SUMMARY: The biomechanics of running in small animals have remained poorly characterized because of the difficulty of recording three-dimensional ground reaction forces. Available techniques limit investigations to animals with a body mass above 1 g. Here we present, for the first time, single-leg ground reaction forces of ants (body mass 10 mg), measured with a custom-built miniature force plate. We investigated forces and high-speed kinematics for straight level runs (average speed: 8.4 cm s–1) of Formica polyctena workers. The major finding was that the time course of ground reaction forces strongly differed from previous observations of larger insects. Maximum vertical force was reached during the first third of the tripod contact phase. During this period the body was decelerated predominantly by the front legs. Subsequently, the front legs pulled and accelerated the body. This `climbing' type of stride may be useful on the bumpy and unstable substrates that the animals face in their natural habitats,and may therefore also occur on level ground. Propulsive forces were generated predominantly by the front and hind legs. Dragging of the gaster on the substrate resulted in a breaking momentum, which was compensated by the legs. Future investigations will reveal, whether the identified pattern is due to specialization.
Many legged animals change gaits when increasing speed. In insects, only one gait change has been documented so far, from slow walking to fast running, which is characterised by an alternating tripod. Studies on some fast-running insects suggested a further gait change at higher running speeds. Apart from speed, insect gaits and leg co-ordination have been shown to be influenced by substrate properties, but the detailed effects of speed and substrate on gait changes are still unclear. Here we investigate high-speed locomotion and gait changes of the cockroach Nauphoeta cinerea, on two substrates of different slipperiness. Analyses of leg co-ordination and body oscillations for straight and steady escape runs revealed that at high speeds, blaberid cockroaches changed from an alternating tripod to a rather metachronal gait, which to our knowledge, has not been described before for terrestrial arthropods. Despite low duty factors, this new gait is characterised by low vertical amplitudes of the centre of mass (COM), low vertical accelerations and presumably reduced total vertical peak forces. However, lateral amplitudes and accelerations were higher in the faster gait with reduced leg synchronisation than in the tripod gait with distinct leg synchronisation. Temporally distributed leg force application as resulting from metachronal leg coordination at high running speeds may be particularly useful in animals with limited capabilities for elastic energy storage within the legs, as energy efficiency can be increased without the need for elasticity in the legs. It may also facilitate locomotion on slippery surfaces, which usually reduce leg force transmission to the ground. Moreover, increased temporal overlap of the stance phases of the legs likely improves locomotion control, which might result in a higher dynamic stability.
16 September 2009 in Journal of Comparative Physiology A
Knowing the functionality and capabilities of masticatory apparatuses is essential for the ecological classification of jawed organisms. Nevertheless insects, especially with their outstanding high species number providing an overwhelming morphological diversity, are notoriously underexplored with respect to maximum bite forces and their dependency on the mandible opening angles. Aiming for a general understanding of insect biting, we examined the generalist feeding cockroach Periplaneta americana, characterized by its primitive chewing mouth parts. We measured active isometric bite forces and passive forces caused by joint resistance over the entire mandibular range with a custom-built 2D force transducer. The opening angle of the mandibles was quantified by using a video system. With respect to the effective mechanical advantage of the mandibles and the cross-section areas, we calculated the forces exerted by the mandible closer muscles and the corresponding muscle stress values. Comparisons with the scarce data available revealed close similarities of the cockroaches’ mandible closer stress values (58 N/cm2) to that of smaller specialist carnivorous ground beetles, but strikingly higher values than in larger stag beetles. In contrast to available datasets our results imply the activity of faster and slower muscle fibres, with the latter becoming active only when the animals chew on tough material which requires repetitive, hard biting. Under such circumstances the coactivity of fast and slow fibres provides a force boost which is not available during short-term activities, since long latencies prevent a specific effective employment of the slow fibres in this case.
Spiders are an old yet very successful predatory group of arthropods. Their locomotor system differs from those of most other arthropods by the lack of extensor muscles in two major leg joints. Though specific functional characteristics can be expected regarding the locomotion dynamics of spiders, this aspect of movement physiology has been only scarcely examined so far. This study presents extensive analyses of a large dataset concerning global kinematics and the implications for dynamics of adult female specimens of the large Central American spider Cupiennius salei (Keyserling). The experiments covered the entire speed-range of straight runs at constant speeds. The analyses revealed specific characteristics of velocity dependent changes in the movements of the individual legs, as well as in the translational and rotational degrees of freedom of both the centre of mass and the body. In contrast to many other fast moving arthropods, C. salei avoid vertical fluctuations of their centre of mass during fast locomotion. Accordingly, aerial phases were not observed here. This behaviour is most likely a consequence of optimising energy expenditure with regard to the specific requirements of spiders' leg anatomy. A strong synchronisation of two alternating sets of legs appears to play only a minor role in the locomotion of large spiders. Reduced frequency and low centre of mass amplitudes as well as low angular changes of the body axes, in turn, seems to be the result of relatively low leg coordination.
16 September 2009 in Journal of Comparative Physiology B