Walking and Running Require Greater Effort from the Ankle than the Knee Extensor Muscles :

The amount of energy used to run a mile is nearly the same whether it is run at top speed or at a leisurely pace (although it is used more rapidly at the higher speed). This puzzling independence of energy cost and speed is found generally among running animals, although, on a per gram basis, cost is much higher for smaller animals. Running involves little work against the environment; work is done by muscles and tendons to lift and accelerate the body and limbs. Some of the work is recovered from muscle-tendon springs without metabolic cost and work rate does not parallel metabolic rate with either speed or size. Regardless of the amount of work muscles do, they must be activated and develop force to support the weight of the body. Load-carrying experiments have shown that the cost of supporting an extra newton of load is the same as the weight-specific cost of running. Size differences in cost are proportional to stride frequency at equivalent speeds, suggesting that the time available for developing force is important in determining cost. We report a simple inverse relationship between the rate of energy used for running and the time the foot applies force to the ground during each stride. These results support the hypothesis that it is primarily the cost of supporting the animal's weight and the time course of generating this force that determines the cost of running.

We twice tested the hypothesis that top running speeds are determined by the amount of force applied to the ground rather than how rapidly limbs are repositioned in the air. First, we compared the mechanics of 33 subjects of different sprinting abilities running at their top speeds on a level treadmill. Second, we compared the mechanics of declined (-6 degrees ) and inclined (+9 degrees ) top-speed treadmill running in five subjects. For both tests, we used a treadmill-mounted force plate to measure the time between stance periods of the same foot (swing time, t(sw)) and the force applied to the running surface at top speed. To obtain the force relevant for speed, the force applied normal to the ground was divided by the weight of the body (W(b)) and averaged over the period of foot-ground contact (F(avge)/W(b)). The top speeds of the 33 subjects who completed the level treadmill protocol spanned a 1.8-fold range from 6.2 to 11.1 m/s. Among these subjects, the regression of F(avge)/W(b) on top speed indicated that this force was 1.26 times greater for a runner with a top speed of 11.1 vs. 6.2 m/s. In contrast, the time taken to swing the limb into position for the next step (t(sw)) did not vary (P = 0.18). Declined and inclined top speeds differed by 1.4-fold (9.96+/-0.3 vs. 7.10+/-0.3 m/s, respectively), with the faster declined top speeds being achieved with mass-specific support forces that were 1.3 times greater (2.30+/- 0.06 vs. 1.76+/-0.04 F(avge)/ W(b)) and minimum t(sw) that were similar (+8%). We conclude that human runners reach faster top speeds not by repositioning their limbs more rapidly in the air, but by applying greater support forces to the ground.