Crossbridge properties investigated by fast ramp stretching of activated frog muscle fibres

Very fast ramp stretches at 9.5–33 sarcomere lengths s−1 (l0s−1) stretching speed, 16–25 nm per half-sarcomere (nm hs−1) amplitude were applied to activated intact frog muscle fibres at tetanus plateau, during the tetanus rise, during the isometric phase of relaxation and during isotonic shortening. Stretches produced an almost linear tension increase above the isometric level up to a peak, and fell to a lower value in spite of continued stretching, indicating that the fibre became suddenly very compliant. This suggests that peak tension (critical tension, Pc) represents the tension at which crossbridges are forcibly detached by the stretch. The ratio of Pc to the isometric tension at tetanus plateau (P0) was 2.37 ± 0.12 (s.e.m.). This ratio did not change significantly at lower tension (P) during the tetanus rise but decreased with time during the relaxation and increased with speed during isotonic shortening. At tetanus plateau Pc occurred when sarcomere elongation attained a critical length (Lc) of 10.98 ± 0.13 nm hs−1, independently of the stretching speed. Lc remained constant during the tetanus rise but decreased on the relaxation and increased during isotonic shortening. Length-clamp experiments on the relaxation showed that the lower values of Pc/P ratio and Lc, were both due to the slow sarcomere stretching occurring during this phase. Our data show that Pc can be used as a measure of crossbridge number, while Lc is a measure of crossbridge mean extension. Accordingly, for a given tension, crossbridges on the isometric relaxation are fewer than during the rise, develop a greater individual force and have a greater mean extension, while during isotonic shortening crossbridges are in a greater number but develop a smaller individual force and have a smaller extension.