Dynamic simulation of insect walking.

Insect walking relies on a complex interaction between the environment, body segments, muscles and the nervous system. For the stick insect in particular, previous investigations have highlighted the role of specific sensory signals in the timing of activity of central neural networks driving the individual leg joints. The objective of the current study was to relate specific sensory and neuronal mechanisms, known from experiments on reduced preparations, to the generation of the natural sequence of events forming the step cycle in a single leg. We have done this by simulating a dynamic 3D-biomechanical model of the stick insect coupled to a reduced model of the neural control system, incorporating only the mechanisms under study. The neural system sends muscle activation levels to the biomechanical system, which in turn provides correctly timed propriosensory signals back to the neural model. The first simulations were designed to test if the currently known mechanisms would be sufficient to explain the coordinated activation of the different leg muscles in the middle leg. Two experimental situations were mimicked: restricted stepping where only the coxa-trochanteral joint and the femur-tibia joint were free to move, and the unrestricted single leg movements on a friction-free surface. The first of these experimental situations is in fact similar to the preparation used in gathering much of the detailed knowledge on sensory and neuronal mechanisms. The simulations show that the mechanisms included can indeed account for the entire step cycle in both situations. The second aim was to test to what extent the same sensory and neuronal mechanisms would be adequate also for controlling the front and hind legs, despite the large differences in both leg morphology and kinematic patterns. The simulations show that front leg stepping can be generated by basically the same mechanisms while the hind leg control requires some reorganization. The simulations suggest that the influence from the femoral chordotonal organs on the network controlling levation-depression may have a reversed effect in the hind legs as compared to the middle and front legs. This, and other predictions from the model will have to be confirmed by additional experiments.