The high energetic cost of rapid force development in muscle

Muscles consume metabolic energy for active movement, particularly when performing mechanical work or producing force. Less appreciated is the cost for activating muscle quickly, which adds considerably to the overall cost of cyclic force production. However, the cost magnitude relative to the cost of mechanical work, which features in many movements, is unknown. We therefore tested whether fast activation is costly compared with performing work or producing isometric force. We hypothesized that metabolic cost would increase with a proposed measure termed force rate (rate of increase in muscle force) in cyclic tasks, separate from mechanical work or average force level. We tested humans (N=9) producing cyclic knee extension torque against an isometric dynamometer (torque 22 N m, cyclic waveform frequencies 0.5-2.5 Hz), while also quantifying quadriceps muscle force and work against series elasticity (with ultrasonography), along with metabolic rate through respirometry. Net metabolic rate increased by more than four-fold (10.5 to 46.8 W) with waveform frequency. At high frequencies, the hypothesized force-rate cost accounted for nearly half (40%) of energy expenditure. This exceeded the cost for average force (17%) and was comparable to the cost for shortening work (43%). The force-rate cost is explained by additional active calcium transport necessary for producing forces at increasing waveform frequencies, owing to rate-limiting dynamics of force production. The force-rate cost could contribute substantially to the overall cost of movements that require cyclic muscle activation, such as locomotion.