Do muscle synergies reduce the dimensionality of behavior?

The muscle synergy hypothesis is an archetype of the notion of Dimensionality Reduction (DR) occurring in the central nervous system due to modular organisation. Towards validating this hypothesis, it is however important to understand if muscle synergies can reduce the state-space dimensionality while suitably achieving task control. In this paper we present a scheme for investigating this reduction, utilising the temporal muscle synergy formulation. Our approach is based on the observation that constraining the control input to a weighted combination of temporal muscle synergies instead constrains the dynamic behaviour of a system in trajectory-specific manner. We compute this constrained reformulation of system dynamics and then use the method of system balancing for quantifying the DR; we term this approach as Trajectory Specific Dimensionality Analysis (TSDA). We then use this method to investigate the consequence of minimisation of this dimensionality for a given task. These methods are tested in simulation on a linear (tethered mass) and a nonlinear (compliant kinematic chain) system; dimensionality of various reaching trajectories is compared when using idealised temporal synergies. We show that as a consequence of this Minimum Dimensional Control (MDC) model, smooth straight-line Cartesian trajectories with bell-shaped velocity profiles are obtained as the solution to reaching tasks in both of the test systems. We also investigate the effect on dimensionality due to adding via-points to a trajectory. The results indicate that a synergy basis and trajectory-specific DR of motor behaviours results from usage of muscle synergy control. The implications of these results for the synergy hypothesis, optimal motor control, developmental skill acquisition and robotics are then discussed

Muscle synergies in neuroscience and robotics: from input-space to task-space perspectives

In this paper we review the works related to muscle synergies that have been carried-out in neuroscience and control engineering. In particular, we refer to the hypothesis that the central nervous system (CNS) generates desired muscle contractions by combining a small number of predefined modules, called muscle synergies. We provide an overview of the methods that have been employed to test the validity of this scheme, and we show how the concept of muscle synergy has been generalized for the control of artificial agents. The comparison between these two lines of research, in particular their different goals and approaches, is instrumental to explain the computational implications of the hypothesized modular organization. Moreover, it clarifies the importance of assessing the functional role of muscle synergies: although these basic modules are defined at the level of muscle activations (input-space), they should result in the effective accomplishment of the desired task. This requirement is not always explicitly considered in experimental neuroscience, as muscle synergies are often estimated solely by analyzing recorded muscle activities. We suggest that synergy extraction methods should explicitly take into account task execution variables, thus moving from a perspective purely based on input-space to one grounded on task-space as well

On Neuromechanical Approaches for the Study of Biological Grasp and Manipulation

Biological and robotic grasp and manipulation are undeniably similar at the level of mechanical task performance. However, their underlying fundamental biological vs. engineering mechanisms are, by definition, dramatically different and can even be antithetical. Even our approach to each is diametrically opposite: inductive science for the study of biological systems vs. engineering synthesis for the design and construction of robotic systems. The past 20 years have seen several conceptual advances in both fields and the quest to unify them. Chief among them is the reluctant recognition that their underlying fundamental mechanisms may actually share limited common ground, while exhibiting many fundamental differences. This recognition is particularly liberating because it allows us to resolve and move beyond multiple paradoxes and contradictions that arose from the initial reasonable assumption of a large common ground. Here, we begin by introducing the perspective of neuromechanics, which emphasizes that real-world behavior emerges from the intimate interactions among the physical structure of the system, the mechanical requirements of a task, the feasible neural control actions to produce it, and the ability of the neuromuscular system to adapt through interactions with the environment. This allows us to articulate a succinct overview of a few salient conceptual paradoxes and contradictions regarding under-determined vs. over-determined mechanics, under- vs. over-actuated control, prescribed vs. emergent function, learning vs. implementation vs. adaptation, prescriptive vs. descriptive synergies, and optimal vs. habitual performance. We conclude by presenting open questions and suggesting directions for future research. We hope this frank assessment of the state-of-the-art will encourage and guide these communities to continue to interact and make progress in these important areas

Challenges and New Approaches to Proving the Existence of Muscle Synergies of Neural Origin

Muscle coordination studies repeatedly show low-dimensionality of muscle activations for a wide variety of motor tasks. The basis vectors of this low-dimensional subspace, termed muscle synergies, are hypothesized to reflect neurally-established functional muscle groupings that simplify body control. However, the muscle synergy hypothesis has been notoriously difficult to prove or falsify. We use cadaveric experiments and computational models to perform a crucial thought experiment and develop an alternative explanation of how muscle synergies could be observed without the nervous system having controlled muscles in groups. We first show that the biomechanics of the limb constrains musculotendon length changes to a low-dimensional subspace across all possible movement directions. We then show that a modest assumption—that each muscle is independently instructed to resist length change—leads to the result that electromyographic (EMG) synergies will arise without the need to conclude that they are a product of neural coupling among muscles. Finally, we show that there are dimensionality-reducing constraints in the isometric production of force in a variety of directions, but that these constraints are more easily controlled for, suggesting new experimental directions. These counter-examples to current thinking clearly show how experimenters could adequately control for the constraints described here when designing experiments to test for muscle synergies—but, to the best of our knowledge, this has not yet been done

In silico case studies of compliant robots: AMARSI deliverable 3.3

In the deliverable 3.2 we presented how the morphological computing ap- proach can significantly facilitate the control strategy in several scenarios, e.g. quadruped locomotion, bipedal locomotion and reaching. In particular, the Kitty experimental platform is an example of the use of morphological computation to allow quadruped locomotion. In this deliverable we continue with the simulation studies on the application of the different morphological computation strategies to control a robotic system

Probabilistic Models of Motor Production

N. Bernstein defined the ability of the central neural system (CNS) to control many degrees of freedom of a physical body with all its redundancy and flexibility as the main problem in motor control. He pointed at that man-made mechanisms usually have one, sometimes two degrees of freedom (DOF); when the number of DOF increases further, it becomes prohibitively hard to control them. The brain, however, seems to perform such control effortlessly. He suggested the way the brain might deal with it: when a motor skill is being acquired, the brain artificially limits the degrees of freedoms, leaving only one or two. As the skill level increases, the brain gradually "frees" the previously fixed DOF, applying control when needed and in directions which have to be corrected, eventually arriving to the control scheme where all the DOF are "free". This approach of reducing the dimensionality of motor control remains relevant even today. One the possibles solutions of the Bernstetin's problem is the hypothesis of motor primitives (MPs) - small building blocks that constitute complex movements and facilitite motor learnirng and task completion. Just like in the visual system, having a homogenious hierarchical architecture built of similar computational elements may be beneficial. Studying such a complicated object as brain, it is important to define at which level of details one works and which questions one aims to answer. David Marr suggested three levels of analysis: 1. computational, analysing which problem the system solves; 2. algorithmic, questioning which representation the system uses and which computations it performs; 3. implementational, finding how such computations are performed by neurons in the brain. In this thesis we stay at the first two levels, seeking for the basic representation of motor output. In this work we present a new model of motor primitives that comprises multiple interacting latent dynamical systems, and give it a full Bayesian treatment. Modelling within the Bayesian framework, in my opinion, must become the new standard in hypothesis testing in neuroscience. Only the Bayesian framework gives us guarantees when dealing with the inevitable plethora of hidden variables and uncertainty. The special type of coupling of dynamical systems we proposed, based on the Product of Experts, has many natural interpretations in the Bayesian framework. If the dynamical systems run in parallel, it yields Bayesian cue integration. If they are organized hierarchically due to serial coupling, we get hierarchical priors over the dynamics. If one of the dynamical systems represents sensory state, we arrive to the sensory-motor primitives. The compact representation that follows from the variational treatment allows learning of a motor primitives library. Learned separately, combined motion can be represented as a matrix of coupling values. We performed a set of experiments to compare different models of motor primitives. In a series of 2-alternative forced choice (2AFC) experiments participants were discriminating natural and synthesised movements, thus running a graphics Turing test. When available, Bayesian model score predicted the naturalness of the perceived movements. For simple movements, like walking, Bayesian model comparison and psychophysics tests indicate that one dynamical system is sufficient to describe the data. For more complex movements, like walking and waving, motion can be better represented as a set of coupled dynamical systems. We also experimentally confirmed that Bayesian treatment of model learning on motion data is superior to the simple point estimate of latent parameters. Experiments with non-periodic movements show that they do not benefit from more complex latent dynamics, despite having high kinematic complexity. By having a fully Bayesian models, we could quantitatively disentangle the influence of motion dynamics and pose on the perception of naturalness. We confirmed that rich and correct dynamics is more important than the kinematic representation. There are numerous further directions of research. In the models we devised, for multiple parts, even though the latent dynamics was factorized on a set of interacting systems, the kinematic parts were completely independent. Thus, interaction between the kinematic parts could be mediated only by the latent dynamics interactions. A more flexible model would allow a dense interaction on the kinematic level too. Another important problem relates to the representation of time in Markov chains. Discrete time Markov chains form an approximation to continuous dynamics. As time step is assumed to be fixed, we face with the problem of time step selection. Time is also not a explicit parameter in Markov chains. This also prohibits explicit optimization of time as parameter and reasoning (inference) about it. For example, in optimal control boundary conditions are usually set at exact time points, which is not an ecological scenario, where time is usually a parameter of optimization. Making time an explicit parameter in dynamics may alleviate this

Common synaptic input, synergies and size principle: Control of spinal motor neurons for movement generation

Understanding how movement is controlled by the CNS remains a major challenge, with ongoing debate about basic features underlying this control. In current established views, the concepts of motor neuron recruitment order, common synaptic input to motor neurons and muscle synergies are usually addressed separately and therefore seen as independent features of motor control. In this review, we analyse the body of literature in a broader perspective and we identify a unified approach to explain apparently divergent observations at different scales of motor control. Specifically, we propose a new conceptual framework of the neural control of movement, which merges the concept of common input to motor neurons and modular control, together with the constraints imposed by recruitment order. This framework is based on the following assumptions: (1) motor neurons are grouped into functional groups (clusters) based on the common inputs they receive; (2) clusters may significantly differ from the classical definition of motor neuron pools, such that they may span across muscles and/or involve only a portion of a muscle; (3) clusters represent functional modules used by the CNS to reduce the dimensionality of the control; and (4) selective volitional control of single motor neurons within a cluster receiving common inputs cannot be achieved. Here, we discuss this framework and its underlying theoretical and experimental evidence

A Framework for Human Motion Strategy Identification and Analysis

The human body has many biomechanical degrees of freedom and thus multiple movement strategies can be employed to execute any given task. Automated identification and classification of these movement strategies have potential applications in various fields including sports performance research, rehabilitation, and injury prevention. For example, in the field of rehabilitation, the choice of movement strategy can impact joint loading patterns and risk of injury. The problem of identifying movement strategies is related to the problem of classifying variations in the observed motions. When differences between two movement trajectories performing the same task are large, they are considered to be different movement strategies. Conversely, when the differences between observed movements are small, they are considered to be variations of the same movement strategy. In the simplest scenario a movement strategy can represent a cluster of similar movement trajectories, but in more complicated scenarios differences in movements could also lie on a continuum. The goal of this thesis is to develop a computational framework to automatically recognize different movement strategies for performing a task and to identify what makes each strategy different. The proposed framework utilizes Gaussian Process Dynamical Models (GPDM) to convert human motion trajectories from their original high dimensional representation to a trajectory in a lower dimensional space (i.e. the latent space). The dimensionality of the latent space is determined by iteratively increasing the dimensionality until the reduction in reconstruction error between iterations becomes small. Then, the lower dimensional trajectories are clustered using a Hidden Markov Model (HMM) clustering algorithm to identify movement strategies in an unsupervised manner. Next, we introduce an HMM-based technique for detecting differences in signals between two HMM models. This technique is used to compare latent space variables between the low-dimensional trajectory models as well as differences in degrees-of-freedom (DoF) between the corresponding high-dimensional (original) trajectory models. Then, through correlating latent variable and DoF differences movement synergies are discovered. To validate the proposed framework, it was tested on 3 different datasets – a synthetic dataset, a real labeled motion capture dataset, and an unlabeled motion capture dataset. The proposed framework achieved higher classification accuracy against competing algorithms (Joint Component Vector and Kinematic Synergies) where labels were known apriori. Additionally, the proposed algorithm showed that it was able to discover strategies that were not known apriori and how the strategies differed

Neural bases of hand synergies

abstract: The human hand has so many degrees of freedom that it may seem impossible to control. A potential solution to this problem is “synergy control” which combines dimensionality reduction with great flexibility. With applicability to a wide range of tasks, this has become a very popular concept. In this review, we describe the evolution of the modern concept using studies of kinematic and force synergies in human hand control, neurophysiology of cortical and spinal neurons, and electromyographic (EMG) activity of hand muscles. We go beyond the often purely descriptive usage of synergy by reviewing the organization of the underlying neuronal circuitry in order to propose mechanistic explanations for various observed synergy phenomena. Finally, we propose a theoretical framework to reconcile important and still debated concepts such as the definitions of “fixed” vs. “flexible” synergies and mechanisms underlying the combination of synergies for hand control.View the article as published at http://journal.frontiersin.org/article/10.3389/fncom.2013.00023/ful