The acquisition of bimanual coordination is mediated by anisotropic coupling between the hands

The present study was designed to test two predictions from the coupled oscillator model of multifrequency coordination. First, it was predicted that multifrequency tasks that match the inherent manual asymmetry (i.e., the preferred hand assigned to the faster tempo) would be easier to learn than tasks that do not match the inherent dynamics (i.e., the non-preferred hand assigned to the faster tempo). Second, in the latter case acquisition of the multifrequency coordination would involve a reorganisation of the coupling dynamics such that the faster hand would exert a greater influence on the slower hand than vice versa. Sixteen right-handed volunteers received extensive training on a 2:1 coordination pattern involving a bimanual forearm pronation-supination task. Participants were randomly assigned to one of two groups: 1L:2R in which the preferred right hand performed the higher frequency, or 2L:1R in which the non-preferred left hand performed the higher frequency. The dynamic stability of the patterns was assessed by the ability of participants to maintain the coordination pattern as movement frequency was increased. Changes in the directional coupling between the hands was assessed by transition pathways and lead-lag relationship evident in a 1:1 anti-phase frequency-scaled coordination task performed prior to and following three practice sessions of the 2:1 task. The predicted differential stability between the multifrequency patterns was evident in the initial acquisition sessions but by the end of training the two patterns evidenced equivalent stability. Unexpectedly, for both groups the fast hand displayed greater variability in amplitude and movement frequency than the slow hand perhaps reflecting anchoring afforded to the slow hand by synchronising movement endpoints with the auditory pacing metronome. Analysis of pre- to post-training changes to the coupling dynamics in the 1:1 anti-phase task support the hypothesis that acquisition of the 2L:1R pattern involved reorganisation of the inherent dynamics.     

Laterally focused attention modulates asymmetric coupling in rhythmic interlimb coordination

Peters (J Motor Behav 21:151-155, 1989; Interlimb coordination: neural, dynamical and cognitive constraints, Academic, Orlando, pp 595-615, 1994) suggested that expressions of handedness in bimanual coordination may be reflections of an inherent attentional bias. Indeed, previous results indicated that focusing attention on one of the limbs affected the relative phasing between the limbs in a manner comparable to the effects of hand dominance. The present study extended the comparison between the effects of attentional focus and handedness by testing their impact on the interactions between the limbs. Both left-handed and right-handed participants performed rhythmic bimanual coordination tasks (in-phase and antiphase coordination), while directing attention to either limb. Using brief mechanical perturbations, the degree to which the limbs were influenced by each other was determined. The results revealed that the non-dominant limb was more strongly affected by the dominant limb than vice versa and that, in line with Peters' proposition, this handedness-related asymmetry in coupling strength was reduced when attention was focused on the non-dominant limb, thereby highlighting the potential relation between inherent (handedness-related) asymmetries and voluntary attentional asymmetries. In contrast to previous findings, the (commonly observed) phase lead of the dominant limb was attenuated (rather than accrued) when attention was focused on this limb. This unexpected result was explained in terms of the observed attention-related difference in amplitude between the limbs.     

The Dynamics of Bimanual Circle Drawing

A bimanual circle drawing task was employed to elucidate the dynamics of intralimb and interlimb coordination. Right-handed subjects were required to produce circles with both hands in either a symmetrical (mirror) mode (i.e. one hand moving clockwise, the other counter-clockwise) or in an asymmetrical mode (i.e. both hands moving clockwise or counter-clockwise). The frequency of movement was scaled by an auditory metronome from 1.50 Hz to 3.25 Hz in8 (8-sec) steps.In the asymmetrical mode,distortions ofthe movement trajectories, transient departures from the target pattern of coordination, and phase wandering were evident as movement frequency was increased. These features suggested loss of stability. Deviations from circular trajectories were most prominent for movements of the left hand. Transient departures from the required mode of coordination were also largely precipitated by the left hand. The results are discussed with reference to manual asymmetries and mechanisms of interlimb and intersegmental coordination.     

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     

Dynamics of 1:2 Coordination: Generalizing Relative Phase to n:m Rhythms

Interlimb rhythmic movements can be modeled as coupled oscillators, with stable performance characterized by the relative phase between the limbs. In the present study, that modeling strategy, verified previously for 1:1 coordination, was generalized to 1:2 coordination with a view to n:m coordination. The generalized model predicted interactions between coordination (specifically, 1:1 vs. 1:2) and the frequency asymmetry between the limbs determining mean relative phase and its variability. The predicted interactions were evaluated with bimanual 1:2 and 1:1 rhythmic tasks in which participants (N = 8) oscillated hand-held pendulums whose uncoupled frequencies could be adjusted so that different interlimb asymmetries were produced. The authors needed new analytic procedures to verify stable 1:2 coordination and to resolve stochastic and deterministic sources of variability in the component oscillations. The major expectations from the generalized model were confirmed, and the implications of additional but unpredicted findings for the modeling of multifrequency behavior are discussed.     

Asymmetries in Coupling Dynamics of Perception and Action

The influence of information-based dynamics on coordination dynamics of rhythmic movement was examined with special reference to the expression of asymmetries. In Experiment 1, right-handed subjects performed unimanual, rhythmical movements in coordination with either a discrete or continuous visual display. The right hand-visual display system defined a more stable perception-action collective than the left, particularly when continuous visual information was available. In Experiment 2, the same subjects performed rhythmic bimanual movements in coordination with a continuous visual display. The action collective was inherently more stable than the perception-action collective, although similar patterns were observed at both levels. Importantly, the dynamics of the perception-action collective impinged upon the dynamics of the action collective in terms of stability. Asymmetries remained evident between limbs in the bimanual preparations, with the left hand exhibiting greater limit-cycle variability and also a tendency to more often effect transitions at the action couple. Features of dynamical models that capture characteristics of manual asymmetries are discussed.     

Hemispheric asymmetries in attentional control: implications for hand preference in sensorimotor tasks

Liepmann (1908) proposed that handedness reflects the greater capacity of one hemisphere to learn the execution of skilled movements. Although asymmetries in motor control are an important basis for hand asymmetries, recent studies have suggested that handedness may be determined by multiple factors. In the present study, we examine how attentional asymmetries may contribute to hand preferences. Right-handed subjects participated in a reaction time task in which they were given preliminary information about where a target stimulus would occur (selective attention) or which hand to use for responding (selective intention). Our findings indicate that these processes influence each other reciprocally and favor a state of optimal attentional and intentional preparation of the right hand. We suggest that these hemispheric asymmetries in attentional control contribute to hand preferences in certain sensorimotor tasks.     

EEG correlates of behavioural laterality: right-handedness

Among right-handers, the magnitude of differences in proficiency between the left and right hands varies considerably. Yet significance of the extent of right-handedness is still a controversial issue. To examine whether individual differences in asymmetry of hand skill can partly be attributed to individual differences in asymmetrical hemispheric activation, handedness and electroencephalographic (EEG) laterality were correlated in two large samples (ns = 60 and 128). Analysis indicated that part of the variability in right-handedness may arise from activation asymmetries in the cortex, but whether this relation becomes apparent depends on the cortical area examined and on the experimental condition under which the EEG measures are taken.     

Attention as a mediating variable in the dynamics of bimanual coordination

The influence of focal attention on the coordination dynamics in a bimanual circle drawing task was investigated. Six right-handed and seven left-handed subjects performed bimanual circling movements, in two modes of coordination, symmetrical or asymmetrical. The frequency of movement was scaled by an auditory metronome from 1.50 Hz to 3.00 Hz in 7 steps. On each trial, subjects were required to attend either to the dominant hand, to a neutral position, or to the nondominant hand.The uniformity of the relative tangential angle was lower in asymmetrical than in symmetrical conditions, but was not influenced by the direction of attention. In the asymmetrical mode, shifts in RTA relations, suggestive of loss of stability, were evident as the movement frequency was increased. Typically, these shifts were mediated by distortions of the trajectory of the nondominant limb. When the nondominant hand was the focus of attention, movements of this hand were more circular, and temporal variability was reduced, at the cost of a greater deviation from the target frequency. Movements of the dominant hand were not affected by the direction of attention. The findings show that although directed attention acts to modify the coordination dynamics, it does so primarily at the level of the individual hands, rather then in terms of the relation between them.     

A dynamical systems approach to manual tracking performance

Manual tracking performance was investigated from the perspective of dynamical systems theory. The authors manipulated the type of visual display, the control system dynamics, and the frequency of the sinusoidal input signal to examine couplings with various phases between the visual signal and control movements. Analyses of the system output amplitude ratio and relative phase showed that participants (N = 24) performed poorly with 90 degrees relative phase coupling. All the couplings became less stable as the movement frequency increased. The authors developed an adaptive oscillator model with linear damping to describe the coupled system consisting of the human performer, the visual display, and the control system dynamics. A geometric account of the stability of performance at different relative phases is also presented.     

Maternal cradling bias in baboons: The first environmental factor affecting early infant handedness development?

The most emblematic behavioral manifestation of human brain asymmetries is handedness. While the precise mechanisms behind the development of handedness are still widely debated, empirical evidences highlight that besides genetic factors, environmental factors may play a crucial role. As one of these factors, maternal cradling behavior may play a key role in the emergence of early handedness in the offspring. In the present study we followed 41 Papio anubis infants living in social groups with their mother for which direction (e.g., left- or right-arm) and degree of maternal cradling-side bias were available from a previous published study. We assessed hand preferences for an unimanual grasping task at three developmental stages: (A) 0–4, (B) 4—6, and (C) 9–10 months of age. We found that individual hand preferences for grasping exist as soon as the first months of age, with a population-level left-handedness predominance, being stable until 6 months; to wit the period during which juveniles are mainly carried by their mothers. More importantly, this early postnatal handedness is positively correlated with maternal cradling lateralization. Interestingly, hand preferences assessed later in the development, once juveniles are no longer carried (i.e., from 9 to 10 months of age), are less dependent from the maternal cradling bias and less consistent with the earlier developmental stages, especially in infants initially cradled on the right maternal side. Our findings suggest that the ontogenetic dynamics of the infant's hand preference and its changes might ultimately rely on the degree of infant dependence from the mother across development.     

The distribution and development of handedness for manual gestures in captive chimpanzees (Pan troglodytes)

This article describes the distribution and development of handedness for manual gestures in captive chimpanzees. Data on handedness for unimanual gestures were collected in a sample of 227 captive chimpanzees. Handedness for these gestures was compared with handedness for three other measures of hand use: tool use, reaching, and coordinated bimanual actions. Chimpanzees were significantly more right-handed for gestures than for all other measures of hand use. Hand use for simple reaching at 3 to 4 years of age predicted hand use for gestures 10 years later. Use of the right hand for gestures was significantly higher when gestures were accompanied by a vocalization than when they were not. The collective results suggest that left-hemisphere specialization for language may have evolved initially from asymmetries in manual gestures in the common ancestor of chimpanzees and humans, rather than from hand use associated with other, non-communicative motor actions, including tool use and coordinated bimanual actions, as has been previously suggested in the literature.     

Coordination Between Arm and Leg Movements During Locomotion

To evaluate the contrasting dynamical and biomechanical interpretations of the 2:1 frequency coordination between arm and leg movements that occurs at low walking velocities and the 1:1 frequency coordination that occurs at higher walking velocities, the authors conducted an experiment in which they quantified the effect of walking velocity on the stability of the frequency and phase coordination between the individual limb movements. Spectral analyses revealed the presence of 2:1 frequency coordination as a consistent feature of the data in only 3 out of 8 participants at walking velocities ranging from 1.0 to 2.0 km/h, in spite of the fact that the eigenfrequencies of the arms were rather similar across participants. The degree of interlimb coupling, as indexed by weighted coherence and variability of relative phase, was lower for the arm movements and for ipsilateral and diagonal combinations of arm and leg movements than for the leg movements. Furthermore, the coupling between all pairs of limb movements was found to increase with walking velocity, whereas no clear signs were observed that the switches from 2:1 to 1:1 frequency coordination and vice versa were preceded by loss of stability. Therefore, neither a purely biomechanical nor a purely dynamical model is optimally suited to explain these results. Instead, an integrative model involving elements of both approaches seems to be required.     

Coordination between arm and leg movements during locomotion

To evaluate the contrasting dynamical and biomechanical interpretations of the 2:1 frequency coordination between arm and leg movements that occurs at low walking velocities and the 1:1 frequency coordination that occurs at higher walking velocities, the authors conducted an experiment in which they quantified the effect of walking velocity on the stability of the frequency and phase coordination between the individual limb movements. Spectral analyses revealed the presence of 2:1 frequency coordination as a constant feature of the data in only 3 out of 8 participants at walking velocities ranging from 1.0 to 2.0 km/h, in spite of the fact that the eigenfrequencies of the arms were rather similar across participants. The degree of interlimb coupling, as indexed by weighted coherence and variability of relative phase, was lower for the arm movements and for ipsilateral and diagonal combinations of arm and leg movements than for the leg movements. Furthermore, the coupling between all pairs of limb movements was found to increase with walking velocity, whereas no clear signs were observed that the switches from 2:1 to 1:1 frequency coordination and vice versa were preceded by loss of stability. Therefore, neither a purely biomechanical nor a purely dynamical model is optimally suited to explain these results. Instead, an integrative model involving elements of both approaches seems to be required.     

Dynamics of 1:2 Coordination: Temporal Scaling, Latent 1:1, and Bistability

The simplest interlimb multifrequency coordination of 1:2 can be performed at different speeds and in at least two different styles or modes. The effects of speed and mode (in-phase or antiphase) were evaluated in a bimanual 1:2 rhythmic task in which participants (N = 8) oscillated hand-held pendulums with identical or different uncoupled frequencies. A motion equation in relative phase that captures the asymmetries of components and task predicted the 1:2 coordination equilibria resulting from temporal scaling. According to the experimental results, both coordination modes proved to be equally stable. More detailed analyses of individual trials showed signs that the more fundamental 1:1 coordination intruded into the 1:2 coordination.     

Dynamics of 1:2 Coordination: Sources of Symmetry Breaking

Three asymmetries in the dynamics of 1:2 interlimb coordination were examined: the asymmetry in uncoupled frequencies, the asymmetry in coupled frequencies, and the left-right functional asymmetry of the body. In a bimanual 1:2 task, participants (N = 8) oscillated hand-held pendulums whose uncoupled frequencies were adjusted so that the first kind of asymmetry could be manipulated. For any given pendulum pair, the pendulum assuming the faster motion in the 1:2 coordination was oscillated in the right and the left hands. By assigning combinations of uncoupled eigenfrequencies and coupled task-specified frequencies across hands, the authors studied the interaction of all 3 asymmetries. The results confirm the appropriateness of generalized relative phase as a collective variable for 1:2 coordination. Additionally, they suggest that the generalized form of the detuning parameter represents the first asymmetry and that the coupling function expresses the second asymmetry. In 1:2 coordination, the body's functional asymmetry plays a limited role.     

A Dynamical Systems Approach to Manual Tracking Performance

Manual tracking performance was investigated from the perspective of dynamical systems theory. The authors manipulated the type of visual display, the control system dynamics, and the frequency of the sinusoidal input signal to examine couplings with various phases between the visual signal and control movements. Analyses of the system output amplitude ratio and relative phase showed that participants (N = 24) performed poorly with 90° relative phase coupling. All the couplings became less stable as the movement frequency increased. The authors developed an adaptive oscillator model with linear damping to describe the coupled system consisting of the human performer, the visual display, and the control system dynamics. A geometric account of the stability of performance at different relative phases is also presented.     

Dynamics of 1:2 Coordination: Temporal Scaling,Latent 1:1, and Bistability

The simplest interlimb multifrequency coordination of 1:2 can be performed at different speeds and in at least two different styles or modes. The effects of speed and mode (in-phase or antiphase) were evaluated in a bimanual 1:2 rhythmic task in which participants (N = 8) oscillated hand-held pendulums with identical or different uncoupled frequencies. A motion equation in relative phase that captures the asymmetries of components and task predicted the 1:2 coordination equilibria resulting from temporal scaling. According to the experimental results, both coordination modes proved to be equally stable. More detailed analyses of individual trials showed signs that the more fundamental 1:1 coordination intruded into the 1:2 coordination.     

Performance Asymmetries in Computer Mouse Control of Right-Handers, and Left-Handers with Left- and Right-Handed Mouse Experience

Precision and general computer mouse aiming performance by right-handers (RH-RM) and left-handers with right-handed mouse experience (LH-RM) and by left-handers with left-handed mouse experience (LH-LM) were compared. A number of performance measures, such as reaction time, time to reach a target, time to click on target, and cursor trajectory, were analyzed. Superficially, specific hand experience seemed to dictate performance asymmetries, but a closer look revealed interactions between hand preference and hand performance. That finding has implications for theories of handedness. In addition, precision and general directional aiming with the mouse cursor showed a clear right hand superiority in reaction time in both RH-RM and LH-RM subjects, whereas LH-LM subjects showed no lateral asymmetries. Finally, the overall time taken for the task, averaged across groups and conditions, favored the experienced hand by some 180 ms. In practical terms, that is not a large difference, especially because the difference will be reduced with practice. Thus, the use of the inexperienced hand can be advocated when there is a need to forestall or ameliorate repetitive stress in the experienced hand.     

Eye-Hand Coordination in Rhythmical Pointing

The authors investigated the relation between hand kinematics and eye movements in 2 variants of a rhythmical Fitts's task in which eye movements were necessary or not necessary. P. M. Fitts's (1954) law held in both conditions with similar slope and marginal differences in hand-kinematic patterns and movement continuity. Movement continuity and eye—hand synchronization were more directly related to movement time than to task index of difficulty. When movement time was decreased to fewer than 350 ms, eye—hand synchronization switched from continuous monitoring to intermittent control. The 1:1 frequency ratio with stable π/6 relative phase changed for 1:3 and 1:5 frequency ratios with less stable phase relations. The authors conclude that eye and hand movements in a rhythmical Fitts's task are dynamically synchronized to produce the best behavioral performance.