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The Journal of Neurophysiology Vol. 87 No. 6 June 2002, pp. 2760-2769
Copyright ©2002 by the American Physiological Society
Department of Biology, National Science Foundation Center for Biological Timing, University of Virginia, Charlottesville, Virginia 22904-4328
Cang, Jianhua and
W. Otto Friesen.
Model for Intersegmental Coordination of Leech Swimming: Central
and Sensory Mechanisms. J. Neurophysiol. 87: 2760-2769, 2002. Sensory feedback as well as the coupling
signals within the CNS are essential for leeches to produce
intersegmental phase relationships in body movements appropriate for
swimming behavior. To study the interactions between the central
pattern generator (CPG) and peripheral feedback in controlling
intersegmental coordination, we have constructed a computational model
for the leech swimming system with physiologically realistic
parameters. First, the leech swimming CPG is simulated by a chain of
phase oscillators coupled by three channels of coordinating signals.
The activity phase, the projection direction, and the phase response
curve (PRC) of each channel are based on the identified intersegmental
interneuron network. Output of this largely constrained model produces
stable coordination in the simulated CPG with average phase lags of
8-10°/segment in the period range from 0.5 to 1.5 s, similar to
those observed in isolated nerve cords. The model also replicates the
experimental finding that shorter chains of leech nerve cords have
larger phase lags per segment. Sensory inputs, represented by stretch
receptors, were subsequently incorporated into the CPG model. Each
stretch receptor with its associated PRC, which was defined to mimic
the experimental results of phase-dependent phase shifts of the central oscillator by the ventral stretch receptor, can alter the phase of the
local central oscillator. Finally, mechanical interactions between the muscles from neighboring segments were simulated by PRCs
linking adjacent stretch receptors. This model shows that interactions
between neighboring muscles could globally increase the phase lags to
the larger value required for the one-wavelength body form observed in
freely swimming leeches. The full model also replicates the
experimental observation that leeches with severed nerve cords have
larger intersegmental phase lags than intact animals. The similarities
between physiological and simulation results demonstrate that we have
established a realistic model for the central and peripheral control of
intersegmental coordination of leech swimming.
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