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Journal of Neuroscience, Vol 10, 1495-1512, Copyright © 1990 by Society for Neuroscience
Neuromodulation of the crab pyloric central pattern generator by serotonergic/cholinergic proprioceptive afferents
PS Katz and RM Harris-Warrick
Section of Neurobiology and Behavior, Cornell University, Ithaca, New York 14853.
In the stomatogastric nervous system of the crab, Cancer borealis, a set of
4 serotonergic/cholinergic proprioceptive neurons, called gastropyloric
receptor (GPR) cells, have effects on the pyloric motor pattern. In a
semi-intact foregut preparation, the GPR cells are not activated by
movements of the pyloric filter; instead they respond to the slower
movements of the gastric mill (Katz et al., 1989). Thus, their activity is
not synchronized to the pyloric motor pattern. However, when the GPR cells
are stimulated in an in vitro preparation in a manner that resembles their
normal firing pattern, they produce dramatic effects on the pyloric motor
pattern. These effects include: (1) a prolonged increase in the pyloric
cycle frequency, (2) a momentary pause in the motor pattern, (3) transient
inhibition of some motor neurons, (4) strong excitation of other motor
neurons, and (5) altered phase relationships of the different components of
the motor pattern. These changes in the motor pattern are due to direct
effects of the GPR cells on neurons in the pyloric central pattern
generator (CPG). All of the cells in the pyloric circuit appear to receive
GPR input. However, only 2 neurons receive detectable rapid nicotinic
synaptic potentials. The other neurons receive only slower neuromodulatory
input from GPR stimulation. The neuromodulatory effects include burst
enhancement, plateau potential enhancement, excitation, and inhibition.
These modulatory effects are largely mimicked by bath- applied serotonin
(5-HT). Thus, primary sensory neurons can alter the production of motor
patterns by a CPG through a phase-independent mechanism; these
proprioceptors do not need to fire at a precise time in the cycle to be
effective because their effects are mediated through the slower actions of
the neuromodulator 5-HT.
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