Maximum single leg force production: cockroaches righting on photoelastic gelatin

Abstract

Integrating studies of mechanics, neural control and isolated muscle function are possible using arthropod legs. To evaluate leg performance, we measured the ground reaction forces generated by individual legs of the six-legged cockroach Blaberus discoidalis (3.1 g), during an emergency behavior, righting or over-turning. We used a photoelastic method to measure the forces generated by individual legs simultaneously. A gelatin track placed between crossed polarizing filters was illuminated from below, and a high-speed video camera recorded the stress-induced optical signals from above. The size and skew of the optical patterns were found to be related to the magnitude and direction of the force. We discovered that the ground reaction forces generated during the righting behavior of the death-head cockroach were eight times greater than those observed during high-speed running, supporting the possibility that relative leg forces (leg force per unit body weight) during running and maximal leg activity differ more in small arthropods than in larger vertebrates. Non-geometric scaling of relative leg force (i.e. scaling to less than body mass-0.33), along with the reduced force-generating ability of a single leg in animals with many legs, may help to explain why the maximum relative leg force production by six-legged cockroaches, as well as by some other small insects, can be similar to the relative single leg forces produced by two- and four-legged vertebrates that are almost 1000 times more massive. Leg number and body mass alone, however, appear to be insufficient to explain the variation observed in relative leg force production at a given body mass, because enormous diversity in musculo-skeletal parameters exists. The maximal relative leg force of the cockroach B. discoidalis during righting was at the low end of a 100-fold variation observed for smaller insects wedging (pushing through a small crevice) and pulling loads. Thus, this cockroach can be characterized as a moderately strong insect with the capacity for relatively high speed. Results from the present study question the predictive strength of the simple geometric scaling arguments involving a strength:weight ratio as they are applied to small arthropods and encourage further consideration of the importance of leg number, muscle force production and mechanical advantage in the derivation of general principles of leg performance.