Muscle activation during unilateral stepping occurs in the nonstepping limb of humans with clinically complete spinal cord injury.

STUDY DESIGN: Comparison of different kinematic and loading conditions on muscle activation in clinically complete spinal cord-injured subjects stepping unilaterally with manual assistance. OBJECTIVE: To determine if rhythmic lower limb loading or movement could produce rhythmic muscle activation in the nonstepping limb of subjects with clinically complete spinal cord injury (SCI). SETTING: Human Locomotion Research Center, Department of Neurology, University of California, Los Angeles, USA. METHODS: We recorded electromyography, joint kinematics, and vertical ground reaction forces as four subjects with clinically complete SCI stepped with manual assistance and partial bodyweight support. For all trials, one limb continuously stepped while the other limb underwent different conditions, including rhythmic lower limb loading in an extended position without limb movement, rhythmic lower limb movement similar to stepping without limb loading, and no lower limb loading or movement with the leg in an extended or flexed position. RESULTS: Three subjects displayed rhythmic muscle activity in the nonstepping limb for trials with rhythmic limb loading, but no limb movement. One subject displayed rhythmic muscle activity in the nonstepping limb for trials without ipsilateral limb loading or movement. The rhythmic muscle activity in the nonstepping limb was similar to the rhythmic muscle activity during bilateral stepping. CONCLUSIONS: The human spinal cord can use sensory information about ipsilateral limb loading to increase muscle activation even when there is no limb movement. The results also indicate that movement and loading in one limb can produce rhythmic muscle activity in the other limb even when it is stationary and unloaded. These findings emphasize the importance of optimizing load-related and contralateral sensory input during gait rehabilitation after SCI.

Soleus H-reflex gain in humans walking and running under simulated reduced gravity.

The Hoffmann (H-) reflex is an electrical analogue of the monosynaptic stretch reflex, elicited by bypassing the muscle spindle and directly stimulating the afferent nerve. Studying H-reflex modulation provides insight into how the nervous system centrally modulates stretch reflex responses.A common measure of H-reflex gain is the slope of the relationship between H-reflex amplitude and EMG amplitude. To examine soleus H-reflex gain across a range of EMG levels during human locomotion, we used simulated reduced gravity to reduce muscle activity. We hypothesised that H-reflex gain would be independent of gravity level.We recorded EMG from eight subjects walking (1.25 m s-1) and running (3.0 m s-1) at four gravity levels (1.0, 0.75, 0.5 and 0.25 G (Earth gravity)). We normalised the stimulus M-wave and resulting H-reflex to the maximal M-wave amplitude (Mmax) elicited throughout the stride to correct for movement of stimulus and recording electrodes relative to nerve and muscle fibres. Peak soleus EMG amplitude decreased by ~30% for walking and for running over the fourfold change in gravity. As hypothesised, slopes of linear regressions fitted to H-reflex versus EMG data were independent of gravity for walking and running (ANOVA, P > 0.8). The slopes were also independent of gait (P > 0.6), contrary to previous studies. Walking had a greater y-intercept (19.9% Mmax) than running (-2.5% Mmax; P < 0.001). At all levels of EMG, walking H-reflex amplitudes were higher than running H-reflex amplitudes by a constant amount. We conclude that the nervous system adjusts H-reflex threshold but not H-reflex gain between walking and running. These findings provide insight into potential neural mechanisms responsible for spinal modulation of the stretch reflex during human locomotion.

Runners adjust leg stiffness for their first step on a new running surface.

Human runners adjust the stiffness of their stance leg to accommodate surface stiffness during steady state running. This adjustment allows runners to maintain similar center of mass movement (e.g., ground contact time and stride frequency) regardless of surface stiffness. When runners encounter abrupt transitions in the running surface, they must either make a rapid adjustment or allow the change in the surface stiffness to disrupt their running mechanics. Our goal was to determine how quickly runners adjust leg stiffness when they encounter an abrupt but expected change in surface stiffness that they have encountered previously. Six human subjects ran at 3 m s(-1) on a rubber track with two types of rubber surfaces: a compliant "soft" surface (ksurf = 21.3 kN m(-1) and a non-compliant "hard" surface (ksurf = 533 kN m(-1). We found that runners completely adjusted leg stiffness for their first step on the new surface after the transition. For example, runners decreased leg stiffness by 29% between the last step on the soft surface and the first step on the hard surface (from 10.7 kN m(-1) to 7.6 kN m(-1), respectively). As a result, the vertical displacement of the center of mass during stance ( approximately 7 cm) did not change at the transition despite a reduction in surface compression from 6 cm to less than 0.25 cm. By rapidly adjusting leg stiffness, each runner made a smooth transition between surfaces so that the path of the center of mass was unaffected by the change in surface stiffness.

Running in the real world: adjusting leg stiffness for different surfaces.

A running animal coordinates the actions of many muscles, tendons, and ligaments in its leg so that the overall leg behaves like a single mechanical spring during ground contact. Experimental observations have revealed that an animal's leg stiffness is independent of both speed and gravity level, suggesting that it is dictated by inherent musculoskeletal properties. However, if leg stiffness was invariant, the biomechanics of running (e.g. peak ground reaction force and ground contact time) would change when an animal encountered different surfaces in the natural world. We found that human runners adjust their leg stiffness to accommodate changes in surface stiffness, allowing them to maintain similar running mechanics on different surfaces. These results provide important insight into mechanics and control of animal locomotion and suggest that incorporating an adjustable leg stiffness in the design of hopping and running robots is important if they are to match the agility and speed of animals on varied terrain.

Effect of reduced gravity on the preferred walk-run transition speed.

We investigated the effect of reduced gravity on the human walk-run gait transition speed and interpreted the results using an inverted-pendulum mechanical model. We simulated reduced gravity using an apparatus that applied a nearly constant upward force at the center of mass, and the subjects walked and ran on a motorized treadmill. In the inverted pendulum model for walking, gravity provides the centripetal force needed to keep the pendulum in contact with the ground. The ratio of the centripetal and gravitational forces (mv2/L)/(mg) reduces to the dimensionless Froude number (v2/gL). Applying this model to a walking human, m is body mass, v is forward velocity, L is leg length and g is gravity. In normal gravity, humans and other bipeds with different leg lengths all choose to switch from a walk to a run at different absolute speeds but at approximately the same Froude number (0.5). We found that, at lower levels of gravity, the walk-run transition occurred at progressively slower absolute speeds but at approximately the same Froude number. This supports the hypothesis that the walk-run transition is triggered by the dynamics of an inverted-pendulum system.

Interaction of leg stiffness and surfaces stiffness during human hopping.

When mammals run, the overall musculoskeletal system behaves as a single linear "leg spring". We used force platform and kinematic measurements to determine whether leg spring stiffness (k(leg)) is adjusted to accommodate changes in surface stiffness (ksurf) when humans hoop in place, a good experimental model for examining adjustments to k(leg) in bouncing gaits. We found that k(leg) was greatly increased to accommodate surfaces of lower stiffnesses. The series combination of k(leg) and ksurf [total stiffness (ktot)] was independent of ksurf at a given hopping frequency. For example, when humans hopped at a frequency of 2 Hz, they tripled their k(leg) on the least stiff surface (ksurf = 26.1 kN/m; k(leg) = 53.3 kN/m) compared with the most stiff surface (ksurf = 35,000 kN/m; k(leg) = 17.8 kN/m). Values for ktot were not significantly different on the least stiff surface (16.7 kN/m) and the most stiff surface (17.8 kN/m). Because of the k(leg) adjustment, many aspects of the hopping mechanics (e.g., ground-contact time and center of mass vertical displacement) remained remarkably similar despite a > 1,000-fold change in ksurf. This study provides insight into how k(leg) adjustments can allow similar locomotion mechanics on the variety of terrains encountered by runners in the natural world.