Changes in the maximum speed of shortening of frog muscle fibres early in a tetanic contraction and during relaxation

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Abstract
1. Isotonic shortening velocities at very light loads were examined in single fibres of the anterior tibialis muscle of the frog, Rana temporaria, using load-clamp recording and slack tests (temperature, 1-3 degrees C; initial sarcomere length, 2.25 microns). 2. Shortening velocities at very light loads (force-clamp recording) were found to be higher early in the rise of a tetanic contraction than during the plateau of the contraction. The upper limit of the load at which there was elevated shortening velocity early in the contraction was 1.5-5.4% of the maximum tetanic tension (Fo) depending on the particular fibre. 3. The maximum shortening velocity determined using the slack test method (Vo) was as much as 30% greater early in a contraction than at the tetanic plateau. Vo was elevated above the plateau level up to about 30 ms after the end of the latent period, which is equivalent to the time required for the force in an isometric contraction to rise to about 30% of Fo. Vo is depressed below the plateau value during relaxation at the cessation of stimulation. 4. Stimulation studies show that the cross-bridge model of Huxley (1957) predicts the maximum shortening velocity to be greater early in a contraction, when new actin binding sites are becoming activated and new cross-bridge connections are being formed rapidly, than during steady-state contraction. The elevated shortening velocity in the model is a consequence of new cross-bridges being formed in the pulling configuration, and there being a delay before the newly added bridges are dragged beyond their equilibrium position so they begin to retard shortening. The model also predicts that maximum shortening velocity should be depressed below the plateau level during early relaxation as cross-bridge binding sites are rapidly removed from the active population.