An analysis of the temperature dependence of force, during steady shortening at different velocities, in (mammalian) fast muscle fibres.

We examined, over a wide range of temperatures (10-35 degrees C), the isometric tension and tension during ramp shortening at different velocities (0.2-4 L(0)/s) in tetanized intact fibre bundles from a rat fast (flexor hallucis brevis) muscle; fibre length (L(0)) was 2.2 mm and sarcomere length approximately 2.5 microm. During a ramp shortening, the tension change showed an initial inflection of small amplitude (P(1)), followed by a larger exponential decline towards an approximate steady level; the tension continued to decline slowly afterwards and the approximate steady tension at a given velocity was estimated as the tension (P(2)) at the point of intersection between two linear slopes, as previously described (Roots et al. 2007). At a given temperature, the tension P(2) declined to a lower level and at a faster rate (from an exponential curve fit) as the shortening velocity was increased; the temperature sensitivity of the rate of tension decline during ramp shortening at different velocities was low (Q(10) 0.9-1.5). The isometric tension and the P(2) tension at a given shortening velocity increased with warming so that the relation between tension and (reciprocal) temperature was sigmoidal in both. In isometric muscle, the temperature T(0.5) for half-maximal tension was approximately 10 degrees C, activation enthalpy change (DeltaH) was approximately 100 kJ mol(-1) and entropy change (DeltaS) approximately 350 J mol(-1) K(-1). In shortening, these were increased with increase of velocity so that at a shortening velocity (approximately 4 L(0)/s) producing maximal power at 35 degrees C, T(0.5) was approximately 28 degrees C, DeltaH was approximately 200 kJ mol(-1) and DeltaS approximately 700 J mol(-1) K(-1); the same trends were seen in the tension data from isotonic release experiments on intact muscle and in ramp shortening experiments on maximally Ca-activated skinned fibres. In general, our findings show that the sigmoidal relation between force and temperature can be extended from isometric to shortening muscle; the implications of the findings are discussed in relation to the crossbridge cycle. The data indicate that the endothermic, entropy driven process that underlies crossbridge force generation in isometric muscle (Zhao and Kawai 1994; Davis, 1998) is even more pronounced in shortening muscle, i.e. when doing external work.