Measured and modeled properties of mammalian skeletal muscle: III. the effects of stimulus frequency on stretch-induced force enhancement and shortening-induced force depression

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Abstract
Stretch-induced force enhancement and shortening-induced force depression were examined in fast-twitch feline caudofemoralis muscle at 37∘C. These phenomena were induced by applying ramp length changes during the first 100–200 ms of an otherwise isometric contraction. The effects of various stimulus frequencies ranging from 30 to 120 pps were investigated over lengths ranging from 0.85 to 1.15 L0. Distributed asynchronous stimulation of bundles of ventral roots was employed to produce smooth contractions at sub-tetanic stimulus frequencies in whole muscle. Of the two components of force enhancement identified by Noble (1992) we observed only the transient component that decays with time; we did not observe residual force enhancement. The force depression that we observed was symmetrical in almost all respects to the transient force enhancement, and was unlike the shortening-induced de-activation and residual force depression identified by Edman (Edman, 1975; Edman et al., 1993). Both transient force enhancement and depression were independent of work, load and activation. Reversals in the direction of ramp length changes following either an initial stretch or initial shortening were shown to cancel the effects of both transient force enhancement and transient force depression. The distances over which these cancellations could be achieved were different for the lengthening and shortening effects. This asymmetry can be reconciled with the predictions of Huxley's original cross-bridge mechanism by incorporating the recent suggestion that myosin heads can interact with multiple actin binding sites during a single ‘working’ stroke. We conclude that the types of force enhancement/depression that are most likely to be encountered under physiological conditions are the transient effects observed here, but that even these will have relatively little effect on force production during most natural behaviors.