Adaptation of muscle size and myofascial force transmission: a review and some new experimental results

This paper considers the literature and some new experimental results important for adaptation of muscle fiber cross-sectional area and serial sarcomere number. Two major points emerge: (1) general rules for the regulation of adaptation (for in vivo immobilization, low gravity conditions, synergist ablation, tenotomy and retinaculum trans-section experiments) cannot be derived. As a consequence, paradoxes are reported in the literature. Some paradoxes are resolved by considering the interaction between different levels of organization (e.g. muscle geometrical effects), but others cannot. (2) An inventory of signal transduction pathways affecting rates of muscle protein synthesis and/or degradation reveals controversy concerning the pathways and their relative contributions. A major explanation for the above is not only the inherently limited control of the experimental conditions in vivo, but also of in situ experiments. Culturing of mature single Xenopus muscle fibers at high and low lengths (allowing longitudinal study of adaptation for periods up to 3 months) did not yield major changes in the fiber cross-sectional area or the serial sarcomere number. This is very different from substantial effects (within days) of immobilization in vivo. It is concluded that overall strain does not uniquely regulate muscle fiber size. Force transmission, via pathways other than the myotendinous junctions, may contribute to the discrepancies reported: because of substantial serial heterogeneity of sarcomere lengths within muscle fibers creating local variations in the mechanical stimuli for adaptation. For the single muscle fiber, mechanical signalling is quite different from the in vivo or in vitro condition. Removal of tensile and shear effects of neighboring tissues (even of antagonistic muscle) modifies or removes mechanical stimuli for adaptation. It is concluded that the study of adaptation of muscle size requires an integrative approach taking into account fundamental mechanisms of adaptation, as well as effects of higher levels of organization. More attention should be paid to adaptation of connective tissues within and surrounding the muscle and their effects on muscular properties.