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Volume 17, Number 20,
Issue of October 15, 1997
pp. 7703-7713
Copyright ©1997 Society for Neuroscience
Removal of Spike Frequency Adaptation via Neuromodulation
Intrinsic to the Tritonia Escape Swim Central Pattern
Generator
Received May 27, 1997; revised Aug. 5, 1997; accepted Aug. 6, 1997.
Paul S. Katz1 and
William N. Frost2
1 Department of Biology, Georgia State University,
Atlanta, Georgia 30303, and 2 Department of Neurobiology
and Anatomy, University of Texas Medical School, Houston, Texas 77030
For the mollusc Tritonia diomedea to generate its
escape swim motor pattern, interneuron C2, a crucial member of the
central pattern generator (CPG) for this rhythmic behavior, must fire repetitive bursts of action potentials. Yet, before swimming, repeated
depolarizing current pulses injected into C2 at periods similar those
in the swim motor program are incapable of mimicking the firing rate
attained by C2 on each cycle of a swim motor program. This resting
level of C2 inexcitability is attributable to its own inherent spike
frequency adaptation (SFA). Clearly, this property must be altered for
the swim behavior to occur. The pathway for initiation of the swimming
behavior involves activation of the serotonergic dorsal swim
interneurons (DSIs), which are also intrinsic members of the swim CPG.
Physiologically appropriate DSI stimulation transiently decreases C2
SFA, allowing C2 to fire at higher rates even when repeatedly
depolarized at short intervals. The increased C2 excitability caused by
DSI stimulation is mimicked and occluded by serotonin application.
Furthermore, the change in excitability is not caused by the
depolarization associated with DSI stimulation or serotonin application
but is correlated with a decrease in C2 spike afterhyperpolarization.
This suggests that the DSIs use serotonin to evoke a neuromodulatory
action on a conductance in C2 that regulates its firing rate. This
modulatory action of one CPG neuron on another is likely to play a role
in configuring the swim circuit into its rhythmic pattern-generating
mode and maintaining it in that state.
Key words:
intrinsic neuromodulation;
serotonin;
nudibranch mollusc;
repetitive firing;
central pattern generator;
after-
hyperpolarization
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