Gain changes in sensorimotor pathways of the locust leg

Abstract

Feedback systems that control the leg joints of animals must be highly flexible in adapting to different behavioural tasks. One manifestation of such flexibility is changes in the gain of joint control networks. The femur- tibia (FT) control network of the locust leg is one of the feedback systems most thoroughly studied with regard to its neural circuitry. Despite excellent information concerning network topology, however, actual gain changes and their underlying mechanisms have not yet been examined because of the marked spontaneous variations in the action of the control network for this joint. We describe a behavioural situation and a preparation in which the locust (Locusta migratoria L.) FT control network exhibits reproducible changes in gain, allowing investigation of the neuronal basis of gain control. After ('fictive') flight motor activity, the gain of resistance reflexes in the FT joint of the locust middle leg is significantly decreased, with the flexor tibiae muscles being affected more strongly than the extensor muscles. Immediately after flight motor activity, the gain may be as low as 30 % of pre-flight levels. It returns to pre-flight values in under 150s. The decrease in gain following flight motor activity is due to a decrease in motoneurone recruitment in the resistance reflex elicited by stimulation of the appropriate mechanoreceptor, the femoral chordotonal organ. Motoneurone recruitment is changed as a result of a drastic decline in the stimulus- related synaptic input to the motoneurones, which appears to be produced exclusively at the level of the pre-motor network. Two factors led to this conclusion: first, we found no indication of changes in membrane potential or membrane conductance of the tibia flexor and extensor motoneurones; second, recording from identified pre-motor nonspiking interneurones demonstrated that these may be involved in the observed gain changes. The putative behavioural relevance is discussed.