Biaxial strain and variable stiffness in aponeuroses

The elastic structures of many muscles include both an extramuscular free tendon as well as a sheet-like aponeurosis. An important distinguishing feature of aponeuroses is that these tendinous structures function as the attachment and insertion surfaces of muscle fascicles and therefore surround a substantial portion of the muscle belly. As a result, aponeuroses must expand both parallel (longitudinal) and perpendicular (transverse) to a muscle's line of action when contracting muscles bulge to maintain a constant volume. In this study, we use biplanar high-speed fluoroscopy to track the strain patterns of the turkey lateral gastrocnemius aponeurosis during active and passive force production in situ. We find that the behaviour of the aponeurosis during passive force production is consistent with uniaxial loading, as aponeuroses stretch only in the longitudinal direction. By contrast, our results show that aponeuroses are stretched in both longitudinal and transverse directions during active force production and that transverse strains are on average 4 times greater than longitudinal strains. Biaxial loading of aponeuroses appears to effectively modulate longitudinal stiffness, as we find the measured stiffness in the longitudinal direction varies in proportion to transverse strain. We conclude that biaxial strain during active force production distinguishes aponeuroses from free tendons and may function to dynamically modulate stiffness along the axis of muscle force production. It is likely that consideration of strains measured only in the longitudinal direction result in an underestimation of aponeurosis stiffness as well as its capacity for elastic energy storage.