Moving the Arm at Different Rates: Slow Movements are Avoided

Qualitative and quantitative changes characterize locomotion and rhythmic interlimb coordination at different speeds. Legs and hands do not move more or less quickly; they also adopt different relative coordination patterns. In the present article, the authors asked whether similar transitions occur for unimanual hand movements when speed is slowed below the preferred speed. Participants moved a handheld dowel back and forth between 2 large circular targets in time with a metronome at periods between 370 ms and 1667 ms. The authors analyzed the kinematics of participants’ movements at each period and found that proportional dwell time and number of peaks in the velocity profile increased as driving periods increased. Path lengths and peak velocities remained relatively constant for driving periods exceeding 800 ms. Participants made only gradual changes to their movement parameters, so that they went from a continuous mode to a more discrete mode of behavior for longer driving periods. Thus, unlike for rhythmic bimanual movements or locomotory patterns, there are quantitative but no clear qualitative changes for unimanual movements. The results suggest that participants tried to move close to their preferred tempo at different rates, and that they avoided moving slowly.     

The temporal representation of in-phase and anti-phase movements

We have proposed that the stability of bimanual coordination is influenced by the complexity of the representation of the task goals. Here, we present two experiments to explore this hypothesis. First, we examined whether a temporal event structure is present in continuous movements by having participants vocalize while producing bimanual circling movements. Participants tended to vocalize once per movement cycle when moving in-phase. In contrast, vocalizations were not synchronized with anti-phase movements. While the in-phase result is unexpected, the latter would suggest anti-phase continuous movements lack an event structure. Second, we examined the event structure of movements marked by salient turn-around points. Participants made bimanual wrist flexion movements and were instructed to move 'in synchrony' with a metronome, without specifying how they should couple the movements to the metronome. During in-phase movements, participants synchronized one hand cycle with every metronome beat; during anti-phase movements, participants synchronized flexion of one hand with one metronome beat and extension of the other hand with the next beat. The results are consistent with the hypothesis that the instability of anti-phase movements is related to their more complex (or absent) event representation relative to that associated with in-phase movements.     

Coordination dynamics of rhythmical and discrete prehension movements: Implications of the scanning procedure and individual differences

The aim of this study was to compare the coordination dynamics of discrete and rhythmical reaching and grasping movements from a dynamical systems perspective. Previous research from this theoretical perspective had focused on rhythmical actions and it is unclear to what extent discrete movements are amenable to a similar dynamical systems analysis. Six adult subjects performed prehension in two conditions: a discrete, non-continuous mode and a rhythmical, continuous mode. A `scanning procedure' was implemented between pre- and post-tests in which the required time of final relative hand closure (T_rfc) was systematically varied. It was shown that the error in the reaching and grasping pattern was least at an attractor region and systematically increased with deviation from the attractor. Results also indicated that there were no differences between condition or trial block for the group. However, there were several within-subject effects of interest. The validity of the scanning procedure was found to be questionable in the discrete condition, where four subjects showed differences in T_rfc between pre- or post-test and the predicted T_rfc of the scanning procedure. Four out of six subjects also had different preferred T_rfc values for discrete and rhythmical movement, indicating that individual specific models might need to be constructed for future dynamical modelling of discrete movement.PsycINFO classification: 2330     

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.     

GPS analysis of human locomotion: further evidence for long-range correlations in stride-to-stride fluctuations of gait parameters

During free walking, gait is automatically adjusted to provide optimal mechanical output and minimal energy expenditure; gait parameters, such as cadence, fluctuate from one stride to the next around average values. It was described that this fluctuation exhibited long-range correlations and fractal-like patterns. In addition, it was suggested that these long-range correlations disappeared if the participant followed the beep of metronome to regulate his or her pace. Until now, these fractal fluctuations were only observed for stride interval, because no technique existed to adequately analyze an extended time of free walking. The aim of the present study was to measure walking speed (WS), step frequency (SF) and step length (SL) with high accuracy (<1 cm) satellite positioning method (global positioning system or GPS) in order to detect long-range correlations in the stride-to-stride fluctuations. Eight participants walked 30 min under free and constrained (metronome) conditions. Under free walking conditions, DFA (detrended fluctuation analysis) and surrogate data tests showed that the fluctuation of WS, SL and SF exhibited a fractal pattern (i.e., scaling exponent alpha: 0.5 < alpha < 1) in a large majority of participants (7/8). Under constrained conditions (metronome), SF fluctuations became significantly anti-correlated (alpha < 0.5) in all participants. However, the scaling exponent of SL and WS was not modified. We conclude that, when the walking pace is controlled by an auditory signal, the feedback loop between the planned movement (at supraspinal level) and the sensory inputs induces a continual shifting of SF around the mean (persistent anti-correlation), but with no effect on the fluctuation dynamics of the other parameters (SL, WS).     

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.     

Space-time behavior of single and bimanual rhythmical movements: data and limit cycle model

How do space and time relate in rhythmical tasks that require the limbs to move singly or together in various modes of coordination? And what kind of minimal theoretical model could account for the observed data? Earlier findings for human cyclical movements were consistent with a nonlinear, limit cycle oscillator model (Kelso, Holt, Rubin, & Kugler, 1981) although no detailed modeling was performed at that time. In the present study, kinematic data were sampled at 200 samples/second, and a detailed analysis of movement amplitude, frequency, peak velocity, and relative phase (for the bimanual modes, in phase and antiphase) was performed. As frequency was scaled from 1 to 6 Hz (in steps of 1 Hz) using a pacing metronome, amplitude dropped inversely and peak velocity increased. Within a frequency condition, the movement's amplitude scaled directly with its peak velocity. These diverse kinematic behaviors were modeled explicitly in terms of low-dimensional (nonlinear) dissipative dynamics, with linear stiffness as the only control parameter. Data and model are shown to compare favorably. The abstract, dynamical model offers a unified treatment of a number of fundamental aspects of movement coordination and control.     

The distinction between tapping and circle drawing with and without tactile feedback: an examination of the sources of timing variance

An internal clock-like process has been implicated in the control of rhythmic movements performed for short (250-2,000 ms) time scales. However, in the past decade, it has been claimed that a clock-like central timing mechanism is not required for smooth cyclical movements. The distinguishing characteristic delineating clock-like (event) from non-clock-like (emergent) timing is thought to be the kinematic differences between tapping (discrete-like) and circle drawing (smooth). In the archetypal event-timed task (tapping), presence of perceptual events is confounded with the discrete kinematics of movement (table contact). Recently, it has been suggested that discrete perceptual events help participants synchronize with a metronome. However, whether discrete tactile events directly elicit event timing has yet to be determined. In the present study, we examined whether a tactile event inserted into the circle drawing timing task could elicit event timing in a self-paced (continuation) timing task. For a majority of participants, inserting an event into the circle drawing task elicited timing behaviour consistent with the idea that an internal timekeeper was employed (a correlation of circle drawing with tapping). Additionally, some participants exhibited characteristics of event timing in the typically emergently timed circle drawing task. We conclude that the use of event timing can be influenced by the insertion of perceptual events, and it also exhibits persistence over time and over tasks within certain individuals.     

The distinction between tapping and circle drawing with and without tactile feedback: An examination of the sources of timing variance

An internal clock-like process has been implicated in the control of rhythmic movements performed for short (250–2,000 ms) time scales. However, in the past decade, it has been claimed that a clock-like central timing mechanism is not required for smooth cyclical movements. The distinguishing characteristic delineating clock-like (event) from non-clock-like (emergent) timing is thought to be the kinematic differences between tapping (discrete-like) and circle drawing (smooth). In the archetypal event-timed task (tapping), presence of perceptual events is confounded with the discrete kinematics of movement (table contact). Recently, it has been suggested that discrete perceptual events help participants synchronize with a metronome. However, whether discrete tactile events directly elicit event timing has yet to be determined. In the present study, we examined whether a tactile event inserted into the circle drawing timing task could elicit event timing in a self-paced (continuation) timing task. For a majority of participants, inserting an event into the circle drawing task elicited timing behaviour consistent with the idea that an internal timekeeper was employed (a correlation of circle drawing with tapping). Additionally, some participants exhibited characteristics of event timing in the typically emergently timed circle drawing task. We conclude that the use of event timing can be influenced by the insertion of perceptual events, and it also exhibits persistence over time and over tasks within certain individuals.     

Cancelling discrete and stopping ongoing rhythmic movements: Do they involve the same process of motor inhibition?

Motor inhibition is considered to be an important process of executive control and to be implicated in numerous activities in order to cancel prepared actions and, supposedly, to suppress ongoing ones. Usually, it is evaluated using a “stop-signal task” in which participants have to inhibit prepared discrete movements. However, it is unknown whether other movement types involve the same inhibition process. We therefore investigated whether the inhibition process for discrete movements is involved in stopping ongoing rhythmic movements as well.Twenty healthy adults performed two counterbalanced tasks. The first task was used to estimate the stop-signal reaction time (SSRTd) needed to inhibit prepared discrete key-pressing movements. In the second task, participants drew graphic patterns on a tablet and had to stop the movement when a stop-signal occurred. We calculated the rhythmic stop signal-reaction time as the time needed to initiate stopping such ongoing rhythmic movement (SSRTr) and the same latency relative to the period of the rhythmic movement (relSSRTr). We measured these delays under different movement frequencies and motor coordination conditions and further investigated whether they varied as a function of several parameters of the rhythmic movements (speed, mean and variance of the relative phase, and movement phase at several time events).We found no correlation between inhibition measures in the two tasks. In contrast, generalized linear models showed a moderate yet significant influence of the motion parameters on the inhibition of ongoing rhythmic movements. We therefore conclude that the motor inhibition processes involved in cancelling prepared discrete movements and stopping ongoing rhythmic movements are dissimilar.     

Comfortable synchronization of cyclic drawing movements with a metronome

Continuous circle drawing is considered a paragon of emergent timing, whereas the timing of finger tapping is said to be event-based. Synchronization with a metronome, however, must to some extent be event-based for both types of movement. Because the target events in the movement trajectory are more poorly defined in circle drawing than in tapping, circle drawing shows more variable asynchronies with a metronome than does tapping. One factor that may have contributed to high variability in past studies is that circle size, drawing direction, and target point were prescribed and perhaps outside the comfort range. In the present study, participants were free to choose most comfortable settings of these parameters for two continuously drawn shapes, circles and infinity signs, while synchronizing with a regular or intermittently perturbed metronome at four different tempi. Results showed that preferred circle sizes were generally smaller than in previous studies but tended to increase as tempo decreased. Synchronization results were similar for circles and infinity signs, and similar to earlier results for circles drawn within a fixed template (Repp & Steinman, 2010). Comparison with tapping data still showed drawing to exhibit much greater variability and persistence of asynchronies as well as slower phase correction in response to phase shifts in the metronome. With comfort level ruled out as a factor, these differences can now be attributed more confidently to differences in event definition and/or movement dynamics.     

Interlimb coordination following stroke

Studies investigating whether simultaneous bilateral movements can facilitate performance of the impaired limb(s) of stroke patients have returned mixed results. In the present study we compared unilateral limb performance (amplitude, cycle duration) with performance during an interlimb coordination task involving both homologous (both arms, both legs) and non-homologous (one arm, one leg) limbs in stroke participants (n=7) and healthy age-matched controls (n=7). In addition, the effect of on-line augmented visual feedback on interlimb coordination was investigated. Participants performed cyclical flexion-extension movements of the arms and legs in the sagittal plane paced by an auditory metronome (1 Hz). Movement amplitudes were larger and cycle durations shorter during homologous limb coordination than non-homologous coordination. Compared with unilateral movements both groups had reduced movement amplitudes and the stroke group increased cycle duration when interlimb coordination tasks were performed. These effects were most evident during non-homologous (arm and leg) coordination. No evidence of facilitation of the impaired limb(s) was found in any of the interlimb coordination conditions. Augmented visual feedback had minimal effect on the movements of control participants but lead to an increase of cycle duration for stroke participants.     

Parameter value switching in discrete and continuous aiming movements

The effect of practice variations on spatial and temporal accuracy was investigated in both discrete and continuous aiming movements in the preferred hand of college-aged participants (N=25). In a completely within-subject design, participants made rapid reversal movements with a lightweight lever in the sagittal plane, practicing 20 degrees and 60 degrees movements in repeated (same distance) and alternating (switching between 20 degrees and 60 degrees) conditions. Movements were also made one at a time (discretely) or in sequences of 20 movements (continuously). Spatial constant error, spatial variable error, spatial overall error, the coefficient of variation, movement time, and the relative timing were calculated for each set of 20 movements and analyzed by within-subject analyses of variance. Movements in the repeated conditions for both discrete and continuous movements were more accurate and consistent compared to the alternating condition where the short movements were overshot and the long movements were undershot. Discrete movements were more spatially and temporally variable than continuous movements. The discrete and continuous movements showed different relative timing patterns, suggesting that the temporal structure of the motor program is affected by task characteristics.     

Hand path priming in manual obstacle avoidance: evidence for abstract spatiotemporal forms in human motor control

Previous research suggests that motor equivalence is achieved through reliance on effector-independent spatiotemporal forms. Here the authors report a series of experiments investigating the role of such forms in the production of movement sequences. Participants were asked to complete series of arm movements in time with a metronome and, on some trials, with an obstacle between 1 or more of the target pairs. In moves following an obstacle, participants only gradually reduced the peak heights of their manual jumping movements. This hand path priming effect, scaled with obstacle height, was preserved when participants cleared the obstacle with 1 hand and continued with the other, and it was modulated by future task demands. The results are consistent with the hypothesis that the control of movement sequences relies on abstract spatiotemporal forms. The data also support the view that motor programming is largely achieved by changing just those features that distinguish the next movement to be made from the movement that was just made.     

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

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

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.     

The coupled oscillator model of between-hand coordination in alternate-hand tapping: a reappraisal

Single and alternating hand tapping were compared to test the hypothesis that coordination during rhythmic movements is mediated by the control of specific time intervals. In Experiment 1, an auditory metronome was used to indicate a set of timing patterns in which a 1-s interval was divided into 2 subintervals. Performance, measured in terms of the deviation from the target patterns and variability, was similar under conditions in which the finger taps were made with 1 hand or alternated between the 2 hands. In Experiment 2, the modality of the metronome (auditory or visual) was found to influence the manner in which the produced intervals deviated from the target patterns. These results challenge the notion that bimanual coordination emerges from coupling constraints intrinsic to the 2-hand system. They are in accord with a framework that emphasizes the control of specific time intervals to form a series of well-defined motor events.     

Changes in movement variables associated with transient overshoot of the final position

Transient overshoot (TO), which is assessed as the distance between the movement amplitude and the final position, was measured in a series of rapid, discrete elbow flexion movements performed under different distance and loading conditions by 7 participants. A positive relationship was found between kinematic variables (peak velocity, peak acceleration and deceleration, and the symmetry ratio) and the magnitude of TO, particularly in short movements performed against a light load. The relationships between TO and electromyographic (EMG) variables were low and mainly insignificant. Thus, TO contributes to the variability of rapid, discrete movements and therefore should be taken into account as an additional parameter in studies of the scaling of movement variables with movement mechanical conditions. TO could also represent a consequence of mechanical properties of the single-joint system rather than an independently programmed primary submovement.     

Changes in Movement Variables Associated With Transient Overshoot of the Final Position

Transient overshoot (TO), which is assessed as the distance between the movement amplitude and the final position, was measured in a series of rapid, discrete elbow flexion movements performed under different distance and loading conditions by 7 participants. A positive relationship was found between kinematic variables (peak velocity, peak acceleration and deceleration, and the symmetry ratio) and the magnitude of TO, particularly in short movements performed against a light load. The relationships between TO and electromyographic (EMG) variables were low and mainly insignificant. Thus, TO contributes to the variability of rapid, discrete movements and therefore should be taken into account as an additional parameter in studies of the scaling of movement variables with movement mechanical conditions. TO could also represent a consequence of mechanical properties of the single-joint system rather than an independently programmed primary submovement.     

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.