Patterns of Human Interlimb Coordination Emerge from the Properties of Non-Linear, Limit Cycle Oscillatory Processes

Abstract

The present article represents an initial attempt to offer a principled solution to a fundamental problem of movement identified by Bernstein (1967), namely, how the degrees of freedom of the motor system are regulated. Conventional views of movement control focus on motor programs or closed-loop devices and have little or nothing to say on this matter. As an appropriate conceptual framework we offer Iberall and his colleagues' physical theory of homeokinetics first elaborated for movement by Kugler, Kelso, and Turvey (1980). Homeokinetic theory characterizes biological systems as ensembles of non-linear, limit cycle oscillatory processes coupled and mutually entrained at all levels of organization. Patterns of interlimb coordination may be predicted from the properties of non-linear, limit cycle oscillators. In a set of experiments and formal demonstrations we show that cyclical, two-handed movements maintain fixed amplitude and frequency (a stable limit cycle organization) under the following conditions: (a) when brief and constantly applied load perturbations are imposed on one hand or the other, (b) regardless of the presence or absence of fixed mechanical constraints, and (c) in the face of a range of external driving frequencies from a visual source. In addition, we observe a tight phasic relationship between the hands before and after perturbations (quantified by cross-correlation techniques), a tendency of one limb to entrain the other (mutual entrainment) and that limbs cycling at different frequencies reveal non-arbitrary, sub-harmonic relationships (small integer, subharmonic entrainment). In short, all the above patterns of interlimb coordination fall out of a non-linear oscillatory design. Discussion focuses on the compatibility of these results with past and present neurobiological work, and the theoretical insights into problems of movement offered by homeokinetic physics. Among these are, we think, the beginnings of a principled solution to the degrees of freedom problem, and the tentative claim that coordination and control are emergent consequences of dynamical interactions among non-linear, limit cycle oscillatory processes.