Kinetic attraction during bimanual coordination

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

Kinetic Attraction During Bimanual Coordination

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

Generation of Bimanual Trajectories of Disparate Eccentricity: Levels of Interference and Spontaneous Changes Over Practice

The authors examined intermanual interactions of 2 hands that were required to concurrently follow trajectories that differed in eccentricity. Ten healthy participants attempted to learn to trace 2 figures, a circle and an ellipse, with bilaterally isochronous (1:1) timing demands. Initial unimanual trials were followed by bilateral practice comprising 750 movement cycles. Two objectives were addressed: The authors' primary aim was to determine if kinematic interlimb interference is evident independent of spatial and temporal interference and to observe the potential practice-related changes in the nature of that interference. That test was afforded by participants' natural tendency to draw a circle with a relatively constant tangential velocity and an ellipse with a systematically varying velocity. A second aim was to observe the nature of spontaneous changes in the performance of each individual effector, and in the relationship between effectors, across practice. Those objectives were specifically addressed in a context in which augmented feedback was not available to direct the learners' attention to a particular feature of performance. The results suggested that interlimb assimilation of spatial features is the primary source of interference for that task and that apparent effects at the kinematic level are the secondary, indirect product of spatial coupling. Those results were found across blocks of practice. With respect to nondirected performance changes, substantially less improvement was evident in the performance of each individual effector than in the reduction of interlimb interference. Specifically, no practice-related changes in temporal variability or velocity bias, and minimal changes in trajectory smoothness, were evident in individual limbs. Conversely, significant reductions were observed in the variability of relative phase between limbs and in the magnitude of interlimb phase lag.     

Generation of bimanual trajectories of disparate eccentricity: levels of interference and spontaneous changes over practice

The authors examined intermanual interactions of 2 hands that were required to concurrently follow trajectories that differed in eccentricity. Ten healthy participants attempted to learn to trace 2 figures, a circle and an ellipse, with bilaterally isochronous (1:1) timing demands. Initial unimanual trials were followed by bilateral practice comprising 750 movement cycles. Two objectives were addressed: The authors' primary aim was to determine if kinematic interlimb interference is evident independent of spatial and temporal interference and to observe the potential practice-related changes in the nature of that interference. That test was afforded by participants' natural tendency to draw a circle with a relatively constant tangential velocity and an ellipse with a systematically varying velocity. A second aim was to observe the nature of spontaneous changes in the performance of each individual effector, and in the relationship between effectors, across practice. Those objectives were specifically addressed in a context in which augmented feedback was not available to direct the learners' attention to a particular feature of performance. The results suggested that interlimb assimilation of spatial features is the primary source of interference for that task and that apparent effects at the kinematic level are the secondary, indirect product of spatial coupling. Those results were found across blocks of practice. With respect to nondirected performance changes, substantially less improvement was evident in the performance of each individual effector than in the reduction of interlimb interference. Specifically, no practice-related changes in temporal variability or velocity bias, and minimal changes in trajectory smoothness, were evident in individual limbs. Conversely, significant reductions were observed in the variability of relative phase between limbs and in the magnitude of interlimb phase lag.     

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.     

Coordination Between Arm and Leg Movements During Locomotion

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

Coordination between arm and leg movements during locomotion

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

Effects of velocity and limb loading on the coordination between limb movements during walking

The authors investigated the effects of velocity (increasing from 0.5 to 5.0 km/hr in steps of 0.5 km/hr) and limb loading on the coordination between arm and leg movements during treadmill walking in 7 participants. Both the consistency of the individual limb movements and the stability of their coordination increased with increasing velocity; the frequency coordination between arm and leg movements was 2:1 at the lower velocities and 1:1 at the higher velocities. The mass manipulation affected the individual limb movements but not their coordination, indicating that a stable walking pattern was preserved. The results differed qualitatively from those obtained in studies on bimanual interlimb coordination, implying that the dynamical principles identified therein are not readily applicable to locomotion.     

Proprioceptive feedback as a mediator in interlimb timing

Prior findings regarded as evidence for proprioceptive feedback as a mediator in interlimb timing can also be interpreted as evidence for motor outflow because they came from research that had subjects make voluntary movements, and such movements allow for both feedback and outflow to operate. The present study was designed to resolve this controversy by determining if these findings could be replicated with passive movements which allow for feedback, but not outflow, to operate. The interlimb timing task studied was one where subjects made the timing response with their right hand while moving their left arm during the 1.5-sec interval to be timed. Three groups of 16 male college students performed 50 trials of the right-hand response with knowledge of results, under one of three left-arm conditions: (a) passive movement, (b) voluntary movement, and (c) no movement. The results indicated that the findings were replicated with passive movements and this was interpreted as support for the involvement of proprioceptive feedback in interlimb timing.     

Enhanced arm swing alters interlimb coordination during overground walking in individuals with traumatic brain injury

The current study investigated interlimb coordination in individuals with traumatic brain injury (TBI) during overground walking. The study involved 10 participants with coordination, balance, and gait abnormalities post-TBI, as well as 10 sex- and age-matched healthy control individuals. Participants walked 12 m under two experimental conditions: 1) at self-selected comfortable walking speeds; and 2) with instructions to increase the amplitude and out-of-phase coordination of arm swinging. The gait was assessed with a set of spatiotemporal and kinematic parameters including the gait velocity, step length and width, double support time, lateral displacement of the center of mass, the amplitude of horizontal trunk rotation, and angular motions at shoulder and hip joints in sagittal plane. Interlimb coordination (coupling) was analyzed as the relative phase angles between the left and right shoulders, hips, and contralateral shoulders and hips, with an ideal out-of-phase coupling of 180° and ideal in-phase coupling of 0°. The TBI group showed much less interlimb coupling of the above pairs of joint motions than the control group. When participants were required to increase and synchronize arm swinging, coupling between shoulder and hip motions was significantly improved in both groups. Enhanced arm swinging was associated with greater hip and shoulder motion amplitudes, and greater step length. No other significant changes in spatiotemporal or kinematic gait characteristics were found in either group. The results suggest that arm swinging may be a gait parameter that, if controlled properly, can improve interlimb coordination during overground walking in patients with TBI.     

Dependence of Finger Flexion Force on the Posture of the Nonperforming Fingers During Key Pressing Tasks

In visuomotor tasks that involve accuracy demands, small directional changes in the trajectories have been taken as evidence of feedback-based error corrections. In the present study variability, or intermittency, in visuomanual tracking of sinusoidal targets was investigated. Two lines of analyses were pursued: First, the hypothesis that humans fundamentally act as intermittent servo-controllers was re-examined, probing the question of whether discontinuities in the movement trajectory directly imply intermittent control. Second, an alternative hypothesis was evaluated: that rhythmic tracking movements are generated by entrainment between the oscillations of the target and the actor, such that intermittency expresses the degree of stability. In 2 experiments, participants (N = 6 in each experiment) swung 1 of 2 different hand-held pendulums, tracking a rhythmic target that oscillated at different frequencies with a constant amplitude. In 1 line of analyses, the authors tested the intermittency hypothesis by using the typical kinematic error measures and spectral analysis. In a 2nd line, they examined relative phase and its variability, following analyses of rhythmic interlimb coordination. The results showed that visually guided corrective processes play a role, especially for slow movements. Intermittency, assessed as frequency and power components of the movement trajectory, was found to change as a function of both target frequency and the manipulandum's inertia. Support for entrainment was found in conditions in which task frequency was identical to or higher than the effector's eigenfrequency. The results suggest that it is the symmetry between task and effector that determines which behavioral regime is dominant.     

The Dynamics of Bimanual Circle Drawing

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

Bimanual circle drawing in children with spastic hemiparesis: effect of coupling modes on the performance of the impaired and unimpaired arms

The present study examined the effect of interlimb coupling on the performance of the impaired and unimpaired arm in children with spastic hemiparesis during bimanual circle drawing. The following questions were addressed: (1) does coupling positively influence the performance of the impaired arm compared to single-hand performance and (2) is such an effect dependent on mode of coordination (i.e., symmetric versus asymmetric). Twelve children with spastic hemiparesis produced circle drawings on a digitizer under different task conditions. Spatiotemporal characteristics and quality of movement of pen trajectories of the individual limbs as well as interlimb relative phase were analysed. Coupling in a symmetric coordination mode resulted in a decrease of temporal variability and an increase of smoothness of circle drawing movements in the impaired arm compared to single-handed performance. Coupling in an asymmetric coordination mode resulted in an increase of spatial and temporal variability in the unimpaired arm. It is concluded that coupling may enhance the performance of the impaired arm in children with spastic hemiparesis, but only during symmetric bimanual coordination. A possible underlying neural mechanism that might explain these findings is discussed.     

Hemispace Asymmetries and Laterality Effects in Arm Positioning

Hemispace asymmetries and laterality effects were examined on an arm positioning reproduction task. Sixteen male subjects were asked to reproduce both abductive and adductive positioning movements with the left or right arm within either the left or the right hemispace. Hemispace was manipulated using a 90 degrees head-rotation paradigm. A left hemispace advantage in positioning accuracy was predicted for both left and right arm movements on the grounds that the perceptual-motor control of positioning movements made in left hemispace is primarily mediated by the right hemisphere which is known to be advantageous for tasks which are spatial in nature (Heilman, Bowers, & Watson, 1984). No arm laterality effects were predicted to occur because the proximal musculature involved in the control of arm movements is innervated from both contralateral and ipsilateral cerebral hemispheres (Brinkman & Kuypers, 1973). Results showed that the predicted left hemispace advantage was evident for the right arm on the positioning variability measure alone, whereas it was absent for all other possible conditions on all error measures. Laterality (arm) effects were absent as predicted. The experiment also demonstrated a greater degradation of reproduction performance under the ′crossed" arm-hemispace conditions than under the ′uncrossed" conditions. A plausible explanation for the uncrossed advantage for the task is that under normal conditions, a single hemisphere is primarily responsible for both controlling the contralateral arm and directing attention to the contralateral hemispace, and consequently potential interhemispheric interference is minimized. A clear response bias effect in movement reproduction was also evident as a function of the direction of concurrent arm movement and head rotation. Arm movements made in the same direction as head rotation were systematically undershot in reproduction to a much greater degree than arm movements made in the opposite direction to head rotation.     

Wrist and arm movements of varying difficulties

Although a great deal of experimental attention has been directed at understanding Fitts' law, only a limited number of experiments have attempted to determine if performance differs across effectors for a given movement difficulty. In three experiments reciprocal wrist and arm movements were compared at IDs of 1.5, 3, 4.5 and 6. When absolute movement requirements and visual display were constant, participants' movement times and response characteristics for the arm and wrist were remarkably similar (Experiment 1). However, when amplitude for wrist movements was reduced to 8° and the gain (4×) for the visual display increased participants' movement time, defined on the basis of kinematic markers (movement onset − movement termination), was increasingly shorter relative to arm movements as movement difficulty was increased (Experiment 2). Experiment 3 where the arm was tested at 32° and 8° with the 8° movements provided the same gain (4×) that was used for the 8° wrist movements in Experiment 2, no advantage was observed for the arm at the shorter amplitude. The results are interpreted in terms of the advantages afforded by the increased gain of the visual display, which permitted the wrist, but not the arm, to more effectively preplan and/or correct ongoing movements to achieve the required accuracy demands. It was also noted that while the wrist was more effective during the actual movement production this was accompanied by an offsetting increase in dwell time which presumably is utilized to dissipate the forces accrued during movement production and plan the subsequent movement segment.     

Dynamic dominance persists during unsupported reaching

Previous studies examining lateralization of arm movements focused on supported movements in the horizontal plane, removing the effects of gravity. The authors hypothesized that interlimb differences in free reaching would be consistent with the differences shown during supported reaching. Kinematic and kinetic data were collected for the forearm and upper arm segments in a 3-direction reaching task. Results showed lateralization of coordination, reflected by initial movement direction and trajectory curvature. The nondominant arm showed increased initial direction errors, and path curvature associated with a timing deficit between elbow and shoulder peak torques. These coordination deficits did not disrupt final position accuracy. The authors conclude that nondominant arm coordination deficits are similar to those reported previously for horizontal plane movements.     

Interlimb coordination in prosthetic walking: effects of asymmetry and walking velocity

The present study focuses on interlimb coordination in walking with an above-knee prosthesis using concepts and tools of dynamical systems theory (DST). Prosthetic walkers are an interesting group to investigate from this theory because their locomotory system is inherently asymmetric, while, according to DST, coordinative stability may be expected to be reduced as a function of the asymmetry of the oscillating components. Furthermore, previous work on locomotion motivated from DST has shown that the stability of interlimb coordination increases with walking velocity, leading to the additional expectation that the anticipated destabilizing effect of the prosthesis-induced asymmetry may be diminished at higher walking velocities. To examine these expectations, an experiment was conducted aimed at comparing interlimb coordination during treadmill walking between seven participants with an above-knee prosthesis and seven controls across a range of walking velocities. The observed gait patterns were analyzed in terms of standard gait measures (i.e., absolute and relative swing, stance and step times) and interlimb coordination measures (i.e., relative phase and frequency locking). As expected, the asymmetry brought about by the prosthesis led to a decrease in the stability of the coordination between the legs as compared to the control group, while coordinative stability increased with increasing walking velocity in both groups in the absence of a significant interaction. In addition, the 2:1 frequency coordination between arm and leg movements that is generally observed in healthy walkers at low walking velocities was absent in the prosthetic walkers. Collectively, these results suggest that both stability and adaptability of coordination are reduced in prosthetic walkers but may be enhanced by training them to walk at higher velocities.     

Tactile suppression in goal-directed movement

Sharing numerous characteristics with suppression in the other senses, tactile suppression is a reliable phenomenon that accompanies movement. By investigating the simplest of movements (e.g., finger flexions), early research tried to explain the origins of the phenomenon in terms of motor command generation together with sensory reafference. Here, we review recent research that has delved into (naturalistic) goal-directed movements. In connection with goal-directed movement, tactile suppression is evident as a decrease in behavioural performance measured shortly prior to, and during, movement execution. It is also reflected in a consistent response bias highlighting the (perceptual) uncertainty of the movement. Goal-directed movement supports the forward model and establishes contextual influences as the defining influences on tactile suppression. Depending on the task at hand, people prioritize a certain percept during movement. Future research, we argue, should focus on studying naturalistic movements, or sequences of movements, that share a common meaning or goal.     

Attentional loads associated with interlimb interactions underlying rhythmic bimanual coordination

Studies of rhythmic bimanual coordination under dual-task conditions revealed (1) a dependence of secondary task performance on the stability of coordinative tasks, in that secondary task performance was better during in-phase than antiphase coordination, and (2) a shift in the mean relative phasing between the limbs compared to single-task conditions. The present study aimed to account for these phenomena by dissociating three qualitatively different interactions between the limbs that govern this motor behavior, related to movement planning, error correction, and interlimb reflex activity. The experiment probed the cognitive demands associated with each interlimb interaction by examining the attentional load under dual-task conditions, indexed by reaction times of the secondary task and kinematic changes in the coordinative tasks relative to single-task conditions. First, only in the condition that involved interlimb interactions at the level of movement planning reaction times were shorter for in-phase than for antiphase coordination, highlighting an intimate relation between movement planning and attentional processes. Second, under dual-task conditions a shift in the mean relative phase was observed relative to single-task conditions, but only for the interlimb interactions that depend directly on sensory feedback (error correction and interlimb reflex activity). These observations qualified the effects of attentional load reported in previous studies. Third, reaction times varied systematically over the movement cycle. These variations revealed a dynamical signature of the attentional load that differed between the three interlimb interactions.