Effect of knee joint angle on individual hamstrings morphology quantified using free-hand 3D ultrasonography

Exercise responses and injury rates differ between individual hamstrings and this may be linked with their morphology. The aim of this study was to compare muscle length and tendon dimensions between the individual hamstrings at two knee joint angles using free hand three-dimensional ultrasound (3D US). Muscle-tendon length and distal tendon cross-sectional area (CSA), volume, length and echogenicity of biceps femoris long (BFlh) and short (BFsh) head, semimembranosus (SM) and semitendinosus (ST) of 16 individuals were measured using free-hand 3D US at 0° (full extension) and 45° of knee flexion. ST showed the greatest length than all muscles and BFsh the lowest (p < 0.05). No difference was observed between SM and BFlh length (p > 0.05). Of the four muscles, ST tendon was longer, with less volume and CSA but greater echogenicity than the other tendons. In contrast, SM and BFlh showed shorter tendons and lower echogenicity but a greater volume and CSA than ST (p < 0.05). Muscle and tendon lengthened from 45° to 0° knee flexion angle (p < 0.05) but this change was not statistically different between individual hamstrings (p > 0.05). Freehand 3D US indicated that hamstring muscle length and distal tendon dimensions differ between individual hamstrings. All muscles and tendons lengthened as the knee was extended but this change was similar for all individual hamstrings.     

Validity of architectural properties of the hamstring muscles: Correlation of ultrasound findings with cadaveric dissection

The purpose of this study was to compare the architectural parameters of the long head of biceps femoris (BFlh) and semitendinosus (ST) muscles by comparing measurements from ultrasound (US) with those obtained from direct dissection. The BFlh and ST architectures were examined bilaterally in 6 legs from 3 male cadavers. The fascicle length, pennation angle, muscle thickness and muscle and tendon length were obtained from direct measurement and US scans along each muscle. Intraclass correlation coefficients between the two methods ranged from 0.905 to 0.913 for the BFlh variables and from 0.774 to 0.974 for the ST parameters. Compared with the direct measurements, the US method showed a mean typical error of 0.09–0.14 cm for muscle thickness, 1.01–1.31° for the pennation angle, 0.92–1.71 cm for fascicle length and muscle–tendon length measurements. The US method is a valid alternative tool for assessing basic architectural parameters of ST and BFlh components of the hamstring muscles.     

Musculotendon variability influences tissue strains experienced by the biceps femoris long head muscle during high-speed running

The hamstring muscles frequently suffer injury during high-speed running, though the factors that make an individual more susceptible to injury remain poorly understood. The goals of this study were to measure the musculotendon dimensions of the biceps femoris long head (BFlh) muscle, the hamstring muscle injured most often, and to use computational models to assess the influence of variability in the BFlh’s dimensions on internal tissue strains during high-speed running. High-resolution magnetic resonance (MR) images were acquired over the thigh in 12 collegiate athletes, and musculotendon dimensions were measured in the proximal free tendon/aponeurosis, muscle and distal free tendon/aponeurosis. Finite element meshes were generated based on the average, standard deviation and range of BFlh dimensions. Simulation boundary conditions were defined to match muscle activation and musculotendon length change in the BFlh during high-speed running. Muscle and connective tissue dimensions were found to vary between subjects, with a coefficient of variation (CV) of 17±6% across all dimensions. For all simulations peak local strain was highest along the proximal myotendinous junction, which is where injury typically occurs. Model variations showed that peak local tissue strain increased as the proximal aponeurosis width narrowed and the muscle width widened. The aponeurosis width and muscle width variation models showed that the relative dimensions of these structures influence internal muscle tissue strains. The results of this study indicate that a musculotendon unit’s architecture influences its strain injury susceptibility during high-speed running.     

Superimposing hip extension on knee flexion evokes higher activation in biceps femoris than knee flexion alone

Hamstring muscle function during knee flexion has been linked to hamstring injury and performance. However, it is unclear whether knee flexion alone (KF) requires similar hamstring electromyography (EMG) activity pattern to simultaneous hip extension and knee flexion (HE-KF), a combination that occurs in the late swing phase of sprinting. This study examined whether HE-KF maximal voluntary isometric contraction (MVIC) evokes higher (EMG) activity in biceps femoris long head (BFlh) and semitendinosus (ST) than KF alone. Effects of shank rotation angles were also tested. Twenty-one males performed the above-mentioned MVICs while EMG activity was measured along ST and BFlh. Conditions were compared using a one-way mixed functional ANOVA model under a fully Bayesian framework. Higher EMG activity was found in HE-KF in all shank rotation positions than in KF in the middle region of BFlh (highest in the 9th channel, by 0.022 mV [95%CrI 0.014 to 0.030] in neutral shank position). For ST, this was only observed in the neutral shank position and in the most proximal channel (by 0.013 mV [95%CrI 0.001 to 0.025]). We observed muscle- and region-specific responses to HE-KF. Future studies should examine whether hamstring activation in this task is related to injury risk and sprint performance.     

Fascicle length of gastrocnemius muscles in monozygous twins

A large inter-individual variation is seen in muscle fascicle length of the athletes but the reasons for this phenomenon are unclear. The purpose of this study was to determine whether genetic factors contribute to the variances in muscle architectural characteristics. Nine monozygous twin pairs (3 males and 6 females), mean age 23 years (range 17-40) were studied. Fascicle length, pennation angle, and muscle thickness of the medial (MG) and lateral (LG) gastrocnemius muscles were measured in vivo by B-mode ultrasound. In the LG muscle intrapair resemblance (P < 0.01) for fascicle length (r = 0.98), pennation angle (r = 0.94) and muscle thickness (r = 0.86) were observed. In MG muscle, however, there was no intrapair resemblance for fascicle length (r = 0.66, P > 0.05), although pennation angle (r = 0.73, P < 0.05) and muscle thickness (r = 0.86, P < 0.01) were significant. Mean percent intrapair difference in LG and MG muscles were 1.8% and 5.1% for fascicle length, 11.3% and 12.3% for pennation angle and 12.4% and 9.9% for muscle thickness, respectively. There is intrapair difference between muscle thickness and pennation angle in both MG (r = 0.69, P < 0.05) and LG (r = 0.70, P < 0.05) muscles. However, no significant correlation was observed for intrapair difference between muscle thickness and fascicle length in both muscles (MG, r = 0.46; LG, r = 0.40). It appears that genetic predisposition is the predominant factor for the determination of muscle fascicle length. However, a lack of intrapair resemblance in MG fascicle length raises the possibility that fascicle length may be further influenced by external environmental factors such as physical training.     

Hamstring muscle architecture and myotonometer measurements in elite professional football players with a prior strained hamstring

The purpose of this study was to compare the fascicle length, angle pennation and mechanical properties of the biceps femoris long head (BFlh) in dominant and non-dominant limbs in previously injured and uninjured professional football players. Fifteen professional football players were recruited to participate in this study. Seven players had suffered a BFlh injury during the previous season. Myotonometry mechanical properties were measured in the proximal, common tendon and distal BFlh using MyotonPRO, and angle pennation and fascicle length were also measured. We observed significantly higher distal BFlh frequency, stiffness, decrement, relaxation and creep than in the common tendon and proximal BFlh. The previously injured players showed significantly higher frequency and stiffness, and lower relaxation and creep in the dominant BFlh than did uninjured players. There were no significant differences between the fascicle length and angle pennation in previously injured and uninjured BFlh. Myotonometric measurement provides a quick and inexpensive way to check the properties of the BFlh in professional football players. Professional football players with previous BFlh injury showed higher intrinsic tension and a poorer capacity to deform than did players with no injury to the BFlh.     

The isometric functional capacity of muscles that cross the elbow

We hypothesized that muscles crossing the elbow have fundamental differences in their capacity for excursion, force generation, and moment generation due to differences in their architecture, moment arm, and the combination of their architecture and moment arm. Muscle fascicle length, sarcomere length, pennation angle, mass, and tendon displacement with elbow flexion were measured for the major elbow muscles in 10 upper extremity specimens. Optimal fascicle length, physiological cross-sectional area (PCSA), moment arm, operating range on the force-length curve, and moment-generating capacity were estimated from these data. Brachioradialis and pronator teres had the longest (17.7cm) and shortest (5.5cm) fascicles, respectively. Triceps brachii (combined heads) and brachioradialis had the greatest (14.9cm(2)) and smallest (1.2cm(2)) PCSAs, respectively. Despite a comparable fascicle length, long head of biceps brachii operates over a broader range of the force-length curve (length change=56% of optimal length, 12.8cm) than the long head of triceps brachii (length change=28% of optimal length, 12. 7cm) because of its larger moment arm (4.7cm vs. 2.3cm). Although brachioradialis has a small PCSA, it has a relatively large moment-generating capacity (6.8cm(3)) due to its large moment arm (average peak=7.7cm). These results emphasize the need to consider the interplay of architecture and moment arm when evaluating the functional capabilities of a muscle.     

Sprint performance is related to muscle fascicle length in male 100-m sprinters.

The purpose of this study was to investigate the relationship between muscle fascicle length and sprint running performance in 37 male 100-m sprinters. The sample was divided into two performance groups by the personal-best 100-m time: 10.00-10.90 s (S10; n = 22) and 11.00-11.70 s (S11; n = 15). Muscle thickness and fascicle pennation angle of the vastus lateralis and gastrocnemius medialis and lateralis muscles were measured by B-mode ultrasonography, and fascicle length was estimated. Standing height, body weight, and leg length were similar between groups. Muscle thickness was similar between groups for vastus lateralis and gastrocnemius medialis, but S10 had a significantly greater gastrocnemius lateralis muscle thickness. S10 also had a greater muscle thickness in the upper portion of the thigh, which, given similar limb lengths, demonstrates an altered "muscle shape." Pennation angle was always less in S10 than in S11. In all muscles, S10 had significantly greater fascicle length than did S11, which significantly correlated with 100-m best performance (r values from -0.40 to -0.57). It is concluded that longer fascicle length is associated with greater sprinting performance.     

Relationship between sprint performance and muscle fascicle length in female sprinters

The purpose of this study was to investigate the relationship between sprint performance and architectural characteristics of leg muscles in 26 female 100-m sprinters. Pennation angle and muscle thickness of the vastus lateralis (VL) and gastrocnemius medialis (GM) and lateralis (GL) muscles were measured by B-mode ultrasonography, and fascicle length was estimated. Sprinters had a significantly lower VL pennation angle, but GM and GL pennation angle was similar between sprinters and female control subjects (N = 22). There was no significant correlation between pennation angle and 100-m personal best performance. Sprinters had significantly greater absolute fascicle length in VL and GL than controls, which significantly correlated to 100-m best-record (r = -0.51 and r = -0.44, respectively). Relative fascicle length (VL and GL) were also significantly greater in sprinters than controls. However, there were no significant correlation between relative fascicle length and 100-m best-record (r = -0.36 and r = -0.29, respectively). No relationship was found between the sprint performance and fat-free mass (r = -0.26) or body mass index (r = -0.03). However, there was a significant correlation between percent (%) body fat and 100-m best-record (r = 0.62, p < 0.01). Adjusting the confounding effect of % fat, significant correlations were seen between relative fascicle length and 100-m best-record (VL; r = -0.39 and GL; r = -0.40). Absolute and relative fascicle length were similar in elite female sprinters compared with previous reported values for elite male sprinters (Kumagai et al., 2000). It was concluded that longer fascicle length is associated with greater sprinting performance in sprinters, but there is no gender differences in fascicle length for elite sprinters.     

In vivo fascicle behavior of synergistic muscles in concentric and eccentric plantar flexions in humans.

Ultrasonography was used to directly measure in vivo fascicle behavior of the medial gastrocnemius (MG) and soleus (SOL) muscles while the subjects (n=6 men) performed maximal voluntary concentric and eccentric plantar flexions at 60, 120, 180 and 240 deg/s. Fascicle shortening and lengthening velocities of MG, obtained from fascicle length changes over time, were significantly higher than those of SOL at +/-120, +/-180 and +240 deg/s, possibly reflecting physiological and mechanical differences between these muscles. On the other hand, the effective fascicle shortening and lengthening velocities, defined as the velocities in the longitudinal direction of muscle belly, were not significantly different between MG and SOL. This could be due to difference in fascicle architecture and/or the existence of mechanical linkages between these muscles. Moreover, when the contribution of tendinous tissues to muscle-tendon complex length change was determined from fascicle length, pennation angle, moment arm and joint angle, it accounted for approximately 50% in both concentric and eccentric trials, but showed considerable intra-subject variations. This result quantifiably demonstrates the importance of tendinous tissues in isokinetically controlled joint movements.     

Acute effect of muscle stretching on the steadiness of sustained submaximal contractions of the plantar flexor muscles

This paper examines the acute effect of a bout of static stretches on torque fluctuation during an isometric torque-matching task that required subjects to sustain isometric contractions as steady as possible with the plantar flexor muscles at four intensities (5, 10, 15, and 20% of maximum) for 20 s. The stretching bout comprised five 60-s passive stretches, separated by 10-s rest. During the torque-matching tasks and muscle stretching, the torque (active and passive) and surface electromyogram (EMG) of the medial gastrocnemius (MG), soleus (Sol), and tibialis anterior (TA) were continuously recorded. Concurrently, changes in muscle architecture (fascicle length and pennation angle) of the MG were monitored by ultrasonography. The results showed that during stretching, passive torque decreased and fascicle length increased gradually. Changes in these two parameters were significantly associated (r(2) = 0.46; P < 0.001). When data from the torque-matching tasks were collapsed across the four torque levels, stretches induced greater torque fluctuation (P < 0.001) and enhanced EMG activity (P < 0.05) in MG and TA muscles with no change in coactivation. Furthermore, stretching maneuvers produced a greater decrease (～15%; P < 0.001) in fascicle length during the torque-matching tasks and change in torque fluctuation (CV) was positively associated with changes in fascicle length (r(2) = 0.56; P < 0.001), MG and TA EMG activities, and coactivation (r(2) = 0.35, 0.34, and 0.35, respectively; P < 0.001). In conclusion, these observations indicate that repeated stretches can decrease torque steadiness by increasing muscle compliance and EMG activity of muscles around the joint. The relative influence of such adaptations, however, may depend on the torque level during the torque-matching task.     

Reliability and robustness of muscle architecture measurements obtained using diffusion tensor imaging with anatomically constrained tractography

For detailed analyses of muscle adaptation mechanisms during growth, ageing or disease, reliable measurements of muscle architecture are required. Diffusion tensor imaging (DTI) and DTI tractography have been used to reconstruct the architecture of human muscles in vivo. However, muscle architecture measurements reconstructed with conventional DTI techniques are often anatomically implausible because the reconstructed fascicles do not terminate on aponeuroses, as real muscle fascicles are known to do. In this study, we tested the reliability of an anatomically constrained DTI-based method for measuring three-dimensional muscle architecture. Anatomical magnetic resonance images and diffusion tensor images were obtained from the left legs of eight healthy participants on two occasions one week apart. Muscle volumes, fascicle lengths, pennation angles and fascicle curvatures were measured in the medial and lateral gastrocnemius, soleus and the tibialis anterior muscles. Averaged across muscles, the intraclass correlation coefficient was 0.99 for muscle volume, 0.81 for fascicle length, 0.73 for pennation angle and 0.76 for fascicle curvature. Measurements of muscle architecture obtained using conventional DTI tractography were highly sensitive to variations in the stopping criteria for DTI tractography. The application of anatomical constraints reduced this sensitivity significantly. This study demonstrates that anatomically constrained DTI tractography can provide reliable and robust three-dimensional measurements of whole-muscle architecture. The algorithms used to constrain tractography have been made publicly available.     

American Society of Biomechanics Journal of Biomechanics Award 2017: High-acceleration training during growth increases optimal muscle fascicle lengths in an avian bipedal model

Sprinters have been found to possess longer muscle fascicles than non-sprinters, which is thought to be beneficial for high-acceleration movements based on muscle force-length-velocity properties. However, it is unknown if their morphology is a result of genetics or training during growth. To explore the influence of training during growth, thirty guinea fowl (Numida meleagris) were split into exercise and sedentary groups. Exercise birds were housed in a large pen and underwent high-acceleration training during their growth period (age 4–14 weeks), while sedentary birds were housed in small pens to restrict movement. Morphological analyses (muscle mass, PCSA, optimal fascicle length, pennation angle) of a hip extensor muscle (ILPO) and plantarflexor muscle (LG), which differ in architecture and function during running, were performed post-mortem. Muscle mass for both ILPO and LG was not different between the two groups. Exercise birds were found to have ∼12% and ∼14% longer optimal fascicle lengths in ILPO and LG, respectively, than the sedentary group despite having ∼3% shorter limbs. From this study we can conclude that optimal fascicle lengths can increase as a result of high-acceleration training during growth. This increase in optimal fascicle length appears to occur irrespective of muscle architecture and in the absence of a change in muscle mass. Our findings suggest high-acceleration training during growth results in muscles that prioritize adaptations for lower strain and shortening velocity over isometric strength. Thus, the adaptations observed suggest these muscles produce higher force during dynamic contractions, which is beneficial for movements requiring large power outputs.     

Estimation of cat medial gastrocnemius fascicle lengths during dynamic contractions

In typical muscle models, it is often assumed that the contractile element (fascicle) length depends exclusively on the instantaneous muscle-tendon length and the instantaneous muscle force. In order to test whether the instantaneous fascicle length during dynamic contractions can be predicted from muscle-tendon length and force, fascicle lengths, muscle-tendon lengths, and muscle forces were directly measured in cat medial gastrocnemii during isometric and dynamic contractions. Two theoretical muscle models were developed: model A was based on force-time data obtained during the activation phase and model D on force-time data obtained during the deactivation phase of isometric contractions. To test the models, instantaneous fascicle lengths were predicted from muscle-tendon lengths and forces during dynamic contractions that simulated cat locomotion for speeds ranging from 0.4 to 1.6m/s. The theoretically predicted fascicle lengths were compared with the experimentally measured fascicle lengths. It was found that fascicle lengths were not uniquely associated with muscle-tendon lengths and forces; that is, for a given muscle-tendon length and force, fascicle lengths varied depending on the contractile history. Consequently, models A and D differed in fascicle length predictions; model D (maximum average error=8.5%) was considerably better than model A (maximum average error=22.3%). We conclude from this study that it is not possible to predict the exact fascicle lengths from muscle-tendon lengths and forces alone, however, adequate predictions seem possible based on such a model. The relationship between fascicle length and muscle force and muscle-tendon length is complex and highly non-linear, thus, it appears unlikely that accurate fascicle length predictions can be made without some reference contractions in which fascicle length, muscle-tendon length, and force are measured simultaneously.     

Changes in fascicle length from rest to maximal voluntary contraction affect the assessment of voluntary activation

The purpose of this study was to investigate the effect of the differences between the actual fascicle length during a voluntary contraction and the fascicle length at rest of the triceps surae muscle on the determination of the voluntary activation (VA) by using the interpolated twitch technique. Twelve participants performed isometric voluntary maximal (MVC) and submaximal (20%, 40%, 60% and 80% MVC) contractions at two different ankle angles (75 degrees and 90 degrees ) under application of the interpolated twitch technique. Two ultrasound probes were used to determine the fascicle length of soleus, gastrocnemius medialis and gastrocnemius lateralis muscles. Further, the MVCs and the twitches were repeated for six more ankle angles (85 degrees , 95 degrees , 100 degrees , 105 degrees , 110 degrees and 115 degrees ). The VA of the triceps surae muscle were calculated (a) using the rest twitch force (RTF) measured during the same trial as the interpolated twitch force (ITF; traditional method) and (b) using the RTF at an ankle angle where the fascicle length showed similar values between ITF and RTF (fascicle length consideration method). The continuous changes in fascicle length from rest to MVC affect the accuracy of the assessment of the VA. The traditional method overestimates the assessment of the VA on average 4% to 12%, especially at 90 degrees ankle angle (i.e. short muscle length). The reason for this influence is the unequal force-length potential of the muscle at twitch application by the measure of ITF and RTF. These findings provide evidence that the fascicle length consideration method permits a more precise prediction (an improvement of 4-12%) of the voluntary contraction compared to the traditional method.     

Skeletal Muscle Fascicle Arrangements Can Be Reconstructed Using a Laplacian Vector Field Simulation

Skeletal muscles are characterized by a large diversity in anatomical architecture and function. Muscle force and contraction are generated by contractile fiber cells grouped in fascicle bundles, which transmit the mechanical action between origin and insertion attachments of the muscle. Therefore, an adequate representation of fascicle arrangements in computational models of skeletal muscles is important, especially when investigating three-dimensional muscle deformations in finite element models. However, obtaining high resolution in vivo measurements of fascicle arrangements in skeletal muscles is currently still challenging. This motivated the development of methods in previous studies to generate numerical representations of fascicle trajectories using interpolation templates. Here, we present an alternative approach based on the hypothesis of a rotation and divergence free (Laplacian) vector field behavior which reflects observed physical characteristics of fascicle trajectories. To obtain this representation, the Laplace equation was solved in anatomical reconstructions of skeletal muscle shapes based on medical images using a uniform flux boundary condition on the attachment areas. Fascicle tracts were generated through a robust flux based tracing algorithm. The concept of this approach was demonstrated in two-dimensional synthetic examples of typical skeletal muscle architectures. A detailed evaluation was performed in an example of the anatomical human tibialis anterior muscle which showed an overall agreement with measurements from the literature. The utility and capability of the proposed method was further demonstrated in other anatomical examples of human skeletal muscles with a wide range of muscle shapes and attachment morphologies.     

Effects of 20 days bed rest on muscle morphology

Using ultrasound, muscle thickness and fascicle angles from aponeurosis were evaluated before, during and after 20 days bed rest (BR). Subjects were healthy adults (4 women and 4 men). Measurements were carried out before and after BR and after 10 weeks of recovery, respectively. Muscle measurements were taken at nine sites in trunk and upper and lower extremities, respectively. For the m. triceps brachii, m. vastus lateralis, and m. gastrocnemius medialis, fascicle angles from the aponeurosis as well as muscle thickness were measured. There was a high statistical significant correlation between muscle thickness and cross-sectional area for quadriceps muscles, suggesting applicability of muscle thickness for evaluation of muscle size. Muscle thickness decreased in muscles of the lower extremity by 2.1-4.4 % after bed rest. In triceps brachii and vastus lateralis muscles, there were no prominent changes in muscle thickness and fascicle angles. It was concluded that muscle morphology deteriorates with changes in muscle architecture by bed rest but the response is small and muscle-specific. It was also suggested that bed rest affects not only muscle mass but muscle tone as well.     

Hamstring muscle kinematics and activation during overground sprinting

Hamstring muscle strain injury is one of the most commonly seen injuries in sports such as track and field, soccer, football, and rugby. The purpose of this study was to advance our understanding of the mechanisms of hamstring muscle strain injuries during over ground sprinting by investigating hamstring muscle-tendon kinematics and muscle activation. Three-dimensional videographic and electromyographic (EMG) data were collected for 20 male runners, soccer or lacrosse players performing overground sprinting at their maximum effort. Hamstring muscle-tendon lengths, elongation velocities, and linear envelop EMG data were analyzed for a running gait cycle of the dominant leg. Hamstring muscles exhibited eccentric contractions during the late stance phase as well as during the late swing phase of overground sprinting. The peak eccentric contraction speeds of the hamstring muscles were significantly greater during the late swing phase than during the late stance phase (p=0.001) while the hamstring muscle-tendon lengths at the peak eccentric contraction speeds were significantly greater during the late stance phase than during the late swing phase (p=0.001). No significant differences existed in the maximum hamstring muscle-tendon lengths between the two eccentric contractions. The potential for hamstring muscle strain injury exists during the late stance phase as well as during the late swing phases of overground sprinting.     

Biceps femoris and semitendinosus tendon/aponeurosis strain during passive and active (isometric) conditions

The purpose of this study was to quantify strain and elongation of the long head of the biceps femoris (BFlh) and the semitendinosus (ST) tendon/aponeurosis. Forty participants performed passive knee extension trials from 90° of knee flexion to full extension (0°) followed by ramp isometric contractions of the knee flexors at 0°, 45° and 90° of knee flexion. Two ultrasound probes were used to visualize the displacement of BFlh and ST tendon/aponeurosis. Three-way analysis of variance designs indicated that: (a) Tendon/aponeurosis (passive) elongation and strain were higher for the BFlh than the ST as the knee was passively extended (p < 0.05), (b) contraction at each angular position was accompanied by a smaller BFlh tendon/aponeurosis (active) strain and elongation than the ST at higher levels of effort (p < 0.05) and (c) combined (passive and active) strain was significantly higher for the BFlh than ST during ramp contraction at 0° but the opposite was observed for the 45° and 90° flexion angle tests (p < 0.05). Passive elongation of tendon/aponeurosis has an important effect on the tendon/aponeurosis behavior of the hamstrings and may contribute to a different loading of muscle fibers and tendinous tissue between BFlh and ST.