Synaptic inputs to motor neurons underlying muscle coactivation for functionally different tasks have different spectral characteristics.

The central nervous system (CNS) may produce the same endpoint trajectory or torque profile with different muscle activation patterns. What differentiates these patterns is the presence of cocontraction, which does not contribute to effective torque generation but allows to modulate joints' mechanical stiffness. Although it has been suggested that the generation of force and the modulation of stiffness rely on separate pathways, a characterization of the differences between the synaptic inputs to motor neurons (MNs) underlying these tasks is still missing. In this study, participants coactivated the same pair of upper-limb muscles, i.e., the biceps brachii and the triceps brachii, to perform two functionally different tasks: limb stiffness modulation or endpoint force generation. Spike trains of MNs were identified through decomposition of high-density electromyograms (EMGs) collected from the two muscles. Cross-correlogram showed a higher synchronization between MNs recruited to modulate stiffness, whereas cross-muscle coherence analysis revealed peaks in the ß-band, which is commonly ascribed to a cortical origin. These peaks did not appear during the coactivation for force generation, thus suggesting separate cortical inputs for stiffness modulation. Moreover, a within-muscle coherence analysis identified two subsets of MNs that were selectively recruited to generate force or regulate stiffness. This study is the first to highlight different characteristics, and probable different neural origins, of the synaptic inputs driving a pair of muscles under different functional conditions. We suggest that stiffness modulation is driven by cortical inputs that project to a separate set of MNs, supporting the existence of a separate pathway underlying the control of stiffness.NEW & NOTEWORTHY The characterization of the pathways underlying force generation or stiffness modulation are still unknown. In this study, we demonstrated that the common input to motor neurons of antagonist muscles shows a high-frequency component when muscles are coactivated to modulate stiffness but not to generate force. Our results provide novel insights on the neural strategies for the recruitment of multiple muscles by identifying specific spectral characteristics of the synaptic inputs underlying functionally different tasks.

Muscle synergies for multidirectional isometric force generation during maintenance of upright standing posture.

Muscle synergies are defined as coordinated recruitment of groups of muscles with specific activation balances and time profiles aimed at generating task-specific motor commands. While muscle synergies in postural control have been investigated primarily in reactive balance conditions, the neuromechanical contribution of muscle synergies during voluntary control of upright standing is still unclear. In this study, muscle synergies were investigated during the generation of isometric force at the trunk during the maintenance of standing posture. Participants were asked to maintain the steady-state upright standing posture while pulling forces of different magnitudes were applied at the level at the waist in eight horizontal directions. Muscle synergies were extracted by nonnegative matrix factorization from sixteen lower limb and trunk muscles. An average of 5-6 muscle synergies were sufficient to account for a wide variety of EMG waveforms associated with changes in the magnitude and direction of pulling forces. A cluster analysis partitioned the muscle synergies of the participants into a large group of clusters according to their similarity, indicating the use of a subjective combination of muscles to generate a multidirectional force vector in standing. Furthermore, we found a participant-specific distribution in the values of cosine directional tuning parameters of synergy amplitude coefficients, suggesting the existence of individual neuromechanical strategies to stabilize the whole-body posture. Our findings provide a starting point for the development of novel diagnostic tools to assess muscle coordination in postural control and lay the foundation for potential applications of muscle synergies in rehabilitation.

Transferring Sensor-Based Assessments to Clinical Practice: The Case of Muscle Synergies.

Sensor-based assessments in medical practice and rehabilitation include the measurement of physiological signals such as EEG, EMG, ECG, heart rate, and NIRS, and the recording of movement kinematics and interaction forces. Such measurements are commonly employed in clinics with the aim of assessing patients' pathologies, but so far some of them have found full exploitation mainly for research purposes. In fact, even though the data they allow to gather may shed light on physiopathology and mechanisms underlying motor recovery in rehabilitation, their practical use in the clinical environment is mainly devoted to research studies, with a very reduced impact on clinical practice. This is especially the case for muscle synergies, a well-known method for the evaluation of motor control in neuroscience based on multichannel EMG recordings. In this paper, considering neuromotor rehabilitation as one of the most important scenarios for exploiting novel methods to assess motor control, the main challenges and future perspectives for the standard clinical adoption of muscle synergy analysis are reported and critically discussed.

Persistent changes in motor adaptation strategies after perturbations that require exploration of novel muscle activation patterns.

Motor skill learning requires the acquisition of novel muscle patterns and a new control policy-a process that requires time. In contrast, motor adaptation often requires only the adjustment of existing muscle patterns-a fast process. By altering the mapping of muscle activations onto cursor movements in a myoelectrically controlled virtual environment, we are able to create perturbations that require either the recombination of existing muscle synergies (compatible virtual surgery) or the learning of novel muscle patterns (incompatible virtual surgery). We investigated whether adaptation to a compatible surgery is affected by prior exposure to an incompatible surgery, i.e., a motor skill learning task. We found that adaptation to a compatible surgery was characterized by a decrease in the quality of muscle pattern reconstructions using the original synergies and an increase in reaction times only after exposure to an incompatible surgery. In contrast, prior exposure to a compatible surgery did not affect the learning process required to overcome an incompatible surgery. The fact that exposure to an incompatible surgery had a profound effect on the muscle patterns during the adaptation to a subsequent compatible surgery and not vice versa suggests that null space exploration, possibly combined with an explicit exploration strategy, is engaged during exposure to an incompatible surgery and remains enhanced during a new adaptation episode. We conclude that motor skill learning, requiring novel muscle activation patterns, leads to changes in the exploration strategy employed during a subsequent perturbation.NEW & NOTEWORTHY Motor skill learning requires the acquisition of novel muscle patterns, whereas motor adaptation requires adjusting existing ones. We wondered whether training a new motor skill affects motor adaptation strategies. We show that learning an incompatible perturbation, a complex skill requiring new muscle synergies, affects the muscle patterns observed during adaption to a compatible perturbation, which requires adjusting the existing synergies. Our results suggest that motor skill learning results in persistent changes in the exploration strategy.

Distinct locomotor precursors in newborn babies.

Mature locomotion involves modular spinal drives generating a set of fundamental patterns of motoneuron activation, each timed at a specific phase of locomotor cycles and associated with a stable muscle synergy. How locomotor modules develop and to what extent they depend on prior experience or intrinsic programs remains unclear. To address these issues, we herein leverage the presence at birth of two types of locomotor-like movements, spontaneous kicking and weight-bearing stepping. The former is expressed thousands of times in utero and postnatally, whereas the latter is elicited de novo by placing the newborn on the ground for the first time. We found that the neuromuscular modules of stepping and kicking differ substantially. Neonates kicked with an adult-like number of temporal activation patterns, which lacked a stable association with systematic muscle synergies across movements. However, on the ground neonates stepped with fewer temporal patterns but all structured in stable synergies. Since kicking and ground-stepping coexist at birth, switching between the two behaviors may depend on a dynamic reconfiguration of the underlying neural circuits as a function of sensory feedback from surface contact. We tracked the development of ground-stepping in 4- to 48-mo-old infants and found that, after the age of 6 mo, the number of temporal patterns increased progressively, reaching adult-like conformation only after independent walking was established. We surmise that mature locomotor modules may derive by combining the multiple patterns of repeated kicking, on the one hand, with synergies resulting from fractionation of those revealed by sporadic weight-bearing stepping, on the other hand.

Virtual Stiffness: A Novel Biomechanical Approach to Estimate Limb Stiffness of a Multi-Muscle and Multi-Joint System.

In recent years, different groups have developed algorithms to control the stiffness of a robotic device through the electromyographic activity collected from a human operator. However, the approaches proposed so far require an initial calibration, have a complex subject-specific muscle model, or consider the activity of only a few pairs of antagonist muscles. This study described and tested an approach based on a biomechanical model to estimate the limb stiffness of a multi-joint, multi-muscle system from muscle activations. The "virtual stiffness" method approximates the generated stiffness as the stiffness due to the component of the muscle-activation vector that does not generate any endpoint force. Such a component is calculated by projecting the vector of muscle activations, estimated from the electromyographic signals, onto the null space of the linear mapping of muscle activations onto the endpoint force. The proposed method was tested by using an upper-limb model made of two joints and six Hill-type muscles and data collected during an isometric force-generation task performed with the upper limb. The null-space projection of the muscle-activation vector approximated the major axis of the stiffness ellipse or ellipsoid. The model provides a good approximation of the voluntary stiffening performed by participants that could be directly implemented in wearable myoelectric controlled devices that estimate, in real-time, the endpoint forces, or endpoint movement, from the mapping between muscle activation and force, without any additional calibrations.

Mixed matrix factorization: a novel algorithm for the extraction of kinematic-muscular synergies.

Synergistic models have been employed to investigate motor coordination separately in the muscular and kinematic domains. However, the relationship between muscle synergies, constrained to be non-negative, and kinematic synergies, whose elements can be positive and negative, has received limited attention. Existing algorithms for extracting synergies from combined kinematic and muscular data either do not enforce non-negativity constraints or separate non-negative variables into positive and negative components. We propose a mixed matrix factorization (MMF) algorithm based on a gradient descent update rule that overcomes these limitations. It allows to directly assess the relationship between kinematic and muscle activity variables, by enforcing the non-negativity constrain on a subset of variables. We validated the algorithm on simulated kinematic-muscular data generated from known spatial synergies and temporal coefficients, by evaluating the similarity between extracted and ground truth synergies and temporal coefficients when the data are corrupted by different noise levels. We also compared the performance of MMF to that of non-negative matrix factorization applied to separate positive and negative components (NMFpn). Finally, we factorized kinematic and electromyographic data collected during upper-limb movements to demonstrate the potential of the algorithm. MMF achieved almost perfect reconstruction on noiseless simulated data. It performed better than NMFpn in recovering the correct spatial synergies and temporal coefficients with noisy simulated data. It also allowed to correctly select the original number of ground truth synergies. We showed meaningful applicability to real data; MMF can also be applied to any multivariate data that contain both non-negative and unconstrained variables.NEW & NOTEWORTHY The mixed matrix factorization (MMF) is a novel method for extracting kinematic-muscular synergies. The previous state of the art algorithm (NMFpn) factorizes separately positive and rectified negative waveforms; the MMF instead employs a gradient descent method to factorize mixed kinematic unconstrained data and muscular non-negative data. MMF achieves perfect reconstruction on noiseless data, improving the NMFpn. MMF shows promising applicability on real data.

Task space exploration improves adaptation after incompatible virtual surgeries.

Humans have a remarkable capacity to learn new motor skills, a process that requires novel muscle activity patterns. Muscle synergies may simplify the generation of muscle patterns through the selection of a small number of synergy combinations. Learning of new motor skills may then be achieved by acquiring novel muscle synergies. In a previous study, we used myoelectric control to construct virtual surgeries that altered the mapping from muscle activity to cursor movements. After compatible virtual surgeries, which could be compensated by recombining subject-specific muscle synergies, participants adapted quickly. In contrast, after incompatible virtual surgeries, which could not be compensated by recombining existing synergies, participants explored new muscle patterns but failed to adapt. Here, we tested whether task space exploration can promote learning of novel muscle synergies required to overcome an incompatible surgery. Participants performed the same reaching task as in our previous study but with more time to complete each trial, thus allowing for exploration. We found an improvement in trial success after incompatible virtual surgeries. Remarkably, improvements in movement direction accuracy after incompatible surgeries occurred faster for corrective movements than for the initial movement, suggesting that learning of new synergies is more effective when used for feedback control. Moreover, reaction time was significantly higher after incompatible than compatible virtual surgeries, suggesting an increased use of an explicit adaptive strategy to overcome incompatible surgeries. Taken together, these results indicate that exploration is important for skill learning and suggest that human participants, with sufficient time, can learn new muscle synergies.NEW & NOTEWORTHY Motor skill learning requires the acquisition of novel muscle patterns, a slow adaptive process. Here we show that learning to control a cursor after an incompatible virtual surgery, a complex skill requiring new muscle synergies, is possible when enough time for task space exploration is provided. Our results suggest that learning new synergies is related to the exceptional human capacity to acquire a wide variety of novel motor skills with practice.

Evaluation of Spatiotemporal Patterns of the Spinal Muscle Coordination Output during Walking in the Exoskeleton.

Recent advances in the performance and evaluation of walking in exoskeletons use various assessments based on kinematic/kinetic measurements. While such variables provide general characteristics of gait performance, only limited conclusions can be made about the neural control strategies. Moreover, some kinematic or kinetic parameters are a consequence of the control implemented on the exoskeleton. Therefore, standard indicators based on kinematic variables have limitations and need to be complemented by performance measures of muscle coordination and control strategy. Knowledge about what happens at the spinal cord output level might also be critical for rehabilitation since an abnormal spatiotemporal integration of activity in specific spinal segments may result in a risk for abnormalities in gait recovery. Here we present the PEPATO software, which is a benchmarking solution to assess changes in the spinal locomotor output during walking in the exoskeleton with respect to reference data on normal walking. In particular, functional and structural changes at the spinal cord level can be mapped into muscle synergies and spinal maps of motoneuron activity. A user-friendly software interface guides the user through several data processing steps leading to a set of performance indicators as output. We present an example of the usage of this software for evaluating walking in an unloading exoskeleton that allows a person to step in simulated reduced (the Moon's) gravity. By analyzing the EMG activity from lower limb muscles, the algorithms detected several performance indicators demonstrating differential adaptation (shifts in the center of activity, prolonged activation) of specific muscle activation modules and spinal motor pools and increased coactivation of lumbar and sacral segments. The software is integrated at EUROBENCH facilities to benchmark the performance of walking in the exoskeleton from the point of view of changes in the spinal locomotor output.

Does the cerebellum shape the spatiotemporal organization of muscle patterns? Insights from subjects with cerebellar ataxias.

The role of the cerebellum in motor control has been investigated extensively, but its contribution to the muscle pattern organization underlying goal-directed movements is still not fully understood. Muscle synergies may be used to characterize multimuscle pattern organization irrespective of time (spatial synergies), in time irrespective of the muscles (temporal synergies), and both across muscles and in time (spatiotemporal synergies). The decomposition of muscle patterns as combinations of different types of muscle synergies offers the possibility to identify specific changes due to neurological lesions. In this study, we recorded electromyographic activity from 13 shoulder and arm muscles in subjects with cerebellar ataxias (CA) and in age-matched healthy subjects (HS) while they performed reaching movements in multiple directions. We assessed whether cerebellar damage affects the organization of muscle patterns by extracting different types of muscle synergies from the muscle patterns of each HS and using these synergies to reconstruct the muscle patterns of all other participants. We found that CA muscle patterns could be accurately captured only by spatial muscle synergies of HS. In contrast, there were significant differences in the reconstruction R2 values for both spatiotemporal and temporal synergies, with an interaction between the two synergy types indicating a larger difference for spatiotemporal synergies. Moreover, the reconstruction quality using spatiotemporal synergies correlated with the severity of impairment. These results indicate that cerebellar damage affects the temporal and spatiotemporal organization, but not the spatial organization, of the muscle patterns, suggesting that the cerebellum plays a key role in shaping their spatiotemporal organization.NEW & NOTEWORTHY In recent studies, the decomposition of muscle activity patterns has revealed a modular organization of the motor commands. We show, for the first time, that muscle patterns of subjects with cerebellar damage share with healthy controls spatial, but not temporal and spatiotemporal, modules. Moreover, changes in spatiotemporal organization characterize the severity of the subject's impairment. These results suggest that the cerebellum has a specific role in shaping the spatiotemporal organization of the muscle patterns.

A bang-bang control model predicts the triphasic muscles activity during hand reaching.

There are numerous ways to reach for an apple hanging from a tree. Yet, our motor system uses a specific muscle activity pattern that features activity bursts and silent periods. We suggest that these bursts are an evidence against the common view that the brain controls the commands to the muscles in a smooth continuous manner. Instead, we propose a model in which a motor plan is transformed into a piecewise-constant control signal that is low-pass filtered and transmitted to the muscles with different muscle-specific delays. We use a Markov chain Monte Carlo (MCMC) method to identify transitions in the state of the muscles following initial activation and show that fitting a bang-bang control model to the kinematics of movement predicts these transitions in the state of the muscles. Such a bang-bang controller suggests that the brain reduces the complexity of the problem of ballistic movements control by sending commands to the muscles at sparse times. Identifying this bang-bang controller can be useful to develop efficient controllers for neuroprostheses and other physical human-robot interaction systems.NEW & NOTEWORTHY While ballistic hand reaching movements are characterized by smooth position and velocity signals, the activity of the muscles exhibits bursts and silent periods. Here, we propose that a model based on bang-bang control provides the link between the abrupt changes in the muscle activity and the smooth reaching trajectory. Using bang-bang control instead of continuous control may simplify the design of prostheses and other physical human-robot interaction systems.

Contraction level, but not force direction or wrist position, affects the spatial distribution of motor unit recruitment in the biceps brachii muscle.

PURPOSE: Different motor units (MUs) in the biceps brachii (BB) muscle have been shown to be preferentially recruited during either elbow flexion or supination. Whether these different units reside within different regions is an open issue. In this study, we tested wheter MUs recruited during submaximal isometric tasks of elbow flexion and supination for two contraction levels and with the wrist fixed at two different angles are spatially localized in different BB portions. METHODS: The MUs' firing instants were extracted by decomposing high-density surface electromyograms (EMG), detected from the BB muscle of 12 subjects with a grid of electrodes (4 rows along the BB longitudinal axis, 16 columns medio-laterally). The firing instants were then used to trigger and average single-differential EMGs. The average rectified value was computed separately for each signal and the maximal value along each column in the grid was retained. The center of mass, defined as the weighted mean of the maximal, average rectified value across columns, was then consdiered to assess the medio-lateral changes in the MU surface representation between conditions. RESULTS: Contraction level, but neither wrist position nor force direction (flexion vs. supination), affected the spatial distribution of BB MUs. In particular, higher forces were associated with the recruitment of BB MUs whose action potentials were represented more medially. CONCLUSION: Although the action potentials of BB MUs were represented locally across the muscle medio-lateral region, dicrimination between elbow flexion or supination seems unlikely from the surface representation of MUs action potentials.

A whole body characterization of individual strategies, gender differences, and common styles in overarm throwing.

Overarm throwing is a fundamental human skill. Since paleolithic hunter-gatherer societies, the ability of throwing played a key role in brain and body co-evolution. For decades, throwing skill acquisition has been the subject of developmental and gender studies. However, due to its complex multijoint nature, whole body throwing has found little space in quantitative studies of motor behavior. In this study we examined how overarm throwing varies within and between individuals in a sample of untrained adults. To quantitatively compare whole body kinematics across throwing actions, we introduced a new combination of spatiotemporal principal component, linear discrimination, and clustering analyses. We found that the identity and gender of a thrower can be robustly inferred by the kinematics of a single throw, reflecting the characteristic features in individual throwing strategies and providing a quantitative ground for the well-known differences between males and females in throwing behavior. We also identified four main classes of throwing strategies, stable within individuals and resembling the main stages of throwing proficiency acquisition during motor development. These results support earlier proposals linking interindividual and gender differences in throwing, with skill acquisition interrupted at different stages of the typical developmental trajectory of throwing motor behavior.NEW & NOTEWORTHY Unconstrained throwing, because of its complexity, received little attention in quantitative motor control studies. By introducing a new approach to analyze whole body kinematics, we quantitatively characterized gender effects, interindividual differences, and common patterns in nontrained throwers. The four throwing styles identified across individuals resemble different stages in the acquisition of throwing skills during development. These results advance our understanding of complex motor skills, bridging the gap between motor control, motor development, and sport science.

Intercepting virtual balls approaching under different gravity conditions: evidence for spatial prediction.

To accurately time motor responses when intercepting falling balls we rely on an internal model of gravity. However, whether and how such a model is also used to estimate the spatial location of interception is still an open question. Here we addressed this issue by asking 25 participants to intercept balls projected from a fixed location 6 m in front of them and approaching along trajectories with different arrival locations, flight durations, and gravity accelerations (0g and 1g). The trajectories were displayed in an immersive virtual reality system with a wide field of view. Participants intercepted approaching balls with a racket, and they were free to choose the time and place of interception. We found that participants often achieved a better performance with 1g than 0g balls. Moreover, the interception points were distributed along the direction of a 1g path for both 1g and 0g balls. In the latter case, interceptions tended to cluster on the upper half of the racket, indicating that participants aimed at a lower position than the actual 0g path. These results suggest that an internal model of gravity was probably used in predicting the interception locations. However, we found that the difference in performance between 1g and 0g balls was modulated by flight duration, the difference being larger for faster balls. In addition, the number of peaks in the hand speed profiles increased with flight duration, suggesting that visual information was used to adjust the motor response, correcting the prediction to some extent.NEW & NOTEWORTHY Here we show that an internal model of gravity plays a key role in predicting where to intercept a fast-moving target. Participants also assumed an accelerated motion when intercepting balls approaching in a virtual environment at constant velocity. We also show that the role of visual information in guiding interceptive movement increases when more time is available.

Representation of Muscle Synergies in the Primate Brain.

Evidence suggests that the CNS uses motor primitives to simplify movement control, but whether it actually stores primitives instead of computing solutions on the fly to satisfy task demands is a controversial and still-unanswered possibility. Also in contention is whether these primitives take the form of time-invariant muscle coactivations ("spatial" synergies) or time-varying muscle commands ("spatiotemporal" synergies). Here, we examined forelimb muscle patterns and motor cortical spiking data in rhesus macaques (Macaca mulatta) handling objects of variable shape and size. From these data, we extracted both spatiotemporal and spatial synergies using non-negative decomposition. Each spatiotemporal synergy represents a sequence of muscular or neural activations that appeared to recur frequently during the animals' behavior. Key features of the spatiotemporal synergies (including their dimensionality, timing, and amplitude modulation) were independently observed in the muscular and neural data. In addition, both at the muscular and neural levels, these spatiotemporal synergies could be readily reconstructed as sequential activations of spatial synergies (a subset of those extracted independently from the task data), suggestive of a hierarchical relationship between the two levels of synergies. The possibility that motor cortex may execute even complex skill using spatiotemporal synergies has novel implications for the design of neuroprosthetic devices, which could gain computational efficiency by adopting the discrete and low-dimensional control that these primitives imply. SIGNIFICANCE STATEMENT: We studied the motor cortical and forearm muscular activity of rhesus macaques (Macaca mulatta) as they reached, grasped, and carried objects of varied shape and size. We applied non-negative matrix factorization separately to the cortical and muscular data to reduce their dimensionality to a smaller set of time-varying "spatiotemporal" synergies. Each synergy represents a sequence of cortical or muscular activity that recurred frequently during the animals' behavior. Salient features of the synergies (including their dimensionality, timing, and amplitude modulation) were observed at both the cortical and muscular levels. The possibility that the brain may execute even complex behaviors using spatiotemporal synergies has implications for neuroprosthetic algorithm design, which could become more computationally efficient by adopting the discrete and low-dimensional control that they afford.

Modularity for Motor Control and Motor Learning.

How the central nervous system (CNS) overcomes the complexity of multi-joint and multi-muscle control and how it acquires or adapts motor skills are fundamental and open questions in neuroscience. A modular architecture may simplify control by embedding features of both the dynamic behavior of the musculoskeletal system and of the task into a small number of modules and by directly mapping task goals into module combination parameters. Several studies of the electromyographic (EMG) activity recorded from many muscles during the performance of different tasks have shown that motor commands are generated by the combination of a small number of muscle synergies, coordinated recruitment of groups of muscles with specific amplitude balances or activation waveforms, thus supporting a modular organization of motor control. Modularity may also help understanding motor learning. In a modular architecture, acquisition of a new motor skill or adaptation of an existing skill after a perturbation may occur at the level of modules or at the level of module combinations. As learning or adapting an existing skill through recombination of modules is likely faster than learning or adapting a skill by acquiring new modules, compatibility with the modules predicts learning difficulty. A recent study in which human subjects used myoelectric control to move a mass in a virtual environment has tested this prediction. By altering the mapping between recorded muscle activity and simulated force applied on the mass, as in a complex surgical rearrangement of the tendons, it has been possible to show that it is easier to adapt to a perturbation that is compatible with the muscle synergies used to generate hand force than to a similar but incompatible perturbation. This result provides direct support for a modular organization of motor control and motor learning.

Differences in adaptation rates after virtual surgeries provide direct evidence for modularity.

Whether the nervous system relies on modularity to simplify acquisition and control of complex motor skills remains controversial. To date, evidence for modularity has been indirect, based on statistical regularities in the motor commands captured by muscle synergies. Here we provide direct evidence by testing the prediction that in a truly modular controller it must be harder to adapt to perturbations that are incompatible with the modules. We investigated a reaching task in which human subjects used myoelectric control to move a mass in a virtual environment. In this environment we could perturb the normal muscle-to-force mapping, as in a complex surgical rearrangement of the tendons, by altering the mapping between recorded muscle activity and simulated force applied on the mass. After identifying muscle synergies, we performed two types of virtual surgeries. After compatible virtual surgeries, a full range of movements could still be achieved recombining the synergies, whereas after incompatible virtual surgeries, new or modified synergies would be required. Adaptation rates after the two types of surgery were compared. If synergies were only a parsimonious description of the regularities in the muscle patterns generated by a nonmodular controller, we would expect adaptation rates to be similar, as both types of surgeries could be compensated with similar changes in the muscle patterns. In contrast, as predicted by modularity, we found strikingly faster adaptation after compatible surgeries than after incompatible ones. These results indicate that muscle synergies are key elements of a modular architecture underlying motor control and adaptation.

Myoelectric control and virtual reality to enhance motor rehabilitation after stroke.

Effective upper-limb rehabilitation for severely impaired stroke survivors is still missing. Recent studies endorse novel motor rehabilitation approaches such as robotic exoskeletons and virtual reality systems to restore the function of the paretic limb of stroke survivors. However, the optimal way to promote the functional reorganization of the central nervous system after a stroke has yet to be uncovered. Electromyographic (EMG) signals have been employed for prosthetic control, but their application to rehabilitation has been limited. Here we propose a novel approach to promote the reorganization of pathological muscle activation patterns and enhance upper-limb motor recovery in stroke survivors by using an EMG-controlled interface to provide personalized assistance while performing movements in virtual reality (VR). We suggest that altering the visual feedback to improve motor performance in VR, thereby reducing the effect of deviations of the actual, dysfunctional muscle patterns from the functional ones, will actively engage patients in motor learning and facilitate the restoration of functional muscle patterns. An EMG-controlled VR interface may facilitate effective rehabilitation by targeting specific changes in the structure of muscle synergies and in their activations that emerged after a stroke-offering the possibility to provide rehabilitation therapies addressing specific individual impairments.

SynergyAnalyzer: A Matlab toolbox implementing mixed-matrix factorization to identify kinematic-muscular synergies.

BACKGROUND AND OBJECTIVE: A new direction in the study of motor control was opened about two decades ago with the introduction of a model for the generation of motor commands as combination of muscle synergies. Muscle synergies provide a simple yet quantitative framework for analyzing the hierarchical and modular architecture of the human motor system. However, to gain insights on the functional role of muscle synergies, they should be related to the task space. The recently introduced mixed-matrix factorization (MMF) algorithm extends the standard approach for synergy extraction based on non-negative matrix factorization (NMF) allowing to factorize data constituted by a mixture of non-negative variables (e.g. EMGs) and unconstrained variables (e.g. kinematics, naturally including both positive and negative values). The kinematic-muscular synergies identified by MMF provide a direct link between muscle synergies and the task space. In this contribution, we support the adoption of MMF through a Matlab toolbox for the extraction of kinematic-muscular synergies and a set of practical guidelines to allow biomedical researchers and clinicians to exploit the potential of this novel approach. METHODS: MMF is implemented in the SynergyAnalyzer toolbox using an object-oriented approach. In addition to the MMF algorithm, the toolbox includes standard methods for synergy extraction (NMF and PCA), as well as methods for pre-processing EMG and kinematic data, and for plotting data and synergies. RESULTS: As an example of MMF application, kinematic-muscular synergies were extracted from EMG and kinematic data collected during reaching movements towards 8 targets on the sagittal plane. Instructions and command lines to achieve such results are illustrated in detail. The toolbox has been released as an open-source software on GitHub under the GNU General Public License. CONCLUSIONS: Thanks to its ease of use and adaptability to a variety of datasets, SynergyAnalyzer will facilitate the adoption of MMF to extract kinematic-muscular synergies from mixed EMG and kinematic data, a useful approach in biomedical research to better understand and characterize the functional role of muscle synergies.