A synergetic theory of quadrupedal gaits and gait transitions

We present a theoretical analysis of the patterns of interlimb co-ordination in the gaits of quadrupedal locomotion. Introducing as collective variables a set of relative phases that describe the co-ordination patterns, we classify gaits by their symmetry properties, which can be expressed as invariances under groups of transformations. We define dynamics of the collective variables, on which we impose symmetry restrictions. The stable observable gait patterns correspond to atractors of these dynamics. A non-trivial consequence of this theoretical viewpoint is that gait transitions can take the form of non-equilibrium phase transitions that are accompanied by loss of stability. We show how various types of such phase transitions involving hysteresis, slowing down and fluctuation enhancement can occur. Also the difference between smooth and abrupt transitions is given theoretical foundation. While existing experimental evidence is consistent with the theory developed here, we propose new experimental measures that can serve to test the present theoretical framework. Finally, the influence of underlying symmetries of the dynamics on the nature of the gait patterns and their stability is analyzed. For example, breaking of a front-hind symmetry can lead to a change from absolute to relative co-ordination in the sense of von Holst (1939, Ergebnisse der Physiologie 42, 228). Also, differential stability of straight and reverse gaits results from thus lowering the symmetry.