Patterned activation of unpaired median neurons during fictive crawling in manduca sexta larvae

The unpaired median neurons are common to the segmental ganglia of many insects. Although some of the functional consequences of their activation, among them the release of octopamine to modulate muscle contraction, have been described, less is understood about how and when these neurons are recruited during movement. The present study demonstrates that peripherally projecting unpaired median neurons in the abdominal and thoracic ganglia of the larval tobacco hornworm Manduca sexta are recruited rhythmically during the fictive crawling motor activity that is produced by the isolated central nervous system in response to pilocarpine. Regardless of the muscles to which they project, the efferent unpaired median neurons in all segmental ganglia are depolarized together during the phase of the crawling cycle when the thoracic leg levator motoneurons are active. During fictive crawling, therefore, the unpaired median neurons are not necessarily active in synchrony with the muscles to which they project. The rhythmical synaptic drive of the efferent unpaired median neurons is derived, at least in part, from a source within the subesophageal ganglion, even when the motor pattern is evoked by exposing only the more posterior ganglia to pilocarpine. In pairwise intracellular recordings from unpaired median neurons in different ganglia, prominent excitatory postsynaptic potentials, which occur with an anterior-to-posterior delay in both neurons, are seen to underlie the rhythmic depolarizations. One model consistent with these findings is that one or more neurons within the subesophageal ganglion, which project posteriorly to the segmental ganglia and ordinarily provide unpatterned synaptic inputs to all efferent unpaired median neurons, become rhythmically active during fictive crawling in response to ascending information from the segmental pattern-generating network.