Does the Motor Cortex Want the Full Story? The Influence of Sentence Context on Corticospinal Excitability in Action Language Processing.

The reading of action verbs has been shown to activate motor areas, whereby sentence context may serve to either globally strengthen this activation or to selectively sharpen it. To investigate this issue, we manipulated the presence of manual actions and sentence context, assessing the level of corticospinal excitability by means of transcranial magnetic stimulation. We hypothesized that context would serve to sharpen the neural representation of the described actions in the motor cortex, reflected in context-specific modulation of corticospinal excitability. Participants silently read manual action verbs and non-manual verbs, preceded by a full sentence (rich context) or not (minimal context). Transcranial magnetic stimulation pulses were delivered at rest or shortly after verb presentation. The coil was positioned over the cortical representation of the right first dorsal interosseous (pointer finger). We observed a general increase of corticospinal excitability while reading both manual action and non-manual verbs in minimal context, whereas the modulation was action-specific in rich context: corticospinal excitability increased while reading manual verbs, but did not differ from baseline for non-manual verbs. These findings suggest that sentence context sharpens motor representations, activating the motor cortex when relevant and eliminating any residual motor activation when no action is present.

Evidence for subjective values guiding posture and movement coordination in a free-endpoint whole-body reaching task.

When moving, humans must overcome intrinsic (body centered) and extrinsic (target-related) redundancy, requiring decisions when selecting one motor solution among several potential ones. During classical reaching studies the position of a salient target determines where the participant should reach, constraining the associated motor decisions. We aimed at investigating implicit variables guiding action selection when faced with the complexity of human-environment interaction. Subjects had to perform whole body reaching movements towards a uniform surface. We observed little variation in the self-chosen motor strategy across repeated trials while movements were variable across subjects being on a continuum from a pure 'knee flexion' associated with a downward center of mass (CoM) displacement to an 'ankle dorsi-flexion' associated with an upward CoM displacement. Two optimality criteria replicated these two strategies: a mix between mechanical energy expenditure and joint smoothness and a minimization of the amount of torques. Our results illustrate the presence of idiosyncratic values guiding posture and movement coordination that can be combined in a flexible manner as a function of context and subject. A first value accounts for the reach efficiency of the movement at the price of selecting possibly unstable postures. The other predicts stable dynamic equilibrium but requires larger energy expenditure and jerk.

Direction-dependent activation of the insular cortex during vertical and horizontal hand movements.

The planning of any motor action requires a complex multisensory processing by the brain. Gravity - immutable on Earth - has been shown to be a key input to these mechanisms. Seminal fMRI studies performed during visual perception of falling objects and self-motion demonstrated that humans represent the action of gravity in parts of the cortical vestibular system; in particular, the insular cortex and the cerebellum. However, little is known as to whether a specific neural network is engaged when processing non-visual signals relevant to gravity. We asked participants to perform vertical and horizontal hand movements without visual control, while lying in a 3T-MRI scanner. We highlighted brain regions activated in the processing of vertical movements, for which the effects of gravity changed during execution. Precisely, the left insula was activated in vertical movements and not in horizontal movements. Moreover, the network identified by contrasting vertical and horizontal movements overlapped with neural correlates previously associated to the processing of simulated self-motion and visual perception of the vertical direction. Interestingly, we found that the insular cortex activity is direction-dependent which suggests that this brain region processes the effects of gravity on the moving limbs through non-visual signals.

Effect of mental fatigue on speed-accuracy trade-off.

The aim of this study was to investigate the effects of mental fatigue on the duration of actual and imagined goal-directed arm movements involving speed-accuracy trade-off. Ten participants performed actual and imagined point-to-point arm movements as accurately and as fast as possible, before and after a 90-min sustained cognitive task inducing mental fatigue, and before and after viewing a neutral control task (documentary movie) that did not induce mental fatigue. Target width and center-to-center target distance were varied, resulting in five different indexes of difficulty. Prior to mental fatigue, actual and imagined movement duration increased with the difficulty of the task, as predicted by Fitts' law. Mental fatigue task induced a 4.1±0.7% increase in actual movement duration and a 9.6±1.1% increase in imagined movement duration, independently of the index of difficulty. The trial-by-trial evolution of actual and imagined movement duration remained stable with mental fatigue. The control task did not induce any change in actual and imagined movement duration. The results suggested that movement was slowed in the presence of mental fatigue, maybe due to proactive changes occurring during the preparatory state of the movement, to preserve task success.

Laterality effects in motor learning by mental practice in right-handers.

Converging evidences suggest that mental movement simulation and actual movement production share similar neurocognitive and learning processes. Although a large body of data is available in the literature regarding mental states involving the dominant arm, examinations for the nondominant arm are sparse. Does mental training, through motor-imagery practice, with the dominant arm or the nondominant arm is equally efficient for motor learning? In the current study, we investigated laterality effects in motor learning by motor-imagery practice. Four groups of right-hander adults mentally and physically performed as fast and accurately as possible (speed/accuracy trade-off paradigm) successive reaching movements with their dominant or nondominant arm (physical-training-dominant-arm, mental-training-dominant-arm, physical-training-nondominant-arm, and mental-training-nondominant-arm groups). Movement time was recorded and analyzed before, during, and after the training sessions. We found that physical and mental practice had a positive effect on the motor performance (i.e., decrease in movement time) of both arms through similar learning process (i.e., similar exponential learning curves). However, movement time reduction in the posttest session was significantly higher after physical practice than motor-imagery practice for both arms. More importantly, motor-imagery practice with the dominant arm resulted in larger and more robust improvements in movement speed compared to motor-imagery practice with the nondominant arm. No such improvements were observed in the control group. Our results suggest a superiority of the dominant arm in motor learning by mental practice. We discussed these findings from the perspective of the internal models theory.

Modulation of spinal excitability by a sub-threshold stimulation of M1 area during muscle lengthening.

It is well known that the H-reflex amplitude decreases during passive muscle lengthening in comparison with passive shortening. However, this decrease in spinal synaptic efficacy observed during passive lengthening seems to be lesser during eccentric voluntary contraction. The aim of the present study was to examine whether spinal excitability during lengthening condition could be modulated by magnetic brain stimulation. H reflexes of the triceps surae muscles were elicited on 10 young healthy subjects, and conditioned by a sub-threshold transcranial magnetic stimulation (TMS). The conditioning stimulation was applied over the M1 area of triceps surae muscles at an intensity below motor threshold with a conditioning-test interval of 5ms. Conditioned and non-conditioned H-reflexes were elicited at rest, during passive lengthening and shortening, and during submaximal contractions (concentric, eccentric and isometric). During passive and active lengthening, H reflexes conditioned by a sub-threshold TMS pulse increased on average by 50% compared with non-conditioned responses. No significant effect was found during isometric and concentric conditions. Activation of the corticospinal pathway would partially cancel inhibitions caused by muscle stretch, and according to the time-delayed effect, this result suggested the existence of a specific polysynaptic pathway. In additional experiments, H responses were conditioned by cervico-medullary stimulations, showing that the modulation described by the previous results involves subcortical mechanisms. This study provides further evidences that the modulation of the final cortico-spinal command reaching the muscle depends on a central mechanism that controls peripheral input, such as Ia afference discharge during lengthening.

Elderly adults delay proprioceptive reweighting during the anticipation of collision avoidance when standing.

The ability to reweight visual and proprioceptive information is critical for maintaining postural stability in a dynamic environment. In this study, we examined whether visual anticipation of collision avoidance (AV) while standing could facilitate the down-weighting of altered proprioception in young and elderly adults. Twelve young (24.91±6.44years) and 12 elderly (74.8±6.42years) participants stood upright for 180s under two task conditions: (a) quiet stance (QS) and (b) standing while anticipating virtual objects to be avoided. In order to disrupt the accuracy of proprioceptive input participants were exposed to bilateral Achilles tendon vibration during the middle 60s of standing in both tasks. Visual field dependence was assessed using the Rod and Frame Test (RFT). Elderly demonstrated significantly higher visual field dependence compared to the young participants. Analysis of the normalized Root Mean Square (RMS) of the Center of Pressure velocity (dCoP) revealed that young participants immediately reduced the sway velocity variability induced by tendon vibration during the anticipation of collision AV compared to the QS task. In the elderly, however, the modulating influence of visual anticipation was delayed and became significant only in the last two time intervals of the vibration phase. These results suggest that volitionally shifting reliance on vision when anticipating a collision AV event facilitates the down-weighting of altered proprioception. Elderly adults seem to be unable to dynamically exploit visual anticipation in order to down weight the altered proprioception possibly as a result of their more permanent up-weighting of the visual modality. Sensory reweighting seems to be a more time consuming process in aging which may have important clinical implications for falling.

A model of the cerebellar sensory--motor control applied to fast human forearm movements.

To address the problem of how the cerebellum processes the premotor orders that control fast movements of the forearm, a model of the cerebellar control is proposed: a cybernetic circuit composed of a model of the cerebellar premotor pathways driving a biomechanical model of the human forearm. Experiments consist of recording electromyographic (EMG) activities and cinematic variables of the human forearm during fast, single joint, point-to-point movements performed in horizontal and vertical directions with and without mass. The biomechanical model of the forearm is first validated by comparing actual movements and movements simulated by using, as inputs to this model, the synthesized EMG signals and of real EMG activities recorded during the experiments. Then the entire control model is validated by comparing actual movements to the desired ones simulated by the model of the cerebellar pathways whose inputs are velocity signals with Gaussian time-courses. The results show that approximate inverse functions can be computed by means of inner models of direct functions placed in feedback loops, and suggest that the orientation of any member segment with respect to gravity is computed as a cinematic variable in the Central Nervous System (CNS).

Motor planning of arm movements is direction-dependent in the gravity field.

In the present study we analyzed kinematic and dynamic features of arm movements in order to better elucidate how the motor system integrates environmental constraints (gravity) into motor planning and control processes. To reach this aim, we experimentally manipulated the mechanical effects of gravity on the arm while maintaining arm inertia constant (i.e. the distribution of the mass around the shoulder joint). Six subjects performed single-joint arm movements (rotation around the shoulder joint) in both sagittal (upward, U, versus downward, D) and horizontal (left, L, versus right, R) planes, at different amplitudes and from different initial positions. Under these conditions, shoulder gravitational torques (SGTs) significantly varied when arm movements were performed in the sagittal but not in the horizontal plane. Contrary to SGTs, arm inertia remained constant and similar for both horizontal and sagittal planes since subjects performed arm movements with only one degree of freedom. All subjects, whatever the movement direction, appropriately scaled shoulder joint kinematic parameters according to movement amplitude. Furthermore, peak velocity and movement duration were equivalent for both horizontal and sagittal planes. Interestingly, some kinematic parameters significantly differed according to U/D but not L/R directions. Specifically, acceleration duration was greater for D than U movements, while the opposite was true for peak acceleration. Consequently, although vertical and horizontal arm movements shared a general common strategy (i.e. scaling law), the kinematic asymmetries between U and D arm movements, especially those that reflect central planning process (i.e. peak acceleration), indicated different motor intentions regarding the direction of the upcoming movement. These findings indicate that the interaction of the arm with the dynamics of the environment is internally represented during the generation of arm trajectories.

WITHDRAWN: A model of the cerebellar sensory-motor control applied to the fast human forearm movements.

This article has been withdrawn consistent with Elsevier Policy on Article Withdrawal (http://www.elsevier.com/locate/withdrawalpolicy). The Publisher apologizes for any inconvenience this may cause.

Improvement and generalization of arm motor performance through motor imagery practice.

This study compares the improvement and generalization of arm motor performance after physical or mental training in a motor task requiring a speed-accuracy tradeoff. During the pre- and post-training sessions, 40 subjects pointed with their right arm as accurately and as fast as possible toward targets placed in the frontal plane. Arm movements were performed in two different workspaces called right and left paths. During the training sessions, which included only the right path, subjects were divided into four training groups (n = 10): (i) the physical group, subjects overtly performed the task; (ii) the mental group, subjects imagined themselves performing the task; (iii) the active control group, subjects performed eye movements through the targets, (iv) the passive control group, subjects did not receive any specific training. We recorded movement duration, peak acceleration and electromyographic signals from arm muscles. Our findings showed that after both physical and mental training on the right path (training path), hand movement duration and peak acceleration respectively decreased and increased for this path. However, motor performance improvement was greater after physical compared with mental practice. Interestingly, we also observed a partial learning generalization, namely an enhancement of motor performance for the left path (non-training path). The amount of this generalization was roughly similar for the physical and mental groups. Furthermore, while arm muscle activity progressively increased during the training period for the physical group, the activity of the same muscles for the mental group was unchanged and comparable with that of the rest condition. Control groups did not exhibit any improvement. These findings put forward the idea that mental training facilitates motor learning and allows its partial transfer to nearby workspaces. They further suggest that motor prediction, a common process during both actual and imagined movements, is a fundamental operation for both sensorimotor control and learning.

Comparison of motor strategies in sit-to-stand and back-to-sit motions between healthy and Alzheimer's disease elderly subjects.

We studied the kinematics of shoulder displacement during sit-to-stand and back-to-sit in 6 healthy elderly subjects and six elderly subjects with mild to moderate Alzheimer's disease in order to elucidate the impact of Alzheimer's disease on motor planning and control processes. During sit-to-stand, Alzheimer's disease subjects reduced their forward displacement and started their upward displacement earlier than healthy elderly subjects. Furthermore, shoulder path curvatures were more pronounced for upward compared with downward displacement in healthy elderly group, in contrast with Alzheimer's disease group. Temporal analysis found that: 1) for both groups, profiles of velocity of sit-to-stand and back-to-sit showed two peaks corresponding respectively to forward/upward and to downward/backward displacements, 2) peaks of velocity were almost comparable between the two groups, 3) duration of sit-to-stand was shorter than duration of back-to-sit in the two groups and 4) duration of sit-to-stand and back-to-sit was shorter in Alzheimer's disease group than in healthy elderly group. However, dissimilarities were observed for transition and deceleration phases during sit-to-stand, and for acceleration and transition phases during back-to-sit, between the two groups. Interestingly, while sit-to-stand and back-to-sit differed in healthy elderly subjects during transition and deceleration phases, such a difference was not observed for Alzheimer's disease subjects. So, our study showed that invariant spatio-temporal movement parameters in the two groups differed, while non-invariant parameters did not, and suggests that higher level motor process of whole body motions are affected by Alzheimer's disease, while lower level motor features remain intact.

Kinematic and dynamic processes for the control of pointing movements in humans revealed by short-term exposure to microgravity.

The generation of accurate motor commands requires implicit knowledge of both limb and environmental dynamics. The action of gravity on moving limb segments must be taken into account within the motor command, and may affect the limb trajectory chosen to accomplish a given motor task. Exactly how the CNS deals with these gravitoinertial forces remains an open question. Does the CNS measure gravitational forces directly, or are they accommodated in the motor plan by way of internal models of physical laws? In this study five male subjects participated. We measured kinematic and dynamic parameters of upward and downward arm movements executed at two different speeds, in both normal Earth gravity and in the weightless conditions of parabolic flight. Exposure to microgravity affected velocity profiles for both directions and speeds. The shape of velocity profiles (the ratio of maximum to mean velocity) and movement duration both showed transient perturbations initially in microgravity, but returned to normal gravity values with practice in 0 x g. Differences in relative time to peak velocity between upward versus downward movements, persisted for all trial performed in weightlessness. These differences in kinematic profiles and in the torque profiles used to produce them, diminished, however, with practice in 0 x g. These findings lead to the conclusion that the CNS explicitly represents gravitational and inertial forces in the internal models used to generate and execute arm movements. Furthermore, the results suggest that the CNS adapts motor plans to novel environments on different time scales; dynamics adapt first to reproduce standard kinematics, and then kinematics patterns are adapted to optimize dynamics.

Similar planning strategies for whole-body and arm movements performed in the sagittal plane.

The present paper looks for kinematic similarities between whole-body and arm movements executed in the sagittal plane. Eight subjects performed sit-to-stand (STS) and back-to-sit (BTS) movements at their preferred speed in the sagittal plane. Kinematics analysis focused on shoulder motion revealed that STS was composed of a straight, forward displacement followed by a curved, upward displacement while BTS was characterized by a curved, downward and straight, backward displacement. Curvature of the upward displacement was significantly greater than the downward one. Analysis of shoulder-velocity profiles showed that movement duration was significantly longer for BTS compared with STS and that the shape of the velocity profiles changed when subjects performed an STS compared with a BTS movement. Velocity profiles of the upward and downward displacements also differed; the relative acceleration duration (acceleration duration divided by movement duration during the vertical motion) was smaller for the upward compared with the downward displacement. The present results are in accordance with previous findings concerning the execution of vertical arm movements and suggest that the CNS uses similar motor plans for the performance of arm and whole-body movements in the sagittal plane.

Investigating centre of mass stabilisation as the goal of posture and movement coordination during human whole body reaching.

In the light of experimental results showing significant forward centre of mass (CoM) displacements within the base of support, this study investigated if whole body reaching movements can be executed whilst keeping the CoM fixed in the horizontal axis. Using kinematic simulation techniques, angular configurations were recreated from experimental data imposing two constraints: a constant horizontal position of the CoM and an identical trajectory of the hand to grasp an object. The comparison between recorded and simulated trials showed that stabilisation of the CoM was associated with greater backward hip displacements, which became more marked with increasing object distance. This was in contrast to recorded trials showing reductions in backward hip displacements with increasing distance. Results also showed that modifications to angular displacements were necessary only at the shoulder and hip joints, but that these modifications were within the limits of joint mobility. The analysis of individual joint torques revealed that the pattern and timing of simulated trials were similar to those recorded experimentally. Peak joint torque values showed particularly that keeping the CoM at a constant horizontal position resulted in significantly smaller ankle peak flexor and extensor torques. It may be concluded from this study that 'stabilising' the CoM during human whole body reaching represents a feasible strategy, but not the one chosen by subjects under experimental conditions. Our results also do not support the idea of the CoM as the stabilised reference value for the coordination between posture and goal-directed movements.

The sensorimotor and cognitive integration of gravity.

In order to demonstrate that gravity is not only a load acting locally and continuously on the body limbs, but is also used by higher levels of the nervous system as a dynamic orienting reference for the elaboration of the motor act, a review of several experiments conducted both in 1 g and 0 g are presented. During various locomotor tasks, the strategy that consists of stabilizing the head with respect to gravity illustrates one of the solutions used by the CNS to optimize the control of dynamic equilibrium. A question which remains to be solved when considering experimental results obtained in weightlessness concerns, however, the maintenance of motor schema that has evolved under normal gravity. Results have suggested that the concept of conservative processes, that would adapt postural control to weightlessness by using previously learned innate strategies, must be reconsidered during goal-oriented tasks. In fact, it is proposed that when conservative processes and existing solutions derived from a repertoire of terrestrial postural strategies do not provide efficient output, the CNS has to create novel strategies through a slow learning process. As with the study of postural control, three-dimensional arm reaching movements also illustrate the central representation of gravity. Indeed, gravity can be regarded as either initiating or braking arm movements and, consequently, may be represented in the motor command at the planning level. Finally, from a prospective point of view, there is a need to determine new experimental paradigms in order to study the specific motor control of man in space. It is suggested that the formulation of experimental paradigms should not consider man in space simply as a terrestrial biped.

Effects of movement direction upon kinematic characteristics of vertical arm pointing movements in man.

Vertical arm pointing movements in two directions (upwards and downwards), imposing two different loads (unload and 0.5 kg) and speeds (normal and fast) have been studied in six subjects. Movements were recorded using an optoelectronic system. Data analysis concentrated upon finger-tip kinematics. Significant effects of movement direction were recorded upon velocity profiles. The acceleration time, computed relative to total movement time, was greater for downward movements than for upward movements. In contrast however, no effects of load or speed were observed. Movement time was not affected by movement direction or load, for both speeds tested. These results suggest different planning processes, for movements with and against gravity and indicate that gravitational force influences the processes controlling movement execution.

[Drawing movements and gravitational force: central or peripheral regulation?]. / Mouvements graphiques et force gravitaire: traitement central ou périphérique?

Drawing arm movements in four different directions: a) upward vertical (0 degree), b) upward oblique (45 degrees), c) downward vertical (180 degrees) and d) downward oblique (135 degrees), and at two different speeds, normal and fast, were executed by eight subjects. Movements of the arm were recorded using an optoelectronic (2 TV, 100 Hz) system which allowed the computer reconstruction of joint motion. Analyses focused upon pen kinematics in the frontal plane. Velocity profiles were unimodal for all conditions. The ratio of acceleration time to total movement time changed significantly as a function of the direction and the speed of the movement. Movement time and was not affected by movement direction and consequently changes in gravitational torques, for both speeds tested. Results from this study provide indirect evidence that the CNS executes movements by taking advantage of gravitational force.

Hand trajectories of vertical arm movements in one-G and zero-G environments. Evidence for a central representation of gravitational force.

The purpose of the present experiment was to study the way in which the central nervous system (CNS), represents gravitational force during vertical arm pointing movements. Movements in upward (against gravity) and downward (with gravity) directions, with two different mass loads (hand empty and with a hand-held 0.5-kg weight) were executed by eight subjects in a normal gravitational environment. Movements by two cosmonauts, in the two directions, were also tested in a state of weightlessness. Analyses focused upon finger trajectories in the sagittal plane. Subjects in a normal gravitational environment showed curved paths for both directions and weight conditions. In addition, downward movements showed significantly smaller curvatures than upward movements. Movement times were approximately the same for all the experimental conditions. Curvature differences between upward and downward movements persisted during space flight and immediately postflight. Movement times from both cosmonauts increased slightly during flight, but returned to normal immediately on reentry in a one-G environment. Results from the present study provide evidence that gravity is centrally represented in an anticipatory fashion as a driving force during vertical arm movement planning.

The representation of gravitational force during drawing movements of the arm.

The purpose of the present experiment was to study the way in which the central nervous system (CNS) represents gravitational force (GF) during vertical drawing movements of the arm. Movements in four different directions: (a) upward vertical (0 degrees), (b) upward oblique (45 degrees), (c) downward vertical (180 degrees) and (d) downward oblique (135 degrees), and at two different speeds, normal and fast, were executed by nine subjects. Data analysis focused upon arm movement kinematics in the frontal plane and gravitational torques (GTs) exerted around the shoulder joint. Regardless of movement direction, subjects showed straight-line paths for both speed conditions. In addition, movement time and peak velocity were not affected by movement direction and consequently changes in GT, for both speeds tested. Movement timing (evaluated through the ratio of acceleration time to total time) changed significantly, however, as a function of movement direction and speed. Upward movements showed shorter acceleration times when compared with downward movements. Concerning the four directions, movements made at 0 degrees and 45 degrees differed significantly from those made at 135 degrees and 180 degrees. Drawing movements executed at rapid speed presented similar acceleration and deceleration times compared with movements executed at normal speed, which showed greater acceleration than deceleration times. In addition, the form of velocity profiles (assessed through the ratio of maximum to mean velocities), was significantly modified only with movement speed. Results from the present study suggest that GF is efficiently incorporated into internal dynamic models that the brain builds up for the execution of arm movements. Furthermore, it seems that GF not only is a mechanical parameter to be overcome by the motor system but also constitutes a reference (vertical direction), both of which are represented by the CNS during inverse kinematic and dynamic processes.