Effect of Training-Induced Changes in Achilles Tendon Stiffness on Muscle-Tendon Behavior During Landing.
ABSTRACT: 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 (n = 11) or to a non-training control group (n = 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:Background:During the stance phase of running, the elasticity of the Achilles tendon enables the utilisation of elastic energy and allows beneficial contractile conditions for the triceps surae muscles. However, the effect of changes in tendon mechanical properties induced by chronic loading is still poorly understood. We tested the hypothesis that a training-induced increase in Achilles tendon stiffness would result in reduced tendon strain during the stance phase of running, which would reduce fascicle strains in the triceps surae muscles, particularly in the mono-articular soleus. Methods:Eleven subjects were assigned to a training group performing isometric single-leg plantarflexion contractions three times per week for ten weeks, and another ten subjects formed a control group. Before and after the training period, Achilles tendon stiffness was estimated, and muscle-tendon mechanics were assessed during running at preferred speed using ultrasonography, kinematics and kinetics. Results:Achilles tendon stiffness increased by 18% (P < 0.01) in the training group, but the associated reduction in strain seen during isometric contractions was not statistically significant. Tendon elongation during the stance phase of running was similar after training, but tendon recoil was reduced by 30% (P < 0.01), while estimated tendon force remained unchanged. Neither gastrocnemius medialis nor soleus fascicle shortening during stance was affected by training. Discussion:These results show that a training-induced increase in Achilles tendon stiffness altered tendon behaviour during running. Despite training-induced changes in tendon mechanical properties and recoil behaviour, the data suggest that fascicle shortening patterns were preserved for the running speed that we examined. The asymmetrical changes in tendon strain patterns supports the notion that simple in-series models do not fully explain the mechanical output of the muscle-tendon unit during a complex task like running.
Project description: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 (<i>n</i> = 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 (<i>n</i> = 13, <i>p</i> < 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%, <i>p</i> < 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:Aim: Stretching is often used to increase/maintain muscle length and improve joint range of motion (ROM) in children with cerebral palsy (CP). However, outcomes at the muscle (remodeling) and resulting function appear to be highly variable and often unsatisfactory. During passive joint rotation, the Achilles tendon lengthens more than the in-series medial gastrocnemius muscle in children with CP, which might explain the limited effectiveness of stretching interventions. We aimed to ascertain whether increasing tendon stiffness, by performing resistance training, improves the effectiveness of passive stretching, indicated by an increase in medial gastrocnemius fascicle length. Methods: Sixteen children with CP (Age median [IQR]: 9.6 [8.6, 10.5]) completed the study. Children were randomly assigned to a combined intervention of stretching and strengthening of the calf muscles (n = 9) or a control (stretching-only) group (n = 7). Medial gastrocnemius fascicle length at a resting ankle angle, lengthening during passive joint rotations, and tendon stiffness were assessed by combining dynamometry and ultrasound imaging. The study was registered on clinicaltrials.gov (NCT02766491). Results: Resting fascicle length and tendon stiffness increased more in the intervention group compared to the control group (median [95% CI] increase fascicle length: 2.2 [1.3, 4.3] mm; stiffness: 13.6 [9.9, 17.7] N/mm) Maximum dorsiflexion angle increased equally in both groups. Conclusion: This study provides proof of principle that a combined resistance and stretching intervention can increase tendon stiffness and muscle fascicle length in children with CP. This demonstrates that remodeling of muscle structure is possible with non-invasive interventions in spastic CP.
Project description:After surgical repair of traumatically severed peripheral nerves, associated muscles are paralyzed for weeks. Little is known about fascicle length changes in paralyzed muscles during locomotion. The aim of this study was to investigate to what extent, if any, muscle fascicles of denervated feline soleus (SO) change length during stance of walking when intact SO synergists are actively contracting. Hindlimb kinematics, SO fascicle and muscle-tendon unit (MTU) length, and EMG activity of SO, lateral gastrocnemius (LG) and medial gastrocnemius (MG) were measured during level and slope walking in adult cats. Measurements were taken before and 1-2 weeks following SO-LG denervation. Unexpectedly, SO fascicle lengthening and shortening during stance in all walking conditions were evident after denervation. The greatest SO fascicle shortening (17.3 ± 2.2% of a reference length) and least fascicle lengthening (1.5 ± 0.8%) after denervation were found during upslope walking, where MG EMG activity was greatest across slopes (P < 0.05) and greatest discrepancies between post denervation SO fascicle and MTU length changes occurred. These findings suggest that myofascial linkages between denervated SO and its active synergists might affect its fascicle length changes. Further studies are needed to directly test this suggestion.
Project description:Differences in muscle and tendon responsiveness to mechanical stimuli and time courses of adaptive changes may disrupt the interaction of the musculotendinous unit (MTU), increasing the risk for overuse injuries. We monitored training-induced alterations in muscle and tendon biomechanical properties in elite jumpers over 4 years of athletic training to detect potential non-synchronized adaptations within the triceps surae MTU. A combined cross-sectional and longitudinal investigation over 4 years was conducted by analyzing triceps surae MTU mechanical properties in both legs via dynamometry and ultrasonography in 67 elite track and field jumpers and 24 age-matched controls. Fluctuations in muscle and tendon adaptive changes over time were quantified by calculating individual residuals. The cosine similarity of the relative changes of muscle strength and tendon stiffness between sessions served as a measure of uniformity of adaptive changes. Our cross-sectional study was unable to detect clear non-concurrent differences in muscle strength and tendon stiffness in elite jumpers. However, when considering the longitudinal data over several years of training most of the jumpers demonstrated greater fluctuations in muscle strength and tendon stiffness and hence tendon strain compared to controls, irrespective of training period (preparation vs. competition). Moreover, two monitored athletes with chronic Achilles tendinopathy showed in their affected limb lower uniformity in MTU adaptation as well as higher fluctuations in tendon strain over time. Habitual mechanical loading can affect the MTU uniformity in elite jumpers, leading to increased mechanical demand on the tendon over an athletic season and potentially increased risk for overuse injuries.
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:Current debate exists around whether a presumed eccentric exercise, the Nordic hamstring exercise (NHE), actually causes active hamstring muscle lengthening. This is because of the decoupling that can occur between the muscle fascicle and muscle-tendon unit (MTU) length changes in relatively compliant human lower-limb MTUs, which results in MTU lengthening not necessarily causing muscle fascicle lengthening. This missing knowledge complicates the interpretation of why the NHE is effective at reducing running-related hamstring muscle injury risk in athletes previously unfamiliar with performing this exercise. The purpose of the study was therefore to investigate if the most-commonly injured hamstring muscle, the biceps femoris long head (BF), exhibits active muscle lengthening (i.e. an eccentric muscle action) during the NHE up until peak force in Nordic novices. External reaction force at the ankle, knee flexion angle, and BF and semitendinosus muscle activities were recorded from the left leg of 14 participants during the NHE. Simultaneously, BF muscle architecture was imaged using B-mode ultrasound imaging, and muscle architecture changes were tracked using two different tracking algorithms. From ~85 to 100% of peak NHE force, both tracking algorithms detected that BF muscle fascicles (<i>n</i> = 10) significantly lengthened (<i>p</i> < 0.01) and had a mean positive lengthening velocity (<i>p</i> ≤ 0.02), while knee extension velocity remained positive (17°·s<sup>-1</sup>) over knee flexion angles from 53 to 37° and a duration of 1.6 s. Despite some individual cases of brief isometric fascicle behavior and brief fascicle shortening during BF MTU lengthening, the predominant muscle action was eccentric under a relatively high muscle activity level (59% of maximum). Eccentric hamstring muscle action therefore does occur during the NHE in relatively strong (429 N) Nordic novices, which might contribute to the increase in resting BF muscle fascicle length and reduction in running-related injury risk, which have previously been reported following NHE training. Whether an eccentric BF muscle action occurs in individuals accustomed to the NHE remains to be tested.
Project description:The purpose of this study was to investigate Achilles tendon (AT) length changes during a series of tasks that involved combinations of higher/lower force, and larger/smaller length changes of the medial gastrocnemius muscle-tendon unit (MTU). We sought to determine if common ultrasound-based estimates of AT length change were consistent with expectations for a passive elastic tendon acting in series with a muscle. We tested 8 healthy individuals during restricted joint calf contractions (high force, low displacement), ankle dorsi-/plantar-flexion (DF/PF) with the foot in the air (low force, high displacement), and heel raises (high force, high displacement). We experimentally estimated AT length change using two ultrasound methods, one based on muscle-tendon junction (MTJ) tracking and one based on muscle fascicle (MF) tracking. Estimates of AT length change were consistent with model expectations during restricted calf contractions, when the MTU underwent minimal length change. However, estimates of AT length changes were inconsistent with model expectations during the ankle DF/PF and heel raise tasks. Specifically, the AT was estimated to shorten substantially, often 10-20 mm, when the ankle plantarflexed beyond neutral position, despite loading conditions in which a passive, stiff spring would be expected to either lengthen (under increasing force) or maintain its length (under low force). These unexpected findings suggest the need for improvements in how we conceptually model and/or experimentally estimate MTU dynamics in vivo during motion analysis studies, particularly when the ankle plantarflexes beyond neutral.
Project description:In recent years, increasing the midsole bending stiffness (MBS) of running shoes by embedding carbon fibre plates in the midsole resulted in many world records set during long-distance running competitions. Although several theories were introduced to unravel the mechanisms behind these performance benefits, no definitive explanation was provided so far. This study aimed to investigate how the function of the gastrocnemius medialis (GM) muscle and Achilles tendon is altered when running in shoes with increased MBS. Here, we provide the first direct evidence that the amount and velocity of GM muscle fascicle shortening is reduced when running with increased MBS. Compared to control, running in the stiffest condition at 90% of speed at lactate threshold resulted in less muscle fascicle shortening (p?=?0.006, d?=?0.87), slower average shortening velocity (p?=?0.002, d?=?0.93) and greater estimated Achilles tendon energy return (p???0.001, d?=?0.96), without a significant change in GM fascicle work (p?=?0.335, d?=?0.40) or GM energy cost (p?=?0.569, d?=?0.30). The findings of this study suggest that running in stiff shoes allows the ankle plantarflexor muscle-tendon unit to continue to operate on a more favourable position of the muscle's force-length-velocity relationship by lowering muscle shortening velocity and increasing tendon energy return.
Project description:The <i>in vivo</i> characterization of the passive mechanical properties of the human triceps surae musculotendinous unit is important for gaining a deeper understanding of the interactive responses of the tendon and muscle tissues to loading during passive stretching. This study sought to quantify a comprehensive set of passive muscle-tendon properties such as slack length, stiffness, and the stress-strain relationship using a combination of ultrasound imaging and a three-dimensional motion capture system in healthy adults. By measuring tendon length, the cross-section areas of the Achilles tendon subcompartments (i.e., medial gastrocnemius and soleus aspects), and the ankle torque simultaneously, the mechanical properties of each individual compartment can be specifically identified. We found that the medial gastrocnemius (GM) and soleus (SOL) aspects of the Achilles tendon have similar mechanical properties in terms of slack angle (GM: -10.96° ± 3.48°; SOL: -8.50° ± 4.03°), moment arm at 0° of ankle angle (GM: 30.35 ± 6.42?mm; SOL: 31.39 ± 6.42?mm), and stiffness (GM: 23.18 ± 13.46?Nmm<sup>-1</sup>; SOL: 31.57 ± 13.26?Nmm<sup>-1</sup>). However, maximal tendon stress in the GM was significantly less than that in SOL (GM: 2.96 ± 1.50?MPa; SOL: 4.90 ± 1.88?MPa, <i>p</i> = 0.024), largely due to the higher passive force observed in the soleus compartment (GM: 99.89 ± 39.50?N; SOL: 174.59 ± 79.54?N, <i>p</i> = 0.020). Moreover, the tendon contributed to more than half of the total muscle-tendon unit lengthening during the passive stretch. This unequal passive stress between the medial gastrocnemius and the soleus tendon might contribute to the asymmetrical loading and deformation of the Achilles tendon during motion reported in the literature. Such information is relevant to understanding the Achilles tendon function and loading profile in pathological populations in the future.