Spinal Mechanisms May Provide a Combination of Intermittent and Continuous Control of Human Posture: Predictions from a Biologically Based Neuromusculoskeletal Model

Abstract

Several models have been employed to study human postural control during upright quiet stance. Most have adopted an inverted pendulum approximation to the standing human and theoretical models to account for the neural feedback necessary to keep balance. The present study adds to the previous efforts in focusing more closely on modelling the physiological mechanisms of important elements associated with the control of human posture. This paper studies neuromuscular mechanisms behind upright stance control by means of a biologically based large-scale neuromusculoskeletal (NMS) model. It encompasses: i) conductance-based spinal neuron models (motor neurons and interneurons); ii) muscle proprioceptor models (spindle and Golgi tendon organ) providing sensory afferent feedback; iii) Hill-type muscle models of the leg plantar and dorsiflexors; and iv) an inverted pendulum model for the body biomechanics during upright stance. The motor neuron pools are driven by stochastic spike trains. Simulation results showed that the neuromechanical outputs generated by the NMS model resemble experimental data from subjects standing on a stable surface. Interesting findings were that: i) an intermittent pattern of muscle activation emerged from this posture control model for two of the leg muscles (Medial and Lateral Gastrocnemius); and ii) the Soleus muscle was mostly activated in a continuous manner. These results suggest that the spinal cord anatomy and neurophysiology (e.g., motor unit types, synaptic connectivities, ordered recruitment), along with the modulation of afferent activity, may account for the mixture of intermittent and continuous control that has been a subject of debate in recent studies on postural control. Another finding was the occurrence of the so-called “paradoxical” behaviour of muscle fibre lengths as a function of postural sway. The simulations confirmed previous conjectures that reciprocal inhibition is possibly contributing to this effect, but on the other hand showed that this effect may arise without any anticipatory neural control mechanism.

Author Summary

The control of upright stance is a challenging task since the objective is to maintain the equilibrium of an intrinsically unstable biomechanical system. Somatosensory information is used by the central nervous system to modulate muscle contraction, which prevents the body from falling. While the visual and vestibular systems also provide important additional sensory information, a human being with only somatosensory inputs is able to maintain an upright stance. In this study, we used a biologically-based large-scale neuromusculoskeletal model driven only by somatosensory feedback to investigate human postural control from a neurophysiological point of view. No neural structures above the spinal cord were included in the model. The results showed that the model based on a spinal control of posture can reproduce several neuromechanical outcomes previously reported in the literature, including an intermittent muscle activation. Since this intermittent muscular recruitment is an emergent property of this spinal-like controller, we argue that the so-called intermittent control of upright stance might be produced by an interplay between spinal cord properties and modulated sensory inflow.