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1 Department of Integrative Physiology, University of Colorado, Boulder, CO, USA; Department of Integrative Biology, University of California, Berkeley, CA`, USA
2 Department of Integrative Physiology, University of Colorado, Boulder, CO, USA
* To whom correspondence should be addressed. E-mail: chet.moritz{at}colorado.edu.
Humans running and hopping maintain similar center of mass motions despite large changes in surface stiffness and damping. The goal of this study was to determine the contributions of anticipation and reaction when human hoppers encounter surprise, expected and random changes from a soft elastic surface (27 kN/m) to a hard surface (411 kN/m). Subjects encountered the expected hard surface on every fourth hop and the random hard surface on an average of 25% of the hops in a trial. When hoppers on a soft surface were surprised by a hard surface, the ankle and knee joints were forced into greater flexion by passive interaction with the hard surface. Within 52 ms after landing on the surprise hard surface, joint flexion increased, and the legs became less stiff than on the soft surface. These mechanical changes occurred before EMG first changed 68-188 ms after landing. Due to the fast mechanical reaction to the surprise hard surface, center of mass displacement and average leg stiffness were the same as on expected and random hard surfaces. This similarity is striking because subjects anticipated the expected and random hard surfaces by landing with their knees more flexed. Subjects also anticipated the expected hard surface by increasing the level of EMG by 24-76% during the 50 ms before landing. These results show that passive mechanisms alter leg stiffness for unexpected surface changes before muscle EMG changes and may be critical for adjustments to variable terrain encountered during locomotion in the natural world.
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