Muscle stiffness in human ankle dorsiflexors: intrinsic and reflex components

1. The purpose of this study was to evaluate the mechanical response to stretch in normal human ankle dorsiflexors at different levels of voluntary contraction. In an active muscle, the total mechanical response is the sum of the intrinsic response from the contractile apparatus, the response from passive tissues, and the reflex mediated response. Each of these components was investigated. 2. The total incremental stiffness was defined as the ratio between the torque increment and the amplitude of the stretch. In 14 subjects the total stiffness increased from approximately 0.6 N.m/deg to approximately 2.5 N.m/deg at 50% of MVC and remained constant (+/- 10%) from 30 to 80% of MVC. 3. The contribution to incremental stiffness from intrinsic muscle properties was measured during electrical stimulation of the deep peroneal nerve at 7-50 Hz. Intrinsic stiffness increased linearly with torque from approximately 0.5 N.m/deg to approximately 2.5 N.m/deg at 80% of MVC. 4. The reflex component (total minus intrinsic stiffness) had a maximum of 0.5-1.5 N.m/deg at 30-50% of MVC and was approximately zero at no and maximal contraction. For intermediate levels of contraction the reflex increased the stiffness with 40-100% of the intrinsic stiffness in this flexor muscle. 5. The reflex contribution to total stiffness began approximately 50 ms after onset of stretch and peaked 150-300 ms after onset of stretch. 6. Total, intrinsic, and reflex mediated stiffness were all nearly independent of the amplitude of stretch in the range from 2 to 7 degrees. The higher stiffness observed for 1 degree stretches could be due to "short range stiffness" of the cross bridges. 7. Stretching of a contracting muscle generates large force increments even for moderate amplitudes of stretch. Approximately half of this force increment is due to the stretch reflex, which makes the muscle stiffer than predicted from the intrinsic stiffness. These findings in human flexor muscles are surprisingly similar to previous findings in extensor muscles of the decerebrate cat.