Abstract

Lateral balance is a critical factor in keeping the human body upright during walking. Two important mechanisms for balance control are the stepping strategy, in which the foot placement is changed in the direction of a sensed fall to modulate how the gravitational force acts on the body, and the lateral ankle strategy, in which the body mass is actively accelerated by an ankle torque. Currently, there is minimal evidence about how these two strategies complement one another to achieve upright balance during locomotion. We use Galvanic vestibular stimulation (GVS) to induce the sensation of a fall at heel-off during gait initiation. We found that young healthy adults respond to the illusory fall using both the lateral ankle strategy and the stepping strategy. The stance foot center of pressure (CoP) is shifted in the direction of the perceived fall by ≈2.5 mm, starting ≈247 ms after stimulus onset. The foot placement of the following step is shifted by ≈15 mm in the same direction. The temporal delay between these two mechanisms suggests that they independently contribute to upright balance during locomotion, potentially in a serially coordinated manner. Modeling results indicate that without the lateral ankle strategy, a much larger step width is required to maintain upright balance, suggesting that the small but early CoP shift induced by the lateral ankle strategy is critical for upright stability during locomotion. The relative importance of each mechanism and how neurological disorders may affect their implementation remain an open question.

Highlights

  • Upright balance is a critical factor of everyday human life, forming the basis for functional behavior like tool use

  • The neural control of balance has largely been studied under conditions of standing, with much of the analysis focusing on the different channels of sensory information detecting deviations in the kinematic state of the body and the modulation of the center of pressure (CoP) to correct them [1]

  • The control problem in posture has often been simplified to a single degree of freedom in the inverted pendulum approximation, it has become clear in recent years that even in PLOS ONE | DOI:10.1371/journal.pone

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Summary

Introduction

Upright balance is a critical factor of everyday human life, forming the basis for functional behavior like tool use. Later work by Hof showed that humans respond to mechanical lateral perturbations by changing the foot placement in the direction of the push [15]. This response is preceded by activity in hip abductor muscles [16], suggesting an actively generated response. In a study of unperturbed gait, Wang and Srinivasan observed a correlation between the position and velocity of the pelvis in mid-stance and the following foot placement, indicating that such “stepping in the direction of the fall” is a regular mechanism of balance control in normal gait [17]

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