Scaling of the hydrostatic skeleton in the earthworm Lumbricus terrestris

The structural and functional consequences of changes in size or scale have been well studied in animals with rigid skeletons, but relatively little is known about scale effects in animals with hydrostatic skeletons. We used glycol methacrylate histology and microscopy to examine the scaling of mechanically important morphological features of the earthworm Lumbricus terrestris over an ontogenetic size range from 0.03 to 12.89 g. We found that L. terrestris becomes disproportionately longer and thinner as it grows. This increase in the length to diameter ratio with size means that, when normalized for mass, adult worms gain ~117% mechanical advantage during radial expansion, compared with hatchling worms. We also found that the cross-sectional area of the longitudinal musculature scales as body mass to the ~0.6 power across segments, which is significantly lower than the 0.66 power predicted by isometry. The cross-sectional area of the circular musculature, however, scales as body mass to the ~0.8 power across segments, which is significantly higher than predicted by isometry. By modeling the interaction of muscle cross-sectional area and mechanical advantage, we calculate that the force output generated during both circular and longitudinal muscle contraction scales near isometry. We hypothesize that the allometric scaling of earthworms may reflect changes in soil properties and burrowing mechanics with size.