The functional sense of central oscillations in walking

Abstract.

 Rhythmic motor output is generally assumed to be produced by central pattern generators or, more specific, central oscillators, the rhythmic output of which can be entrained and modulated by sensory input and descending control. In the case of locomotor systems, the output of the central system, i.e., the output obtained after deafferentation of sensory feedback, shows many of the temporal characteristics of real movements. Therefore the term fictive locomotion has been coined. This article concentrates on a specific locomotor behavior, namely walking; in particular walking in invertebrates. In contrast to the traditional view, an alternative hypothesis is formulated to interpret the functional sense of these central oscillations which have been found in many cases. It is argued that the basic function of the underlying circuit is to avoid cocontraction of antagonistic muscles. Such a system operates best with an inherent period just above the maximum period observed in real walking. The circuit discussed in this article (Fig. 2) shows several properties in common with results described as “fictive walking”. It furthermore could explain a number of properties observed in animals walking in different situations. According to this hypothesis, the oscillations found after deafferentation are side effects occurring in specific artificial situations. If, however, a parameter called central excitation is large enough, the system can act as a central oscillator that overrides the sensory input completely.