The Role of Heterarchical Control in the Evolution of Central Pattern Generators

The acceptance of the concept of central pattern generators (CPGs) led to the perception that descending inputs initiate stereotyped movements, such as locomotion, but play relatively minor roles after the movement begins. Sensory input could entrain the CPG, and the CPG was responsive to the proper inputs for switching, etc. Evidence is here presented that the influences of both descending and sensory inputs are two-way. Descending inputs are shown to be involved in an ongoing manner during locomotion, as it has been found that CPGs are phasically driving the same descending systems that themselves activate the CPGs. Similarly, sensory inputs are being actively processed by the CPG and, here again, produce a two-way interaction between sensory input and CPGs. Finally, mechanical factors are shown to be major contributors to the form of the movement. Thus, overall the CPG can only be considered as one of several contributors to any movement; all concurrently process the flow of information. Control is viewed as distributed, that is, as heterarchical as opposed to hierarchical. Because of the complexity of interaction between the levels of the system, it is argued that any change in the body will propagate through the system and effect the final output regardless of where the change originates. Thus, ontogenetic and phylogenetic changes must have influences felt throughout the entire system. Examples are presented to demonstrate that this is, indeed, the case. The effect of these changes will primarily be manifest in the interface between the layers, that is, in the interface between the descending and sensory inputs and the CPG, so that the changes can be adaptively accommodated on a moment by moment basis when necessary.