Biomechanical limits to soil penetration by earthworms: direct measurements of hydroskeletal pressures and peristaltic motions.

Burrows resulting from earthworm activity are important for supporting various physical and ecological soil processes. Earthworm burrowing activity is quantified using models for earthworm penetration and cavity expansion that consider soil moisture and mechanical properties. Key parameters in these models are the maximal pressures exerted by the earthworm's hydroskeleton (estimated at 200 kPa). We designed a special pressure chamber that directly measures the pressures exerted by moving earthworms under different confining pressures to delineate the limits of earthworm activity in soils at different mechanical and hydration states. The chamber consists of a Plexiglas prism fitted with inner flexible tubing that hosts the earthworm. The gap around the tubing is pressurized using water, and the earthworm's peristaltic motion and concurrent pressure fluctuations were recorded by a camera and pressure transducer. A model that links the earthworm's kinematics with measured pressure fluctuations was developed. Resulting maximal values of radial pressures for anecic and endogeic earthworms were 130 kPa and 195 kPa, respectively. Mean earthworm peristaltic frequencies were used to quantify burrowing rates that were in agreement with previous results. The study delineates mechanical constraints to soil bioturbation by earthworms by mapping the elastic behaviour in the measurement chamber onto the expected elasto-viscoplastic environment of natural soils.