The generation of rhythmic activity in a distributed motor system.

Rhythmic activity that is distributed to the brain and buccal ganglia and which underlies several types of behaviour, can be evoked from isolated nervous systems of Pleurobranchaea californica by tonic nerve stimulation. The experiments presented here were designed to test whether this rhythmic activity is produced by independent neuronal oscillators located in each ganglion or whether the rhythmic activity arises from a single oscillatory locus in the buccal ganglion and is transmitted passively to the brain. By interrupting the conduction of activity in the cerebrobuccal connectives (CBC) between brain and buccal ganglia we show that motor output from the brain depends on sustained, cycle to cycle input from the buccal ganglion and cannot be reset with respect to the buccal activity. The production of rhythmic activity in the brain depends on the generation of rhythmic activity in the buccal ganglia whether the rhythms are activated by stimulation of buccal roots or paracerebral command cells in the brain. Simultaneous intracellular recordings from brain motoneurones and buccal interneurones which project to the brain indicate that these interneurones provide both the drive and the pattern for rhythmic motor output in the brain. Tonic stimulation of the CBC can produce rhythmic activity in isolated brains in which all nerve roots and connectives have been cut. This can be explained by the fact that tonic stimulation of the connectives is transformed into phasic activity by the axons within the connective. We conclude therefore, that rhythmic, coordinated activity in the brain and buccal ganglia of Pleurobranchaea arises from oscillatory circuits that are located only in the buccal ganglia.