Components of activation heat in skeletal muscle

Activation heat (q_A) production by muscle is the thermal accompaniment of the release of Ca²⁺ from the sarcoplasmic reticulum (SR) into the cytoplasm, its interactions with regulatory proteins and other cytoplasmic Ca²⁺ buffers and its return to the SR. The contribution of different Ca²⁺-related reactions to q_A is difficult to determine empirically and therefore, for this study, a mathematical model was developed to describe Ca²⁺ movements and accompanying thermal changes in muscle fibres in response to stimulation. The major sources of heat within a few milliseconds of the initiation of Ca²⁺ release are Ca²⁺ binding to Tn and Pv. Ca²⁺ binding to ATP produces a relatively small amount of heat. Ca²⁺ dissociation from ATP and Tn, with heat absorption, are of similar time course to the decline of force. In muscle lacking Pv (e.g. mouse soleus), Ca²⁺ is then rapidly pumped into the SR. In muscles with Pv, Ca²⁺ that dissociates from Tn and ATP binds to Pv and then dissociates slowly (over 10 s of seconds) and is then pumped into the SR; the net effect of these two processes is heat absorption. It is proposed that this underlies Hill's "negative delayed heat". After all the Ca²⁺ is returned to the SR, q_A is proportional to the amount of Ca²⁺ released into the cytoplasm. In muscles with Pv this is 20-60 s after Ca²⁺ release; in muscles without Pv, all Ca²⁺ is returned to the SR soon after the end of force relaxation.