Reduce torques and stick the landing: limb posture during landing in toads

A controlled landing, where an animal does not crash or topple, requires enough stability to allow muscles to effectively dissipate mechanical energy. Toads (Rhinella marina) are exemplary models for understanding the mechanics and motor control of landing given their ability to land consistently during bouts of continuous hopping. Previous studies in anurans have shown that ground reaction forces (GRFs) during landing are significantly higher compared with takeoff and can potentially impart large torques about the center of mass (COM), destabilizing the body at impact. We predict that in order to minimize such torques, toads will align their COM with the GRF vector during the aerial phase in anticipation of impact. We combined high-speed videography and force-plate ergometry to quantify torques at the COM and relate the magnitude of torques to limb posture at impact. We show that modulation of hindlimb posture can shift the position of the COM by about 20% of snout-vent length. Rapid hindlimb flexion during the aerial phase of a hop moved the COM anteriorly and reduced torque by aligning the COM with the GRF vector. We found that the addition of extrinsic loads did not significantly alter landing behavior but did change the torques experienced at impact. We conclude that anticipatory hindlimb flexion during the aerial phase of a hop is a critical feature of a mechanically stable landing that allows toads to quickly string together multiple, continuous hops.