Transmission of forces within mammalian skeletal muscles

Most models of in vivo musculoskeletal function fail to take into account the diversity of force trajectories defined by muscle fiber architecture. It has been shown for many muscles, across species, that muscle fibers commonly end within muscle fascicles without reaching a myotendinous junction, and that many of these fibers show a progressive decline in cross-sectional area along the length of the muscle. The significance of these anatomical observations is that the tapering would seem to preclude forces generated at the largest cross-sectional area of the fibers being transmitted to the sarcomeres toward the ends of the tapered fiber. If all of the forces are transmitted via the sarcomeres arranged in series, those few sarcomeres at the smaller ends of the fibers must tolerate the stress exerted by the more numerous sarcomeres arranged in parallel at the portions of the fiber with larger cross-sectional areas. A logical alternative would be for forces to be transmitted laterally along the length of a fiber to the cell membrane and the extracellular matrix. Such a structural arrangement would permit an alternative force transmission vector and minimize the necessity for a precise level of force to be generated along the entire length of a fiber. There are cytoarchitectural and biochemical data demonstrating the presence of a subcellular network which is appropriately located to transmit forces from the active intracellular contractile elements to the extracellular intramuscular connective tissues. However, to fully comprehend how forces are transmitted from individual cross bridges to the tendon, it will be necessary to understand the interactions of all of the components of the muscle–tendon complex from the molecular to the multicellular level. It is insufficient to know the physiology of the individual components in a restricted experimental paradigm and assume that these conditions account for the functional characteristics in vivo. Thus, the challenge is to understand how the sarcomeres and all of the associated structures transmit the forces of the whole muscle to its attachments.