Control of reflex reversal in stick insect walking: effects of intersegmental signals, changes in direction, and optomotor-induced turning.

In many animals, the effects of sensory feedback on motor output change during locomotion. These changes can occur as reflex reversals in which sense organs that activate muscles to counter perturbations in posture control instead reinforce movements in walking. The mechanisms underlying these changes are only partially understood. As such, it is unclear whether reflex reversals are modulated when locomotion is adapted, such as during changes in walking direction or in turning movements. We investigated these questions in the stick insect Carausius morosus, where sensory signals from the femoral chordotonal organ are known to produce resistance reflexes at rest but assistive movements during walking. We studied how intersegmental signals from neighboring legs affect the generation of reflex reversals in a semi-intact preparation that allows free leg movement during walking. We found that reflex reversal was enhanced by stepping activity of the ipsilateral neighboring rostral leg, whereas stepping of contralateral legs had no effect. Furthermore, we found that the occurrence of reflex reversals was task-specific: in the front legs of animals with five legs walking, reflex reversal was generated only during forward and not backward walking. Similarly, during optomotor-induced curved walking, reflex reversal occurred only in the middle leg on the inside of the turn and not in the contralateral leg on the outside of the turn. Thus our results show for the first time that the nervous system modulates reflexes in individual legs in the adaptation of walking to specific tasks.