Premotor interneurons in the local control of stepping motor output for the stick insect single middle leg.

In insect walking systems, nonspiking interneurons (NSIs) play an important role in the control of posture and movement. As such NSIs are known to contribute to state-dependent modifications in processing of proprioceptive signals from the legs. For example, NSIs process a flexion of the femur-tibia (FTi) joint signaled by the femoral chordotonal organ (fCO) such that the stance phase motor output is reinforced in the active locomotor system. This phenomenon representing a reflex reversal is the first part of the "active reaction" (AR) and was hypothesized to functionally represent a major control feature by which sensory feedback supports stance generation. As NSIs are known to contribute to the AR, the question arises, whether they serve similar functions during stepping and whether the AR is generally part of the control system for walking. We studied these issues in vivo, in a single leg preparation of Carausius morosus with the leg kinematics being confined to changes in one plane, along the coxa-trochanteral and the FTi-joint. Following kinematic analysis, identified NSIs (E1-E8, I1, I2, and I4) were recorded intracellularly during single leg stepping at different velocities. We detected clear similarities between the activity pattern of NSIs during single leg stepping and their responses to fCO-stimulation during the generation of the AR. This strongly supports the notion that the motor output generated during the AR reflects part of the control regime for stepping. Furthermore, our experiments revealed that alterations in stepping velocity result from modifications in the activity of the premotor NSIs involved in stance phase generation.