Fifty years of CPGs: two neuroethological papers that shaped the course of neuroscience

Half a century ago, two independent papers that described unexpected results of experiments on locomotion in insects and crayfish appeared almost simultaneously. Together these papers demonstrated that an animal's central nervous system (CNS) was organized to produce behaviorally important motor output without the need for constant sensory feedback. These results contradicted the established line of thought that was based on interpretations of reflexes and ablation experiments, and established that in these animals the CNS contained neural circuits that could produce complex, periodic, multisegmental patterns of activity. These papers stimulated a flowering of research on central pattern-generating mechanisms that displaced reflex-based thinking everywhere except in medical physiology texts. Here we review these papers and their influence on thinking in the 1960s, 1970s, and today. We follow the development of ideas about central organization and control of expression of motor patterns, the roles of sensory input to central pattern-generating circuits, and integration of continuous sensory signals into a periodic motor system. We also review recent work on limb coordination that provides detailed cellular explanations of observations and speculations contained in those original papers.