Composition and decomposition in bimanual dynamic learning.

Our ability to skillfully manipulate an object often involves the motor system learning to compensate for the dynamics of the object. When the two arms learn to manipulate a single object they can act cooperatively, whereas when they manipulate separate objects they control each object independently. We examined how learning transfers between these two bimanual contexts by applying force fields to the arms. In a coupled context, a single dynamic is shared between the arms, and in an uncoupled context separate dynamics are experienced independently by each arm. In a composition experiment, we found that when subjects had learned uncoupled force fields they were able to transfer to a coupled field that was the sum of the two fields. However, the contribution of each arm repartitioned over time so that, when they returned to the uncoupled fields, the error initially increased but rapidly reverted to the previous level. In a decomposition experiment, after subjects learned a coupled field, their error increased when exposed to uncoupled fields that were orthogonal components of the coupled field. However, when the coupled field was reintroduced, subjects rapidly readapted. These results suggest that the representations of dynamics for uncoupled and coupled contexts are partially independent. We found additional support for this hypothesis by showing significant learning of opposing curl fields when the context, coupled versus uncoupled, was alternated with the curl field direction. These results suggest that the motor system is able to use partially separate representations for dynamics of the two arms acting on a single object and two arms acting on separate objects.

Reflex contributions to the directional tuning of arm stiffness

It has been shown that during arm movement, humans selectively change the endpoint stiffness of their arm to compensate for the instability in an unstable environment. When the direction of the instability is rotated with respect to the direction of movement, it was found that humans modify the antisymmetric component of their endpoint stiffness. The antisymmetric component of stiffness arises due to reflex responses suggesting that the subjects may have tuned their reflex responses as part of the feedforward adaptive control. The goal of this study was to examine whether the CNS modulates the gain of the reflex response for selective tuning of endpoint impedance. Subjects performed reaching movements in three unstable force fields produced by a robotic manipulandum, each field differing only in the rotational component. After subjects had learned to compensate for the field, allowing them to make unperturbed movements to the target, the endpoint stiffness of the arm was estimated in the middle of the movements. At the same time electromyographic activity (EMG) of six arm muscles was recorded. Analysis of the EMG revealed differences across force fields in the reflex gain of these muscles consistent with stiffness changes. This study suggests that the CNS modulates the reflex gain as part of the adaptive feedforward command in which the endpoint impedance is selectively tuned to overcome environmental instability. © 2008 Springer-Verlag Berlin Heidelberg.

Task-dependent coordination of rapid bimanual motor responses.

Optimal feedback control postulates that feedback responses depend on the task relevance of any perturbations. We test this prediction in a bimanual task, conceptually similar to balancing a laden tray, in which each hand could be perturbed up or down. Single-limb mechanical perturbations produced long-latency reflex responses ("rapid motor responses") in the contralateral limb of appropriate direction and magnitude to maintain the tray horizontal. During bimanual perturbations, rapid motor responses modulated appropriately depending on the extent to which perturbations affected tray orientation. Specifically, despite receiving the same mechanical perturbation causing muscle stretch, the strongest responses were produced when the contralateral arm was perturbed in the opposite direction (large tray tilt) rather than in the same direction or not perturbed at all. Rapid responses from shortening extensors depended on a nonlinear summation of the sensory information from the arms, with the response to a bimanual same-direction perturbation (orientation maintained) being less than the sum of the component unimanual perturbations (task relevant). We conclude that task-dependent tuning of reflexes can be modulated online within a single trial based on a complex interaction across the arms.

A Self-Organizing Neural Model of Motor Equivalent Reaching and Tool Use by a Multijoint Arm

This paper describes a self-organizing neural model for eye-hand coordination. Called the DIRECT model, it embodies a solution of the classical motor equivalence problem. Motor equivalence computations allow humans and other animals to flexibly employ an arm with more degrees of freedom than the space in which it moves to carry out spatially defined tasks under conditions that may require novel joint configurations. During a motor babbling phase, the model endogenously generates movement commands that activate the correlated visual, spatial, and motor information that are used to learn its internal coordinate transformations. After learning occurs, the model is capable of controlling reaching movements of the arm to prescribed spatial targets using many different combinations of joints. When allowed visual feedback, the model can automatically perform, without additional learning, reaches with tools of variable lengths, with clamped joints, with distortions of visual input by a prism, and with unexpected perturbations. These compensatory computations occur within a single accurate reaching movement. No corrective movements are needed. Blind reaches using internal feedback have also been simulated. The model achieves its competence by transforming visual information about target position and end effector position in 3-D space into a body-centered spatial representation of the direction in 3-D space that the end effector must move to contact the target. The spatial direction vector is adaptively transformed into a motor direction vector, which represents the joint rotations that move the end effector in the desired spatial direction from the present arm configuration. Properties of the model are compared with psychophysical data on human reaching movements, neurophysiological data on the tuning curves of neurons in the monkey motor cortex, and alternative models of movement control.

Influence of predominant patterns of coordination on the exploitation of interaction torques in a two-joint rhythmic arm movement

In this study we investigate the coordination between rhythmic flexion-extension (FE) and supination-pronation (SP) movements at the elbow joint-complex, while manipulating the intersegmental dynamics by means of a 2-degrees of freedom (df) robot arm. We hypothesized that constraints imposed by the structure of the neuromuscular-skeletal system would (1) result in predominant pattern(s) of coordination in the absence of interaction torques and (2) influence the capabilities of participants to exploit artificially induced interaction torques. Two experiments were conducted in which different conditions of interaction torques were applied on the SP-axis as a function of FE movements. These conditions promoted different patterns of coordination between the 2-df. Control trials conducted in the absence of interaction torques revealed that both the in-phase (supination synchronized with flexion) and the anti-phase (pronation synchronized with flexion) patterns were spontaneously established by participants. The predominance of these patterns of coordination is explained in terms of the mechanical action of bi-articular muscles acting at the elbow joint-complex, and in terms of the reflexes that link the activity of the muscles involved. Results obtained in the different conditions of interaction torques revealed that those neuromuscular-skeletal constraints either impede or favor the exploitation of intersegmental dynamics depending on the context. Interaction torques were indeed found to be exploited to a greater extent in conditions in which the profiles of interaction torques favored one of the two predominant patterns of coordination (i.e., in-phase or anti-phase) as opposed to other patterns of coordination (e.g., 90 degrees or 270 degrees). Those results are discussed in relation to recent studies reporting exploitation of interaction torques in the context of rhythmic movements.

Neuromuscular-skeletal origins of predominant patterns of coordination in a rhythmic two-joint arm movement

The authors tested for predominant patterns of coordination in the combination of rhythmic flexion-extension (FE) and supination-pronation (SP) at the elbow-joint complex. Participants (N = 10) spontaneously established in-phase (supination synchronized with flexion) and antiphase (pronation synchronized with flexion) patterns. In addition, the authors used a motorized robot arm to generate involuntary SP movements with different phase relations with respect to voluntary FE. The involuntarily induced in-phase pattern was accentuated and was more consistent than other patterns. That result provides evidence that the predominance of the in-phase pattern originates in the influence of neuro-muscular-skeletal constraints rather than in a preference dictated by perceptual-cognitive factors implicated in voluntary control. Neuromuscular-skeletal constraints involved in the predominance of the in-phase and the antiphase patterns are discussed.

Visual feedback alters the variations in corticospinal excitability that arise from rhythmic movements of the opposite limb

Augmented visual feedback can have a profound bearing on the stability of bimanual coordination. Indeed, this has been used to render tractable the study of patterns of coordination that cannot otherwise be produced in a stable fashion. In previous investigations (Carson et al. 1999), we have shown that rhythmic movements, brought about by the contraction of muscles on one side of the body, lead to phase-locked changes in the excitability of homologous motor pathways of the opposite limb. The present study was conducted to assess whether these changes are influenced by the presence of visual feedback of the moving limb. Eight participants performed rhythmic flexion-extension movements of the left wrist to the beat of a metronome (1.5 Hz). In 50% of trials, visual feedback of wrist displacement was provided in relation to a target amplitude, defined by the mean movement amplitude generated during the immediately preceding no feedback trial. Motor potentials (MEPs) were evoked in the quiescent muscles of the right limb by magnetic stimulation of the left motor cortex. Consistent with our previous observations, MEP amplitudes were modulated during the movement cycle of the opposite limb. The extent of this modulation was, however, smaller in the presence of visual feedback of the moving limb (FCR omega(2) =0.41; ECR omega(2)=0.29) than in trials in which there was no visual feedback (FCR omega(2)=0.51; ECR omega(2)=0.48). In addition, the relationship between the level of FCR activation and the excitability of the homologous corticospinal pathway of the opposite limb was sensitive to the vision condition; the degree of correlation between the two variables was larger when there was no visual feedback of the moving limb. The results of the present study support the view that increases in the stability of bimanual coordination brought about by augmented feedback may be mediated by changes in the crossed modulation of excitability in homologous motor pathways.

Bimanual coordination: constraints imposed by the relative timing of homologous muscle activation

It has often been supposed that patterns of rhythmic bimanual coordination in which homologous muscles are engaged simultaneously, are performed in a more stable manner than those in which the same muscles are activated in an alternating fashion. In order to assess the efficacy of this constraint, the present study investigated the effect of forearm posture (prone or supine) on bimanual abduction-adduction movements of the wrist in isodirectional and non-isodirectional modes of coordination. Irrespective of forearm posture, non-isodirectional coordination was observed to be more stable than isodirectional coordination. In the latter condition, there was a more severe deterioration of coordination accuracy/stability as a function of cycling frequency than in the former condition. With elevations in cycling frequency, the performers recruited extra mechanical degrees of freedom, principally via flexion-extension of the wrist, which gave rise to increasing motion in the vertical plane. The increases in movement amplitude in the vertical plane were accompanied by decreasing amplitude in the horizontal plane. In agreement with previous studies, the present findings confirm that the relative timing of homologous muscle activation acts as a principal constraint upon the stability of interlimb coordination. Furthermore, it is argued that the use of manipulations of limb posture to investigate the role of other classes of constraint (e.g. perceptual) should be approached with caution because such manipulations affect the mapping between muscle activation patterns, movement dynamics and kinematics.

Interaction of directional, neuromuscular and egocentric constraints on the stability of preferred bimanual coordination patterns

We investigated how the relative direction of limb movements in external space (iso- and non-isodirectionality), muscular constraints (the relative timing of homologous muscle activation) and the egocentric frame of reference (moving simultaneously toward/away the longitudinal axis of the body) contribute to the stability of coordinated movements. In the first experiment, we attempted to determine the respective stability of isodirectional and non-isodirectional movements in between-persons coordination. In a second experiment, we determined the effect of the relative direction in external space, and of muscular constraints, on pattern stability during a within-person bimanual coordination task. In the third experiment we dissociated the effects on pattern stability of the muscular constraints, relative direction and egocentric frame of reference. The results showed that (1) simultaneous activation of homologous muscles resulted in more stable performance than simultaneous activation of non-homologous muscles during within-subject coordination, and that (2) isodirectional movements were more stable than non-isodirectional movements during between-persons coordination, confirming the role of the relative direction of the moving limbs in the stability of bimanual coordination. Moreover, the egocentric constraint was to some extent found distinguishable from the effect of the relative direction of the moving limbs in external space, and from the effect of the relative timing of muscle activation. In summary, the present study showed that relative direction of the moving limbs in external space and muscular constraints may interact either to stabilize or destabilize coordination patterns. (C) 2003 Published by Elsevier B.V.

The effect of volition on the stability of bimanual coordination

The authors investigated how the intention to passively perform a behavior and the intention to persist with a behavior impact upon the spatial and temporal properties of bimanual coordination. Participants (N = 30) were asked to perform a bimanual coordination task that demanded the continuous rhythmic extension-flexion of the wrists. The frequency of movement was scaled by an auditory metronome beat from 1.5 Hz, increasing to 3.25 Hz in .25-Hz increments. The task was further defined by the requirement that the movements be performed initially in a prescribed pattern of coordination (in-phase or antiphase) while the participants assumed one of two different intentional states: stay with the prescribed pattern should it become unstable or do not intervene should the pattern begin to change. Transitions away from the initially prescribed pattern were observed only in trials conducted in the antiphase mode of coordination. The time at which the antiphase pattern of coordination became unstable was not found to be influenced by the intentional state. In addition, the do-not-intervene set led to a switch to an in-phase pattern of coordination whereas the stay set led to phase wandering. Those findings are discussed within the framework of a dynamic account of bimanual coordination.

The acquisition of movement skills: Practice enhances the dynamic stability of bimanual coordination

During bimanual movements, two relatively stable

Bimanual coordination in chronic schizophrenia

Anomalies of movement are observed both clinically and experimentally in schizophrenia. While the basal ganglia have been implicated in its pathogenesis, the nature of such involvement is equivocal. The basal ganglia may be involved in bimanual coordination through their input to the supplementary motor area (SMA). While a neglected area of study in schizophrenia. a bimanual movement task may provide a means of assessing the functional integrity of the motor circuit. Twelve patients with chronic schizophrenia and 12 matched control participants performed a bimanual movement task on a set of vertically mounted cranks at different speeds (1 and 2 Hz) and phase relationships. Participants performed in-phase movements (hands separated by 0 degrees) and out-of-phase movements (hands separated by 180 degrees) at both speeds with an external cue on or off. All participants performed the in-phase movements well. irrespective of speed or cueing conditions. Patients with schizophrenia were unable to perform the out-of-phase movements, particularly at the faster speed, reverting instead to the in-phase movement. There was no effect of external cueing on any of the movement conditions. These results suggest a specific problem of bimanual coordination indicative of SMA dysfunction per se and/or faulty callosal integration. A disturbance in the ability to switch attention during the out-of-phase task may also be involved. (C) 2001 Academic Press.

Interval timing and trajectory in unequal amplitude movements

The Wing-Kristofferson (WK) model of movement timing emphasises the separation of central timer and motor processes. Several studies of repetitive timing have shown that increase in variability at longer intervals is attributable to timer processes; however, relatively little is known about the way motor aspects of timing are affected by task movement constraints. In the present study, we examined timing variability in finger tapping with differences in interval to assess central timer effects, and with differences in movement amplitude to assess motor implementation effects. Then, we investigated whether effects of motor timing observed at the point of response (flexion offset/tap) are also evident in extension, which would suggest that both phases are subject to timing control. Eleven participants performed bimanual simultaneous tapping, at two target intervals (400, 600 ms) with the index finger of each hand performing movements of equal (3 or 6 cm) or unequal amplitude (left hand 3, right hand 6 cm and vice versa). As expected, timer variability increased with the mean interval but showed only small, non-systematic effects with changes in movement amplitude. Motor implementation variability was greater in unequal amplitude conditions. The same pattern of motor variability was observed both at flexion and extension phases of movement. These results suggest that intervals are generated by a central timer, triggering a series of events at the motor output level including flexion and the following extension, which are explicitly represented in the timing system.

Evidence for a reference frame transformation of vestibular contributions to voluntary reaching movements

Les estimations des mouvements de soi provenant des signaux vestibulaires contribuent à la planification et l’exécution des mouvements volontaires du bras lorsque le corps se déplace. Cependant, comme les senseurs vestibulaires sont fixés à la tête alors que le bras est fixé au corps, les signaux vestibulaires doivent être transformés d’un système de référence centré sur la tête à un système centré sur le corps pour pouvoir contribuer de façon appropriée au contrôle moteur du bras. Le but premier de ce travail était d’étudier l’évidence d’une telle transformation. La stimulation galvanique vestibulaire (SGV) a été utilisée pour activer les afférences vestibulaires et simuler une rotation autour d’un axe naso-occipital fixe pendant que des sujets humains faisaient des mouvements du bras dans le plan horizontal, avec la tête dans différentes orientations. Une transformation des signaux vestibulaires implique que la SVG devrait simuler une rotation autour d’un axe horizontal lorsque la tête est droite et autour d’un axe vertical lorsque la tête est en flexion antérieure. La SGV devrait ainsi perturber les mouvements du bras en fonction de l’orientation de la tête. Nos résultats démontrent que les signaux vestibulaires contribuant aux mouvements d’atteinte sont effectivement transformés en un système de référence centrée sur le corps. Le deuxième but de ce travail était d’explorer les mécanismes utilisant ces signaux vestibulaires transformés. En comparant les effets de la SGV appliquée avant ou pendant les mouvements d’atteinte nous avons montré que les signaux vestibulaires transformés contribuent à des mécanismes de compensation distincts durant la planification des mouvements d’atteinte comparativement à l’exécution.

Midpoint perturbation response in haptically-guided movements

This study investigates the human response to impulse perturbations at the midpoint of a haptically-guided straight-line point-to-point movement. Such perturbation response may be used as an assessment tool during robot-mediated neuro-rehabilitation therapy. Subjects show variety in their perturbation responses. Movements with a lower perturbation displacement exhibit high frequency oscillations, indicative of increased joint stiffness. Equally, movements with a high perturbation displacement exhibit lower frequency oscillations with higher amplitude and a longer settling time. Some subjects show unexpected transients during the perturbation impulse, which may be caused by complex joint interactions in the hand and arm.