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A modular network for legged locomotion. (English) Zbl 1039.92009

Summary: We use symmetry methods to study networks of coupled cells, which are models for central pattern generators (CPGs). In these models the cells obey identical systems of differential equations and the network specifies how cells are coupled. Previously, J. J. Collins and I. Stewart [Biol. Cybern. 68, 287–298 (1993; Zbl 0767.92015); J. Nonlinear Sci. 3, 349–392 (1993; Zbl 0808.92012); Biol. Cybern. 71, 95–103 (1994; Zbl 0804.92009)] showed that the phase relations of many of the standard gaits of quadrupeds and hexapods can be obtained naturally via Hopf bifurcation in small networks. For example, the networks they used to study quadrupeds all had four cells, with the understanding that each cell determined the phase of the motion of one leg.
However, in their work it seemed necessary to employ several different four-oscillator networks to obtain all of the standard quadrupedal gaits. We show that this difficulty with four-oscillator networks is unavoidable, but that the problems can be overcome by using a larger network. Specifically, we show that the standard gaits of a quadruped, including walk, trot and pace, cannot all be realized by a single four-cell network without introducing unwanted conjugacies between trot and pace-conjugacies that imply a dynamic equivalence between these gaits that seems inconsistent with observations. In this sense a single network with four cells cannot model the CPG of a quadruped.
We also introduce a single eight-cell network that can model all of the primary gaits of quadrupeds without these unwanted conjugacies. Moreover, this network is modular in that it naturally generalizes to provide models of gaites in hexapods, centipeds and millipedes. The analysis of models for many-legged animals shows that wave-like motions, similar to those obtained by N. Kopell and G. B. Ermentrout [see Math. Biosci. 90, 87–109 (1988; Zbl 0649.92009); SIAM J. Appl. Math. 54, 478–507 (1994; Zbl 0811.92004)], can be expected. However, our network leads to a prediction that the wavelength of the wave motion will divide twice the length of the animal. Indeed, we reproduce illustrations of wave-like motions in centipedes where the animal is approximately one-and-a-half wavelengths long-motions that are consistent with this prediction. We discuss the implications of these results for the development of modular control networks for adaptive legged robots.

MSC:

92C15 Developmental biology, pattern formation
92D50 Animal behavior
34C60 Qualitative investigation and simulation of ordinary differential equation models
93C95 Application models in control theory
94C99 Circuits, networks
Full Text: DOI

References:

[1] Alexander, R. McN., Terrestrial locomotion, (Alexander, R. McN.; Goldspink, J. M., Mechanics and Energetics of Animal Locomotion (1977), Chapman and Hall: Chapman and Hall London), 168-203
[2] Cohen, A. H.; Holmes, P. J.; Rand, R. H., The nature of the coupling between segmental oscillators of the lamprey spinal generator for locomotion: a mathematical model, J. Math. Biol., 13, 345-369 (1982) · Zbl 0476.92003
[3] Collins, J. J.; Chow, C. C.; Capela, A. C.; Imhoff, T. T., Aperiodic stochastic resonance, Phys. Rev. E, 54, 5575-5584 (1996)
[4] Collins, J. J.; Richmond, S. A., Hard-wired central pattern generators for quadrupedal locomotion, Biol. Cybern., 71, 375-385 (1994) · Zbl 0800.92086
[5] Collins, J. J.; Stewart, I., Symmetry-breaking bifurcation: a possible mechanism for 2:1 frequency-locking in animal locomotion, J. Math. Biol., 30, 827-838 (1992) · Zbl 0757.92011
[6] Collins, J. J.; Stewart, I., Coupled nonlinear oscillators and the symmetries of animal gaits, J. Nonlin. Sci., 3, 349-392 (1993) · Zbl 0808.92012
[7] Collins, J. J.; Stewart, I., Hexapodal gaits and coupled nonlinear oscillator models, Biol. Cybern., 68, 287-298 (1993) · Zbl 0767.92015
[8] Collins, J. J.; Stewart, I., A group-theoretic approach to rings of coupled biological oscillators, Biol. Cybern., 71, 95-103 (1994) · Zbl 0804.92009
[9] Dionne, B.; Golubitsky, M.; Stewart, I., Coupled cells with internal symmetry. Part I: wreath products, Nonlinearity, 9, 559-574 (1996) · Zbl 0894.58049
[10] Dionne, B.; Golubitsky, M.; Stewart, I., Coupled cells with internal symmetry, Part II: direct products, Nonlinearity, 9, 575-599 (1996) · Zbl 0894.58050
[11] Epstein, I. R.; Golubitsky, M., Symmetric patterns in linear arrays of coupled cells, Chaos, 3, 1, 1-5 (1993) · Zbl 1055.80505
[12] Ermentrout, G. B.; Kopell, N., Multiple pulse interactions and averaging in systems of coupled neural oscillators, J. Math. Biol., 29, 195-217 (1991) · Zbl 0718.92004
[13] Gambaryan, P. P., How Mammals Run: Anatomical Adaptations (1974), Wiley: Wiley New York
[14] Golubitsky, M.; Stewart, I. N., Hopf bifurcation in the presence of symmetry, Arch. Rational Mech. Anal., 87, 2, 107-165 (1985) · Zbl 0588.34030
[15] Golubitsky, M.; Stewart, I. N., Hopf bifurcation with dihedral group symmetry: coupled nonlinear oscillators, (Golubitsky, M.; Guckenheimer, J., Multiparameter Bifurcation Theory. Multiparameter Bifurcation Theory, Providence, RI, 1986. Multiparameter Bifurcation Theory. Multiparameter Bifurcation Theory, Providence, RI, 1986, Contemporary Mathematics, 56 (1986)), 131-173 · Zbl 0602.58038
[16] Golubitsky, M.; Stewart, I. N.; Schaeffer, D. G., (Singularities and Groups in Bifurcation Theory, Vol II (1988), Springer: Springer New York), Applied Mathematical Sciences, vol. 69 · Zbl 0691.58003
[17] Grillner, S., Locomotion in vertebrates: central mechanisms and reflex interaction, Physiol. Rev., 55, 247-304 (1975)
[18] Grillner, S., Control of locomotion in bipeds, tetrapods and fish, (Brooks, V. B., Handbook of Physiology, Section 1: The Nervous System, Vol. II: Motor Control (1981), American Physiological Society: American Physiological Society Bethesda), 1179-1236
[19] Grillner, S., Neurobiological bases of rhythmic motor acts in vertebrates, Science, 228, 143-149 (1985)
[20] Grillner, S., Neural networks for vertebrate locomotion, Sci. Am., 274, 1, 48-53 (1996)
[21] Kopell, N.; Ermentrout, G. B., Symmetry and phaselocking in chains of weakly coupled oscillators, Comm. Pure Appl. Math., 39, 623-660 (1986) · Zbl 0596.92011
[22] Kopell, N.; Ermentrout, G. B., Coupled oscillators and the design of central pattern generators, Math. Biosci., 89, 14-23 (1988) · Zbl 0649.92009
[23] Kopell, N.; Ermentrout, G. B., Phase transitions and other phenomena in chains of oscillators, SIAM J. Appl. Math., 50, 1014-1052 (1990) · Zbl 0711.34029
[24] Pearson, K. G., Common principles of motor control in vertebrates and invertebrates, Annu. Rev. Neurosci., 16, 265-297 (1993)
[25] Rand, R. H.; Cohen, A. H.; Holmes, P. J., Systems of coupled oscillators as models of central pattern generators, (Cohen, A. H.; Rossignol, S.; Grillner, S., Neural Control of Rhythmic Movements in Vertebrates (1988), Wiley: Wiley New York), 333-367
[26] Schöner, G.; Jiang, W. Y.; Kelso, J. A.S., A synergetic theory of quadrupedal gaits and gait transitions, J. Theor. Biol., 142, 359-391 (1990)
[27] Shik, M. L.; Orlovsky, G. N., Neurophysiology of locomotor automatism, Physiol. Rev., 56, 465-501 (1976)
[28] Smale, S., A mathematical model of two cells via Turing’s equation, (Cowan, J. D., Some Mathematical Questions in Biology V. Some Mathematical Questions in Biology V, Amer. Math. Soc. Lecture Notes on Mathematics in the Life Sciences, vol. 6 (1974), AMS: AMS Providence, RI), 15-26 · Zbl 0333.92002
[29] Stein, P. S.G., Motor systems, with specific reference to the control of locomotion, Annu. Rev. Neurosci., 1, 61-81 (1978)
[30] Williams, T. L.; Sigvardt, K. A.; Kopell, N.; Ermentrout, G. B.; Remler, M. P., Forcing of coupled nonlinear oscillators: studies of intersegmental coordination in the lamprey locomotor central pattern generator, J. Neurophysiol., 64, 862-871 (1990)
[31] Yuasa, H.; Ito, M., Coordination of many oscillators and generation of locomotory patterns, Biol. Cybern., 63, 177-184 (1990) · Zbl 0706.92012
[32] Yuasa, H.; Ito, M., Generation of locomotive patterns and self-organization, J. Robot. Mechatron., 4, 142-147 (1992)
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