SUMMARYWe investigated the mechanical function of two ankle extensor muscles, the lateral gastrocnemius (LG) and peroneus longus (PL), in wild turkeys Meleagris gallopavo during steady speed running. We hypothesized that mechanical work output of the LG and PL during running parallels the demand for mechanical work on the body. The turkeys ran on level, inclined (+6°,+12°) and declined (–6°, –12°) treadmills to change the demand for mechanical work. Simultaneous measurements of muscle length(from sonomicrometry) and muscle force (from tendon strain gauges) were used to calculate mechanical work output.During level running at a speed of 2 m s–1, the LG and PL were both active in stance but produced peak force at different times, at approximately 21% of stance duration for the LG and 70% for the PL. The LG and PL also had different length patterns in stance during level running. The LG underwent little shortening during force production, resulting in negligible net positive work (2.0±0.8 J kg–1). By contrast, the PL produced force across a stretch–shorten cycle in stance and did significant net positive work (4.7±1.6 J kg–1). Work outputs for both the LG and PL were directly proportional to running slope. When we increased the demand for net positive work by running the turkeys on an incline, the LG and PL increased stance net positive work output in direct proportion to slope (P<0.05). Stance net positive work output increased to 7.0±1.3 J kg–1 for the LG and 8.1±2.9 J kg–1 for the PL on the steepest incline. Increases in stance net positive work for the LG and PL were associated with increases in net shortening strain and average shortening velocity, but average force in stance remained constant. The LG and PL muscles were also effective energy absorbers during decline running, when there is demand for net negative work on the body. During decline running at 2 m s–1 on the steepest slope, the LG absorbed 4.6±2.2 J kg–1 of net work in stance and the PL absorbed 2.4±0.9 J kg–1 of net work. Shifts in muscle mechanical function from energy production during incline running to energy absorption during decline running were observed over a range of running speeds from 1–3 m s–1 for both the LG and PL.Two fundamentally different mechanisms for changing work output were apparent in the mechanical behavior of the LG and PL. The LG simply altered its length pattern; it actively shortened during incline running to produce mechanical energy and actively lengthened during decline running to absorb mechanical energy. The PL changed mechanical function by altering its length pattern and by shifting the timing of force production across its stretch–shorten cycle. During incline running, the PL produced force during late stance shortening for positive work, but during decline running,the timing of force production shifted into early stance, to align with lengthening for negative work. In addition, during decline running, the PL greatly reduced or eliminated late stance shortening, thus reducing the potential for positive work.Our results show that the changing demands for whole body work during steady speed running are met, at least in part, by an ability of single muscles to shift mechanical function from net energy production to net energy absorption.

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. The consistent patterns of strain changes with running slope are evidence that strain pattern is modulated to meet the changes in demand for net mechanical work. The relatively poor relationship between strain and running speed may reflect the fact that changes in running speed during level running are not associated with a change in demand for net mechanical work. Taken together, the speed and slope results suggest that the demand for mechanical work is an important determinant of muscle length patterns in running and walking.

SUMMARYLittle is known about how in vivo muscle efficiency, that is the ratio of mechanical and metabolic power, is affected by changes in locomotory tasks. One of the main problems with determining in vivo muscle efficiency is the large number of muscles generally used to produce mechanical power. Animal flight provides a unique model for determining muscle efficiency because only one muscle, the pectoralis muscle, produces nearly all of the mechanical power required for flight. In order to estimate in vivo flight muscle efficiency, we measured the metabolic cost of flight across a range of flight speeds (6-13ms -1 ) using masked respirometry in the cockatiel (Nymphicus hollandicus) and compared it with measurements of mechanical power determined in the same wind tunnel. Similar to measurements of the mechanical power-speed relationship, the metabolic power-speed relationship had a U-shape, with a minimum at 10ms -1 . Although the mechanical and metabolic power-speed relationships had similar minimum power speeds, the metabolic power requirements are not a simple multiple of the mechanical power requirements across a range of flight speeds. The pectoralis muscle efficiency (estimated from mechanical and metabolic power, basal metabolism and an assumed value for the 'postural costs' of flight) increased with flight speed and ranged from 6.9% to 11.2%. However, it is probable that previous estimates of the postural costs of flight have been too low and that the pectoralis muscle efficiency is higher.

The force-velocity properties of skeletal muscle have an important influence on locomotor performance. All skeletal muscles produce less force the faster they shorten and typically develop maximal power at velocities of approximately 30% of maximum shortening velocity (V(max)). We used direct measurements of muscle mechanical function in two ankle extensor muscles of wild turkeys to test the hypothesis that during level running muscles operate at velocities that favor force rather than power. Sonomicrometer measurements of muscle length, tendon strain-gauge measurements of muscle force, and bipolar electromyographs were taken as animals ran over a range of speeds and inclines. These measurements were integrated with previously measured values of muscle V(max) for these muscles to calculate relative shortening velocity (V/V(max)). At all speeds for level running the V/V(max) values of the lateral gastrocnemius and the peroneus longus were low (<0.05), corresponding to the region of the force-velocity relationship where the muscles were capable of producing 90% of peak isometric force but only 35% of peak isotonic power. V/V(max) increased in response to the demand for mechanical power with increases in running incline and decreased to negative values to absorb energy during downhill running. Measurements of integrated electromyograph activity indicated that the volume of muscle required to produce a given force increased from level to uphill running. This observation is consistent with the idea that V/V(max) is an important determinant of locomotor cost because it affects the volume of muscle that must be recruited to support body weight.

Muscle produces force by forming cross-bridges, using energy released from ATP. While the magnitude and duration of force production primarily determine the energy requirement, nearly a century ago Fenn observed that muscle shortening or lengthening influenced energetic cost of contraction. When work is done by the muscle, the energy cost is increased and when work is done on the muscle the energy cost is reduced. However, the magnitude of the 'Fenn effect' and its mirror ('negative Fenn effect') have not been quantitatively resolved. We describe a new technique coupling magnetic resonance spectroscopy with an in vivo force clamp that can directly quantify the Fenn effect [E=I+W, energy liberated (E) equals the energy cost of isometric force production (I ) plus the work done (W )] and the negative Fenn effect (E=I−W ) for one muscle, the first dorsal interosseous (FDI). ATP cost was measured during a series of contractions, each of which occurred at a constant force and for a constant duration, thus constant force-time integral (FTI). In all subjects, as the FTI increased with load, there was a proportional linear increase in energy cost. In addition, the cost of producing force greatly increased when the muscle shortened, and was slightly reduced during lengthening contraction. These results, though limited to a single muscle, contraction velocity and muscle length change, do quantitatively support the Fenn effect. We speculate that they also suggest that an elastic element within the FDI muscle functions to preserve the force generated within the cross-bridges.

Ca2+ movements during intercall intervals in superfast toadfish swimbladder muscle fibers depend on the accumulation of Ca2+ on parvalbumin and a consequent slow rate of Ca2+ pumping.

SummaryCan human muscle be highly efficient in vivo? Animal muscles typically show contraction-coupling efficiencies <50% in vitro but a recent study reports that the human first dorsal interosseous (FDI) muscle of the hand has an efficiency value in vivo of 68%. We examine two key factors that could account for this apparently high efficiency value: (1) transfer of cross-bridge work into mechanical work and (2) the use of elastic energy to do external work. Our analysis supports a high contractile efficiency reflective of nearly complete transfer of muscular to mechanical work with no contribution by recycling of elastic energy to mechanical work. Our survey of reported contraction-coupling efficiency values puts the FDI value higher than typical values found in small animals in vitro but within the range of values for human muscle in vivo. These high efficiency values support recent studies that suggest lower Ca 2+ cycling costs in working contractions and a decline in cost during repeated contractions. In the end, our analysis indicates that the FDI muscle may be exceptional in having an efficiency value on the higher end of that reported for human muscle. Thus, the FDI muscle may be an exception both in contraction-coupling efficiency and in Ca 2+ cycling costs, which makes it an ideal muscle model system offering prime conditions for studying the energetics of muscle contraction in vivo.