Humans use multi-objective control to regulate lateral foot placement when walking.

Abstract

A fundamental question in human motor neuroscience is to determine how the nervous system generates goal-directed movements despite inherent physiological noise and redundancy. Walking exhibits considerable variability and equifinality of task solutions. Existing models of bipedal walking do not yet achieve both continuous dynamic balance control and the equifinality of foot placement humans exhibit. Appropriate computational models are critical to disambiguate the numerous possibilities of how to regulate stepping movements to achieve different walking goals. Here, we extend a theoretical and computational Goal Equivalent Manifold (GEM) framework to generate predictive models, each posing a different experimentally testable hypothesis. These models regulate stepping movements to achieve any of three hypothesized goals, either alone or in combination: maintain lateral position, maintain lateral speed or “heading”, and/or maintain step width. We compared model predictions against human experimental data. Uni-objective control models demonstrated clear redundancy between stepping variables, but could not replicate human stepping dynamics. Most multi-objective control models that balanced maintaining two of the three hypothesized goals also failed to replicate human stepping dynamics. However, multi-objective models that strongly prioritized regulating step width over lateral position did successfully replicate all of the relevant step-to-step dynamics observed in humans. Independent analyses confirmed this control was consistent with linear error correction and replicated step-to-step dynamics of individual foot placements. Thus, the regulation of lateral stepping movements is inherently multi-objective and balances task-specific trade-offs between competing task goals. To determine how people walk in their environment requires understanding both walking biomechanics and how the nervous system regulates movements from step-to-step. Analogous to mechanical “templates” of locomotor biomechanics, our models serve as “control templates” for how humans regulate stepping movements from each step to the next. These control templates are symbiotic with well-established mechanical templates, providing complimentary insights into walking regulation.