Modulation of joint moments and work in the goat hindlimb with locomotor speed and surface grade

Goats and other quadrupeds must modulate the work output of their muscles to accommodate the changing mechanical demands associated with locomotion in their natural environments. This study examined which hindlimb joint moments goats use to generate and absorb mechanical energy on level and sloped surfaces over a range of locomotor speeds. Ground reaction forces and the three-dimensional locations of joint markers were recorded as goats walked, trotted and galloped over 0, +15 and -15 deg sloped surfaces. Net joint moments, powers and work were estimated at the goats' hip, knee, ankle and metatarsophalangeal joints throughout the stance phase via inverse dynamics calculations. Differences in locomotor speed on the level, inclined and declined surfaces were characterized and accounted for by fitting regression equations to the joint moment, power and work data plotted versus non-dimensionalized speed. During level locomotion, the net work generated by moments at each of the hindlimb joints was small (less than 0.1 J kg(-1) body mass) and did not vary substantially with gait or locomotor speed. During uphill running, by contrast, mechanical energy was generated at the hip, knee and ankle, and the net work at each of these joints increased dramatically with speed (P<0.05). The greatest increases in positive joint work occurred at the hip and ankle. During downhill running, mechanical energy was decreased in two main ways: goats generated larger knee extension moments in the first half of stance, absorbing energy as the knee flexed, and goats generated smaller ankle extension moments in the second half of stance, delivering less energy. The goats' hip extension moment in mid-stance was also diminished, contributing to the decrease in energy. These analyses offer new insight into quadrupedal locomotion, clarifying how the moments generated by hindlimb muscles modulate mechanical energy at different locomotor speeds and grades, as needed to accommodate the demands of variable terrain.