Rhombencephalic pathways and neurotransmitters controlling deglutition

Information from neuronal pathway tracing and pharmacologic microstimulation studies, in conjunction with electrophysiological data, has begun to coalesce into a coherent, if still incomplete, picture of the brain stem circuitry responsible for generating the motor patterns underlying deglutition and esophageal peristalsis in the rat. The intermediate, interstitial, and ventral subnuclei of the solitarius complex appear to play a pivotal part, as evidenced by their viscerosensory inputs and extensive projections to the parvicellular intermediate reticular formation of the medulla, to the brain stem deglutitive motor neuron pools, and to general visceral efferent preganglionic neurons controlling the upper alimentary tract striated and smooth musculature, respectively. The dense projections of the solitarial central subnucleus form a separate subcircuit controlling esophageal, and also some aspects of gastric, motility. Although not extensive, direct connections between the latter subnucleus and interneurons coordinating the buccopharyngeal stage of swallowing appear to exist. In both subcircuits, fast information transfer uses excitatory amino acidergic transmission by means of several glutamate-receptor subtypes. Release from tonic GABAergic inhibition exerted by local solitarial interneurons may provide a mechanism for triggering deglutitive premotoneuronal activity. Local or reticular cholinergic neurons are implicated in pharyngoesophageal coupling and the generation of propulsive esophagomotor output. The solitary interneurons under investigation engage in complex local dendritic and axonal projections within the solitarius complex. Further analysis of these local circuits and their transmitters should yield essential clues regarding the mechanisms underlying deglutitive motor pattern generation.