rapid communication
Determination of fascicle length and pennation
in a contracting human muscle in vivo
TETSUO FUKUNAGA, YOSHIHO ICHINOSE, MASAMITSU ITO,
YASUO KAWAKAMI, AND SENSHI FUKASHIRO
Department of Life Sciences, The University of Tokyo,
Komaba 3–8–1, Meguro, Tokyo 153, Japan
Fukunaga, Tetsuo, Yoshiho Ichinose, Masamitsu Ito,
Yasuo Kawakami, and Senshi Fukashiro. Determination
of fascicle length and pennation in a contracting human
muscle in vivo. J. Appl. Physiol. 82(1): 354–358, 1997.—We
have developed a technique to determine fascicle length in
human vastus lateralis muscle in vivo by using ultrasonogra-
phy. When the subjects had the knee fully extended passively
from a position of 110° flexion (relaxed condition), the fascicle
length decreased from 133 to 97 mm on average. During static
contractions at 10% of maximal voluntary contraction strength
(tensed condition), fascicle shortening was more pronounced
(from 126 to 67 mm), especially when the knee was closer to
full extension. Similarly, as the knee was extended, the angle
of pennation (fascicle angle, defined as the angle between
fascicles and aponeurosis) increased (relaxed, from 14 to 18°;
tensed, from 14 to 21°), and a greater increase in the
pennation angle was observed in the tensed than in the
relaxed condition when the knee was close to extension
(,40°). We conclude that there are differences in fascicle
lengths and pennation angles when the muscle is in a relaxed
and isometrically tensed conditions and that the differences
are affected by joint angles, at least at the submaximal
contraction level.
vastus lateralis muscle; ultrasound; pennation angle; muscle
contraction
THE FORCE EXERTED BY MUSCLE fibers is modified by their
geometric arrangement, structures of the joint, and the
angle and location of the tendon in respect to the bone,
before it appears at the joint as moments ( joint torque).
Thus the sole observation of joint actions gives little
information on muscle actions within. For instance, the
relationship between joint position and torque does not
disclose the length-force characteristics of the muscle.
This is especially true for pennate muscles in which
short fibers are arranged at an angle to the line of
action of the muscle, and the angle as well as muscle
fiber length change during contraction.
Knowledge of the geometric arrangement of muscle
fibers, i.e., muscle architecture, is therefore important
when studying muscle functions and the resultant joint
actions. To date, parameters such as the length, cross-
sectional area, and pennation angles of fibers have been
investigated (1–3, 6, 7, 10, 11, 15, 19, 20, 23), and it has
been shown that the muscle architecture considerably
affects the manner in which muscle force is transmitted
354 0161-7567/97 $5.00 Copyright r 1997 the American Physiological Society
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to the tendons and bones (11, 15). Many of the previous
studies on human muscle architecture have been done
through a direct dissection of cadaver specimens. How-
ever, muscles in the embalmed cadavers have been
reported to change their morphological characteristics
because of factors such as shrinkage (7). Besides, this
method does not allow to study the effect of muscle
contraction and changes in joint position on muscle
architecture.
The purpose of the present study was to determine
architecture of a human muscle, both at rest and
during contractions. We employed real-time ultrasonog-
raphy to visualize fascicles in vivo (12, 13). The relation-
ships between knee joint angles and fascicle lengths as
well as angles of pennation for the vastus lateralis
muscle were investigated.
METHODS
Six healthy men (age 25 6 1 yr, body height 174.5 6 2.6 cm,
body mass 72.1 6 4.4 kg) participated as subjects. The nature
and possible consequences of the study were explained to
each subject, and informed consent was obtained before-
hand.
A real-time ultrasonic apparatus (SSD-2000, 7.5 MHz,
Aloka, Japan) was used. Precision and linearity of the image
have been confirmed by the authors elsewhere (12). The level
of 50% of the thigh length (50% of the distance from the
greater trochanter to the lateral epicondyle of the femur) was
determined. At this level, the width of the vastus lateralis
muscle was measured by the use of the ultrasound apparatus,
and the position of one-half of the width was marked with a
pen. At this position, a longitudinal ultrasonic image was
obtained, and fascicles were visualized. One of the fascicles in
the image was carefully traced by moving the scanning head
on the dermal surface proximally and distally, and ultrasound
images containing the whole fascicle from the proximal and
distal ends were obtained. From them, the length of the
fascicle was measured by the use of a digital curvimeter
(Comcurve-8, Koizumi-Sokki, Japan). The measurement was
repeated five times for each subject. The coefficients of
variation ranged from 0.5 to 4.3%. The pennation angles
(fascicle angles), defined as the angles between the echoes of
the deep aponeurosis of vastus lateralis and the echoes from
interspaces among fascicles (9, 12, 17), were also measured.
During measurement, the subject was seated on a test
bench of an electric dynamometer (MYORET, Asics, Japan)
with a backrest and was secured by using straps around the
waist, breast, and the knee. The axis of the lever arm of the
 FASCICLE LENGTH AND PENNATION IN CONTRACTING HUMAN MUSCLE
dynamometer was visually aligned with the estimated center
of the knee joint. An ankle cuff on the lever arm was attached
proximal to the lateral malleolus. The knee joint angles were
changed every 10° from full extension to flexion at 110°, and,
at each joint angle, ultrasonic measurement was performed.
The subject relaxed knee extensor muscles during measure-
ment.
After the measurement in the relaxed condition, measure-
ments were carried out while the muscles were in a tensed
condition. After the subject familiarized himself with the
testing procedures and after a warm-up session of several
submaximal and maximal exertions, the subject was encour-
aged to perform maximal isometric knee extension at the
joint angles from flexion at 110° to full extension, with an
interval of 10°, and the peak torque during exertion was
recorded. Rest periods were .1 min between contractions to
prevent fatigue and ensure a maximal contraction at each
joint angle. After the maximal torque measurement, 10% of
the maximal torque was calculated for each joint angle. At
respective joint angles, the subject exerted knee extension
torque at 10% of the maximum, with a visual aid of the torque
displayed with the target torque on a monitor screen. The
ultrasonic measurement was done after confirmation that
torque became stable at the target level.
The lengths and angles of the fascicles in the relaxed and
tensed conditions were measured on two different occasions.
Reproducibility was tested by computing coefficients of varia-
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355
Fig. 1. Ultrasonic longitudinal image of vastus
lateralis muscle. Ultrasonic transducer was
placed on skin over the muscle at 50% distance
from greater trochanter to lateral epicondyle of
femur. Fascicle length (fL) was determined as
length of a line drawn along ultrasonic echo
parallel to fascicle. Fascicle angle (u) was deter-
mined as angle between echoes obtained from
fascicles and deep aponeurosis in ultrasonic
image. k, Distal end of a fascicle.
tion (CV) according to the following equation
CV 5 (SD/ xc) 3 100
where SD is the square root of the between-test variance
obtained from the one-way repeated-measures analysis of
variance, and xc is the combined mean of the two measure-
ments (5). In addition, a paired t-test was performed for two
measurements in each condition and at each joint angle to
test their statistical difference. To investigate the effect of the
conditions (relaxed and tensed) and knee joint angles on the
lengths and angles of the fascicles, a two-way analysis of
variance with repeated measures was used (2 3 12, condi-
tions 3 knee joint angles). For those variables for which a
significant effect was found, a Tukey post hoc test was used. A
P , 0.05 level of confidence was set for all analyses.
RESULTS
In the ultrasonic image obtained from vastus latera-
lis muscle of one subject (Fig. 1), one can clearly observe
the echoes reflected from fascicles. The CV of repeated
measurements of the fascicle lengths ranged from 0 to
6.8% (relaxed, mean: 2.1%) and from 0 to 6.1% (tensed,
mean: 3.0%) for the knee angles tested; CV values of the
fascicle angles ranged from 0 to 2.0% (relaxed, mean:
0.8%) and from 0 to 3.8% (tensed, mean: 1.7%). The
356 FASCICLE LENGTH AND PENNATION IN CONTRACTING HUMAN MUSCLE
Fig. 2. Changes in fascicle lengths at various angles of knee joint.
Measurements were done in 2 conditions: a relaxed (r) condition
with no muscle contraction and a tensed condition (s), where subject
performed static knee extensions at 10% of maximal voluntary
strength using an electrical dynamometer. Error bars indicate 1 SE
between subjects. * Significantly different between relaxed and tensed
conditions, P , 0.05.
variability of CV was not affected by the knee joint
angles. Also, there were no significant differences be-
tween mean values of the repeated measurements both
for the lengths and angles of the fascicles.
When the knee was fully extended passively from the
flexed position (110°, full extension corresponds to 0°;
relaxed condition), the fascicle length decreased by 27%
from 132.9 6 8.3 to 96.7 6 6.9 (SE) mm (Fig. 2). When
static voluntary contractions were performed at several
joint angles at 10% of the maximal voluntary contrac-
tion (MVC) strength, fascicle lengths decreased, and
the magnitude was greater (30%) when the knee was
closer to full extension than when the knee was flexed
(5%) (Fig. 2). The effect of different conditions was
significant at each joint angle except at 110°.
Fig. 3. Changes in fascicle angles as a function of knee angles.
Significantly larger fascicle angles in tensed than in relaxed condi-
tions were observed at extended positions (i.e., knee angle ,30°).
Error bars indicate 1 SE between subjects. * Significantly different
between relaxed (r) and tensed (s) conditions, P , 0.05.
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The fascicle angle increased from 14 to 18° during
knee extension (Fig. 3). At the same joint angle, larger
fascicle angles were observed in tensed than in relaxed
conditions, especially when the knee was in extended
positions with joint angles ,40° (statistically signifi-
cant at 0, 10, 20, and 30°). Angles of pennation for
vastus lateralis muscle have been reported to be 5–20°
in cadavers (23) and 6–27° in healthy subjects (9, 17).
These results are in good agreement with our measure-
ments.
The relationships between fascicle angles, fascicle
lengths, and knee joint angles differed between relaxed
and tensed conditions. When the knee was flexed
beyond 90°, fascicle length in the tensed condition was
slightly but significantly shorter than in the relaxed
condition, with no apparent difference in fascicle angles.
When the knee joint angle was between 90 and 40°,
shorter fascicle lengths were more pronounced in the
tensed condition, with fascicle angles being somewhat
smaller than in the relaxed condition. When the knee
was extended ,40°, fascicle lengths were much shorter
in the tensed condition, with a large significant incre-
ment in fascicle angles.
DISCUSSION
The muscle architecture is an intermediate level that
affects conversion of the force and excursion of muscle
fibers into joint actions. Understanding of this level is
important when we estimate events that are happening
in the muscle from observations at the joint, but
analysis of this level is often ignored (8). It has been
shown that architectural parameters such as muscle
fiber length and pennation angles affect functional
characteristics of a muscle (e.g., maximal shortening
velocity and maximal tension) as well as intrinsic ones,
such as fiber composition (4, 18). In some previous
studies, human muscle architecture has been observed
in cadaver specimens and related to functional charac-
teristics (2, 3, 6, 10, 21), but very little data are
available in living human muscles. In the present
study, fascicle length and pennation angles were deter-
mined in vivo in a contracting human muscle, which is
the first attempt as long as the authors’ knowledge can
reach. We considered that the arrangement of a fas-
cicle, which is connected to proximal and distal ten-
dons, is important when one studies how muscle force is
transmitted to tendons and bones.
Changing knee position passively from 110° to full
extension resulted in change of fascicle length from 133
to 97 mm on average. When the muscle was actively
contracting in an isometric manner, the change in knee
position by the same amount corresponded to fascicle
shortening from 126 to 67 mm. The larger shortening
would have been caused by taking up of the elongated
series elastic component. The 30-mm difference be-
tween the relaxed and tensed conditions is as much as
one-half of the optimal fiber length of the vastus
lateralis muscle (64–67 mm; Ref. 20). The results also
show that the range of the fascicle length change is
beyond the optimal length of the vastus lateralis muscle
 FASCICLE LENGTH AND PENNATION IN CONTRACTING HUMAN MUSCLE
(20). This could mean that this muscle uses a descend-
ing limb of the length-tension relationship within the
physiological range, which agrees with a previous
report on hand muscles (16).
Wickiewicz et al. (21) investigated in vivo force-
velocity characteristics of muscle fibers from torque
and angle of the knee joint. In their study, fiber lengths
of the knee extensor muscles were derived from cadav-
ers. They measured knee extension torque using an
isokinetic dynamometer at an angle of 30° from full
extension. However, as shown in the present study, at
this knee position, the length of the fascicle between
relaxed and tensed conditions differs greatly (110 vs. 92
mm). In fact, the researchers estimated that the muscle
fiber length change for 90° knee excursion was 15–23%,
which was comparable to the present result for the
relaxed condition (23%, Fig. 2), but in the tensed
condition the change was much greater (43%). There-
fore, the determination by Wickiewicz et al. of shorten-
ing velocity of the muscle fibers could have been
underestimated. If the present results were interpo-
lated to their data, the predicted maximal shortening
velocity of fibers would be 589 not 315 mm/s (21).
Because the force applied to the tendon by the
contraction of muscle fibers is reduced by a factor of
cosu (where u 5 pennation angle) from the sum of the
forces of the individual fibers, the larger the pennation
angles, the greater the reduction in force development
(11, 12, 14, 15). Larger fascicle angles at the extended
positions could, therefore, cause more mechanical disad-
vantage in the force transmission. However, present
results show that the largest fascicle angle is 21° in the
tensed condition; thus the loss of force during transmis-
sion would be 7% (5 cos21°) at the most. Considering
also the accuracy of the present fascicle angle measure-
ment, one could not put physiological significance on
such a small result from the present study. Similarly, in
the stance phase of walking, the knee joint moves
between 40 and 0° of flexion (22). According to the
present results, in this range of motion, the fascicle
angle of the vastus lateralis muscle changes from 17 to
18° and from 18 to 21° in the relaxed and tensed
conditions, respectively. The cosine of these angles then
ranges from 0.93 to 0.96. Thus the expected loss of the
force exerted by muscle fibers when it is transmitted to
tendon would be ,4–7%. Considering, again, the accu-
racy of present measurements of fascicle angle, the
impact of pennation angles on force transmission and
the difference between the relaxed and tensed condi-
tions would not be significant. However, the fascicle
length showed larger variance between relaxed and
tensed conditions, and this variance, possibly that of
fascicle angles as well, would be larger if muscle
activation were greater (e.g., MVC), in which case the
difference between the two conditions could have func-
tional importance. Care must be taken when estimat-
ing changes in muscle fiber length from the joint angle
changes by using the data of the embalmed cadavers (2,
6, 15).
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357
In the present study, the length and angle of fascicles
were determined from the midbelly of the vastus latera-
lis muscle at the level of 50% of the thigh length.
Although it has been shown that there is marked
uniformity of fiber length throughout a muscle (20),
some studies have reported heterogeneity of fascicle
angles along the length of the muscle (10, 19). Thus the
relationships between joint angles and fascicle arrange-
ment might differ in the different portions of the
muscle, which remain to be determined. Besides, in
this study, only the muscle activity at 10% MVC was
tested in the tensed condition. Architectural behavior of
fascicles would change at different activation levels, a
fact that also deserves further studies.
To date, architectural data of human skeletal muscles
have been derived mainly from cadaver specimens
(1–3, 6, 7, 11, 15, 20, 21, 23). The present results clearly
show that the architecture of actively contracting muscle
fibers differs considerably from that in which the
movement is passively induced. Although caution is
certainly merited in interpreting the present results, it
should be pointed out that the use of the architectural
data obtained in vivo would greatly contribute to the
understanding of contracting human muscles.
Address for reprint requests: Y. Kawakami, Dept. of Life Science
(Sports Science), The Univ. of Tokyo, Komaba 3–8–1, Meguro, Tokyo
153, Japan (E-mail: kawakami@idaten.c.u-tokyo.ac.jp).
Received 16 January 1996; accepted in final form 10 September 1996.
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