Interlimb Interactions Research Articles

Distinct contribution of interneuronal circuits in the human motor cortex during interlimb interactions: A transcranial magnetic stimulation study

Distinct contribution of interneuronal circuits in the human motor cortex during interlimb interactions: A transcranial magnetic stimulation study

Characterization of interlimb interaction via transcutaneous spinal stimulation of cervical and lumbar spinal enlargements.

The use of transcutaneous electrical spinal stimulation (TSS) to modulate sensorimotor networks after neurological insult has garnered much attention from both researchers and clinicians in recent years. Although many different stimulation paradigms have been reported, the interlimb effects of these neuromodulation techniques have been little studied. The effects of multisite TSS on interlimb sensorimotor function are of particular interest in the context of neurorehabilitation, as these networks have been shown to be important for functional recovery after neurological insult. The present study utilized a condition-test paradigm to investigate the effects of interenlargement TSS on spinal motor excitability in both cervical and lumbosacral motor pools. Additionally, comparison was made between the conditioning effects of lumbosacral and cervical TSS and peripheral stimulation of the fibular nerve and ulnar nerve, respectively. In 16/16 supine, relaxed participants, facilitation of spinally evoked motor responses (sEMRs) in arm muscles was seen in response to lumbosacral TSS or fibular nerve stimulation, whereas facilitation of sEMRs in leg muscles was seen in response to cervical TSS or ulnar nerve stimulation. The decreased latency between TSS- and peripheral nerve-evoked conditioning implicates interlimb networks in the observed facilitation of motor output. The results demonstrate the ability of multisite TSS to engage interlimb networks, resulting in the bidirectional influence of cervical and lumbosacral motor output. The engagement of interlimb networks via TSS of the cervical and lumbosacral enlargements represents a feasible method for engaging spinal sensorimotor networks in clinical populations with compromised motor function.NEW & NOTEWORTHY Bidirectional interlimb modulation of spinal motor excitability can be evoked by transcutaneous spinal stimulation over the cervical and lumbosacral enlargements. Multisite transcutaneous spinal stimulation engages spinal sensorimotor networks thought to be important in the recovery of function after spinal cord injury.

Foot clearance when crossing obstacles of different heights with the lead and trail limbs.

In order to predict and prevent falls and fall-related injuries, it is crucial to understand the motor control for crossing obstacles. In real life, since obstacles do not always take regular shapes like rectangles, the lead and trail limbs sometimes need to negotiate different obstacle heights. The interlimb interaction in this process has remained unknown, since obstacle crossing studies commonly use a single-obstacle paradigm in which the obstacle height is the same for the lead and trail limbs. We used a dual-obstacle paradigm to test whether the foot clearance over one obstacle was influenced by the contralateral obstacle's height. Sixteen healthy young male and female participants (age: 22.5 ± 1.9 years) crossed over two obstacles placed side by side. Four obstacle conditions were made by combining obstacles of two heights (low, L, 9.0 cm; high, H, 22.5 cm) of the obstacles.In the LL condition, both obstacles were low, and in the LH condition, there was a low obstacle for the lead limb and a high one for the trail limb. Similarly, we also arranged HL and HH conditions. Each subject performed 20 trials per condition. We compared the vertical foot clearance, prestep distance, and poststep distance between the conditions. The foot trajectory to step over the obstacles were affected by the contralateral obstacle's height. The vertical foot clearance of the trail limb was greater in the HL condition than in the LL condition. The vertical foot clearance of the lead limb was greater in the LH condition than in the LL condition. The results suggest that the foot trajectory was not determined exclusively by the obstacle to be crossed. Instead, comprehensive information, including the height of the obstacle for the other limb, might be used for motor control during obstacle crossing.

Cortical and Subcortical Neural Interactions Between Trunk and Upper-limb Muscles in Humans

Activities of daily living require simultaneous and coordinated activation of trunk and upper-limb segments, which involves complex interlimb interaction within the central nervous system. Although many studies have reported associations between activity of trunk and limb muscles during functional tasks, evidence on cortical and subcortical contributions to trunk-limb neural interactions is still not fully clear. Therefore, the aim of this study was to examine interactions between trunk and upper-limb muscles in the: (i) corticospinal circuits by using motor evoked potential (MEP) elicited through transcranial magnetic stimulation; and (ii) subcortical circuits by using cervicomedullary motor evoked potential (CMEP) elicited through cervicomedullary junction magnetic stimulation. Responses were evoked in the erector spinae (trunk) and flexor carpi radialis (upper-limb) muscles in twelve able-bodied individuals: (1) while participants were relaxed; (2) during trunk muscle contractions while arms were at rest; and (3) during upper-limb muscle contractions while the trunk was at rest. Our results showed that trunk muscle CMEP responses were not affected by upper-limb muscle contractions, while MEP responses were modulated. This indicates that at least the subcortical circuits may not attribute to facilitation of the trunk muscles during upper-limb contractions. On the other hand, in the upper-limb muscles, both CMEP and MEP responses were modulated during trunk contractions. These results indicate that cortical and subcortical mechanisms attributed to facilitation of upper-limb muscles during trunk contractions. In conclusion, our study demonstrated evidence that trunk-limb neural interactions may be attributed to cortical and/or subcortical mechanisms depending on the contracted muscle.

Contralateral seventh cervical nerve transfer can affect the pennation angle of the lower limb in spastic hemiplegia patients: An observational case series study.

IntroductionWe previously reported transferring seventh cervical (C7) nerve from unaffected side to affected side in patients with spastic hemiplegia due to chronic cerebral injury, to improve function and reduce spasticity of paralyzed upper limb. In the clinics, some patients also reported changes of spasticity in their lower limb, which could not be detected by routine physical examinations. Pennation angle of muscle can indirectly reflect the condition of spasticity. The purpose of this study was to evaluate whether this upper limb procedure may affect spasticity of lower limb, using ultrasonography to detect changes of muscle pennation angle (PA).MethodsTwelve spastic hemiplegia patients due to cerebral injury including stroke, cerebral palsy, and traumatic brain injury, who underwent C7 nerve transfer procedure, participated in this study. B‐mode ultrasonography was used to measure PA of the gastrocnemius medialis (GM) muscle at rest preoperatively and postoperatively. The plantar load distribution of the lower limbs was evaluated using a Zebris FDM platform preoperatively and postoperatively.ResultsThe PA of the GM was significantly smaller on the affected side than that of unaffected side before surgery. On the affected side, the postoperative PA was significantly larger than preoperative PA. On the unaffected side, the postoperative PA was not significantly different compared to preoperative PA. The postoperative plantar load distribution of the affected forefoot was significantly smaller than preoperative load distribution, which was consistent with ultrasonography results.ConclusionsThis study indicates that C7 nerve transfer surgery for improving upper limb function can also affect muscle properties of lower limb in spastic hemiplegia patients, which reveals a link between the upper and lower limbs. The interlimb interactions should be considered in rehabilitation physiotherapy, and the regular pattern and mechanism need to be further studied.

Evidence for existence of trunk-limb neural interaction in the corticospinal pathway

In humans, trunk muscles have an essential role in postural control as well as walking. However, little is known about the mechanisms of interaction with different muscles, especially related to how trunk muscles interact with the limbs. Contraction of muscles can modulate the corticospinal excitability not only of the contracted muscle, but also of other muscles even in the remote segments of the body. However, “remote effect” mechanism has only been examined for inter-limb interactions. The aim of our current study was to test if there are trunk-limb interactions in the corticospinal pathways. We examined corticospinal excitability of: (a) trunk muscles at rest when hands, legs and jaw muscles were contracted and; (b) hand, leg, and jaw muscles at rest when trunk muscles were contracted. We measured motor evoked potentials elicited using transcranial magnetic stimulation in the rectus abdominis, flexor digitorum superficialis, masseter, tibialis anterior muscles under the following experimental conditions: (1) participants remained relaxed (Rest); (2) during trunk contraction (Trunk); (3) during bilateral hand clenching (Hands); (4) during jaw clenching (Jaw); and (5) during bilateral ankle dorsiflexion (Legs). Each condition was performed at three different stimulation intensities and conditions were randomized between participants. We found that voluntary contraction of trunk muscle facilitated the corticospinal excitability of upper-limb and lower-limb muscles during rest state. Furthermore, voluntary contraction of upper-limb muscle also facilitated the corticospinal excitability of trunk muscles during rest state. Overall, these results suggest the existence of trunk-limb interaction in the corticospinal pathway, which is likely depended on proximity of the trunk and limb representation in the motor cortex.

Are interlimb interactions disturbed in patients with Parkinson’s disease? A study under unloading conditions

During natural human locomotion neural connections characteristic for the control of quadrupedal walking are activated. The degree of interaction between the neural networks generating rhythmic movements of the upper and lower extremities, depends on the tonic state of each of these networks, adjustable by motor command in the brain. The distortion of such command in patients with Parkinson's disease (PD) may lead to disturbance of interlimb interactions. In conditions of arms and legs unloading in 17 patients with PD and 16 healthy subjects investigated the effect of limb movement of one girdle on the parameters of motor activity of the limbs of the other girdle in their combined cyclic movements. In patients the weakening of influences of active or passive arm movements, and active movements of the distal parts of the arms on voluntary leg movements was shown, while in healthy subjects the effect of arm movements on the parameters of voluntary movements was significant. The effect of arm movements on the possibility of involuntary activation of air- stepping by vibratory stimulation of the leg muscles in patients was absent, while in healthy subjects the motor activity of the arms increased the opportunity of involuntary rhythm activation. The differences in the influence of leg movements to the rhythmic movement of the upper extremities in patients and healthy subjects were revealed. The medication exhibited interlimb interaction, but it was insufficient for normalization of the state of neural networks in patients. It is concluded that in patients with PD, neural networks generating stepping rhythm are in a state of heightened tonic activity. This does not allow not only activating involuntary rhythmic movements, but also showing facilitation effect with arms to legs.

Activation of interlimb interactions increases the motor output in legs of healthy subjects: Study under the conditions of arm and leg unloading

We studied the effect of arm movements and movements of separate arm joints on the electrophysiological and kinematic characteristics of voluntary and vibration-triggered stepping-like leg movements under the conditions of horizontal support of upper and lower limbs. The horizontal support of arms provided a significantly increase in the rate of activation of locomotor automatism by non-invasive impact on tonic sensory inputs. The addition of active arm movements during involuntary rhytmic stepping-like leg movements led to an increase in EMG activity of hip muscles and was accompanied by an increase in the amplitude of hip and shin movements. Passive arm movements had the same effect on induced leg movements. The movement of the shoulder joints led to an increase in the activity of hip muscles and an increase in the amplitude of movements of the knee and hip joints. At the same time, the movement of forearms. and wrists had similar facilitating effect on electrophysiological and kinematic characteristics of rhytmic stepping-like movements, but influenced the distal segments of legs to a greater extent. Under the conditions of sub-threshold vibration of leg muscles, voluntary arm movements led to the activation of involuntary rhytmic stepping movements. During voluntary leg movements, the addition of arm movements had a significantly smaller impact on the parameters of rhytmic stepping than during involuntary leg movements. Thus, the simultaneous movements of upper and lower limbs are an effective method of activation of neural networks connecting the rhythm generators of arms and legs. Under the conditions of arm and leg unloading, the interactions between the cervical and lumbosacral segments of the spinal cord seem to play the major role in the impact of arm movements on the patterns of leg movements. The described methods of activation of interlimb interactions can be used in the rehabilitation of post-stroke patients and patients with spinal cord injuries, Parkinson's disease and other neurological diseases.

Interlimb interactions during cyclic in-phase and antiphase movements of arms and legs and their dependence on afferent influences

Coordinated movements of arms and legs suppose the neural interaction between the generators of the rhythmics of the upper and lower extremities. In the lying position in 10 healthy subjects activity of the muscles of the upper and lower extremities was recorded when separate and joint cyclic movements of the arms and legs with different phase relationships between the movements of the limbs were performed, and under various conditions of the motor task. Antiphased active arm movements were characterized by increased muscle activity than during in-phase mode. The activity of the arm muscles under passive movements, imposed by experimenter, was significantly less than their activity when passive movements of the arms were imposed by the other arm. When loading one arm the muscle activity in the other passively moving arm increased independently from the synergy of arm movements. During motor tasks, implementing joint antiphased movements as the upper and lower extremities, compared to motor task, implementing their joint in-phase movements, there has been a significant increase in activity in the biceps brahii muscle, the tibialis anterior muscle and biceps femoris muscle. Loading of arms in these motor tasks has been accompanied by increased activity in some of the leg muscles. Increasing of frequency of rhythmic movements resulted in a significant growth of the muscle activity of the arms and legs with their cooperative movements with greater rate of rise of flexor muscles activity for arms and legs during joint antiphased movements. Thus, the spatial organization of movements and kind of afferent influences are significant factors of interlimb interaction, which, in turn, determine the type of neural interconnections that are involved in the regulation of movements.

Inter-limb force coupling is resistant to distorted visual feedback in chronic hemiparetic stroke

Interlimb coupling between impaired and non-impaired limbs after stroke has been a common observation. The aim of this study was to examine interlimb interactions in force production in responses to altered visual gain in hemiparetic stroke survivors. prospective clinical study A convenient sample of 7 hemiparetic stroke subjects (3 women and 4 men; mean age 56.0 years (standard errors 12.8) of age; history of stroke: mean duration 61.6 months (standard errors 53.3)) participated in the study. Subjects performed bilateral elbow flexion to varying total force targets from 3% to 60% maximal contraction forces with normal visual gain (1:1) and to a 10% maximal voluntary contraction target with altered visual gains (1/8, 1/4, 1/2, 2, 4, and 8) for the force of the less-impaired, ipsilesional side. Across all conditions, the forces produced by both impaired and non-impaired limb changed proportionally to their maximal voluntary contraction force, such that relative contributions of each limb's force to the total force remained unchanged. In conditions with altered visual gain, high and low, the total force showed errors in the direction of under-shooting. Our findings indicate that there is a strong interlimb force coupling in hemiparetic stroke, resistant to distorted visual feedback. It may reflect a default sharing pattern dominant after stroke.

Interlimb interactions during bilateral voluntary elbow flexion tasks in chronic hemiparetic stroke

The purpose was to systematically investigate interlimb interactions in chronic hemiparetic stroke. Fourteen poststroke hemiparetic subjects (>1 year) performed maximum voluntary contraction (MVC) elbow flexion tasks without visual feedback with one (unilateral) and two limbs simultaneously (bilateral). At submaximal levels, subjects produced force to a visual target reflecting 20%, 40%, 60%, and 80% of corresponding MVC in unilateral tasks, and of summated unilateral MVCs in bilateral tasks. Elbow flexion force and biceps surface electromyogram (EMG) were measured bilaterally. Proportionally increased EMG activity on the contralateral limb (motor overflow) was observed during unilateral tasks of the nonimpaired limb but not of the impaired limb. During bilateral tasks at submaximal levels, the impaired limb produced less force (i.e., force deficit [FD]) as compared to expected forces based upon its unilateral MVC. Force deficit on the impaired limb was compensated by greater force production on the nonimpaired limb such that the visual target was reached. However, force contribution to the total force progressively decreased from the nonimpaired side, when the level of submaximal contractions increased. During bilateral MVC tasks, there was no FD on the impaired limb, but FD was observed on the nonimpaired limb. A net result of a small bilateral deficit in force with parallel changes in EMG was observed. These novel findings of activation level–dependent interactions and asymmetrical contralateral motor overflow provide new insights that, among other compensatory mechanisms, ipsilateral corticospinal projections from the nonlesioned hemisphere play an important role in interlimb interactions in chronic stroke, in addition to unbalanced interhemispheric inhibition.

Learning a New Bimanual Coordination Pattern: Interlimb Interactions, Attentional Focus, and Transfer

ABSTRACT Because bimanual coordinative stability is governed by interlimb coupling, we examined how learning a new pattern (90°) was reflected in changes in the underlying interlimb interactions. Three interlimb interaction sources were distinguished: integrated timing of feedforward control signals, error corrections based on perceived relative phase, and phase entrainment by contralateral afference. By comparing 4 tasks that involve these interactions to a different extent, changes in the stabilizing contributions of these coupling sources could be studied. Furthermore, we studied how the learning process and changes in the underlying interactions were influenced by attentional focus (internal vs. external), and we examined retention of the learned pattern and transfer to the mirror-symmetrical pattern (270°). Results showed that stability and accuracy of the new pattern increased significantly with learning, due to improved integrated timing and error correction. Integrated timing improved first, possibly providing a reference frame for the error corrections that subsequently became more effective. Despite some qualitative differences in the learning process, neither performance of the learned pattern nor the underlying interlimb interactions was influenced by attentional focus. Whereas the learned pattern improved directly after practice, transfer followed later, suggesting that a more general representation was formed at a slower rate after practice.

Development of Temporal and Spatial Bimanual Coordination During Childhood

Developmental changes in bimanual coordination were examined in four age groups: 6/7, 10/11, 14/15 years, and young adults. Temporal coupling was assessed through the stabilizing contributions of interlimb interactions related to planning, error correction, and reflexes during rhythmic wrist movements, by comparing various unimanual and bimanual tasks involving passive and active movements. Spatial coupling was assessed via bimanual line-circle drawing. With increasing age, temporal stability improved. Relative contributions of planning and reflex interactions to the achieved stability did not change, whereas error correction improved. In-phase and antiphase coordination developed at similar rates; implications of this result were discussed in terms of mirror-activity inhibition. Overall spatial drawing performance (circularity, variability, smoothness) improved with age, and spatial interference was smaller in adults than children. Whereas temporal coupling increased from 6/7 years to adulthood, spatial coupling changed mainly after 14/15 years. This difference in the development of temporal and spatial coupling corresponds to the anterior-posterior direction of corpus callosum myelination as reported in the literature.

Abnormalities of mutual influences of the upper and lower limbs after a stroke

The common pattern of muscle activation and specifics of interlimb neuronal connections during the performance of rhythmic separate and simultaneous arm and leg movements in the lying position in healthy subjects, which reflected functionally significant interlimb neuronal interactions, were shown. The study was designed to investigate these mutual influences of the upper and lower limbs during the performance of similar motor tasks by stroke patients. Sixteen poststroke patients with different degrees of hemiparesis performed active and passive arm movements simultaneously with stepping leg movements or without them while lying supine. It was demonstrated that the patients had a disordered common pattern of distribution of muscle activity when they performed voluntary cyclic movements with both arms. Passive movements of both paretic and nonparetic arms led to different degrees of activation of their muscles, depending on the degree of paresis: in patients with mild paresis, muscle activation was similar to that in healthy subjects; in patients with severe paresis, it was insignificant. The loading of the nonparetic arm resulted in an increase in the activity in the paretic arm shoulder flexor muscles in patients with mild paresis (which was typical of healthy subjects), while loading did not influence significantly patients with severe paresis. The combination of cyclic arm movements and stepping leg movements in diagonal synergy decreased the activity in the proximal muscles of both arms, irrespective of the degree of paresis, as it was observed in healthy subjects. Simultaneous arm and leg movements did not change the muscle activity in nonparetic legs in either groups of patients, but the activity in the paretic leg muscles even decreased. The results obtained revealed important features of poststroke motor disturbances, which caused changes in interlimb interactions and largely depended on the degree of paresis. The data could be useful for developing new methods for the performance of rehabilitative procedures in poststroke patients.

What Has Transcranial Magnetic Stimulation Taught Us About Neural Adaptations To Strength Training? A Brief Review

The evidence for neural mechanisms underpinning rapid strength increases has been investigated and discussed for over 30 years using indirect methods, such as surface electromyography, with inferences made toward the nervous system. Alternatively, electrical stimulation techniques such as the Hoffman reflex, volitional wave, and maximal wave have provided evidence of central nervous system changes at the spinal level. For 25 years, the technique of transcranial magnetic stimulation (TMS) has allowed for noninvasive supraspinal measurement of the human nervous system in a number of areas such as fatigue, skill acquisition, clinical neurophysiology, and neurology. However, it has only been within the last decade that this technique has been used to assess neural changes after strength training. The aim of this brief review is to provide an overview of TMS, discuss specific strength training studies that have investigated changes, after short-term strength training in healthy populations in upper and lower limbs, and conclude with further research suggestions and the application of this knowledge for the strength and conditioning coach.

Frequency-induced changes in interlimb interactions: Increasing manifestations of closed-loop control

In bimanual coordination, interactions between the limbs result in attraction to in-phase and antiphase coordination. Increasing movement frequency leads to decreasing stability of antiphase coordination, often resulting in a transition to the more stable in-phase pattern. It is unknown, however, how this frequency-induced loss of stability is engendered in terms of the interlimb interactions underwriting bimanual coordination. The present study was conducted to help resolve this issue. Using an established method (based on comparison of various unimanual and bimanual tasks involving both passive and active movements), three sources of interlimb interaction were dissociated: (1) integrated timing of feedforward signals, (2) afference-based correction of relative phase errors, and (3) phase entrainment by contralateral afference. Results indicated that phase entrainment strength remained unaffected by frequency and that the stabilizing effects of error correction and integrated timing decreased with increasing frequency. Their contributions, however, reflected an interesting interplay as frequency increased. For moderate frequencies coordinative stability was predominantly secured by integrated timing processes. However, at high frequencies, the stabilization of the antiphase pattern required combined contributions of both integrated timing and error correction. In sum, increasing frequency was found to induce a shift from predominantly open-loop control to more closed-loop control. The results may be accounted for by means of an internal forward model for sensorimotor integration in which the sensory signals are compared to values predicted on the basis of efference copies.

Sensory afferent inhibition within and between limbs in humans

To examine the distribution and inter-limb interaction of short-latency afferent inhibition (SAI) in the arm and leg. Motor evoked potentials (MEPs) in distal and proximal arm, shoulder and leg muscles induced with ranscranial magnetic stimulation (TMS) were conditioned by painless electrical stimuli applied to the index finger (D2) and great toe (T1) at interstimulus intervals (ISIs) of 15, 25-35, 80 ms (D2) and 35, 45, 55, 65 and 100 ms (T1) in 27 healthy human subjects. TMS was delivered over primary motor cortex (M1) arm and leg areas. Electrical stimulus intensities were varied between 1 and 3 times the sensory perception thresholds. We also tested effects of posterior cutaneous brachial nerve (PCBN) stimulation on MEPs in arm muscles at ISIs of 18 and 28 ms. D2 but not PCBN electrical conditioning reduced MEP amplitudes in upper limb muscles at ISIs of 25 and 35 ms. SAI was more pronounced in distal as compared to proximal arm muscles. Also, SAI following D2 stimulation increased with higher conditioning intensities. D2 stimulation did not change lower limb muscles MEPs. In contrast, T1 stimulation did not induce SAI in any muscles but caused MEP facilitation in a foot muscle at an ISI of 55 ms and in upper limb muscles at ISIs of 35 and 55 ms. Short interval intracortical inhibition (SICI) and intracortical facilitation (ICF) were not affected by electrical T1 conditioning. D2 stimulation causes segmental SAI in upper limb muscles with a distal to proximal attenuation without affecting leg muscles. In contrast, toe stimulation facilitates motor output both in foot and upper arm muscles. Our data suggest that cutaneo-motor pathways in arms and legs are functionally organized in a different way with cutaneo-motor interactions induced by toe stimulation probably relayed at a thalamic level. Abnormal cutaneo-motor interactions following electrical toe stimulation may serve as an electrophysiological marker of thalamic dysfunction, e.g. in neurodegenerative diseases.

Attentional loads associated with interlimb interactions underlying rhythmic bimanual coordination

Studies of rhythmic bimanual coordination under dual-task conditions revealed (1) a dependence of secondary task performance on the stability of coordinative tasks, in that secondary task performance was better during in-phase than antiphase coordination, and (2) a shift in the mean relative phasing between the limbs compared to single-task conditions. The present study aimed to account for these phenomena by dissociating three qualitatively different interactions between the limbs that govern this motor behavior, related to movement planning, error correction, and interlimb reflex activity. The experiment probed the cognitive demands associated with each interlimb interaction by examining the attentional load under dual-task conditions, indexed by reaction times of the secondary task and kinematic changes in the coordinative tasks relative to single-task conditions. First, only in the condition that involved interlimb interactions at the level of movement planning reaction times were shorter for in-phase than for antiphase coordination, highlighting an intimate relation between movement planning and attentional processes. Second, under dual-task conditions a shift in the mean relative phase was observed relative to single-task conditions, but only for the interlimb interactions that depend directly on sensory feedback (error correction and interlimb reflex activity). These observations qualified the effects of attentional load reported in previous studies. Third, reaction times varied systematically over the movement cycle. These variations revealed a dynamical signature of the attentional load that differed between the three interlimb interactions.

Interlimb coupling strength scales with movement amplitude

The relation between movement amplitude and the strength of interlimb interactions was examined by comparing bimanual performance at different amplitude ratios (1:2, 1:1, and 2:1). For conditions with unequal amplitudes, the arm moving at the smaller amplitude was predicted to be more strongly affected by the contralateral arm than vice versa. This prediction was based on neurophysiological considerations and the HKB model of coupled oscillators. Participants performed rhythmic bimanual forearm movements at prescribed amplitude relations. After a brief mechanical perturbation of one arm, the relaxation process back to the initial coordination pattern was examined. This analysis focused on phase adaptations in the unperturbed arm, as these reflect the degree to which the movements of this arm were affected by the coupling influences stemming from the contralateral (perturbed) arm. The thus obtained index of coupling (IC) reflected the relative contribution of the unperturbed arm to the relaxation process. As predicted IC was larger when the perturbed arm moved at a larger amplitude than did the unperturbed arm, indicating that coupling strength scaled with movement amplitude. This result was discussed in relation to previous research regarding sources of asymmetry in coupling strength and the effects of amplitude disparity on interlimb coordination.

Unraveling Interlimb Interactions Underlying Bimanual Coordination

Three sources of interlimb interactions have been postulated to underlie the stability characteristics of bimanual coordination but have never been evaluated in conjunction: integrated timing of feedforward control signals, phase entrainment by contralateral afference, and timing corrections based on the perceived error of relative phase. In this study, the relative contributions of these interactions were discerned through systematic comparisons of five tasks involving rhythmic flexion-extension movements about the wrist, performed bimanually (in-phase and antiphase coordination) or unimanually with or without comparable passive movements of the contralateral hand. The main findings were the following. 1) Contralateral passive movements during unimanual active movements induced phase entrainment to interlimb phasing of either 0 degrees (in-phase) or 180 degrees (antiphase). 2) Entrainment strength increased with the passive movements' amplitude, but was similar for in-phase and antiphase movements. 3) Coordination of unimanual active movements with passive movements of the contralateral hand (kinesthetic tracking) was characterized by similar bilateral EMG activity as observed in active bimanual coordination. 4) During kinesthetic tracking the timing of the movements of the active hand was modulated by afference-based error corrections, which were more pronounced during in-phase coordination. 5) Indications of in-phase coordination being more stable than antiphase coordination were most prominent during active bimanual coordination and marginal during kinesthetic tracking. Together the results indicated that phase entrainment by contralateral afference contributed equally to the stability of in-phase and antiphase coordination, and that differential stability of these patterns depended predominantly on integrated timing of feedforward signals, with only a minor role for afference-based error corrections.