Patterns of human interlimb coordination emerge from the properties of non-linear, limit cycle oscillatory processes: theory and data

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). Homeo kinetic theory characterizes biological systems as ensembles of non-linear, limit cycle oscillatory processes couple and mutually entrained at all the 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 interaction among non-linear, limit cycle oscillatory processes.     

Spontaneous transitions and symmetry: Pattern dynamics in human four-limb coordination

Transitions between the coordinative patterns of rhythmically moving human arms and legs were studied to test the predictions of a four-component model (Schöner, Jiang and Kelso, 1990). Based upon results from previous two-component experiments (Kelso and Jeka, 1992), three assumptions were made about the four-limb system: (1) all limb pairs produce stable in-phase and anti-phase patterns; (2) the coupling between homologous limbs (i.e., right and left arms or right and left legs) is appreciably stronger than the coupling between nonhomologous limbs (i.e., arm and leg); and (3) right-left symmetry. An analysis of a four-component model (Jeka, Kelso and Kiemel, 1993) led to the prediction of four attracting invariant circles, each with two stable patterns in the state space of four-limb dynamics. In an experiment to test this prediction, subjects were required to cycle all four limbs in one of the eight patterns to the beat of an auditory metronome whose frequency was systematically increased. All subjects demonstrated spontaneous transitions corresponding to pathways along the invariant circles. Pre-transition relative phase variability increased with required frequency up to the transition, suggesting that loss of pattern stability induced the observed transitions. Thus, despite a large number of potential transitions, differential coupling between limb pairs and symmetry of the pattern dynamics restricts the behavior of the human four-limb system to a limited area of its state space.     

Steady-state and perturbed rhythmical movements: a dynamical analysis.

This study examined rhythmic finger movements in the steady state and when momentarily perturbed in order to derive their qualitative dynamical properties. Movement frequency, amplitude, and peak velocity were stable under perturbation, signaling the presence of an attractor, and the topological dimensionality of that attractor was approximately equal to one. The strength of the attractor was constant with increasing movement frequency, and the Fourier spectra of the steady-state trials showed an alternating harmonic pattern. These results are consistent with a previously derived nonlinear oscillator model. However, the oscillation was phase advanced by perturbation overall, and a consistent phase-dependent, phase-shift pattern occurred, which is inconsistent with the model. The overall phase advance also shows that any central pattern generator responsible for generating the rhythm must be nontrivially modulated by the limb being controlled.     

Disaster rescue work: The consequences for the family

This paper reports on the findings of a pilot study investigating the impact of disaster relief work on the wives and family members of the relief workers. The study revealed that the wives did experience stress, although the nature of the problems identifed changed with each phase of disaster involvement. Children may also be affected. The age of the child may be an important determinant of the nature and severity of their problems. Interventions that could be adopted to deal with these concerns are outlined.     

Tracking simple and complex sequences

We address issues of synchronization to rhythms of musical complexity. In two experiments, synchronization to simple and more complex rhythmic sequences was investigated. Experiment 1 examined responses to phase and tempo perturbations within simple, structurally isochronous sequences, presented at different base rates. Experiment 2 investigated responses to similar perturbations embedded within more complex, metrically structured sequences; participants were explicitly instructed to synchronize at different metrical levels (i.e., tap at different rates to the same rhythmic patterns) on different trials. We found evidence that (1) the intrinsic tapping frequency adapts in response to temporal perturbations in both simple (isochronous) and complex (metrically structured) rhythms, (2) people can synchronize with unpredictable, metrically structured rhythms at different metrical levels, with qualitatively different patterns of synchronization seen at higher versus lower levels of metrical structure, and (3) synchronization at each tapping level reflects information from other metrical levels. The latter finding provides evidence for a dynamic and flexible internal representation of the sequence's metrical structure.     

Coordination dynamics of learning and transfer across different effector systems

If different effector systems share a common task-specific coordination dynamics, transfer and generalization of sensorimotor learning are predicted. Subjects learned a visually specified phase relationship with either the arms or the legs. Coordination tendencies in both effector systems were evaluated before and after practice to detect attractive states of the coordination dynamics. Results indicated that learning a novel relative phase with a single effector system spontaneously transferred to the other, untrained effector system. Transfer was revealed not only as improvements in performance but also as modifications of each system's initial (prelearning) coordinative landscape. What is learned, appears to be a high-level but neurally instantiated dynamic representation of skilled behavior that proves to be largely effector independent, at least across anatomically symmetric limbs.     

Intentional Switching Between Patterns of Bimanual Coordination Depends on the Intrinsic Dynamics of the Patterns

Two predictions arising from previous theoretical and empirical work which demonstrated that spontaneous changes of bimanual coordination patterns result from a loss of pattern stability (i.e., a nonequilibrium phase transition) were tested: (a) that the time it takes to intentionally switch from one pattern to another depends on the differential stability of the patterns themselves; and (b) that an intention, defined as an intended behavioral pattern, can change the dynamical characteristics, e.g., the overall stability of the behavioral patterns. Subjects moved both index fingers rhythmically at one of six movement frequencies while performing either an in-phase or antiphase pattern of finger coordination. On cue from an auditory signal, subjects switched from the ongoing pattern to the other pattern. The relative phase of movement between the two fingers was used to characterize the ongoing coordinative pattern. The time taken to switch between patterns, or switching time, and relative phase fluctuations were used to evaluate the modified pattern dynamics resulting from a subject's intention to change patterns. Switching from the in-phase to the antiphase pattern was significantly slower than switching in the opposite direction for all subjects. Both the mean and distribution of switching times in each direction were found to be in agreement with model predictions. Movement frequency had little effect on switching time, a finding that is also consistent with the model. Relative phase fluctuations were significantly larger when moving in the antiphase pattern at the highest movement frequencies studied. The results show that, although intentional influences act to modify a coordinative pattern's intrinsic dynamics, the influence of these dynamics on the resulting behavior is always present and is particularly strong at high movement frequencies.     

Reconciling symbolic and dynamic aspects of language: Toward a dynamic psycholinguistics

The present paper examines natural language as a dynamical system. The oft-expressed view of language as “a static system of symbols” is here seen as an element of a larger system that embraces the mutuality of symbols and dynamics. Following along the lines of the theoretical biologist H.H. Pattee, the relation between symbolic and dynamic aspects of language is expressed within a more general framework that deals with the role of information in biological systems. In this framework, symbols are seen as information-bearing entities that emerge under pressures of communicative needs and that serve as concrete constraints on development and communication. In an attempt to identify relevant dynamic aspects of such a system, one has to take into account events that happen on different time scales: evolutionary language change (i.e., a diachronic aspect), processes of communication (language use) and language acquisition. Acknowledging the role of dynamic processes in shaping and sustaining the structures of natural language calls for a change in methodology. In particular, a purely synchronic analysis of a system of symbols as “meaning-containing entities” is not sufficient to obtain answers to certain recurring problems in linguistics and the philosophy of language. A more encompassing research framework may be the one designed specifically for studying informationally based coupled dynamical systems (coordination dynamics) in which processes of self-organization take place over different time scales.