The influence of perceptual and motor factors on bimanual coordination in a polyrhythmic tapping task

Summary In this study the role of perceptual and motor factors on the motor organization (integrated versus parallel) adopted by musically skilled and unskilled subjects in a polyrhythmic tapping task was investigated. Subjects tapped a 3:2 polyrhythm to match the timing of two isochronous tone trains, one tone train for each hand. Perceptual factors were examined by the manipulation of the frequency difference between the tone trains to produce either an integrated or a streamed percept. Motor factors were examined by comparison of performance on two versions of the 3:2 polyrhythm. In one (simultaneous) version, each cycle of the polyrhythm began with a simultaneous left- and right-hand tap. In the other (shifted) version a 100-ms interval was introduced between the initial left and right taps in each cycle. Examination of the pattern of variances and covariances among intertap intervals suggested that most of the subjects in this study adopted an integrated motor organization that involved interleaving the timing of the two hands. Further analysis revealed that a serial chained model described the pattern of covariances best for the simultaneous pattern, whereas a hierarchical organization described the pattern of covariances for the shifted pattern best. The finding that performance was more accurate with integrated tones than with streamed tones provides some support for a perceptual-motor facilitation hypothesis.     

Effects of movement frequency and joint kinetics on the joint coordination underlying bimanual circle drawing

Ten healthy participants performed bimanual symmetric and asymmetric circle drawing at 4 frequencies. The authors partitioned the variance of the joint configuration across repetitions into 1 component representing equivalent joint configurations with respect to achieving stability of the mean hand path (i.e., goal-equivalent variance, GEV) and 1 component leading to a variable hand path (non-goal-equivalent variance, NGEV) across cycles. Higher frequencies led to increased NGEV related to control of the nondominant hand and to the relative position and orientation between the hands during asymmetric drawing. The results were related to differences in muscle and interaction moments between the arms, and they suggest a possible relationship between the ability to use intersegmental forces and the stability of interlimb synergy.     

Timing goals in bimanual coordination

Phase coupling between movement trajectories has been proposed as the basic mechanism of hand coordination in the production of bimanual rhythmic movements with a 1:2 frequency ratio. Here a central temporal coupling view is proposed as an alternative. Extending previous models of two-handed synchronic and alternate-hand tapping, we hypothesized that 1:2 tapping is performed under the control of a single internal timekeeper set at the frequency required for the fast hand. The fast hand is assumed to use every signal and the slow hand every other signal of the timekeeper, to produce actions coordinated in time. The model's predictions for the variance-covariance pattern of tap timing within and across hands were tested in an experiment that required tapping with both hands with 1:1 or 1:2 frequency ratio. The finger contact on the response plate was to be short or long, according to instruction. Prolonged finger contact entailed profound modifications in the movement trajectories but failed to modify the variance-covariance pattern of the tap timing. This pattern proved to conform to predictions under both the short and the long contact conditions, thus supporting the central temporal coupling hypothesis.     

Kinetic attraction during bimanual coordination

Two experiments examined the effects of independent variations in kinetic and kinematic requirements on interlimb coupling during a bimanual task. The goal of the investigation was to provide preliminary evidence regarding one general class of physical variables that constrains discrete bimanual movements. Subjects attempted to execute a smooth unidirectional movement with the left arm, along with a three-segment reversal movement with the right arm. The first experiment manipulated the torque required to produce the reversal action, while movement duration and average angular velocity were held constant for both limbs. Several indications of increased interlimb coupling, due to the kinetic variation, were evident. The converse manipulation was used in the second experiment, with movement time and kinematics (velocity, acceleration) changed independently of joint torque requirements for the reversal limb. No clear effect of kinematics on coupling strength was noted. The results suggest that one variable influencing interlimb attraction toward common spatiotemporal trajectories may be kinetic in nature.     

Anchoring in a novel bimanual coordination pattern

Anchoring in cyclical movements has been defined as regions of reduced spatial or temporal variability [Beek, P. J. (1989). Juggling dynamics. PhD thesis. Amsterdam: Free University Press] that are typically found at movement reversal points. For in-phase and anti-phase movements, synchronizing reversal points with a metronome pulse has resulted in decreased anchor point variability and increased pattern stability [Byblow, W. D., Carson, R. G., & Goodman, D. (1994). Expressions of asymmetries and anchoring in bimanual coordination. Human Movement Science, 13, 3-28; Fink, P. W., Foo, P., Jirsa, V. K., & Kelso, J. A. S. (2000). Local and global stabilization of coordination by sensory information. Experimental Brain Research, 134, 9-20]. The present experiment examined anchoring during acquisition, retention, and transfer of a 90 degrees phase-offset continuous bimanual coordination pattern (whereby the right limb lags the left limb by one quarter cycle), involving horizontal flexion about the elbow. Three metronome synchronization strategies were imposed: participants either synchronized maximal flexion of the right arm (i.e., single metronome), both flexion and extension of the right arm (i.e., double metronome within-limb), or flexion of each arm (i.e., double metronome between-limb) to an auditory metronome. In contrast to simpler in-phase and anti-phase movements, synchronization of additional reversal points to the metronome did not reduce reversal point variability or increase pattern stability. Furthermore, practicing under different metronome synchronization strategies did not appear to have a significant effect on the rate of acquisition of the pattern.     

Encoding and retrieval during bimanual rhythmic coordination

In 2 experiments, bimanual 1:1 rhythmic coordination was performed concurrently with encoding or retrieval of word lists. Effects of divided attention (DA) on coordination were indexed by changes in mean relative phase and recurrence measures of shared activity between the 2 limbs. Effects of DA on memory were indexed by deficits in recall relative to baseline. For DA at both encoding and retrieval, the equilibrium values of relative phase were shifted and the degree of shared activity between left and right rhythmic motions was reduced. Recall was reduced, however, only for DA at encoding. The results corroborate and extend those obtained with more conventional secondary tasks (e.g., visual reaction time), suggesting attention dissimilarities between episodic encoding and retrieval.     

Deterministic variability and stability in detuned bimanual rhythmic coordination

We examined the effects of crossing different degrees of cooperation and competition on inphase and antiphase 1:1 frequency locked coordination of left- and right-hand-oscillated pendulums. Degree of cooperation was manipulated through the joint frequency of oscillation specified by a metronome (the higher the frequency, the weaker the cooperation), and degree of competition by length (and, therefore, preferred frequency) differences between the two pendulums (the greater the difference, the stronger the competition). Increasing competition was accompanied by either decreasing cooperation (for six participants) or increasing cooperation (for six different participants). On each trial, a participant attempted to produce a steady-state phase relation phi for a given combination of competition and cooperation. Numerical simulations of the extended Haken-Kelso-Bunz (HKB) equation were used to predict (a) the patterns of shift in phi from either 0 or pi radians due to the different competition-cooperation relations and (b) the patterns of variability in phi. It was expected that the HKB equation would be successful in respect to (a), which it was, but not in respect to (b). The observed failure to confirm (b) was expected from the variability due to the different nonharmonic dynamics of the component oscillators, a source of variability not included in the HKB equation. The experimental results together with simulations and analyses of the phase-plane trajectories of the component oscillators suggest the operation of deterministic in addition to stochastic variability in the phase relation of contralateral limbs.     

Torso movement constraint in stability of bimanual coordination

This study investigated the relation between postural movement and upper-limb coordination stability. Adults produced bimanual circles using in-phase and anti-phase coordination patterns in time to an increasing rate metronome (i.e., movement-time instruction) in the horizontal (e.g., tabletop) and vertical (e.g., "wall" perpendicular to body) planes. All participants produced the instructed in- and anti-phase patterns. Coordination stability (i.e., SD of relative phase) was larger for anti-phase than in-phase patterns in both planes; however, anti-phase coordination stability was lower in the vertical plane than in the horizontal plane. Torso movement was larger during anti-phase coordination patterns in the horizontal plane, whereas it was larger during in-phase coordination patterns in the vertical plane. These results indicate that different orientations of the same task can produce different results for stability of coordination. This information may be important for performing and learning complex motor-coordination movements (e.g., playing musical instruments).     

Goal congruency without stimulus congruency in bimanual coordination

Two experiments explored the role of stimulus congruency and goal congruency for the generation of object-oriented actions with two hands. Participants had to place two objects into either parallel or opposite orientations by carrying out either symmetrical or asymmetrical forearm rotations. Performance was superior when the required object orientations were identical rather than different, almost independent of the symmetry of the required arm movements. In extending previous research, goal congruency effects ensued even under conditions in which congruency of imperative response signals could not have contributed to goal congruency effects, either because only a single stimulus was used to indicate the required goals in an individual trial (Experiment 1) or such stimuli were absent at all (Experiment 2). The results thus confirm the importance of goal codes for the reconcilableness of bimanual actions, and rule out accounts in terms of stimulus-related processes.     

The interplay of attention and bimanual coordination dynamics

Despite their common origin, studies on motor coordination and on attentional load have developed into separate fields of investigation, bringing out findings, methods, and theories which are diverse if not mutually exclusive. Sitting at the intersection of these two fields, this article addresses the issue of behavioral flexibility by investigating how intention modifies the stability of existing patterns of coordination between moving limbs. It addresses the issue, largely ignored until now, of the attentional cost incurred by the central nervous system (CNS) in maintaining a coordination pattern at a given level of stability, in particular under different attentional priority requirements. The experimental paradigm adopted in these studies provides an original mix of a classical measure of attentional load, namely, reaction time, and of a dynamic approach to coordination, most suitable for characterizing the dynamic properties of coordinated behavior and behavioral change. Findings showed that central cost and pattern stability covary, suggesting that bimanual coordination and the attentional activity of the CNS involved in maintaining such a coordination bear on the same underlying dynamics. Such a conclusion provides a strong support to a unified approach to coordination encompassing a conceptualization in terms of information processing and another, more recent framework rooted in self-organization theories and dynamical systems models     

Stabilisation of bimanual coordination through visual coupling

Eight right-handed participants performed a bilateral circle tracing task in symmetric or asymmetric patterns. Circle tracing was performed in synchrony with an auditory metronome and a visual display at, or comfortably below, each participant's transition frequency. The visual display consisted of a row of five light-emitting diodes (LEDs) arranged between the two circles (hands). Bimanual pattern stability was examined under conditions where the direction of illumination of the visual stimuli was compatible or incompatible with the hand direction. Symmetric patterns maintained stability for both movement rates whereas asymmetric patterns exhibited loss of stability at the transition frequency. Spontaneous reversals in circling direction occurred predominantly (94%) through the nondominant hand. Laterality effects were also evident in the aspect ratio (circularity of trajectories) and limb frequency variation, particularly in asymmetric patterns at the transition frequency. Compatibility between the stimulus direction and circling direction served to: stabilise symmetric patterns; stabilise asymmetric patterns by delaying the onset of transition; and stabilise the individual limb dynamics when the direction of the dominant side was compatible with the visual stimulus. The data from this multisegmental task lend support to a model of coupled oscillators whereby the coupling strength is anisotropic between the dominant and nondominant side, and lend further support for an account of manual asymmetries by way of a preferential perception–action coupling through the dominant limb. PsycINFO Classification: 2320     

Inter-limb coupling in bimanual rhythmic coordination in Parkinson's disease

Recently, it has been shown that rhythmic inter-limb coordination is disturbed in patients with Parkinson's disease (PD). The present study aims to investigate whether this coordination deficit is primarily the result of an impaired coupling, related to hypoactivation of the supplementary motor area (SMA), or primarily the indirect result of an asymmetrical distribution of PD symptoms over the left and right limbs (a peripheral process). Thirty PD patients and 30 matched control participants tapped with the index fingers anti-phase and left and right leading gallop patterns in four visual feedback conditions. Symmetrically affected participants performed significantly worse than asymmetrically affected and control participants in the gallop patterns. This result suggested that the central deficit has a stronger effect on inter-limb coupling in PD than the neuromuscular and biomechanical asymmetry between the limbs. Detailed analysis of inter-tap intervals (variability and correlation) suggested that this deficit leads to a compensatory asymmetrical inter-limb coupling in the primarily right-affected patient group, and under specific circumstances also in the primarily left-affected patient group. The difference in coordination strategy between left- and right-affected patients suggested that pre-morbid hand preference is an important structural constraint on the coupling strategies available to the participants.     

Visual and tactile action effects determine bimanual coordination performance

Effect-based models of motor control assign a crucial role to anticipated perceptual feedback in action planning. Two experiments were conducted to test the validity of this proposal for discrete bimanual key press responses. The results revealed that the normally observed performance advantage for the preparation of two responses with homologous rather than non-homologous fingers becomes inverted when homologous fingers produce non-identical visual effects, and non-homologous fingers produce identical visual effects. In the second experiment the finger homology effect was strongly reduced when homologous fingers produced non-identical tactile feedback. The results show that representations of to-be-produced visual and tactile action effects both contribute to action planning, though possibly to a varying degree. Implications of these results for effect-based models of motor control are considered.     

Anchoring strategies for learning a bimanual coordination pattern

Anchoring has been defined as synchronizing a point in a movement cycle with an external stimulus (W. D. Byblow, R. G. Carson, & D. Goodman, 1994). Previously, investigators have examined anchoring during in-phase and antiphase movements. The present authors examined anchoring during acquisition of a novel bimanual coordination pattern. Participants performed a 90 degrees pattern at 1 Hz, with a 2- or 4-Hz metronome. No group differences were found in pattern performance; however, the 4-Hz group developed more consistent anchoring relative to the metronome. Mechanical anchor-point variability differed by hand, position (midpoint vs. endpoint), and direction (flexion vs. extension) but not by metronome frequency. Those results support and extend previous findings but leave unanswered questions regarding the benefits and effectiveness of anchoring during a 90 degrees pattern.     

Perception and action influences on discrete and reciprocal bimanual coordination

For nearly four decades bimanual coordination, “a prototype of complex motor skills” and apparent “window into the design of the brain,” has been intensively studied. Past research has focused on describing and modeling the constraints that allow the production of some coordination patterns while limiting effective performance of other bimanual coordination patterns. More recently researchers have identified a coalition of perception-action constraints that hinder the effective production of bimanual skills. The result has been that given specially designed contexts where one or more of these constraints are minimized, bimanual skills once thought difficult, if not impossible, to effectively produce without very extensive practice can be executed effectively with little or no practice. The challenge is to understand how these contextual constraints interact to allow or inhibit the production of complex bimanual coordination skills. In addition, the factors affecting the stability of bimanual coordination tasks needs to be re-conceptualized in terms of perception-related constraints arising from the environmental context in which performance is conducted and action constraints resident in the neuromotor system.     

The control of amplitude and direction in a bimanual coordination task

Bimanual coordination requires task-specific control of the spatial and temporal characteristics of the movements of both hands. The present study focused on the spatial relationship between hand movements when their amplitude and direction were manipulated. In the experiment in question, participants were instructed to draw two lines simultaneously. These two lines were instructed to be drawn in mirror symmetric or perpendicular directions of each other while the length was instructed to be the same or different. The coordinative quality of amplitude control was compared when the task required symmetric and asymmetric bimanual spatial coordination patterns. Results showed that the amplitude accuracy decreased when different amplitudes and/or directions had to be generated simultaneously. The coordinative quality of direction was also compared when the task required symmetric and asymmetric bimanual spatial coordination patterns. Unlike amplitude, the direction accuracy was largely independent of coordination symmetry/asymmetry of direction or amplitude. The results suggest that the coordinative quality of amplitude control does not only interfere with amplitude asymmetry, but it also interferes with direction asymmetry. Moreover, in bimanual coordination amplitude control is more vulnerable to the influence of direction control demands than vice versa.     

Interference effects in bimanual coordination are independent of movement type

Simultaneously executed limb movements interfere with each other. Whereas the interference between discrete movements is examined mostly from a cognitive perspective, that between rhythmic movements is studied mainly from a dynamical systems perspective. As the tools and concepts developed by both communities are limited in their applicability to the other domain, it remains unclear if a common cause underlies motor interference in both domains. We investigated the interference between simultaneously executed discrete and rhythmic wrist movements. The discrete movements' reaction time and movement time decreased with increasing rhythmic movement frequency. The discrete movements accelerated or decelerated the rhythmic movements in a manner that depended on movement frequency and the discrete movement's initiation phase. The acceleration/deceleration profile was bimodal at low frequencies and unimodal at high frequencies, mimicking the hallmark feature of rhythmic-rhythmic coordination, thus suggesting that interference between movements may be invariant across different movement types.     

Attending to the non-preferred hand improves bimanual coordination in children

The effect of attentional focus in bimanual coordination was investigated from a developmental perspective by examining performance of right- and left-handed children, 5-8-years and 9-12-years old, on bimanual reciprocal tapping tasks. Attentional focus was either specified, by asking the children to attend to the preferred or to the non-preferred hand, or unspecified for the execution of the tasks. When attention was oriented to the non-preferred hand we found a reduced movement time and a lower frequency of errors. Performance differences for handedness and age-groups were observed when the children were oriented to attend to the preferred hand or when there was no instruction regarding attention. These differences in performance were eliminated when attention was oriented to the non-preferred hand.     

Effects of task instructions and oscillation frequency on bimanual coordination

Increases in the oscillation frequency of bimanual movements produce a switch from an anti-phase (180° relative phase) to an in-phase (0° relative phase) coordination pattern. This finding is observed when subjects are instructed not to intervene when they feel themselves slipping out of the anti-phase pattern. The question addressed in this study concerned how performance would be affected if subjects were instructed to try to maintain the pattern at all times. This issue was addressed using two separate groups of subjects: one group was given the do not intervene instructions, the other group was told to try to stay with the pattern at all times. Forearm rotations were tested in 15 s trials, paced by an auditory metronome set at 1.0, 1.5, 2.0, 2.5, and 3.0 Hz. Frequency distributions of the point estimates of relative phase were analyzed. The Do not Intervene group replicated previous findings, as indicated by the development of a bimodal histogram of relative phase distributions with increases in oscillation frequency. However, a very different pattern of findings emerged with increases in oscillation frequency for the group told to stay with the anti-phase pattern. Rather than a bimodal distribution being developed, the data maintained 180° as its central tendency — no secondary distribution developed around 0° relative phase. These data suggest that volitional control can over-ride the inherent dynamical tendencies of the motor system.     

Practice and training in bimanual coordination tasks: strategies and constraints.

There are a number of structural constraints on the coordination of the two hands. Understanding bimanual coordination involves not only identification of these constraints but also how they can be overcome or modulated. The performance of polyrhythms has been used to examine these issues. In this article the constraints on coordination are outlined and research on the acquisition of polyrhythms is reviewed. It is suggested that the constraints on the production of multifrequency ratios are overcome by integrating the timing of the two hands. Oscillator models of coordination are then considered and a two-process model of motor timing is suggested.