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J Neurophysiol 98: 2983-2991, 2007. First published August 29, 2007; doi:10.1152/jn.00407.2007
0022-3077/07 $8.00
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A Central Pattern Generator Producing Alternative Outputs: Phase Relations of Leech Heart Motor Neurons With Respect to Premotor Synaptic Input

Brian J. Norris1,2,*, Adam L. Weaver1,*, Angela Wenning1, Paul S. García1 and Ronald L. Calabrese1

1Department of Biology, Emory University, Atlanta, Georgia; and 2Department of Biological Sciences, California State University, San Marcos, California

Submitted 10 April 2007; accepted in final form 27 August 2007

The central pattern generator (CPG) for heartbeat in leeches consists of seven identified pairs of segmental heart interneurons and one unidentified pair. Four of the identified pairs and the unidentified pair of interneurons make inhibitory synaptic connections with segmental heart motor neurons. The CPG produces a side-to-side asymmetric pattern of intersegmental coordination among ipsilateral premotor interneurons corresponding to a similarly asymmetric fictive motor pattern in heart motor neurons, and asymmetric constriction pattern of the two tubular hearts: synchronous and peristaltic. Using extracellular techniques, we recorded, in 61 isolated nerve cords, the activity of motor neurons in conjunction with the phase reference premotor heart interneuron, HN(4), and another premotor interneuron that allowed us to assess the coordination mode. These data were then coupled with a previous description of the temporal pattern of premotor interneuron activity in the two coordination modes to synthesize a global phase diagram for the known elements of the CPG and the entire motor neuron ensemble. These average data reveal the stereotypical side-to-side asymmetric patterns of intersegmental coordination among the motor neurons and show how this pattern meshes with the activity pattern of premotor interneurons. Analysis of animal-to-animal variability in this coordination indicates that the intersegmental phase progression of motor neuron activity in the midbody in the peristaltic coordination mode is the most stereotypical feature of the fictive motor pattern. Bilateral recordings from motor neurons corroborate the main features of the asymmetric motor pattern.


Address for reprint requests and other correspondence: R. L. Calabrese, Dept. of Biology, Emory University, 1510 Clifton Road N.E., Atlanta, GA 30322 (E-mail: ronald.calabrese{at}emory.edu)







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