The adaptation to sensory information in the production of bimanual movement patterns

Bimanual in-phase and anti-phase patterns were performed in the transverse plane under optimal and degraded proprioceptive conditions, i.e., without and with tendon vibration. Moreover, proprioceptive information was changed midway into each trial to examine on-line reorganization. In addition to the proprioceptive perturbation, the availability of visual information was manipulated to study to which degree sensory information from different modalities interact. Movement patterns performed under identical sensory conditions were compared, i.e., the first 15 s (control) and the 15 s following a change in afferent input (transfer). In the control and transfer conditions, movements with vibrations were less accurate than those without vibrations indicating the influence of optimal proprioceptive information in the calibration and recalibration of intrinsic bimanual movement patterns. Furthermore, pattern stability was affected by the nature of the transfer condition. This indicated that the degree of fluctuations in a sensory transfer situation depended upon the quality of the proprioceptive information experienced in the initial conditions. The influence of visual information was not without importance, although the nature of the coordination mode must be taken into account. In the control conditions, in-phase movements were less stable when vision was absent, whereas anti-phase movements were more stable when vision was not present. This observation was made independent of the available proprioceptive information revealing differences in visual guidance between both coordination modes. In the transfer conditions, pattern stability was similar during the vision and no-vision conditions suggesting a limited influence of visual information in the recalibration process.     

Interaction of neuromuscular, spatial and visual constraints on hand-foot coordination dynamics

In the present study, we investigated the contributions of motor and perceptual processes to directional constraints as observed during hand-foot coordination. Participants performed cyclical flexion-extension movements of the right hand and foot under two coordination modes: in-phase (isodirectional) and antiphase (non-isodirectional). Those tasks were performed either with full vision or no vision of the limbs. Depending on the position of the forearm (prone or supine), the coordination patterns were performed with similar and dissimilar neuro-muscular coupling with respect to their phylogenetic origin as antigravity muscles. Results showed that the antiphase pattern was more difficult to maintain than the in-phase pattern and that neuro-muscular coupling significantly influenced the coordination dynamics. Moreover, the effect of vision differed as a function of both neuro-muscular coupling and coordination mode. Under dissimilar neuro-muscular coupling, the presence of visual feedback stabilized the in-phase pattern and destabilized the antiphase pattern. In contrast, visual feedback did not influence pattern stability during conditions of similar neuro-muscular coupling. These results shed light on the complex interactions between motor and perceptual (visual) constraints during the production of hand-foot coordination patterns.     

Effects of correct and transformed visual feedback on rhythmic visuo-motor tracking: tracking performance and visual search behavior

The effects of correct and transformed visual feedback on rhythmic unimanual visuo-motor tracking were examined, focusing on tracking performance (accuracy and stability) and visual search behavior. Twelve participants (reduced to 9 in the analyses) manually tracked an oscillating visual target signal in phase (by moving the hand in the same direction as the target signal) and in antiphase (by moving the hand in the opposite direction), while the frequency of the target signal was gradually increased to probe pattern stability. Besides a control condition without feedback, correct feedback (representing the actual hand movement) or mirrored feedback (representing the hand movement transformed by 180 degrees) were provided during tracking, resulting in either in-phase or antiphase visual motion of the target and feedback signal, depending on the tracking mode performed. The quality (accuracy and stability) of in-phase tracking was hardly affected by the two forms of feedback, whereas antiphase tracking clearly benefited from mirrored feedback but not from correct feedback. This finding extends previous results indicating that the performance of visuo-motor coordination tasks is aided by visual feedback manipulations resulting in coherently grouped (i.e., in-phase) visual motion structures. Further insights into visuo-motor tracking with and without feedback were garnered from the visual search patterns accompanying task performance. Smooth pursuit eye movements only occurred at lower oscillation frequencies and prevailed during in-phase tracking and when target and feedback signal moved in phase. At higher frequencies, point-of-gaze was fixated at a location that depended on the feedback provided and the resulting visual motion structures. During in-phase tracking the mirrored feedback was ignored, which explains why performance was not affected in this condition. Point-of-gaze fixations at one of the end-points were accompanied by reduced motor variability at this location, reflecting a form of visuo-motor anchoring that may support the pick up of discrete information as well as the control of hand movements to a desired location.     

The metabolic and cognitive energy costs of stabilising a high-energy interlimb coordination task

Kinematic (relative phase error), metabolic (oxygen consumption, heart rate) and attentional (baseline and cycling reaction times) variables were measured while participants practised a high energy-demanding, intrinsically unstable 90 degrees relative phase coordination pattern on independent bicycle ergometers. The variables were found to be strongly inter-correlated, suggesting a link between emerging performance stability with practice and minimal metabolic and attentional cost. The effects of practice of 90 degrees relative phase coordination on the performance of in-phase (0 degrees-phase) and antiphase (180 degrees-phase) coordination were investigated by measuring the relative phase attractor layouts and recording the metabolic and attentional cost of the three coordination patterns before and after practice. The attentional variables did not differ significantly between coordination patterns and did not change with practice. Before practice, the coordination performance was most accurate and stable for in-phase cycling, with antiphase next and 90 degrees-phase the poorest. However, metabolic cost was lower for antiphase than either in-phase or 90 degrees-phase cycling, and the pre-practice attractor layout deviated from that predicted on the basis of dynamic stability as an attractor state, revealing an attraction to antiphase cycling. After practice of 90 degrees-phase cycling, in-phase cycling remained the most accurate and stable, with 90 degrees-phase next and antiphase the poorest, but antiphase retained the lowest metabolic energy cost. The attractor layout had changed, with new attractors formed at the practised 90 degrees-phase pattern and its symmetrical partner of 270 degrees-phase. Considering both the pre- and post-practice results, attractors were formed at either a low metabolic energy cost but less stable (antiphase) pattern or at a more stable but higher metabolic energy cost (90 degrees-phase) pattern, but in neither case at the most stable and accurate (in-phase) pattern. The results suggest that energetic factors affect coordination dynamics and that coordination modes lower in metabolic energy expenditure may compete with dynamically stable modes.     

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

The authors tested for predominant patterns of coordination in the combination of rhythmic flexion-extension (FE) and supination- (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. The result provides evidence that the predominance of the in-phase pattern originates in the influence of neuromuscular-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.     

Phase Transitions and Critical Fluctuations in Rhythmic Coordination of Ipsilateral Hand and Foot

Four subjects performed rhythmic movements of the ankle and the wrist in time with an auditory metronome, in two modes of coordination, antiphase and in-phase. The forearm was placed in either a prone or a supine position. When movements were prepared in the antiphase mode, spontaneous transitions to the in-phase mode, or to phase wandering were observed as metronome frequency was increased. When prepared in the in-phase mode, transitions between in-phase modes or to phase wandering were occasionally observed. Predicted signature features of nonequilbrium phase transitions were noted, including loss of stability and critical fluctuations. The stability of the movement patterns was determined by spatial (dependent upon the direction of movement) rather than anatomical (dependent on the coupling of specific muscle groups) constraints. The position of the forearm had no consistent bearing upon the variability of the phase relations between the limbs, the frequency of phase transitions, or the time of onset of transitions. These results are discussed with reference to the coordination dynamics (e.g., multistability, loss of stability) of multijoint movements.     

Intentional switching between patterns of bimanual coordination depends on the intrinsic dynamics of the patterns

Two predictions arising from previous theoretical and empirical work which demonstrated that spontaneous changes of bimanual coordination patterns result from a loss of pattern stability (i.e., a nonequilibrium phase transition) were tested: (a) that the time it takes to intentionally switch from one pattern to another depends on the differential stability of the patterns themselves; and (b) that an intention, defined as an intended behavioral pattern, can change the dynamical characteristics, e.g., the overall stability of the behavioral patterns. Subjects moved both index fingers rhythmically at one of six movement frequencies while performing either an in-phase or antiphase pattern of finger coordination. On cue from an auditory signal, subjects switched from the ongoing pattern to the other pattern. The relative phase of movement between the two fingers was used to characterize the ongoing coordinative pattern. The time taken to switch between patterns, or switching time, and relative phase fluctuations were used to evaluate the modified pattern dynamics resulting from a subject's intention to change patterns. Switching from the in-phase to the antiphase pattern was significantly slower than switching in the opposite direction for all subjects. Both the mean and distribution of switching times in each direction were found to be in agreement with model predictions. movement frequency had little effect on switching time, a finding that is also consistent with the model. Relative phase fluctuations were significantly larger when moving in the antiphase pattern at the highest movement frequencies studied. The results show that, although intentional influences act to modify a coordinative pattern's intrinsic dynamics, the influence of these dynamics on the resulting behavior is always present and is particularly strong at high movement frequencies.     

The integration of temporally shifted visual feedback in a synchronization task: The role of perceptual stability in a visuo-proprioceptive conflict situation

The present study examined whether the beneficial role of coherently grouped visual motion structures for performing complex (interlimb) coordination patterns can be generalized to synchronization behavior in a visuo-proprioceptive conflict situation. To achieve this goal, 17 participants had to synchronize a self-moved circle, representing the arm movement, with a visual target signal corresponding to five temporally shifted visual feedback conditions (0%, 25%, 50%, 75%, and 100% of the target cycle duration) in three synchronization modes (in-phase, anti-phase, and intermediate). The results showed that the perception of a newly generated perceptual Gestalt between the visual feedback of the arm and the target signal facilitated the synchronization performance in the preferred in-phase synchronization mode in contrast to the less stable anti-phase and intermediate mode. Our findings suggest that the complexity of the synchronization mode defines to what extent the visual and/or proprioceptive information source affects the synchronization performance in the present unimanual synchronization task.     

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.     

Intentional Switching Between Patterns of Bimanual Coordination Depends on the Intrinsic Dynamics of the Patterns

Two predictions arising from previous theoretical and empirical work which demonstrated that spontaneous changes of bimanual coordination patterns result from a loss of pattern stability (i.e., a nonequilibrium phase transition) were tested: (a) that the time it takes to intentionally switch from one pattern to another depends on the differential stability of the patterns themselves; and (b) that an intention, defined as an intended behavioral pattern, can change the dynamical characteristics, e.g., the overall stability of the behavioral patterns. Subjects moved both index fingers rhythmically at one of six movement frequencies while performing either an in-phase or antiphase pattern of finger coordination. On cue from an auditory signal, subjects switched from the ongoing pattern to the other pattern. The relative phase of movement between the two fingers was used to characterize the ongoing coordinative pattern. The time taken to switch between patterns, or switching time, and relative phase fluctuations were used to evaluate the modified pattern dynamics resulting from a subject's intention to change patterns. Switching from the in-phase to the antiphase pattern was significantly slower than switching in the opposite direction for all subjects. Both the mean and distribution of switching times in each direction were found to be in agreement with model predictions. Movement frequency had little effect on switching time, a finding that is also consistent with the model. Relative phase fluctuations were significantly larger when moving in the antiphase pattern at the highest movement frequencies studied. The results show that, although intentional influences act to modify a coordinative pattern's intrinsic dynamics, the influence of these dynamics on the resulting behavior is always present and is particularly strong at high movement frequencies.     

Diffusive,Synaptic, and Synergetic Coupling: An Evaluation Through In-Phase and Antiphase Rhythmic Movements

The in-phase and antiphase patterns of interlimb l:1 frequency locking were contrasted with respect to models of coordination dynamics in biological movement systems that are based on diffusive coupling, synaptic coupling, and synergetic principles. Predictions were made from each model concerning the stable relative phase phi between the rhythmic units, its standard deviation SDphi and the self-chosen coupled frequency omegasubc;. The experimental task involved human subjects oscillating two handheld pendulums either in-phase or antiphase. The eigenfrequencies of the two hand-pendulum systems were manipulated by varying the length and mass of each pendulum individually. Relative to an eigenfrequency difference of Delta equal to zero, |Deltaomega| > 0 displaced phi from phi = 0 and phi = pi, and amplified SDphi. omegasubc; decreased with |Deltaomega|. Both the displacement of phi and SDphi were greater in the antiphase mode. Additionally, the displacement of phi increased more sharply with |Delta| for antiphase than for in-phase coordination. In contrast, omegasubc; was identical for the two coordination modes. Of the models of interlimb coordination dynamics, the synergetic model was the most successful in addressing the pattern of dependencies of phi and SDphi. The specific forms of the functions relating omegasubc; and phi to Deltaomega pose challenges for all three models, however     

Rocking together: dynamics of intentional and unintentional interpersonal coordination

The current study investigated the interpersonal coordination that occurred between two people when sitting side-by-side in rocking chairs. In two experiments participant pairs rocked in chairs that had the same or different natural periods. By instructing pairs to coordinate their movements inphase or antiphase, Experiment 1 investigated whether the stable patterns of intentional interpersonal coordination were consistent with the dynamics of within person interlimb coordination. By instructing the participants to rock at their own preferred tempo, Experiment 2 investigated whether the rocking chair movements of visually coupled individuals would become unintentionally coordinated. The degree to which the participants fixated on the movements of their co-actor was also manipulated to examine whether visual focus modulates the strength of interpersonal coordination. As expected, the patterns of coordination observed in both experiments demonstrated that the intentional and unintentional interpersonal coordination of rocking chair movements is constrained by the self-organizing dynamics of a coupled oscillator system. The results of the visual focus manipulations indicate that the stability of a visual interpersonal coupling is mediated by attention and the degree to which an individual is able to detect information about a co-actor's movements.     

Examination of reactive motor responses to Achilles tendon vibrations during an inhibitory stepping reaction time task

Inhibition is known to influence balance, step initiation and gait control. A specific subcomponent of inhibition, the perceptual inhibition process, has been suggested to be specifically involved in the integration of proprioceptive information that is necessary for efficient postural responses. This study aimed to investigate the inhibition requirements of planning and executing a choice step initiation task in young adults following experimental perturbation of proprioceptive information using Achilles tendon vibrations. We developed an inhibitory stepping reaction time task in which participants had to step in response to visual arrows that manipulated specific perceptual or motor inhibition according to two proprioceptive configurations: without or with application of vibrations. Performance of twenty-eight participants (mean age 21 years) showed that Achilles tendon vibrations induced an increase in attentional demands (higher reaction time and longer motor responses). Further, this increase in attentional demands did not affect specifically the different inhibitory processes tested in this reactive stepping task. It suggests that attentional demands associated with the vibratory perturbation to postural control do not lead to a shift from automatic to more attentional inhibition processes, at least in young adults.     

Intentional Switching Between Bimanual Coordination Patterns

Previous theoretical and empirical work indicates that intentional changes in a bimanual coordination pattern depends on the stability of the bimanual coordination pattern (Kelso, Schotz, & Schöner, 1988; Scholz & Kelso, 1990). The present experiments retest this notion when online Lissajous displays are provided. Switching to and from in-phase and antiphase and to and from 90° and 270° were tested in Experiment 1. Participants were able to very effectively produce the 180°, 90°, and 270° coordination patterns although performance of the in-phase coordination task was even more stable. The data indicated that switching to in-phase from antiphase was more rapid than vice versa and that switching times between 90° to 270° were similar. Experiment 2 investigated switching between 1:2 and 2:1 bimanual coordination patterns. The results indicated that switching time was similar between the 2:1 and 1:2 coordination tasks and that increases in stability over practice resulted in additional decreases in switching times. This provides additional evidence that the attractor landscape is fundamentally different dependent on the type of information provided the performer. What remains to be done is to reconcile these results with the various theories/perspectives currently used to describe and explain bimanual coordination.     

Bimanual Finger Tapping: Effects of Frequency and Auditory Information on Timing Consistency and Coordination

The authors' goal in this study was to probe the basis for an earlier, unexpected finding that preferred-frequency finger tapping tends to have higher frequencies and to be less stable for in-phase than for antiphase tasks. In follow-up experiments, 3 protocols were employed: a preferred-frequency replication in both coordination modes, a metronome-driven matching of the preferred frequencies to each of the coordination modes, and a frequency scaling of both modes. The original findings were affirmed for preferred frequency. Tapping to a metronome had a differential effect on in-phase and antiphase: A more stable coupling across frequencies was exhibited during in-phase. Under frequency scaling, the antiphase pattern decomposed at lower frequencies than did in-phase, but no phase transitions were observed. The loss of stable coordination in both modes was attended by sudden increases in frequency differences between fingers and by phase wandering. The emergence of those effects is discussed in light of asymmetric modifications to the Haken-Kelso-Bunz model (H. Haken, J. A. S. Kelso, & H. Bunz, 1985) and the task constraints of tapping.     

Coordination Dynamics of the Bipedal Galloping Pattern

A motion equation in relative phase was developed that incorporates the spatial-temporal pattern of the bipedal gallop along with the more commonplace patterns of the bipedal jump and walk-run. In 3 experiments, human participants (N = 6 per experiment) simulated the bipedal gait patterns through the rhythmic motions of hand-held pendulums. Predictions of the motion equation for coordination equilibria and their respective degrees of stability were confirmed. In particular, the gallop pattern was less stable than the fundamental in-phase and antiphase patterns but changed in qualitatively similar ways to those gaits as a function of limb asymmetry and movement frequency. The relation between the modeled coordination dynamics and the kinematic characteristics of real bipedal galloping is discussed.     

Bimanual finger tapping: effects of frequency and auditory information on timing consistency and coordination

The authors' goal in this study was to probe the basis for an earlier, unexpected finding that preferred-frequency finger tapping tends to have higher frequencies and to be less stable for in-phase than for antiphase tasks. In follow-up experiments, 3 protocols were employed: a preferred-frequency replication in both coordination modes, a metronome-driven matching of the preferred frequencies to each of the coordination modes, and a frequency scaling of both modes. The original findings were affirmed for preferred frequency. Tapping to a metronome had a differential effect on in-phase and antiphase: A more stable coupling across frequencies was exhibited during in-phase. Under frequency scaling, the antiphase pattern decomposed at lower frequencies than did in-phase, but no phase transitions were observed. The loss of stable coordination in both modes was attended by sudden increases in frequency differences between fingers and by phase wandering. The emergence of those effects is discussed in light of asymmetric modifications to the Haken-Kelso-Bunz model (H. Haken, J. A. S. Kelso, & H. Bunz, 1985) and the task constraints of tapping.     

Cathodal tDCS of the Left Posterior Parietal Cortex Increases Proprioceptive Drift

In aiming movements the limb position drifts away from the defined target after some trials without visual feedback, a phenomenon defined as proprioceptive drift (PD). There are no studies investigating the association between the posterior parietal cortex (PPC) and PD in aiming movements. Therefore, cathodal and sham transcranial direct current stimulation (tDCS) were applied to the left PPC concomitantly with the performance of movements with or without vision. Cathodal tDCS applied without vision produced a higher level of PD and higher rates of drift accumulation while it decreased peak velocity and maintained the number of error corrections, not affecting movement amplitude. The proprioceptive information seems to produce an effective reference to movement, but with PPC stimulation it causes a negative impact on position.     

To Switch or Not to Switch: Recruitment of Degrees of Freedom Stabilizes Biological Coordination

Recruitment and suppression processes were studied in the swinging-pendulum paradigm (cf. P. N. Kugler & M. T. Turvey, 1987). The authors pursued the hypothesis that active recruitment of previously unmeasured degrees of freedom serves to stabilize an antiphase bimanual coordination pattern and thereby obviates the need for pattern switching from an antiphase to an in-phase coordination pattern, a key prediction of the H. Haken, J. A. S. Kelso, and H. Bunz (1985) model. In Experiment 1, 7 subjects swung single hand-held pendulums in time with an auditory metronome whose frequency increased. Pendulum motion changed from planar (2D) to elliptical (3D), and forearm motion (produced by elbow flexion-extension) was recruited with increasing movement rate for cycling frequencies typically above the pendulum's eigenfrequency. In Experiment 2, 7 subjects swung paired pendulums in either an in-phase or an antiphase coordinative mode as movement rate was increased. With the systematic increase in movement rate, the authors attempted to induce transitions from the antiphase to the in-phase coordinative pattern, with loss of stability the key mechanism of pattern change. Transitions from the antiphase to the in-phase coordinative mode were not observed. Pattern stability, as defined by the variability of the phase relation between the pendulums, was affected only a little by increasing movement rate. As in the single-pendulum case, pendulum motion changed from planar to elliptical, and forearm motion was recruited with increasing cycling frequency. Those results reveal a richer dynamics than previously observed in the pendulum paradigm and support the hypothesis that recruitment processes stabilize coordination in biomechanically redundant systems, thereby reducing the need for pattern switching.     

Age-related differences and the role of augmented visual feedback in learning a bimanual coordination pattern

The purpose of this study was to investigate the effects of aging and the role of augmented visual information in the acquisition of a new bimanual coordination pattern, namely a 90° relative phase pattern. In a pilot study, younger and older adults received augmented visual feedback in the form of a real-time orthogonal display of both limb movements after every fifth trial. Younger adults acquired this task over three days of practice and retained the task well over periods of one week and one month of no practice while the older adults showed no improvement at all on the task. It was hypothesized that the amount of augmented information was not sufficient for the older adults to overcome the strong tendency to perform natural, intrinsically stable coordination patterns, which consequently prevented them from learning the task. The present study evaluated the age-related role of augmented visual feedback for learning the new pattern. Participants were randomly assigned within age groups to receive either concurrent or terminal visual feedback after every trial in acquisition. In contrast to the pilot study, all of the older adults learned the pattern, although not to the same level as the younger adults. Both younger and older adults benefitted from concurrent visual feedback, but the older adults gained more from the concurrent feedback than the younger adults, relative to terminal feedback conditions. The results suggest that when learning bimanual coordination patterns, older adults are more sensitive to the structure of the practice conditions, particularly the availability of concurrent visual information. This greater sensitivity to the learning environment may reflect a diminished capacity for inhibitory control and a decreased ability to focus attention on the salient aspects of learning the task.