Muscle, the motor of movement: properties in function, experiment and modelling

The purpose of this paper is to review exemplary aspects of different views of skeletal muscle characteristics. A classical view of muscle characteristics plays a very important role in modelling of muscles and movement. However, it often also pervades concepts on which our understanding of muscle function is based. In this view length effects, velocity effects and effects of degrees of activation and recruitment are distinguished and, often implicitly, assumed to be independent effects. It will be illustrated that using the classical approach many valuable things may be learned about muscle function and adaptation. At the same time we should realize that such a classical approach is too limited for use in generating knowledge about properties of muscles during daily use. The use of scaling of force to estimate muscular properties during submaximal activity on the basis of properties during maximal activation is shown to be very inadequate. An alternative view is described and particular examples are provided of changes in length-force characteristics as a consequence of submaximal activation, previous length change, as well as the effect of short-term histories of these variables. In addition, effects of inhomogeneities of muscle in morphology as well as physiological properties are considered. It is concluded that length-velocity-force characteristics are not unique properties of a muscle, and that these characteristics are not only strongly influenced by actual effects of recruitment, firing frequency, shortening performed and actual velocity of shortening but also by the short time history of these factors. Therefore, length, velocity and activation cannot be considered as independent determinants of muscle functioning. It is also shown that we are confronted with many indications of physiological individuality regarding these phenomena.