Ground Reaction Forces and Energy Exchange During Underwater Walking.

Underwater walking was a crucial step in the evolutionary transition from water to land. Underwater walkers use fins and/or limbs to interact with the benthic substrate and produce propulsive forces. The dynamics of underwater walking remain poorly understood due to the lack of a sufficiently sensitive and waterproof system to measure substrate reaction forces (SRFs). Using an underwater force plate (described in our companion paper), we quantify SRFs during underwater walking in axolotls (Ambystoma mexicanum) and Spot prawn (Pandalus platyceros), synchronized with videography. The horizontal propulsive forces were greater than the braking forces in both species to overcome hydrodynamic drag. In axolotls, potential energy (PE) fluctuations were far smaller than kinetic energy (KE) fluctuations due to high buoyant support (97%), whereas the magnitudes were similar in the prawn due to lower buoyant support (93%). However, both species show minimal evidence of exchange between KE and PE, which, along with the effects of hydrodynamic drag, is incompatible with inverted pendulum dynamics. Our results show that, despite their evolutionary links, underwater walking has fundamentally different dynamics compared with terrestrial walking and emphasize the substantial consequences of differences in body plan in underwater walking.