Charalambos Papaxanthis

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.

Motor Learning Without Doing: Trial-by-Trial Improvement in Motor Performance During Mental Training

Although there is converging experimental and clinical evidences suggesting that mental training with motor imagery can improve motor performance, it is unclear how humans can learn movements through mental training despite the lack of sensory feedback from the body and the environment. In a first experiment, we measured the trial-by-trial decrease in durations of executed movements (physical training group) and mentally simulated movements (motor-imagery training group), by means of training on a multiple-target arm-pointing task requiring high accuracy and speed. Movement durations were significantly lower in posttest compared with pretest after both physical and motor-imagery training. Although both the posttraining performance and the rate of learning were smaller in motor-imagery training group than in physical training group, the change in movement duration and the asymptotic movement duration after a hypothetical large number of trials were identical. The two control groups (eye-movement training and rest groups) did not show change in movement duration. In the second experiment, additional kinematic analyses revealed that arm movements were straighter and faster both immediately and 24 h after physical and motor-imagery training. No such improvements were observed in the eye-movement training group. Our results suggest that the brain uses state estimation, provided by internal forward model predictions, to improve motor performance during mental training. Furthermore, our results suggest that mental practice can, at least in young healthy subjects and if given after a short bout of physical practice, be successfully substituted to physical practice to improve motor performance.

Does order and timing in performance of imagined and actual movements affect the motor imagery process? The duration of walking and writing task

The purpose of the present study was to investigate the effects on the duration of imagined movements of changes in timing and order of performance of actual and imagined movement. Two groups of subjects had to actually execute and imagine a walking and a writing task. The first group first executed 10 trials of the actual movements (block A) and then imagined the same movements at different intervals: immediately after actual movements (block I-1) and after 25 min (I-2), 50 min (I-3) and 75 min (I-4) interval. The second group first imagined and then actually executed the tasks. The duration of actual and imagined movements, recorded by means of an electronic stopwatch operated by the subjects, was analysed. The duration of imagined movements was very similar to those of actual movements, for both tasks, regardless of either the interval elapsed from the actual movements (first group) or the order of performance (second group). However, the variability of imagined movement duration was significantly increased compared to variability of the actual movements, for both motor tasks and groups. The findings give evidence of similar cognitive processes underlying both imagination and actual performance of movement.

Inertial properties of the arm are accurately predicted during motor imagery

In the present study, using the mental chronometry paradigm, we examined the hypothesis that during motor imagery the brain uses a forward internal model of arm inertial properties to predict the motion of the arm in different dynamic states. Seven subjects performed overt and covert arm movements with one (motion around the shoulder joint) and two (motion around both the shoulder and elbow joints) degrees of freedom in the horizontal plane. Arm movements were executed under two loading conditions: without and with an added mass (4kg) attached to the subjects right wrist. Additionally, movements were performed in two different directions, condition which implies changes in the arm inertia due to the inertial anisotropy of the arm. Our analysis was focused on the timing features of overt and covert movements measured by means of an electronic stopwatch. Durations of right-direction arm movements (low inertial resistance) were smaller compared to durations of left-direction arm movements (high inertial resistance). Additionally, loading the arm with an added mass of 4kg significantly changed the dynamics of motion: movements were indeed more prolonged under loaded conditions. In both cases, the duration of simulated movements mirrored that of overtly executed movements. Therefore, neither the inertial anisotropy of the arm nor the addition of an external mass affected the timing correspondence between overt and covert movement execution. These findings suggest that the brain internally represents the inertial properties of the arm and makes use of it both for sensorimotor control and for the generation of motor images.

The inactivation principle: mathematical solutions minimizing the absolute work and biological implications for the planning of arm movements.

An important question in the literature focusing on motor control is to determine which laws drive biological limb movements. This question has prompted numerous investigations analyzing arm movements in both humans and monkeys. Many theories assume that among all possible movements the one actually performed satisfies an optimality criterion. In the framework of optimal control theory, a first approach is to choose a cost function and test whether the proposed model fits with experimental data. A second approach (generally considered as the more difficult) is to infer the cost function from behavioral data. The cost proposed here includes a term called the absolute work of forces, reflecting the mechanical energy expenditure. Contrary to most investigations studying optimality principles of arm movements, this model has the particularity of using a cost function that is not smooth. First, a mathematical theory related to both direct and inverse optimal control approaches is presented. The first theoretical result is the Inactivation Principle, according to which minimizing a term similar to the absolute work implies simultaneous inactivation of agonistic and antagonistic muscles acting on a single joint, near the time of peak velocity. The second theoretical result is that, conversely, the presence of non-smoothness in the cost function is a necessary condition for the existence of such inactivation. Second, during an experimental study, participants were asked to perform fast vertical arm movements with one, two, and three degrees of freedom. Observed trajectories, velocity profiles, and final postures were accurately simulated by the model. In accordance, electromyographic signals showed brief simultaneous inactivation of opposing muscles during movements. Thus, assuming that human movements are optimal with respect to a certain integral cost, the minimization of an absolute-work-like cost is supported by experimental observations. Such types of optimality criteria may be applied to a large range of biological movements.

Mentally represented motor actions in normal aging: I. Age effects on the temporal features of overt and covert execution of actions

The present study examines the temporal features of overt and covert actions as a function of normal aging. In the first experiment, we tested three motor tasks (walking, sit-stand-sit, arm pointing) that did not imply any particular spatiotemporal constraints, and we compared the duration of their overt and covert execution in three different groups of age (mean ages: 22.5, 66.2 and 73.4 years). We found that the ability of generating motor images did not differentiate elderly subjects from young subjects. Precisely, regarding overt and covert durations, subjects presented similarities for the walking and pointing tasks and dissimilarities for the stand-sit-stand task. Furthermore, the timing variability of imagined movements was always greater compared to actual movements and was of the same amount in the three groups of age. In the second experiment, we investigated the effect of age (three groups with mean ages: 22, 64.8 and 73.2 years) upon temporal characteristics of covert and overt movements involving strong spatiotemporal constraints (speed/accuracy trade-off paradigm). During overt execution young and elderly subjects respected Fittss law despite the fact that movement speed progressively decreased with age. Thus, while execution is deteriorated, the motor preparation process is still intact in old age, and follows well-known laws of biological motions. For covert execution, movement speed progressively decreased with age but elderly subjects did not respect Fittss law. This suggests that the generation and control of motor intentions that consciously do not come to execution, particularly those concerning complex motor actions are progressively perturbed in the aging brain.

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.

Coordination between equilibrium and hand trajectories during whole body pointing movements

We examined the coordination between equilibrium and voluntary pointing movements executed from the standing position, using the whole body. It has previously been shown that trunk movement has little effect upon kinematic characteristics of hand pointing when movements are executed in the sitting position. The present study asked if elements of hand trajectory are modified by requirements of large trunk displacements and fine equilibrium control when pointing movements are executed from the standing position. To achieve this, center of pressure (CoP) and center of mass (CoM) displacements were analyzed along with the kinematics of the pointing hand. Results showed that the CoM was not stabilized (it displaced between 23% and 61±21% of the foot’s length), confirming that instead of a compensation of mechanical perturbations due to arm and trunk movements, the present equilibrium strategy consisted of controlling CoM acceleration towards the target. Hand paths were curved and were not distance or speed invariant. Rather than simple inefficiencies in programming or execution, path curvature suggested that different hand movement strategies were chosen as a function of equilibrium constraints. In light of these results, we hypothesize that postural stability may play a role in the generation of hand trajectory for complex, whole-body pointing movements, in addition to constraints placed upon end-effector kinematics or the dynamic optimization of upper-limb movements. A dependent regulation of equilibrium and spatial components of the movement is proposed.

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.

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

Abstract 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 saggital 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.