From Stimulation to Undulation: A Neuronal Pathway for the Control of Swimming in the Leech

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Abstract
Initiation and performance of the swimming movement in the leech (Hirudo medicinalis) are controlled by neurons organized at at least four functional levels—sensory neurons, gating neurons, oscillator neurons, and motor neurons. A paired neuron, designated as Trl, in the subesophageal ganglion of the leech has now been shown to define a fifth level, interposed between sensory and gating neurons. Cell Trl is activated by pressure and nociceptive mechanosensory neurons, which mediate bodywall stimulus-evoked swimming activity in intact leeches. In the isolated leech nervous system, brief stimulation of cell Trl elicits sustained activation of the gating neurons and triggers the onset of swimmning activity. The synaptic interactions between all five levels of control are direct. Discovery of the Trl cells thus completes the identification of a synaptic pathway by which mechanosensory stimulation leads to the swimming movements of the leech.