Compliant tendons permit mechanically unfavourable fascicle dynamics during fixed-end contractions. The purpose of this study was to reduce the effective compliance of tendon and investigate how small reductions in active shortening affect twitch kinetics and contractile performance in response to a second stimulus. The series elastic element (SEE) of the human triceps surae (N=15) was effectively stiffened by applying a 55 ms rotation to the ankle, through a range of 5 deg, at the onset of twitch and doublet [interstimulus interval (ISI) of 80 ms] stimulation. Ultrasonography was employed to quantify lateral gastrocnemius and soleus fascicle lengths. Rotation increased twitch torque (40-75%), rate of torque development (RTD; 124-154%) and torque-time integral (TTI; 70-110%) relative to constant-length contractions at the initial and final joint positions, yet caused only modest reductions in shortening amplitude and velocity. The torque contribution of the second pulse increased when stimulation was preceded by rotation, a finding unable to be explained on the basis of fascicle length or SEE stiffness during contraction post-rotation. A further increase in torque contribution was not demonstrated, nor was an increase in doublet TTI, when the second pulse was delivered during rotation and shortly after the initial pulse (ISI of 10 ms). The depressant effect of active shortening on subsequent torque generation suggests that compliant tendons, by affording large length changes, may limit torque summation. Our findings indicate that changes in tendon compliance shown to occur in response to resistance training or unloading are likely sufficient to considerably alter contractile performance, particularly maximal RTD.
Lateral connections between muscles provide pathways for myofascial force transmission. To elucidate whether these pathways have functional roles in vivo, we examined whether activation could alter the shear between the soleus (SOL) and lateral gastrocnemius (LG) muscles. We hypothesized that selective activation of LG would decrease the stretch-induced shear between LG and SOL. Eleven volunteers underwent a series of knee joint manipulations where plantar flexion force, LG, and SOL muscle fascicle lengths and relative displacement of aponeuroses between the muscles were obtained. Data during a passive full range of motion were recorded, followed by 20° knee extension stretches in both passive conditions and with selective electrical stimulation of LG. During active stretch, plantar flexion force was 22% greater (P < 0.05) and relative displacement of aponeuroses was smaller than during passive stretch (P < 0.05). Soleus fascicle length changes did not differ between passive and active stretches but LG fascicles stretched less in the active than passive condition when the stretch began at angles of 70° and 90° of knee flexion (P < 0.05). The activity-induced decrease in the relative displacement of SOL and LG suggests stronger (stiffer) connectivity between the two muscles, at least at flexed knee joint angles, which may serve to facilitate myofascial force transmission.
There are high mechanical demands placed on skeletal muscles in movements requiring rapid acceleration of the body or its limbs. Tendons are responsible for transmitting muscle forces, but, because of their elasticity, can manipulate the mechanics of the internal contractile apparatus. Shortening of the contractile apparatus against the stretch of tendon affects force generation according to known mechanical properties; however, the extent to which differences in tendon compliance alter force development in response to a burst of electrical impulses is unclear. To establish the influence of series compliance on force summation, we studied electrically evoked doublet contractions in the cane toad peroneus muscle in the presence and absence of a compliant artificial tendon. Additional series compliance reduced tetanic force by two-thirds, a finding predicted based on the force-length property of skeletal muscle. Doublet force and force-time integral expressed relative to the twitch were also reduced by additional series compliance. Active shortening over a larger range of the ascending limb of the force-length curve and at a higher velocity, leading to a progressive reduction in forcegenerating potential, could be responsible. Muscle-tendon interaction may also explain the accelerated time course of force relaxation in the presence of additional compliance. Our findings suggest that a compliant tendon limits force summation under constant-length conditions. However, high series compliance can be mechanically advantageous when a muscle-tendon unit is actively stretched, permitting muscle fibres to generate force almost isometrically, as shown during stretch-shorten cycles in locomotor activities. Restricting active shortening would likely favour rapid force development.
Activation of skeletal muscle twice in quick succession results in nonlinear force summation (i.e., doublet potentiation). The force contributed by a second activation is typically of augmented amplitude, longer in duration, and generated at a greater rate. The purpose of this study was to examine force summation in a muscle attached to a compliant tendon, where considerable internal shortening occurs during a fixed-end contraction. The triceps surae of 21 (Experiment 1) and 9 (Experiment 2) young adults were maximally activated with doublet stimulation of different interstimulus intervals (ISIs) (5-100 ms) at several muscle lengths. Ultrasound images acquired from lateral gastrocnemius and soleus muscles allowed quantification of dynamic fascicle behavior. Force summation was muscle length dependent. Force augmentation was limited to a short muscle length. Lateral gastrocnemius and soleus fascicles underwent large amounts of active shortening and achieved high velocities in response to doublet stimulation, dynamics unfavorable for force production. Summation amplitude and the sensitivity of summation to ISI were dramatically depressed in the triceps surae after comparison to muscles with less fixed-end compliance. We propose that the internal shortening permitted by high series compliance limited force augmentation by offsetting and/or interfering with activation and cross-bridge processes driving augmentation. High series compliance may also reduce the sensitivity of the summated response to ISI, an assertion supported by predictions from a Hill-type muscle model. These muscles may exhibit greater force augmentation during more accustomed stretch-shorten tasks (i.e., hopping), where the compliance of the Achilles tendon actually enables near-isometric fascicle behavior.
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