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PubMed2000

Age-related deterioration of coordinated interlimb behavior.

Younger and older participants performed two-limb coordination patterns of homologous (similar) and nonhomologous (dissimilar) effectors during 1:1 synchronization, according to the in-phase or anti-phase mode. The aim of the study was to examine age-related changes during the production of these basic movement patterns and their relative stability difference. The findings revealed that the aging process modulated the coordination dynamics as a function of effector system characteristics. Whereas the homologous system was resistant to age-related deficits, movements of the nonhomologous system showed coordinative degradation that was most apparent during execution of the anti-phase mode. The latter performance regression is argued to be an expression of age-dependent declines in cognitive regulation and afferent information processing. This implies that deterioration in coordinated behavior across the life span may be strongly task dependent because of a combined effect of cognitive and sensory components.

Adult↗
PubMed2003

Coordinated interlimb compensatory responses to electrical stimulation of cutaneous nerves in the hand and foot during walking.

It has been shown that stimulation of cutaneous nerves innervating the hand (superficial radial, SR) and foot (superficial peroneal, SP) elicit widespread reflex responses in many muscles across the body. These interlimb reflex responses were suggested to be functionally relevant to assist in motor coordination between the arms and legs during motor tasks such as walking. The experiments described in this paper were conducted to test the hypothesis that interlimb reflexes were phase-dependently modulated and produced functional kinematic changes during locomotion. Subjects walked on a treadmill while electromyographic (EMG) activity was collected continuously from all four limbs, and kinematic recordings were made of angular changes across the ankle, knee, elbow, and shoulder joints. Cutaneous reflexes were evoked by delivering trains of electrical stimulation pseudorandomly to the SP nerve or SR nerves in separate trials. Reflexes were phase-averaged according to the time of occurrence in the step cycle, and phasic amplitudes and latencies were calculated. For both nerves, significant phase-dependent modulation (including reflex reversals) of interlimb cutaneous reflex responses was seen in most muscles studied. Both SR and SP nerve stimulation resulted in significant alteration in ankle joint kinematics. The results suggest coordinated and functionally relevant reflex pathways from the SP and SR nerves onto motoneurons innervating muscles in nonstimulated limbs during walking, thus extending observations from the cat to that of the bipedal human.

Adolescent↗
PubMed1979

Interlimb coordination during stepping in the cat: an electromyographic analysis.

1. Simultaneous electromyographic (EMG) records were obtained from a single-joint extensor muscle of each of the four limbs of intact cats during repeated overground stepping trials. 2. In each limb, the temporal spacing of step cycles was determined by measurements of the intervals between consecutive terminations of EMG activity, since this occurs in a consistent relationship to the removal of the limb from the ground. By measuring the latencies between step cycles so determined, the temporal spacing of step cycles between limbs was determined. Each latency was expressed as a function of step duration or as a phase interval. 3. Analysis of the cooordination of step cycles of both homologous limb pairs (the forelimbs and hindlimbs), both homolateral limb pairs (the fore- and hindlimb on the right and left sides), and both sets of diagonal limbs suggest that the step cycles of the four limbs are coordinated according to a few frequently occurring patterns. However, the representation of a large number of phase intervals between these preferred patterns indicates a substantial amount of variability in interlimb coupling. 4. Analysis of the interaction of different interlimb-coupling patterns indicates that during alternate coordination of hindlimbs, coupling of the other limbs is fairly predictable. The step cycles of the forelimbs and hindlimbs are spaced according to a trotting form of coupling. During in-phase coordination of hindlimbs, the patterns of coordination of the other limbs are more diffuse. Forelimbs step cycles are coupled via a number of different modes, as are those of the forelimbs and hindlimbs. 5. It is concluded that the step cycles of different limbs are coordinated, but the association of observed patterns of coordination with any known neural pathways or the interaction of neural pathways should be approached with caution. The variability about the frequently occurring patterns is interpreted as an expression of the faculatative capabilities of the neural mechanisms controlling locomotion. Thus, these data favor a model of interlimb control during stepping, which recognizes preferred patterns of coordination and the variability about these patterns.

Animals↗
PubMed1985

[Studies on interlimb coordination of the cat during locomotion].

Experiments were performed on cats with chronic spinal lesions, in order to decide which systems either interlimb-propriospinal systems or direct descending systems, plays important roles for coordination of overground locomotion of the cat. The phase relationships between homolateral, crossed, bilateral forelimbs and hindlimbs were analysed during locomotion. For 27 cats a single hemisection was applied at the level of C1 or T12. In 19 cats, the spinal cord was firstly hemisected at around T12, then 5-183 days after, contralateral thoracic cord around T8 was hemisected (T-T; thoracic-thoracic bilateral serial hemisection). Therefore in this group, only crossed descending propriospinal systems were preserved and all other propriospinal systems and direct descending systems were severed. In 9 cats following spinal cord hemisection at around C1, at interval of 11-71 days the second hemisection was carried out at thoracic cord about T12 (C-T; cervico-thoracic bilateral serial hemisection). In this group homolateral and crossed propriospinal systems were left intact. In the single hemisection group, the phase relationships of all the six combinations of limbs were quite similar to normal group. Hence for the control of the phase relation during locomotion, only onesided neuronal circuits were needed. In both T-T and C-T group, polar diagram analysis revealed that the phase relationships were completely disturbed in any combination of the forelimbs and hindlimbs. It was concluded that the direct descending systems rather than propriospinal systems play important roles in control of phase relationship during locomotion.

Afferent Pathways↗
PubMed1982

Interlimb coordination in cat locomotion investigated with perturbation. I. Behavioral and electromyographic study on symmetric limbs of decerebrate and awake walking cats.

During locomotion of decerebrate and awake walking cats, perturbation (mechanical tap) was applied to the paw dorsum of the left forelimb (LF), and the responses of both forelimbs were recorded cinematographically and electromyographically (EMG). When the tap was applied during the LF stance phase, the duration of the ongoing LF stance was shortened by 10%; in the right forelimb (RF), the duration of the concomitant swing was shortened by 32%. A tap during the LF swing phase prolonged the duration of the ongoing LF swing phase and the concomitant RF stance phase by 55 and 15%, respectively. Analysis of RF joint angle excursions showed that the shortening of the RF swing phase was related mainly to acceleration of extension movement in the late swing phase; the prolongation of the RF stance phase was related to prolonged extension movement in the late stance phase. While EMG activities were relevant to these limb movements, a notable observation was that, by tapping the LF during the LF stance phase, EMG activity in the RF extensor started well before onset of the elbow extension movement to place down the limb; without the tap, the extensor activity started shortly after onset of the extension. Closely related to changes in phase durations of each forelimb, the period of bisupport phase where both forelimbs were in stance, was retained for more than 40% of that of unperturbed steps, even when the RF or LF made the first touchdown after the tap. The rostrocaudal level at RF touchdown after the tap was comparable to unperturbed steps. These findings on interlimb relation suggest that neural control ensures coordinated movements between symmetric limbs during locomotion.

Animals↗
PubMed1982

Interlimb coordination during stepping in the cat: in-phase stepping and gait transitions.

The coordination of step cycles between all 4 limbs during in-phase stepping and during transitions to and from alternate stepping was studied in 12 adult cats during repeated overground stepping trials. The temporal spacing of step cycles of the different limbs was determined from analysis of electromyographic (EMG) activity in a single extensor muscle of each limb. Patterns of coordination of the different limbs were established on the basis of the frequency with which phase values separating step cycles were encountered. Steps in which the phasing of step cycles of the two hindlimbs were closer to true in-phase coordination than true alternation (phase between 270 degrees and 90 degrees) and where similar coupling was found in both the preceding and following steps were defined as steady state conditions. Distinct patterns of coordination of forelimb-forelimb and forelimb-hindlimb step cycles were noted under steady state conditions. During stepping sequences which include transitions either to or from alternate stepping, both gradual and abrupt phase changes were found. The changes in both forelimb-forelimb and forelimb-hindlimb phase relationships were more often gradual than abrupt. Where abrupt changes were encountered in the change in phase relationships between one such limb pair the phase change in the other pair was gradual. Changes in hindlimb-hindlimb phase relationships during transitions were nearly always abrupt. It is concluded that the 4 limbs are coordinated during in-phase stepping according to a few patterns, but that the variability about these patterns makes their association with simple neural circuitry rather speculative. The finding that transitions were most often gradual is interpreted in terms of a state-dependent model of interlimb control, in which the type of transition utilized depends on the strength of neural coupling of step cycles of all 4 limbs at the time that the transition is initiated.

Animals↗
PubMed1986

Spatiotemporal linkage in infant interlimb coordination.

Timing and direction of movement of the arms and hands of 7- and 11-month-old infants were examined during bimanual reaching for a transparent cube. Videotape analysis of frames from release of the hands to initial contact of one hand with the cube revealed that hand movement was more often temporally linked for 7-month infants than for 11-month infants. Also, at 7 months, the hands moved toward the cube in the same direction, whereas at 11 months, they moved in opposite (complementary) directions. Thus, although both 7- and 11-month infants perform bimanual reaching, at 7 months the 2 hands are linked both in timing and direction of movement in a fashion different than at 11 months.

Child Development↗
PubMed1992

Interlimb coordination in Parkinson's disease.

This study examined the degree to which Parkinson's disease (PD) patients could "spatially link" the upper limbs to facilitate the performance of bimanual simultaneous movements. Six right-handed PD patients, and seven normal age- and sex-matched controls performed three different tasks: (a) an isotonic elbow flexion as rapidly as possible through an angle of 30 degrees; (b) an isometric contraction of the flexor muscles at the elbow joint to 40% and 60% of maximal volitional force (MVF) for a period of 5 s; (c) an isometric contraction for 2.5 s with one limb, then simultaneously performing an isotonic flexion with the contralateral limb while maintaining the isometric contraction for 2.5 s more. As expected, PD patients were significantly slower in performing the isotonic movement and produced lower peak velocities than the controls. More importantly, the two groups were differentially affected during the bimanual condition. In normals, movement time decreased and peak velocity increased in the bimanual condition. In contrast, PD patients showed increased movement times and sometimes decreased peak velocities in the bimanual condition. The results suggest that normal subjects utilize bilateral outflow to symmetrical muscle groups to synchronize the two limbs in the bimanual task, whereas PD patients dissociate the two limbs.

Aged↗
PubMed2002

Modeling rhythmic interlimb coordination: beyond the Haken-Kelso-Bunz model.

Although the Haken-Kelso-Bunz (HKB) model was originally formulated to account for phase transitions in bimanual movements, it evolved, through experimentation and conceptual elaboration, into a fundamental formal construct for the experimental study of rhythmically coordinated movements in general. The model consists of two levels of formalization: a potential defining the stability properties of relative phase and a system of coupled limit cycle oscillators defining the individual limb movements and their interactions. Whereas the empirical validity of the potential is well established, the validity of the formalization in terms of coupled oscillators is questionable, both with regard to the assumption that individual limb movements are limit cycle oscillators with (only) two active degrees of freedom and with regard to the postulated coupling. To remedy these limitations a more elaborate system of coupled oscillators is outlined, comprising two coupled limit cycle oscillators at the neural level, each of which is coupled to a linearly damped oscillator, representing the end-effectors.

Extremities↗
PubMed1982

Interlimb coordination in cat locomotion investigated with perturbation. II. Correlates in neuronal activity of Deiter's cells of decerebrate walking cats.

The effects of mechanical stimulation (tap) on single unit activity of Deiter's neurons were analysed in walking cats decerebrated at the premammillary level. Deiters' neurons projecting to the ipsilateral cervical, but not to the lumbosacral, spinal cord (C-Deiters' neurons) were identified by antidromic activation, cerebellar stimulation, and localization of the neurons. During each unperturbed cycle of quadrupedal locomotion, most C-Deiters' neurons showed two frequency modulation peaks in their impulse discharges: one (A peak) in the late swing (E1) or the early stance (E2) phase, the other (B peak) in the late stance (E3) or the early swing (F) phase, of the ipsilateral forelimb. The A peak started to rise shortly before the ipsilateral forelimb was placed. When mechanical perturbation was applied during locomotion to the paw dorsum of the left forelimb (LF) in its stance phase, the ongoing LF stance phase shortened and the simultaneous swing phase of the right forelimb (RF) shortened. Accordingly, in the RF, extensor activity in the swing phase to place down the limb occurred earlier than in unperturbed step cycles. The same LF tap induced a marked enhancement of impulse discharges in C-Deiters' neurons on the right side (with a magnitude of 20-100 imp/s, and the shortest latency of 25 ms). This enhancement was more pronounced than that induced when the perturbation was applied to the LF during its swing phase. The latency manifested a close time relation to the RF extensor activity supporting the postulate that the increased C-Deiters' activity in the RF swing phase contributes to the earlier onset of RF extensor activity which plays an important role in maintaining alternating footfalls after perturbation.

Animals↗
PubMed2003

A coupled oscillator model of disordered interlimb coordination in patients with Parkinson's disease.

Coordination between the left and right limbs during cyclic movements, which can be characterized by the amplitude of each limb's oscillatory movement and relative phase, is impaired in patients with Parkinson's disease (PD). A pedaling exercise on an ergometer in a recent clinical study revealed several types of coordination disorder in PD patients. These include an irregular and burst-like amplitude modulation with intermittent changes in its relative phase, a typical sign of chaotic behavior in nonlinear dynamical systems. This clinical observation leads us to hypothesize that emergence of the rhythmic motor behaviors might be concerned with nonlinearity of an underlying dynamical system. In order to gain insight into this hypothesis, we consider a simple hard-wired central pattern generator model consisting of two identical oscillators connected by reciprocal inhibition. In the model, each oscillator acts as a neural half-center controlling movement of a single limb, either left or right, and receives a control input modeling a flow of descending signals from higher motor centers. When these two control inputs are tonic-constant and identical, the model has left-right symmetry and basically exhibits ordered coordination with an alternating periodic oscillation. We show that, depending on the intensities of these two control inputs and on the difference between them that introduces asymmetry into the model, the model can reproduce several behaviors observed in the clinical study. Bifurcation analysis of the model clarifies two possible mechanisms for the generation of disordered coordination in the model: one is the spontaneous symmetry-breaking bifurcation in the model with the left-right symmetry. The other is related to the degree of asymmetry reflecting the difference between the two control inputs. Finally, clinical implications by the model's dynamics are briefly discussed.

Ataxia↗
PubMed2000

Relative phase dynamics in perturbed interlimb coordination: stability and stochasticity.

Various stability features of bimanual rhythmic coordination, including phase transitions, have been modeled successfully by means of a one-dimensional equation of motion for relative phase obeying a gradient dynamics, the Haken-Kelso-Bunz model. The present study aimed at assessing pattern stability for stationary performance and estimating the model parameters (a, b, and Q) for the stochastic extension of this model. Estimates of a and b allowed for reconstruction of the potential defining the gradient dynamics. Two coordination patterns between the forearms (in-phase, antiphase) were performed at seven different frequencies. Model parameters were estimated on the basis of an exponential decay parameter describing the relaxation behavior of continuous relative phase following a mechanical perturbation. Variability of relative phase and relaxation time provided measures of pattern stability. Although the predicted inverse relation between pattern stability and movement frequency was observed for the lower tempo conditions, it was absent for the higher tempos, reflecting the influence of task constraints. No statistically significant differences in stability were observed between the two coordination modes, indicating the influence of intention. The reconstructed potential reflected the observed stability features, underscoring the adequacy of the parameter estimations. The relaxation process could not be captured adequately by means of a simple exponential decay function but required an additional oscillatory term. In accordance with previous assumptions, noise strength Q did not vary as a function of movement frequency. However, systematic differences in Q were observed between the two coordination modes. The advantages and (potential) pitfalls of using stationary performance of single patterns to examine the stability features of a bistable potential were discussed.

Computer Simulation↗
PubMed1998

Interactive processes during interlimb coordination: combining movement patterns with different frequency ratios.

The present study examined the formation of a movement pattern that was added to an ongoing coordinative regime across different limb combinations. It was hypothesized that the addition of the secondary mode would perturb the ongoing primary mode by adding rhythmic complexity to the task requirements. Furthermore, the formation of the secondary mode was predicted to be affected by the ongoing coordination pattern. In Exp. 1, a primary multifrequency mode (2:1 ratio) was performed while a secondary isofrequency mode (1:1 ratio) was initiated midway into the trials, whereas the reversed dual-pattern conditions were examined in Exp. 2. The results from both experiments showed that the multifrequency mode deteriorated across limb combination under dual-pattern as compared to single-pattern conditions. The isofrequency mode was also affected under combined pattern conditions, but its degradation was a function of the limb combination under consideration. In particular, the non-homologous limbs, which demonstrated less stable behavior than the homologous limbs under single-pattern conditions, were affected most strongly when confronted with the simultaneous production of the multifrequency mode. In addition, anti-phase movements deteriorated more than in-phase movements, supporting indirectly the contention that afferent feedback monitoring complexity differs for the two movement configurations. The findings of this study suggest that manipulation of task requirements can be used to examine pattern durability and formation in view of dynamical perturbations.

Adult↗
PubMed1979

Experimental modification of interlimb coordination during locomotion of a Crusacea.

An anlaysis of the phase relationship between the same side legs (homolateral coupling) of Rock Lobster (Crustacea) is performed during sequence of locomtion. On intact animals the phase relationship is similar to the most common insect or mammalian patterns ('alternating model'). After successive autotomy of the walking legs, another coordination pattern occurs progressively; it is an 'in phase model'. From these experiments it can be concluded that there exists a central motor program tightly coordinating each thoracic ganglion and that the alternating pattern could be due to a reorganization of the synchronous system by a peripheral proprioceptive inflow.

Animals↗
PubMed1990

Interlimb coordination of stance in children: divergent modulation of spinal reflex responses and cerebral evoked potentials in terms of age.

EMG responses in the gastrocnemius (GM) and tibialis anterior muscles (TA) of both legs together with cerebral evoked potentials (CP), were recorded following perturbations of stance on a treadmill with split belts, in two age groups of children. Unilateral displacements were followed by ipsilateral short latency and bilateral long latency EMG responses. The CP was similar in both tasks. When displacements were simultaneously induced in opposite directions, a significant reduction in the long latency components of EMG responses occurred, while the amplitude of the CP was maximal in this condition. In the older children the CP and long latency EMG responses were larger and the short latency reflex potentials smaller in all conditions compared to the younger children. It is concluded that (1) CP and EMG responses reflect a divergent modulation of a given somatosensory input; (2) developmental changes are reflected in alterations in the amplitude of CP and EMG responses; (3) there is no evidence of transcortically mediated muscle responses.

Aging↗
PubMed1995

Leg muscle activation during gait in Parkinson's disease: adaptation and interlimb coordination.

Adaptation in leg muscle activity and coordination between lower limbs were studied during walking on a treadmill with split belts in one group of parkinsonian patients and one of age-matched healthy subjects. Four different belt speeds (0.25/0.5/0.75/1.0 m/sec) were applied in selected combinations to the left and right leg. While these walking conditions were easily tolerated by the healthy subjects, the parkinsonian patients usually reached the limits of their walking capabilities. Both groups adapted automatically to a change in belt speed within approximately 20 stride cycles. Healthy subjects adapted by reorganizing their stride cycle with a relative shortening of duration of support and lengthening of the swing phase of the "fast" leg and vice versa on the "slow" leg. The patients showed a restricted range of stride frequencies for the various belt speeds during normal and split-belt walking with consequent deviations in the reorganization of the stride cycle. In both healthy subjects and patients, ipsilateral gastrocnemius and contralateral tibialis anterior electromyographic (EMG) activity increased predominantly with an ipsilateral increase in belt speed. Two main differences were observed in the EMG patterns: (1) In the patients leg muscle EMG activity was less modulated and gastrocnemius EMG amplitude was small during normal and split-belt walking. However, there was no significant difference between the two groups in respect to the reorganization of the EMG pattern required for the various split-belt walking conditions. (2) The amount of co-activation of antagonistic leg muscles during the support phase of the stride cycle was greater in the patients compared to the healthy subjects during normal and split-belt walking. It is suggested that reduced EMG modulation and recruitment in the leg extensors may contribute to the impaired walking of the patients. This in turn is a result of an impaired proprioceptive feedback from extensor load receptors. This defective control is partially compensated for in parkinsonian patients by a greater amount of leg flexor activation which leads to a higher degree of co-activation. Visual input plays a role in the control of this increased activation.

Adaptation, Physiological↗
PubMed2005

Hierarchical organisation of neuro-anatomical constraints in interlimb coordination.

Based on the observation that bimanual finger tapping movements tend toward mirror symmetry with respect to the body midline, despite the synchronous activation of non-homologous muscles, F. Mechsner, D. Kerzel, G. Knoblich, and W. Prinz (2001) [Perceptual basis of bimanual coordination. Nature, 414, 69-73] suggested that the basis of rhythmic coordination is purely spatial/perceptual in nature, and independent of the neuro-anatomical constraints of the motor system. To investigate this issue further, we employed a four finger tapping task similar to that used by F. Mechsner and G. Knoblich (2004) [Do muscle matter in bimanual coordination? Journal of Experimental Psychology: Human Perception and Performance, 30, 490-503] in which six male participants were required to alternately tap combinations of adjacent pairs of index (I), middle (M) and ring (R) fingers of each hand in time with an auditory metronome. The metronome pace increased continuously from 1 Hz to 3 Hz over the course of a 30-s trial. Each participant performed three blocks of trials in which finger combination for each hand (IM or MR) and mode of coordination (mirror or parallel) were presented in random order. Within each block, the right hand was placed in one of three orientations; prone, neutral and supine. The order of blocks was counterbalanced across the six participants. The left hand maintained a prone position throughout the experiment. On the basis of discrete relative phase analyses between synchronised taps, the time at which the initial mode of coordination was lost was determined for each trial. When the right hand was prone, transitions occurred only from parallel symmetry to mirror symmetry, regardless of finger combination. In contrast, when the right hand was supine, transitions occurred only from mirror symmetry to parallel but no transitions were observed in the opposite direction. In the right hand neutral condition, mirror and parallel symmetry are insufficient to describe the modes of coordination since the hands are oriented orthogonally. When defined anatomically, however, the results in each of the three right hand orientations are consistent. That is, synchronisation of finger tapping is determined by a hierarchy of control of individual fingers based on their intrinsic neuro-mechanical properties rather than on the basis of their spatial orientation.

Adult↗