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Brain lesion characteristics (timing, location, and extent) and the type of corticospinal tract (CST) wiring have been proposed as determinants of upper limb (UL) motor function in unilateral cerebral palsy (uCP), yet an investigation of the relative combined impact of these factors on both motor and sensory functions is still lacking. Here, we first investigated whether structural brain lesion characteristics could predict the underlying CST wiring and we explored the role of CST wiring and brain lesion characteristics to predict UL motor and sensory functions in uCP. Fifty-two participants with uCP (mean age (SD): 11 y and 3 m (3 y and 10 m)) underwent a single-pulse Transcranial Magnetic Stimulation session to determine CST wiring between the motor cortex and the more affected hand ( n = 17 contralateral, n = 19 ipsilateral, and n = 16 bilateral) and an MRI to determine lesion timing ( n = 34 periventricular (PV) lesion, n = 18 corticosubcortical (CSC) lesion), location, and extent. Lesion location and extent were evaluated with a semiquantitative scale. A standardized protocol included UL motor (grip strength, unimanual capacity, and bimanual performance) and sensory measures. A combination of lesion locations (damage to the PLIC and frontal lobe) significantly contributed to differentiate between the CST wiring groups, reclassifying the participants in their original group with 57% of accuracy. Motor and sensory functions were influenced by each of the investigated neurological factors. However, multiple regression analyses showed that motor function was predicted by the CST wiring (more preserved in individuals with contralateral CST ( p < 0.01)), lesion extent, and damage to the basal ganglia and thalamus. Sensory function was predicted by the combination of a large and later lesion and an ipsilateral or bilateral CST wiring, which led to increased sensory deficits ( p < 0.05). These novel insights contribute to a better understanding of the underlying pathophysiology of UL function and may be useful to delineate individualized treatment strategies.

Recent theories propose that movement observation is not a "passive mirror" of ongoing actions but might induce anticipatory activity when predictable movements are observed, e.g., because the action goal is known. Here we investigate this mechanism in a series of 3 experiments, by applying transcranial magnetic stimulation (TMS) to primary motor cortex (M1) while subjects observed either whole hand or precision grasping performed by an actor. We show that corticomotor excitability changes in a grip-specific manner but only once the grip can be decoded based on the observed kinematic cues (Exp. 1). By contrast, presenting informative contextual precues evokes anticipatory modulations in M1 already during the reach phase, i.e., well before the grip type could be observed, a finding in line with a predictive coding account (Exp. 2). Finally, we used paired-pulse (PP) TMS to show that ventral premotor cortex (PMv) facilitates grip-specific representations in M1 but only while grip formation is observed. These findings suggest that PMv and M1 interact temporarily and mainly when motor aspects of hand-object interactions are extracted from visual information. By contrast, no sustained input from PMv to M1 seems to be required to maintain action representations that are anticipated based on contextual information or once the grip is formed (Exp. 3)., (Copyright © 2015 Elsevier Ltd. All rights reserved.)

In young adults, canceling an initiated action depends on the right inferior frontal cortex (IFC), presupplementary motor area (preSMA), and the basal ganglia. Older adults show response inhibition deficits, but how this relates to functional brain activation remains unclear. Using event-related functional magnetic resonance imaging, we tested whether older adults (N = 20) exhibit overactivation during stop-signal inhibition as shown for attentional control tasks, or reduced activity compared with young adults (N = 20). We used a modified stop-signal task involving coupled bimanual responses and manipulated whether both or just one hand was cued to stop. Stop-task difficulty was matched across groups. We found a group by condition interaction in supramarginal gyrus, anterior insula, rIFC, and preSMA, with activation increasing for successful Stop versus Go trials in the young adults only. Comparing the groups on Stop trials revealed preSMA and striatum hypoactivity for older adults. White matter tracts connecting rIFC, preSMA, and the subthalamic nuclei were associated with stronger activation of preSMA in older adults, suggesting that maintenance of the brain's structure has positive implications for brain function., (© The Author 2014. Published by Oxford University Press. All rights reserved. For Permissions, please e-mail: journals.permissions@oup.com.)

When a stable memory is reactivated it becomes transiently labile and requires restabilization, a process known as reconsolidation. Animal studies have convincingly demonstrated that during reconsolidation memories are modifiable and can be erased when reactivation is followed by an interfering intervention. Few studies have been conducted in humans, however, and results are inconsistent regarding the extent to which a memory can be degraded. We used a motor sequence learning paradigm to show that the length of reactivation constitutes a crucial boundary condition determining whether human motor memories can be degraded. In our first experiment, we found that a short reactivation (less than 60 sec) renders the memory labile and susceptible to degradation through interference, while a longer reactivation does not. In our second experiment, we reproduce these results and show a significant linear relationship between the length of memory reactivation and the detrimental effect of the interfering task performed afterwards, i.e., the longer the reactivation, the smaller the memory loss due to interference. Our data suggest that reactivation via motor execution activates a time-dependent process that initially destabilizes the memory, which is then followed by restabilization during further practice.

During movement observation, corticomotor excitability of the observer's primary motor cortex (M1) is modulated according to the force requirements of the observed action. Here, we explored the time course of observation-induced force encoding. Force-related changes in M1-excitability were assessed by delivering transcranial magnetic stimulations at distinct temporal phases of an observed reach-grasp-lift action. Temporal changes in force-related electromyographic activity were also assessed during active movement execution. In observation conditions in which a heavy object was lifted, M1-excitability was higher compared to conditions in which a light object was lifted. Both during observation and execution, differential force encoding tended to gradually increase from the grasping phase until the late lift phase. Surprisingly, however, during observation, force encoding was already present at the early reach phase: a time point at which no visual cues on the object's weight were available to the observer. As the observer was aware that the same weight condition was presented repeatedly, this finding may indicate that prior predictions concerning the upcoming weight condition are reflected by M1 excitability. Overall, findings may provide indications that the observer's motor system represents motor predictions as well as muscular requirements to infer the observed movement goal.

Diffusion weighted imaging (DWI) studies in humans have shown that seniors exhibit reduced white matter integrity compared with young adults, with the most pronounced change occurring in frontal white matter. It is generally assumed that this structural deterioration underlies inhibitory control deficits in old age, but specific evidence from a structural neuroscience perspective is lacking. Cognitive action control is thought to rely on an interconnected network consisting of right inferior frontal cortex (r-IFC), pre-supplementary motor area (preSMA), and the subthalamic nucleus (STN). Here we performed probabilistic DWI tractography to delineate this cognitive control network and had the same individuals (20 young, 20 older adults) perform a task probing both response inhibition and action reprogramming. We hypothesized that structural integrity (fractional anisotropy) and connection strength within this network would be predictive of individual and age-related differences in task performance. We show that the integrity of r-IFC white matter is an age-independent predictor of stop-signal reaction time (SSRT). We further provide evidence that the integrity of white matter projecting to STN predicts both outright stopping (SSRT) and transient braking of response initiation to buy time for action reprogramming (stopping interference effects). These associations remain even after controlling for Go task performance, demonstrating specificity to the Stop component of this task. Finally, a multiple regression analysis reveals bilateral preSMA-STN tract strength as a significant predictor of SSRT in older adults. Our data link age-related decline in inhibitory control with structural decline of STN projections.

Practicing a motor task can induce neuroplastic changes in the human primary motor cortex (M1) that are subsequently consolidated, leading to a stable memory trace. Currently, little is known whether early consolidation, tested several minutes after skill acquisition, can be improved by behavioral interventions. Here we test whether movement observation, known to evoke similar neural responses in M1 as movement execution, can benefit the early consolidation of new motor memories. We show that observing the same type of movement as that previously practiced (congruent movement stimuli) substantially improves performance on a retention test 30 min after training compared with observing either an incongruent movement type or control stimuli not showing biological motion. Differences in retention following observation of congruent, incongruent, and control stimuli were not found when observed 24 h after initial training and neural evidence further confirmed that, unlike motor practice, movement observation alone did not induce plastic changes in the motor cortex. This time-specific effect is critical to conclude that movement observation of congruent stimuli interacts with training-induced neuroplasticity and enhances early consolidation of motor memories. Our findings are not only of theoretical relevance for memory research, but also have great potential for application in clinical settings when neuroplasticity needs to be maximized.

Language and certain aspects of motor control are typically served by the left hemisphere, whereas visuospatial and attentional control are lateralized to the right. Here a (visuo)motor tracing task was used to identify hemispheric lateralization beyond the general, contralateral organization of the motor system. Functional magnetic resonance imaging (fMRI) was applied in 40 male right-handers (19-30 yrs) during line tracing with dominant and nondominant hand, with and without visual guidance. Results revealed a network of areas activating more in the right than left hemisphere, irrespective of the effector. Inferior portions of frontal gyrus and parietal lobe overlapped largely with a previously described ventral attention network responding to unexpected or behaviourally relevant stimuli. This demonstrates a hitherto unreported functionality of this circuit that also seems to activate when spatial information is continuously exploited to adapt motor behaviour. Second, activation of left dorsal premotor and postcentral regions during tracing with the nondominant left hand was more pronounced than that in their right hemisphere homologues during tracing with the dominant right hand. These activation asymmetries of motor areas ipsilateral to the moving hand could not be explained by asymmetries in skill performance, the degree of handedness, or interhemispheric interactions. The latter was measured by a double-pulse transcranial magnetic stimulation paradigm, whereby a conditioning stimulus was applied over one hemisphere and a test stimulus over the other. We propose that the left premotor areas contain action representations strongly related to movement implementation which are also accessed during movements performed with the left body side., (Copyright © 2010 Wiley-Liss, Inc.)

Transcranial magnetic stimulation (TMS) is increasingly used to perturb targeted human brain sites non-invasively, to test for causal effects on performance of cognitive tasks. TMS might also be used in non-human primates to complement invasive work and compare with human studies. Here, we targeted the frontal eye fields (FEF) in two macaques with a continuous theta-burst (cTBS) protocol, testing the impact on visually guided saccades. After unilateral cTBS over the FEF in either hemisphere, a small (mean 7 ms) but highly consistent decrease in saccadic reaction times (RTs) was observed. Lower latencies arose for saccades both contra- and ipsilateral to the stimulated FEF after cTBS. These results provide the first demonstration that TMS can be used to affect saccadic behavior in non-human primates. The unexpectedly bilateral impact on RTs may reflect an impact on 'fixation' neurons in the FEF and/or transcallosal modulation of both FEFs induced by unilateral cTBS. In either case, this study demonstrates a clear behavioral effect induced by TMS in awake behaving monkeys performing a cognitive task. This opens new opportunities for investigating the causal roles of targeted brain areas in behavior, for measuring physiological consequences of TMS in the primate brain, and ultimately for human-monkey comparisons.

Seeing or hearing manual actions activates the mirror neuron system, that is, specialized neurons within motor areas which fire when an action is performed but also when it is passively perceived. Using TMS, it was shown that motor cortex of typically developed subjects becomes facilitated not only from seeing others' actions, but also from merely hearing action-related sounds. In the present study, TMS was used for the first time to explore the "auditory" and "visual" responsiveness of motor cortex in individuals with congenital blindness or deafness. TMS was applied over left primary motor cortex (M1) to measure cortico-motor facilitation while subjects passively perceived manual actions (either visually or aurally). Although largely unexpected, congenitally blind or deaf subjects displayed substantially lower resonant motor facilitation upon action perception compared to seeing/hearing control subjects. Moreover, muscle-specific changes in cortico-motor excitability within M1 appeared to be absent in individuals with profound blindness or deafness. Overall, these findings strongly argue against the hypothesis that an increased reliance on the remaining sensory modality in blind or deaf subjects is accompanied by an increased responsiveness of the "auditory" or "visual" perceptual-motor "mirror" system, respectively. Moreover, the apparent lack of resonant motor facilitation for the blind and deaf subjects may challenge the hypothesis of a unitary mirror system underlying human action recognition and may suggest that action perception in blind and deaf subjects engages a mode of action processing that is different from the human action recognition system recruited in typically developed subjects.

Structural and neurochemical changes in frontostriatal circuits are thought to underlie age-related behavioral deficits on cognitive tasks. Here, we test the hypothesis that age-related motor switching deficits are associated with reduced basal ganglia (BG) function. Right-handed volunteers (15 Old, and 15 Young) made spatially and temporally coupled bimanual circular motions during event-related FMRI. A visual cue signaled the right hand to Switch or Continue its circling direction. Switching from mirror symmetric to asymmetric (SW»ASYMM) took longer and resulted in more contralateral (left-) hand disruptions than vice versa. These effects were more pronounced in the elderly, showing that the ability to suppress and flexibly adapt motor behavior (agility) declines with age. For both groups, switching activated the BG and a typical network for task-set implementation, including dorsal anterior cingulate cortex/supplementary motor area (pre-SMA, SMA-proper) and anterior insula/inferior frontal gyrus. A region of interest analysis revealed significantly reduced SW»ASYMM activation in bilateral subthalamic nucleus and right globus pallidus, only in the elderly. Age-related behavioral deficits may be related to inefficient recruitment of cortico-BG loops to suppress undesired movements. The elderly may use an alternative strategy to select the required movement pattern as indicated by increased activation of prefrontal cortex.

Coordinated hand use is an essential component of many activities of daily living. Although previous studies have demonstrated age-related behavioral deficits in bimanual tasks, studies that assessed the neural basis underlying such declines in function do not exist. In this fMRI study, 16 old and 16 young healthy adults performed bimanual movements varying in coordination complexity (i.e., in-phase, antiphase) and movement frequency (i.e., 45, 60, 75, 90% of critical antiphase speed) demands. Difficulty was normalized on an individual subject basis leading to group performances (measured by phase accuracy/stability) that were matched for young and old subjects. Despite lower overall movement frequency, the old group "overactivated" brain areas compared with the young adults. These regions included the supplementary motor area, higher order feedback processing areas, and regions typically ascribed to cognitive functions (e.g., inferior parietal cortex/dorsolateral prefrontal cortex). Further, age-related increases in activity in the supplementary motor area and left secondary somatosensory cortex showed positive correlations with coordinative ability in the more complex antiphase task, suggesting a compensation mechanism. Lastly, for both old and young subjects, similar modulation of neural activity was seen with increased movement frequency. Overall, these findings demonstrate for the first time that bimanual movements require greater neural resources for old adults in order to match the level of performance seen in younger subjects. Nevertheless, this increase in neural activity does not preclude frequency-induced neural modulations as a function of increased task demand in the elderly., (2010 Wiley-Liss, Inc.)

Cerebral patterns of activity elicited by dual-task performance throughout the learning of a complex bimanual coordination pattern were addressed. Subjects (N=12) were trained on the coordination pattern and scanned using fMRI at early (PRE) and late (POST) learning stages. During scanning, the coordination pattern was performed either as a single task or in concurrence with a simultaneous visual search task (i.e. dual task). Kinematics data revealed a significant performance improvement as a result of learning. In PRE-scanning, the dual-task condition induced deterioration of motor performance, relative to the single-task condition. Activity in lateral frontal and parietal regions involved in both visual search and motor coordination tasks (i.e. 'overlapping' regions) was reduced when the tasks were performed simultaneously. In POST-scanning, kinematics performance was equivalent under single- and dual-task conditions, suggesting automaticity of the coordination pattern. Furthermore, overlap between regions involved in visual search and motor tasks was reduced, and dual-task performance was no longer associated with reduction of frontal and parietal activity. Our results suggest that behavioral interference between a complex motor coordination task and a simple simultaneous visual search task occurs when both tasks recruit overlapping regions in the frontal and parietal cortices. This may provide a neural basis for dissipation of dual-task interference following extensive motor practice, which is a traditional behavioral marker of motor automaticity., (Copyright (c) 2010 Elsevier Ltd. All rights reserved.)

The cerebral hemispheres of humans exhibit functional asymmetries. It is generally thought that the left hemisphere contributes to higher order planning of demanding motor tasks, while the right hemisphere plays an important role in processing visual or proprioceptive stimuli and controls spatial attention. Few studies have directly investigated which aspects of motor control increase the involvement of right-lateralized areas. We used fMRI to examine hemispheric lateralization during unilateral motor coordination of the wrist and ankle performed either with the left or right body side, and either with or without visual guidance. Visual guidance was provided such that the spatial position of a cursor directly informed subjects about the mode and quality of the coordination pattern. Activation was only considered lateralized for a specific condition if it was significantly stronger in one hemisphere than the other, independent of which body side performed the task. We found that task performance with visual guidance mainly engaged a right-lateralized occipital-temporoparietal network and the inferior frontal gyrus, a circuit known to integrate visual and proprioceptive information to guide movements in space. Importantly, this lateralized activation was only observed when visual guidance was provided, but not when movements were performed without visual guidance or when subjects passively watched a similar visual stimulus without moving their limbs. We argue that the functional lateralization of right visuomotor areas was a direct consequence of performing this motor task in the presence of visual guidance, i.e., visuospatial information was integrated with somatosensory guidance to produce well coordinated hand-foot movements., (2010 Elsevier Inc. All rights reserved.)

Here we review behavioral and brain imaging results on stimulus-response selection in the context of bimanual movements, which is a prototypical paradigm frequently used to investigate the coordination of complex motor behavior. We propose that stimulus-response selection is constrained at the motor, perceptual, and cognitive levels, with the relative importance of each dependent on the task context. Motor constraints seem to dominate when response selection requirements are low, whereas perceptual and cognitive constraints become increasingly important when the appropriate movement has to be associated with a visual cue. We argue that certain cue features determine how task goals are conceptualized, which influences how a particular motor response is selected and implemented by the nervous system.

Functional imaging studies have shown that seniors exhibit more elaborate brain activation than younger controls while performing motor tasks. Here, we investigated whether this age-related overactivation reflects compensation or dedifferentiation mechanisms. "Compensation" refers to additional activation that counteracts age-related decline of brain function and supports successful performance, whereas "dedifferentiation" reflects age-related difficulties in recruiting specialized neural mechanisms and is not relevant to task performance. To test these predictions, performance on a complex interlimb coordination task was correlated with brain activation. Findings revealed that coordination resulted in activation of classical motor coordination regions, but also higher-level sensorimotor regions, and frontal regions in the elderly. Interestingly, a positive correlation between activation level in these latter regions and motor performance was observed in the elderly. This performance enhancing additional recruitment is consistent with the compensation hypothesis and characterizes neuroplasticity at the systems level in the aging brain.

It is known that, in macaques, movements guided by somatosensory information engage anterior parietal and posterior precentral regions. Movements performed with both visual and somatosensory feedback additionally activate posterior parietal and anterior precentral areas. It remains unclear whether the human parieto-frontal circuits exhibit a similar functional organization. Here, we employed a directional interference task requiring a continuous update of sensory information for the on-line control of movement direction, while brain activity was measured by functional magnetic resonance imaging (fMRI). Directional interference arises when bimanual movements occur along different directions in joint space. Under these circumstances, the presence of visual information does not substantially alter performance, such that we could vary the amount and type of sensory information used during on-line guidance of goal-directed movements without affecting motor output. Our results confirmed that in humans, as in macaques, movements guided by somatosensory information engages anterior parietal and posterior precentral regions, while movements performed with both visual and somatosensory information activate posterior parietal and anterior precentral areas. We provide novel evidence on how the interaction of specific portions of the dorsal parietal and precentral cortex in the right hemisphere might generate spatial representations by integrating different sensory modalities during goal-directed movements.

Current behavioural theories consider that during motor learning, an effector-independent memory representation of the acquired skill is built up. Using a transfer paradigm, we addressed the nature of the memory representation for a 2:1 multifrequency co-ordination task, requiring, for example, the left arm to cycle twice as fast as the right. After learning this 2:1 pattern, transfer to its converse pattern (i.e., the right arm cycles twice as fast as the left) revealed powerful evidence for negative transfer. The converse task arrangement revealed similar effects. These observations suggest a reconsideration of current viewpoints on movement representations, which emphasize effector independence. Based on the present findings, we propose a new model of motor memory, consisting of an abstract, effector-independent and an effector-specific layer. The abstract code is hypothesized to represent general spatiotemporal movement features, whereas the specific representation refers to effector-related movement commands. This concept is consistent with recent neuroscientific evidence in animal and human species, and invites a reconsideration of current behavioural theories of motor learning and memory.

Bimanual interference emerges when spatial features, such as movement direction or amplitude, differ between limbs, as indicated by a mutual bias of limb trajectories. Although first insights into the neural basis of directional interference have been revealed recently, little is known about the neural network associated with amplitude interference. We investigated whether amplitude versus directional interference activates differential networks. Functional magnetic resonance imaging (fMRI) was applied while subjects performed cyclical, bimanual joystick movements with either the same vs. different amplitudes, directions, or both. The kinematic analysis confirmed that subjects experienced amplitude interference when they moved with different as compared to the same amplitude, and directional interference when they moved along different as compared to the same direction. On the brain level, amplitude and directional interference both resulted in activation of a bilateral superior parietal-premotor network, which is known to contribute to sensorimotor transformations during goal-directed movements. Interestingly, amplitude but not directional interference exclusively activated a bilateral network containing the dorsolateral prefrontal cortex, anterior cingulate, and supramarginal gyrus, which was shown previously to contribute to executive functions. Even though the encoding of amplitude and directional information converged and activated the same neural substrate, our data thus show that additional and partly independent mechanisms are involved in bimanual amplitude as compared to that in directional control., ((c) 2005 Wiley-Liss, Inc.)

The present study addressed the nature of the memory representation for interlimb coordination tasks. For this purpose, the acquisition of a multifrequency (2:1) task with the ipsilateral limbs and transfer to the ipsilateral and contralateral body side was examined. In particular, subjects practiced a 2:1 coordination pattern whereby the right arm moved twice as fast as the right leg, or vice versa. Subsequently, they transferred the practiced 2:1 task to three different conditions: (1) the converse partner (i.e., the slow-moving limb had to move fast, and vice versa) at the ipsilateral body side, and (2) the identical and (3) converse 2:1 pattern at the contralateral body side. Findings revealed positive transfer of the identical and converse 2:1 pattern to the contralateral body side. However, no transfer of the learned pattern to its converse partner at the same body side was revealed. We propose a new memory representation model for coordination patterns, composed of an effector-independent and effector-specific component (dual-layer model). It is hypothesized that the general movement goal (i.e., moving one limb twice as fast as the other) constitutes the abstract, higher-level representation that may account for positive contralateral transfer. Conversely, the effector-specific component contains task-specific lower-level muscle synergies that are acquired through practice, prohibiting positive transfer when shifting task allocation within the same effectors. These findings are consistent with recent neuroscientific evidence for neuroplastic changes in distributed brain areas.

Behavioral studies in humans have shown that bimanual coordination imposes specific demands on the central nervous system that exceed unimanual task control. In the present study we used functional magnetic resonance imaging to investigate the neural correlate of this additional coordination effort, i.e. regions responding more strongly to bimanual movements than inferred from summing up the responses to the unimanual subtasks. Subjects were scanned while performing movements along different directions, either uni- or bimanually. During the bimanual condition, trajectories of movement of the left and right hand were spatially incompatible, such that additional effort was required to break away from intrinsically favored mirror-movements and to integrate movements of both limbs into a new spatial pattern. Our main finding was that the execution of spatially complex bimanual coordination as compared with the unimanual subtasks activated the anterior cingulate cortex (posterior part) as well as the dorso-anterior precuneus. We hypothesize that the anterior cingulate exerts its modulatory effect on other motor areas, such as the primary motor cortex and the supplementary motor area, in order to suppress intrinsically favored coordination tendencies. Conversely, the precuneus is likely to be involved in shifting attention between different locations in space, which was necessary for monitoring the trajectories of the left and right wrist when both limbs moved in parallel. Our findings suggest that the coordination effort during bimanual and perhaps other modes of coordinated behavior is mediated by regions contributing to higher order functions, which form an interface between cognition and action.

Little is known about activation changes reflecting overlearning, i.e., extensive motor training beyond asymptotic performance. Here we used functional magnetic resonance imaging to trace the neural shifts from an initial to a skilled (learning) and finally overlearned stage (automatization). Scanning occurred before training (PRE) and after 1 (MID) and 2 weeks (POST) of intensive practice on a new bimanual coordination task (>10,500 cycles). Kinematics revealed major improvements between PRE and MID sessions, whereas MID to POST session performance leveled off, indicative of learning and automatization, respectively. Imaging findings showed that activation decreased in bilateral opercular areas, bilateral ventrolateral prefrontal cortex, the right ventral premotor and supramarginal gyrus, and the anterior cingulate sulcus during the learning stage and in the supplementary motor area during the automatization stage. These changes are hypothesized to reflect decreases in attention-demanding sensory processing, as well as suppression of preferred coordination tendencies as a prelude to acquiring new coordination modes. Conversely, learning-related increases were observed in the primary motor cortex (M1), posterior cingulate zone (PCZ), putamen, and right anterior cerebellum. Importantly, both M1 and PCZ activation decreased again to initial level (PRE) during automated performance (POST). Only the putamen and anterior cerebellum remained more activated across both learning and automatization stages, supporting their crucial role in long-term motor memory formation for coordination tasks.

The present study examined whether the ability to dissociate bimanual limb movements following learning of a new coordination task (i.e. star-line drawing paradigm) can be generalised to different effector systems, as expressed by inter- and intralimb transfer. In Experiment 1, subjects practised the 'Line-Star' task (i.e. left arm traced the line/right arm traced the star) and then transferred this pattern to its symmetry partner: the 'Star-Line' task (left arm star/right arm line). In Experiment 2, intralimb transfer from the shoulder-elbow (proximal) to the wrist-finger joints (distal), and vice versa, was investigated. Results revealed positive interlimb transfer among symmetry partners of the star-line movement. Moreover, learning the star-line task spontaneously transferred from the trained to the untrained effector system whereby proximal to distal transfer was larger than vice versa. It is concluded that learning to spatially dissociate the movements of both limbs is generalisable to different motor conditions even though transfer to some conditions is suboptimal. It is hypothesised that the nature of the representation of the spatial interference task is largely effector independent.

Bimanual coordination, a prototype of a complex motor skill, has recently become the subject of intensive investigation. Whereas past research focused mainly on the identification of the elementary coordination constraints that limit performance, the focus is now shifting towards overcoming these coordination constraints by means of task symbolization or perceptual transformation rules that promote the integration of the task components into a meaningful "gestalt". The study of these cognitive penetrations into action will narrow the brain-mind gap and will facilitate the development of a cognitive neuroscience perspective on bimanual movement control.

When performing movements with different spatial trajectories in both upper limbs simultaneously, patterns of interference emerge that can be overcome with practice. Even though studies on the role of augmented feedback in motor learning have been abundant, it still remains to be discovered how overcoming such specific patterns of spatial interference can be optimized by instructional intervention. In the present study, one group acquired a bimanual movement with normal vision, whereas a second group received augmented feedback of the obtained trajectories on a computer screen in real time. Findings revealed that, relative to normal vision, the augmented feedback hampered skill learning and transfer to different environmental conditions. These observations are discussed in view of the benefits and pitfalls of augmented feedback in relation to task context and instructional condition.

It is commonly agreed that a functional dissociation with respect to the internal vs external control of movements exists for several brain regions. This has, however, only been tested in relation to the timing and preparation of motor responses, but not to ongoing movement control. Using functional magnetic resonance imaging (fMRI), the present study addressed the neuroanatomical substrate of the internal-external control hypothesis by comparing regional brain activation for cyclical bimanual movements performed in the presence or absence of augmented visual feedback. Subjects performed a bimanual movement pattern, either with the help of on-line visual feedback of the movements (externally guided coordination) or with the eyes closed on the basis of an internal representation of the movement pattern (internally generated coordination). Visual control and baseline rest conditions were also added. Results showed a clear functional dissociation within the network involved in movement coordination. The hMT/V5+, the superior parietal cortex, the premotor cortex, the thalamus, and cerebellar lobule VI showed higher activation levels when movements were guided by visual feedback. Conversely, the basal ganglia, the supplementary motor area, cingulate motor cortex, the inferior parietal, frontal operculum, and cerebellar lobule IV-V/dentate nucleus showed higher involvement when movements were internally generated. Consequently, the present findings suggest the existence of distinct cortico-cortical and subcortico-cortical neural pathways for externally (augmented feedback) and internally guided cyclical bimanual movements. This provides a neurophysiological account for the beneficial effect of providing augmented visual feedback to optimize movements in normal and motor disordered patients.

In the present study, we investigated the influence of external force manipulations on movements in different directions, while keeping the amplitude invariant. Subjects ( n=10) performed a series of cyclical anteroposterior, mediolateral, and oblique line-drawing movements (star drawing task) with their dominant limb in the horizontal plane. To dissociate kinematics from the underlying patterns of muscle activation, spring loading was applied to the forearm of the moving limb. Whereas spring loading of the arm resulted in considerable changes in the overall amount of muscle activation in the elbow and shoulder muscles, invariance was largely maintained at the kinematic level. Subjects produced the required movement directions and amplitudes of the star drawing largely successfully, irrespective of the force bias induced by the spring. These observations demonstrate motor equivalence and strengthen the notion that the spatial representation of drawing movements is encoded in the higher brain regions in a rather abstract form that is dissociated from the concrete muscle activation patterns underlying a particular movement direction. To achieve this goal, the central nervous system shifted between two or more muscle grouping strategies to overcome modulations in the interaction among posture-dependent (joint stiffness), dynamic (inertial), and elastic (spring) torque components in the joints. Spring loading induced general changes in the overall amount of EMG activity, which was largely muscle but not direction specific, presumably to represent the posture-dependent biasing force of the spring. Loading was mainly shown to increase muscle coactivation in the elbow joint. This indicates that the subjects tended to increase stiffness in the elbow to compensate for changes in the spring bias forces in order to minimize trajectory errors. Changes in muscle grouping of the shoulder antagonists were mainly a consequence of movement direction but were also affected partly by loading, presumably reflecting the influence of dynamic force components. Taken together, the results confirmed the hypothesis that changes of movement direction and direction of force in the end-effector generated specific sets of muscle grouping to overcome the dynamic requirements in the joints while keeping the kinematics largely unchanged. This suggests that directional tuning in muscle activity and changes in muscle grouping reflects the formation of appropriate internal models in the CNS that give rise to motor equivalence.