Asymmetry in the Human Motor Cortex and Handedness

Handedness is one of the most obvious functional asymmetries, but its relation to an anatomical asymmetry of the hand representation area in the motor cortex has not been demonstrated. This would be a crucial test for the hypothesis of structure–function correlation in cortical motor organization. Using magnetic resonance morphometry, we show for the first time that the depth of the central sulcus is related to handedness. In right-handers, the left central sulcus is deeper than the right, and vice versa in left-handers. Macrostructural asymmetry is complemented by a microstructural left-larger-than-right asymmetry in neuropil volume (i.e., tissue compartment containing dendrites, axons, and synapses) in Brodmann's area 4. These asymmetries suggest that hand preference is associated with increased connectivity (demonstrated by an increased neuropil compartment in left area 4) and an increased intrasulcal surface of the precentral gyrus in the dominant hemisphere.     

The effect of handedness on cortical motor activation during simple bilateral movements

The neuronal correlates of handedness are still poorly understood. Here we used event-related functional magnetic resonance imaging to investigate the impact of handedness on neuronal activation of the primary sensorimotor cortex, supplementary motor area and dorsal premotor cortex during simple unilateral and bilateral finger movements. In 16 right-handed and 16 left-handed individuals, we mapped changes in regional neuronal activity while participants responded to four symbolic cues presented in a pseudorandom order. According to pre-specified cues, they pressed a button with their right, left or both index fingers or withheld a response. For unilateral right index finger button presses, reaction times, motor and premotor activity were the same for both right- and left-handers. Compared with right-handers, left-handers had shorter reaction times with unilateral left index finger button presses, along with greater activation of the supplementary motor area and right frontal opercular cortex. Simultaneous bilateral compared with unilateral button presses led to a relative increase of activity in the right and left dorsal premotor cortex and the right primary sensorimotor cortex in right but not left-handers. Neither right nor left-handers showed any tendency during bilateral button presses towards faster responses with the dominant hand and the reaction times were equal in the two groups. Therefore, we conclude that the relative increase of activity in dorsal premotor and right primary sensorimotor cortices in right-handers represents a genuine difference in bimanual motor control related to handedness.     

Handedness-dependent and -independent cerebral asymmetries in the anterior intraparietal sulcus and ventral premotor cortex during grasp planning

When planning grasping actions, right-handers show left-lateralized responses in the anterior intraparietal sulcus (aIPS) and ventral premotor cortex (vPMC), two areas that are also implicated in sensorimotor control of grasp. We investigated whether a similar cerebral asymmetry is evident in strongly left-handed individuals. Fourteen participants were trained to grasp an object appearing in a variety of orientations with their left and right hands and with a novel mechanical tool (operated with either hand). BOLD fMRI data were then acquired while they decided prospectively whether an over- or under-hand grip would be most comfortable for grasping the same stimulus set while remaining still. Behavioral performances were equivalent to those recorded previously in right-handers and indicated reliance on effector-specific internal representations. In left-handers, however, grip selection decisions for both sides (left, right) and effectors (hand, tool) were associated with bilateral increases in activity within aIPS and vPMC. A direct comparison between left- and right-handers did reveal equivalent increases in left vPMC regardless of hand dominance. By contrast, aIPS and right vPMC activity were dependent on handedness, showing greater activity in the motor-dominant hemisphere. Though showing bilateral increases in both left- and right-handers, greater increases in the motor dominant hemisphere were also detected in the caudal IPS (cIPS), superior parietal lobule (SPL) and dorsal premotor cortex (dPMC). These findings provide further evidence that regions involved in the sensorimotor control of grasp also participate in grasp planning, and that for certain areas hand dominance is a predictor of the cerebral organization of motor cognitive functions.     

Hemispheric asymmetries in goal-directed hand movements are independent of hand preference

Asymmetries in the kinematics and neural substrates of voluntary right and left eye-hand coordinated movements have been accredited to differential hemispheric specialization. An alternative explanation for between-hand movement differences could result from hand preference related effects. To test both assumptions, an experiment was conducted with left- and right-handers performing goal-directed movements with either hand paced by a metronome. Spatiotemporal accuracy was comparable between hands, whereas hand peak velocity was reached earlier when moving with the left compared to the right hand. The underlying brain activation patterns showed that both left- and right-handers activated more areas involved in visuomotor attention and saccadic control when using their left compared to the right hand. Altogether, these results confirm a unique perceptuomotor processing specialization of the left brain/right hand system that is independent of hand preference.     

Association between language and spatial laterality and cognitive ability: an fMRI study

The interaction between language and spatial laterality and its association with cognitive ability was explored in a group of 42 right-handers and 40 left-handers using functional magnetic resonance imaging. Cognitive ability measures including working memory, verbal comprehension and perceptual organisation were assessed using the Wechsler Adult Intelligence Scale (version III). Left-handers show lower working memory scores than right-handers. Increased rightward language laterality is also associated with decreased working memory performance, which we suggest is related to the involvement of the left inferior frontal gyrus in subvocal rehearsal during working memory tasks. The interaction between language and spatial laterality is associated with performance on verbal comprehension and perceptual organisation, such that when language and spatial laterality are dissociated between the hemispheres a significant increase in verbal comprehension and perceptual organisation performance is found. There is a decrease in performance on the verbal comprehension and perceptual organisation subtests when language and spatial processing are associated to the same hemisphere (i.e. both lateralised to the right hemisphere or both lateralised to the left). This interaction is interpreted in relation to the 'hemispheric crowding' hypothesis, which proposes increased cognitive ability when language and spatial laterality are dissociated.     

Localization of Language-Related Cortex withO-Labeled Water PET in Patients with Gliomas

Measurement of relative cerebral blood flow (CBF) with¹⁵O-labeled water PET has been widely used for brain mapping experiments on language functions in normal volunteers and patients with epilepsy. We focused on the question of whether PET during speech activation is an appropriate method for noninvasive determination of language-related cortex in patients with brain tumors. Furthermore, the suitability of the method for determination of hemispheric language dominance was examined and compared to the results of the Edinburgh Handedness Inventory. Ten right-handed and six left-handed patients with gliomas were examined prior to surgery while repeatedly performing word repetition and verb generation tasks. A set of volumes of interest (VOIs) was drawn on coregistered MRI in order to account for anatomic variability as well as anatomical alterations due to tumor mass effect. Repetition of nouns did not produce significant hemispheric differences. During stimulation by verb generation, reliable lateralized activations of Broca's area and supplementary motor area were detected in all right-handed patients. Of the left-handed patients, two showed clear right lateralization, two activated Broca's area bilaterally, and two had a pattern similar to that of right-handers. Patients with bilateral activations showed the strongest tendency toward bihandedness according to the handedness inventory. Lateralization of supplementary motor area in left-handers corresponded to lateralized activity in Broca's area. Tumors in the vicinity of language-related regions did not alter activation responses. In conclusion, measurement of CBF changes during verb generation permits identification of language-related areas in patients with gliomas with strong lateralization related to hemispheric dominance. These findings may be of particular clinical interest for left-handed patients.     

Effects of unilateral brain damage on contralateral and ipsilateral upper extremity function in hemiplegia

This article describes the long-term effects of unilateral penetrating hemispheric lesions on contralateral and ipsilateral upper extremity motor performance and functional outcome. Activities-of-daily-living skill and gross motor performance contralateral to the lesions were compared among 32 left-sided and 19 right-sided hemiplegic subjects using analysis of variance and chi-square techniques. Ipsilateral to the damaged hemisphere, fine motor tasks of simple visual motor reaction time, grip and pinch strength, finger tapping, and Purdue Pegboard performance were tested. Analysis of covariance compared each ipsilateral task to performance in the corresponding hand of 70 matched controls. Results indicate similar long-term functional ADL outcome in right and left hemisphere-damaged subjects, despite more severe contralateral functional motor deficits following lesions of the left hemisphere. Right hemisphere lesions led to ipsilateral decrements in reaction time, and lesions of either hemisphere diminished grip or pinch strength, finger tapping, and pegboard performance ipsilaterally. These results demonstrate that unilateral brain damage involving the motor areas of either hemisphere has detrimental effects on ipsilateral upper extremity motor function. Findings are discussed and related to the concept that the left hemisphere is specialized or has greater neuronal representation for bilateral motor processes. Physical therapists involved in the treatment of patients with hemiplegia should be aware that motor functions of the ipsilateral, nonparetic upper extremity may also be affected adversely by unilateral brain lesions.     

Left-hemisphere-dominant SII activation after bilateral median nerve stimulation

We used bilateral median nerve stimuli to find out possible hemispheric dominance in the activation of the second somatosensory cortex, SII. Somatosensory evoked fields (SEFs) were recorded from 14 healthy adults (7 right-handed, 7 left-handed) with a 306-channel neuromagnetometer. Electrical stimuli were applied once every 3 s simultaneously either to the left and right median nerves at the wrists or to the palmar skin of both thumbs. Sources of SEFs were modeled with four current dipoles, located in the SI and SII cortices of both hemispheres. The SI activation strengths did not differ between the hemispheres, whereas the SII responses were significantly stronger in the left than in the right hemisphere. In right-handers, the left/right SII ratios were 1.9 and 1.8 for wrist and thumb stimuli, respectively. The corresponding values were 1.5 and 1.7 in left-handers. The results indicate handedness-independent functional specialization of the human SII cortices.     

中央区脑膜瘤患者的功能性近红外光学成像分析及文献回顾

目的评价中央区脑膜瘤患者和对照组手指运动相关的脑激活部位之区别,探讨脑功能重组的机制。方法对1例脑膜瘤患者和2例对照进行近红外脑功能成像检查。试验采用模块设计、单手对指任务,经数据处理、统计和脑功能区定位,比较患者和对照激活区域的差别。结果对照组进行单手运动时对侧大脑半球运动区激活,脑膜瘤患者右手运动时左侧初级感觉运动皮层激活,而左手运动时右侧半球（患侧）在肿瘤后外侧初级感觉运动皮层的通道出现较弱激活,而左侧半球（健侧）初级感觉运动皮层、补充运动区、运动前区出现较强激活。结论功能性近红外脑功能成像检查可检测中央区脑肿瘤患者手部运动相关的脑功能重组。     

Zones of bimanual and unimanual preference within human primary sensorimotor cortex during object manipulation

We asked which brain areas are engaged in the coordination of our hands in dexterous object manipulations where they cooperate for achieving a common goal. Well-trained right-handers steered a cursor on a screen to hit successively displayed targets by applying isometric forces and torques to a rigid tool. In two bimanual conditions, the object was held freely in the air and the hands thus generated coupled opposing forces. Yet, depending on the mapping rule linking hand forces and cursor movements, all subjects selected either the left or the right hand as prime actor. In two unimanual conditions, the subjects performed the same task with either the left or the right hand operating on a fixed tool. Functional magnetic resonance imaging revealed common activation across all four conditions in a dorsal fronto-parietal network biased to the left hemisphere and in bilateral occipitotemporal cortex. Contrary to the notion that medial wall premotor areas are especially active in complex bimanual actions, their activation depended on acting hand (left, right) rather than on grip type (bimanual, unimanual). We observed effects of grip type only in the primary sensorimotor cortex (SMC). In particular, with either hand as prime actor, bimanual actions preferentially activated subregions of the SMC contralateral to the acting hand. A sizeable subregion with preference for unimanual activity was found only in the left SMC in our right-handed subjects. Collectively, these results indicate a hemispheric asymmetry for the SMC and that partially different neural populations support the control of bimanual versus unimanual object manipulations.     

Morphologic asymmetry of the human anterior cingulate cortex

The anterior cingulate cortex (ACC) is thought to play a major role in executive processes. Studies assessing neuroanatomical attributes of this region report a high degree of morphological variability. Recent theories consider the fissurization of the cortex to be a product of gross mechanical processes related to cortical growth and local cytoarchitectural characteristics. Hence, local sulcal patterning and gray matter volume are supposed to be associated. ACC fissurization was quantified in left- and right-handers of both sexes by recording the presence and extension of the paracingulate sulcus (PCS). Differences between groups regarding local gray matter volume were assessed by means of optimized voxel-based morphometry (oVBM) including additional modulation. Overall, the PCS occurred more often and was more pronounced in the left as compared to the right anterior cingulate region, although hemispheric differences were less pronounced in male left- and female right-handers. These discrepancies between groups seem to stem from variations of cingulate morphology in the left rather than the right hemisphere. The pattern of relevant comparisons in the oVBM analysis indicated a similar interaction. Therefore, evidence was found for discrepancies between groups and hemispheres on the macrostructural level.     

The effect of handedness on the shape of the central sulcus

Sinistrals differ from dextrals in the size of certain cortical folds. For instance, handedness has an impact on central sulcus surface area: the sulcus is larger in the dominant left hemisphere of dextrals and vice versa for sinistrals. However, the impact of handedness on the shape of the central sulcus is largely unexplored. In this paper, we propose first an original strategy based on manifold learning to quantify the shape of the central sulcus. Using this approach we show that the "hand knob", a major landmark of the hand motor representation, is sited more dorsally in the left hemisphere in dextrals than in sinistrals. Sinistrals forced to write with their non-preferred right hand display a pattern of central sulcus size asymmetry which is typical of dextrals, yet forced dextrality does not shift the handedness-specific location of the "hand knob". Hence, cortical morphology in adults holds an accumulated record of both innate biases and early developmental experience. Characterizing normal variation of cortical morphology provides a means of systematically correlating behavior with cortical development.     

Right parietal dominance in spatial egocentric discrimination

Egocentric tactile perception is crucial for skilled hand motor control. In order to better understand the brain functional underpinnings related to this basic sensorial perception, we performed a tactile perception functional magnetic resonance imaging (fMRI) experiment with two aims. The first aim consisted of characterizing the neural substrate of two types of egocentric tactile discrimination: the spatial localization (SLD) and simultaneity succession discrimination (SSD) in both hands to define hemispheric dominance for these tasks. The second goal consisted of characterizing the brain activation related to the spatial attentional load, the functional changes and their connectivity patterns induced by the psychometric performance (PP) during SLD. We used fMRI in 25 right-handed volunteers, applying pairs of sinusoidal vibratory stimuli on eight different positions in the palmar surface of both hands. Subjects were required either to identify the stimulus location with respect to an imaginary midline (SLD), to discriminate the simultaneity or succession of a stimuli pair (SSD) or to simply respond to stimulus detection. We found a fronto-parietal network for SLD and frontal network for SSD. During SLD we identified right hemispheric dominance with increased BOLD activation and functional interaction of the right supramarginal gyrus with contralateral intra-parietal sulcus for right and left hand independently. Brain activity correlated to spatial attentional load was found in bilateral structures of intra-parietal sulcus, precuneus extended to superior parietal lobule, pre-supplementary motor area, frontal eye fields and anterior insulae for both hands. We suggest that the right supramarginal gyrus and its interaction with intra-parietal lobule may play a pivotal role in the phenomenon of tactile neglect in right fronto-parietal lesions.     

The cerebral oscillatory network associated with auditorily paced finger movements

Motor tasks involve neural activity in a spatially distributed network. It is assumed that coherent activity between these brain structures reflects functional connectivity. The aim of the present study was to investigate brain areas associated with a unimanual auditorily paced finger-tapping task and to characterize their dynamic interplay. We examined cerebromuscular and cerebrocerebral coupling in 10 right-handed subjects using recordings of continuous brain activity with a 122-channel whole-head neuromagnetometer while subjects performed the task with both hands consecutively. Additionally, surface EMG of the first dorsal interosseus was measured. Our data demonstrate that an oscillatory network composed of primary sensorimotor cortex, lateral as well as mesial premotor areas, the posterior parietal cortex and thalamus contralateral, and cerebellum and primary auditory cortex ipsilateral to the tapping hand subserves task execution. Connectivity between these areas and direction of coupling agree well with anatomical findings. During the right-hand condition, additional oscillatory activity in the primary sensorimotor cortex ipsilateral to the tapping hand was evident. This result suggests an asymmetric motor control in right-handers. Cerebrocerebral coupling predominantly occurs at 8-12 Hz. Therefore, our data support the hypothesis that coupling at 8-12 Hz in a cerebello-thalamic-cortical network represents a fundamental characteristic of the motor system and provides evidence for the significance of 8-12 Hz oscillations in a large scale network during the execution of simple motor tasks.     

The role of the medial wall and its anatomical variations for bimanual antiphase and in-phase movements

The medial wall of the frontal cortex is thought to play an important role for bimanual coordination. However, there is uncertainty regarding the exact neuroanatomical regions involved. We compared the activation patterns related to bimanual movements using functional magnetic resonance imaging in 12 healthy right-handed subjects, paying special attention to the anatomical variability of the frontal medial wall. The subjects performed unimanual right and left and bimanual antiphase and in-phase flexion and extension movements of the index finger. Activation of the right supplementary motor area (SMA) proper, right and left caudal cingulate motor area (CMA), and right and left premotor cortices was significantly stronger during bimanual antiphase than bimanual in-phase movements, indicating an important function of these areas with bimanual coordination. A frequent anatomical variation is the presence of the paracingulate sulcus (PCS), which might be an anatomical landmark to determine the location of activated areas. Seven subjects had a bilateral, three a unilateral right, and two a unilateral left PCS. Because the area around the PCS is functionally closer coupled to the CMA than to the SMA, activation found in the area around the PCS should be attributed to the CMA. With anatomical variations such as the presence of a PCS or a vertical branch of the cingulate sulcus, normalization and determination of the activation with the help of stereotaxic coordinates can cause an incorrect shift of CMA activation to the SMA. This might explain some of the discrepancies found in previous studies.     

原发性三叉神经痛灰质体积变化的磁共振形态学研究

目的应用基于体素的形态学分析方法检测原发性三叉神经痛患者是否存在灰质体积改变,探索中枢神经系统在其疼痛处理中的作用。材料与方法对19例右侧三叉神经痛患者和22例年龄性别匹配的正常对照进行磁共振三维T1结构像扫描(3D-T1BRAVO),应用基于体素形态学分析(voxel-based morphometry,VBM)检测三叉神经痛患者全脑灰质体积的变化,探索各变化的中枢神经系统脑区在疼痛处理中的作用,并分别分析其体积变化与疼痛程度及病程的相关性。结果与正常对照组相比,病例组左侧前扣带回灰质体积明显减少;右侧额上回、右侧岛叶、右侧壳核、双侧小脑前叶灰质体积明显增加,左侧前扣带回体积变化与疼痛病程呈负相关性。结论三叉神经痛患者存在中枢神经系统灰质体积变化,左侧前扣带回体积与疼痛病程呈负相关性,这些存在差异的中枢神经系统脑区在三叉神经痛患者疼痛的形成及慢性状态的维持中可能有重要作用,提示探索三叉神经痛的中枢处理机制有益于新的治疗方法的研发。     

Where does your own action influence your perception of another person's action in the brain?

Activation of premotor cortex during the observation and imitation of human actions is now increasingly accepted, but it remains unclear how the CNS is able to resolve potential conflicts between the observation of another person's action and the ongoing control of one's own action. Recent data suggest that this overlap leads to a systematic bias, where lifting a box influences participant's perceptual judgments of the weight of a box lifted by another person. We now investigate the neural basis of this bias effect using fMRI. Seventeen participants performed a perceptual weight judgment task or two control conditions while lifting a light box, a heavy box or no box during scanning. Brain regions related to perceptual bias were localized by correlating individual differences in bias with BOLD signal. Five regions were found to show correlations with psychophysical bias: left inferior frontal gyrus, left central sulcus, left extrastriate body area, left lingual gyrus and right intraparietal sulcus. The cluster in primary motor cortex was also activated by box lifting, and the cluster in extrastriate body area by the observation of hand actions and the weight judgment task. We suggest that these brain areas are part of a network where motor processing modulates perceptual judgment of observed human actions, and thus visual and motor processes cannot be thought of as two distinct systems, but instead interact at many levels.     

Posture influences motor imagery: an fMRI study

Motor imagery is widely used to study cognitive aspects of the neural control of action. However, what is exactly simulated during motor imagery is still a matter of debate. On the one hand, it is conceivable that motor imagery is an embodied cognitive process, involving a simulation of movements of one's own body. The alternative possibility is that, although motor imagery relies on knowledge of the motor processes, it does not entail an actual motor simulation that is influenced by the physical configuration of one's own body. Here we discriminate between these two hypotheses, in the context of an established motor imagery task: laterality judgments of rotated hand drawings. We found that reaction times of hand laterality judgments followed the biomechanical constraints of left or right hand movements. Crucially, the position of subjects' own left and right arm influenced laterality judgments of left and right hands. In neural terms, hand laterality judgments activated a parieto-frontal network. The activity within this network increased with increasing biomechanical complexity of the imagined hand movements, even when the amount of stimulus rotation was identical. Moreover, activity in the intraparietal sulcus was modulated by subjects' own hand position: a larger incongruence in orientation between the subjects' hand and the stimulus hand led to a selective increase in intraparietal activity. Our results indicate that motor imagery generates motor plans that depend on the current configuration of the limbs. This motor plan is calculated by a parieto-frontal network. Within this network, the posterior parietal cortex appears to incorporate proprioceptive information related to the current position of the body into the motor plan.     

Comparison of Muscle Activation during Dominant Hand Wrist Flexion when Writing

[Purpose] This study investigated the difference in muscle activation of the dominant upper extremity in right-handed and left-handed persons during writing. [Subjects] There were 36 subjects (16 left- handers/ 20 right- handers), and the study was conducted from 03/01/2012 to 30/3/2012. [Methods] Six electrodes were attached to the FCU (flexor carpi ulnaris), FCR (flexor carpi radialis), ECU (extensor carpi ulnaris), ECR (extensor carpi radialis), and both UT (upper trapezius) muscles. [Results] FCU muscle activation was 16.77±9.12% in left-handers and 10.29±4.13% (%MVIC) in right-handers. FCR muscle activation was 19.09±9.43% in left-handers and 10.64±5.03% in right-handers. In addition, the UT muscle activation on the writing hand side was 11.91±5.79% in left-handers and 1.66±1.19% in right-handers. [Conclusion] As a result of this study, it was discovered that left-handers used more wrist flexion in performance of the writing task with the dominant upper extremity than right-handers, and that the left-handers activated the wrist and shoulder muscles more than the right-handers. These results indicate a potential danger of musculoskeletal disease in left-hander.Key words: Hand function, Left-hander, Muscle activation     

利用fMRI和双手交替运动模式研究脑肿瘤所致的运动功能重组

目的应用fMRI研究双手交替运动模式下中央沟附近脑肿瘤患者运动功能重组的方式及特征.方法6名正常受试者和14名脑肿瘤患者采用双手交替对指运动模式行fMRI扫描,比较正常受试者与脑肿瘤患者脑激活的异同.结果正常人单手对指运动主要激活运动手对侧的大脑半球和同侧小脑半球.脑肿瘤患者非受累手运动所致的激活与正常受试者基本相同;而当受累手运动时,则出现运动功能的重组,包括肿瘤对侧正常半球内M1区的代偿性激活、肿瘤侧半球MI活动的减弱、双侧SMA等次级运动中枢激活区的增大以及双侧小脑半球的激活.结论采取双手交替运动模式,fMRI不仅显示了受累侧运动区的变形与移位,而且揭示了一种新的功能重组模式,即运动功能重组可能涉及分布于全脑的整个神经网络.