The synaptic activities underlying the uropod steering behaviour of crayfish evoked by tilting the substratum beneath the legs have been studied intracellularly in unanaesthetized animals standing or walking on a treadmill. The uropod motoneurones showed little or no synaptic response when the treadmill was tilted while the animal was in a quiescent state and the membrane potential was at its resting value. When the same stimulus was given while the animal was walking or in an active stance on the treadmill, the motoneurones showed transient much-enhanced excitatory or inhibitory responses to tilt, depending on the tilt direction. These responses were superimposed on a sustained level of background excitation so that the spike activity of the motoneurones either increased or decreased. Premotor nonspiking interneurones also showed little or no synaptic response to the tilt stimulus while the animal was resting, but greatly enhanced responses, in either a depolarizing or a hyperpolarizing direction, while the animal was walking or in the active-standing state. The results indicate that the proprioceptor inputs converging onto the uropod motoneurones, either directly or through premotor nonspiking interneurones, are gated not only in the uropod motor system in the terminal abdominal ganglion but also at as yet unidentified sites upstream in anterior ganglia, thus suggesting multiple gate control of the descending proprioceptor pathway.

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