieval was slow, the left hemisphere's error-proneness increased compared to the fast retrieval condition, while the right hemisphere's error-proneness remained the same. These results suggest that the right hemisphere's errors are largely due to activation, while the left hemisphere's errors are influenced by both activation and monitoring.     

Learning in the visual association of novice and expert designers

Designers are adept at determining similarities between previously seen objects and new creations using visual association. However, extant research on the visual association of designers and the differences between expert and novice designers when they engage in the visual association task are scant. Using electroencephalography (EEG), this study attempted to narrow this research gap. Sixteen healthy designers—eight experts and eight novices—were recruited, and asked to perform visual association while EEG signals were acquired, subsequently analysed using independent component analysis. The results indicated that strong connectivity was observed among the prefrontal, frontal, and cingulate cortices, and the default mode network. The experts used both hemispheres and executive functions to support their association tasks, whereas the novices mainly used their right hemisphere and memory retrieval functions. The visual association of experts appeared to be more goal-directed than that of the novices. Accordingly, designing and implementing authentic and goal-directed activities for improving the executive functions of the prefrontal cortex and default mode network are critical for design educators and creativity researchers.     

Activation of right parietal cortex during memory retrieval of nonlinguistic auditory stimuli

In neuroimaging studies, the left ventral posterior parietal cortex (PPC) is particularly active during memory retrieval. However, most studies have used verbal or verbalizable stimuli. We investigated neural activations associated with the retrieval of short, agrammatical music stimuli (Blackwood, 2004), which have been largely associated with right hemisphere processing. At study, participants listened to music stimuli and rated them on pleasantness. At test, participants made old/new recognition judgments with high/low confidence ratings. Right, but not left, ventral PPC activity was observed during the retrieval of these music stimuli. Thus, rather than indicating a special status of left PPC in retrieval, both right and left ventral PPC participate in memory retrieval, depending on the type of information that is to be remembered.