Effects of movement stability and congruency on the emergence of spontaneous interpersonal coordination

The current project evaluated the relationship between the stability of intrapersonal coordination and the emergence of spontaneous interpersonal coordination. Participants were organized into pairs, and each participant was instructed to produce either an inphase or antiphase pattern of intrapersonal bimanual coordination using two hand-held pendulums, while simultaneously performing an interpersonal puzzle task. At issue was whether the emergence and stability of spontaneous interpersonal rhythmic coordination is influenced by ("Experiment 1") the stability of the intrapersonal coordination patterns produced by co-actors and ("Experiment 2") the congruency of the intrapersonal coordination patterns produced by co-actors. The stability of intrapersonal movement coordination did not affect the emergence of spontaneous interpersonal coordination. The degree of interpersonal coordination observed was similar when both participants in a pair produced either inphase or antiphase patterns of intrapersonal bimanual coordination. Moreover, the congruency of the intrapersonal coordination patterns only slightly affected the emergence of interpersonal coordination, with only marginally lower inphase interpersonal entrainment when participants produced incongruent patterns of intrapersonal coordination (e.g., inphase-antiphase). Interestingly, movement observation and the emergence of interpersonal coordination did not affect the stability of intrapersonal bimanual coordination. The results suggest that interlimb rhythmic bimanual coordination reflects a single intrapersonal perceptual-motor synergy and that these bimanual synergies (not individual limbs) are what become spontaneously entrained interpersonally.     

Binding of movement, sound and touch: multimodal coordination dynamics

Very little is known about the coordination of movement in combination with stimuli such as sound and touch. The present research investigates the hypothesis that both the type of action (e.g., a flexion or extension movement) and the sensory modality (e.g., auditory or tactile) determine the stability of multimodal coordination. We performed a parametric study in which the ability to synchronize movement, touch and sound was explored over a broad range of stimulus frequencies or rates. As expected, synchronization of finger movement with external auditory and tactile stimuli was successfully established and maintained across all frequencies. In the key experimental conditions, participants were instructed to synchronize peak flexion of the index finger with touch and peak extension with sound (and vice-versa). In this situation, tactile and auditory stimuli were delivered counter-phase to each other. Two key effects were observed. First, switching between multimodal coordination patterns occurred, with transitions selecting one multimodal pattern (flexion with sound and extension with touch) more often than its partner. This finding indicates that the stability of multimodal coordination is influenced by both the type of action and the stimulus modality. Second, at higher rates, transitions from coherent to incoherent phase relations between touch, movement and sound occurred, attesting to the breakdown of multimodal coordination. Because timing errors in multimodal coordination were systematically altered when compared to unimodal control conditions we are led to consider the role played by time delays in multimodal coordination dynamics.     

Phase transitions and postural deviations during bimanual kinesthetic tracking

Upper limb coordination was studied by examining pattern stability of between-hand rhythmical coordination. In the first of two experiments, relative phase of rhythmical wrist flexion-extension was examined within a kinesthetic tracking paradigm. Eight right-handed subjects actively tracked a driven hand being flexed and extended by a computer-controlled AC servo-motor. Hand movements were constrained in flexion or extension. The simultaneous contraction of wrist flexors and extensors was defined as inphase (IP) and the alternating contraction of wrist flexors and extensors as antiphase (AP). Phase transitions (from AP to IP) were observed in 16% of trials prepared in AP. Fewer phase transitions occurred when the right wrist was constrained in flexion, and also when the left wrist was constrained in extension. IP patterns were performed with greater stability than AP patterns. These effects were explored further in a second experiment with the addition of a secondary probe reaction time task to assess demands on central capacity, and the analysis of wrist flexor and extensor electromyographic activity. Subjects returned longer reaction times for AP than IP movement, suggesting the AP movement pattern placed a greater demand on central capacity than the IP movement pattern. During this kinesthetic tracking task, similar dynamic principles emerged as those observed during bilaterally active bimanual rhythmical coordination. The greater stability of the hand-posture combination where the driven left hand was constrained in extension and the active right hand was constrained in flexion may be a demonstration of unique central control of coupled activity. Electronic Publication     

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.     

Cortical and behavioral adaptations in response to short-term inphase versus antiphase bimanual movement training

Bimanual movement training (BMT) may be an effective rehabilitative protocol for movement-related deficits following a stroke; however, it is unclear how varying types of BMT induce cortical adaptations in the healthy population. Moreover, we lack a methodology to measure cortical adaptations in response to modes of movement training. Therefore, the present study measured the cued movement-related potential (MRP) to investigate cortical adaptations during cued inphase versus antiphase BMT that transferred to a unimanual task and how cortical modulations related to behavior. Three specific hypotheses were investigated: (1) cued inphase BMT would induce cortical adaptations within regions subserving motor preparation and movement execution, (2) repetitive cued unimanual training would induce cortical activity modulations associated with motor execution, and (3) increased cortical activity would be associated with enhanced performance. On three separate days, EEG was recorded from 22 electrodes during three types of cued movement training: inphase BMT, antiphase BMT and repetitive unimanual movement, in addition to pre- and post-training unimanual movement trials involving cued right wrist flexion. The MRP was measured for each repetition during each trial. Results showed a significant training-related increase in preparatory activation correlated with a behavioral enhancement following cued inphase BMT. This effect was not attributable to a change in arousal. No significant training-related modulation occurred in response to cued antiphase BMT or repetitive unimanual movement training. These results suggest that cortical adaptations in relation to the preparation of a cued movement enhance in response to cued inphase BMT, and the MRP is an effective measurement tool to assess training-related adaptations in response to inphase BMT specifically.     

Time dependence of coupling in frequency-scaled bimanual coordination

Prior research has shown that fluctuations in the relative phase of bimanual coordination do not reflect a white Gaussian noise process. The present study furthered the examination of time-dependent properties in bimanual coordination by comparing the magnitude of relative phase variability and the degree of effector independence within the time domain. The original Kelso (1984) [10] bimanual frequency-scaling protocol was reproduced in which phase transitions from antiphase to in-phase were induced with increasing movement frequency. The results showed that as movement frequency was scaled-up the amount of relative phase variability increased and the effector movements became more dependent prior to the transition. This is consistent with previous modeling showing that stronger effector coupling can prevent the occurrence of phase transitions when long range correlations in relative phase are present. It appears that, as movement frequency is scaled up, increases in effector coupling strength minimize loss of pattern stability and delay the onset of phase transitions.     

Keeping with the beat: movement trajectories contribute to movement timing

Previous studies of paced repetitive movements with respect to an external beat have either emphasised (a) the form of movement trajectories or (b) timing errors made with respect to the external beat. The question of what kinds of movement trajectories assist timing accuracy has not previously been addressed. In an experiment involving synchronisation or syncopation with an external auditory metronome we show that the nervous system produces trajectories that are asymmetric with respect to time and velocity in the out and return phases of the repeating movement cycle. This asymmetry is task specific and is independent of motor implementation details (finger flexion vs. extension). Additionally, we found that timed trajectories are less smooth (higher mean squared jerk) than unpaced ones. The degree of asymmetry in the flexion and extension movement times is positively correlated with timing accuracy. Negative correlations were observed between synchronisation timing error and the movement time of the ensuing return phase, suggesting that late arrival of the finger is compensated by a shorter return phase and conversely for early arrival. We suggest that movement asymmetry in repetitive timing tasks helps satisfy requirements of precision and accuracy relative to a target event.     

Informational and Neuromuscular Contributions to Anchoring in Rhythmic Wrist Cycling

Continuous rhythmic movements are often geared toward particular points in the movement cycle, as evidenced by a local reduction in trajectory variability. These so-called anchor points provide a window into motor control, since changes in the degree of anchoring may reveal how informational and/or neuromuscular properties are exploited in the organization of rhythmic movements. The present experiment examined the relative contributions of informational timing (metronome beeps) and neuromuscular (wrist postures) constraints on anchoring by systematically varying both factors at movement reversal points. To this end, participants cycled their right wrist in a flexed, neutral, or extended posture, either self-paced or synchronized to a metronome pacing peak flexion, peak extension, or both peak flexion and extension. The effects of these manipulations were assessed in terms of kinematics, auditory-motor coordination, and muscle activity. The degree of anchoring seen at the reversal points depended on the degree of compatibility of the prevailing configuration of neuromuscular and informational timing constraints, which had largely independent effects. We further observed systematic changes in muscular activity, which revealed distinct contributions of posture- and muscle-dependent neuromuscular properties to motor control. These findings indicate that the anchor-based discretization of the control of continuous rhythmic wrist movements is determined by both informational timing and neuromuscular constraints in a task-specific manner with subtle interactions between the two, and exemplify how movement variability may be exploited to gain such insights.     

Location but not amount of stimulus occlusion influences the stability of visuo-motor coordination

The current study examined whether the amount and location of available movement information influenced the stability of visuo-motor coordination. Participants coordinated a hand-held pendulum with an oscillating visual stimulus in an inphase and antiphase manner. The effects of occluding different amounts of phase at different phase locations were examined. Occluding the 0°/180° phase locations (end-points) significantly increased the variability of the visuo-motor coordination. The amount of occlusion had little or no affect on the stability of the coordination. We concluded that the end-points of a visual rhythm are privileged and provide access to movement information that ensures stable coordination. The results are discussed with respect to the proposal of Bingham and colleagues (e.g., Bingham GP. Ecol Psychol 16:45–53, 2004a; Wilson AD, Collins DR, Bingham GP. Exp Brain Res 165:351–361, 2005a) that the relevant information for rhythmic visual coordination is relative direction information.     

The relative effects of external spatial and motoric factors on the bimanual coordination of discrete movements

The ability to coordinate the two hands effectively is a fundamental requirement for many everyday tasks. To investigate how bimanual coordination is achieved we asked subjects to perform discrete bimanual key-press responses under conditions in which the motoric (i.e., muscles employed) and external spatial (i.e., direction of movement in external space) relationships between the actions of the left and right index fingers were systematically varied. Subjects made simultaneous right and left index finger key-presses in response to an auditory tone. The right finger always made downward flexion movements whilst the left finger either flexed in a downward/upward direction, or extended in a downward/upward direction. Unimanual control trials of each movement type for both hands were also performed. Reaction times for each hand (RTs) and the inter-response interval (IRI) were recorded . Right hand RTs were significantly affected only when the left finger performed motorically different actions, but were unaffected by the external spatial direction in which the left hands actions were made. The IRI results followed a similar pattern with the worst coordination (highest IRI) occurring when the left finger performed different motor actions to the right finger regardless of the direction of the left hand movement. In contrast to other recent results from experiments examining oscillatory tasks (e.g., Mechsner et al. 2001), our results suggest that in discrete tasks there is a dominance of the motor relationship between the hands over the external spatial relationship.     

Bimanual coordination between isometric contractions and rhythmic movements: an asymmetric coupling

 Interactions between rhythmically moving limbs typically result in attraction to a limited number of coordination modes, which are distinguished in terms of their stability. In addition, the stability of coordination typically decreases with elevations in movement frequency. To gain more insight into the neurophysiological mechanisms underlying these stability characteristics, the effects of phasic voluntary muscle activation onto the movement pattern of the contralateral limb as well as onto the stability of interlimb coordination were examined. This was done in circumstances in which a minimal degree of movement-elicited afferent information was available to mediate the coupling influences. The task involved rhythmic application of isometric torque by one hand, while the other hand was moving rhythmically with unconstrained amplitude. The effects of two levels of applied torque, two coordination patterns (inphase and antiphase), and two movement frequencies were determined, both at the behavioural level (movement kinematics and kinetics) and the neuromuscular level (EMG). The isometric applications of torque clearly influenced the muscle-activation profile and movement pattern of the other limb, affecting both temporal variability and amplitude. Surprisingly, there were no differences between the two coordination patterns or between the tempo conditions. As such, the results did not conform to the Haken-Kelso-Bunz model for rhythmic movement coordination. These data suggest that the archetypal differences in stability of rhythmic bimanual coordination are contingent upon a correspondence between the limbs in terms of their respective tasks. This interpretation is elaborated in terms of the role of sensory feedback and the functional specificity of motor unit recruitment in rhythmic interlimb coordination. Received: 6 November 1998 / Accepted: 7 July 1999     

Visuomotor transformations affect bimanual coupling

Interactions between bimanual movements may occur at two different levels: at a visually based level, where movement trajectories are programmed within the visually perceived external space, and at the executional level, through crosstalk of sensorimotor signals arising during movement execution. In order to distinguish between these sources of interactions, we investigated bimanual reversal movements under different conditions of visual feedback. A visuomotor transformation dissociated movement execution from visual appearance on a computer screen. The transformation we used made movements of the same amplitude evoke different excursions, and made movements of different amplitudes entail matched excursions on the screen. The transformed conditions allowed us to study which parameters of bimanual coupling were related to the way movements were executed and which correlated with the visual movement display. We found a clear dissociation between execution-related and visually related bimanual interactions. The assimilation of movement amplitudes was completely execution-related. Whenever movements of different amplitudes were generated, the shorter movement was lengthened, irrespective of how the movements appeared on the feedback screen. In contrast, temporal coordination at the point of movement reversal, as well as trial-by-trial correlations of movement amplitudes, also showed significant effects of the visuomotor transformation, suggesting that these parameters are influenced by visually perceived effects of movements. This dissociation confirms the idea of separate pathways for bimanual interactions and shows that a specific set of bimanual interactions occur at least partly within a visually based external reference frame. Electronic Publication     

Intermanual interactions during initiation and production of rhythmic and discrete movements in individuals lacking a corpus callosum

Three individuals lacking a corpus callosum, two due to callosotomy and one agenesis, and three age-matched healthy controls were tested on a bimanual task in which a discrete or rhythmic arm movement was initiated following a visual signal while the other arm produced continuous, rhythmic movements. The control participants initiated the secondary, rhythmic movement in phase with the ongoing rhythmic base movement and the two limbs were coupled in an inphase mode across the duration of the trial. In contrast, the acallosal individuals failed to show phase entrainment at the initiation of the secondary, rhythmic movements. Moreover, the callosotomy patients exhibited weak coupling between the rhythmically moving limbs while the individual with callosal agenesis consistently synchronized in an antiphase mode. The control participants exhibited increased perturbation of the ongoing base movement when initiating a discrete movement; for the acallosal participants, the base movement was similarly perturbed in both secondary movement conditions. These results are consistent with the hypothesis that intermanual interactions observed during bimanual movements arise from various levels of control, and that these are distinct for discrete and rhythmic movements. Temporal coupling during rhythmic movements arises in large part from transcallosal interactions between the two hemispheres. The imposition of a secondary movement may transiently disrupt an ongoing rhythmic movement even in the absence of the corpus callosum. This may reflect subcortical interactions associated with response initiation, or, due to dual task demands, a transient shift in attentional resources.     

Unraveling interlimb interactions underlying bimanual coordination

Three sources of interlimb interactions have been postulated to underlie the stability characteristics of bimanual coordination but have never been evaluated in conjunction: integrated timing of feedforward control signals, phase entrainment by contralateral afference, and timing corrections based on the perceived error of relative phase. In this study, the relative contributions of these interactions were discerned through systematic comparisons of five tasks involving rhythmic flexion-extension movements about the wrist, performed bimanually (in-phase and antiphase coordination) or unimanually with or without comparable passive movements of the contralateral hand. The main findings were the following. 1) Contralateral passive movements during unimanual active movements induced phase entrainment to interlimb phasing of either 0 degrees (in-phase) or 180 degrees (antiphase). 2) Entrainment strength increased with the passive movements' amplitude, but was similar for in-phase and antiphase movements. 3) Coordination of unimanual active movements with passive movements of the contralateral hand (kinesthetic tracking) was characterized by similar bilateral EMG activity as observed in active bimanual coordination. 4) During kinesthetic tracking the timing of the movements of the active hand was modulated by afference-based error corrections, which were more pronounced during in-phase coordination. 5) Indications of in-phase coordination being more stable than antiphase coordination were most prominent during active bimanual coordination and marginal during kinesthetic tracking. Together the results indicated that phase entrainment by contralateral afference contributed equally to the stability of in-phase and antiphase coordination, and that differential stability of these patterns depended predominantly on integrated timing of feedforward signals, with only a minor role for afference-based error corrections.     

Location but not amount of stimulus occlusion influences the stability of visuomotor coordination

The current study examined whether the amount and location of available movement information influenced the stability of visuomotor coordination. Participants coordinated a handheld pendulum with an oscillating visual stimulus in an inphase and antiphase manner. The effects of occluding different amounts of phase at different phase locations were examined. Occluding the 0°/180° phase locations (end-points) significantly increased the variability of the visuomotor coordination. The amount of occlusion had little or no affect on the stability of the coordination. We concluded that the end-points of a visual rhythm are privileged and provide access to movement information that ensures stable coordination. The results are discussed with respect to the proposal of Bingham (Ecol Psychol 16:45–43, 2004) and Wilson et al. (Exp Brain Res 165:351–361, 2005) that the relevant information for rhythmic visual coordination is relative direction information.     

Instabilities during antiphase bimanual movements: are ipsilateral pathways involved?

The spatial and temporal coupling between the hands is known to be very robust during movements which use homologous muscles (in-phase or symmetric movements). In contrast, movements using nonhomologous muscles (antiphase or asymmetric movements) are less stable and exhibit a tendency to undergo a phase transition to in-phase movements as movement frequency increases. The instability during antiphase movements has been modeled in terms of signal interference mediated by the ipsilateral corticospinal pathways. In this study we report that participants in whom distal ipsilateral motor-evoked potentials could be elicited with transcranial magnetic stimulation (TMS), exhibited higher variability during a bimanual circling task than participants whose ipsilateral pathways could not be transcranially activated. These results suggest that ipsilateral control of the limb affects the level of bimanual coupling, and may contribute to uncoupling phenomena observed during asymmetric coordination.     

Neural networks for the coordination of the hands in time

Without practice, bimanual movements can typically be performed either in phase or in antiphase. Complex temporal coordination, e.g., during movements at different frequencies with a noninteger ratio (polyrhythms), requires training. Here, we investigate the organization of the neural control systems for in-phase, antiphase, and polyrhythmic coordination using functional magnetic resonance imaging (fMRI). Brisk rhythmic tapping with the index fingers was used as a model behavior. We demonstrate different patterns of brain activity during in-phase and antiphase coordination. In-phase coordination was characterized by activation of the right anterior cerebellum and cingulate motor area (CMA). Antiphase coordination was accompanied by extensive fronto-parieto-temporal activations, including the supplementary motor area (SMA), the preSMA, and the bilateral inferior parietal gyri, premotor cortex, and superior temporal gyri. When contrasting polyrhythmic tapping with in-phase tapping, activity was seen in the same set of brain regions, and in the posterior cerebellum and the CMA. Antiphase and polyrhythmic coordination may thus to a large extent use common neural control circuitry. In a separate experiment, we analyzed the neural control of the rhythmic structure and the serial order of finger movements during polyrhythmic tapping. Polyrhythmic tapping was compared with an isochronous coordination pattern that retained the same serial order of finger movements as the polyrhythm. This experiment showed that the preSMA and the bilateral superior temporal gyri may be crucial for the rhythmic control of polyrhythmic tapping, while the cerebellum, the CMA, and the premotor cortices presumably are more involved in the ordinal control of the sequence of finger movements.     

Functional synchronization in repetitive bimanual prehension movements

To examine the mechanisms of functional bimanual synchronization in goal-directed movements, we studied the movement kinematics of motorically unimpaired subjects while they performed repetitive prehension movements (either unimanually or bimanually) to small food items. Compared to unimanual conditions, bimanual movement execution yielded a significantly prolonged mouth contact phase. We hypothesized that this threefold prolongation led to a proper functional synchronization of the movement onsets of both hands at the beginning of each new movement cycle. That these temporal adjustments occurred in the movement phase with maximal haptic input points to the importance of sensory feedback for bimanual coordination. These results are discussed with respect to the important role of sensory feedback in the timing of coordinated bimanual movements. Furthermore, we propose that time-based coordinating schemas, which are implemented by the cerebellum and the posterior parietal cortex using sensory feedback, underlie functional inter-limb coordination.     

The effect of postural stability and spatial orientation of the upper limbs on interlimb coordination

It has recently been reported that the spatial orientation of two moving limbs has a determining influence on the relative accuracy and stability of coordination patterns. The purpose of the present experiments was to test perceptual and neuromuscular explanations of these spatial orientation effects. Experiment 1 was an initial test of the hypotheses and an extension of a previous study [Lee et al. (2002) Exp Brain Res 146:205-212] that required participants to coordinate inphase and antiphase movement patterns in four spatial orientations: two symmetric orientations (90 degrees and 180 degrees separation between the limbs) and two asymmetric orientations (90 degrees and 135 degrees separation between the limbs). Results of Experiment 1 suggest that the symmetry of movement may be a key factor influencing spatial orientation effects observed during interlimb coordination. In Experiment 2, participants again performed inphase and antiphase movement patterns in symmetric and asymmetric spatial orientations. However, one-half of the participants in Experiment 2 were provided with mechanical constraints during the performance of the desired coordination patterns. The mechanical constraints provided postural support but did not influence the visual experience. Results showed that the addition of the postural support improved performance. These findings suggest that neuromuscular, and perhaps biomechanical, constraints contribute more to the influence of spatial orientation than visual-perceptual constraints.     

Effect of transcranial magnetic stimulation on bimanual movements

Transcranial magnetic stimulation (TMS) of the motor cortex can interrupt voluntary contralateral rhythmic limb movements. Using the method of "resetting index" (RI), our study investigated the TMS effect on different types of bimanual movements. Six normal subjects participated. For unimanual movement, each subject tapped either the right or left index finger at a comfortable rate. For bimanual movement, index fingers of both hands tapped in the same (in-phase) direction or in the opposite (antiphase) direction. TMS was applied to each hemisphere separately at various intensities from 0.5 to 1.5 times motor threshold (MT). TMS interruption of rhythm was quantified by RI. For the unimanual movements, TMS disrupted both contralateral and ipsilateral rhythmic hand movements, although the effect was much less in the ipsilateral hand. For the bimanual in-phase task, TMS could simultaneously reset the rhythmic movements of both hands, but the effect on the contralateral hand was less and the effect on the ipsilateral hand was more compared with the unimanual tasks. Similar effects were seen from right and left hemisphere stimulation. TMS had little effect on the bimanual antiphase task. The equal effect of right and left hemisphere stimulation indicates that neither motor cortex is dominant for simple bimanual in-phase movement. The smaller influence of contralateral stimulation and the greater effect of ipsilateral stimulation during bimanual in-phase movement compared with unimanual movement suggest hemispheric coupling. The antiphase movements were resistant to TMS disruption, and this suggests that control of rhythm differs in the 2 tasks. TMS produced a transient asynchrony of movements on the 2 sides, indicating that both motor cortices might be downstream of the clocking command or that the clocking is a consequence of the 2 hemispheres communicating equally with each other.