Enthalpy efficiency of the soleus muscle contributes to improvements in running economy.
ABSTRACT: During human running, the soleus, as the main plantar flexor muscle, generates the majority of the mechanical work through active shortening. The fraction of chemical energy that is converted into muscular work (enthalpy efficiency) depends on the muscle shortening velocity. Here, we investigated the soleus muscle fascicle behaviour during running with respect to the enthalpy efficiency as a mechanism that could contribute to improvements in running economy after exercise-induced increases of plantar flexor strength and Achilles tendon (AT) stiffness. Using a controlled longitudinal study design (n = 23) featuring a specific 14-week muscle-tendon training, increases in muscle strength (10%) and tendon stiffness (31%) and reduced metabolic cost of running (4%) were found only in the intervention group (n = 13, p < 0.05). Following training, the soleus fascicles operated at higher enthalpy efficiency during the phase of muscle-tendon unit (MTU) lengthening (15%) and in average over stance (7%, p < 0.05). Thus, improvements in energetic cost following increases in plantar flexor strength and AT stiffness seem attributed to increased enthalpy efficiency of the operating soleus muscle. The results further imply that the soleus energy production in the first part of stance, when the MTU is lengthening, may be crucial for the overall metabolic energy cost of running.
Project description:Running is thought to be an efficient gait due, in part, to the behavior of the plantar flexor muscles and elastic energy storage in the Achilles tendon. Although plantar flexor muscle mechanics and Achilles tendon energy storage have been explored during rearfoot striking, they have not been fully characterized during forefoot striking. This study examined how plantar flexor muscle-tendon mechanics during running differs between rearfoot and forefoot striking. We used musculoskeletal simulations, driven by joint angles and electromyography recorded from runners using both rearfoot and forefoot striking running patterns, to characterize plantar flexor muscle-tendon mechanics. The simulations revealed that foot strike pattern affected the soleus and gastrocnemius differently. For the soleus, forefoot striking decreased tendon energy storage and fiber work done while the muscle fibers were shortening compared to rearfoot striking. For the gastrocnemius, forefoot striking increased muscle activation and fiber work done while the muscle fibers were lengthening compared to rearfoot striking. These changes in gastrocnemius mechanics suggest that runners planning to convert to forefoot striking might benefit from a progressive eccentric gastrocnemius strengthening program to avoid injury.
Project description:The human ankle joint and plantar flexor muscle-tendon unit play an important role in endurance running. It has been assumed that muscle and tendon interactions and their biomechanical behaviours depend on their morphological and architectural characteristics. We aimed to study how plantar flexor muscle characteristics influence marathon running performance and to determine whether there is any difference in the role of the soleus and gastrocnemii. The right lower leg of ten male distance runners was scanned with magnetic resonance imagining. The cross-sectional areas of the Achilles tendon, soleus, and lateral and medial gastrocnemius were measured, and the muscle volumes were calculated. Additional ultrasound scanning was used to estimate the fascicle length of each muscle to calculate the physiological cross-sectional area. Correlations were found between marathon running performance and soleus volume (r?=?0.55, p?=?0.048), soleus cross-sectional area (r?=?0.57, p?=?0.04), soleus physiological cross-sectional area (PCSA-IAAF r?=?0.77, p?<?0.01, CI± 0.28 to 0.94), Achilles tendon thickness (r?=?0.65, p?<?0.01), and soleus muscle-to-tendon ratio (r?=?0.68, p?=?0.03). None of the gastrocnemius characteristics were associated with marathon performance. We concluded that a larger soleus muscle with a thicker Achilles tendon is associated with better marathon performance. Based on these results, it can be concluded the morphological characteristics of the lower leg muscle-tendon unit correlate with running performance.
Project description:Many studies have shown that connective tissue linkages can transmit force between synergistic muscles and that such force transmission depends on the position of these muscles relative to each other and on properties of their intermuscular connective tissues. Moving neighboring muscles has been reported to cause longitudinal deformations within passive muscles held at a constant muscle-tendon unit (MTU) length (e.g., soleus [SO]), but muscle forces were not directly measured. Deformations do not provide a direct measure of the force transmitted between muscles. We combined two different muscle preparations to assess whether myofascial loads exerted by neighboring muscles result in length changes of SO fascicles. We investigated the effects of proximal MTU length changes of two-joint gastrocnemius (GA) and plantaris (PL) muscles on the fascicle length of the one-joint SO muscle within (1) an intact muscle compartment and (2) a disrupted compartment that allowed measurements of fascicle length and distal tendon force of SO simultaneously. SO muscle bellies of Wistar rats (n?=?5) were implanted with sonomicrometry crystals. In three animals, connectivity between SO and GA+PL was enhanced. Measurements were performed before and during maximal excitation of all plantar flexor muscles. In both setups, MTU length of GA+PL did not affect the length of SO fascicles, neither during passive nor active conditions. However, lengthening the MTU of GA+PL increased distal tendon force of SO by 43.3-97.8% (P?<?0.001) and 27.5-182.6% (P?<?0.001), respectively. This indicates that substantial myofascial force transmission between SO and synergistic muscle can occur via a connective tissue network running parallel to the series of SO sarcomeres without substantial length changes of SO fascicles.
Project description:The aim of this study was to compare young and adult sprinters on several biomechanical parameters that were previously highlighted as performance-related and to determine the behaviour of several muscle-tendon units (MTU) in the first stance phase following a block start in sprint running. The ground reaction force (GRF) and kinematic data were collected from 16 adult and 21 young well-trained sprinters. No difference between the groups was found in some of the previously highlighted performance-related parameters (ankle joint stiffness, the range of dorsiflexion and plantar flexor moment). Interestingly, the young sprinters showed a greater maximal and mean ratio of horizontal to total GRF, which was mainly attributed to a greater horizontal GRF relative to body mass and resulted in a greater change in horizontal centre of mass (COM) velocity during the stance phase in the young compared with the adult sprinters. Results from the MTU length analyses showed that adult sprinters had more MTU shortening and higher maximal MTU shortening velocities in all plantar flexors and the rectus femoris. Although previously highlighted performance-related parameters could not explain the greater 100?m sprinting times in the adult sprinters, differences were found in the behaviour of the MTU of the plantar flexors and rectus femoris during the first stance phase. The pattern of length changes in these MTUs provides ideal conditions for the use of elastic energy storage and release for power enhancement.
Project description:Previous studies of human locomotion indicate that foot and ankle structures can interact in complex ways. The structure of the foot defines the input and output lever arms that influences the force-generating capacity of the ankle plantar flexors during push-off. At the same time, deformation of the foot may dissipate some of the mechanical energy generated by the plantar flexors during push-off. We investigated this foot-ankle interplay during walking by adding stiffness to the foot through shoes and insoles, and characterized the resulting changes in in vivo soleus muscle-tendon mechanics using ultrasonography. Added stiffness decreased energy dissipation at the foot (p < 0.001) and increased the gear ratio (i.e., ratio of ground reaction force and plantar flexor muscle lever arms) (p < 0.001). Added foot stiffness also altered soleus muscle behaviour, leading to greater peak force (p < 0.001) and reduced fascicle shortening speed (p < 0.001). Despite this shift in force-velocity behaviour, the whole-body metabolic cost during walking increased with added foot stiffness (p < 0.001). This increased metabolic cost is likely due to the added force demand on the plantar flexors, as walking on a more rigid foot/shoe surface compromises the plantar flexors' mechanical advantage.
Project description:During rapid deceleration of the body, tendons buffer part of the elongation of the muscle-tendon unit (MTU), enabling safe energy dissipation via eccentric muscle contraction. Yet, the influence of changes in tendon stiffness within the physiological range upon these lengthening contractions is unknown. This study aimed to examine the effect of training-induced stiffening of the Achilles tendon on triceps surae muscle-tendon behavior during a landing task. Twenty-one male subjects were assigned to either a 10-week resistance-training program consisting of single-leg isometric plantarflexion (<i>n</i> = 11) or to a non-training control group (<i>n</i> = 10). Before and after the training period, plantarflexion force, peak Achilles tendon strain and stiffness were measured during isometric contractions, using a combination of dynamometry, ultrasound and kinematics data. Additionally, testing included a step-landing task, during which joint mechanics and lengths of gastrocnemius and soleus fascicles, Achilles tendon, and MTU were determined using synchronized ultrasound, kinematics and kinetics data collection. After training, plantarflexion strength and Achilles tendon stiffness increased (15 and 18%, respectively), and tendon strain during landing remained similar. Likewise, lengthening and negative work produced by the gastrocnemius MTU did not change detectably. However, in the training group, gastrocnemius fascicle length was offset (8%) to a longer length at touch down and, surprisingly, fascicle lengthening and velocity were reduced by 27 and 21%, respectively. These changes were not observed for soleus fascicles when accounting for variation in task execution between tests. These results indicate that a training-induced increase in tendon stiffness does not noticeably affect the buffering action of the tendon when the MTU is rapidly stretched. Reductions in gastrocnemius fascicle lengthening and lengthening velocity during landing occurred independently from tendon strain. Future studies are required to provide insight into the mechanisms underpinning these observations and their influence on energy dissipation.
Project description:Deficits in ankle muscle strength and ankle stiffness may be present in those subjects who underwent surgical treatment for an Achilles tendon rupture. The presence of these long-term deficits may contribute to a lower performance during daily activities and may be linked to future injuries.To compare the ankle passive stiffness and the plantar flexor muscle performance in patients who underwent unilateral surgical treatment of Achilles tendon rupture with nonsurgical subjects.Twenty patients who underwent unilateral surgical treatment of Achilles tendon rupture [surgical (SU) group], and twenty nonsurgical subjects [non-surgical (NS) group] participated in this study. The ankle passive stiffness was evaluated using a clinical test. The concentric and eccentric plantar flexors performance (i.e. peak torque and work) was evaluated using an isokinetic dynamometer at 30°/s.The surgical ankle of the surgical group presented lower stiffness compared to the non-surgical ankle (mean difference=3.790; 95%CI=1.23-6.35) and to the non-dominant ankle of the non-surgical group (mean difference=-3.860; 95%CI=-7.38 to -0.33). The surgical group had greater absolute asymmetry of ankle stiffness (mean difference=-2.630; 95%CI=-4.61 to -0.65) and greater absolute asymmetry of concentric (mean difference=-8.3%; 95%CI=-13.79 to -2.81) and eccentric (mean difference=-6.9%; 95%CI=-12.1 to -1.7) plantar flexor work compared to non-surgical group. There was no other difference in stiffness and plantar flexor performance.Patients who underwent surgical repair of the Achilles tendon presented with long-term (1 year or more) deficits of ankle stiffness and asymmetries of ankle stiffness and plantar flexor work in the affected ankle compared to the uninjured side in the surgical group and both sides on the nonsurgical group.
Project description:Human running features a spring-like interaction of body and ground, enabled by elastic tendons that store mechanical energy and facilitate muscle operating conditions to minimize the metabolic cost. By experimentally assessing the operating conditions of two important muscles for running, the soleus and vastus lateralis, we investigated physiological mechanisms of muscle work production and muscle force generation. We found that the soleus continuously shortened throughout the stance phase, operating as work generator under conditions that are considered optimal for work production: high force-length potential and high enthalpy efficiency. The vastus lateralis promoted tendon energy storage and contracted nearly isometrically close to optimal length, resulting in a high force-length-velocity potential beneficial for economical force generation. The favorable operating conditions of both muscles were a result of an effective length and velocity-decoupling of fascicles and muscle-tendon unit, mostly due to tendon compliance and, in the soleus, marginally by fascicle rotation.
Project description:Experiments have shown that elastic ankle exoskeletons can be used to reduce ankle joint and plantar-flexor muscle loading when hopping in place and, in turn, reduce metabolic energy consumption. However, recent experimental work has shown that such exoskeletons cause less favourable soleus (SO) muscle-tendon mechanics than is observed during normal hopping, which might limit the capacity of the exoskeleton to reduce energy consumption. To directly link plantar-flexor mechanics and energy consumption when hopping in exoskeletons, we used a musculoskeletal model of the human leg and a model of muscle energetics in simulations of muscle-tendon dynamics during hopping with and without elastic ankle exoskeletons. Simulations were driven by experimental electromyograms, joint kinematics and exoskeleton torque taken from previously published data. The data were from seven males who hopped at 2.5 Hz with and without elastic ankle exoskeletons. The energetics model showed that the total rate of metabolic energy consumption by ankle muscles was not significantly reduced by an ankle exoskeleton. This was despite large reductions in plantar-flexor force production (40-50%). The lack of larger metabolic reductions with exoskeletons was attributed to increases in plantar-flexor muscle fibre velocities and a shift to less favourable muscle fibre lengths during active force production. This limited the capacity for plantar-flexors to reduce activation and energy consumption when hopping with exoskeleton assistance.
Project description:Tendon elastic strain energy is the dominant contributor to muscle-tendon work during steady-state running. Does this behaviour also occur for sprint accelerations? We used experimental data and computational modelling to quantify muscle fascicle work and tendon elastic strain energy for the human ankle plantar flexors (specifically soleus and medial gastrocnemius) for multiple foot contacts of a maximal sprint as well as for running at a steady-state speed. Positive work done by the soleus and medial gastrocnemius muscle fascicles decreased incrementally throughout the maximal sprint and both muscles performed more work for the first foot contact of the maximal sprint (FC1) compared with steady-state running at 5 m s(-1) (SS5). However, the differences in tendon strain energy for both muscles were negligible throughout the maximal sprint and when comparing FC1 to SS5. Consequently, the contribution of muscle fascicle work to stored tendon elastic strain energy was greater for FC1 compared with subsequent foot contacts of the maximal sprint and compared with SS5. We conclude that tendon elastic strain energy in the ankle plantar flexors is just as vital at the start of a maximal sprint as it is at the end, and as it is for running at a constant speed.