The history-dependent features of muscle force production: A challenge to the cross-bridge theory and their functional implications.

The cross-bridge theory predicts that muscle force is determined by muscle length and the velocity of active muscle length changes. However, before the formulation of the cross-bridge theory, it had been observed that the isometric force at a given muscle length is enhanced or depressed depending on active muscle length changes before that given length is reached. These enhanced and depressed force states are termed residual force enhancement (rFE) and residual force depression (rFD), respectively, and together they are known as the history-dependent features of muscle force production. In this review, we introduce early attempts in explaining rFE and rFD before we discuss more recent research from the past 25 years which has contributed to a better understanding of the mechanisms underpinning rFE and rFD. Specifically, we discuss the increasing number of findings on rFE and rFD which challenge the cross-bridge theory and propose that the elastic element titin plays a role in explaining muscle history-dependence. Accordingly, new three-filament models of force production including titin seem to provide better insight into the mechanism of muscle contraction. Complementary to the mechanisms behind muscle history-dependence, we also show various implications for muscle history-dependence on in-vivo human muscle function such as during stretch-shortening cycles. We conclude that titin function needs to be better understood if a new three-filament muscle model which includes titin, is to be established. From an applied perspective, it remains to be elucidated how muscle history-dependence affects locomotion and motor control, and whether history-dependent features can be changed by training.