How muscles deal with real-world loads: the influence of length trajectory on muscle performance.

The performance of skeletal muscles in vivo is determined by the feedback received when the muscle interacts with the external environment via various morphological structures. This interaction between the muscle and the 'real-world load' forces us to reconsider how muscles are adapted to suit their in vivo function. We must consider the co-evolution of the muscles and the morphological structures that 'create' the load in concert with the properties of the external environment. This complex set of interactions may limit muscle performance acutely and may also constrain the evolution of morphology and physiology. The performance of skeletal muscle is determined by the length trajectory during movement and the pattern of stimulation. Important features of the length trajectory include its amplitude, frequency, starting length and shape (velocity profile). Many of these parameters interact. For example, changing the velocity profile during shortening may change the optimum values of the other parameters. The length trajectory that maximizes performance depends on the task to be performed. During cyclical work, muscles benefit from using asymmetric cycles with longer shortening than lengthening phases. Modifying this 'sawtooth' cycle by increasing the velocity during shortening may further increase power by augmenting force output and speeding deactivation. In contrast, when accelerating an inertial load, as in jumping, the predicted 'optimal' velocity profile has two peak values, one early and one late in shortening. During level running at constant speed, muscles perform tasks other than producing work and power. Producing force to support the body weight is performed with nearly isometric contractions in some of the limb muscles of vertebrates. Muscles also play a key role in producing stability during running, and the intrinsic properties of the musculoskeletal system may be particularly important in stabilizing rapid running. Recently, muscles in running invertebrates and vertebrates have been described that routinely absorb large amounts of work during running. These muscles are hypothesized to play a key role in stability.