Eur J Appl Physiol

DOI 10.1007/s00421-016-3349-3

ORIGINAL ARTICLE

Muscle‑specific acute changes in passive stiffness of human
triceps surae after stretching
Kosuke Hirata1 · Eri Miyamoto‑Mikami1,2 · Hiroaki Kanehisa1 · Naokazu Miyamoto1 

Received: 6 December 2015 / Accepted: 24 February 2016
© Springer-Verlag Berlin Heidelberg 2016

Abstract 
Purpose  It remains unclear whether the acute effect of
stretching on passive muscle stiffness differs among the
synergists. We examined the muscle stiffness responses of
the medial (MG) and lateral gastrocnemii (LG), and soleus
(Sol) during passive dorsiflexion before and after a static
stretching by using ultrasound shear wave elastography.
Methods  Before and after a 5-min static stretching by
passive dorsiflexion, shear modulus of the triceps surae
and the Achilles tendon (AT) during passive dorsiflexion
in the knee extended position were measured in 12 healthy
subjects.
Results  Before the static stretching, shear modulus was
the greatest in MG and smallest in Sol. The stretching
induced significant reductions in shear modulus of MG, but
not in shear modulus of LG and Sol. The slack angle was
observed at more plantar flexed position in the following
order: AT, MG, LG, and Sol. After the stretching, the slack
angles of each muscle and AT were significantly shifted to
more dorsiflexed positions with a similar extent. When considering the shift in slack angle, the change in MG shear
modulus became smaller.
Conclusion  The present study indicates that passive
muscle stiffness differs among the triceps surae, and that
the acute effect of a static stretching is observed only in
the stiff muscle. However, a large part of the reduction of

Communicated by Olivier Seynnes.
* Naokazu Miyamoto
miyamoto@nifs‑k.ac.jp
1

National Institute of Fitness and Sports in Kanoya,
1 Shiromizu, Kanoya, Kagoshima 891‑2311, Japan

2

Japan Society for the Promotion of Science, Tokyo, Japan

passive muscle stiffness at a given joint angle could be due
to an increase in the slack length.
Keywords  Ultrasound shear wave elastography · Shear
modulus · Gastrocnemius · Soleus · Ankle joint
Abbreviations
ANOVA Analysis of variance
AT Achilles tendon
EMG Electromyography
LG Lateral gastrocnemius
MG Medial gastrocnemius
MTU Muscle–tendon unit
MVC Maximal voluntary contraction
RMS Root mean square
ROM Range of motion
Sol Soleus

Introduction
Pre-exercise stretching is usually performed for decreasing passive muscle stiffness. Understanding the effect
of stretching on muscle stiffness has long been a topic of
interest for not only athletes and coaches but also clinicians and researchers. In human experiments, passive muscle stiffness has been widely evaluated from the relationship between joint angle and passive joint torque (Gajdosik
2001; Magnusson 1998; Magnusson et al. 1997). However,
the joint torque is not a measure specific to the responses
of individual muscles, because the joint torque results from
a composite of contractile (i.e., synergists and antagonists)
and non-contractile tissues such as tendon, skin, ligament and articular structures. Based on the change in joint
torque, therefore, it is difficult to determine alteration in the

13

Nakamura et al. 1973). Each subject completed an informed consent form. 2010. at an angular velocity of 1°/s. the subjects were requested to relax completely and to not resist the movement of the footplate. previous studies have examined changes in muscle fascicle length and behavior of tendinous tissues during passive lengthening of the MTU (Abellaneda et al. 1998). Additionally. The Ethics Committee on Human Research of National Institute of Fitness and Sports in Kanoya approved the investigation. the muscle stiffness of the medial gastrocnemius (MG) has been reported to be reduced (Morse et al. Because of this limitation. Mutungi and Ranatunga 1996. 2013. Muscle shear modulus measured by ultrasound shear wave 13 Eur J Appl Physiol elastography is shown to be highly correlated with Young’s modulus obtained via traditional materials testing (Eby et al. This is a recently developed imaging technique that can quantify localized tissue stiffness along the principal axis of the probe. 2013). 61. Morse et al.. For example. By the use of this technique. 2002. The rotation axes of the ankle and the footplate were aligned as closely as possible. 20. and Sol during passive dorsiflexion before and after an acute stretching intervention. Using B-mode ultrasonography. 2008). all reported angle measurements refer to the ankle joint angle assessed with the goniometer. Konrad and Tilp 2014b).5 cm. previous studies which examined passive lengthening of the plantar flexors (i. The very slow angular velocity was used to obtain a better resolution for the muscle stiffness via ultrasound shear wave elastography and to avoid or minimize the stretch reflex (Hirata et al.7 ± 8. perhaps because of the difficulty to properly quantify the elongation of other muscles such as the lateral gastrocnemius (LG) and soleus (Sol) by using B-mode ultrasonography.9 years. The development of imaging techniques such as ultrasonography allows us to evaluate underlying architectures of the muscle–tendon unit (MTU) in humans in vivo. all subjects were fully informed of the purpose and experimental procedures. 2008). the present study aimed to test the hypotheses mentioned above by measuring the muscle stiffness responses of MG. In order to familiarize the subjects to the procedure and to ensure that they were as relaxed as possible. The right foot was tightly fixed to the dynamometer’s footplate. muscle fiber type composition has been shown to be different among the synergists (Johnson et al. Biometrics. the sole of the foot at right angles to the tibia axis) defined as 0°. 2008. Thus. None of the subjects reported any ongoing neuromuscular diseases or musculoskeletal injuries specific to the ankle or knee joints. They were asked to refrain from strenuous exercise 48 h before testing. As one of the reasons for the inconsistent conclusions over studies. Materials and methods Subjects Twelve healthy subjects (eight men and four women) participated in the present study (168. Morse et al. Herbert et al. methodological issue for evaluating the muscle stiffness may be involved. Morse et al. 2011) or not be affected (Kato et al. UK) fixed to the ankle joint with double-sided adhesive tape. type I muscle fibers exhibit greater passive stiffness (Kovanen et al. the conclusions of previous studies using B-mode ultrasonography on the effect of stretching on muscle stiffness have not reached a consensus. 2011).4 ± 2. The ankle joint angle was passively dorsiflexed from 50° plantar flexion [neutral position (i.8 ± 7. 2009. Before participation. The angular displacement of the ankle joint was measured with an electronic goniometer (SG110/A.. 2015. However. PHYSIOMED. not about mechanical properties such as stiffness. In the human triceps surae. 2008. One of the methods to resolve the aforementioned concerns is to use ultrasound shear wave elastography. Palmeri et al. Nakamura et al. 1973). Throughout the passive lengthening.9 kg. and all of the experimental procedures were performed in accordance with the Declaration of Helsinki. it remains unknown whether the acute effect of stretching on passive muscle stiffness differs among the synergists. passive dorsiflexion) have very often investigated only MG (Abellaneda et al. other methods are required to more directly and properly assess the passive stiffness of individual muscles in human in vivo. B-mode ultrasonography per se can provide information only about architecture such as muscle– tendon junction displacement and fascicle length. Blazevich et al. not the angle of the dynamometer footplate. mean ± SD). 2008). Muscle stiffness is influenced by the muscle fiber type composition. 2004. stiffness of individual muscles. 2009. a familiarization session of . In addition to the fact that the passive torque is related to the viscoelastic properties of the entire musculoarticular complex.e. Blazevich et al. which could stiffen the muscles. Germany) bed with their right knees fully extended. Experimental setup and protocol Subjects lay prone on a dynamometer (CON-TREX MJ.e. 1984b. 2012. with larger values for dorsiflexion] to the end ROM which was defined as the angle that the subjects felt pain. Therefore. LG. it is hypothesized that the responses in passive muscle stiffness to passive ankle dorsiflexion and their changes induced by an acute bout of stretching is greater in Sol composed of 70–100 % type I muscle fibers than in MG and LG composed of >50 % type II muscle fibers (Johnson et al. 2012. Konrad and Tilp 2014b. In this study. based on the propagation speed of remotely induced shear wave (Bercoff et al. Hug et al.

FA-DL-141. these data were manually synchronized with elastography recording (sampling frequency = 1 Hz). Maïsetti et al. LG. data from 50° plantarflexion up to 25° dorsiflexion were used for the analyses. For elastographic data. Therefore. The reference electrode was placed over the left medial malleolus. inter-electrode distance: 12 mm. This familiarization session also has a role to avoid a conditioning effect of passive lengthening on the stiffness of muscle and tendon (Hirata et al. and Sol. 2) were 0. the data processing was 13 .01x − 5. the measurements were performed during the 3rd cycle and later (PRE).97 ± 0. surface EMG signals during the prescribed tasks were obtained from MG.00 ± 0. along the fascicle direction of each muscle. Konrad and Tilp 2014a. and 0. the same passive lengthening procedure was applied (POST) to the pre-determined end ROM. and cleaning with alcohol. Australia) with a sampling frequency of 1 kHz. Also. Data analysis The passive torque was corrected for the gravity effect of the footplate.3 kPa.Eur J Appl Physiol two cycles was performed followed by the start of the testing session. Throughout the stretching.2 for MG. joint angle. over the medial or lateral aspect of Sol distal to muscle-tendon junction of MG or LG. France) with a 50 mm linear array probe (SL15-4. Yoshitake et al. ADInstrument. 4-assist. bandpass filtering: 5–450 Hz) bipolar active surface electrodes (electrode shape: parallel-bar. Then. respectively. 6 and Ver. Sol. The Young’s modulus measured by shear wave elastography was highly correlated with measured values of Young’s modulus obtained via traditional materials testing (r > 0. and Sol. Static stretching maneuver Static stretching was administered to the right lower leg of the subject. The torque. or over AT. Each ultrasound probe was placed over the bellies of MG and LG. Validity and repeatability of shear wave elastography measurements The validity of measurement of shear modulus by using the ultrasound shear wave elastography has been described in detail in previous reports (Miyamoto et al. 2015a). and EMG data were simultaneously stored on a personal computer for later analyses using a 16 bit analogue-to-digital converter (PowerLab/16SP. the validity was evaluated in tissue-mimicking phantoms which were calibrated by a conventional stress–strain test. The order of testing for MG. Supersonic Imagine.1. LG. The determination of probe location was performed prior to the placement of electromyography (EMG) electrodes. respectively. Supersonic Imagine. The optimal probe location was determined for subject-by-subject to identify several fascicles without interruption across the B-mode image in a certain plane and get good-looking images. 2014). the passive lengthening as mentioned above was repeated over the same ROM for assessing the other targets. The repeatability during the passive dorsiflexion was evaluated in three subjects for each muscle. b). and AT within the two passive lengthening was randomly assigned and counterbalanced across subjects.01. rubbing with sandpaper. the root mean square values (RMSEMG) were computed over 0. Additionally. LG. the RMS-EMG value of each muscle was normalized to that obtained during isometric maximal voluntary contraction (MVC) which performed at 0° of the ankle joint. Care was taken not to press and deform the muscles while scanning.00. previous studies have suggested that shear modulus measured by ultrasound shear wave elastography could be considerably affected by pennation angle (Gennisson et al. The inter-trial absolute difference was 0. 7.5 s period at each joint angle for each muscle. Then. France) with musculoskeletal preset were used to quantify the passive stiffness of each muscle and the Achilles tendon (AT). After shaving. However. 2010. 2015. Absolute differences were 1. size: 1 mm width × 8 mm length. LG. pre-amplified (gain: 500×.99 ± 0. 2012). more recently we have shown that the effect of pennation angle on the measured shear modulus was negligibly small (<1. along the fascicle direction. Since four targets (three muscles and one tendon) were investigated with two systems in the present study.2 ± 0.99). as all subjects could reach this ankle angle before and after a 5-min static stretching. Briefly. The ankle joint angle was held at the end ROM of dorsiflexion for 5 min.8 ± 0.3. 0. and Sol.3 %) (Miyamoto et al. immediately after a 5-min static stretching (see below). Japan) were placed next to each ultrasound position of MG.45) close to the identity line. and 0. 1. Ultrasound shear wave elastography Two ultrasound shear wave elastography systems (Aixplorer Ver.7 ± 0. LG. All passive lengthening procedures used for measurements of muscle stiffness were performed up to the end ROM. The intra-class correlation coefficients (1. The posture of the subject and setup were similar to those for the muscle stiffness measurement as mentioned above. with the regression line (y = 1. For EMG data. EMG In order to ensure that the lengthening procedures were conducted under passive condition. However. and Sol. 2015b. the subjects were requested to relax completely and not offer any voluntary resistance.3 kPa for MG.

Results ROM. a circular area as large as possible with exclusion of aponeurosis and subcutaneous adipose tissue (Fig.2 N m.9 ± 8. Due to the saturation limit of shear modulus for the elastography system. based on the assumption of an isotropic material. and AT.0 % MVC) at plantar flexed positions whereas they were slightly increased toward the end ROM (<3. 2).. Fig. the shear modulus of AT were saturated at about −25° (i. passive torque and EMG After 5 min static stretching.3° ± 2. Actually. Hence. IBM Japan. Japan).05. shear modulus measurements of AT were used only for the determination of the tendon’s slack angle.6° to 31. we calculated shear modulus by dividing the obtained Young’s modulus by 3. The significance level for all comparisons was set at P < 0. Sol. Thus.003). AT)]. which is obviously not true for the muscle. Data are reported as means and SD.9 ± 4. soleus (Sol.e.2 ± 5. For RMS-EMG data. a three-way ANOVA revealed that there was only a significant main effect of angle (P = 0.2 % MVC) for each muscle. POST) × muscle × angle] with repeated measures were used. The EMG activities during the passive lengthening were relatively small (<1.. For the RMS-EMG and shear modulus data. Eur J Appl Physiol performed by using software of the ultrasound scanner for every 1° of the ankle joint. triangles). a paired t test was performed. 0°. Arrows in shear modulus-joint angle curves indicate the slack angles of MG. and 20° of dorsiflexion angle. Depending on the subject and target. LG.. The colored region represents the shear modulus map with the scale to the right of the figure.e. ROM was significantly increased from 28. circles) and lateral gastrocnemius (LG. 1b). and Achilles tendon (AT. the reduction at 25° was 6. No value in the circular region reached the saturation limit of shear modulus for the elastography system (267 kPa) except for AT. and time (i. Based on the shear modulus-ankle joint relationships.e. the software provides the Young’ modulus by multiplying the shear modulus by the constant 3. Circle is the region of interest for determination of shear modulus (color figure online) flexion torque increased exponentially with an increase in the ankle joint angle (Fig. .0°) (P  = 0. The passive torque was reduced after a 5-min static stretching. PRE and POST) information. which reached significance at 10°–25° (P < 0. Two-way ANOVAs with repeated measures were used for the torque (time × angle) and slack angle [time × target (i. When appropriate. For maximal ROM data. 1  Typical example of responses in passive plantar flexion torque (a) and shear modulus (b) of the medial (MG.014) without any interactions. This determination of the slack angle was visually conducted (Hirata et al. 2015. squares) during passive lengthening before a static stretching. diamonds).8° (ΔROM  = 2. Lacourpaille et al. despite the subjects being asked to relaxed during the passive lengthening. PRE. MG. LG. The bottom images (c) are typical examples of shear modulus measurements of MG obtained at −40°. Pearson’s product-moment correlation coefficient (r) was determined to assess the correlation between changes in passive torque and shear modulus. The mean value of the three experimenters was used for further analyses. 1c) was chosen as the region of interest for the shear modulus calculation. Passive plantar 13 Fig. The statistical analyses were performed by using statistical software (SPSS Statistics 21. Statistics Statistical analyses were performed for the values at every 5° of the ankle joint angle. Sol..e. the slack angles of each muscle were determined as the first increase above the variation in shear modulus at the plantarflexed position (Fig. 1a). 25° of plantar flexion). target.05. 2014) by three experimenters who were blinded to subject identification. additional analyses (Tukey and paired t-tests with Bonferroni correction) were performed. separate three-way analyses of variance (ANOVAs) [time (i.

001) without time × target interaction.3 ± 8. filled) and after (POST.001) without main effect of time or time × angle interaction for LG and Sol. the slack angle differed among the three muscles.001) and target (P < 0. P = 0. Table 1 shows the slack angles of the triceps surae and AT before and after the static stretching. A three-way ANOVA showed a significant time × muscle × angle interaction (P  = 0. filled) and after (POST.001).022).008.8°. 2  Changes in passive plantar flexion torque during passive lengthening before (PRE. Follow-up analyses demonstrated that shear modulus of MG after the static stretching was significantly smaller than that before the stretching at 30° and 35° from the slack angle (P  = 0. Additionally. circles) and lateral gastrocnemius (LG. The relative change [i. the shear modulus of MG after the static stretching was significantly smaller than that before the stretching at 10°–25° of the ankle joint angle (Fig. 3)..e. showing that the slack angles of each muscle and AT were significantly shifted toward more dorsiflexed positions after the static stretching. diamonds) during passive lengthening before (PRE. showing the characteristic curvilinear relationships with shear modulus increasing steeply toward the end ROM. *Significantly different between PRE and POST (P < 0.Eur J Appl Physiol Fig. especially in MG.4 ± 17.2 kPa (20. 1b.031). Separate twoway ANOVAs for each muscle showed that a significant time  × angle interaction (P  = 0. open) a 5-min static stretching. Follow-up two-way ANOVAs revealed that a significant time × angle interaction was observed only for MG shear modulus whereas there was only a significant main effect of Angle (P < 0. As observed in Fig.e. triangles). A two-way ANOVA showed significant main effects of time (P  = 0. The mean difference in slack angle between PRE and POST across Target was 1. Figure 3 shows pooled data of shear modulus of each muscle during passive lengthening before and after the static stretching. respectively) although the magnitude of the reduction in shear modulus after the stretching was considerably smaller 13 . the reduction at 25° was 29.05) Shear modulus Slack angle Figure  1b represents examples of ultrasound images and of the shear modulus-ankle joint angle relationship. more plantar flexed position) in the following order: AT < MG < LG < Sol (P < 0. and soleus (Sol. open) a 5-min static stretching.9 % of PRE value). Tukey post hoc tests demonstrated that shear modulus was significantly greater in MG than in LG and Sol above −5° and greater in LG than Sol above 5° both before and after the stretching (P < 0.. A Tukey post hoc test revealed that the slack angle was significantly smaller (i.620. Correlation analysis was further performed to examine a potential association between changes in passive torque and shear modulus of MG.003) was observed only for MG whereas there was only a significant main effect of angle without main effect of time or time × angle interaction for LG and Sol. *Significantly different between PRE and POST in MG (P < 0.045 and P  = 0. 3  Changes in shear modulus of the medial (MG.05) Fig. According to further analyses. (POST value − PRE value)/PRE value] in passive torque was significantly correlated to that in shear modulus of MG across subjects (r = 0. Effect of slack angle on shear modulus Figure  4 shows the relationships between shear modulus and ankle angle from slack for each muscle.05).

circles) and lateral gastrocnemius (LG. and Achilles tendon before and after a static stretching Dorsiflexion angle at slack (°) MG LG Sol AT Before After* −19. AT achilles tendon * Significant main effect of time without time × target interaction (P < 0. Theoretically. the present result that regarding RMS-EMG only a significant main effect of Angle was observed without any interactions indicates that the muscle activities during the .9 −14. open) a 5-min static stretching. triangles).2 ± 5. (2) the effect of a static stretching on shear modulus was significant only in MG. Thus. However. muscle stiffness at a given ankle joint angle was greater in MG and LG than Sol.3 ± 2. *Significantly different between PRE and POST in MG (P < 0. 1998). 1991) and/or in content of collagen which composes intramuscular connective such as the perimysium and endomysium (Kovanen et al. the reflexive muscle activities would likely have contributed significantly to the muscle stiffness during “passive” lengthening. Eur J Appl Physiol Table 1  Slack angles of the medial and lateral gastrocnemius. soleus.0 ± 7. MG.1 ± 4. it was expected that. contrary to the expectation. LG.3 −44. Sol composed of 70–100 % type I muscle fibers exhibits greater muscle stiffness than MG and LG composed of >50 % type II muscle fibers (Johnson et al. Intermuscle differences in passive stiffness Previous in vitro studies have shown that type I muscle fibers have greater passive stiffness compared with type II muscle fibers (Mutungi and Ranatunga 1996. the EMG activities during the lengthening procedure were negligible at plantar flexed positions (<1 % MVC) whereas the EMG activities were slightly increased toward the end ROM for each muscle (<~3 % MVC). 2013).6 kPa. Sol soleus. Zimmerman et al. 1973). However.8 ± 2. diamonds) before (PRE. the present findings suggest that passive muscle stiffness differs among the triceps surae.0 ± 5.5 −2.0 −17. 1993). However. 3). In this study.5 ± 5. 2003.8 −4. Nevertheless. filled) and after (POST. and soleus (Sol. it is possible that the passive stiffness of the individual muscles of the triceps surae is strongly influenced by architectural characteristics such as sarcomere length and corresponding slack length. Friden and Lieber 2002).05). At the start of this study.8 % of PRE) compared with that based on the shear modulus-joint angle relationship (Fig. Kawakami et al. Thus. passive stretching elicits the stretch reflex. a large part of the reduction of passive muscle stiffness at a given joint angle could be due to an increase in the slack length. Discussion The major findings of the present study were that (1) shear modulus during passive lengthening of the triceps surae was the greatest in MG and smallest in Sol. MG). LG lateral gastrocnemius.. Shear modulus measured by ultrasound shear wave elastography has been shown to be highly correlated with the material stiffness (Young’s modulus) of a 13 muscle measured by a conventional stress–strain test (Eby et al. therefore.1 ± 2. This result suggests that factors other than muscle fiber type composition should be involved to determine the intermuscle differences in passive stiffness among the triceps surae. and Sol. which would stiffen the lengthened muscles. 4  Relationships between shear modulus and ankle angle from slack for the medial (MG. 6.05) (4. and that the acute effect of a static stretching is observed only in the stiff muscle (i. during passive lengthening of the triceps surae. the slack angle was observed at more plantar flexed position in the following order: AT. and (3) the magnitude of the reduction of MG shear modulus after the static stretching became smaller when considering the shift in slack angle induced by the stretching.0 −12.8 MG medial gastrocnemius. the rate of lengthening was set at a sufficiently slow angular velocity (1°/s) to avoid or minimize the stretch reflex.e. (2000) have reported that the sarcomere length of MG is longer than that of Sol at a given ankle joint angle in the knee extended position. possibly due to differences in titin isoform within each fiber (Wang et al. Taking these observations into account together with the finding that passive force generally develops at or near the optimal length (Davis et al. 1984a.3 −43. Values are mean ± SD Fig. Additionally.

13 . A limitation of the present study is that passive lengthening and stretching were performed only in the knee extended position. Another limitation is that shear modulus measurement was performed at only one region for each muscle. Sol) in the knee extended position can be observed in the knee flexed position. in the present study. Especially. the static stretching could not alter the passive muscle stiffness of LG and Sol.g. Acute effect of static stretching on MG passive stiffness The stretching maneuver adopted here increased ROM and decreased passive torque at a given ankle joint angle. proximal vs. Morse et al. Therefore. 2005).Eur J Appl Physiol lengthening procedure were not different among the three muscles and between before and after the static stretching. They concluded that thixotropic properties of a muscle are unlikely to be the mechanism leading to increased flexibility as a result of static stretching. However. Taking these observations into account together with the fact that the triceps surae and AT are serially connected. and aponeurosis from the analyses of shear wave elastographic data. the connective tissue. However. muscle stiffness was reduced only in MG.and post-stretching. (2008) have subjected the plantar flexors including MG to a number of quick stretches that would be expected to abolish the short range elastic component or possibly change the conformation of titin. After the stretching. 2003. not to that of the epimysium. particularly the perimysium. As mentioned above. 3). perimysium and endomysium) could influence passive stiffness. Additionally. it is possible that the reduction of MG muscle stiffness at the dorsiflexed positions after the stretching is attributed mainly to the shift of slack angle of AT. the extent of the substantial reduction of MG passive stiffness was considerably smaller (7 %) when considering the shift in slack angles after the stretching (Fig.g. Gajdosik (2001) suggested that lengthening deformation of the connective tissues within the muscle belly (e. Moreover. a large part of the reduction of passive muscle stiffness at a given joint angle could be due to an increase in the slack length. These results indicate that the stretching maneuver could be effective to increase the flexibility of the entire plantar flexor muscle group. passive muscle stiffness was significantly reduced only in MG. the 2° shift of the slack angle in AT necessarily induces the same degree of the slack angles in the triceps surae. where shear modulus of LG and Sol was less than half of that of MG (Fig. Thus. distal. the present study has shown that passive muscle stiffness during passive lengthening of the triceps surae was the greatest in MG and smallest in Sol. Therefore. Compliance with ethical standards  Conflict of interest  The authors declare that they have no conflict of interest.. in spite of the fact that MG and LG are bi-articular muscles crossing for the knee and ankle joints. Thus. Thixotropic property of a muscle has been proposed as one of the possible mechanisms for the reduced muscle stiffness after the static stretching (Rassier et al. Another mechanism is decreased stiffness of intramuscular connective tissues (Morse et al. (2015) have revealed that MG passive stiffness measured by ultrasound shear wave elastography was reduced only when sufficient mechanical stress was imposed on the muscle during stretching. However. it is unclear whether the current findings hold true for other regions (e. The muscle stiffness of MG was significantly reduced by 20 % at 25° after the static stretching (Fig. In conclusion. lateral) within each muscle. However. it is reasonable that the stretching effect was observed only in MG. In other words. 4). Freitas et al. The significance remained even when considering the shift in slack angle induced by the static stretching (Fig. after a 5-min static stretching of the triceps surae. the slack angles of each muscle and AT were significantly shifted toward more dorsiflexed positions with a similar extent.. Further investigation is warranted to clarify these. The ultrasound shear wave elastography scanner used in the present study can quantify localized tissue stiffness and we have taken care to exclude the outside connective tissues such as epimysium. the reduction in passive muscle stiffness of MG after static stretching observed in the present study could be due to the deformation of the perimysium and/or endomysium. it is well known that muscle stiffness of this nature can be reduced by relatively small amounts of movement (Campbell and Lakie 1998). is considered to be a major extracellular contributor to passive stiffness (Purslow 1989). fascia. 2006). it is unlikely that the muscle activities during passive lengthening are responsible for the observed differences in passive stiffness between muscles and between pre. 4). Additionally. the ankle joint angle was held at the end ROM of dorsiflexion. Thus. Acknowledgments  This work was supported by JSPS KAKENHI Grant Number 25702038. the slack angle was significantly observed at more plantar flexed position in the following order: AT < MG < LG < Sol. Hodgson et al. after the static stretching. During the static stretching in the present study. 3). Thus. it is unclear whether the differences in passive muscle stiffness and slack angle among triceps surae (MG or LG vs. 2008). medial vs. Sol has a complex multipennate arrangement of the muscle fibers (Agur et al. These findings suggest that the acute effect of a static stretching on passive muscle stiffness is muscle-specific and occurs only in the stiff muscle.

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