Context effects recognition of the emotional facial expression

Influence of additional working memory load on emotional face recognition was studied in healthy adults. Visual set to emotional face expression was experimentally formed, and two types of additional task--visual-spatial or semantic--were embedded in the experiment. Additional task caused less plastic set, i.e., a slower set-shifting. This effect displayed itself in an increase of erroneous facial expression perceptions. The character of these erroneous perceptions (assimilative or contrast or visual illusions) depended on the type of the additional task. Pre-stimulus EEG coherence across experimental trials in theta (4-7), low alpha (8-10 Hz) and beta (14--20) bands was analysed. Data of low-alpha and beta-coherence supported the hypothesis that increased memory load caused less involvement of frontal lobes in selective attention mechanisms that are associated with set-forming. This results in a slower set-shifting. Increased memory load also led to a growth of theta-band coherence in the left hemisphere and its decrease in the right hemisphere. The account of theta-coherence decrease in the right hemisphere between prefrontal and temporal areas for a slower set-shifting is discussed.     

Attentional set-shifting deficit in Parkinson's disease is associated with prefrontal dysfunction: an FDG-PET study

The attentional set-shifting deficit that has been observed in Parkinson's disease (PD) has long been considered neuropsychological evidence of the involvement of meso-prefrontal and prefrontal-striatal circuits in cognitive flexibility. However, recent studies have suggested that non-dopaminergic, posterior cortical pathologies may also contribute to this deficit. Although several neuroimaging studies have addressed this issue, the results of these studies were confounded by the use of tasks that required other cognitive processes in addition to set-shifting, such as rule learning and working memory. In this study, we attempted to identify the neural correlates of the attentional set-shifting deficit in PD using a compound letter task and 18F-fluoro-deoxy-glucose (FDG) positron emission tomography during rest. Shift cost, which is a measure of attentional set-shifting ability, was significantly correlated with hypometabolism in the right dorsolateral prefrontal cortex, including the putative human frontal eye field. Our results provide direct evidence that dysfunction in the dorsolateral prefrontal cortex makes a primary contribution to the attentional set-shifting deficit that has been observed in PD patients.     

Social Suppressive Behavior Is Organized by the Spatiotemporal Integration of Multiple Cortical Regions in the Japanese Macaque

Under social conflict, monkeys develop hierarchical positions through social interactions. Once the hierarchy is established, the dominant monkey dominates the space around itself and the submissive monkey tries not to violate this space. Previous studies have shown the contributions of the frontal and parietal cortices in social suppression, but the contributions of other cortical areas to suppressive functions remain elusive. We recorded neural activity in large cortical areas using electrocorticographic (ECoG) arrays while monkeys performed a social food-grab task in which a target monkey was paired with either a dominant or a submissive monkey. If the paired monkey was dominant, the target monkey avoided taking food in the shared conflict space, but not in other areas. By contrast, when the paired monkey was submissive, the target monkey took the food freely without hesitation. We applied decoding analysis to the ECoG data to see when and which cortical areas contribute to social behavioral suppression. Neural information discriminating the social condition was more evident when the conflict space was set in the area contralateral to the recording hemisphere. We found that the information increased as the social pressure increased during the task. Before food presentation, when the pressure was relatively low, the parietal and somatosensory–motor cortices showed sustained discrimination of the social condition. After food presentation, when the monkey faced greater pressure to make a decision as to whether it should take the food, the prefrontal and visual cortices started to develop buildup responses. The social representation was found in a sustained form in the parietal and somatosensory–motor regions, followed by additional buildup form in the visual and prefrontal cortices. The representation was less influenced by reward expectation. These findings suggest that social adaptation is achieved by a higher-order self-regulation process (incorporating motor preparation/execution processes) in accordance with the embodied social contexts.     

第一和第二语言Stroop任务中EEG同步化分析

采用基于多元自回归的瞬时EEG相干方法研究了十位汉英双语者执行Stroop任务时脑神经电活动及其功能皮层区的协同作用。结果显示：在β1(13-18Hz)频段,无论是汉语(第一语言,L1)还是英语(第二语言,L2)呈现的刺激,不一致条件的EEG相干值明显大于一致条件的EEG相干值,表明β1频段对刺激类型敏感;与L2相比,L1的Stroop任务中,额一顶区的相干值显著增强。EEG相干值反映了不同脑皮层间的相互作用强度。因此研究结果表明：判断和处理冲突信息(如Stroop的不一致条件)时脑功能皮层区之间的协同作用增强;相对于第二语言,第一语言处理过程中额一顶区之间的通信协作增加。     

An Event-Related fMRI Study of Phonological Verbal Working Memory in Schizophrenia

Background

While much is known about the role of prefrontal cortex (PFC) in working memory (WM) deficits of schizophrenia, the nature of the relationship between cognitive components of WM and brain activation patterns remains unclear. We aimed to elucidate the neural correlates of the maintenance component of verbal WM by examining correct and error trials with event-related fMRI.

Methodology/Findings

Twelve schizophrenia patients (SZ) and thirteen healthy control participants (CO) performed a phonological delayed-matching-to-sample-task in which a memory set of three nonsense words was presented, followed by a 6-seconds delay after which a probe nonsense word appeared. Participants decided whether the probe matched one of the targets, and rated the confidence of their decision. Blood-oxygen-level-dependent (BOLD) activity during WM maintenance was analyzed in relation to performance (correct/error) and confidence ratings. Frontal and parietal regions exhibited increased activation on correct trials for both groups. Correct and error trials were further segregated into true memory, false memory, guess, and true error trials. True memory trials were associated with increased bilateral activation of frontal and parietal regions in both groups but only CO showed deactivation in PFC. There was very little maintenance-related cortical activity during guess trials. False memory was associated with increased left frontal and parietal activation in both groups.

Conclusion

These findings suggest that a wider network of frontal and parietal regions support WM maintenance in correct trials compared with error trials in both groups. Furthermore, a more extensive and dynamic pattern of recruitment of the frontal and parietal networks for true memory was observed in healthy controls compared with schizophrenia patients. These results underscore the value of parsing the sources of memory errors in fMRI studies because of the non-linear nature of the brain-behavior relationship, and suggest that group comparisons need to be interpreted in more specific behavioral contexts.     

Origins of spatial working memory deficits in schizophrenia: an event-related FMRI and near-infrared spectroscopy study

Abnormal prefrontal functioning plays a central role in the working memory (WM) deficits of schizophrenic patients, but the nature of the relationship between WM and prefrontal activation remains undetermined. Using two functional neuroimaging methods, we investigated the neural correlates of remembering and forgetting in schizophrenic and healthy participants. We focused on the brain activation during WM maintenance phase with event-related functional magnetic resonance imaging (fMRI). We also examined oxygenated hemoglobin changes in relation to memory performance with the near-infrared spectroscopy (NIRS) using the same spatial WM task. Distinct types of correct and error trials were segregated for analysis. fMRI data indicated that prefrontal activation was increased during WM maintenance on correct trials in both schizophrenic and healthy subjects. However, a significant difference was observed in the functional asymmetry of frontal activation pattern. Healthy subjects showed increased activation in the right frontal, temporal and cingulate regions. Schizophrenic patients showed greater activation compared with control subjects in left frontal, temporal and parietal regions as well as in right frontal regions. We also observed increased 'false memory' errors in schizophrenic patients, associated with increased prefrontal activation and resembling the activation pattern observed on the correct trials. NIRS data replicated the fMRI results. Thus, increased frontal activity was correlated with the accuracy of WM in both healthy control and schizophrenic participants. The major difference between the two groups concerned functional asymmetry; healthy subjects recruited right frontal regions during spatial WM maintenance whereas schizophrenic subjects recruited a wider network in both hemispheres to achieve the same level of memory performance. Increased "false memory" errors and accompanying bilateral prefrontal activation in schizophrenia suggest that the etiology of memory errors must be considered when comparing group performances. Finally, the concordance of fMRI and NIRS data supports NIRS as an alternative functional neuroimaging method for psychiatric research.     

Functional alterations in neural substrates of geometric reasoning in adults with high-functioning autism

Individuals with autism spectrum condition (ASC) are known to excel in some perceptual cognitive tasks, but such developed functions have been often regarded as "islets of abilities" that do not significantly contribute to broader intellectual capacities. However, recent behavioral studies have reported that individuals with ASC have advantages for performing Raven's (Standard) Progressive Matrices (RPM/RSPM), a standard neuropsychological test for general fluid intelligence, raising the possibility that ASC's cognitive strength can be utilized for more general purposes like novel problem solving. Here, the brain activity of 25 adults with high-functioning ASC and 26 matched normal controls (NC) was measured using functional magnetic resonance imaging (fMRI) to examine neural substrates of geometric reasoning during the engagement of a modified version of the RSPM test. Among the frontal and parietal brain regions involved in fluid intelligence, ASC showed larger activation in the left lateral occipitotemporal cortex (LOTC) during an analytic condition with moderate difficulty than NC. Activation in the left LOTC and ventrolateral prefrontal cortex (VLPFC) increased with task difficulty in NC, whereas such modulation of activity was absent in ASC. Furthermore, functional connectivity analysis revealed a significant reduction of activation coupling between the left inferior parietal cortex and the right anterior prefrontal cortex during both figural and analytic conditions in ASC. These results indicate altered pattern of functional specialization and integration in the neural system for geometric reasoning in ASC, which may explain its atypical cognitive pattern, including performance on the Raven's Matrices test.     

EEG correlates of individual differences in performance efficiency of students during examination stress

Performance of cognitive tests and EEG spectral power were evaluated in 39 students aged from 19-21 years in two conditions: during common educational process and immediately before examination (stress condition). Before examination, the performance was better in subjects with low level of spectral density in the delta band (in the occipital, parietal, central and frontal cortical areas) and high level of the alpha-rhythm spectral density in all the cortical areas, A decrease in performance scores before examination was correlated with an increase in the delta activity (in the right frontal and temporal cortical areas) and rise of the anxiety level (tested by Spielberger).     

An Operant Intra-/Extra-dimensional Set-shift Task for Mice

Alterations in executive control and cognitive flexibility, such as attentional set-shifting abilities, are core features of several neuropsychiatric diseases. The most widely used neuropsychological tests for the evaluation of attentional set-shifting in human subjects are the Wisconsin Card Sorting Test (WCST) and the CANTAB Intra-/Extra-dimensional set shift task (ID/ED). These tasks have proven clinical relevance and have been modified and successfully adapted for research in animal models. However, currently available tasks for rodents present several limitations, mainly due to their manual-based testing procedures, which are hampering translational advances in psychiatric medicine. To overcome these limitations and to better mimic the original version in primates, we present the development of a novel operant-based two-chamber ID/ED "Operon" task for rodents. We demonstrated the effectiveness of this novel task to measure different facets of cognitive flexibility in mice including attentional set formation and shifting, and reversal learning. Moreover, we show the high flexibility of this task in which three different perceptual dimensions can be manipulated with a high number of stimuli cues for each dimension. This novel ID/ED Operon task can be an effective preclinical tool for drug testing and/or large genetic screening relevant to the study of executive dysfunction and cognitive symptoms found in psychiatric disorders.     

The mapping of brain biopotentials during the solving of a verbal task]

Reverse averaging of cortical potentials from the moment of the motor response followed the verbal task solving (anagram riddle) revealed some brain potentials correlations with the process of a decision making. In the case of task solving the negative frontal wave with the latency 900-400 ms from the motor response was recorded. Intracortical interaction mapping of this potential showed the regular patterns of cortical functional connections in different frequency ranges (alpha, beta). Successful solving of the task was characterized with predominant interaction foci topography in the frontal and left-temporal cortical areas in alpha band and parietal zones in beta. The absence of the task solution was characterized with the parieto-occipital interaction foci in alpha band and their frontal localization in beta.     

Changes in emotional face expression recognition caused by an additional visuospatial task

Changes in the recognition of facial expression and spatial synchronization of the cortical electrical activity of the θ- and α-potentials caused by load on working memory were studied in healthy adults by introducing an additional semantic or visuospatial task into the context of experiment with a visual set. An increase in the number of erroneous recognitions of facial stimuli in the form of assimilative illusions was revealed in both types of the additional task. The analysis of the function of coherence of the low-frequency α-potentials indicates (8–10 Hz) a decrease in this situation in the number of connections in the frontal cortical divisions with other cortical zones, which is regarded as a lesser involvement of the frontal system of selective attention in set-forming and set-shifting for an emotionally negative facial expression. Spatial synchronization of the θ-activity (4–7 Hz) with an increase in the load on working memory changes ambiguously in different cortical structures: it decreases in the system of the fronto-temporal connections of the right hemisphere; in the other cortical areas, especially in the left hemisphere, and in the system of interhemispheric connections it substantially increases. The facts confirming the hypothesis that the fronto-thalamic and cortico-hippocampal systems are the two key formations involved in changes in the plasticity of cognitive sets for facial expression are discussed.     

Cortical functional connectivity during the retention of affective pictures in working memory: EEG-source theta coherence analysis

The pattern of cortical functional connectivity in the source space was studied in a group of righthanded adult participants (N = 44:17 women, 27 men, aged M = 29.61 ± 6.45 years). Participants retained the traces of realistic pictures of positive, neutral, and negative emotional valences in their working memory (WM) while performing the same-different task. Within the framework of this task, participants had to compare the initial picture against a target picture that followed after a specified delay. The coherence (COH) between the pairs of cortical sources chosen in advance according to fMRI data was estimated in the theta frequency range for the period preceding the initial stimulus, during the retention of the initial stimulus in WM, and during the rest interval between successive trials. Two distinct sets of functional links were found. The links of the first type that presumably reflected the involvement of sustained attention were between the dorsal anterior cingulate cortex, the prefrontal areas, and temporal areas of the right hemispheres. When compared to the rest period, the links of this type showed strengthening not only during the retention period but also during the period preceding the initial picture. The links of the second type presumably reflected a progressive neocortex-to-hippocampus functional integration with increasing memory load and strengthened exclusively during the retention period. These links were between the parietal, temporal and prefrontal cortices in the lateral surface of both hemispheres with the additional inclusion of the posterior cingulate cortex and the medial parietal cortex in the left hemisphere. The impact of emotional valence on the strength and topography of the functional links of the second type was found. In the left hemisphere, the increase of strength of cortical interaction was more pronounced for the pictures of positive valence than for the pictures of either neutral or negative valences. When compared to the pictures of neutral valence, the retention of pictorial information of both positive and negative valence showed some extraneous integration of the cortical areas for the theta rhythm. This finding might be related to the additional load exerted by emotionally colored pictures onto the mechanisms of short-time retention of visual information.     

Topography of the chimpanzee corpus callosum

The corpus callosum (CC) is the largest commissural white matter tract in mammalian brains, connecting homotopic and heterotopic regions of the cerebral cortex. Knowledge of the distribution of callosal fibers projecting into specific cortical regions has important implications for understanding the evolution of lateralized structures and functions of the cerebral cortex. No comparisons of CC topography in humans and great apes have yet been conducted. We investigated the topography of the CC in 21 chimpanzees using high-resolution magnetic resonance imaging (MRI) and diffusion tensor imaging (DTI). Tractography was conducted based on fiber assignment by continuous tracking (FACT) algorithm. We expected chimpanzees to display topographical organization similar to humans, especially concerning projections into the frontal cortical regions. Similar to recent studies in humans, tractography identified five clusters of CC fibers projecting into defined cortical regions: prefrontal; premotor and supplementary motor; motor; sensory; parietal, temporal and occipital. Significant differences in fractional anisotropy (FA) were found in callosal regions, with highest FA values in regions projecting to higher-association areas of posterior cortical (including parietal, temporal and occipital cortices) and prefrontal cortical regions (p<0.001). The lowest FA values were seen in regions projecting into motor and sensory cortical areas. Our results indicate chimpanzees display similar topography of the CC as humans, in terms of distribution of callosal projections and microstructure of fibers as determined by anisotropy measures.     

The importance of timing in segregated theta phase-coupling for cognitive performance

Functional cortical circuits for central executive functions have been shown to emerge by theta (~6 Hz) phase-coupling of distant cortical areas. It has been repeatedly shown that frontoparietal theta coupling at ~0° relative phase is associated with recognition, encoding, short-term retention, and planning; however, a causal link has not been demonstrated so far. Here we used transcranial alternating current stimulation simultaneously applied at 6 Hz over left prefrontal and parietal cortices with a relative 0° ("synchronized" condition) or 180° ("desynchronized" condition) phase difference or a placebo stimulation condition, whereas healthy subjects performed a delayed letter discrimination task. We show that exogenously induced frontoparietal theta synchronization significantly improves visual memory-matching reaction times as compared to placebo stimulation. In contrast, exogenously induced frontoparietal theta desynchronization deteriorates performance. The present findings provide for the first time evidence of causality of theta phase-coupling of distant cortical areas for cognitive performance in healthy humans. Moreover, the results demonstrate the suitability of transcranial alternating current stimulation to artificially induce coupling or decoupling of behaviorally relevant brain rhythms between segregated cortical regions.     

A preliminary study suggests that nicotine and prefrontal dopamine affect cortico‐striatal areas in smokers with performance feedback

Nicotine and tonic dopamine (DA) levels [as inferred by catechol‐O‐methyl tranferase (COMT) Val158Met genotype] interact to affect prefrontal processing. Prefrontal cortical areas are involved in response to performance feedback, which is impaired in smokers. We investigated whether there is a nicotine × COMT genotype interaction in brain circuitry during performance feedback of a reward task. We scanned 23 healthy smokers (10 Val/Val homozygotes, 13 Met allele carriers) during two fMRI sessions while subjects were wearing a nicotine or placebo patch. A significant nicotine × COMT genotype interaction for BOLD signal during performance feedback in cortico‐striatal areas was seen. Activation in these areas during the nicotine patch condition was greater in Val/Val homozygotes and reduced in Met allele carriers. During negative performance feedback, the change in activation in error detection areas such as anterior cingulate cortex (ACC)/superior frontal gyrus on nicotine compared to placebo was greater in Val/Val homozygotes compared to Met allele carriers. With transdermal nicotine administration, Val/Val homozygotes showed greater activation with performance feedback in the dorsal striatum, area associated with habitual responding. In response to negative feedback, Val/Val homozygotes had greater activation in error detection areas, including the ACC, suggesting increased sensitivity to loss with nicotine exposure. Although these results are preliminary due to small sample size, they suggest a possible neurobiological mechanism underlying the clinical observation that Val/Val homozygotes, presumably with elevated COMT activity compared to Met allele carriers and therefore reduced prefrontal DA levels, have poorer outcomes with nicotine replacement therapy .     

Temporal and topographic characteristics of evoked potentials in the conflict of two consecutive visual stimuli in a working memory task

Behavioral reactions and brain mechanisms involved in processing two matching or mismatching (conflicting) visual stimuli were studied in healthy subjects (mean age 22.57 ± 0.46 years). Line orientations (vertical, horizontal, or 45°) were used as stimuli and were presented with an interval of 1500–1800 ms. The reaction time was shown to increase in the case of a conflict of two orientations as compared with matching orientations. The reaction time depended on the orientation of the reference stimulus and was minimal when a vertical line was used as a reference. An increase in N2 negativity (time window 200–280 ms) in the frontal and parietal cortical areas was identified as an informative indicator of a conflict between the current orientation and the orientation stored in working memory. The dipole sources of N2 were localized to the prefrontal cortex (middle frontal gyrus, frontal pole, and pars orbitalis). The N2 amplitude was found to depend on the orientation of the first stimulus in a pair, being higher in the case of a 45° orientation. The visual areas were shown to play a role in detecting a conflict of two consecutive signals because the early sensory components increased in amplitude. The results implicate cortical structures, including the sensory-specific visual, parietal, and prefrontal areas, in comparing consecutive visual signals and detecting their conflict.     

Interaction between the Frontal and Left Parietotemporal Areas in Verbal Thinking

The synchronization of the rhythmical components of evoked potentials (EP) was studied during verbal-task solving. A novel method of the calculation of Wavelet curve correlation was used to reveal synchronization between the evoked rhythmical components in short time intervals. This method was applied to earlier EP records, which were conducted during the search for verbal associations and revealed the successive activation of the frontal and left parietal cortical areas. Two stages of task solving were identified. Independently of the task type, the first stage was characterized by a diffuse synchronization in a broad frequency band below 22 Hz immediately after the stimulus presentation. This stage results in a realization of the verbal stimulus. The second stage was manifested in a localized synchronization between the frontal and left temporal (Wernicke's) areas in the narrow frequency band about 17 Hz only during search for associations. This specific and local synchronization took place earlier than the diffuse activation of the left temporal cortex. This stage appears to reflect the information transmission from the frontal cortex to the left parietotemporal area.     

参与不同难度数字计算的脑区——脑功能磁共振成像研究

本研究用功能磁共振成像技术观察了人脑进行不同难度数字加减计算时的脑区激活情况,并探讨大脑皮层和皮层下结构在数字计算中的作用.用Siemens 1.5 Tesla磁共振机对16名右利手健康志愿者进行简单及复杂数字加减任务的fMRI扫描.实验采用组块设计.刺激任务分为简单加减计算任务、复杂加减计算任务和基线任务.用SPM99软件进行数据分析和脑功能区定位.分别比较同一任务各个脑区平均激活强度和同一脑区在两种任务中的激活强度.结果显示,简单及复杂加减计算激活的被试者的脑区基本相同,激活的皮层区主要见于额叶、顶叶、枕叶、扣带回、丘脑及小脑;简单及复杂加减计算激活的皮层下结构包括两侧尾状核、左纹状体边缘区等基底核结构和丘脑.在简单及复杂计算中,纹状体与皮质结构(额叶、顶叶)间激活强度均无显著性差异.简单计算与复杂计算比较,右顶叶,在复杂任务时出现激活,在简单任务时未出现激活.上述结果提示,完成数字计算任务的脑区除了额叶、顶叶、扣带回等皮层结构外,大脑皮层下的一些结构如纹状体、纹状体边缘区,也是参与数字计算的重要部位.皮层下结构纹状体和优势半球的纹状体边缘区参与了数字工作记忆,可能是进行数字计算神经环路的重要组成部位.右项叶(缘上回)只在复杂任务出现激活,该区可能是视空间记忆和加工的重要部位.     

The brain network of expectancy and uncertainty processing

Background

The Stimulus Preceding Negativity (SPN) is a non-motor slow cortical potential elicited by temporally predictable stimuli, customarily interpreted as a physiological index of expectancy. Its origin would be the brain activity responsible for generating the anticipatory mental representation of an expected upcoming event. The SPN manifests itself as a slow cortical potential with negative slope, growing in amplitude as the stimulus approximates. The uncertainty hypothesis we present here postulates that the SPN is linked to control-related areas in the prefrontal cortex that become more active before the occurrence of an upcoming outcome perceived as uncertain.

Methods/Findings

We tested the uncertainty hypothesis by using a repeated measures design in a Human Contingency Learning task with two levels of uncertainty. In the high uncertainty condition, the outcome is unpredictable. In the mid uncertainty condition, the outcome can be learnt to be predicted in 75% of the trials. Our experiment shows that the Stimulus Preceding Negativity is larger for probabilistically unpredictable (uncertain) outcomes than for probabilistically predictable ones. sLoreta estimations of the brain activity preceding the outcome suggest that prefrontal and parietal areas can be involved in its generation. Prefrontal sites activation (Anterior Cingulate and Dorsolateral Prefrontal Cortex) seems to be related to the degree of uncertainty. Activation in posterior parietal areas, however, does not correlates with uncertainty.

Conclusions/Significance

We suggest that the Stimulus Preceding Negativity reflects the attempt to predict the outcome, when posterior brain areas fail to generate a stable expectancy. Uncertainty is thus conceptualized, not just as the absence of learned expectancy, but as a state with psychological and physiological entity.     

Motor learning in man: A review of functional and clinical studies

This chapter reviews results of clinical and functional imaging studies which investigated the time-course of cortical and subcortical activation during the acquisition of motor a skill. During the early phases of learning by trial and error, activation in prefrontal areas, especially in the dorsolateral prefrontal cortex, is has been reported. The role of these areas is presumably related to explicit working memory and the establishment of a novel association between visual cues and motor commands. Furthermore, motor associated areas of the right hemisphere and distributed cerebellar areas reveal strong activation during the early motor learning. Activation in superior-posterior parietal cortex presumably arises from visuospatial processes, while sensory feedback is coded in the anterior-inferior parietal cortex and the neocerebellar structures. With practice, motor associated areas of the left-hemisphere reveal increased activity. This shift to the left hemisphere has been observed regardless of the hand used during training, indicating a left-hemispheric dominance in the storage of visuomotor skills. Concerning frontal areas, learned actions of sequential character are represented in the caudal part of the supplementary motor area (SMA proper), whereas the lateral premotor cortex appears to be responsible for the coding of the association between visuo-spatial information and motor commands. Functional imaging studies which investigated the activation patterns of motor learning under implicit conditions identified for the first, a motor circuit which includes lateral premotor cortex and SMA proper of the left hemisphere and primary motor cortex, for the second, a cognitive loop which consists of basal ganglia structures of the right hemisphere. Finally, activity patterns of intermanual transfer are discussed. After right-handed training, activity in motor associated areas maintains during performance of the mirror version, but is increased during the performance of the original-oriented version with the left hand. In contrary, increased activity during the mirror reversed action, but not during the original-oriented performance of the untrained right hand is observed after left-handed training. These results indicate the transfer of acquired right-handed information which reflects the mirror symmetry of the body, whereas spatial information is mainly transferred after left-handed training. Taken together, a combined approach of clinical lesion studies and functional imaging is a promising tool for identifying the cerebral regions involved in the process of motor learning and provides insight into the mechanisms underlying the generalisation of actions.