Determinants of aponeurosis shape change during muscle contraction

Aponeuroses are sheet-like elastic tendon structures that cover a portion of the muscle belly and act as insertion sites for muscle fibers while free tendons connect muscles to bones. During shortening contractions, free tendons are loaded in tension and lengthen due to the force acting longitudinally along the muscle׳s line of action. In contrast, aponeuroses increase in length and width, suggesting that aponeuroses are loaded in directions along and orthogonal to the muscle׳s line of action. Because muscle fibers are isovolumetric, they must expand radially as they shorten, potentially generating a force that increases aponeurosis width. We hypothesized that increases in aponeurosis width result from radial expansion of shortening muscle fibers. We tested this hypothesis by combining in situ muscle-tendon measurements with high-speed biplanar fluoroscopy measurements of the turkey׳s lateral gastrocnemius (n=6) at varying levels of isotonic muscle contractions. The change in aponeurosis width during periods of constant force depended on both the amount of muscle shortening and the magnitude of force production. At low to intermediate forces, aponeurosis width increased in direct proportion to fiber shortening. At high forces, aponeurosis width increased to a lesser extent or in some cases, decreased slightly during fiber shortening. Our results demonstrate that forces generated from radial expansion of shortening muscle fibers tend to drive increases in aponeurosis width, whereas longitudinal forces tend to decrease aponeurosis width. Ultimately, it is these two opposing forces that drive changes in aponeurosis width and alter series elastic stiffness during a muscle contraction.