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.

Electromyography is often used to infer the pattern of production of force by skeletal muscles. The interpretation of muscle function from the electromyogram (EMG) is challenged by the fact that factors such as type of muscle fiber, muscle length, and muscle velocity can all influence the relationship between electrical and mechanical activity of a muscle. Simultaneous measurements of EMG, muscle force, and fascicle length in hindlimb muscles of wild turkeys allow us to probe the quantitative link between force and EMG. We examined two features of the force-EMG relationship. First, we measured the relaxation electromechanical delay (r-EMD) as the time from the end of the EMG signal to time of the end of force. This delay varied with locomotor speed in the lateral gastrocnemius (LG); it was longer at slow walking speeds than for running. This variation in r-EMD was not explained by differences in muscle length trajectory, as the magnitude of r-EMD was not correlated with the velocity of shortening of the muscle during relaxation. We speculate that the longer relaxation times at slow walking speeds compared with running may reflect the longer time course of relaxation in slower muscles fibers. We also examined the relationship between magnitude of force and EMG across a range of walking and running speeds. We analyzed the force-EMG relationship during the swing phase separately from the force-EMG relationship during stance phase. During stance, force amplitude (average force) was linearly related to mean EMG amplitude (average EMG). Forces during swing phase were lower than predicted from the stance phase force-EMG relationship. The different force-EMG relationships during the stance and swing phases may reflect the contribution of passive structures to the development of force, or a nonlinear force-EMG relationship at low levels of muscle activity. Together the results suggest that any inference of force from EMG must be done cautiously when a broad range of activities is considered.

No abstract