Coordination between arm and leg movements during locomotion

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

Coordination Between Arm and Leg Movements During Locomotion

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

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.     

Short-term limb immobilization affects motor performance

C. Ghez, J. Gordon, and M. R Ghilardi (1995; J. Gordon, M. R Ghilardi, & C. Ghez, 1995; R. L. Sainburg, M. R Ghilardi, H. Poizner, & C. Ghez, 1995) have found that proprio-ceptive deafferentation impairs feedforward and feedback mechanisms that control reaching movements. In the present study, the authors found immobilization-induced changes in limb kinematics, including joint motion, in 32 healthy participants who performed out-and-back movements before and after 0, 6, or 12 hr of immobilization of the left arm. Control participants did not undergo the arm immobilization procedure. Immobilization for 12 hr, but not 6 hr, caused trajectories with increased hand-path areas and altered interjoint coordination. The abnormalities were smaller in amplitude but similar in quality to those reported in deafferented patients (R. L. Sainburg et al.). In addition, movement onset point significantly drifted after immobilization. Thus, short-term limb disuse can affect interjoint coordination by acting on feedforward mechanisms. These behavioral alterations are potentially related to cortical plastic changes.     

Initial-state dependency of learning in young infants

With the aim of investigating the effect of the initial state of pre-learning on subsequent infant learning (i.e., the initial-state dependency), we observed the limb movements in 3-month-old infants in the course of a motor learning task. The session comprised 2-min pre-learning and 4-min learning periods, and the infants learned to move a toy using a string attached to either an arm (arm-based learning, Experiment 1) or a leg (leg-based learning, Experiment 2). Infants were assigned to low- and high-state groups in the initial-state condition according to the average velocity of the arm (Experiment 1) or leg (Experiment 2) movements during the pre-learning period. The results revealed that, during the learning period, infants in the low-state group increased the movement of their limbs, whereas those in the high-state group showed no significant changes in the movement of most of their limbs. These results suggest that infants demonstrating a low average velocity of movement in the initial state easily observed and learned the circular causality between self-produced movements and environmental changes. On the other hand, it seemed that infants demonstrating a high average velocity of movement in the initial state could not or did not need to increase their limb movements (the toy would already be shaking enough to form striking movements).     

Perception and action in swimming: Effects of aquatic environment on upper limb inter-segmental coordination

This study assessed perception–action coupling in expert swimmers by focusing on their upper limb inter-segmental coordination in front crawl. To characterize this coupling, we manipulated the fluid flow and compared trials performed in a swimming pool and a swimming flume, both at a speed of 1.35 m s⁻¹. The temporal structure of the stroke cycle and the spatial coordination and its variability for both hand/lower arm and lower arm/upper arm couplings of the right body side were analyzed as a function of fluid flow using inertial sensors positioned on the corresponding segments. Swimmers’ perceptions in both environments were assessed using the Borg rating of perceived exertion scale. Results showed that manipulating the swimming environment impacts low-order (e.g., temporal, position, velocity or acceleration parameters) and high-order (i.e., spatial-temporal coordination) variables. The average stroke cycle duration and the relative duration of the catch and glide phases were reduced in the flume trial, which was perceived as very intense, whereas the pull and push phases were longer. Of the four coordination patterns (in-phase, anti-phase, proximal and distal: when the appropriate segment is leading the coordination of the other), flume swimming demonstrated more in-phase coordination for the catch and glide (between hand and lower arm) and recovery (hand/lower arm and lower arm/upper arm couplings). Conversely, the variability of the spatial coordination was not significantly different between the two environments, implying that expert swimmers maintain consistent and stable coordination despite constraints and whatever the swimming resistances. Investigations over a wider range of velocities are needed to better understand coordination dynamics when the aquatic environment is modified by a swimming flume. Since the design of flumes impacts significantly the hydrodynamics and turbulences of the fluid flow, previous results are mainly related to the characteristics of the flume used in the present study (or a similar one), and generalization is subject to additional investigations.     

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.     

Adaptation to rearranged eye-foot coordination

The generality of adaptation following three types of movements during prism exposure was investigated. The three exposure conditions consisted of (1) walking with prisms, (2) viewing leg movements through prisms, and (3) viewing arm movements through prisms. The results showed that changes in eye-foot coordination and egocentric localization occur following both (1) and (2). Exposure Conditions 2 and 3 both produce changes in eye-hand coordination; however, eye-foot coordination was found unaltered following (3).     

Paradoxical decrease of imitation performance with age in children

Imitation development was studied in a cross-sectional design involving 174 primary-school children (aged 6–10), focusing on the effect of actions' complexity and error analysis to infer the underlying cognitive processes. Participants had to imitate the model's actions as if they were in front of a mirror (‘specularly’). Complexity varied across three levels: movements of a single limb; arm and leg of the same body side; or arm and leg of opposite body sides. While the overall error rate decreased with age, this was not true of all error categories. The rate of ‘side’ errors (using a limb of the wrong body side) paradoxically increased with age (from 9 years). However, with increasing age, the error rate also became less sensitive to the complexity of the action. This pattern is consistent with the hypothesis that older children have the working memory (WM) resources and the body knowledge necessary to imitate ‘anatomically’, which leads to additional side errors. Younger children might be paradoxically free from such interference because their WM and/or body knowledge are insufficient for anatomical imitation. Yet, their limited WM resources would prevent them from successfully managing the conflict between spatial codes involved in complex actions (e.g. moving the left arm and the right leg). We also found evidence that action side and content might be stored in separate short-term memory (STM) systems: increasing the number of sides to be encoded only affected side retrieval, but not content retrieval; symmetrically, increasing the content (number of movements) of the action only affected content retrieval, but not side retrieval. In conclusion, results suggest that anatomical imitation might interfere with specular imitation at age 9 and that STM storages for side and content of actions are separate.     

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.     

Postural fluctuations during pointing from a unilateral or bilateral stance

An experiment was conducted to compare the effects of bilateral and unilateral stance on postural fluctuations and intralimb coordination during active balance control. Fifteen participants stood bilaterally and unilaterally while conducting a pointing task with an outstretched arm. Excursion of center of foot pressure (CoP) and limb movements were recorded with a force plate and eight dual-axis accelerometers, respectively. Compared to bilateral stance, unilateral stance resulted in wider CoP trajectories and greater postural fluctuations, especially in the lower limbs. The limb-dependent postural fluctuations during unilateral stance were associated with an increased coupling between the upper limb segments and a decreased coupling between the segments of the stance leg. Unilateral stance further resulted in greater regularity and spectral changes in postural fluctuations of the trunk and lower limb due to increased central oscillations (8-15 Hz). The observed structural differences in postural fluctuations between unilateral and bilateral stance strongly suggested that the postural control system modulates joint stiffness in a stance-dependent manner. Probably, in unilateral stance, attentive control was shifted to the stance leg at the expense of increasing arm stiffness to reduce movement redundancy.     

Differences in spatial-temporal parameters and arm-leg coordination in butterfly stroke as a function of race pace, skill and gender

Spatial-temporal parameters (velocity, stroke rate, stroke length) and arm-leg coordination in the butterfly stroke were studied as a function of race pace, skill (due to technical level, age, and experience) and gender. Forty swimmers (ten elite men, ten elite women, ten less-skilled men, and ten less-skilled women) performed the butterfly stroke at four velocities corresponding to the appropriate paces for the 400-m, 200-m, 100-m, and 50-m, respectively. Arm and leg stroke phases were identified by video analysis and used to calculate four time gaps (T1: the time difference between the start of the arms' catch phase and the start of the legs' downward phase of the first leg kick; T2: the time difference between the start of the arms' pull phase and the start of the legs' upward phase of the first leg kick; T3: the time difference between the start of the arms' push phase and the start of the legs' downward phase of the second leg kick; and T4: the time difference between the start of the arms' recovery and the start of the legs' upward phase of the second leg kick) and the total time gap (TTG), i.e., the sum of the four discrete time gaps. These values described the changing coupling of arm to leg actions over an entire stroke cycle. A significant race pace effect indicated that the synchronization between the key motor points of the arms and legs, which determine the starts and ends of the arm and leg stroke phases, increased with pace for all participants. A significant skill effect indicated that the elite swimmers had greater velocity, stroke length, and stroke rate and stronger synchronization of the arm and leg stroke phases than the less-skilled swimmers, due to smaller T2 and T3 and greater T1. A significant gender effect revealed greater velocity and stroke length for the men, and smaller T1 for the less-skilled women. These time gap differences between skill levels were related to the capacity of elite swimmers to assume a more streamlined position of trunk, head and upper limbs during leg actions, adopt a shorter glide and higher stroke rate to overcome great forward resistance, and generate higher forces and use better technique during the arm pull. Thus, coaches are advised to begin monitoring arm-leg coordination earlier in swimmers' careers to ensure that they attain their highest possible skill levels.     

Saccadic output is influenced by limb kinetics during eye-hand coordination

In several recent studies, saccadic eye movements were found to be influenced by concurrent reaching movements. The authors investigated whether that influence originates in limb kinematic or kinetic signals. To dissociate those 2 possibilities, the authors required participants (N = 6) to generate pointing movements with a mass that either resisted or assisted limb motion. With practice, participants were able to generate pointing responses with very similar kinematics but whose kinetics varied in a systematic manner. The results showed that saccadic output was altered by the amount of force required to move the arm, consistent with an influence from limb kinetic signals. Because the interaction occurred before the pointing response began, the authors conclude that a predictive signal related to limb kinetics modulates saccadic output during tasks requiring eye-hand coordination.     

Horizontal plane head stabilization during locomotor tasks

Frequency characteristics of head stabilization were examined during locomotor tasks in healthy young adults(N = 8) who performed normal walking and 3 walking tasks designed to produce perturbations primarily in the horizontal plane. In the 3 walking tasks, the arms moved in phase with leg movement, with abnormally large amplitude, and at twice the frequency of leg movement. Head-in-space angular velocity was examined at the predominant frequencies of trunk motion. Head movements in space occurred at low frequencies (< 4.0 Hz) in all conditions and at higher frequencies (> 4.0 Hz) when the arms moved at twice the frequency of the legs. Head stabilization strategies were determined from head-on-trunk with respect to trunk frequency profiles derived from angular velocity data. During natural walking at low frequencies (< 3.0 Hz), head-on-trunk movement was less than trunk movement. At frequencies 3.0 Hz or greater, equal and opposite compensatory movement ensured head stability. When arm swing was altered, compensatory movement guaranteed head stability at all frequencies. Head stabilization was successful for frequencies up to 10.0 Hz during locomotor tasks. Maintaining head stability at high frequencies during voluntary tasks suggests that participants used feedforward mechanisms to coordinate head and trunk movements. Maintenance of head stability during dynamic tasks allows optimal conditions for vestibulo-ocular reflex function.     

Development of upper limb proprioceptive accuracy in children and adolescents

This study was designed to determine differences in the proprioceptively guided movements of children (8-10 years) and adolescents (16-18 years). Participants were blindfolded and asked to actively match passively determined target positions of the elbow joint under three matching conditions. Overall, children were less accurate than adolescents in all matching tasks and utilized different kinematic strategies for making the matching movements. Specifically, children made larger absolute errors and utilized matching movements which, compared to adolescents, were of shorter duration and less irregular in terms of their velocity profiles. An assessment of limb asymmetry was also performed revealing a non-dominant arm matching advantage but only for children and only in the task requiring interhemispheric transfer of a memory-based model of limb position. The proprioceptive differences observed in this study are likely the result of experience-driven refinement in the utilization of somatosensory feedback throughout childhood and into adolescence.     

Rapid aimed limb movements: differential effects of practice on component submovements

Three experiments are reported in which subjects practiced rapid aimed limb movements (arm pointing and wrist rotation) toward a visible target region. Subjects were required to minimize their movement durations while still landing in the target. The movement trajectories were examined to assess the effects of practice on separate component submovements of the limb movements. The results revealed that practice improved primarily temporal, not spatial, aspects of performance. Practice reduced the overall movement durations, but had different effects on the individual submovements. Practice allowed subjects to reduce the amount of time spent performing final corrective submovements, but actually increased slightly the time needed to produce the initial ballistic submovement. The results suggest that practice in the present task primarily enhanced the ability to use feedback information, but there was also some evidence of changes in the ballistic, preprogrammed portion of the movements. The results demonstrate that analysis of submovements can reveal important details of the underlying motor control processes.     

Adaptation to visual displacement with active and passive limb movements: effect of movement frequency and predictability of movement

Three experiments were performed to evaluate the influence of active and passive limb movements on adaptation to visual displacement. Over a wide frequency range (0·5-1·25 Hz) with constant amplitude, 30°, significant adaptation was achieved with active and passive movements. When arm movement frequency was constant at 1·0 Hz but amplitude of movement was varied, less adaptation was achieved for both active and passive movements than when amplitude was held constant. Even at a frequency above that of most naturally occurring limb movements, 1·67 Hz, and with variable amplitude motion, significant adaptation was achieved with active and passive limb movements. These findings emphasize the importance of visual-proprioceptive discordances for adaptation to visual displacement when only sight of the hand is permitted. Significant differences did not appear between the active and passive movement conditions in any of the experiments.     

Upper limb movement analysis during gait in multiple sclerosis patients

PurposeGait disorders in multiple sclerosis (MS) are well studied; however, no previous study has described upper limb movements during gait. However, upper limb movements have an important role during locomotion and can be altered in MS patients due to direct MS lesions or mechanisms of compensation. The aim of this study was to describe the arm movements during gait in a population of MS patients with low disability compared with a healthy control group.MethodsIn this observational study we analyzed the arm movements during gait in 52 outpatients (mean age: 39.7 ± 9.6 years, female: 40%) with relapsing-remitting MS with low disability (mean EDSS: 2 ± 1) and 25 healthy age-matched controls using a 3-dimension gait analysis.ResultsMS patients walked slower, with increased mean elbow flexion and decreased amplitude of elbow flexion (ROM) compared to the control group, whereas shoulder and hand movements were similar to controls. These differences were not explained by age or disability.ConclusionUpper limb alterations in movement during gait in MS patients with low disability can be characterized by an increase in mean elbow flexion and a decrease in amplitude (ROM) for elbow flexion/extension. This upper limb movement pattern should be considered as a new component of gait disorders in MS and may reflect subtle motor deficits or the use of compensatory mechanisms.     

Horizontal Plane Head Stabilization During Locomotor Tasks

Frequency characteristics of head stabilization were examined during locomotor tasks in healthy young adults (N = 8) who performed normal walking and 3 walking tasks designed to produce perturbations primarily in the horizontal plane. In the 3 walking tasks, the arms moved in phase with leg movement, with abnormally large amplitude, and at twice the frequency of leg movement. Head-in-space angular velocity was examined at the predominant frequencies of trunk motion. Head movements in space occurred at low frequencies (< 4.0 Hz) in all conditions and at higher frequencies (> 4.0 Hz) when the arms moved at twice the frequency of the legs. Head stabilization strategies were determined from head-on-trunk with respect to trunk frequency profiles derived from angular velocity data. During natural walking at low frequencies (< 3.0 Hz), head-on-trunk movement was less than trunk movement. At frequencies 3.0 Hz or greater, equal and opposite compensatory movement ensured head stability. When arm swing was altered, compensatory movement guaranteed head stability at all frequencies. Head stabilization was successful for frequencies up to 10.0 Hz during locomotor tasks Maintaining head stability at high frequencies during voluntary tasks suggests that participants used feedforward mechanisms to coordinate head and trunk movements. Maintenance of head stability during dynamic tasks allows optimal conditions for vestibulo-ocular reflex function.     

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.