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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.

Journal Article↗

Exploring interlimb constraints during bimanual graphic performance: effects of muscle grouping and direction.

Past studies on bimanual coordination have revealed a general preference to move the limbs in a symmetrical fashion, also denoted as the in-phase mode. Its counterpart, the asymmetrical or anti-phase mode, is performed with lower degrees of accuracy and stability. This ubiquitous tendency to activate the homologous muscle groups is referred to as the muscle grouping constraint (egocentric constraint). The present study confirmed the generalizability of this constraint across various coordination patterns, performed in the horizontal plane. In addition, evidence was generated that movement direction in extrinsic space also constrains bimanual coordination (allocentric constraint). Overall, the present observations suggest that direction is an important movement parameter that is encoded in the central nervous system and that is subject to interactions between the neural specifications of both limbs.

Adult↗

Handedness-related asymmetry in coupling strength in bimanual coordination: furthering theory and evidence.

The effects of handedness on bimanual isofrequency coordination (e.g., phase advance of the dominant limb) have been suggested to result from an asymmetry in interlimb coupling strength, with the non-dominant limb being more strongly influenced by the dominant limb than vice versa. A formalized version of this hypothesis was tested by examining the phase adjustments in both limbs in response to mechanical perturbation of the bimanual coordination pattern and during frequency-induced phase transitions, for both right- and left-handed participants. In both situations, the phase adaptations were made predominantly by the non-dominant limb in right-handers, whereas this effect failed to reach significance in left-handers. Thus, the asymmetry in coupling strength was less pronounced in the latter group. In addition, the degree of asymmetry depended on movement frequency. The observed asymmetry was discussed in relation to pertinent neurophysiological findings.

Adult↗

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.

Adolescent↗

Sensorimotor state of the contralateral leg affects ipsilateral muscle coordination of pedaling.

The objective of this study was to determine if independent central pattern generating elements controlling the legs in bipedal and unipedal locomotion is a viable theory for locomotor propulsion in humans. Coordinative coupling of the limbs could then be accomplished through mechanical interactions and ipsilateral feedback control rather than through central interlimb neural pathways. Pedaling was chosen as the locomotor task to study because interlimb mechanics can be significantly altered, as pedaling can be executed with the use of either one leg or two legs (cf. walking) and because the load on the limb can be well-controlled. Subjects pedaled a modified bicycle ergometer in a two-legged (bilateral) and a one-legged (unilateral) pedaling condition. The loading on the leg during unilateral pedaling was designed to be identical to the loading experienced by the leg during bilateral pedaling. This loading was achieved by having a trained human "motor" pedal along with the subject and exert on the opposite crank the torque that the subject's contralateral leg generated in bilateral pedaling. The human "motor" was successful at reproducing each subject's one-leg crank torque. The shape of the motor's torque trajectory was similar to that of subjects, and the amount of work done during extension and flexion was not significantly different. Thus the same muscle coordination pattern would allow subjects to pedal successfully in both the bilateral and unilateral conditions, and the afferent signals from the pedaling leg could be the same for both conditions. Although the overall work done by each leg did not change, an 86% decrease in retarding (negative) crank torque during limb flexion was measured in all 11 subjects during the unilateral condition. This corresponded to an increase in integrated electromyography of tibialis anterior (70%), rectus femoris (43%), and biceps femoris (59%) during flexion. Even given visual torque feedback in the unilateral condition, subjects still showed a 33% decrease in negative torque during flexion. These results are consistent with the existence of an inhibitory pathway from elements controlling extension onto contralateral flexion elements, with the pathway operating during two-legged pedaling but not during one-legged pedaling, in which case flexor activity increases. However, this centrally mediated coupling can be overcome with practice, as the human "motor" was able to effectively match the bilateral crank torque after a longer practice regimen. We conclude that the sensorimotor control of a unipedal task is affected by interlimb neural pathways. Thus a task performed unilaterally is not performed with the same muscle coordination utilized in a bipedal condition, even if such coordination would be equally effective in the execution of the unilateral task.

Adult↗

The integration of cortical and behavioural dynamics during initial learning of a motor task.

Here we test the hypothesis that frequency and topographically specific changes in the strength of functional cortico-cortical coupling occur during the acquisition of a completely new task. To this end we studied the behavioural and cortical dynamics of a bimanual multifrequency coordination pattern during which one hand moved at twice the frequency of the other hand. This pattern represents a noninherent assignment and necessitates training before appropriate interlimb decoupling takes place. Results showed that acquisition of the multifrequency task was associated with an improved behavioural output that matched specific changes in the electroencephalogram dynamics. In particular, practice of the coordination pattern was accompanied by a decrease in coherence between the primary sensorimotor regions, and over the midline area in the alpha and beta bands, respectively, along with an increase in functional interhemispheric coupling between the prefrontal areas in the gamma band. These data suggest that the strength of cortico-cortical connectivity is adaptively modified across regions and across frequencies during early learning as the functional couplings are created and optimized for the purpose of movement execution.

Cerebral Cortex↗

Symmetry, broken symmetry, and handedness in bimanual coordination dynamics.

The symmetrical dynamics of 1:1 rhythmic bimanual coordination may be specified by an order parameter equation involving the relative phase between rhythmic components, and an interlimb coupling which determines the relative attractiveness of in-phase and anti-phase patterns. Symmetry breaking of these dynamics can occur via the difference in the natural frequencies, delta omega, of the left and right rhythmic components, or by the intrinsic asymmetrical dynamics of the body. The latter is captured by additional terms that render the symmetrical coupling slightly anisotropic. A major prediction resulting from this step is that although delta omega = 0, as the frequency of coordination is increased, the asymmetrical coupling will increase and the symmetrical coupling will decrease. This results in a greater left-limb bias in left-handers and right-limb bias in right-handers. This "increased handedness" prediction was confirmed in an experiment in which 20 left-handed and 20 right-handed individuals performed 1:1 coordination with hand-held rigid pendulums. Manipulations of left and right pendulum lengths controlled delta omega, and the coupled frequency was determined by a metronome. Also confirmed was the prediction that the small shift in equilibria from in-phase and anti-phase due to the intrinsic asymmetry should be amplified in left-handers when delta omega > 0 and in right-handers when delta omega < 0. Further, the bias in left-handers was more consistent than the bias in right-handers, and a subgroup of right-handers was identified who performed similarly to left-handers. The coordination dynamics of functional asymmetry provides insights into the elementary synergy between the limbs, the dynamical mechanism that modulates it, and the nature of the asymmetry in left-handed and right-handed individuals.

Adult↗

Descending lumbosacral cord potentials (DLCP) evoked by stimulation of the median nerve.

In 22 normal human subjects, descending lumbosacral cord potentials (DLCP) were recorded intrathecally after stimulation of the median nerve at the elbow. The onset of DLCP is very short in latency (mean 12.1 ms) with a prominent sharp early positive peak (mean latency 13.7 ms) followed by a sharp negative peak (mean 17.6 ms). The amplitude of the first part of DLCP varied between 0.6 and 6.7 microV (mean 2.3 microV). The response was recorded most easily when the tip of intrathecal electrode was posterolaterally positioned. The threshold of the response was above or around the excitation threshold of the motor nerve fibers and it could not be produced by pure skin nerve stimulation. It resisted to subtetanic peripheral shocks. Mean peripheral conduction velocity responsible for the response was about 60.8 m/s. Some late and slower deflections appeared in many cases. It was concluded that the DLCP must have originated from the descending and very fast conducting propriospinal pathways located within the anterolateral funiculus which has an oligosynaptic anatomical organization. This response seemed to be the first direct evidence of interlimb reflex action between the arm and leg in man which is important in the coordination of movements and posture.

Arm↗

Interlimb reflexes evoked in human arm muscles by ankle displacement.

Interlimb reflexes evoked by ankle displacements were studied in arm muscles of 6 normal subjects. EMGs from grastrocnemius (G), tibialis anterior (TA), biceps brachii (BB), and triceps brachii (TB) were amplified, rectified and low-pass filtered before recording. Averaging and Wiener filtering were used to detect changes in tonic EMG activity evoked by dorsiflexing or plantarflexing displacements of the ankle. A consistent pattern of response was observed in all subjects. In the leg muscles, the responses to stretch were consistent with previous reports. In the arm muscles, the response of TB was dominant. Dorsiflexing displacements of the ankle evoked a small excitation followed by a more marked decrease in TB activity but had no effect on BB. In contrast, plantarflexing displacements of the ankle resulted in a large, early period of excitation followed by a decreased level of activity in TB. A similar but smaller pattern of activity was observed in BB. It is notable that the TB responses to displacement were sizable, often modulating the tonic EMG activity by as much as 80%. Interlimb reflexes evoked by ankle displacement were larger and of shorter latency than those evoked by cutaneous electrical stimulation of the foot reported previously. This suggests that proprioceptive afferents may have stronger and more direct interlimb reflex connections than cutaneous afferents and may therefore play an important role in the coordination of movement.

Adult↗

A higher-order mechanism overrules the automatic grip-load force constraint during bimanual asymmetrical movements.

The aim of the present study was to examine grip-load force regulation during unimanual and bimanual movements. Two protocols were included which manipulated the object's weight and covered distance. Results showed that grip-load ratio was adapted to the task requirements. During unimanual and bimanual symmetrical movements, an increased grip-load force ratio for long versus short amplitude movements as well as for light versus heavy weight movements was noted. These findings could be related to the observed movement speed variations associated with the tasks. During bimanual asymmetrical movements, the grip-load force ratio became comparable for both sides. When transporting different object's weights to constant distances, the grip-load force ratio of light weight movements decreased towards that of heavy weight movements. As movement speed was reduced, it indicates that grasping forces were adapted accordingly. When transporting constant object's weights to different distances, the grip-load force ratio of short amplitude movements increased towards that of long amplitude movements. Since movement speed was decreased, it suggests that a bimanual coordinative command overruled the automatic grip-load coupling. In conclusion, these data show that interlimb coupling induced a rescaling towards a common control structure, leading to similar grasping forces during bimanual movements with dissimilar actions.

Functional Laterality↗

Kinematic comparison of the leading and trailing fore- and hindlimbs at the canter.

The canter is a 3 beat asymmetrical gait with a difference in timing between left and right limbs. To evaluate intralimb asymmetry at the canter, a group of 24 Dutch Warmbloods was evaluated on a treadmill (7 m/s) using a modified CODA-3 optoelectronic gait analysis system. Thirteen horses cantered in the left lead ('leading limb' group) and 11 in the right lead ('trailing limb' group) during left forelimb recordings, while 11 horses were at the left and 13 were at the right lead during left hindlimb recordings. Kinematic differences between horses from the 'leading limb' and 'trailing limb' group were statistically evaluated at a significance level of P<0.05. Stride, stance and swing duration were similar between the 2 groups. The pelvis rotation, angle of maximal protraction and total range of maximal pro- and retraction were larger in the 'leading limb' group, while the scapula rotation, and the angle of maximal retraction were larger in the 'trailing limb' group. The elbow and hip joints were more flexed at impact, at maximal extension and at maximal flexion of the leading limb, whereas the stifle joint was more extended at impact. Furthermore, the leading tarsal joint was more maximally flexed in stance and swing phase, whereas the carpal joint was more flexed only in the swing phase of the leading limb. However, during the stance phase the maximal fetlock extension of the trailing fore- and hindlimbs were significantly larger. Apparently, horses move at the canter with a more protracted leading limb by more flexing the elbow, carpal, hip and tarsal joints. In the trailing limb, however, the scapula is more rotated, and the tarsal and fetlock joints are more loaded. In conclusion, the difference in interlimb timing between left and right limbs at canter also leads to an asymmetry in intralimb coordination of these limbs.

Analysis of Variance↗

Coping with systematic bias during bilateral movement.

The present studies examined the nature of kinematic interlimb interference during bilateral elbow movements of 1:1, 2:1 and 3:1 frequency ratios and the manner in which subjects cope with coordination bias. Analysis of movement trajectories in the first experiment indicated progressively greater angular velocity assimilation across 2:1 and 3:1 conditions. The desired temporal relationship was maintained by slowing or pausing the low-frequency movement at peak extension while the high-frequency arm produced intervening cycles. An increase in amplitude was also evident for concurrent, homologous cycles. Movement smoothness was emphasized and additional practice was provided in a second experiment. This resulted in dissociated peak angular velocity between limbs and eliminated hesitations and amplitude effects. Bias was still evident, however, as an intermittent approach toward a 1:1 ratio within each cycle. This systematic tendency was somewhat greater at the lower of two absolute frequency combinations but was not influenced by the role of each arm in producing the higher or lower frequency movement. The findings from the first experiment suggest that subjects initially accommodate interlimb kinematic assimilation, while producing the intended timing ratio, by intermittently slowing or pausing the lower-frequency movement. This attenuates the need for bilaterally-disparate movement parameters and provides additional time for organizing residual kinematic differences, perhaps reducing "transient coupling." Evidence from the second experiment indicates that subtle relative motion preferences are still evident following sufficient practice to perform the movements smoothly. The within-cycle locations of the points of greatest interlimb bias for the 2:1 rhythms were positively displaced from those previously observed for 1:1 oscillations. The persistent coordination tendencies noted in both experiments perhaps reflect an assimilation/compensation cycle and constitute one potential source of the systematic error that often emerges during the acquisition of complex skills.

Adult↗

Modulations of interlimb and intralimb cutaneous reflexes during simultaneous arm and leg cycling in humans.

OBJECTIVE: We investigated to what extent intralimb and interlimb cutaneous reflexes are altered while simultaneously performing arm and leg cycling (AL cycling) under different kinematic and postural conditions. METHODS: Eleven subjects performed AL cycling under conditions in which the arm and leg crank ipsilateral to the stimulation side were moved synchronously (in-phase cycling) or asynchronously (anti-phase cycling) while sitting or standing. Cutaneous reflexes following superficial radial or superficial peroneal nerve stimulation (2.0-2.5 times radiating threshold, 5 pulses at 333 Hz) were recorded at 4 different pedal positions from 12 muscles in the upper and lower limbs. Cutaneous reflexes with a peak latency of 80-120 ms were then analyzed. RESULTS: The magnitude of interlimb and intralimb cutaneous reflexes in the arm and leg muscles was significantly modulated depending on the crank position for the relevant limb (phase-dependent modulation). A significant correlation between the magnitude of the cutaneous reflex and background EMG was observed in the majority of muscles during static contraction, but not during AL cycling (task-dependent modulation). No significant difference was found in comparisons of the magnitude of intralimb and interlimb cutaneous reflexes obtained during in- and anti-phase AL cycling. Qualitatively, the same results were obtained during AL cycling while sitting or standing. In addition, the modulation of cutaneous reflexes in arm muscles was identical among in-phase, anti-phase and isolated arm cycling. Results were the same for leg muscles. CONCLUSIONS: Cutaneous reflexes in arm muscles are little influenced by rhythmic movement of the legs and vice versa during AL cycling. It is likely that neural components that control interlimb reflexes are loosely coupled during AL cycling while sitting or standing. SIGNIFICANCE: Our results provide a better understanding of the coordination between the upper and lower limbs during rhythmic movement.

Adult↗

Spatial conceptual influences on the coordination of bimanual actions: when a dual task becomes a single task.

When the left and right hands produce 2 different rhythms simultaneously, coordination of the hands is difficult unless the rhythms can be integrated into a unified temporal pattern. In the present study, the authors investigated whether a similar account can be applied to the spatial domain. Participants (N = 8) produced a movement trajectory of semicircular form in single-limb and bimanual conditions. In the bimanual tasks, 1 limb moved above the other in the frontal plane. Bimanual unified tasks were constructed so that the spatial paths to be produced by the 2 limbs could be easily conceptualized as parts of a unified circle pattern. Bimanual distinct tasks availed a less obvious spatial pattern that would unify the 2 tasks, despite similar demands placed on the coordination dynamics in the 2 cases (e.g., the phase relations). The authors conclude that a dual task becomes a single task, and interlimb interference is reduced, when the spatial patterns produced by the 2 hands form a geometric arrangement that can be conceptualized as a unified representation.

Adolescent↗

Influence of task dynamics on the organization of interlimb responses accompanying standing human leg flexion movements.

The organization of electromyographic (EMG) and kinetic (ground reaction forces) responses in postural (single stance limb) and primary task (flexing limb) components was examined in standing human subjects during single leg flexion movements over a range of speeds. Interlimb responses occurred coincidently prior to unloading and were scaled as a function of movement speed, suggesting a centrally organized, synchronous mode of coordination which reflected the mechanical constraints related to stance support function.

Adult↗

Interlimb transfer of visuomotor rotations: independence of direction and final position information.

Previous findings from our laboratory support the idea that the dominant arm is more proficient than the non-dominant arm in coordinating intersegmental dynamics for specifying trajectory direction and shape during multijoint reaching movements. We also showed that adaptation of right and left arms to novel visuomotor rotations was equivalent, suggesting that this process occurs upstream to processes that distinguish dominant and non-dominant arm performance. Because of this, we speculate that such visuomotor adaptations might transfer to subsequent performance during adaptation with the other arm. We now examine whether opposite arm training to novel visuomotor rotations transfers to affect adaptation using the right and left arms. Two subject groups, RL and LR, each comprising seven right-handed subjects, adapted to a 30 degrees counterclockwise rotation in the visual display during a center-out reaching task performed in eight directions. Each group first adapted using either the right (RL) or left (LR) arm, followed by opposite arm adaptation. In order to assess transfer, we compared the same side arm movements (either right or left) following opposite arm adaptation to those performed prior to opposite arm adaptation. Our findings indicate unambiguous transfer of learning across the arms. Different features of movement transferred in different directions: Opposite arm training improved the initial direction of right arm movements under the rotated visual condition, whereas opposite arm training improved the final position accuracy, but not the direction of left arm movements. These findings confirm that transfer of training was not due to a general cognitive strategy, since such an effect should influence either hand equally. These findings support the hypothesis that each arm controller has access to information learned during opposite arm training. We suggest that each controller uses this information differently, depending on its proficiency for specifying particular features of movement. We discuss evidence that these two aspects of control are differentially mediated by the right and left cerebral hemispheres.

Adolescent↗

Visual perception of the relative phasing of human limb movements.

Studies of bimanual coordination have found that only two stable relative phases (0 degree and 180 degrees) are produced when a participant rhythmically moves two joints in different limbs at the same frequency. Increasing the frequency of oscillation causes an increase in relative phase variability in both of these phase modes. However, relative phasing at 180 degrees is more variable than relative phasing at 0 degree, and when the frequency of oscillation reaches a critical frequency, a transition to 0 degree occurs. These results have been replicated when 2 people have coordinated their respective limb movements using vision. This inspired us to investigate the visual perception of relative phase. In Experiment 1, recordings of human interlimb oscillations exhibiting different frequencies, mean relative phases, and different amounts of phase variability were used to generate computer displays of spheres oscillating either side to side in a frontoparallel plane or in depth. Participants judged the stability of relative phase. Judgments covaried with phase variability only when the mean phase was 0 degree or 180 degrees. Otherwise, judgments covaried with mean relative phase, even after extensive instruction and demonstration. In Experiment 2, mean relative phase and phase variability were manipulated independently via simulations, and participants were trained to perceive phase variability in testing sessions in which mean phase was held constant. The results of Experiment 1 were replicated. The HKB model was fitted to mean judgment standard deviations.

Adolescent↗

The coordination of limb movements with different kinematic patterns.

The principles underlying the coordination of limb movements with different spatiotemporal features were explored. After an initial training session in which the same unidirectional movement had to be performed with both upper limbs, subjects attempted to coordinate two different movements in a second session, i.e., the learned unidirectional movement in the left limb and a new double reversal movement in the right limb. The findings uncovered a wide variety in patterns of interlimb dependence among and within subjects, going from a high degree of dependence to relative independence. The relationship between limbs was studied by means of a detailed analysis of the displacement and acceleration patterns and the electromyographic activity of the major muscles involved. The general underlying principle that appeared to account for the diversity in movement organization was this: higher independence between limb movements is achieved when subjects initiate the movements to be coordinated successively. This asynchrony in movement onset can possibly be viewed as an attempt to safeguard against interference.

Acceleration↗