In terrestrial locomotion, sensory feedback from load sensors is important for altering ongoing motor output on a step-by-step basis. We investigated the influence of load signals from the leg on motoneuron pools of the thorax-coxa (ThC) joint in the stick insect walking system. Load sensors were stimulated during rhythmic, alternating activity in protractor coxae (ProCx) and retractor coxae (RetCx) motoneuron pools. Alternating activity in the segment of interest was induced by mechanical stimulation of the animal or pharmacological activation of the isolated thoracic ganglia. Load signals from the legs altered the timing of ThC motoneuron activity by resetting and entraining the activity of the central rhythm generating network of the ThC joint. In the front and middle legs, load signals induced or promoted RetCx activity and decreased or terminated ProCx activity. In the hindleg, reverse transitions were elicited, with increasing load terminating RetCx and initiating ProCx activity. Studies in semi-intact walking animals showed that the effect of load on the ThC-joint motoneurons depended on walking direction, with increased load promoting the functional stance phase motoneuron pool (in forward walking, RetCx activity; in backward walking, ProCx activity). Thus, we show that modifications of sensory feedback in a locomotor system are related to walking direction. In a final set of ablation experiments, we show that the load influence is mediated by the three groups of trochanteral campaniform sensilla.

Locomotion requires the coordination of movements across body segments, which in walking animals is expressed as gaits. We studied the underlying neural mechanisms of this coordination in a semi-intact walking preparation of the stick insect Carausius morosus. During walking of a single front leg on a treadmill, leg motoneuron (MN) activity tonically increased and became rhythmically modulated in the ipsilateral deafferented and deefferented mesothoracic (middle leg) ganglion. The pattern of modulation was correlated with the front leg cycle and specific for a given MN pool, although it was not consistent with functional leg movements for all MN pools. In an isolated preparation of a pair of ganglia, where one ganglion was made rhythmically active by application of pilocarpine, we found no evidence for coupling between segmental central pattern generators (CPGs) that could account for the modulation of MN activity observed in the semi-intact walking preparation. However, a third preparation provided evidence that signals from the front leg's femoral chordotonal organ (fCO) influenced activity of ipsilateral MNs in the adjacent mesothoracic ganglion. These intersegmental signals could be partially responsible for the observed MN activity modulation during front leg walking. While afferent signals from a single walking front leg modulate the activity of MNs in the adjacent segment, additional afferent signals, local or from contralateral or posterior legs, might be necessary to produce the functional motor pattern observed in freely walking animals.