Abstract

In many situations, the human movement system has more degrees of freedom than needed to achieve a given movement task. Martin et al. (Neural Comput 21(5):1371–1414, 2009) accounted for signatures of such redundancy like self-motion and motor equivalence in a process model in which a neural oscillator generated timed end-effector virtual trajectories that a neural dynamics transformed into joint virtual trajectories while decoupling task-relevant and task-irrelevant combinations of joint angles. Neural control of muscle activation and the biomechanical dynamics of the arm were taken into account. The model did not address the main signature of redundancy, however, the UCM structure of variance: Many experimental studies have shown that across repetitions, variance of joint configuration trajectories is structured. Combinations of joint angles that affect task variables (lying in the uncontrolled manifold, UCM) are much more variable than combinations of joint angles that do not. This finding has been robust across movement systems, age, and tasks and is often preserved in clinical populations as well. Here, we provide an account for the UCM structure of variance by adding four types of noise sources to the model of Martin et al. (Neural Comput 21(5):1371–1414, 2009). Comparing the model to human pointing movements and systematically examining the role of each noise source and mechanism, we identify three causes of the UCM effect, all of which, we argue, contribute: (1) the decoupling of motor commands across the task-relevant and task-irrelevant subspaces together with “neural” noise at the level of these motor commands; (2) “muscle noise” combined with imperfect control of the limb; (3) back-coupling of sensed joint configurations into the motor commands which then yield to the sensed joint configuration within the UCM.

Highlights

  • How are the many kinematic degrees of freedom (DoF) of the human motor system and the muscles that actuate them harnessed to achieve a particular movement goal? This is Communicated by Benjamin Lindner.the classical “degree of freedom problem” that has been a major theme in motor control at least since Bernstein (1967)

  • In a previous theoretical analysis (Martin et al 2009), we examined some of the consequences of this hypothesis that the DoF problem is solved through a kinematic transformation

  • These include (1) a process for generating the time course of the movement as a virtual end-effector trajectory, (2) a dynamical system that transforms the virtual end-effector trajectory into motor commands for the muscle-joint systems by decoupling those combinations of degrees of freedom that move the end-effector from those combinations that do not, (3) and a neural dynamics of muscular control that generate torques in response to descending motor commands and proprioceptive feedback

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Summary

Introduction

How are the many kinematic degrees of freedom (DoF) of the human motor system and the muscles that actuate them harnessed to achieve a particular movement goal? This is Communicated by Benjamin Lindner.the classical “degree of freedom problem” that has been a major theme in motor control at least since Bernstein (1967). The problem arises because motor systems are redundant (or abundant, see Gelfand and Latash 1998) for many tasks; that is, there are typically more joints or muscles available than needed to control a set of task variables like the position or orientation of the hand in space. It is not clear, a priori, that this is a problem for the central nervous system (CNS) because, in principle, the neural networks that generate and control movement could be structured such that they provide one particular solution among the many possible ones. This intuition is based on empirical evidence that the CNS is flexible about which particular solution is employed

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