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PubMed2002

Handedness: dominant arm advantages in control of limb dynamics.

Recent findings from our laboratory suggest that a major factor distinguishing dominant from nondominant arm performance is the ability by which the effects of intersegmental dynamics are controlled by the CNS. These studies indicated that the dominant arm reliably used more torque-efficient patterns for movements made with similar speeds and accuracy than nondominant arm movements. Whereas, nondominant hand-path curvatures systematically varied with the amplitude of the interaction torques transferred between the segments of the moving limb, dominant hand-path curvatures did not. However, our previous studies did not distinguish whether dominant arm coordination advantages emerged from more effective control of dynamic factors or were simply a secondary effect of planning different kinematics. The purpose of this study was to further investigate interlimb differences in coordination through analysis of inverse dynamics and electromyography recorded during the performance of reaching movements. By controlling the amplitude of intersegmental dynamics in the current study, we were able to assess whether systematic differences in torque-efficiency exist, even when differences in hand-path shape were minimal. Subject's arms were supported in the horizontal plane by a frictionless air-jet system and were constrained to movements about the shoulder and elbow joints. Two targets were designed, such that the interaction torques elicited at the elbow were either large or small. Our results showed that the former produced large differences in hand-path curvature, whereas the latter did not. Additionally, the movements with small differences in hand-path kinematics showed substantial differences in torque patterns and corresponding EMG profiles which implied a more torque-efficient strategy for the dominant arm. In view of these findings we propose that distinct neural control mechanisms are employed for dominant and nondominant arm movements.

Adult↗

PubMed2001

Evidence for a dynamic-dominance hypothesis of handedness.

Handedness is a prominent behavioral phenomenon that emerges from asymmetrical neural organization of human motor systems. However, the aspects of motor performance that correspond to handedness remain largely undetermined. A recent study examining interlimb differences in coordination of reaching demonstrated dominant arm advantages in controlling limb segment inertial dynamics (Sainburg and Kalakanis 2000). Based on these findings, I now propose the dynamic-dominance hypothesis, which states that the essential factor that distinguishes dominant from nondominant arm performance is the facility governing the control of limb dynamics. The purpose of this study is to test two predictions of this hypothesis: 1) adaptation to novel intersegmental dynamics, requiring the development of new dynamic transforms, should be more effective for the dominant arm; 2) there should be no difference in adapting to visuomotor rotations performed with the dominant as compared with the nondominant arm. The latter prediction is based on the idea that visual information about target position is translated into an internal reference frame prior to transformation of the movement plan into dynamic properties, which reflect the forces required to produce movement. To test these predictions, dominant arm adaptation is compared to nondominant arm adaptation during exposure to novel inertial loads and to novel visuomotor rotations. The results indicate substantial interlimb differences in adaptation to novel inertial dynamics, but equivalent adaptation to novel visuomotor rotations. Inverse dynamic analysis revealed better coordination of dominant arm muscle torques across both shoulder and elbow joints, as compared with nondominant arm muscle torques. As a result, dominant arm movements were produced with a fraction of the mean squared muscle torque computed for nondominant arm movements made at similar speeds. These results support the dynamic-dominance hypothesis, indicating that interlimb asymmetries in control arise downstream to visuomotor transformations, when dynamic variables that correspond to the forces required for motion are specified.

Adaptation, Physiological↗

PubMed1996

Coordination dynamics of trajectory formation.

The present study aims to understand the neurally based coordination dynamics (multistability, loss of stability, transitions, etc.) of trajectory formation in a simple task. Six subjects produced two spatial patterns of coordination in the xy plane by alternating the abduction-adduction and flexion-extension motions of their right index finger. Each pattern was characterized by a unique temporal ratio between the x and y directions of motion: (1) a figure zero, a 1:1 temporal pattern; and (2) a figure eight, a 2:1 temporal pattern. The patterns were produced rhythmically and movement frequency was scaled across ten frequency plateaus, with ten cycles of motion per step. As movement frequency increased, switching from a figure eight to a figure zero was observed at critical cycling frequencies. The switch from pattern (2) to pattern (1) was identified in the spatial trajectory and power spectra of x(t) and y(t). En route to the transition, enhancement of fluctuations was observed in the Fourier amplitudes of x(t) and y(t), specifically at f0 (the metronome frequency) and 2f0 (the first harmonic of f0). Interestingly, there was no difference in the spatial variability of the two patterns. Overall, the data demonstrate that spatial patterns of coordination can be characterized in terms of the temporal relationship between the spatial components of the trajectory itself. We discuss the experimental findings in relation to other end-point planning and multijoint control strategies, as well as the much more general problem of temporal synchronization in many interlimb and intralimb coordination tasks.

Adult↗

PubMed2002

Limb and gender differences in the development of coordination in early infancy.

Young infants produce a variety of spontaneous arm and leg movements in the first few months of life. Coordination of leg joints has been extensively investigated, whereas arm joint coordination has mainly been investigated in the sitting position in the context of early reaching and grasping. The current study investigated arm and leg joint coordination of movements produced in the supine position in 10 fullterm infants aged 6, 12 and 18 weeks. Longitudinal comparisons within limbs (intralimb) as well as between limbs (interlimb, ipsilateral and contralateral) were made as well as an exploration of differences in the development for boys and girls. The relationship between the joint angles was examined by measuring pair-wise cross-correlation functions for the angular displacement curves of the leg (hip, knee and ankle) and arm (shoulder, elbow and wrist) joints of both the right and left side. Both the arms and legs were found to follow a similar pattern of intralimb coordination, although the leg joints were more tightly coupled than the arm joints, particularly the proximal with the middle joint. In support of earlier findings, differences in the development of the right and left side were identified. In addition, gender differences in joint coordination were found for both intralimb and interlimb coordination. This contrasts with the view that gender differences in motor development may be primarily a result of environmental influences.

Age Factors↗

PubMed1996

Self-Optimization of Walking in Nondisabled Children and Children With Spastic Hemiplegic Cerebral Palsy.

Children voluntarily adopt a frequency and movement pattern for walking. The force-driven harmonic oscillator (FDHO) model was used in this study for accurate prediction of the preferred walking frequency of nondisabled children and children with spastic hemiplegic cerebral palsy. Four potential optimality criteria with which the preferred walking pattern was forced to comply were examined: minimization of physiological costs, maximization of mechanical energy conservation, minimization of asymmetry in lower limb movements and minimization of variability of interlimb and intralimb coordination. Age and gender-matched nondisabled children (n = 6) and children with spastic hemiplegic cerebral palsy (n = 6) were tested under six frequency conditions of walking at a constant speed on a treadmill. For the nondisabled children, the results indicated that their preferred walking frequency could be accurately predicted by the FDHO model. They freely adopted a walking pattern that minimized physiological costs, asymmetry, and variability of inter- and intralimb coordination. For the children with spastic hemiplegic cerebral palsy, the prediction of preferred overground walking frequency required that the FDHO model be modified to account for muscle mass and leg length discrepancies between limbs and increased stiffness. Most of the children achieved the same optimality goals as the nondisabled when walking at the preferred frequency. However, the children were found to use different mechanisms to attain these goals: for example, a steeper increase observed in physiological cost at higher frequencies; a lowered center of gravity of the body, which allowed for angular symmetry; and greater variability of between-joint coordination in the nonaffected limb and less variability in the affected limb.

Journal Article↗

PubMed2005

Performance of locomotion and foot grasping following a unilateral thoracic corticospinal tract lesion in monkeys (Macaca mulatta).

Six adult monkeys (Macaca mulatta) received a unilateral lesion of the lateral corticospinal tract (CST) in the thoracic spinal cord. Prior to surgery, the animals were trained to perform quadrupedal stepping on a treadmill, and item retrieval with the foot. Whole body kinematics and electromyogram (EMG) recordings were made prior to, and at regular intervals over a period of 12 weeks after the CST lesion. After 1 week of recovery, all monkeys were able to walk unaided quadrupedally on the treadmill. The animals, however, dragged the hindpaw ipsilateral to the lesion along the treadmill belt during the swing phase and showed a significant reorganization of the spatiotemporal pattern of hindlimb (HL) and forelimb (FL) displacements. The inability to appropriately trigger the swing phase resulted in an increase in the cycle duration and stride length of both HLs. The stance duration decreased in the ipsilateral HL, and increased in the contralateral HL and both FLs. Consequently, there was a dramatic disruption of interlimb and intralimb coupling that was reflected in the limb kinematic and EMG patterns. The CST lesion completely abolished the ability of the monkeys to retrieve items with the foot ipsilateral to the lesion and significantly disrupted the level of performance of the contralateral HL during the first 2 weeks post-lesion. Interestingly, selected HL muscles remained almost quiescent when the monkeys attempted to retrieve items, but were unsuccessful with the affected foot at 1 week post-lesion, whereas the capacity to activate the same muscles was preserved, although reduced, during stepping. Spatial and temporal parameters of gait, kinematics, and EMG patterns recorded during locomotion generally converged toward control values over time, but significant differences persisted up to 12 weeks post-lesion. Although some control was recovered over the distal foot musculature, fine foot grasping remained significantly impaired at the end of the testing period. These findings demonstrate that the CST pathway from the brain normally makes an important contribution to interlimb and intralimb coordination during basic locomotion, and to muscle activation to produce dexterous foot digit movements in the monkey. Furthermore, the present study indicates that the primate has the ability to rapidly accommodate locomotor performance, and to a lesser degree fine foot motor skills, to a reduction in supraspinal control. Identification of the neural substrates mediating the rapid recovery of motor function following injury to the primate spinal cord could provide insight into developing repair strategies to augment functional recovery from neuromotor impairments.

Animals↗

PubMed2001

Hemiparetic gait parameters in overground versus treadmill walking.

OBJECTIVE: Hemiparetic gait is characterized by high stride-cycle variability, diminished stance time, single-limb stance time, and stance/swing ratio in the paretic limb. Recent studies suggest treadmill (TM) training may improve the motor control underlying these variables, but supporting evidence is sparse. METHODS: This study compared gait patterns of untrained chronic hemiparetic stroke patients (n = 18; mean, 39.5 months poststroke) during overground (OG) and TM walking at matched velocities. Variables included relative stance time, relative single-limb stance time, stance/swing ratio, peak force, and impulse. Within-subject variability of these measures (CV) was used to assess gait pattern stability. RESULTS: OG and TM cycle durations were similar, but CVs differed (TM < OG, p < 0.05). In the paretic limb, differences were seen in relative stance time, relative single-limb stance time, and stance/swing ratio, respectively (TM > OG, p < 0.05). These variables decreased in the nonparetic limb during TM walking (p < 0.05 for all). Improved interlimb symmetry and coordination were evidenced by decreased between-limb differences and improved relative temporal phasing, respectively, in the TM condition (p < 0.05). CONCLUSIONS: Collectively, these results demonstrate that the TM induces an immediate alteration toward a more consistent and symmetric gait pattern. Further investigation is needed to determine whether TM training leads to motor relearning and neuroplasticity in chronic hemiparetic subjects.

Chronic Disease↗

PubMed2001

Proprioceptive regulation of interlimb behavior: interference between passive movement and active coordination dynamics.

The coordination of homolateral effectors (right arm/right leg) according to the in-phase or anti-phase mode was perturbed through passive movement of a third segment (left arm or left leg) imposed by the experimenter. The manipulated parameters of the passive segment were frequency and amplitude along with their degree of scaling. Results showed that passive movement degraded anti-phase patterns more than in-phase patterns. Furthermore, the anti-phase mode deteriorated profoundly during frequency manipulation, but scaling did not induce additional effects, whereas a linear association was observed between anti-phase deterioration and amplitude manipulation. Together, these data indicate that passive movement disturbed the coordination dynamics of an actively performed task. The fact that interference depended on the manipulated parameter suggests a distinction in the degree of intrusiveness of the irrelevant afferent information induced by the passive limb. It is concluded that sensory discrimination between irrelevant and relevant input is critical in performing a coordinated task adequately under perturbed conditions.

Adult↗

PubMed2000

Contralateral movement and extensor force generation alter flexion phase muscle coordination in pedaling.

The importance of bilateral sensorimotor signals in coordination of locomotion has been demonstrated in animals but is difficult to ascertain in humans due to confounding effects of mechanical transmission of forces between the legs (i.e., mechanical interleg coupling). In a previous pedaling study, by eliminating mechanical interleg coupling, we showed that muscle coordination of a unipedal task can be shaped by interlimb sensorimotor pathways. Interlimb neural pathways were shown to alter pedaling coordination as subjects pedaling unilaterally exhibited increased flexion-phase muscle activity compared with bilateral pedaling even though the task mechanics performed by the pedaling leg(s) in the unilateral and bilateral pedaling tasks were identical. To further examine the relationship between contralateral sensorimotor state and ipsilateral flexion-phase muscle coordination during pedaling, subjects in this study pedaled with one leg while the contralateral leg either generated an extensor force or relaxed as a servomotor either held that leg stationary or moved it in antiphase with the pedaling leg. In the presence of contralateral extensor force generation, muscle activity in the pedaling leg during limb flexion was reduced. Integrated electromyographic activity of the pedaling-leg hamstring muscles (biceps femoris and semimembranosus) during flexion decreased by 25-30%, regardless of either the amplitude of force generated by the nonpedaling leg or whether the leg was stationary or moving. In contrast, rectus femoris and tibialis anterior activity during flexion decreased only when the contralateral leg generated high rhythmic force concomitant with leg movement. The results are consistent with a contralateral feedforward mechanism triggering flexion-phase hamstrings activity and a contralateral feedback mechanism modulating rectus femoris and tibialis anterior activity during flexion. Because only muscles that contribute to flexion as a secondary function were observed, it is impossible to know whether the modulatory effect also acts on primary, unifunctional, limb flexors or is specific to multifunctional muscles contributing to flexion. The influence of contralateral extensor-phase sensorimotor signals on ipsilateral flexion may reflect bilateral coupling of gain control mechanisms. More generally, these interlimb neural mechanisms may coordinate activity between muscles that perform antagonistic functions on opposite sides of the body. Because pedaling and walking share biomechanical and neuronal control features, these mechanisms may be operational in walking as well as pedaling.

Adult↗

PubMed1999

Dynamics of 1:2 Coordination: Generalizing Relative Phase to n:m Rhythms.

Interlimb rhythmic movements can be modeled as coupled oscillators, with stable performance characterized by the relative phase between the limbs. In the present study, that modeling strategy, verified previously for 1:1 coordination, was generalized to 1:2 coordination with a view to n:m coordination. The generalized model predicted interactions between coordination (specifically, 1:1 vs. 1:2) and the frequency asymmetry between the limbs determining mean relative phase and its variability. The predicted interactions were evaluated with bimanual 1:2 and 1:1 rhythmic tasks in which participants (N = 8) oscillated hand-held pendulums whose uncoupled frequencies could be adjusted so that different interlimb asymmetries were produced. The authors needed new analytic procedures to verify stable 1:2 coordination and to resolve stochastic and deterministic sources of variability in the component oscillations. The major expectations from the generalized model were confirmed, and the implications of additional but unpredicted findings for the modeling of multifrequency behavior are discussed.

Journal Article↗

PubMed2004

Interlimb influences on paretic leg function in poststroke hemiparesis.

After stroke, paretic leg motor impairment is typically viewed as a unilateral control deficit. However, much of the neural circuitry controlling normal leg function is organized bilaterally to produce coordinated, task-specific activity in the two legs. Thus, as a result of contralesional neural control processes, paretic leg motor pattern generation may be substantially influenced by the nonparetic leg sensorimotor state during bilateral lower limb tasks. Accordingly, we investigated whether different paretic leg motor patterns are observed during mechanically equivalent bilateral and unilateral tasks and, if so, whether nonparetic leg participation improved or exacerbated paretic leg coordination deficits. A pedaling apparatus that mechanically decoupled the legs was used to present subjects with increasingly complex bi- and unilateral motor tasks: isometric force generation, discrete movement, and pedaling. Bilateral electromyographic and pedal force data were collected from 21 persons with chronic poststroke hemiparesis and 11 similarly aged controls. During isometric force generation and discrete movements, nonparetic leg influences on paretic leg coordination were similar and not markedly different from interlimb influences in controls. In bilateral pedaling, however, interlimb influences differed from controls such that paretic leg coordination deficits were exacerbated. During pedaling movements, the suppression of interlimb influences similar to those observed in isometric and discrete movement may occur in controls but may be disrupted in hemiparesis. We suggest that the coupling of pattern generation between the two legs may result in greater, albeit more impaired, paretic leg motor output during bilateral pedaling than during unilateral pedaling.

Aged↗

PubMed2000

Spontaneous kicking behavior in infants: age-related effects of unilateral weighting.

The age-related effects of unilateral weighting on spontaneously generated kicks in 18 healthy, full-term infants were investigated. The main question was whether infants during the first half-year after birth reveal changes in how they adjust to unilateral weighting. At 6 weeks, infants reduced the frequency of the weighted leg and increased that of the unweighted leg whereas at 12 weeks the frequency of kicking increased in both legs. At both ages, unilateral weighting also resulted in differences on a number of kinematic parameters. By 18 and 26 weeks, such frequency and kinematic effects were no longer present. With regard to interlimb couplings, a clear pattern of bilateral coordination was only present at 26 weeks; these results suggest that the effects of unilateral weighting are not directly related to the tightness of interlimb couplings. The implications of these age-related differences for understanding developmental changes in the control of leg movements are discussed. It is suggested that the infants' improved ability to act in a task-specific manner as well as nonlinear changes in the musculo-skeletal system and fine-tuning processes at a neural level might be factors of importance.

Age Factors↗

PubMed1996

Interlimb co-ordination of force and movement-related cortical potentials.

The question we asked in this study was how the cerebral hemispheres are coordinated when the two hands simultaneously generate force at the same level. Interlimb co-ordination of contractions at 20% of the maximal level was investigated during bilateral (BL) and unilateral (UL) handgrip in eight male right-handed subjects. The accuracy as determined by the force error was larger during BL than during UL for the left hand only. The movement-related cortical potential (MRCP) of C3 (left precentral cortex) was studied during UL right and BL handgrip and the C4 (right precentral cortex) during UL left and BL handgrip. At the phase of the readiness potential (-1.0 to -0.6 s prior to the force onset), similar levels of correlation in the C3 (C4) MRCP amplitudes (P < 0.05) between UL right (UL left) and BL handgrip were observed. At the phase of the negative slope (-0.6 to -0.2 s) and the motor potential (-0.2 to -0.05 s), C3 revealed almost the same levels of correlation as for the readiness potential, whereas in C4 a marked decrease was noted. Accordingly, from this study the force error and movement-related cortical potentials would indicate that the left dominant hemisphere is specialized not only for unilateral contractions but also for bilateral contractions.

Adult↗

PubMed2006

Intentional switches between bimanual coordination patterns are primarily effectuated by the nondominant hand.

Based on indications that hand dominance is characterized by asymmetrical interlimb coupling strength (with the dominant hand exerting stronger influences on the nondominant hand than vice versa), intentional switches between rhythmic bimanual coordination patterns were predicted to be mediated primarily by phase adaptations in the movements of the nondominant hand. This hypothesis was supported for both right-handed and left-handed participants who performed voluntary switches from in-phase to antiphase coordination and vice versa, at four different frequencies. In accordance with previous indications that handedness is expressed less consistently in left-handers, the asymmetry between the hands was less pronounced in left-handed than in right-handed participants. The asymmetry was smaller for switches from in-phase to antiphase coordination (i.e., in the direction opposite to spontaneous transitions) than for switches in the reverse direction, suggesting that (the expression of) the handedness-related asymmetry in coupling strength was weakened by intentional processes associated with these switches.

Adolescent↗

PubMed2002

Anisotropic tracking: evidence for automatic synergy formation in a bimanual task.

Investigation of interlimb synergy has become synonymous with the study of coordination dynamics and is largely confined to periodic movement. Based on a computational approach this paper demonstrates a method of investigating the formation of a novel synergy in the context of stochastic, spatially asymmetric movements. Nine right-handed participants performed a two degrees of freedom (2D) "etch-a-sketch" tracking task where the right hand controlled the horizontal position of the response cursor on the display while the left hand controlled the vertical position. In a pre-practice 2D tracking task, measures of phase lag between the irregularly moving target and the response showed that participants controlled left and right hands independently, performance of the right hand being slightly superior to the left. Participants then undertook 4 h 16 min distributed practice of a one degree of freedom etch-a-sketch task where the target was constrained to move irregularly in only the 45 degrees direction on the display. To track such a target accurately participants had to make in-phase coupled stochastic movements of the hands. In a post-practice 2D task, measures of phase lag showed anisotropic improvement in performance, the amount of improvement depending on the direction of motion on the display. Improvement was greatest in the practised 45 degrees and least in the orthogonal 135 degrees direction. Best and worst performances were no longer in the directions associated with right and left hands independently, but in directions requiring coupled movements of the two hands. These data support the proposal that the nervous system can establish a model of novel coupling between the hands and thereby form a task-dependent bimanual synergy for controlling the stochastic coupled movements as an entity.

Adult↗

PubMed1980

Equine locomotion: 2. The analysis of coordination between limbs of trotting standardbreds.

A high-speed cinematographic analysis of interlimb time and linear gait characteristics was carried out at racing speed (12.0 m/sec; 1:23.6 min/km) in 30 clinically sound Standard-bred trotters. In respect of the whole group of horses the racing trot was found to be a very regular and symmetric gait. The variation coefficient between different horses ranged from 8 to 21 per cent depending on the particular characteristic being investigated. The variations recorded in individual animals were about 60 per cent less than those found for the whole group. The most stable characteristics were the duration and length of the step. Some horses exhibited striking gait asymmetries and the possible causes are considered. It is also observed that some trotters adopt a transition of gait that cannot be classified as a trot.

Animals↗

PubMed2006

Context-dependent modulation of interlimb cutaneous reflexes in arm muscles as a function of stability threat during walking.

Cutaneous reflexes evoked in the muscles of the arms with electrical stimulation of nerves of the foot ("interlimb reflexes") are observed during walking. These reflexes have been suggested to coordinate the actions of the legs and arms when walking is disturbed. Recently, we showed that cutaneous reflexes evoked in the leg muscles after stimulation at the foot are modulated according to the level of postural threat during walking. We hypothesized that the amplitude of interlimb cutaneous reflexes would similarly be modulated when subjects walk in unstable environments. Subjects walked on a treadmill under four walking conditions: 1) normal; 2) normal with unpredictable anterior-posterior (AP) perturbations; 3) arms crossed; and 4) arms crossed with unpredictable AP perturbations. Interlimb reflexes evoked from electrical stimulation of the right superficial peroneal or sural nerves were recorded bilaterally, at four points of the step cycle. These reflexes were compared between conditions in which the arms were moving in a similar manner: 1) normal versus AP walking and 2) arms crossed versus arms crossed with AP perturbations. Differences in reflex amplitudes between arms-crossed conditions were observed in most upper limb muscles when subjects were perturbed while walking compared with undisturbed walking. This effect was less apparent when the arms were swinging freely. The results indicate that the strength of interlimb connections is influenced by the level of postural threat (i.e., the context of the behavior), thereby suggesting that these reflexes serve a functional link between the legs and arms during locomotion.

Adult↗

PubMed2003

Intralimb incoordination in patients with ataxia.

Intralimb incoordination needs to be distinguished from interlimb incoordination, which is observed in Parkinson's disease (PD). With this aim, intra and inter-limb coordination impairment in the lower limbs were measured in 13 patients with cerebellar ataxia during pedaling of an ergometer with left and right pedals that can be rotated independently. Intralimb incoordination in patients with cerebellar ataxia was compared with interlimb incoordiation in patients with PD. We concluded that impairment of intralimb coordination in ataxia patients leads to inability to maintain pedaling amplitude and speed, but unlike in PD, right and left coordination was roughly preserved. Our method may be useful for assessing the severity of ataxia and detecting hidden parkinsonism in ataxic patients.

Adult↗