Identified higher-order neurons controlling the feeding motor program of helisoma

A bilaterally symmetrical pair of neurons in the anterior region of the cerebral ganglia of the snail Helisoma trivolvis were found to have excitatory input to the feeding motor program contained in the buccal ganglia. Intracellular microelectrodes were used to stimulate and record from these cells while the motor output of the buccal ganglia was monitored with a combination of intracellular and extracellular recordings. Experimentally evoked tonic activity in an individual cerebral cell could initiate and maintain the patterned motor output from buccal ganglia, characteristic of the activity underlying buccal mass feeding movements. The rate of buccal motor output could be modified directly by varying the firing frequency of the cerebral cell. Cobaltous chloride backfills of cerebrobuccal connectives revealed that these higher-order neurons were the only large cells in the anterior portion of the ganglia to send processes into the connectives. Furthermore, they are the only cells in this region to fluoresce when processed with a sucrose-phosphate-glyoxylic acid solution, indicating that they contain an indoleamine, probably serotonin. Application of low concentrations of serotonin to isolated buccal ganglia or buccal ganglia-buccal mass preparations mimics the effects of the cerebral cells' activity on the buccal motor output, implying that serotonin is a putative transmitter for these cerebral cells.