Haptic information stabilizes and destabilizes coordination dynamics

Goal-directed, coordinated movements in humans emerge from a variety of constraints that range from 'high-level' cognitive strategies based oil perception of the task to 'low-level' neuromuscular-skeletal factors such as differential contributions to coordination from flexor and extensor muscles. There has been a tendency in the literature to dichotomize these sources of constraint, favouring one or the other rather than recognizing and understanding their mutual interplay. In this experiment, subjects were required to coordinate rhythmic flexion and extension movements with an auditory metronome, the rate of which was systematically increased. When subjects started in extension on the beat of the metronome, there was a small tendency to switch to flexion at higher rates, but not vice versa. When subjects: were asked to contact a physical stop, the location of which was either coincident with or counterphase to the auditor) stimulus, two effects occurred. When haptic contact was coincident with sound, coordination was stabilized for both flexion and extension. When haptic contact was counterphase to the metronome, coordination was actually destabilized, with transitions occurring from both extension to flexion on the beat and from flexion to extension on the beat. These results reveal the complementary nature of strategic and neuromuscular factors in sensorimotor coordination. They also suggest the presence of a multimodal neural integration process-which is parametrizable by rate and context - in which intentional movement, touch and sound are bound into a single, coherent unit.

The influence of joint position on the dynamics of perception-action coupling

Six right-handed subjects performed rhythmic flexion and extension movements of the index finger in time with an auditory metronome. On each block of trials, the wrist of the response hand was placed in a extended, neutral or flexed position. In the flex-on-the-beat condition, subjects were instructed to coordinate maximum excursion in the direction of finger flexion with each beat of the metronome. In the extend-on-the-beat condition, subjects were instructed to coordinate maximum excursion in the direction of finger extension with each beat of the metronome. The frequency of the metronome was increased from 2.00 Hz to 3.75 Hz in 8 steps (8 s epochs) of 0.25 Hz. During trials prepared in the extend-on-the-beat pattern, all subjects exhibited transitions to either a flex-on-the-beat pattern or to phase wandering as the frequency of pacing was increased. The time at which these transitions occurred was reliably influenced by the position of the wrist. Four subjects exhibited qualitative departures from the flex-on-the-beat pattern at pacing frequencies that were greater than those at which the extend-on-the-beat pattern could be maintained. The lime at which these departures occurred was not influenced by the position of the wrist. These results are discussed with reference to the constraints imposed on the coordination dynamics by the intrinsic properties of the neuromuscular-skeletal system.

A neuro-mechanical model for interpersonal coordination

The present study investigates the coordination between two people oscillating handheld pendulums, with a special emphasis on the influence of the mechanical properties of the effector systems involved. The first part of the study is an experiment in which eight pairs of participants are asked to coordinate the oscillation of their pendulum with the other participant's in an in-phase or antiphase fashion. Two types of pendulums, A and B, having different resonance frequencies (Freq A=0.98 Hz and Freq B=0.64 Hz), were used in different experimental combinations. Results confirm that the preferred frequencies produced by participants while manipulating each pendulum individually were close to the resonance frequencies of the pendulums. In their attempt to synchronize with one another, participants met at common frequencies that were influenced by the mechanical properties of the two pendulums involved. In agreement with previous studies, both the variability of the behavior and the shift in the intended relative phase were found to depend on the task-effector asymmetry, i.e., the difference between the mechanical properties of the effector systems involved. In the second part of the study, we propose a model to account for these results. The model consists of two cross-coupled neuro-mechanical units, each composed of a neural oscillator driving a wrist-pendulum system. Taken individually, each unit reproduced the natural tendency of the participants to freely oscillate a pendulum close to its resonance frequency. When cross-coupled through the vision of the pendulum of the other unit, the two units entrain each other and meet at a common frequency influenced by the mechanical properties of the two pendulums involved. The ability of the proposed model to address the other effects observed as a function of the different conditions of the pendulum and intended mode of coordination is discussed.

Neuromuscular-skeletal constraints upon the dynamics of unimanual and bimanual coordination

In the first of three experiments, 11 participants generated pronation and supination movements of the forearm, in time with an auditory metronome. The metronome frequency was increased in eight steps (0.25 Hz) from a base frequency of 1.75 Hz. On alternating trials, participants were required to coordinate either maximum pronation or maximum supination with each beat of the metronome. In each block of trials, the axis of rotation was either coincident with the long axis of the forearm, above this axis, or below this axis. The stability of the pronate-on-the-beat pattern, as indexed by the number of pattern changes, and the time of onset of pattern change, was greatest when the axis of rotation of the movement was below the long axis of the forearm. In contrast, the stability of the supinate-on-the-beat pattern was greatest when the axis of rotation of the movement was above the long axis of the forearm. In a second experiment, we examined how changes in the position of the axis of rotation alter the activation patterns of muscles that contribute to pronation and supination of the forearm. Variations in the relative dominance of the pronation and supination phases of the movement cycle across conditions were accounted for primarily by changes in the activation profile of flexor carpi radialis (FCR) and extensor carpi radialis longus (ECR). In the Final experiment we examined how these constraints impact upon the stability of bimanual coordination. Thirty-two participants were assigned at random to one of four conditions, each of which combined an axis of rotation configuration (bottom or top) for each limb. The participants generated both inphase (both limbs pronating simultaneously, and supinating simultaneously) and antiphase (left limb pronating and right limb supinating simultaneously, and vice versa) patterns of coordination. When the position of the axis of rotation was equivalent for the left and the right limb, transitions from antiphase to inphase patterns of coordination were Frequently observed. In marked contrast, when the position of the axis of rotation for the left and right limb was contradistinct, transitions From inphase to antiphase patterns of coordination occurred. The results demonstrated that when movements are performed in an appropriate mechanical context, inphase patterns of coordination are less stable than antiphase patterns.

Effector dynamics of rhythmic wrist activity and its implications for (modeling) bimanual coordination

To examine the role of the effector dynamics of the wrist in the production of rhythmic motor activity, we estimated the phase shifts between the EMG and the task-related output for a rhythmic isometric torque production task and an oscillatory movement, and found a substantial difference (45-52degrees) between the two. For both tasks, the relation between EMG and task-related output (torque or displacement) was adequately reproduced with a physiologically motivated musculoskeletal model. The model simulations demonstrated the importance of the contribution of passive structures to the overall dynamics and provided an account for the observed phase shifts in the dynamic task. Additional simulations of the musculoskeletal model with added load suggested that particular changes in the phase relation between EMG and movement may follow largely from the intrinsic muscle dynamics, rather than being the result of adaptations in the neural control of joint stiffness. The implications of these results are discussed in relation to (models of) interlimb coordination in rhythmic tasks. (C) 2004 Elsevier B.V. All rights reserved.

Let your feet do the walking: constraints on the stability of bipedal coordination

In this paper we consider whether the behaviour of the neural circuitry that controls lower limb movements in humans is shaped primarily by the spatiotemporal characteristics of bipedal gait patterns, or by selective pressures that are sensitive to considerations of balance and energetics. During the course of normal locomotion, the full dynamics of the neural circuitry are masked by the inertial properties of the limbs. In the present study, participants executed bipedal movements in conditions in which their feet were either unloaded or subject to additional inertial loads. Two patterns of rhythmic coordination were examined. In the in-phase mode, participants were required to flex their ankles and extend their ankles in synchrony. In the out-of-phase mode, the participants flexed one ankle while extending the other and vice versa. The frequency of movement was increased systematically throughout each experimental trial. All participants were able to maintain both the in-phase and the out-of-phase mode of coordination, to the point at which they could no longer increase their frequency of movement. Transitions between the two modes were not observed, and the stability of the out-of-phase and in-phase modes of coordination was equivalent at all movement frequencies. These findings indicate that, in humans, the behaviour of the neural circuitry underlying coordinated movements of the lower limbs is not constrained strongly by the spatiotemporal symmetries of bipedal gait patterns.

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.

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.

Hierarchical organisation of neuro-anatomical constraints in interlimb coordination

Based on the observation that bimanual finger tapping movements tend toward mirror symmetry with respect to the body midline, despite the synchronous activation of non-homologous muscles, F. Mechsner, D. Kerzel, G. Knoblich, and W. Prinz (2001) [Perceptual basis of bimanual coordination. Nature, 414, 69-73] suggested that the basis of rhythmic coordination is purely spatial/perceptual in nature, and independent of the neuro-anatomical constraints of the motor system. To investigate this issue further, we employed a four finger tapping task similar to that used by F. Mechsner and G. Knoblich (2004) [Do muscle matter in bimanual coordination? Journal of Experimental Psychology: Human Perception and Performance, 30, 490-503] in which six male participants were required to alternately tap combinations of adjacent pairs of index (1), middle (M) and ring (R) fingers of each hand in time with an auditory metronome. The metronome pace increased continuously from 1 Hz to 3 Hz over the course of a 30-s trial. Each participant performed three blocks of trials in which finger combination for each hand (IM or MR) and mode of coordination (mirror or parallel) were presented in random order. Within each block, the right hand was placed in one of three orientations; prone, neutral and supine. The order of blocks was counterbalanced across the six participants. The left hand maintained a prone position throughout the experiment. On the basis of discrete relative phase analyses between synchronised taps, the time at which the initial mode of coordination was lost was determined for each trial. When the right hand was prone, transitions occurred only from parallel symmetry to mirror symmetry, regardless of finger combination. In contrast, when the right hand was supine, transitions occurred only from mirror symmetry to parallel but no transitions were observed in the opposite direction. In the right hand neutral condition, mirror and parallel symmetry are insufficient to describe the modes of coordination since the hands are oriented orthogonally. When defined anatomically, however, the results in each of the three right hand orientations are consistent. That is, synchronisation of finger tapping is deter-mined by a hierarchy of control of individual fingers based on their intrinsic neuro-mechanical properties rather than on the basis of their spatial orientation. (c) 2005 Elsevier B.V. All rights reserved.

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.

Scaling and non-scaling of muscle activity, kinematics, and dynamics in sit-ups with different degrees of difficulty

The purpose of this study was to investigate how the CNS adjusts motor patterns for variants of a complex axial movement-the situp. Adjustments were induced by changing the support surface contact and mass distribution of the body. Healthy adults performed straight-legged sit-ups, 3 s in duration, with support added to or removed from the lumbar trunk, or with mass added to the head or to the legs. Each of these interventions either increased or decreased the difficulty of the task. The study addressed the extent to which changes in sit-up difficulty are compensated by scaling of muscle activity, kinematics, and dynamics versus the extent to which they are compensated by changing discretely the motor pattern. The analysis of muscle activity, kinematics, and dynamics focused on the first 30-40% of the sit-up-the trunk flexion phase-since this is the most critical part of the movement. Our results demonstrate that, in some respects, sit-up kinematics and dynamics scaled with difficulty, but in other respects, they did not. Muscle activity also scaled, in many respects, but in more difficult sit-ups, abdominal flexor activity decreased instead of increased. Non-scaling changes in these parameters suggest that complex movements, such as the sit-up, may require discrete changes in motor pattern in order to deal with large loads, which challenge the available leverage. (C) 2005 Elsevier Ltd. All rights reserved.

The effects of viscous loading of the human forearm flexors on the stability of coordination

This experiment investigated whether the stability of rhythmic unimanual movements is primarily a function of perceptual/spatial orientation or neuro-mechanical in nature. Eight participants performed rhythmic flexion and extension movements of the left wrist for 30 s at a frequency of 2.25 Hz paced by an auditory metronome. Each participant performed 8 flex-on-the-beat trials and 8 extend-on-the-beat trials in one of two load conditions, loaded and unload. In the loaded condition, a servo-controlled torque motor was used to apply a small viscous load that resisted the flexion phase of the movement only. Both the amplitude and frequency of the movement generated in the loaded and unloaded conditions were statistically equivalent. However, in the loaded condition movements in which participants were required to flex-on-the-beat became less stable (more variable) while extend-on-the-beat movements remained unchanged compared with the unload condition. The small alteration in required muscle force was sufficient to result in reliable changes in movement stability even a situation where the movement kinematics were identical. These findings support the notion that muscular constraints, independent of spatial dependencies, can be sufficiently strong to reliably influence coordination in a simple unimanual task.