The spring-mass model is a model of locomotion aimed at giving the essential mathematical laws of the trajectory of the center of mass of an animal during bouncing gaits, such as hopping (one-dimensional) and running (two-dimensional). This reductionist mechanical system has been extensively investigated for locomotion over horizontal surfaces, whereas it has been largely neglected on other ecologically relevant surfaces, including inclines. For example, how the degree of inclination impacts the dynamics of the center of mass of the spring-mass model has not been investigated thoroughly. In this work, we derive a mathematical model which extends the spring-mass model to inclined surfaces. Among our results, we derive an approximate solution of the system, assuming a small angular sweep of the limb and a small spring compression during stance, and show that this approximation is very accurate, especially for small inclinations of the ground. Furthermore, we derive theoretical bounds on the difference between the Lagrangian and Lagrange equationsÂ of the true and approximate systems, and discuss locomotor stability questions of the approximate solutions. We test our models through a sensitivity analysis using parameters relevant to the locomotion of bipedal animals (quail, pheasant, guinea fowl, turkey, ostrich, and humans) and compare our approximate solution to the numerically derived solution of the exact system. We compare the two-dimensional spring-mass model on inclines with the one-dimensional spring-mass model to which it reduces under the limit of no horizontal velocity; we compare the two-dimensional spring-mass model on inclines with the inverted pendulum model on inclines towards which it converges in the case of high stiffness-to-mass ratio. We include comparisons with historically prevalent no-gravity approximations of these models, as well. The insights we have gleaned through all these comparisons and the ability of our approximation to replicate some of the kinematic changes observed in animals moving on different inclines (e.g., reduction in vertical oscillation of the center of mass and decreased stride length) underline the valuable and reasonable contributions that very simple, reductionist models, like the spring-mass model, can provide.

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