Activation Of Muscle Synergies Research Articles

Psychological stress affects trial-to-trial variability of temporal-spatial gait parameters, but not of muscle synergy activation coefficients

Psychological stress affects trial-to-trial variability of temporal-spatial gait parameters, but not of muscle synergy activation coefficients

Investigation of adaptive muscle synergy modulated motor responses to grasping perturbations

This study investigated how muscle synergies adapt in response to unexpected changes in object weight during lifting tasks. The aim was to discover which motor control strategies individuals use to maintain their grasping performance. Muscle synergies were extracted from the muscle activity of fifteen healthy participants who lifted objects of identical appearance but varying weights in a randomized order, which introduced artificial perturbations. Reaching and manipulation phases of object lifting were analyzed using constrained non-negative matrix factorization and k-means clustering. Participants exhibited a perturbation-independent and thus consistent recruitment of spatial synergy components, while significant adaptations in muscle synergy activation occurred in response to unexpected perturbations. Perturbations caused by unexpectedly heavy objects led to delayed and gradual increases in muscle synergy activation until the force required to lift the object was reached. In contrast, perturbations caused by lighter objects led to reductions in excess muscle synergy activation occurring later. Sensorimotor control maintains the modularity of muscle synergies. Even when external mechanical perturbations occur, the grasping performance is preserved, and control is adapted solely through muscle synergy activation. These results suggest that using pure spatial synergy components as control signals for myoelectric arm prostheses may prevent them from malfunctioning due to external perturbations.

Increased trial-to-trial similarity and reduced temporal overlap of muscle synergy activation coefficients manifest during learning and with increasing movement proficiency

Muscle synergy analyses are used to enhance our understanding of motor control. Spatially fixed synergy weights coordinate multiple co-active muscles through activation commands, known as activation coefficients. To gain a more comprehensive understanding of motor learning, it is essential to understand how activation coefficients vary during a learning task and at different levels of movement proficiency. Participants walked on a line, a beam, and learned to walk on a tightrope—tasks that represent different levels of proficiency. Muscle synergies were extracted from electromyography signals across all conditions and the number of synergies was determined by the knee-point of the total variance accounted for (tVAF) curve. The results indicated that the tVAF of one synergy decreased with task proficiency, with the tightrope task resulting in the highest tVAF compared to the line and beam tasks. Furthermore, with increasing proficiency and after a learning process, trial-to-trial similarity increased and temporal overlap of synergy activation coefficients decreased. Consequently, we propose that precise adjustment and refinement of synergy activation coefficients play a pivotal role in motor learning.

Feasibility of Isokinetic Training to Modify Coupling of Upper Limb Muscle Synergy Activation in Stroke-affected Upper Limb.

Abnormal intermuscular coordination in stroke-affected upper limbs contributes to motor deficits after stroke. In particular, abnormalities in the activation of upper limb muscle synergies after stroke were demonstrated for endpoint force control during isokinetic exercises. This study aimed to investigate the feasibility of isokinetic training to alter these abnormal synergy activations and improve motor control. Muscle synergies and Wolf Motor Function Test Functional Ability Scale (WMFT-FAS) score were compared before and after three weeks of electromyography-based training. The proposed training changed the synergy activation and improved the WMFT-FAS score in a chronic stroke survivor while preserving the muscle weights of the synergies.Clinical Relevance- This study presents the feasibility of neuromuscular training to modify the activation of upper limb muscle synergies against stroke-specific patterns of intermuscular coordination and improve WMFT-FAS score.

Generalizability of muscle synergies in isometric force generation versus point-to-point reaching in the human upper extremity workspace.

Isometric force generation and kinematic reaching in the upper extremity has been found to be represented by a limited number of muscle synergies, even across task-specific variations. However, the extent of the generalizability of muscle synergies between these two motor tasks within the arm workspace remains unknown. In this study, we recorded electromyographic (EMG) signals from 13 different arm, shoulder, and back muscles of ten healthy individuals while they performed isometric and kinematic center-out target matches to one of 12 equidistant directional targets in the horizontal plane and at each of four starting arm positions. Non-negative matrix factorization was applied to the EMG data to identify the muscle synergies. Five and six muscle synergies were found to represent the isometric force generation and point-to-point reaches. We also found that the number and composition of muscle synergies were conserved across the arm workspace per motor task. Similar tuning directions of muscle synergy activation profiles were observed at different starting arm locations. Between the isometric and kinematic motor tasks, we found that two to four out of five muscle synergies were common in the composition and activation profiles across the starting arm locations. The greater number of muscle synergies that were involved in achieving a target match in the reaching task compared to the isometric task may explain the complexity of neuromotor control in arm reaching movements. Overall, our results may provide further insight into the neuromotor compartmentalization of shared muscle synergies between two different arm motor tasks and can be utilized to assess motor disabilities in individuals with upper limb motor impairments.

Muscle coordination patterns in regulation of medial gastrocnemius activation during walking

The ankle plantar flexion in the late stance phase is referred to as the ankle push-off. When the ankle push-off force is enhanced, compensatory adjustments occur in the adjacent phases. The muscle control that achieves these compensatory movements remains unknown, although they are expected to be coordinately regulated across multiple muscles and phases. Muscle synergy is used as a quantification technique for muscle coordination, and this analysis enables the comparison of synchronized activity between multiple muscles. Therefore, this study aimed to elucidate the tuning of muscle synergies in muscle activation adjustment of push-off. It is hypothesized that muscle activation adjustment of push-off is performed in the muscle synergy related to ankle push-off and in the muscle synergy that activates during the adjacent push-off phase. Eleven healthy men participated, and participants manipulated the activity of the medial gastrocnemius during walking through visual feedback. Two conditions were compared as experimental conditions: increasing the muscle activity to 1.6 times that during normal walking (High) and matching it with that during normal walking (Normal). Twelve muscle activities in the trunk and lower limb and kinematic data were recorded. Muscle synergies were extracted by the non-negative matrix factorization. No significant difference was observed in the number of synergies (High: 3.5 ± 0.8, Normal: 3.7 ± 0.9, p = 0.21) and muscle synergy activation timing and duration between the High and Normal conditions (p > 0.27). However, significant differences were observed in the peak muscle activity during the late stance phase of the rectus femoris (RF), biceps femoris (BF) between conditions (RF at High: 0.32 ± 0.21, RF at Normal: 0.45 ± 0.17, p = 0.02; BF at High: 0.16 ± 0.01, BF at Normal: 0.08 ± 0.06 p = 0.02). Although the quantification of force exertion has not been conducted, the modulation of RF and BF activation could have occurred due to the attempts to help knee flexion. Muscle synergies during normal walking are therefore maintained, and slight adjustments in the amplitude of muscle activity occurred for each muscle.

Agonist-antagonist muscle strain in the residual limb preserves motor control and perception after amputation

BackgroundElucidating underlying mechanisms in subject-specific motor control and perception after amputation could guide development of advanced surgical and neuroprosthetic technologies. In this study, relationships between preserved agonist-antagonist muscle strain within the residual limb and preserved motor control and perception capacity are investigated.MethodsFourteen persons with unilateral transtibial amputations spanning a range of ages, etiologies, and surgical procedures underwent evaluations involving free-space mirrored motions of their lower limbs. Research has shown that varied motor control in biologically intact limbs is executed by the activation of muscle synergies. Here, we assess the naturalness of phantom joint motor control postamputation based on extracted muscle synergies and their activation profiles. Muscle synergy extraction, degree of agonist-antagonist muscle strain, and perception capacity are estimated from electromyography, ultrasonography, and goniometry, respectively.ResultsHere, we show significant positive correlations (P < 0.005–0.05) between sensorimotor responses and residual limb agonist-antagonist muscle strain. Identified trends indicate that preserving even 20–26% of agonist-antagonist muscle strain within the residuum compared to a biologically intact limb is effective in preserving natural motor control postamputation, though preserving limb perception capacity requires more (61%) agonist-antagonist muscle strain preservation.ConclusionsThe results suggest that agonist-antagonist muscle strain is a characteristic, readily ascertainable residual limb structural feature that can help explain variability in amputation outcome, and agonist-antagonist muscle strain preserving surgical amputation strategies are one way to enable more effective and biomimetic sensorimotor control postamputation.

Clarify Sit-to-Stand Muscle Synergy and Tension Changes in Subacute Stroke Rehabilitation by Musculoskeletal Modeling.

Post-stroke patients exhibit distinct muscle activation electromyography (EMG) features in sit-to-stand (STS) due to motor deficiency. Muscle activation amplitude, related to muscle tension and muscle synergy activation levels, is one of the defining EMG features that reflects post-stroke motor functioning and motor impairment. Although some qualitative findings are available, it is not clear if and how muscle activation amplitude-related biomechanical attributes may quantitatively reflect during subacute stroke rehabilitation. To better enable a longitudinal investigation into a patient's muscle activation changes during rehabilitation or an inter-subject comparison, EMG normalization is usually applied. However, current normalization methods using maximum voluntary contraction (MVC) or within-task peak/mean EMG may not be feasible when MVC cannot be obtained from stroke survivors due to motor paralysis and the subject of comparison is EMG amplitude. Here, focusing on the paretic side, we first propose a novel, joint torque-based normalization method that incorporates musculoskeletal modeling, forward dynamics simulation, and mathematical optimization. Next, upon method validation, we apply it to quantify changes in muscle tension and muscle synergy activation levels in STS motor control units for patients in subacute stroke rehabilitation. The novel method was validated against MVC-normalized EMG data from eight healthy participants, and it retained muscle activation amplitude differences for inter- and intra-subject comparisons. The proposed joint torque-based method was also compared with the common static optimization based on squared muscle activation and showed higher simulation accuracy overall. Serial STS measurements were conducted with four post-stroke patients during their subacute rehabilitation stay (137 ± 22 days) in the hospital. Quantitative results of patients suggest that maximum muscle tension and activation level of muscle synergy temporal patterns may reflect the effectiveness of subacute stroke rehabilitation. A quality comparison between muscle synergies computed with the conventional within-task peak/mean EMG normalization and our proposed method showed that the conventional was prone to activation amplitude overestimation and underestimation. The contributed method and findings help recapitulate and understand the post-stroke motor recovery process, which may facilitate developing more effective rehabilitation strategies for future stroke survivors.

Alterations in intermuscular coordination underlying isokinetic exercise after a stroke and their implications on neurorehabilitation

BackgroundAbnormal intermuscular coordination limits the motor capability of stroke-affected upper limbs. By evaluating the intermuscular coordination in the affected limb under various biomechanical task constraints, the impact of a stroke on motor control can be analyzed and intermuscular coordination-based rehabilitation strategies can be developed. In this study, we investigated upper limb intermuscular coordination after a stroke during isokinetic movements.MethodsSixteen chronic stroke survivors and eight neurologically intact individuals were recruited. End-point forces and electromyographic activities of the shoulder and elbow muscles were measured while the participants performed isokinetic upper limb movements in a three-dimensional space. Intermuscular coordination of the stroke survivors and the control participants was quantified in the form of muscle synergies. Then, we compared the number, composition, and activation coefficients of muscle synergies and the end-point force between the groups. The correlation between the alteration of muscle synergies and the level of motor impairment was investigated.ResultsFour and five muscle synergies in the stroke and control groups were observed, respectively. The composition of muscle synergies was comparable between the groups, except that the three heads of the deltoid muscle were co-activated and formed one synergy in the stroke group, whereas those muscles formed two synergies in the control group. When the number of muscle synergies between the groups matched, the comparable composition of muscle synergies was observed in both groups. Alternatively, the modulation of synergy activation coefficients was altered after a stroke. The severity of motor impairments was negatively correlated with the similarity of the post-stroke synergies with respect to the mean control synergies.ConclusionsStroke-affected upper limbs seemed to modularize the activation of the shoulder and elbow muscles in a fairly similar way to that of neurologically intact individuals during isokinetic movements. Compared with free (i.e., unconstrained) movement, exercise under biomechanical constraints including the isokinetic constraint might promote the activation of muscle synergies independently in stroke survivors. We postulated the effect of biomechanical constraints on the intermuscular coordination and suggested a possible intermuscular coordination-based rehabilitation protocol that provides the biomechanical constraint appropriate to a trainee throughout the progress of rehabilitation.

Motor Control After Human SCI Through Activation of Muscle Synergies Under Spinal Cord Stimulation.

Spinal cord stimulation (SCS) has enabled motor recovery in paraplegics with motor complete spinal cord injury (SCI). However, the physiological mechanisms underlying this recovery are unknown. This paper analyzes muscle synergies in two motor complete SCI patients under SCS during standing and compares them with muscle synergies in healthy subjects, in order to help elucidate the mechanisms that enable motor control through SCS. One challenge is that standard muscle synergy extraction algorithms, such as non-negative matrix factorization (NMF), fail when applied to SCI patients under SCS. We develop a new algorithm-rShiftNMF-to extract muscle synergies in these cases. We find muscle synergies extracted by rShiftNMF are significantly better at interpreting electromyography (EMG) activity, and resulting synergy features are more physiologically meaningful. By analyzing muscle synergies from SCI patients and healthy subjects, we find that: 1) SCI patients rely significantly on muscle synergy activation to generate motor activity; 2) interleaving SCS can selectively activate an additional muscle synergy that is critical to SCI standing; and 3) muscle synergies extracted from SCI patients under SCS differ substantially from those extracted from healthy subjects. We provide evidence that after spinal cord injury, SCS influences motor function through muscle synergy activation.

Cortical Correlates of Locomotor Muscle Synergy Activation in Humans: An Electroencephalographic Decoding Study

SummaryMuscular control during walking is believed to be simplified by the coactivation of muscles called muscle synergies. Although significant corticomuscular connectivity during walking has been reported, the level at which the cortical activity is involved in muscle activity (muscle synergy or individual muscle level) remains unclear. Here we examined cortical correlates of muscle activation during walking by brain decoding of activation of muscle synergies and individual muscles from electroencephalographic signals. We demonstrated that the activation of locomotor muscle synergies was decoded from slow cortical waves. In addition, the decoding accuracy for muscle synergies was greater than that for individual muscles and the decoding of individual muscle activation was based on muscle-synergy-related cortical information. These results indicate the cortical correlates of locomotor muscle synergy activation. These findings expand our understanding of the relationships between brain and locomotor muscle synergies and could accelerate the development of effective brain-machine interfaces for walking rehabilitation.

Constrained particle filtering for movement identification in forearm prosthesis

We formulate the problem of movement identification for the forearm prosthesis using a nonlinear state-space system and the hypothesis of muscle synergies. The synergy activation coefficients contain task-specific information and can be used to identify limb movements. In the proposed framework, the measurements are EMG data and the system state consists of muscle synergy activation coefficients, which are physiologically constrained to be nonnegative on average. Particle filters are the state-of-the-art techniques for optimal state estimation in nonlinear and non-Gaussian systems. However, the very numerical nature of the particle filters, which constitutes their strength, becomes their major weakness in handling constraints on the state. In this paper, we solve the movement identification problem by introducing a constrained particle filter termed as mean density truncation (MiND). We show that MiND minimally perturbs the unconstrained distribution of the state while simultaneously satisfying the desired constraints on the unknown state. We recorded EMG data from forearm muscles of 12 participants for identification of hand and wrist movements. The proposed particle filtering with MiND provided an accurate stream of synergy activation coefficients (p < 0.001) which were used for movement identification with error rates significantly lower (p < 0.05) than currently used heuristics and Linear Discriminate Analysis.

Visual and Vestibular Inputs Affect Muscle Synergies Responsible for Body Extension and Stabilization in Sit-to-Stand Motion.

The sit-to-stand motion is a common movement in daily life and understanding the mechanism of the sit-to-stand motion is important. Our previous study shows that four muscle synergies can characterize the sit-to-stand motion, and they have specific roles, such as upper body flexion, rising from a chair, body extension, and posture stabilization. The time-varying weight of these synergies are changed to achieve adaptive movement. However, the relationship between sensory input and the activation of the muscle synergies is not completely understood. In this paper, we aim to clarify how vestibular and visual inputs affect the muscle synergy in sit-to-stand motion. To address this, we conducted experiments as follows. Muscle activity, body kinematics, and ground reaction force were measured for the sit-to-stand motion under three different conditions: control, visual-disturbance, and vestibular-disturbance conditions. Under the control condition, the participants stood without any intervention. Under the visual-disturbance condition, the participants wore convex lens glasses and performed the sit-to-stand motion in a dark room. Under the vestibular-disturbance condition, a caloric test was performed. Muscle synergies were calculated for these three conditions using non-negative matrix factorization. We examined whether the same four muscle synergies were employed under each condition, and the changes in the time-varying coefficients were determined. These experiments were conducted on seven healthy, young participants. It was found that four muscle synergies could explain the muscle activity in the sit-to-stand motion under the three conditions. However, there were significant differences in the time-varying weight coefficients. When the visual input was disturbed, a larger amplitude was found for the muscle synergy that activated mostly in the final posture stabilization phase of the sit-to-stand motion. Under vestibular-disturbance condition, a longer activation was observed for the synergies that extended the entire body and led to posture stabilization. The results implied that during human sit-to-stand motion, visual input has less contribution to alter or correct activation of muscle synergies until the last phase. On the other hand, duration of muscle synergies after the buttocks leave are prolonged in order to adapt to the unstable condition in which sense of verticality is decreased under vestibular-disturbance.

Lower Local Dynamic Stability and Invariable Orbital Stability in the Activation of Muscle Synergies in Response to Accelerated Walking Speeds.

In order to achieve flexible and smooth walking, we must accomplish subtasks (e. g., loading response, forward propulsion or swing initiation) within a gait cycle. To evaluate subtasks within a gait cycle, the analysis of muscle synergies may be effective. In the case of walking, extracted sets of muscle synergies characterize muscle patterns that relate to the subtasks within a gait cycle. Although previous studies have reported that the muscle synergies of individuals with disorders reflect impairments, a way to investigate the instability in the activations of muscle synergies themselves has not been proposed. Thus, we investigated the local dynamic stability and orbital stability of activations of muscle synergies across various walking speeds using maximum Lyapunov exponents and maximum Floquet multipliers. We revealed that the local dynamic stability in the activations decreased with accelerated walking speeds. Contrary to the local dynamic stability, the orbital stability of the activations was almost constant across walking speeds. In addition, the increasing rates of maximum Lyapunov exponents were different among the muscle synergies. Therefore, the local dynamic stability in the activations might depend on the requirement of motor output related to the subtasks within a gait cycle. We concluded that the local dynamic stability in the activation of muscle synergies decrease as walking speed accelerates. On the other hand, the orbital stability is sustained across broad walking speeds.

Fatigue Influences the Recruitment, but Not Structure, of Muscle Synergies.

The development of fatigue elicits multiple adaptations from the neuromuscular system. Muscle synergies are common patterns of neuromuscular activation that have been proposed as the building blocks of human movement. We wanted to identify possible adaptations of muscle synergies to the development of fatigue in the upper limb. Recent studies have reported that synergy structure remains invariant during the development of fatigue, but these studies did not examine isolated synergies. We propose a novel approach to characterise synergy adaptations to fatigue by taking advantage of the spatial tuning of synergies. This approach allows improved identification of changes to individual synergies that might otherwise be confounded by changing contributions of overlapping synergies. To analyse upper limb synergies, we applied non-negative matrix factorization to 14 EMG signals from muscles of 11 participants performing isometric contractions. A preliminary multidirectional task was used to identify synergy directional tuning. A subsequent fatiguing task was designed to fatigue the participants in their synergies’ preferred directions. Both tasks provided virtual reality feedback of the applied force direction and magnitude, and were performed at 40% of each participant’s maximal voluntary force. Five epochs were analysed throughout the fatiguing task to identify progressive changes of EMG amplitude, median frequency, synergy structure, and activation coefficients. Three to four synergies were sufficient to account for the variability contained in the original data. Synergy structure was conserved with fatigue, but interestingly synergy activation coefficients decreased on average by 24.5% with fatigue development. EMG amplitude did not change systematically with fatigue, whereas EMG median frequency consistently decreased across all muscles. These results support the notion of a neuromuscular modular organisation as the building blocks of human movement, with adaptations to synergy recruitment occurring with fatigue. When synergy tuning properties are considered, the reduction of activation of muscle synergies may be a reliable marker to identify fatigue.

Speed-Dependent Modulation of Muscle Activity Based on Muscle Synergies during Treadmill Walking.

The regulation of walking speed is easily achieved. However, the central nervous system (CNS) must coordinate numerous muscles in order to achieve a smooth and continuous control of walking speed. To control walking speed appropriately, the CNS may need to utilize a simplified system for the control of numerous muscles. Previous studies have revealed that the CNS may control walking via muscle synergies that simplify the control of muscles by modularly organizing several muscles. We hypothesized that the CNS controls the walking speed by flexibly modulating activation of muscle synergies within one gait cycle. Then, we investigated how the activation of muscle synergies depend on walking speeds using the center of activity (CoA) that indicates the center of the distribution of activation timing within one gait cycle. Ten healthy men walked on a treadmill at 14 different walking speeds. We measured the surface electromyograms (EMGs) and kinematic data. Muscle synergies were extracted using non-negative matrix factorization. Then, we calculated the CoA of each muscle synergy. We observed that the CoA of each specific synergy would shift as the walking speed changed. The CoA that was mainly activated during the heel contact phase (C1) and the activation that contributed to the double support phase (C3) shifted to the earlier phase as the walking speed increased, whereas the CoA that produced swing initiation motion (C4) and the activation that related to the late-swing phase (C5) shifted to the later phase. This shifting of the CoA indicates that the CNS controls intensive activation of muscle synergies during the regulation of walking speed. In addition, shifting the CoA might be associated with changes in kinematics or kinetics depending on the walking speed. We concluded that the CNS flexibly controls the activation of muscle synergies in regulation of walking speed.

EEG topographies provide subject-specific correlates of motor control

Electroencephalography (EEG) of brain activity can be represented in terms of dynamically changing topographies (microstates). Notably, spontaneous brain activity recorded at rest can be characterized by four distinctive topographies. Despite their well-established role during resting state, their implication in the generation of motor behavior is debated. Evidence of such a functional role of spontaneous brain activity would provide support for the design of novel and sensitive biomarkers in neurological disorders. Here we examined whether and to what extent intrinsic brain activity contributes and plays a functional role during natural motor behaviors. For this we first extracted subject-specific EEG microstates and muscle synergies during reaching-and-grasping movements in healthy volunteers. We show that, in every subject, well-known resting-state microstates persist during movement execution with similar topographies and temporal characteristics, but are supplemented by novel task-related microstates. We then show that the subject-specific microstates’ dynamical organization correlates with the activation of muscle synergies and can be used to decode individual grasping movements with high accuracy. These findings provide first evidence that spontaneous brain activity encodes detailed information about motor control, offering as such the prospect of a novel tool for the definition of subject-specific biomarkers of brain plasticity and recovery in neuro-motor disorders.

Action Direction of Muscle Synergies in Voluntary Multi-Directional Postural Control.

A muscle synergy is a coordinative structure of muscles that has been proposed as a strategy to reduce the number of variables that the central nervous system (CNS) has to address in motor tasks. In this article, the mechanical contribution of muscle synergies and coordinative structures of muscles in voluntary multi-directional postural control were investigated. The task for healthy, young subjects was to shift and align their center of pressure (COP) to targets dispersed in 12 different directions in the horizontal plane by leaning their bodies for 10 s. Electromyograms (EMGs) of 18 muscles and COPs were recorded in the experiment. Muscle synergies were extracted using non-negative matrix factorization (NMF), and the structure of coordinative modules to keep the posture leaning toward various directions was disclosed. Then the directional properties, such as the mechanical role (i.e., action directions, we use ADs as abbreviation below), of muscle synergies and muscles were estimated using an electromyogram-weighted averaging (EWA) method, which is based on a cross-correlation between the fluctuations in the activation of muscle synergies and the COP. The results revealed that the ADs of muscle synergies were almost uniformly distributed in the task space in most of the subjects, which indicates that mechanical characteristics reduce the redundancy in postural control. In terms of the composition of muscle synergies and the ADs of individual muscles, we confirmed that muscle synergies in multi-directional postural control comprised a combination of several muscles, including various ADs, that generate torque at different joints.

Shared and Task-Specific Muscle Synergies during Normal Walking and Slipping.

Falling accidents are costly due to their prevalence in the workplace. Slipping has been known to be the main cause of falling. Understanding the motor response used to regain balance after slipping is crucial to developing intervention strategies for effective recovery. Interestingly, studies on spinalized animals and studies on animals subjected to electrical microstimulation have provided major evidence that the Central Nervous System (CNS) uses motor primitives, such as muscle synergies, to control motor tasks. Muscle synergies are thought to be a critical mechanism used by the CNS to control complex motor tasks by reducing the dimensional complexity of the system. Even though synergies have demonstrated potential for indicating how the body responds to balance perturbations by accounting for majority of the data set's variability, this concept has not been applied to slipping. To address this gap, data from 11 healthy young adults were collected and analyzed during both unperturbed walking and slipping. Applying an iterative non-negative matrix decomposition technique, four muscle synergies and the corresponding time-series activation coefficients were extracted. The synergies and the activation coefficients were then compared between baseline walking and slipping to determine shared vs. task-specific synergies. Correlation analyses found that among four synergies, two synergies were shared between normal walking and slipping. However, the other two synergies were task-specific. Both limbs were contributing to each of the four synergies, suggesting substantial inter-limb coordination during gait and slip. These findings stay consistent with previous unilateral studies that reported similar synergies between unperturbed and perturbed walking. Activation coefficients corresponding to the two shared synergies were similar between normal walking and slipping for the first 200 ms after heel contact and differed later in stance, suggesting the activation of muscle synergies in response to a slip. A muscle synergy approach would reveal the used sub-tasks during slipping, facilitating identification of impaired sub-tasks, and potentially leading to a purposeful rehabilitation based on damaged sub-functions.

Common mechanisms of force detection and control in insects.

Information from sense organs that detect forces may be particularly important in neural control of muscles as modular groups (synergists). We have performed comparative studies to characterize force detection in insects using physiological studies in stick insects and cockroaches to gain insight into comparable mechanisms in fruit flies (Drosophila), in which new and powerful genetic tools have been developed. Campaniform sensilla are mechanoreceptors that encode forces as cuticular strains. Previous studies have shown they can reinforce muscle synergies. Receptors fulfill similar functions in different species but the effects of individual groups are determined by the specific forces they encode. Whole mount and dissected preparations were studied using confocal and light microscopy. Sense organs and leg muscles were labeled using diI diffusion from the thorax. Activities in peripheral nerves and muscles were recorded extracellularly. Many receptors are homologous but the structure and responses of the femoral group (fCS) and mobility of the trochanter-femur joint are species specific. In cockroaches and stick insects, sensitivities to forces in the joint plane are correlated with effects on muscle synergies. In Drosophila, the fCS are located on the ventral femur, suggesting that they should be sensitive to forces in the joint plane. We are currently testing motor effects using neurophysiological recordings from larger Calliphorid flies. There are clear homologies in some groups of force receptors. Legs of flies combine elements found in cockroaches and stick insects. Sensory feedback can have similar effects in activation of muscle synergies in flies.