Grounded running in quails: Simulations indicate benefits of observed fixed aperture angle between legs before touch-down

Many birds use grounded running (running without aerial phases) in a wide range of speeds. Contrary to walking and running, numerical investigations of this gait based on the BSLIP (bipedal spring loaded inverted pendulum) template are rare. To obtain template related parameters of quails (e.g. leg stiffness) we used x-ray cinematography combined with ground reaction force measurements of quail grounded running. Interestingly, with speed the quails did not adjust the swing leg's angle of attack with respect to the ground but adapted the angle between legs (which we termed aperture angle), and fixed it about 30 ms before touchdown. In simulations with the BSLIP we compared this swing leg alignment policy with the fixed angle of attack with respect to the ground typically used in the literature. We found symmetric periodic grounded running in a simply connected subset comprising one third of the investigated parameter space. The fixed aperture angle strategy revealed improved local stability and surprising tolerance with respect to large perturbations. Starting with the periodic solutions, after step-down step-up or step-up step-down perturbations of 10% leg rest length, in the vast majority of cases the bipedal SLIP could accomplish at least 50 steps to fall. The fixed angle of attack strategy was not feasible. We propose that, in small animals in particular, grounded running may be a common gait that allows highly compliant systems to exploit energy storage without the necessity of quick changes in the locomotor program when facing perturbations.