It is known that muscular force is reduced in old age. We investigate what are the effects of this phenomenon on the mechanics of running. We hypothesized that the deficit in force would result in a lower push, causing reduced amplitude of the vertical oscillation, with smaller elastic energy storage and increased step frequency. To test this hypothesis, we measured the mechanical energy of the centre of mass of the body during running in old and young subjects. The amplitude of the oscillation is indeed reduced in the old subjects, resulting in an approximately 20% smaller elastic recovery and a greater step frequency (3.7 versus 2.8 Hz, pZ1.9!10 K5 , at 15-17 km h K1 ). Interestingly, the greater step frequency is due to a lower aerial time, and not to a greater natural frequency of the system, which is similar in old and young subjects (3.6 versus 3.4 Hz, pZ0.2). Moreover, we find that in the old subjects, the step frequency is always similar to the natural frequency, even at the highest speeds. This is at variance with young subjects who adopt a step frequency lower than the natural frequency at high speeds, to contain the aerobic energy expenditure. Finally, the external work to maintain the motion of the centre of mass is reduced in the old subjects (0.9 versus 1.2 J kg K1 m K1 , pZ5.1!10 K6 ) due to the lower work done against gravity, but the higher step frequency involves a greater internal work to reset the limbs at each step. The net result is that the total work increases with speed more steeply in the old subjects than in young subjects.

SUMMARYThe landing-take-off asymmetry of running was thought to derive from, or at least to be consistent with, the physiological property of muscle to resist stretching (after landing) with a force greater than it can develop during shortening (before take-off). In old age, muscular force is reduced, but the deficit in force is less during stretching than during shortening. The greater loss in concentric versus eccentric strength with aging led us to hypothesize that older versus younger adults would increase the landing-take-off asymmetry in running. To test this hypothesis, we measured the within-step changes in mechanical energy of the centre of mass of the body in old and young subjects. The difference between the peaks in kinetic energy attained during the fall and during the lift of the centre of mass is greater in the old subjects. The difference between the time to lift and accelerate the centre of mass (positive work) and to absorb the same amount of energy during the downward displacement (negative work) is also greater in the old subjects. Both these findings imply a difference in force between stretching and shortening during the bounce, which is greater in the old subjects than in the young subjects. This is qualitatively consistent with the more asymmetric force-velocity relation found in aged muscle and supports, even if does not prove, the hypothesis that the landing-take-off asymmetry in running derives from the different response of muscle to stretching and shortening.

Human running at low and intermediate speeds is characterized by a greater average force exerted after 'landing', when muscle-tendon units are stretched ('hard landing'), and a lower average force exerted before 'takeoff ', when muscle-tendon units shorten ('soft takeoff '). This landing-takeoff asymmetry is consistent with the force-velocity relation of the 'motor' (i.e. with the basic property of muscle to resist stretching with a force greater than that developed during shortening), but it may also be due to the 'machine' (e.g. to the asymmetric lever system of the foot operating during stance). Hard landing and soft takeoff-never the reverse-were found in running, hopping and trotting animals using diverse lever systems, suggesting that the different machines evolved to comply with the basic force-velocity relation of the motor. Here we measure the mechanical energy of the centre of mass of the body in backward running, an exercise where the normal coupling between motor and machine is voluntarily disrupted, in order to see the relevance of the motor-machine interplay in human running. We find that the landing-takeoff asymmetry is reversed. The resulting 'soft landing' and 'hard takeoff ' are associated with a reduced efficiency of positive work production. We conclude that the landing-takeoff asymmetry found in running, hopping and trotting is the expression of a convenient interplay between motor and machine. More metabolic energy must be spent in the opposite case when muscle is forced to work against its basic property (i.e. when it must exert a greater force during shortening and a lower force during stretching).

The bouncing mechanism of human running is characterized by a shorter duration of the brake after ‘landing’ compared with a longer duration of the push before ‘takeoff’. This landing–takeoff asymmetry has been thought to be a consequence of the force–velocity relation of the muscle, resulting in a greater force exerted during stretching after landing and a lower force developed during shortening before takeoff. However, the asymmetric lever system of the human foot during stance may also be the cause. Here, we measure the landing–takeoff asymmetry in bouncing steps of running, hopping and trotting animals using diverse lever systems. We find that the duration of the push exceeds that of the brake in all the animals, indicating that the different lever systems comply with the basic property of muscle to resist stretching with a force greater than that developed during shortening. In addition, results show both the landing–takeoff asymmetry and the mass-specific vertical stiffness to be greater in small animals than in large animals. We suggest that the landing–takeoff asymmetry is an index of a lack of elasticity, which increases with increasing the role of muscle relative to that of tendon within muscle–tendon units.