Adaptive changes in locust kicking and jumping behaviour during development
The hind, or metathoracic, leg of a locust is specialised, enabling it to store energy that is used to extend the tibia rapidly during kicking and jumping; behaviours which share a common motor pattern. This study describes developmental changes in kicking and jumping behaviour and relates these changes to the development of the exoskeleton and jumping performance. In mature adults and intermoult larvae, the exoskeleton is strong and kicks can readily be elicited. Before and after the adult moult, when the exoskeleton is weak, kicks can be elicited less frequently, thus avoiding skeletal damage. At these times, animals do not generate the adult motor pattern for kicking, so that extension of the tibia is powered by direct muscle contraction, rather than through the release of stored energy. The muscles of all newly moulted animals are capable of generating sufficient force to damage the leg, but 14 days later the muscles can rarely generate sufficient force to damage the leg. To mimic the forces generated during the preparation for a kick, when the flexor and extensor tibiae muscles co-contract, the extensor muscle was stimulated electrically at a range of frequencies and the nature and occurrence of the resulting mechanical damage to components of the skeleton were assessed over a 14 day period following the adult moult. In newly moulted animals, the proximal femur partially collapses and thus protects the leg from damage before the muscles generate sufficient force to damage chronically other components of the leg. This partial collapse of the femur is reversible when the extensor muscle is activated at low frequency, but high frequencies cause permanent damage. The muscles of all animals 1 day after the moult are also capable of generating sufficient force to damage the leg, but the proximal tibia breaks most commonly in the region where the extensor muscle apodeme attaches. 5 days after the moult, the muscles in only 50 % of animals can damage the leg and most commonly the extensor muscle apodeme breaks. In mature animals, the only structure that fails is the extensor muscle apodeme, which fractures close to its point of attachment to the tibia. Damaging a metathoracic leg can significantly decrease the ability of a locust to jump and to compete for mates. Changes in the likelihood of damage to a metathoracic leg occur at predictable stages of development. Locust behaviour is modified during development, avoiding such damage.