Strain of passive elements during force enhancement by stretch in frog muscle fibres

1. The force enhancement during and after stretch (0.15 micron per sarcomere) was studied during fused tetani of single fibres isolated from the anterior tibialis muscle of Rana temporaria (0.5-3.6 degrees C; sarcomere length, 2.05-2.65 microns). Changes in length were recorded simultaneously from the fibre as a whole (puller movement) and from marked segments (approximately 0.5 mm in length) of the same fibre. 2. The residual force enhancement after stretch (recorded at the end of a long tetanus) was found to be linearly related to the slow component of tension rise during the stretch ramp. 3. The fibres were released to shorten against a very small load at different times after stretch (load clamp). The shortening records derived after a preceding stretch exhibited a larger and steeper initial transient than that recorded in an isometric tetanus without stretch. The excess length change (LS; nanometres per half-sarcomere) recorded during the initial transient increased with the amplitude of stretch and was linearly related to the force enhancement produced by the stretch (FE; % of maximum tetanic tension) according to the following regression: LS = 0.200 FE + 8.65 (P < 0.001). The length changes recorded from the whole fibre agreed well with measurements from individual segments. 4. Slack-test measurements confirmed the existence of a large initial transient phase when the fibre was released to shorten after a preceding stretch. The excess length change determined from the slack tests agreed closely with the values derived from load-clamp recordings. 5. The results support the view that stretching a muscle fibre during tetanus leads to strain of elastic elements and, presumably, to variation of filament overlap due to non-uniform distribution of the length change within the fibre volume. Regions with greater filament overlap are likely to generate the long-lasting extra force referred to as 'residual force enhancement after stretch'. The elastic elements recruited during stretch can be presumed to play an essential part in this process by supporting regions in which the filament overlap has been reduced during the stretch ramp. Recoil of these elastic elements is responsible for the excess length change that is recorded during the initial transient after release as described under point 3.