Mechanical transients initiated by ramp stretch and release to P0 in frog muscle fibers

Single fibers from the tibialis muscle of Rana temporaria were subjected to ramp stretches during tetanic stimulation at a sarcomere length of ~2 \u3bcm. Immediately after the stretch, or after different time delays, the active fiber was released against a constant force equal to the isometric force (P0) exerted immediately before the stretch. Four phases were detected after release: 1) an elastic recoil of the fiber's undamped elements, 2) a transient rapid shortening, 3) a marked reduction in the velocity of shortening (often to 0), and 4) an apparently steady shortening (sometimes absent). Increasing the amplitude of the stretch from ~2 to 10% of the fiber rest length led to an increase in phase 2 shortening from ~5 to 10 nm per half-sarcomere. Phase 2 shortening increased further (up to 14 nm per half-sarcomere) if a time interval of 5-10 ms was left between the end of large ramp stretches and release to P0. After 50- to 100-ms time intervals, shortening occurred in two steps of ~5 nm per half-sarcomere each. These findings suggest that phase 2 is due to charging, during and after the stretch, of a damped element, which can then shorten against P0 in at least two steps of ~5 nm/half sarcomere each

Mechanical work, oxygen consumption, and efficiency in isolated frog and rat muscle

The total work done during shortening, in repeated stretch-shortening cycles and the subsequent recovery oxygen consumption were measured in isolated frog (Rana esculenta) sartorius at 12 degrees C and rat (Wistar strain) extensor digitorum longus (EDL) and soleus at 20 degrees C. Two procedures were followed. In the first, the muscles were lengthened in the relaxed state and stimulated isometrically just before and during the first part of shortening. The peak efficiency (positive work done divided by the energetic equivalent of the oxygen consumed) was approximately 25% at 0.75-1.5 muscle lengths/s (Lo/s) in sartorius, 19% at 1.0 Lo/s in EDL, and 15% at 0.5 Lo/s in the soleus. In contrast to the measured efficiency values, the ratio between the tension-time integral and the oxygen consumption (the economy) is greater in soleus than in EDL. In the second procedure, stimulation began before stretching and continued during the first part of shortening. In this case, the efficiency attained values of approximately 35% in sartorius, 50% in EDL, and 40% in soleus. These values are in rough agreement with those measured in vivo during running

Efficiency of vertebrate locomotory muscles

We have examined the efficiency of vertebrate striated muscle at two different organizational levels: whole animals and isolated muscles. Terrestrial locomotion is used as a model of 'normal' muscular contraction; animal size and running speed are used as independent variables in order to change either the metabolic requirements of the muscles or the mechanical power production by the muscles over a wide range of values. The weight-specific metabolic power input to an animal increases nearly linearly with speed and increases with decreasing body size, while the weight-specific mechanical power output increases curvilinearly with speed and is independent of size. Consequently, the efficiency of the muscles in producing positive work increases with speed and the peak efficiency increases with increasing body size, attaining values of over 70% in large animals, but only 7% in small ones. The isolated muscle experiments were performed on frog muscle, and rat 'fast' and 'slow' muscles. We measured the work done, the oxygen consumed during recovery from the stimulation, and calculated the efficiency and the 'economy' (the cost of maintaining tension). The muscles were made to: (i) emulate the contractions seen during locomotion, i.e. shorten after a pre-stretch; or (ii) shorten at the same velocity and from the same muscle length as in (i), but without the pre-stretch. It was found that in mammalian muscles the peak efficiency with a pre-stretch attained high values, approaching the peak efficiencies in large animals. The maximum efficiency (attained at 1 length s-1 in fast muscle and at 0.5 lengths s-1 in slow muscle) did not differ much in the two muscles, whereas economy was greater in the slow muscle than in the fast muscle

Muscle work enhancement by stretch : passive visco-elasticity or cross-bridges?

Tetanized frog muscle fibres subjected to ramp stretches on the plateau of the tension-length relation, followed by an isotonic release against a load equal to the maximum isometric tension (T0), exhibit a well defined transient shortening against T0 which was attributed to the release of mechanical energy stored during stretching within the damped element of the cross-bridges. However, this interpretation has recently been challenged, and 'transient shortening against T0' has instead been attributed to elastic elements strained because of non-uniform distribution of lengthening within the fibre volume. The 'excess length change', resulting from the recoil of these elastic elements, was found i) to increase continuously with stretch amplitude up to 50 nm per h.s. with a 100 nm per h.s. strain, ii) to decrease steadily with the decrease in force during stress relaxation after the ramp stretch, and iii) to increase on the descending limb of the tension-length relation where sarcomere inhomogeneity is greater. In contrast, the transient shortening against T0: i) reaches a plateau at 8 nm per half sarcomere after about 50 nm per half sarcomere strain, ii) remains constant during the temperature dependent, fast phase of stress relaxation, when the excess in force above isometric reduces to about one half, iii) also occurs on the ascending limb of the tension-length relation where sarcomere inhomogeneity is drastically reduced. As a consequence of these differences we conclude that transient shortening and 'excess length change' do not "reflect the same underlying process"

Mechanical work in terrestrial locomotion: two basic mechanisms for minimizing energy expenditure

The work done during each step to lift and to reaccelerate (in the forward direction) and center of mass has been measured during locomotion in bipeds (rhea and turkey), quadrupeds (dogs, stump-tailed macaques, and ram), and hoppers (kangaroo and springhare). Walking, in all animals (as in man), involves an alternate transfer between gravitational-potential energy and kinetic energy within each stride (as takes place in a pendulum). This transfer is greatest at intermediate walking speeds and can account for up to 70% of the total energy changes taking place within a stride, leaving only 30% to be supplied by muscles. No kinetic-gravitational energy transfer takes place during running, hopping, and trotting, but energy is conserved by another mechanism: an elastic "bounce" of the body. Galloping animals utilize a combination of these two energy-conserving mechanisms. During running, trotting, hopping, and galloping, 1) the power per unit weight required to maintain the forward speed of the center of mass is almost the same in all the species studied; 2) the power per unit weight required to lift the center of mass is almost independent of speed; and 3) the sum of these two powers is almost a linear function of speed

Ergonomic evaluation of pathological gait

At each step of walking, the center of gravity of the body moves up and down and accelerates and decelerates forward with a combined movement that allows an appreciable transfer (R) between gravitational potential energy and kinetic energy, as occurs in a pendulum. The positive work and power to lift the center of gravity, to accelerate it forward, and to maintain its motion in a sagittal plane, the amount of R, the maximal height reached during each step by the center of gravity, and the step length and frequency are all determined by a microcomputer a few minutes after a subject walks on a force platform. This method is applied to the analysis of pathological gait in the attempt to measure quantitatively the alteration of the normal locomotory movement of the center of gravity. The strides of the patient are compared with the strides of normal subjects; in addition, the movement of the center of gravity of the patient during the stance on the affected limb is compared with the movement of the center of gravity during the stance on the unaffected limb, thus giving an index of the asymmetry of locomotion

Mechanical transients initiated by ramp stretch and release to Po in frog muscle fibers

Single fibers from the tibialis muscle of Rana temporaria were subjected to ramp stretches during tetanic stimulation at a sarcomere length of approximately 2 microns. Immediately after the stretch, or after different time delays, the active fiber was released against a constant force equal to the isometric force (Po) exerted immediately before the stretch. Four phases were detected after release: an elastic recoil of the fiber's undamped elements, a transient rapid shortening, a marked reduction in the velocity of shortening (often to 0), and an apparently steady shortening (sometimes absent). Increasing the amplitude of the stretch from approximately 2 to 10% of the fiber rest length led to an increase in phase 2 shortening from approximately 5 to 10 nm per half-sarcomere. Phase 2 shortening increased further (up to 14 nm per half-sarcomere) if a time interval of 5-10 ms was left between the end of large ramp stretches and release to Po. After 50- to 100-ms time intervals, shortening occurred in two steps of approximately 5 nm per half-sarcomere each. These findings suggest that phase 2 is due to charging, during and after the stretch, of a damped element, which can then shorten against Po in at least two steps of approximately 5 nm/half sarcomere each

Ergometric evaluation of pathological gait

At each step of walking, the center of gravity of the body moves up and down and accelerates and decelerates forward with a combined movement that allows an appreciable transfer (R) between gravitational potential energy and kinetic energy, as occurs in a pendulum. The positive work and power to lift the center of gravity, to accelerate it forward, and to maintain its motion in a sagittal plane, the amount of R, the maximal height reached during each step by the center of gravity, and the step length and frequency are all determined by a microcomputer a few minutes after a subject walks on a force platform. This method is applied to the analysis of pathological gait in the attempt to measure quantitatively the alteration of the normal locomotory movement of the center of gravity. The strides of the patient are compared with the strides of normal subjects; in addition, the movement of the center of gravity of the patient during the stance on the affected limb is compared with the movement of the center of gravity during the stance on the unaffected limb, thus giving an index of the asymmetry of locomotion