Organization Of Muscle Synergies Research Articles

EFFECTS OF SPINAL TRANSECTION AND LOCOMOTOR SPEED ON MUSCLE SYNERGIES OF THE CAT HINDLIMB.

It was suggested that during locomotion, the nervous system controls movement by activating groups of muscles, or muscle synergies. Analysis of muscle synergies can reveal the organization of spinal locomotor networks and how it depends on the state of the nervous system, such as before and after spinal cord injury, and on different locomotor conditions, including a change in speed. The goal of this study was to investigate the effects of spinal transection and locomotor speed on hindlimb muscle synergies and their time-dependent activity patterns in adult cats. EMG activities of 15 hindlimb muscles were recorded in 9 adult cats of either sex during tied-belt treadmill locomotion at speeds of 0.4, 0.7, and 1.0 m/s before and after recovery from a low thoracic spinal transection. We determined EMG burst groups using cluster analysis of EMG burst onset and offset times and muscle synergies using non-negative matrix factorization. We found five major EMG burst groups and five muscle synergies in each of six experimental conditions (2 states x 3 speeds). In each case, the synergies accounted for at least 90% of muscle EMG variance. Both spinal transection and locomotion speed modified subgroups of EMG burst groups and the composition and activation patterns of selected synergies. However, these changes did not modify the general organization of muscle synergies. Based on the obtained results, we propose an organization for a pattern formation network of a two-level central pattern generator that can be tested in neuromechanical simulations of spinal circuits controlling cat locomotion.

SPATIO-TEMPORAL PATTERNS OF CORTICAL-MUSCULAR INTERACTION DURING LOCOMOTION

The paper considers the synergetic effects manifested at the cortical and muscular levels during locomotor activity showed in conditions of horizontal hanging of the lower extremities. The analysis of the synergies’ spatio-temporal structure was carried out using the data matrix factorization methods. It was found out the control of the movements’ structure is mainly realized through the three muscle synergies. The activity synchronization of the motor, associative, visual and sensorimotor regions of the cortex bilateral part is due to the specifics of performing locomotion in conditions of gravitational unloading and the associated features of receptor signaling. The identified components indicating synchronization of different areas of the right and left cortex may reflect the control processes associated with the control of alternate activation of the flexor and extensor muscles of the contralateral limb during locomotion. The data on the cortical activity spatial-temporal structuring indicate the separate management of muscle synergies through synchronization of cortical commands and the temporary organization of muscle synergies in the frequency ranges 0.30 to 8.00 Hz. Such patterns may reflect the activity of the rhythm-generating mechanism involved in the management of cyclic locomotor activity.

Sex influence on muscle synergies in a ballistic force-velocity test during the delayed recovery phase after a graded endurance run

The acute and delayed phases of the functional recovery pattern after running exercise have been studied mainly in men. However, it seems that women are less fatigable and/or recover faster than men, at least when tested in isometric condition. After a 20 km graded running race, the influence of sex on the delayed phase of recovery at 2–4 days was studied using a horizontal ballistic force-velocity test. Nine female and height male recreational runners performed maximal concentric push-offs at four load levels a week before the race (PRE), 2 and 4 days (D2 and D4) later. Ground reaction forces and surface electromyographic (EMG) activity from 8 major lower limb muscles were recorded. For each session, the mechanical force-velocity-power profile (i.e. theoretical maximal values of force (F¯ 0), velocity (V¯ 0), and power (P¯ max)) was computed. Mean EMG activity of each recorded muscle and muscle synergies (three for both men and women) were extracted. Independently of the testing sessions, men and women differed regarding the solicitation of the bi-articular thigh muscles (medial hamstring muscles and rectus femoris). At mid-push-off, female made use of more evenly distributed lower limb muscle activities than men. No fatigue effect was found for both sexes when looking at the mean ground reaction forces. However, the force-velocity profile varied by sex throughout the recovery: only men showed a decrease of both V¯ 0 (p < 0.05) and P¯ max (p < 0.01) at D2 compared to PRE. Vastus medialis activity was reduced for both men and women up to D4, but only male synergies were impacted at D2: the center of activity of the first and second synergies was reached later. This study suggests that women could recover earlier in a dynamic multi-joint task and that sex-specific organization of muscle synergies may have contributed to their different recovery times after such a race.

Stroke impairs the control of isometric forces and muscle activations in the ipsilesional arm

Stroke often impairs the control of the contralesional arm, thus most survivors rely on the ipsilesional arm to perform daily living activities that require an efficient control of movements and forces. Whereas the ipsilesional arm is often called ‘unaffected’ or ‘unimpaired’, several studies suggested that during dynamic tasks its kinematics and joint torques are altered. Is stroke also affecting the ability of the ipsilesional arm to produce isometric force, as when pushing or pulling a handle? Here, we address this question by analyzing behavioral performance and muscles’ activity when subjects applied an isometric force of 10 N in eight coplanar directions. We found that stroke affected the ability to apply well-controlled isometric forces with the ipsilesional arm, although to a minor extent compared to the contralesional arm. The spinal maps, the analysis of single muscle activities and the organization of muscle synergies highlighted that this effect was mainly associated with abnormal activity of proximal muscles with respect to matched controls, especially when pushing or pulling in lateral directions.

Common and distinct muscle synergies during level and slope locomotion in the cat.

Although it is well established that the motor control system is modular, the organization of muscle synergies during locomotion and their change with ground slope are not completely understood. For example, typical reciprocal flexor-extensor muscle synergies of level walking in cats break down in downslope: one-joint hip extensors are silent throughout the stride cycle, whereas hindlimb flexors demonstrate an additional stance phase-related electromyogram (EMG) burst (Smith JL, Carlson-Kuhta P, Trank TV. J Neurophysiol 79: 1702-1716, 1998). Here, we investigated muscle synergies during level, upslope (27°), and downslope (-27°) walking in adult cats to examine common and distinct features of modular organization of locomotor EMG activity. Cluster analysis of EMG burst onset-offset times of 12 hindlimb muscles revealed five flexor and extensor burst groups that were generally shared across slopes. Stance-related bursts of flexor muscles in downslope were placed in a burst group from level and upslope walking formed by the rectus femoris. Walking upslope changed swing/stance phase durations of level walking but not the cycle duration. Five muscle synergies computed using non-negative matrix factorization accounted for at least 95% of variance in EMG patterns in each slope. Five synergies were shared between level and upslope walking, whereas only three of those were shared with downslope synergies; these synergies were active during the swing phase and phase transitions. Two stance-related synergies of downslope walking were distinct; they comprised a mixture of flexors and extensors. We suggest that the modular organization of muscle activity during level and slope walking results from interactions between motion-related sensory feedback, CPG, and supraspinal inputs.NEW & NOTEWORTHY We demonstrated that the atypical EMG activities during cat downslope walking, silent one-joint hip extensors and stance-related EMG bursts in flexors, have many features shared with activities of level and upslope walking. Majority of EMG burst groups and muscle synergies were shared among these slopes, and upslope modulated the swing/stance phase duration but not cycle duration. Thus, synergistic EMG activities in all slopes might result from a shared CPG receiving somatosensory and supraspinal inputs.

4451 On the loss of individual joint controllability and the organization of muscle synergies in the impaired arm following a stroke: A pilot study

OBJECTIVES/GOALS: Damage to the sensorimotor cortex areas or/and motor/sensory pathways after a stroke could lead the motor system to a loss of controllability for joints. We investigate the loss of individual joint controllability called a loss of individualization during arm movement, which would provide an insight into abnormal motor coordination. METHODS/STUDY POPULATION: We recruit 12 chronic stroke survivors with Fugl-Meyer score between 26 and 50. A robotic exoskeleton with minimum mechanical resistance is equipped to measure the movements of the shoulder, elbow and wrist joints, respectively. Surface EMGs on muscles related to the joints are recorded using 11 wireless pre-amplified electrodes. Participants are asked to move the shoulder, elbow, or wrist joint individually throughout their range of motion, without moving the other joints voluntarily. RESULTS/ANTICIPATED RESULTS: It would be expected that participants show more difficulty in individualization of the distal joint in comparison with the proximal joint. A reduced joint range of motion would be observed in a descending order of the wrist, elbow and shoulder. These results are in line with the proximal-to-distal gradient of motor deficits after a stroke. Intention of moving the distal joint would induce a greater deviation in the position of the proximal joint than that of the distal joint when moving the proximal joint. A non-negative matrix factorization algorithm would reveal a decreased number of muscle synergies in the groups with a loss of individuation in comparison with the groups with no loss. DISCUSSION/SIGNIFICANCE OF IMPACT: We demonstrate that a stroke leads to a lack of individual joint controllability, with a greater deficits on the distal joint, and that it is related to a decreased number of muscle synergies across the corresponding joints. CONFLICT OF INTEREST DESCRIPTION: N/A.

Age-Related Differences in Muscle Synergy Organization during Step Ascent at Different Heights and Directions

The aim of this study was to explore the underlying age-related differences in dynamic motor control during different step ascent conditions using muscle synergy analysis. Eleven older women (67.0 y ± 2.5) and ten young women (22.5 y ± 1.6) performed stepping in forward and lateral directions at step heights of 10, 20 and 30 cm. Surface electromyography was obtained from 10 lower limb and torso muscles. Non-negative matrix factorization was used to identify sets of (n) synergies across age groups and stepping conditions. In addition, variance accounted for (VAF) by the detected number of synergies was compared to assess complexity of motor control. Finally, correlation coefficients of muscle weightings and between-subject variability of the temporal activation patterns were calculated and compared between age groups and stepping conditions. Four synergies accounted for &gt;85% VAF across age groups and stepping conditions. Age and step height showed a significant negative correlation with VAF during forward stepping but not lateral stepping, with lower VAF indicating higher synergy complexity. Muscle weightings showed higher similarity across step heights in older compared to young women. Neuromuscular control of young and community-dwelling older women could not be differentiated based on the number of synergies extracted. Additional analyses of synergy structure and complexity revealed subtle age- and step-height-related differences, indicating that older women rely on more complex neuromuscular control strategies.

Is the Neuromuscular Organization of Throwing Unchanged in Virtual Reality? Implications for Upper Limb Rehabilitation

Virtual reality (VR) is an appealing approach for increasing the engagement and attention of patients during rehabilitation. Understanding how motor control changes in real vs. virtual scenarios is a research challenge in terms of validating its administration. This study evaluates muscle synergies when subjects conduct throwing tasks in virtual reality. Seventeen healthy subjects performed 20 throws both in a virtual environment and in real one as they threw a ball with both dominant and nondominant arms. The electromyography (EMG) signals of 11 muscles of the upper limbs were recorded. Non-negative matrix factorization was used to extract muscle synergies. The cosine similarity was computed to assess the consistence of muscle synergy organization between virtual and real tasks. The same parameter was used to establish the inter-subject similarity. A three-synergy model was selected as the most likely. No effects of virtual reality and arm side on neuromuscular organization were found. Forearm muscles, not necessary for ball holding and release, were comprised in the activation synergies in the virtual reality environment. Finally, the synergies were consistent across subjects, especially during the deceleration phase. Results are encouraging for the application of virtual reality to complement conventional therapy, improve engagement, and facilitate objective measurements of pathology progression.

Modular Organization of Muscle Synergies to Achieve Movement Behaviors.

Muscle synergy has been applied to comprehend how the central nervous system (CNS) controls movements for decades. However, it is not clear about the motion control mechanism and the relationship between motions and muscle synergies. In this paper, we designed two experiments to corroborate the hypothesis: (1) motions can be decomposed to motion primitives, which are driven by muscle synergy primitives and (2) variations of motion primitives in direction and scale are modulated by activation coefficients rather than muscle synergy primitives. Surface electromyographic (EMG) signals were recorded from nine muscles of the upper limb. Nonnegative matrix factorization (NMF) was applied to extract muscle synergy vectors and corresponding activation coefficients. We found that synergy structures of different movement patterns were similar (α=0.05). The motion modulation indexes (MMI) among movement patterns in reaching movements showed apparent differences. Merging coefficients and reconstructed similarity of synergies between simple motions and complex motions were significant. This study revealed the motion control mechanism of the CNS and provided a rehabilitation and evaluation method for patients with motor dysfunction in exercise and neuroscience.

Age-related differences in muscle synergy organization during step ascent across step heights and directions

Recently, the eigenvalue problems formulated with symmetric positive definite bilinear forms have been well investigated with the aim of explicit bounds for the eigenvalues. In this paper, the existing theorems for bounding eigenvalues are further extended to deal with the case of eigenvalue problems defined by positive semi-definite bilinear forms. As an application, the eigenvalue estimation theorems are applied to the error constant estimation for polynomial projections.

Design and effectiveness evaluation of mirror myoelectric interfaces: a novel method to restore movement in hemiplegic patients

The motor impairment occurring after a stroke is characterized by pathological muscle activation patterns or synergies. However, while robot-aided myoelectric interfaces have been proposed for stroke rehabilitation, they do not address this issue, which might result in inefficient interventions. Here, we present a novel paradigm that relies on the correction of the pathological muscle activity as a way to elicit rehabilitation, even in patients with complete paralysis. Previous studies demonstrated that there are no substantial inter-limb differences in the muscle synergy organization of healthy individuals. We propose building a subject-specific model of muscle activity from the healthy limb and mirroring it to use it as a learning tool for the patient to reproduce the same healthy myoelectric patterns on the paretic limb during functional task training. Here, we aim at understanding how this myoelectric model, which translates muscle activity into continuous movements of a 7-degree of freedom upper limb exoskeleton, could transfer between sessions, arms and tasks. The experiments with 8 healthy individuals and 2 chronic stroke patients proved the feasibility and effectiveness of such myoelectric interface. We anticipate the proposed method to become an efficient strategy for the correction of maladaptive muscle activity and the rehabilitation of stroke patients.

Saturday, September 29, 2018 9:00 am–10:00 am Impact of Adult Deformity Correction: 262. The effect of surgical alignment on gait complexity in adult deformity patients: a neuromuscular synergy approach

BACKGROUND CONTEXT Adult degenerative scoliosis (ADS) patients frequently suffer from impairments in mobility. Surgical intervention for ADS mostly improves gait, balance and other health-related quality of life scores. Previous literature demonstrates that the central nervous system (CNS) might use an organization of muscle synergies to control a wide range of activities, for example walking. Instead of controlling each muscle individually, the CNS groups the muscles and activates many them with a single control signal. Hence, neuromuscular synergies may indicate the ability of the CNS in generating independent control signals. Less number of muscle synergies during walking emphasizes inability of the CNS to generate as many independent control signals. The higher number of synergies represents the higher gait complexity since more number of control inputs are required to achieve the same task. PURPOSE To compare number of walking muscle synergies in ADS patients both before and 3 months postsurgery. STUDY DESIGN/SETTING A prospective concurrent control cohort study. PATIENT SAMPLE A total of 13 ADS patients. OUTCOME MEASURES Gait complexity by number of muscle synergies. METHODS Clinical gait analysis was performed one week prior and 3 months postsurgery. Five walking trials were performed at comfort speed. Surface electromyography (EMG) electrodes were placed and recorded bilaterally from 16 trunk and lower extremity muscles: external oblique, gluteus maximus, multifidus, erector spinae, rectus femoris, semitendinosus, tibialis anterior, medial gastrocnemius. EMG data was collected at 2,000 Hz, filtered, rectified, and normalized. The processed EMG was mathematically broken into synergies and their activation ratio (from 100%). The higher number of synergies always reduces the residual error between the reconstructed EMG and the original EMG. The required number of synergies were defined as the minimum number of synergies that could reconstruct EMG signals with subtracting 5% error ([EMGoriginal -EMGreconstructed] = e, e/EMGoriginal RESULTS Clinical gait analysis was performed one week prior and 3 months postsurgery. Five walking trials were performed at comfort speed. Surface electromyography (EMG) electrodes were placed and recorded bilaterally from 16 trunk and lower extremity muscles: external oblique, gluteus maximus, multifidus, erector spinae, rectus femoris, semitendinosus, tibialis anterior and medial gastrocnemius. EMG data was collected at 2,000 Hz, filtered, rectified and normalized. The processed EMG was mathematically broken into synergies and their activation ratio (from 100%). The higher number of synergies always reduces the residual error between the reconstructed EMG and the original EMG. The required number of synergies were defined as the minimum number of synergies that could reconstruct EMG signals with subtracting 5% error {(EMGoriginal -EMGreconstructed) = e, e/EMGoriginal CONCLUSIONS This study shows an increase in gait complexity by increase in number of synergies following a surgical alignment in ADS patients. More number of synergies postsurgery shows a more elaborate gait pattern since more number of synergies (modules) are required to rebuild the EMG signals, in addition to improvements in ability of the CNS in generating more number of independent or rich control signals. As an example, inability of poststroke patients in generating independent control signals for gait makes their CNS to activate numerous muscles with a single signal (at the same time), causing unwanted co-contractions that hinders their normal gait. We recommend that spine care providers use gait analysis as part of their clinical evaluation to provide an objective measure of function and to better understand the effects of the disease and its treatment on their patients’ gait, function, and, ultimately, quality of life. FDA DEVICE/DRUG STATUS This abstract does not discuss or include any applicable devices or drugs.

Evaluating upper limb impairments in multiple sclerosis by exposure to different mechanical environments

Multiple sclerosis is a chronic, autoimmune and neurodegenerative disease affecting multiple functional systems and resulting in motor impairments associated with muscle weakness and lack of movement coordination. We quantified upper limb motor deficits with a robot-based assessment including behavioral and muscle synergy analysis in 11 multiple sclerosis subjects with mild to moderate upper limb impairment (9 female; 50 ± 10 years) compared to 11 age- and gender- matched controls (9 female; 50 ± 9 years). All subjects performed planar reaching tasks by moving their upper limb or applying force while grasping the handle of a robotic manipulandum that generated four different environments: free space, assistive or resistive forces, and rigid constraint. We recorded the activity of 15 upper body muscles. Multiple sclerosis subjects generated irregular trajectories. While activities in isolated arm muscles appeared generally normal, shoulder muscle coordination with arm motions was impaired and there was a marked co-activation of the biceps and triceps in extension movements. Systematic differences in timing and organization of muscle synergies have also been observed. This study supports the definition of new biomarkers and rehabilitative treatments for improving upper limb motor coordination in multiple sclerosis.

Effect of handedness on muscle synergies during upper limb planar movements.

Handedness is a prominent but poorly understood aspect of human motor performances. Despite it is generally accepted that it results from differences in the neural control of the arm, the mechanisms at the origin of the side-difference in motor performances are still unknown. In this work, we propose to deepen this aspect by investigating muscle synergies organization. We obtained muscle synergies through the factorization of the superficial electromyographical (EMG) activity related to fifteen upper limb muscles in the dominant and non-dominant side of 5 healthy young right and left dominant subjects, while executing planar wide and tight circular trajectories. Our preliminary results showed that right and left handed subjects performed the circular trajectories with a different muscle organization. Moreover, a task-related side-difference in muscle synergies was observed. Further investigations in a larger cohort of individuals are necessary to determine the neural mechanisms generating the differences in number and organization of muscle synergies between left and right handed individuals.

Functional muscle synergies constrain force production during postural tasks

We recently demonstrated that a set of five functional muscle synergies were sufficient to characterize both hindlimb muscle activity and active forces during automatic postural responses in cats standing at multiple postural configurations. This characterization depended critically upon the assumption that the endpoint force vector (synergy force vector) produced by the activation of each muscle synergy rotated with the limb axis as the hindlimb posture varied in the sagittal plane. Here, we used a detailed, 3D static model of the hindlimb to confirm that this assumption is biomechanically plausible: as we varied the model posture, simulated synergy force vectors rotated monotonically with the limb axis in the parasagittal plane ( r 2=0.94±0.08). We then tested whether a neural strategy of using these five functional muscle synergies provides the same force-generating capability as controlling each of the 31 muscles individually. We compared feasible force sets (FFSs) from the model with and without a muscle synergy organization. FFS volumes were significantly reduced with the muscle synergy organization ( F=1556.01, p≪0.01), and as posture varied, the synergy-limited FFSs changed in shape, consistent with changes in experimentally measured active forces. In contrast, nominal FFS shapes were invariant with posture, reinforcing prior findings that postural forces cannot be predicted by hindlimb biomechanics alone. We propose that an internal model for postural force generation may coordinate functional muscle synergies that are invariant in intrinsic limb coordinates, and this reduced-dimension control scheme reduces the set of forces available for postural control.

Modular Organization of Finger Movements by the Human Central Nervous System

The motor system may generate automated movements, such as walking, by combining modular spinal motor synergies. However, it remains unknown whether a modular neuronal architecture is sufficient to generate the unique flexibility of human finger movements, which rely on cortical structures. Here we show that finger movements evoked by transcranial magnetic stimulation (TMS) of the primary motor cortex reproduced distinctive features of the spatial representation of voluntary movements as identified in previous neuroimaging studies, consistent with naturalistic activation of neuronal elements. Principal component analysis revealed that the dimensionality of TMS-evoked movements was low. Principal components extracted from TMS-induced finger movements resembled those derived from end-postures of voluntary movements performed to grasp imagined objects, and a small subset of them was sufficient to reconstruct these movements with remarkable fidelity. The motor system may coordinate even the most dexterous movements by using a modular architecture involving cortical components.

Muscle Synergy Organization Is Robust Across a Variety of Postural Perturbations

We recently showed that four muscle synergies can reproduce multiple muscle activation patterns in cats during postural responses to support surface translations. We now test the robustness of functional muscle synergies, which specify muscle groupings and the active force vectors produced during postural responses under several biomechanically distinct conditions. We aimed to determine whether such synergies represent a generalized control strategy for postural control or if they are merely specific to each postural task. Postural responses to multidirectional translations at different fore-hind paw distances and to multidirectional rotations at the preferred stance distance were analyzed. Five synergies were required to adequately reconstruct responses to translation at the preferred stance distance-four were similar to our previous analysis of translation, whereas the fifth accounted for the newly added background activity during quiet stance. These five control synergies could account for > 80% total variability or r2 > 0.6 of the electromyographic and force tuning curves for all other experimental conditions. Forces were successfully reconstructed but only when they were referenced to a coordinate system that rotated with the limb axis as stance distance changed. Finally, most of the functional muscle synergies were similar across all of the six cats in terms of muscle synergy number, synergy activation patterns, and synergy force vectors. The robustness of synergy organization across perturbation types, postures, and animals suggests that muscle synergies controlling task-variables are a general construct used by the CNS for balance control.

Central and Sensory Contributions to the Activation and Organization of Muscle Synergies during Natural Motor Behaviors

Previous studies have suggested that the motor system may simplify control by combining a small number of muscle synergies represented as activation profiles across a set of muscles. The role of sensory feedback in the activation and organization of synergies has remained an open question. Here, we assess to what extent the motor system relies on centrally organized synergies activated by spinal and/or supraspinal commands to generate motor outputs by analyzing electromyographic (EMG) signals collected from 13 hindlimb muscles of the bullfrog during swimming and jumping, before and after deafferentation. We first established that, for both behaviors, the intact and deafferented data sets possess low and similar dimensionalities. Subsequently, we used a novel reformulation of the non-negative matrix factorization algorithm to simultaneously search for synergies shared by, and synergies specific to, the intact and deafferented data sets. Most muscle synergies were identified as shared synergies, suggesting that EMGs of locomotor behaviors are generated primarily by centrally organized synergies. Both the amplitude and temporal patterns of the activation coefficients of most shared synergies, however, were altered by deafferentation, suggesting that sensory inflow modulates activation of those centrally organized synergies. For most synergies, effects of deafferentation on the activation coefficients were not consistent across frogs, indicating substantial interanimal variability of feedback actions. We speculate that sensory feedback might adapt recruitment of muscle synergies to behavioral constraints, and the few synergies specific to the intact or deafferented states might represent afferent-specific modules or feedback reorganization of spinal neuronal networks.

Neuromuscular Adaptation During Skill Acquisition on a Two Degree-of-Freedom Target-Acquisition Task: Isometric Torque Production

In this study we attempted to identify the principles that govern the changes in neural control that occur during repeated performance of a multiarticular coordination task. Eight participants produced isometric flexion/extension and pronation/supination torques at the radiohumeral joint, either in isolation (e.g., flexion) or in combination (e.g., flexion-supination), to acquire targets presented by a visual display. A cursor superimposed on the display provided feedback of the applied torques. During pre- and postpractice tests, the participants acquired targets in eight directions located either 3.6 cm (20% maximal voluntary contraction [MVC]) or 7.2 cm (40% MVC) from a neutral cursor position. On each of five consecutive days of practice the participants acquired targets located 5.4 cm (30% MVC) from the neutral position. EMG was recorded from eight muscles contributing to torque production about the radiohumeral joint during the pre- and posttests. Target-acquisition time decreased significantly with practice in most target directions and at both target torque levels. These performance improvements were primarily associated with increases in the peak rate of torque development after practice. At a muscular level, these changes were brought about by increases in the rates of recruitment of all agonist muscles. The spatiotemporal organization of muscle synergies was not significantly altered after practice. The observed adaptations appear to lead to performances that are generalizable to actions that require both greater and smaller joint torques than that practiced, and may be successfully recalled after a substantial period without practice. These results suggest that tasks in which performance is improved by increasing the rate of muscle activation, and thus the rate of joint torque development, may benefit in terms of the extent to which acquired levels of performance are maintained over time.

EMG responses to maintain stance during multidirectional surface translations.

To characterize muscle synergy organization underlying multidirectional control of stance posture, electromyographic activity was recorded from 11 lower limb and trunk muscles of 7 healthy subjects while they were subjected to horizontal surface translations in 12 different, randomly presented directions. The latency and amplitude of muscle responses were quantified for each perturbation direction. Tuning curves for each muscle were examined to relate the amplitude of the muscle response to the direction of surface translation. The latencies of responses for the shank and thigh muscles were constant, regardless of perturbation direction. In contrast, the latencies for another thigh [tensor fascia latae (TFL)] and two trunk muscles [rectus abdominis (RAB) and erector spinae (ESP)] were either early or late, depending on the perturbation direction. These three muscles with direction-specific latencies may play different roles in postural control as prime movers or as stabilizers for different translation directions, depending on the timing of recruitment. Most muscle tuning curves were within one quadrant, having one direction of maximal activity, generally in response to diagonal surface translations. Two trunk muscles (RAB and ESP) and two lower limb muscles (semimembranosus and peroneus longus) had bipolar tuning curves, with two different directions of maximal activity, suggesting that these muscle can play different roles as part of different synergies, depending on translation direction. Muscle tuning curves tended to group into one of three regions in response to 12 different directions of perturbations. Two muscles [rectus femoris (RFM) and TFL] were maximally active in response to lateral surface translations. The remaining muscles clustered into one of two diagonal regions. The diagonal regions corresponded to the two primary directions of active horizontal force vector responses. Two muscles (RFM and adductor longus) were maximally active orthogonal to their predicted direction of maximal activity based on anatomic orientation. Some of the muscles in each of the synergic regions were not anatomic synergists, suggesting a complex central organization for recruitment of muscles. The results suggest that neither a simple reflex mechanism nor a fixed muscle synergy organization is adequate to explain the muscle activation patterns observed in this postural control task. Our results are consistent with a centrally mediated pattern of muscle latencies combined with peripheral influence on muscle magnitude. We suggest that a flexible continuum of muscle synergies that are modifiable in a task-dependent manner be used for equilibrium control in stance.