This study examined the relationship between ontogenetic increase in body size and the kinematics of peristaltic locomotion by the earthworm Lumbricus terrestris, a soft-bodied organism supported by a hydrostatic skeleton. Whereas the motions of most vertebrates and arthropods are based primarily on the changes in the joint angles between rigid body segments, the motions of soft-bodied organisms with hydrostatic skeletons are based primarily on the changes in dimensions of the deformable body segments themselves. The overall kinematics of peristaltic crawling and the dynamic shape changes of individual earthworm segments were measured for individuals ranging in body mass (mb) by almost three orders of magnitude (0.012-8.5 g). Preferred crawling speed varied both within and among individuals: earthworms crawled faster primarily by taking longer strides, but also by taking more strides per unit time and by decreasing duty factor. On average, larger worms crawled at a greater absolute speed than smaller worms (U p2finity mb0.33) and did so by taking slightly longer strides (l p2finity mb0.41, where l is stride length) than expected by geometric similarity, using slightly lower stride frequencies (f p2finity mb-0.07) and the same duty factor (df p2finity mb-0.03). Circumferential and longitudinal body wall strains were generally independent of body mass, while strain rates changed little as a function of body mass. Given the extent of kinematic variation within and among earthworms, the crawling of earthworms of different sizes can be considered to show kinematic similarity when the kinematic variables are normalized by body length. Since the motions of peristaltic organisms are based primarily on changes in the dimensions of the deformable body wall, the scaling of the material properties of the body wall is probably an especially important determinant of the scaling of the kinematics of locomotion.
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.
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