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Dissociation of muscular and spatial constraints on patterns of interlimb coordination.

Interlimb coordination is subject to constraints. One major constraint has been described as a tendency for homologous muscle groups to be activated simultaneously. Another has been described as a biasing of limb segments to movement in the same direction. In 2 experiments, the 2 constraints were placed in opposition: In-phase or antiphase contraction of homologous muscles of contralateral limbs produced movement that was spatially antiphase or in-phase, respectively. Probability distributions of relative phase were obtained under manipulations of phase detuning and movement speed. They revealed that the equilibrium and stability of coordination were related, respectively, to spatial relative phase and muscular relative phase. Previously observed spatial and muscular constraints reflect a (possibly very general) factorization of attractor location and attractor strength in the dynamics of interlimb coordination.

Humans↗

Interlimb coordination, gait, and neural control of quadrupedalism in chimpanzees.

Interlimb coordination is directly relevant to the understanding of the neural control of locomotion, but few studies addressing this topic for nonhuman primates are available, and no data exist for any hominoid other than humans. As a follow-up to Jungers and Anapol's ([1985] Am. J. Phys. Anthropol. 67:89-97) analysis on a lemur and talapoin monkey, we describe here the patterns of interlimb coordination in two chimpanzees as revealed by electromyography. Like the lemur and talapoin monkey, ipsilateral limb coupling in chimpanzees is characterized by variability about preferred modes within individual gaits. During symmetrical gaits, limb coupling patterns in the chimpanzee are also influenced by kinematic differences in hindlimb placement ("overstriding"). These observations reflect the neurological constraints placed on locomotion but also emphasize the overall flexibility of locomotor neural mechanisms. Interlimb coordination patterns are also species-specific, exhibiting significant differences among primate taxa and between primates and cats. Interspecific differences may be suggestive of phylogenetic divergence in the basic mechanisms for neural control of locomotion, but do not preclude morphological explanations for observed differences in interlimb coordination across species.

Animals↗

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.

Adult↗

Overground locomotion in intact rats: interlimb coordination, support patterns and support phases duration.

The interlimb coordination during overground locomotion was analysed in intact rats, using the method of contact electrodes (Górska et al. 1998). It was found that in animals moving with a speed ranging from 10 to 78 cm/s (step cycles 685 to 215 ms, respectively) the interlimb coordination was characterized by homologous phase shifts close to 0.5 and much shorter diagonal than lateral phase shifts. These features corresponded to symmetrical gait with diagonal sequence and diagonal couplets (Hildebrandt 1976). Shortening the step cycle changed the gait from a walking trot (duty factor > 0.5) into a running trot (duty factor < 0.5). Correspondingly, the support patterns in the four-legged step cycles, i.e., the sequence of phases of support on various limbs changed: the support on diagonal limbs persisted but the three-limb support was replaced by one-limb support and the support on homolateral limbs by phases of flight. For each phase of support the relationship between its absolute and relative durations and the step cycle duration is being described. The paper explains the variability of support patterns described in the literature. The picture of locomotion obtained in intact rats will be used as a template for studying locomotor control deficits after CNS lesions.

Animals↗

A new learning paradigm: adaptive changes in interlimb coordination during perturbed locomotion in decerebrate cats.

Adaptive interlimb coordination was studied during perturbed locomotion in experimental chronically decerebrate cats. Perturbations were applied to the stance phases of the left forelimb using a newly developed treadmill which consisted of three compartments, one each for the left forelimb, the left hindlimb and both right limbs. During the perturbed locomotion, the treadmill belt for the left forelimb was driven at about twice the speed for the other limbs. During the first 1-50 perturbed steps, the step cycles of both forelimbs showed marked fluctuations; thereafter, the animals achieved stable locomotion by slightly shortening step cycle durations, and also by adjusting durations of bisupport phases asymmetrically in the left and right forelimbs. The present method provides a new test paradigm for adaptive interlimb coordination in locomotion, which will be useful in studying neural mechanisms of motor learning involving cerebellar synaptic plasticity.

Adaptation, Psychological↗

The control and learning of patterns of interlimb coordination: past and present issues in normal and disordered control.

The present paper provides a historical note on the evolution of the behavioral study of interlimb coordination and the reasons for its success as a field of investigation in the past decades. Whereas the original foundations for this field of science were laid down back in the seventies, it has steadily grown in the past decades and has attracted the attention of various scientific disciplines. A diversity of topics is currently being addressed and this is also expressed in the present contributions to the special issue. The main theme is centered on the brain basis of interlimb coordination. On the one hand, this pertains to the study of the control and learning of patterns of interlimb coordination in clinical groups. On the other hand, basic neural approaches are being merged together with behavioral approaches to reveal the neural basis of interlimb coordination.

Extremities↗

Interlimb coordination during locomotion: what can be adapted and stored?

Interlimb coordination is critically important during bipedal locomotion and often must be adapted to account for varying environmental circumstances. Here we studied adaptation of human interlimb coordination using a split-belt treadmill, where the legs can be made to move at different speeds. Human adults, infants, and spinal cats can alter walking patterns on a split-belt treadmill by prolonging stance and shortening swing on the slower limb and vice versa on the faster limb. It is not known whether other locomotor parameters change or if there is a capacity for storage of a new motor pattern after training. We asked whether adults adapt both intra- and interlimb gait parameters during split-belt walking and show aftereffects from training. Healthy subjects were tested walking with belts tied (baseline), then belts split (adaptation), and again tied (postadaptation). Walking parameters that directly relate to the interlimb relationship changed slowly during adaptation and showed robust aftereffects during postadaptation. These changes paralleled subjective impressions of limping versus no limping. In contrast, parameters calculated from an individual leg changed rapidly to accommodate split-belts and showed no aftereffects. These results suggest some independence of neural control of intra- versus interlimb parameters during walking. They also show that the adult nervous system can adapt and store new interlimb patterns after short bouts of training. The differences in intra- versus interlimb control may be related to the varying complexity of the parameters, task demands, and/or the level of neural control necessary for their adaptation.

Adaptation, Physiological↗

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.

Adult↗

Rigidity and bradykinesia reduce interlimb coordination in Parkinsonian gait.

OBJECTIVE: To assess the influence of rigidity and bradykinesia and the extent of dopaminergic degeneration on interlimb coordination during walking in early, drug-naive patients with Parkinsons disease (PD). DESIGN: The interlimb coordination was examined during a systematic manipulation of walking speed on a treadmill. The phase relations between arm and leg movements were related to the clinical measures of rigidity and bradykinesia as well as to the extent of dopaminergic degeneration. SETTING: Movement disorders outpatient clinic (including motion analysis laboratory) and a nuclear medicine department of a university hospital. PARTICIPANTS: Twenty-nine early and drug-naive PD patients. INTERVENTIONS: Not applicable. MAIN OUTCOME MEASURES: The interlimb coordination during walking was evaluated by studying the (continuous) relative phase relations between movements of arms and legs. The clinical assessment of rigidity and bradykinesia was performed by using the Unified Parkinson Disease Rating Scale. The dopaminergic degeneration was expressed as striatal 2beta-carboxymethoxy-3beta-(4-iodophenyl) tropane (beta-CIT) single-photon emission computed tomography (SPECT) binding. RESULTS: The mean relative phase between arm and leg movements increased significantly with walking speed in all patients. Significant correlations were found between the rigidity and bradykinesia and the coordination measures ( P </=.007), as well as contralateral striatal [ 123 I]beta-CIT SPECT binding and coordination measures ( P <.001), in terms of asymmetry indices. CONCLUSIONS: Early, drug-naive PD patients in this sample were able to adapt their coordination patterns when walking speed was systematically manipulated. However, bradykinesia and rigidity as well as the extent of degeneration of the dopaminergic system were associated with a limited adaptive ability (flexibility) in movement coordination. The combination of a drug treatment that controls bradykinesia and rigidity and a physical therapy exercise programs possibly using external cues mechanisms are required to obtain relevant effects on gait in PD patients.

Adult↗

Interlimb coordination during fictive locomotion in the thalamic cat.

Efferent discharges in muscle nerves of the four limbs were recorded simultaneously during spontaneous fictive locomotion in thalamic cats with the goal of understanding how the central nervous system controls interlimb coordination during stepping. The onset of the bursts of activity in the nerve of a selected flexor muscle in each limb allowed the temporal and the phase relationships between the fictive step cycle of a pair of limbs to be determined. Our main results are the following: 1) the fictive step cycles of the two forelimbs are always strictly alternated whereas the phasing of the step cycles of either the two hindlimbs or pairs of homolateral or diagonal limbs is more variable; 2) the time interval between the onsets of the flexor bursts of one of the two pairs of diagonal limbs is independent of the step cycle duration; 3) distinct patterns of interlimb coordination exist during fictive locomotion; a small number of patterns of coordination involving all four limbs, which correspond to the walking and the trotting gaits in the intact cat, occur very frequently. The results demonstrate that the central nervous system deprived of phasic afferent inputs from the periphery has the capacity to generate most of the patterns of interlimb coordination which occur during real locomotion. They further support the view that the central pattern of interlimb coordination essentially results from diagonal interaction between a forelimb generator for locomotion and a hindlimb one.

Afferent Pathways↗

Human interlimb coordination: the first 6 months of independent walking.

The organization of interlimb coordination was studied in newly walking infants with and without support and in infants who had been walking 0.5, 1, 3, and 6 months. Analysis of the temporal and distance phase relationships between the limbs revealed that newly walking infants exhibit interlimb coordination similar to that of mature walkers. The coupling between the limbs, however, is loosely constrained, approaching adult-like consistency after 3 months of walking. Providing support for the newly walking infant afforded a more consistent, mature coordination suggesting that neuromuscular development is in advance of its ability to function in the physical world.

Child Development↗

Explanatory limitations of the HKB model: incentives for a two-tiered model of rhythmic interlimb coordination.

The HKB model for rhythmic interlimb coordination has highlighted the importance of coordinative stability and loss of stability, and introduced, with this focus, a new set of explanatory constructs. However, the phenomenological character of both parts of this model (i.e., the potential and the associated system of coupled oscillators) precludes an understanding of how the observed stability characteristics are related to more specific (e.g., biomechanical and neurophysiological) aspects of the movement system. A two-tiered model (involving a distinction between 'neural' and 'effector' dynamics) is discussed that offers handles for addressing such underpinnings of the identified coordination dynamics. The promise of the model in this regard is illustrated by two recent studies showing how explicit accounts of the effector dynamics may help disclose why (and how) particular properties of the peripheral system affect the overall coordination dynamics.

Attention↗

Relative phase dynamics in perturbed interlimb coordination: the effects of frequency and amplitude.

Schöner [Schöner G (1995) Ecol Psychol 7: 291-314] argued that the relative phase dynamics of rhythmic interlimb coordination may be attributed to the timing level in that the stability properties of the relative phase are largely independent of dynamical principles operating at the goal level, such as those related to the maintenance of a particular amplitude or target position. Yet, according to the coupling functions in the coupled oscillator model proposed by Haken et al. [Haken H, Kelso JAS, Bunz H (1985) Biol Cybern 51: 347-356], the effect of frequency on the stability properties of relative phase is either wholly or partially mediated by frequency-induced changes in amplitude, implying that the relative phase dynamics strongly depends on spatial factors. In order to distinguish between these contrasting interpretations of the organizational principles underwriting the phase dynamics of interlimb coordination, an experiment was conducted in which the effects of frequency and amplitude on the stability of relative phase were separated. Six subjects performed both in-phase and antiphase coordination patterns at seven different frequencies and three different amplitudes. Two measures of pattern stability were used, the standard deviation of relative phase and the exponent of the relaxation process following phasic perturbations of relative phase. According to both measures, pattern stability decreased with increasing frequency, whereas the amplitude manipulation only had a significant effect on the standard deviation of relative phase. This result was interpreted to imply that the organizational principles at the (relative) timing level are affected only moderately by task constraints pertaining to the goal level, and that models of interlimb coordination in which amplitude coupling plays a partial or subordinate role should be preferred above models relying solely on amplitude coupling.

Adult↗

Interlimb coordination during stepping in the cat: the role of the dorsal spinocerebellar tract.

The role of the dorsal spinocerebellar tract (DSCT) in the neural control of normal interlimb coordination during overground stepping in adult cats was investigated using select spinal cord lesions. Previously, it had been shown that lesions of the caudal thoracic dorsal columns (DCs) which might involve the DSCT or its afferent fibers resulted in a marked change in the patterns of forelimb-hind limb coupling during locomotion. In the present study, more rostral DC lesions, which probably included the DSCT or its afferent fibers considerably less, resulted in nearly identical changes in the patterns of interlimb coordination during stepping. Lesions of the dorsolateral funniculus (DLF) at similar spinal levels resulted in no significant changes in interlimb coordination. These lesions did destroy the DSCT, since retrograde transport of horseradish peroxidase (HRP) from the anterior cerebellar vermis to the nucleus dorsalis was blocked caudal to the lesion. These results are consistent with the notion that the DSCT plays little if any role in the precise timing of step cycles of the different limbs.

Animals↗

Human neuronal interlimb coordination during split-belt locomotion.

Human interlimb coordination and the adaptations in leg muscle activity were studied during walking on a treadmill with split belts. Four different belt speeds (0.5, 1.0, 1.5, 2.0 m/s) were offered in all possible combinations for the left and right leg. Subjects adapted automatically to a difference in belt speed within 10-20 stride cycles. This adaptation was achieved by a reorganization of the stride cycle with a relative shortening of the duration of the support and lengthening of the swing phase of the "fast" leg and, vice versa, in support and swing duration on the "slow" leg. The electromyogram EMG patterns were characterized by two basic observations: (1) onset and timing of EMG activity were influenced by biomechanical constraints. A shortening of the support phase on the faster side was related to an earlier onset and increase in gastrocnemius activity, while a coactivation pattern in the antagonistic leg muscles was predominant during a prolonged support phase on the slower side. (2) A differential modulation of the antagonistic leg muscles took place. An increase in ipsilateral belt speed in combination with a constant contralateral belt speed was associated with an almost linear increase in ipsilateral gastrocnemius and contralateral tibialis anterior EMG activity, while the contralateral gastrocnemius and ipsilateral tibialis anterior EMG activity were little affected. It is concluded that a modifiable timing within the stride cycle takes place with a coupling between ipsilateral support and contralateral swing phase. The neuronal control of this coupling is obviously based on ipsilateral modulation of leg extensor EMG by proprioceptive feedback and an appropriate central (e.g. spinal) modulation of contralateral tibialis anterior EMG activity.

Adult↗

Brain areas involved in interlimb coordination: a distributed network.

Whereas behavioral studies have made significant contributions toward the identification of the principles governing the coordination of limb movements, little is known about the role of higher brain areas that are involved in interlimb coordination. Functional magnetic resonance imaging (fMRI) was used to reveal the brain areas activated during the cyclical coordination of ipsilateral wrist and foot movements. Six normal subjects performed five different tasks that were presented in a random order, i.e., isolated flexion-extension movements of the right wrist (WRIST) and right foot (FOOT), cyclical coordination of wrist and foot according to the isodirectional (ISODIR) and nonisodirectional (NON-ISODIR) mode, and rest (REST). All movements were auditory paced at 66 beats/min. During the coordination of both limb segments, a distributed network was identified showing activation levels in the supplementary motor area (SMA), cingulate motor cortex (CMC), premotor cortex (PMC), primary sensorimotor cortex (M1/S1), and cerebellum that exceeded the sum of the activations observed during the isolated limb movements. In addition, coordination of the limb movements in different directions was associated with extra activation of the SMA as compared to movements in the same direction. It is therefore concluded that the SMA is substantially involved in the coordination of the nonhomologous limbs as part of a distributed motor network. Accordingly, the long-standing exclusive association that has been made between this medial frontal area and bimanual (homologous) coordination needs to be abandoned and extended towards other forms of interlimb coordination (nonhomologous).

Adult↗

A comparison of intra- and interpersonal interlimb coordination: coordination breakdowns and coupling strength.

Intra- and interpersonal interlimb coordination of pendulums swung from the wrist was investigated. For both kinds of coordination, the steady state and breakdown of bimanual rhythmic coordination as indexed by the time series of the relative phase angle phi were studied under the manipulation of coordination mode, frequency of oscillation, and the difference in the eigenfrequencies (preferred tempos) of the individual oscillating limbs. The properties observed for both intra- and interpersonal coordination were those predicted by a dynamical model of rhythmic coordination that considers the coordinated limbs coupled to be nonlinear oscillators. Using a regression method, the coupling strengths of the coupled system were recovered. As predicted by the dynamical model, the strength of the dynamic was generally greater for the in-phase than the anti-phase mode and decreased with increasing frequency. Further, the strength of the interpersonal interlimb coupling was weaker than that of intrapersonal interlimb coupling.

Analysis of Variance↗

Effect of duration of upper- and lower-extremity rehabilitation sessions and walking speed on recovery of interlimb coordination in hemiplegic gait.

BACKGROUND AND PURPOSE: The effects of different durations of rehabilitation sessions for the upper extremities (UEs) and lower extremities (LEs) on the recovery of interlimb coordination in hemiplegic gait in patients who have had a stroke were investigated. SUBJECTS AND METHODS: Fifty-three subjects who had strokes involving their middle cerebral arteries were assigned to rehabilitation programs with (1) an emphasis on the LEs, (2) an emphasis on the paretic UE, or (3) a condition in which the paretic arm (UE) and leg (LE) were immobilized with an inflatable pressure splint (control treatment). The 3 treatment regimens were applied for 30 minutes, 5 days a week, during the first 20 weeks after onset of stroke. All subjects also participated in a rehabilitation program 5 days a week that consisted of 15 minutes of UE exercises and 15 minutes of LE exercises in addition to a weekly 11/2-hour session of training in activities of daily living. A repeated-measures design was used. Differences among the 3 treatment regimens were evaluated in terms of comfortable and maximal walking speeds. In addition, mean continuous relative phase (CRP) between paretic arm and leg (PAL) movements and nonparetic arm and leg (NAL) movements and standard deviations of CRP of both limb pairs as a measurement of stability (variability) were evaluated. RESULTS: Comfortable walking speed improved in the group that received interventions involving the LEs compared with the group that received interventions involving the UEs and the group that received the control treatment. No differences among the 3 treatment conditions were found for the mean CRP of NAL and PAL as well as the standard deviation of CRP of both limb pairs. DISCUSSION AND CONCLUSION: With the exception of an improved comfortable walking speed as a result of a longer duration of rehabilitation sessions, no differential effects of duration of rehabilitation sessions for the LEs and UEs on the variable we measured related to hemiplegic gait were found. Increasing walking speed, however, resulted in a larger mean CRP for both limb pairs, with increased stability and asymmetry of walking, indicating that walking speed influences interlimb coordination in hemiplegic gait.

Analysis of Variance↗