Ontogenetic scaling of burrowing forces in the earthworm Lumbricus terrestris

In hydrostatic skeletons, it is the internal fluid under pressure surrounded by a body wall in tension (rather than a rigid lever) that enables the stiffening of the organism, the antagonism of muscles and the transmission of force from the muscles to the environment. This study examined the ontogenetic effects of body size on force production by an organism supported with a hydrostatic skeleton. The earthworm Lumbricus terrestris burrows by forcefully enlarging crevices in the soil. I built a force-measuring apparatus that measured the radial forces as earthworms of different sizes crawled through and enlarged pre-formed soil burrows. I also built an apparatus that measured the radial and axial forces as earthworms of different sizes attempted to elongate a dead-end burrow. Earthworms ranging in body mass m(b) from hatchlings (0.012 g) to adults (8.9 g) exerted maximum forces (F, in N) during active radial expansion of their burrows (F=0.32 m(b)(0.43)) and comparable forces during axial elongation of the burrow (F=0.26 m(b)(0.47)). Both these forces were almost an order of magnitude greater than the radial anchoring forces during normal peristalsis within burrows (F=0.04 m(b)(0.45)). All radial and axial forces scaled as body mass raised to the 2/5 power rather than to the 2/3 power expected by geometric similarity, indicating that large worms exert greater forces than small worms on an absolute scale, but the difference was less than predicted by scaling considerations. When forces were normalized by body weight, hatchlings could push 500 times their own body weight, while large adults could push only 10 times their own body weight.