The purpose of this study was to choose between two popular models of skeletal muscle: one with the parallel elastic component in parallel with both the contractile element and the series elastic component (model A), and the other in which it is in parallel with only the contractile element (model B). Passive and total forces were obtained at a variety of muscle lengths for the medial gastrocnemius muscle in anesthetized rats. Passive force was measured before the contraction (passive A) or was estimated for the fascicle length at which peak total force occurred (passive B). Fascicle length was measured with sonomicrometry. Active force was calculated by subtracting passive (A or B) force from peak total force at each fascicle or muscle length. Optimal length, that fascicle length at which active force is maximized, was 13.1 +/- 1.2 mm when passive A was subtracted and 14.0 +/- 1.1 mm with passive B (P < 0.01). Furthermore, the relationship between double-pulse contraction force and length was broader when calculated with passive B than with passive A. When the muscle was held at a long length, passive force decreased due to stress relaxation. This was accompanied by no change in fascicle length at the peak of the contraction and only a small corresponding decrease in peak total force. There is no explanation for the apparent increase in active force that would be obtained when subtracting passive A from the peak total force. Therefore, to calculate active force, it is appropriate to subtract passive force measured at the fascicle length corresponding to the length at which peak total force occurs, rather than passive force measured at the length at which the contraction begins.
Relative force depression associated with muscle fatigue is reported to be greater when assessed at short vs. long muscle lengths. This appears to be due to a rightward shift in the force-length relationship. This rightward shift may be caused by stretch of in-series structures, making sarcomere lengths shorter at any given muscle length. Submaximal force-length relationships (twitch, double pulse, 50 Hz) were evaluated before and after repetitive contractions (50 Hz, 300 ms, 1/s) in an in situ preparation of the rat medial gastrocnemius muscle. In some experiments, fascicle lengths were measured with sonomicrometry. Before repetitive stimulation, fascicle lengths were 11.3 +/- 0.8, 12.8 +/- 0.9, and 14.4 +/- 1.2 mm at lengths corresponding to -3.6, 0, and 3.6 mm where 0 is a reference length that corresponds with maximal active force for double-pulse stimulation. After repetitive stimulation, there was no change in fascicle lengths; these lengths were 11.4 +/- 0.8, 12.6 +/- 0.9, and 14.2 +/- 1.2 mm. The length dependence of fatigue was, therefore, not due to a stretch of in-series structures. Interestingly, the rightward shift that was evident when active force was calculated in the traditional way (subtraction of the passive force measured before contraction) was not seen when active force was calculated by subtracting the passive force that was associated with the fascicle length reached at the peak of the contraction. This calculation is based on the assumption that passive force decreases as the fascicles shorten during a fixed-end contraction. This alternative calculation revealed similar postfatigue absolute active force depression at all lengths. In relative terms, a length dependence of fatigue was still evident, but this was greatly diminished compared with that observed when active force was calculated with the traditional method.
Inhibition of Ca(2+) release alters summation, but repetitive stimulation leading to fatigue changes it more substantially.
The magnitude of fatigue resulting from repeated contractions at a short length has been reported to be less than that which occurs with contractions at a long length. However, there have been what appear to be contradictory reports; the rate of fatigue is greater at a short length. The purpose of this study was to evaluate the impact of length on the magnitude and the rate of fatigue resulting from a series of repetitive stimulations. Experiments were done with anesthetized rats and the medial gastrocnemius muscle was stimulated via the sciatic nerve. Submaximal force-length relationships were obtained prior to and 45 min after repeated contractions (50 Hz, 300 ms) at short or long length. Stimulation was applied at 1 Hz or 0.5 Hz for 5 min at the long length or 1 Hz at the short length (difference = 3.6mm). This approach permitted evaluation of the impact of rate of muscle activation as well as length on subsequent contractile response. Repetitive stimulation at a short length resulted in more potentiation and a greater (relative) rate of fatigue but after 5 min the depression of relative active force was similar between the series at long and short length. The submaximal force-length relationship obtained after 45 min of recovery revealed that depression of force was greater after 1 Hz contractions at the long length. These results are consistent with both sides of the apparent contradiction in the literature; rate of fatigue is greater at a short length and magnitude of fatigue is greater at a long length.
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