Ballistic movements of jumping legs implemented as variable components of cricket behaviour

Abstract

Ballistic accelerations of a limb or the whole body require special joint mechanisms in many animals. Specialized joints can be moved by stereotypic or variable motor control during motor patterns with and without ballistic components. As a model of variable motor control, the specialized femur-tibia (knee) joints of cricket (Acheta domesticus) hindlegs were studied during ballistic kicking, jumping and swimming and in non-ballistic walking. In this joint the tendons of the antagonistic flexor and the extensor muscles attach at different distances from the pivot and the opposed lever arms form an angle of 120 deg. A 10:1 ratio of their effective lever arms at full knee flexion helps to prepare for most ballistic extensions: the tension of the extensor can reach its peak while it is restrained by flexor co-contraction. In kicks, preparatory flexion is rapid and the co-contraction terminates just before knee extensions. Therefore, mainly the stored tension of the extensor muscle accelerates the small mass of the tibia. Jumps are prepared with slower extensor-flexor co-contractions that flex both knees simultaneously and then halt to rotate both legs outward to a near horizontal level. From there, catapult extension of both knees accelerates the body, supported by continued high frequency motor activity to their tibia extensor muscles during the ongoing push-off from the substrate. Premature extension of one knee instantly takes load from the lagging leg that extends and catches up, which finally results in a straight jump. In swimming, synchronous ballistic power strokes of both hindlegs drive the tibiae on a ventral-to-posterior trajectory through the water, well coordinated with the swimming patterns of all legs. In walking, running and climbing the steps of the hindlegs range between 45 deg flexion and 125 deg extension and use non-ballistic, alternating activity of knee flexor and extensor muscles. Steep climbing requires longer bursts from the extensor tibiae muscles when they support the extended hindlegs against gravity forces when the body hangs over. All ballistic movements of cricket knees are elicited by a basic but variable motor pattern: knee flexions by co-contraction of the antagonists prepare catapult extensions with speeds and forces as required in the different behaviours.