Professional Documents
Culture Documents
www.elsevier.com/locate/gaitpost
Abstract
Utilization of elastic energy in the tendinous tissues (TT) of the human skeletal muscle may be task dependent. The present study was
designed to investigate this problem by comparing the fascicle-TT interaction of the medial gastrocnemius muscle (MG) during ground
contact of running and walking. Seven subjects ran and walked with a natural cadence. Ankle and knee joint angular data were recorded by
electrogoniometers for estimating the entire MG muscle-tendon unit (MTU) length, together with the ground reaction forces. The MG fascicle
length was measured by using the high-speed ultrasound image scanning during movements. The results showed that in running, after the
rapid early fascicle stretching (0–10% of the contact period), the fascicles shortened throughout the ground contact while TT was stretched
prior to shortening. In walking, the fascicles shortened initially (0–15% of the contact period) due to sudden plantar-flexion. Thereafter, the
fascicles and TT lengthened slowly until the end of single support (15–70% of the contact period.). The fascicles then shorted during the push-
off phase (70–100% of the contact period). These results demonstrate that the MG fascicles behaved differently between running and walking
and did not follow the length change pattern of the MTU during the ground contact period. The estimated working range of active muscle
fibers in force–length relationship could shift more to an ascending limb (shorter length) phase in running than in walking. These results
suggest that MG fascicles can work within the optimal working range of the sarcomeres in the force–length relation but are responsible for the
effective utilization of the TT elasticity during human running.
# 2006 Elsevier B.V. All rights reserved.
0966-6362/$ – see front matter # 2006 Elsevier B.V. All rights reserved.
doi:10.1016/j.gaitpost.2006.05.002
M. Ishikawa et al. / Gait & Posture 25 (2007) 380–384 381
tasks. Consequently, the issue of how skeletal muscles scanned (96 images s1) during the movements (Fig. 1b).
function might change with different tasks has remained The MG fascicles were identified in each image along their
relatively unexplored. In human drop jump exercises, the length from the superficial and deep aponeuroses. One end
fascicles of the medial gastrocnemius muscle (MG) of the fascicle, which was extended off the acquired
shortened throughout the ground contact [9]. On the other ultrasound image, was sometimes estimated. Surface bipolar
hand, recent human walking studies showed that MG Ag–AgCL electromyographic (EMG) electrodes (5 mm
fascicles contract isometrically [10] or even lengthen [11] diameter with interelectrode distance 20 mm) were used to
during the single support phase (15–70% of the contact record the EMG activities in MG (Glonner electronic,
period) when MTU and TT are stretching. These different Munich, Germany; input impedance >25 MV, common
and perhaps contradicting MG fascicle behaviors during mode rejection ratio >90 dB). Ankle and knee joint
SSC exercises may suggest existence of a movement rotations were recorded by electrogoniometers attached
specific fascicle behavior. Consequently, the fascicle-TT over the right lateral melleoli and popliteal crease,
interaction during human locomotion needs further respectively. These signals were stored simultaneously by
clarification. To begin to address issues relating to a 12-bit A/D converter with a sampling frequency of 1 kHz.
variability in human skeletal muscle function, the purpose
of the present study was to examine how the fascicle-TT 2.1. Analyses
interaction together with muscle activation varies in medial
gastrocnemius muscle (MG) due to the different tasks of To combine the analysis of the various parameters, the
walking and running. MTU length of MG data was re-sampled at 100 Hz and the
fascicle length data at 96 Hz was interpolated at 100 Hz so
that the time scales matched. The regression equations of
2. Methods Hawkins & Hull [12] were used to estimate MTU length in
MG from measurements of ankle and knee joint flexion
Seven healthy male volunteers (age 26.6 2.7 years, angles. The TT length was calculated by subtracting the
height 183.4 6.0 cm, body mass 80.9 15.2 kg; fascicle length multiplied by the cosine of the fascicle angle
means S.D.) participated after being informed all the from the MTU length [9–11]:
risks associated with this study, which was approved by the
LTT ¼ LMTU Lfa cos a
Ethic Committee of the University of Jyväskylä. The
subjects walked with a natural cadence (1.48 0.12 m s1) where LTT is the TT length, LMTU is the muscle-tendon unit
and ran (2.74 0.21 m s1) twice on a 10 m long force length, Lfa is the fascicle length and a is the angle created by
platform system (Raute Inc, Finland) (Fig. 1a). The ground the fascicle line and its insertion into the aponeurosis lines
reaction forces on the right and left sides were recorded (Fig. 1b). The length changes were calculated from the time
separately to define the ground contact phases. Real-time of initial foot contact to the peak length and from the peak
ultrasound devices (SSD-5500, Aloka, Japan) were used to length to the time of toe-off, respectively.
measure the fascicle length of MG during running and After the EMG signals were rectified, 10 step cycles of
walking [9–11]. A 7.5 MHz linear-array ultrasound probe of the signal data (EMG, ground reaction forces and joint angle
6 cm long was put over the MG muscle and fixed securely. data) were averaged for each subject. In the ultrasonographic
The longitudinal ultrasonographic images in MG were data the averaging was done for three consecutive step
Fig. 1. (a) Set-up for the walking and running experiments: the ultrasound images of fascicles and EMG of the medial gastrocnemius muscle (MG) were
recorded simultaneously on a unique 10 m long force plate system, composed of two rows of individual force plates. The ultrasound apparatus was pushed at the
speed of the subject outside the force plate area. The ankle and knee joint angles were also measured simultaneously by using goniometers. (b) The calculation of
the fascicle length (Lfa) and the angle from medial gastrocnemius (MG) muscle (the width and depth scanning areas are 58.5 and 60.0 mm, respectively).
382 M. Ishikawa et al. / Gait & Posture 25 (2007) 380–384
Fig. 2. Records of the measured and calculated parameters during the contact phase of running and walking. Vertical (A; Fz) and horizontal (B; Fy) ground
reaction forces, EMG activities of medial gastrocnemius (C; MG), the length of muscle-tendon unit (D; MTU), fascicle (E) and tendinous tissues (F; TT). The
contact period was normalized as 100%. Bars show the values of the 1 S.E. The vertical lines denote the contact, the transition point from the braking to push-
off phases and the take-off.
cycles. Coefficient of variation for fascicle length and angle tively) than in walking (48.9 2.5 and 44.1 3.8 cm,
of three step data were in the range of 1–6% and 1–7%, respectively). Table 1 shows the length changes from the
respectively. The EMG signals were integrated and then initial ground contact to the peak lengthening point. Both
averaged (aEMG) in the following three phases; preactiva- MTU and TT demonstrate greater stretching amplitudes in
tion, braking and subsequent push-off phases. The pre- running than in walking (Table 1, p < 0.05). However, the
activation was defined as the 100 ms preceding ground MG fascicle behavior was different between running and
contact and the braking and push-off phases were divided by walking (Fig. 2E). In running, after the initial ground contact,
the zero level of the horizontal ground force. the fascicles were lengthened by 0.4 0.3 cm immediately
after ground contact (0–10% of the contact period). They then
2.2. Treatment of data shortened throughout the contact period. Lfa in running was
greater at the initial ground contact (5.9 1.8 cm) than at the
Mean (S.D.) values were calculated for seven subjects toe-off (4.3 1.3 cm) ( p < 0.05). In contrast, the fascicles in
unless otherwise stated. A one-way repeated ANOVA on walking lengthened until the middle of the push-off phase
rank was used to analyze the significant difference of the
EMG activity between the different phases during the
movements. Where ANOVA was significant, a least Table 1
significant difference post hoc test was used to look for Measured and calculated parameters (n = 7).
significant differences between phases. The Wilcoxon Running Walking
signed-rank test was used to test for a difference between Contact time (ms) 296 28.4 629 33.0**
the parameters (EMG, length changes and ground reaction Averaged EMG (mV)
force) of walking and running. We considered p < 0.05 as a Pre-activation
statistically significant difference. Braking phase
Push-off phase
DLMTU (cm)
Contact-peak 2.2 0.9 1.2 0.4*
3. Results Peak-toe-off 4.0 0.3 3.7 0.5*
DLfascicle (cm)
Both running and walking showed the stretching and Contact-peak 0.4 0.3 0.2 0.3
Peak-toe-off 2.0 0.6 1.0 0.4*
subsequent shortening of the MTU and TT during the ground DLTT (cm)
contact period in all subjects (Fig. 2D and F). LMTU and LTT at Contact-peak 3.0 1.2 1.4 0.5*
the initial ground contact and at the toe-off points did not show Peak-toe-off 3.7 0.6 3.2 0.8
any significant difference between running and walking. Values are expressed as means S.D. *, ** Significantly different between
However, LMTU and LTT at the peak lengthening point were running and walking at p < 0.05, p < 0.01, respectively. #, ## Significantly
longer in running (50.4 2.7 and 45.7 3.9 cm, respec- different between phases at p < 0.05, p < 0.01, respectively.
M. Ishikawa et al. / Gait & Posture 25 (2007) 380–384 383
(70% of the contact period) after the initial shortening due to constant length. However, it has been proposed that the
the sudden plantar flexion (10% of the contact period). They concerted fascicle action, where activation can be matched
then shortened during the late push-off phase (70–100% of the to the imposed load in order to keep the length of the
contact period). In comparison between running and walking, contractile component constant [14], can favor the effective
Lfa at the initial ground contact and at the peak lengthening storage of TT elastic energy during the ground contact.
point did not show any significant difference between running These different fascicle length patterns (shortening and/or
(5.9 1.8 and 6.3 1.5 cm, respectively) and walking constant) may also be related to the intensity of the
(6.1 1.4 and 6.3 1.5 cm, respectively). Lfa at the toe- movement [9,15]. It can be speculated that in the MG
off point was greater in walking (5.3 1.3 cm) than in muscle the elastic energy in TT can be utilized due to the
running (4.3 1.3 cm) ( p < 0.05). The magnitudes of the fascicle shortening during the braking phase of the present
fascicle shortening from the peak Lfa until toe-off were greater running task.
in running than in walking ( p < 0.05, Table 1), but the According to the previous human walking studies
stretching magnitudes did not show any difference. The [10,11], the interaction between the fascicles and TT can
magnitudes of the TT shortening were not significantly play an important role in the storage and release of elastic
different between tasks but in running the TT stretching was energy. In walking, however, the elastic energy cannot be
greater than in walking (Table 1). provided directly from initial impact due to the low Achilles
The peak vertical ground reaction force (Fz) was greater tendon force [11,16] but mainly from the muscle action.
in running (1805 282 N) than in walking (978 215 N) Consequently, the MG muscle in walking was not strongly
( p < 0.05). When compared to walking, the MG aEMGs activated during the preactivation and braking phases while
were greater in the preactivation and braking phases of the MG fascicles were lengthened during the braking phase
running ( p < 0.05). This was the opposite in the push-off (Fig. 2E). Both in running and jumping, however, the
phase ( p < 0.05) (Table 1). Thus in running EMG activity of Achilles tendon force (ATF) increases during the braking
MG occurred primarily during the braking phase (Fig. 2C, phase [11]. The elastic energy can come mainly from the
Table 1). In contrast, the greater muscle activation in initial negative work in running due to the strong muscle
walking was observed during the early push-off phase. activation during the preactivation and braking phases. In the
present running condition, the MG fascicles contracted
together with high muscle activation during the preactiva-
4. Discussion tion and braking phases (Fig. 2). Thus, the present results
suggest that the fascicle-TT interaction is movement specific
We investigated the modulation of the MG fascicle-TT in both human walking and running.
interaction in response to changes in mechanical demands It has been suggested that the muscle fiber contraction
associated with tasks. The TT stretching magnitudes of MG occurs around the plateau region of the sarcomere force–
from the contact to peak length was greater in running length curve during human movements [17]. Supporting
(7.4 2.5%) than in walking (4.1 0.9%). A similar their suggestion, the estimation of the force–length relation
difference was observed for the MTU stretching (running of the sarcomeres (Fig. 3) shows that the working range of
4.3 1.7%, walking 2.5 0.6%). In running, however, the
MG fascicles behaved differently from the MTU and TT.
The MG fascicles in running shortened throughout the
ground contact after the rapid fascicle stretching (0–10% of
the contact period). These results can indicate the existence
of the fascicle-TT interaction, which is movement specific in
both human walking and running.
Roberts et al. [8] have shown that in the turkey running,
the tendon stretched and recoiled as the ankle flexed and
extended while the foot was on the ground, but the fascicles
remained almost constant in length. The present results also
demonstrated that TT was stretched during the braking
phase, followed by shortening during the push-off phase in
running. Thus, we confirmed the existence of the elastic Fig. 3. The proposed working area of sarcomere length–force relation of
behavior of TT, which allows for storage of elastic energy in human gastrocnemius muscle during human locomotion ((a): Ref. [10], (b):
the first half of the step and its release in the second half of Ref. [21], (c): Ref. [9]). Average sarcomere lengths, which were estimated
the contact phase [13]. In this TT behavior, the MG fascicles by dividing the fascicle length by the average number of sarcomeres in
shorten throughout the ground contact phase. This MG series in fascicles [22], have been estimated during human locomotion and
superimposed on a force–length relationship for human muscle derived
fascicle shortening during the MTU stretching accompa- from the data of Walker and Schrodt (see Refs. [17] and [20] for details).
nied by the higher EMG activation (Fig. 2) can increase the The shaded areas refer to the estimation which is based on the present
TT stretch as compared to the fascicle lengthening or the results.
384 M. Ishikawa et al. / Gait & Posture 25 (2007) 380–384