The intersection of ground reaction forces near a point above the center of mass has been observed in computer simulation models and human walking experiments. Observed so ubiquitously, the intersection point (IP) is commonly assumed to provide postural stability for bipedal walking. In this study, we challenge this assumption by questioning if walking without an IP is possible. Deriving gaits with a neuromuscular reflex model through multi-stage optimization, we found stable walking patterns that show no signs of the IP-typical intersection of ground reaction forces. The non-IP gaits found are stable and successfully rejected step-down perturbations, which indicates that an IP is not necessary for locomotion robustness or postural stability. A collision-based analysis shows that non-IP gaits feature center of mass (CoM) dynamics with vectors of the CoM velocity and ground reaction force increasingly opposing each other, indicating an increased mechanical cost of transport. Although our computer simulation results have yet to be confirmed through experimental studies, they already indicate that the role of the IP in postural stability should be further investigated. Moreover, our observations on the CoM dynamics and gait efficiency suggest that the IP may have an alternative or additional function that should be considered.

Muscle fibres possess unique visco-elastic properties, which generate a stabilising zero-delay response to unexpected perturbations. This instantaneous response—termed “preflex”—mitigates neuro-transmission delays, which are hazardous during fast locomotion due to the short stance duration. While the elastic contribution to preflexes has been studied extensively, the function of fibre viscosity due to the force–velocity relation remains unknown. In this study, we present a novel approach to isolate and quantify the preflex force produced by the force–velocity relation in musculo-skeletal computer simulations. We used our approach to analyse the muscle response to ground-level perturbations in simulated vertical hopping. Our analysis focused on the preflex-phase—the first 30 ms after impact—where neuronal delays render a controlled response impossible. We found that muscle force at impact and dissipated energy increase with perturbation height, helping reject the perturbations. However, the muscle fibres reject only 15\% of step-down perturbation energy with constant stimulation. An open-loop rising stimulation, observed in locomotion experiments, amplified the regulatory effects of the muscle fibre’s force–velocity relation, resulting in 68\% perturbation energy rejection. We conclude that open-loop neuronal tuning of muscle activity around impact allows for adequate feed-forward tuning of muscle fibre viscous capacity, facilitating energy adjustment to unexpected ground-level perturbations.

Neuromuscular control loops feature substantial communication delays, but mammals run robustly even in the most adverse conditions. In-vivo experiments and computer simulation results suggest that muscles’ preflex—an immediate mechanical response to a perturbation—could be the critical contributor. Muscle preflexes act within a few milliseconds, an order of magnitude faster than neural reflexes. Their short-lasting activity makes mechanical preflexes hard to quantify in-vivo. Muscle models, on the other hand, require further improvement of their prediction accuracy during the non-standard conditions of perturbed locomotion. Additionally, muscles mechanically adapt by increased damping force. Our study aims to quantify the mechanical preflex work and test its mechanical force adaptation. We performed in-vitro experiments with biological muscle fibers under physiological boundary conditions, which we determined in computer simulations of perturbed hopping. Our findings show that muscles initially resist impacts with a stereotypical sti↵ness response—identified as short-range sti↵ness—regardless of the exact perturbation condition. We then observe a velocity adaptation to the force related to the amount of perturbation. The main contributor to the preflex work adaptation is not the force di↵erence but the muscle fiber stretch di↵erence. We find that both muscle sti↵ness and damping are activity-dependent properties. These results indicate that neural control could tune the preflex properties of muscles in expectation of ground conditions leading to previously inexplicable neuromuscular adaptation speeds.