Intersegmental interneurons can control the gain of reflexes in adjacent segments of the locust by their action on nonspiking local interneurons

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Abstract
The gain of local reflexes of one leg of a locust can be altered by mechanosensory inputs generated by movements of or tactile inputs to an adjacent leg. Touching the mesothoracic tarsus, for example, increases the number of spikes that are produced by the metathoracic slow extensor tibiae motor neuron and enhances the depolarization of flexor tibiae motor neuron in response to imposed movements of the chordotonal organ in the ipsilateral hind femur. The sensory information from the middle leg is conveyed directly to nonspiking interneurons and motor neurons controlling the movements of the hindleg by a population of mesothoracic intersegmental interneurons (Laurent and Burrows, 1989). The metathoracic nonspiking interneurons receive direct inputs from receptors on a hindleg and are, therefore, a point of convergence for local and intersegmental inputs. We examine here the role of the connections between mesothoracic intersegmental interneurons and metathoracic nonspiking interneurons in controlling metathoracic local reflexes. The amplitude of synaptic potentials evoked in leg motor neurons by the stimulation of local afferents can be modulated by altering the membrane potential of an interposed nonspiking interneuron with current injected through an intracellular electrode. These imposed voltage changes mimic a mesothoracic input and show that the state of a nonspiking local interneuron is a determining factor in the expression of a local reflex. Inputs from mesothoracic intersegmental interneurons may cause large changes in the input conductance of nonspiking interneurons that can shunt a local afferent input. In some nonspiking interneurons, synaptic potentials caused by mesothoracic interneurons can be recorded, but no underlying conductance change can be detected at the recording site. Similarly, a particular nonspiking interneuron may receive synaptic inputs when two distinct regions of a middle leg are touched, but only one of these intersegmental inputs may be effective in reducing the amplitude of a synaptic potential caused by afferents from the hindleg. These results suggest that nonspiking local interneurons may be compartmentalized, with synaptic inputs and their associated conductance changes restricted to particular branches. In this way, an individual nonspiking neuron could contribute simultaneously to several local circuits. The inputs from different intersegmental interneurons could then modulate these pathways independently.