Neurones in the leech that facilitate an avoidance behaviour following nearfield water disturbances.

1. A multimodel, multisegmental interneurone (Röhde's fibre, RF) and previously identified mechanoreceptors (T-cells) are shown to respond to nearfield disturbances. Both the T-cells and RF can fire for hundreds of milliseconds following a brief stimulus, and both have subthreshold excitatory synapses onto motor neurones that cause longitudinal contraction of the body wall, an avoidance response. 2. Natural stimulation or electrical stimulation of T-cells in one hemiganglion causes synaptic excitation of T-cells in adjacent ipsilateral hemiganglia and re-excitation of T-cells in the hemiganglion stimulated. A model of repetitive T-cell activity that incorporates previously described synapses among T-cells is presented: the T-cells in adjacent ipsilateral hemiganglia form a reverberatory circuit, re-exciting one another via electrical synapses; repetitive firing is terminated by synaptic inhibition onto T-cells provided by an interneurone excited by the T-cells. With repeated stimulation (0.1--0.2 Hz, 0.2 ms pulses) of a segmental root (directly exciting all the T-cells of a hemiganglion), the number of T-cell impulses per stimulus decreases. Facilitation of inhibition may contribute to the response decrement. 3. The T-cell-RF pathway is investigated. T-cell stimulation can elicit RF impulses in the same and in adjacent ganglia. The long delay between mechanoreceptor stimulation and a response in the interneurone suggests that spatial and temporal summation of T-cell inputs may be required to reach firing threshold in the interneurone. 4. The impulse frequency of the RF response was compared for a travelling surface wave that is approaching a segment v. one that is moving away from the segment. It was found that the frequency was greater as the stimulus approaches; this should allow more effective temporal summation of the subthreshold synaptic potentials which RF evokes in motor neurones that cause longitudinal contraction of the body wall. Therefore, the probability of contraction is greater in segments toward which a stimulus is moving.