ABSTRACT: The mechanisms governing the increases in force production

in response to short periods of strength training have yet to be fully eluci-

dated. We examined whether muscle architectural adaptation was a con-
tributing factor. Ultrasound imaging techniques were used to measure quad-
riceps muscle architecture at 17 sites in vivo in trained and untrained legs of
men and women after 2.5 and 5 weeks of unilateral knee extension training,
as well as in a nontraining control group. Despite increases in knee extensor
strength of the trained and untrained (women only) legs, there were no
changes in muscle thickness, fascicle angle, or fascicle length in any of the
muscles tested. The moderate correlation between vastus lateralis thickness
(middle site) and eccentric (r ⫽ 0.55; P ⬍ 0.05) and concentric (r ⫽ 0.46; P ⬍
0.1) torque after, but not before, training is suggestive of neural rather than
architectural adaptations predominating in the early phase of training.
Muscle Nerve 35: 78 – 86, 2007

LACK OF HUMAN MUSCLE ARCHITECTURAL

ADAPTATION AFTER SHORT-TERM
STRENGTH TRAINING
ANTHONY J. BLAZEVICH, PhD,1 NICHOLAS D. GILL, PhD,2
NATHAN DEANS, BSc,3 and SHI ZHOU, PhD3

1
Centre for Sports Medicine and Human Performance, Brunel University,
Kingston Lane, Uxbridge UB8 3PH, United Kingdom
2
School of Sport and Exercise Science, Waikato Institute of Technology,
Hamilton, New Zealand
3
School of Exercise Science and Sport Management,
Southern Cross University, Lismore, Australia

Accepted 9 August 2006

A muscle’s phenotype is strongly governed by its
architecture or fiber geometry. Muscles with long
fibers (or fascicles) generate forces over greater
ranges of length and at higher shortening velocities,
but at greater metabolic cost than muscles with short
fibers. Moreover, muscles whose fibers insert at large
angles onto the tendon or aponeurosis have a large
physiological cross-sectional area relative to their vol-
ume and are therefore capable of generating large
relative peak forces. Using ultrasound and magnetic
resonance imaging techniques to measure muscle
architecture in vivo, researchers have shown that
significant increases in fascicle angle occur with in-
creases in muscle cross-sectional area or thickness in
This article includes Supplementary Material available via the internet at
http://www.mrw.interscience.wiley.com/suppmat/0148-639X/suppmat/
Abbreviations: ICC, intraclass correlations; MANOVA, multiple analyses of
variance; RM ANOVA, repeated-measures multiple analysis of variance
Key words: exercise; fascicle angle; fascicle length; isokinetic; muscle hy-
pertrophy
Correspondence to: A. J. Blazevich; e-mail: anthony.blazevich@brunel.
ac.uk
© 2006 Wiley Periodicals, Inc.
Published online 12 October 2006 in Wiley InterScience (www.interscience.
wiley.com). DOI 10.1002/mus.20666
response to prolonged periods (⬎14 weeks) of
heavy, isotonic/isoinertial weight training in previ-
ously untrained human subjects.1,17 These increases
in fascicle angle allow a greater muscle physiological
cross-sectional area and maximum force capacity;
the increase in fascicle angle reported after pro-
longed heavy strength training is thought to be one
mechanism by which muscle force increases after
training. Thus, muscle architecture appears to be
highly plastic, and these changes are thought to
contribute to the changes in strength expression
seen in weight-trained subjects.
Although early (e.g., ⬍4 weeks) increases in
strength in previously untrained subjects are accom-
panied by significant increases in muscle contractil-
ity, often little or no observable muscle hypertrophy
occurs.3,23,26 This has led to general acceptance of
the concept that short-term strength increases are
neurally mediated. However, muscle architectural
adaptations occur after 5 weeks in athletes who cease
heavy weight training but increase their speed-type
training.5 This result suggests that muscle architec-
ture change may occur rapidly in humans. In ani-
mals, immobilization of muscle or muscle fibers in a

78 Lack of Muscle Adaptation MUSCLE & NERVE January 2007

lengthened position for periods of days to weeks has
resulted in increases in the number of serially ar-
ranged sarcomeres and overall fiber length,28,29,33
whereas moderate stretching performed for only 30
min.day⫺1 was enough to reduce or reverse the fiber
shortening that accompanied short-length immobi-
lization in rat muscle.32 Also, variations in muscle
loading (eccentric vs. concentric muscle actions) in
rats have produced significant changes in the num-
ber of sarcomeres in series within 3 days of a 5-day
exercise stimulus.21 Despite the data demonstrating
a rapid architectural adaptation in animals, studies
investigating muscle architectural adaptation in hu-
mans have not ascertained its temporal response.
Thus, insufficient information is available with which
to build a complete picture of gross muscle adapta-
tion in response to muscle loading in humans. In
order to test the hypothesis that increases in muscle
fascicle angle occur after short-term strength train-
ing in previously non–strength-trained adults, we
measured architectural characteristics at proximal,
middle, and distal sites in the four heads of the
quadriceps femoris in trained legs of men and
women after 2.5 and 5 weeks of isokinetic strength
training.
MATERIALS AND METHODS
Subjects. Thirty-two subjects gave their written in-
formed consent to participate in the study, and 29
subjects consisting of 14 men [age, 20.6 ⫾ 2.6 (SD)
year; height, 1.80 ⫾ 0.09 m; weight, 76.0 ⫾ 1.3 kg]
and 15 women (age, 19.9 ⫾ 3.4 year; height, 1.70 ⫾
0.39 m, weight; 64.6 ⫾ 0.4 kg) completed the study;
3 subjects withdrew for reasons not pertaining to the
study. The research was approved by the institutional
review board. The subjects had no known musculo-
skeletal disorders, inflammatory conditions, or re-
cent injury that would preclude participation. None
of the subjects had performed resistance training
previously, but they were all recreationally active.
Overview. The subjects were randomly assigned to
either training (N ⫽ 15; 7 men and 8 women) or
control (N ⫽ 14; 7 men and 7 women) groups. Prior
to, and after 2.5 and 5 weeks of unilateral isokinetic
knee extension training, maximal concentric and
eccentric knee extensor torque was measured using
an isokinetic dynamometer (Kin-Com, Chattanooga
Group, Hixson, Tennessee), and muscle architec-
ture (muscle thickness, fascicle angle, and fascicle
length) of quadriceps femoris was quantified in both
trained and untrained legs, as described below. Con-
trol subjects performed no training but continued
Lack of Muscle Adaptation
FIGURE 1. Training volume performed in each session over 5
weeks. Vertical line indicates training performed before the mid-
training testing (4 sets ⫻ 6 repetitions) and after (5 sets ⫻ 6
repetitions).
their normal daily activities. They were assessed for
knee extensor strength and muscle architecture on
the same days as the training subjects. All testing was
performed at the same time of day to minimize
diurnal effects.
Training. Maximal isokinetic, concentric– eccentric
knee extension training of the right leg of the train-
ing subjects was performed at a velocity of 60°.s⫺1
three times a week for 5 weeks. Sessions were sepa-
rated by at least 1 day. Allowing for testing days after
2.5 weeks of training, subjects performed 10.5 ⫾ 1.2
of a possible 12 training sessions over the training
period. In the first 2.5 weeks of training (6.0 ⫾ 0.9
sessions), four sets of six maximal concentric– eccen-
tric knee extensions were performed through a
range of motion from 100° of knee flexion to full
extension (⬃0 –10°). In the second 2.5 weeks (4.3 ⫾
0.8 sessions), five sets of six repetitions were com-
pleted. Rest for 1 minute was allowed between sets.
These sessions are similar in intensity and volume to
those used by Kawakami et al.,17 whose subjects
trained at maximum intensity for five sets of eight
repetitions of a “French press” exercise; thus differ-
ences in training are limited largely to the duration
of training (5 weeks vs. 16 weeks). The present sub-
jects always performed a 5-min cycle warm-up with a
0.5 kg (women) or 1 kg (men) load at 60 rev.min⫺1
before training and no stretching was allowed before
or after the training. In order to ensure that maxi-
mum torque was achieved, straps were placed se-
curely across the subjects’ waists and torsos to mini-
mize extraneous movement, and loud verbal
encouragement and performance feedback was
given for all sets. Training volumes for each training
session are shown in Figure 1.
MUSCLE & NERVE January 2007 79
Testing. Knee Extension Torque. Warm-up in-
cluded a 5-min cycle (as above) and three practice
knee extension trials in which subjects were in-
structed to produce torques of approximately 50%,
70%, and 90% of their perceived maximum. After a
1-min rest, maximal concentric and eccentric torque
of the right and left legs of both trained and un-
trained subjects was measured on the isokinetic dy-
namometer at an angular speed of 60°.s⫺1 with the
hip angle at 85°. The knee of the exercising leg was
aligned approximately 2 cm anterior to the axis of
rotation of the dynamometer. This ensured the lat-
eral condyle of the femur was in line during maxi-
mum efforts when the small deformation of the
chair and padding on the attachment occurred. Sub-
ject positioning was adjusted if misalignment of the
lateral condyle and axis of rotation occurred during
the most maximal (i.e., the third) practice trial. After
two to three maximal practice trials, the subjects
were allowed three maximal trials to register concen-
tric or eccentric peak torque with the knee moving
between 100° of flexion and full extension (⬃0 –
10°). Subjects were verbally encouraged during the
testing, and a 2-min rest was allowed between testing
of concentric (always tested first) and eccentric
torque. In order to prevent muscle adaptation prior
to the commencement of the study and to allow task
learning to contribute to overall strength increases,
subjects did not perform familiarization sessions.
Thus, any increases in peak torque in the training or
control subjects could be partly explained by learn-
ing of the movement, whereas any architectural
changes observed will have occurred from a true
“nontraining” starting point. Peak concentric and
eccentric torque was recorded for analysis.
Measurement of Muscle Thickness and Fascicle Angle.
Muscle architecture was examined in vivo using B-
mode ultrasonography (Acuson Sequoia; Acuson,
Mountain View, California) with a 3.8-cm, 7.5-MHz
linear-array probe. Subjects lay supine with knees
flexed (but stationary) to 45°. Pilot testing had
shown that moderate knee flexion reduced fascicle
curvature and improved the reliability of repeated
measurements, but still allowed significant interindi-
vidual variation to be seen. The probe was coated
with a water-soluble transmission gel to aid acoustic
contact and remove the need for the probe to make
contact with the skin. Thus, no pressure was applied
to the muscle that might affect muscle thickness and
fascicle angle measurements. Scans were taken with
the probe oriented along the sagittal plane of the
fascicles and perpendicular to the skin.6 The angle
of the probe relative to the longitudinal axis of the
thigh therefore varied between subjects. Appropriate
80 Lack of Muscle Adaptation
FIGURE 2. Marks were made on the thigh at distances of 5%,
22%, 39%, 56%, and 73% of the distance from the superior
border of the patella to the anterior superior iliac spine with
subjects lying supine. Each muscle on the trained (right) leg was
scanned at three sites. These corresponded to sites A, B, and C
for vastus medialis (VM); B, C, and D for both the vastus lateralis
(VL) and the frontal plane of vastus intermedius (not shown); and
C, D, and E for both the rectus femoris (RF) and the sagittal plane
of vastus intermedius. The untrained (left) leg was scanned at
site C for vastus lateralis and the frontal plane of vastus interme-
dius. TFL, tensor fascia latae; Add., adductor muscles.
probe alignment was achieved when several fascicles
could be traced without interruption across the im-
age (i.e., they did not run out of the plane of the
sonogram). Scans were taken at three sites on each
of vastus lateralis, rectus femoris, and vastus medialis
muscles of the right (trained) leg, and at three sites
on vastus intermedius with the probe placed over
both vastus lateralis (frontal plane scan) and rectus
femoris (sagittal plane scan) of the right (trained)
leg. Scans were also taken of vastus lateralis and
vastus intermedius at a single site on the left (un-
trained) leg with the probe placed over vastus late-
ralis (i.e., frontal plane scan of vastus intermedius).
Scanning sites are shown in Figure 2; scans were
taken from a total of 17 sites in both training and
control subjects (1,479 images). An example scan
image is shown in Figure 3. After pre-training scan-
ning, the ultrasound scanning sites were mapped
onto a flexible, transparent plastic sheet to ensure
accuracy in repeated scanning.
Ultrasound images were imported into image
digitizing software (Peak Motus, Peak Technologies,
Centennial, Colorado) and landmarks pertaining to
the aponeuroses and muscle fascicles were digitized
as shown in Figure 3. For fascicles, the origin of the
fascicle was digitized at a point approximately 3– 4
mm from the deep aponeurosis and the end was
digitized at the mid-level (Fig. 3). Fascicle curvature
was imperceptible throughout most of the muscles,
MUSCLE & NERVE January 2007
FIGURE 3. Sagittal plane ultrasound image of the vastus lateralis/vastus intermedius muscles, as in image (a). Image landmarks
corresponding to aponeuroses (points 1– 4) and fascicles (points 5– 6) were digitized using appropriate software (see Methods), as in
image (b). Muscle thickness was calculated as the mean of the vertical distances between points 1–3 and 2– 4. The fascicle angle,
measured from 3– 4 mm above the deep aponeurosis (dashed line) to mid-muscle was calculated as the angle between points 2– 4 and
5– 6, while aponeurosis angle was calculated as the angle between points 3– 4 and 1–2 (counterclockwise).

but there was a tendency for fascicles to curve at the
insertion onto the deep aponeurosis. In other im-
ages, the insertion point was invisible. Thus, measur-
ing from approximately 3– 4 mm above the aponeu-
rosis allowed accurate delineation of the fascicles.6
Muscle thickness was calculated as the mean of the
distances between superficial and deep aponeuroses
measured at each edge of the ultrasound image.
Fascicle angle was calculated as the positive angle
between the deep aponeurosis and the line of the
fascicle. Each image was digitized twice on separate
occasions and the values for muscle thickness and
fascicle angle compared. Image pairs were re-digi-
tized separately if muscle thickness measures dif-
fered by more than 2 mm or fascicle angle differed
by more than 1.
Fascicle Length Estimation. As fascicles were usu-
ally too long to be fully captured by our probe,
fascicle length was estimated using eq. (1), where
MT is the muscle thickness, ␪ is the fascicle angle,
and ␥ is the aponeurosis angle.
Fascicle length ⫽ sin( ␥ ⫹ 90°) ⫻
MT/sin(180° ⫺ [␥ ⫹ 180° ⫺ ␪])
Fascicle length was only estimated at the mid-section
of each muscle for two reasons: (1) estimates using
this technique assume that no fascicle or aponeuro-
sis curvature exists, which appeared to be true only at
the mid-point of the muscles, and (2) these tech-
niques assume that muscle architecture is relatively
isotropic, which is not true toward the end of mus-
cles where they generally change shape in response
to space constraints. Fascicle length estimates for
vastus medialis proved to be unreliable and were
thus not included in analyses. Similar methods of
estimating fascicle length when the entire fascicle
cannot be visualized have been used previously, with
errors of approximately 2%–7%.11,12 These small er-
rors contribute to a slightly lower reliability of length
measures compared to muscle thickness and fascicle
angle, as described below.
Physiological Muscle Thickness Calculation. As
changes in fascicle angle would alter the distance
between the aponeuroses measured perpendicular
to the fascicles (i.e., the physiological thickness), this
distance was calculated by eq. (2), where MT is the
muscle thickness and ␪ is the fascicle angle.
Physiological thickness
⫽ 公(MT 2 ⫹ [tan ␪ .MT] 2) (2)
Reliability of Muscle Architecture Measurements.
(1) Reliability was determined for: (1) the digitizing of
images at different muscle sites, and (2) the repeat-
ability of the entire procedure, including location of
imaging sites, acquisition and digitization of images,
and calculation of architectural parameters at each
muscle’s midpoint. Data for calculation of digitizing
reliability were available since all images were digi-

Lack of Muscle Adaptation MUSCLE & NERVE January 2007 81

tized twice. For each digitizing site, 50 images were
randomly selected and typical errors (TE; SDdiff/
公2) and intraclass correlations [ICC; eq. (3)] were
calculated, where SDdiff is the standard deviation of
the difference between measurements 1 and 2, and s
is the parameter variance [of trial set 1 (s1) and trial
set 2 (s2), respectively].
ICC ⫽ 1 ⫺ TE 2/([s 1 ⫹ s 2]/2) (3)
Digitization reliability was assessed at each site on all
muscles of the trained leg for muscle thickness and
fascicle angle. Reliability of repeated measurements
of the whole process was calculated using the same
procedures.
Statistical Analysis. Pre-training differences between
training and control groups were analyzed by multiple
analyses of variance (MANOVA). Changes in peak con-
centric and eccentric knee extensor torque, and
changes in muscle architectural measures, were exam-
ined by separate repeated-measures multiple analysis
of variance (RM ANOVA) with muscle site (proximal,
middle, and distal) and time (0, 2.5, and 5 weeks) as
within-subject factors and training group as the be-
tween-subjects factor. Significant main or interaction
effects were further analyzed by Bonferroni univariate
tests. Normality of the data was checked using Kolmog-
orov–Smirnov’s test, and equality of error variance was
assessed and confirmed using Levene’s test of error
variance. The linear relationship between torque and
muscle architecture variables were assessed using Pear-
son’s product moment correlations after curve-fitting
procedures were applied to ensure nonlinear relation-
ships did not exist between the variables. The alpha
level was set at 0.05 for all tests. Values are presented as
means ⫾ SD.
RESULTS
Reliability of Muscle Architecture Measurements.
The reliability of digitizing of repeated images was
high for muscle thickness and fascicle angle. Intra-
class correlations ranged 0.948 –1.000, with some
variation between muscles and muscle sites. Typical
errors ranged 0.08 – 0.63 mm and 0.27– 0.63°, respec-
tively. Repeated measurements incorporating the
mapping of measurement sites, acquisition of images
and digitizing those images were also very reliable
with intertrial reliabilities (ICC) of 0.882– 0.970,
0.899 – 0.991, and 0.758 – 0.863 (typical errors of
0.74 – 0.97 mm, 0.24 –1.22°, and 10.2–19.4 mm) for
muscle thickness, fascicle angle, and fascicle length,
respectively.
82 Lack of Muscle Adaptation
FIGURE 4. Knee extensor torque in men (a), and women (b) for
training and control subjects’ right legs. Trends among men and
women were similar, although men produced more torque both
concentrically and eccentrically. Open circles, concentric training;
filled triangles, eccentric training; solid lines, training group; dot-
ted lines, control group.
Knee Extensor Torque. There were no changes in
knee extensor strength of the control group in ei-
ther right or left legs. In the training group, there
was a moderate but significant increase in concentric
strength of the right (trained) leg (184.1 ⫾ 41.4 to
200.0 ⫾ 34.9 Nm, ⌬ ⫽ 8.7%; P ⬍ 0.05) and a large
and statistically significant change in eccentric
strength (216.4 ⫾ 51.8 to 309.5 Nm ⫾ 94.7 Nm, ⌬ ⫽
42.8%; P ⬍ 0.01). In the left (untrained) leg, there
was no change in concentric strength (182.6 ⫾ 44.9
to 183.7 ⫾ 32.9 Nm, ⌬ ⫽ 0.6%), although there was
a near-significant increase in eccentric strength
(230.3 ⫾ 49.5 to 248.7 ⫾ 68.6 Nm, ⌬ ⫽ 8.0%; P ⬍
0.1).
In order to determine whether gender effects
influenced the results, the strength responses of
men and women were also compared. There were no
significant differences in the overall, or relative,
strength changes between men and women. None-
theless, as shown in Figure 4, the change in concen-
tric strength of the women’s right (trained) leg was
statistically significant (145 ⫾ 30.1 to 159.6 Nm ⫾
20.6, ⌬ ⫽ 9.9%; P ⬍ 0.05), whereas it only ap-
proached significance in men (217.4 ⫾ 54.0 to
235.1 ⫾ 53.3 Nm, ⌬ ⫽ 8.1%; P ⬍ 0.08). In contrast,
eccentric strength improved significantly for both
genders (women, 175.7 ⫾ 35.3 to 256.0 ⫾ 50.2 Nm,
MUSCLE & NERVE January 2007
 Table 1. Vastus lateralis architecture (N ⫽ 29).
Muscle thickness Fascicle angle Fascicle length
Control Training Control Training Control Training
Mean Mean Mean Mean Mean Mean
Week Site (mm) SD (mm) SD (deg) SD (deg) SD (mm) SD (mm) SD
0 Distal 20.9 3.5 21.9 3.4 9.4 4.1 9.1 4.1
Mid 21.9 3.2 23.2 3.5 13.9 1.7 12.8 4.4 91.1 13.1 127.6 66.3
Proximal 14.1 2.9 18.5 5.3 11.7 3.7 12.3 3.7
2.5 Distal 21.9 3.1 21.5 2.9 9.4 4.6 9.1 4.0
Mid 22.3 2.6 23.4 3.5 12.4 3.1 12.6 1.8 108.2 16.8 103.2 27.9
Proximal 16.1 5.6 17.9 4.6 12.2 4.2 13.5 3.6
5 Distal 20.1 2.9 22.4 3.1 9.7 3.1 10.1 3.5
Mid 22.0 4.3 23.9 2.8 13.1 2.4 12.8 2.4 102.7 17.9 108.2 24.6
Proximal 13.7 2.7 17.3 5.4 12.3 3.8 12.4 2.2

⌬ ⫽ 45.7%; men, 252.0 ⫾ 68.0 to 361.9 ⫾ 134.6 Nm,
⌬ ⫽ 43.6%). In the untrained leg, there was no
change in concentric or eccentric knee extensor
torque in men, although there was a trend toward
greater concentric strength after 2.5 weeks but no
further increase by 5 weeks. In women, there was
small and nonsignificant increase in concentric
strength (8.7%; P ⫽ 0.25) after 5 weeks, but a signif-
icant increase in eccentric strength (41.1%; P ⬍
0.01). Thus, the response of men and women was
not the same in the untrained leg.
Muscle Architecture. There were no changes in
muscle size, fascicle angle, or estimated fascicle
length at any of the 15 sites tested on the trained leg,
or in the middle of vastus lateralis or vastus interme-
dius of the untrained leg in trained subjects. Pre- and
post-training muscle architecture results for vastus
lateralis are shown in Table 1 as an example; results
for the other 14 sites are presented as Supplemen-
tary Tables on the Journal’s website. There was a
strong correlation (r2 ⬎ 0.90) between muscle thick-
ness and physiological muscle thickness that did not
change with training; therefore physiological muscle
thickness data have not been presented. Before
training, there were no differences between men
and women for architectural parameters, except that
the fascicle angle in the proximal region of vastus
medialis was slightly greater for men than for women
(pre-training men ⫽ 11.5 ⫾ 2.9°; women ⫽ 8.7 ⫾
2.6°; P ⬍ 0.05). This pre-training difference per-
sisted consistently through the duration of the study,
so our subjects were considered homogeneous for
our purpose.
DISCUSSION
We investigated the effect of 5 weeks of unilateral,
isokinetic strength training on knee extension
torque and quadriceps muscle architecture in
trained and untrained legs. The training group sig-
nificantly improved their knee extensor strength in
the trained leg in response to the 5-week training
program. The improvement in eccentric strength
(42.8%) was greater than concentric (8.7%)
strength. The results are consistent with previous
reports of greater increases in eccentric strength
after periods of isokinetic strength training.19 The
results are suggestive that a greater capacity for ec-
centric adaptation exists, which is borne out during
isokinetic training where both concentric and eccen-
tric contractions are performed maximally. Absolute
and relative strength increases in the trained leg of
men and women were the same, so gender effects
did not influence the results. Strength changes in
the untrained leg were smaller. There was no change
in concentric strength after the 5-week training pe-
riod (⌬ ⫽ 0.6%); a slight mean increase in eccentric
strength did not reach statistical significance (⌬ ⫽
8.0%; P ⬍ 0.1). The responses of men and women
were slightly different, however. There was no
change in knee extensor torque of the untrained leg
in men, whereas in women there was a small and
nonsignificant increase in concentric strength
(8.7%; P ⫽ 0.25) after 5 weeks and a significant
increase in eccentric strength (41.1%; P ⬍ 0.01).
The lack of contralateral strength changes in the
men is not an uncommon finding,24 but the condi-
tions under which this lack of response occurs are
unclear.
However, a major finding of this study was that,
despite significant increases in knee extensor torque
of the trained and untrained (women only) legs, no
changes occurred in muscle thickness, fascicle angle,
or fascicle length at any of the 17 ipsi- and contralat-
eral sites. Thus, the strength changes resulting from
the training cannot be explained by architectural

Lack of Muscle Adaptation MUSCLE & NERVE January 2007 83

adaptation. Although muscle cross-sectional area or
volume was not measured in the present experi-
ments, the lack of change in superficial-to-deep mus-
cle thickness or of any change in fascicle angle or
length suggests that muscle volume changes were
small or zero. Given that increases in superficial-to-
deep muscle thickness have been previously shown
to accompany changes in muscle volume17 and that
no change in muscle thickness was shown at any of
the 17 muscle sites, we feel it is unlikely that any
increases in muscle size occurred. A lack of increase
in muscle size in response to short-term loading in
previously non–strength-trained subjects has been
noted in earlier studies.3,23,25
We also tested the hypothesis that rapid changes
in fascicle angle and length do not occur after short
periods of strength training. This hypothesis has
never been tested in humans despite significant sci-
entific data suggesting the possibility that adapta-
tions might occur rapidly in both humans and other
animals. For example, fascicle angle and length
changes occurred after 5 weeks in athletes who sig-
nificantly altered their training.5 Although the spe-
cific changes in gene expression and protein synthe-
sis that underpin architectural changes are not
known, muscle protein synthesis is upregulated im-
mediately following muscle loading and stretch stim-
uli,8,9,22,31 which occurs subsequent to rapid in-
creases in gene expression.8 Also, rapid muscle-
specific gene expression changes have been shown
after single bouts of eccentric contractions in mice,4
and after 3 days of gastrocnemius overload resulting
from ablation of synergists in rats.7 Furthermore,
animal studies provide compelling evidence that
rapid architectural adaptations are possible. Repeti-
tive stretching of rat gastrocnemius,15 overload of
the plantaris muscles by ablation of synergists,2 and
percutaneous stimulation of tibialis anterior and ex-
tensor digitorum longus during muscle lengthen-
ing27 have resulted in significant increases in the
activity of myogenic regulatory factors within hours.
These reports are consistent with the rapid changes
in fiber lengths that were shown in immobi-
lized20,28,29,33 and passively stretched32 muscles of var-
ious animals. Thus, rapid muscle adaptation very
likely occurs under the appropriate loading condi-
tions, and, at least in animals, these adaptations oc-
cur with a time course similar to changes in muscle
architecture. Regardless of the rapid increases in
protein synthesis in animals and humans, and the
rapid changes in muscle architecture shown in ani-
mals, there were no observable changes in fascicle
angle or length of the trained leg in the present
study. As discussed below, the precision and sensitiv-
84 Lack of Muscle Adaptation
ity of our measurements were such that very small
changes in architecture that were undetectable
would not have been functionally significant. Our
results, therefore, do not support the hypothesis that
fascicle adaptations occur rapidly in response to
strength training.
We also examined changes in the untrained leg.
Unilateral immobilization of limbs in rats13 has been
associated with significant changes in muscle archi-
tecture of nonimmobilized limbs when compared to
control rats where no immobilization was used, al-
though it is possible that altered activity patterns
could account for these changes.14 Also, Yasuda and
Miyamura34 showed a greater muscle blood flow to
the nonexercising arm during an endurance hand-
grip task after 3 and 6 weeks of unilateral training.
Eklund et al.10 had previously shown that the in-
crease in blood flow to a nonexercising limb was not
a result of inadvertent muscle activation. Such a
phenomenon would allow circulating anabolic hor-
mones to perfuse the nonexercised muscle and
make remote muscle adaptation theoretically possi-
ble. Thus, physiological adaptations to unilateral ex-
ercise seem to be worthy of further consideration,
and only one study has examined architecture in a
nontraining limb.16 Nonetheless, we found no evi-
dence of architectural adaptation in the untrained
leg. The lack of change is consistent with the results
of Kim et al.18 who did not find an increase in resting
myofibrillar or mixed protein synthesis rates in vas-
tus lateralis of an untrained leg, despite significant
increases in the trained leg. Our data are also in
agreement with those of Kawakami et al.17 who did
not find changes in the untrained (control) arm in
subjects after 16 weeks of strength training. Thus,
there is presently no evidence that architectural ad-
aptation occurs in an untrained limb after training
of the contralateral limb. As suggested by other au-
thors,35 the small increases in strength of the con-
tralateral limb in the present study possibly resulted
from spinal (e.g., alterations in reflex sensitivity and
afferent feedback) or supraspinal adaptations (e.g.,
diffusion of impulses to the contralateral motor cor-
tex during unilateral training), or improved postural
stabilization and coordination.
Given the lack of architectural change shown
presently, we closely examined the precision of our
measurements. The validity for using ultrasound
techniques to assess in vivo muscle architecture has
been shown previously by comparing ultrasound-
and cadaver-based measurements.16 The precision
and linearity of image reconstruction has also been
shown.15 The sensitivity of measures is difficult to
ascribe, but it is likely that positive changes of greater
MUSCLE & NERVE January 2007
than 1° of fascicle angle and 5 mm of fascicle length
could be detected for the muscles analyzed in this
study, if those changes occurred; changes smaller
than this would not be functionally significant. We
also used strict methodologies including the map-
ping of measurement sites for high accuracy in the
repeated placement of the ultrasound transducer,
double-digitizing of all images to minimize calcula-
tion errors, and the undertaking of all digitizing by a
single operator to remove multiple-operator unreli-
ability. We measured the reliability of both: (1) our
digitizing procedure, and (2) the reliability of the
complete procedure including the mapping of mea-
surement sites, acquisition of images, and digitizing
those images. With respect to digitizing, repeated
measures were highly accurate with intraclass corre-
lations (ICC) of 0.948 –1.000 and typical errors for
muscle thickness and fascicle angle of 0.08 – 0.63 mm
and 0.27– 0.63°, respectively. Moreover, repeated
measurements of the whole procedure were also very
reliable. We have obtained intertrial reliabilities
(ICC) of 0.882– 0.970, 0.899 – 0.991, and 0.758 –
0.863 (typical errors of 0.74 – 0.97 mm, 0.24 –1.22°,
and 10.2–19.4 mm) for muscle thickness, fascicle
angle, and fascicle length, respectively. We believe
these reliabilities are very high, although there is
some reduction in reliability of the fascicle length
estimates, as would be expected given that errors are
a function of the small errors in muscle thickness,
fascicle angle, and aponeurosis angle. Particularly
for muscle thickness and fascicle angle measure-
ments, errors of the magnitude of our typical errors
are not likely to be functionally significant, so the
lack of change in muscle architecture cannot be
explained by measurement unreliability. Thus, the
strength changes observed in our trained subjects
must have resulted from mechanisms other than
architectural adaptation.
In order to further explore the mechanisms re-
sponsible for the rapid increases in strength, the
relationships between initial muscle architecture
variables and the magnitude of strength change were
examined. Given that relative strength increases in
the trained subjects were similar for men and
women, and that there were no between-gender dif-
ferences in muscle thickness, fascicle angle, or fasci-
cle length [with the single exception that fascicle
angle at the proximal part of vastus medialis was
greater in men (11.5 ⫾ 2.9°) than women (8.7 ⫾
2.6°)], the data for men and women were pooled for
further analysis. Before training, there was no differ-
ence in muscle thickness, fascicle angle, or fascicle
length of the trained leg in subjects with greater-
than-median or less-than-median knee extensor
Lack of Muscle Adaptation
strength, and no correlation between strength and
muscle architecture measures. After training, how-
ever, there was a stronger correlation between vastus
lateralis thickness at mid-muscle (i.e., 39% of dis-
tance from lateral condyle) and torque developed
eccentrically (r ⫽ 0.55; P ⬍ 0.05) and concentrically
(r ⫽ 0.46; P ⬍ 0.1). Thus, subjects who had greater
thickness of the vastus lateralis muscle prior to train-
ing produced greater knee extensor torque after
training. Given that vastus lateralis accounts for a
substantial proportion of total quadriceps femoris
volume (e.g., 36%30), it seems possible that intersub-
ject size differences might have impacted on force
development. It is possible then that strength in-
creases occurred largely by an alteration in muscle
recruitment strategies where those with larger vastus
lateralis muscles benefited most clearly. Although
muscle activation was not measured in the present
study, rapid alterations in muscle recruitment in
response to resistance training have been shown pre-
viously.23
In summary, significant increases in knee exten-
sor strength of previously untrained ipsi- and con-
tralateral (women only) legs occurred without
changes in muscle architecture. These data form an
important part of the picture in understanding the
early adaptations to muscle loading in humans.
Given that there was a stronger correlation between
vastus lateralis thickness and knee extensor torque
after training, the rapid increase in strength can
probably be explained by alterations in muscle re-
cruitment strategies, with those subjects having
larger muscles prior to training realizing the greatest
strength improvements after training.
The authors thank the staff at North Coast Radiology, New South
Wales, Australia for their assistance through the duration of the
study.
REFERENCES
1. Aagaard P, Andersen J, Dyhre-Poulsen P, Leffers A-M, Wagner
A, Magnusson SP, et al. A mechanism for increased contrac-
tile strength of human pennate muscle in response to
strength training: changes in muscle architecture. J Physiol
(Lond) 2001;534:613– 623.
2. Adams GR, Haddad F, Baldwin KM. Time course of changes
in markers of myogenesis in overloaded rat skeletal muscles.
J Appl Physiol 1999;87:1705–1712.
3. Akima H, Takahashi H, Kuno SY, Masuda K, Masuda T,
Shimajo H, et al. Early phase adaptations of muscle use and
strength to isokinetic training. Med Sci Sports Exerc 1999;31:
588 –594.
4. Barash IA, Mathew L, Ryan AF, Chen J, Lieber RL. Rapid
muscle-specific gene expression changes after a single bout of
eccentric contractions in the mouse. Am J Physiol 2004;286:
C355–C364.
MUSCLE & NERVE January 2007 85
 5. Blazevich AJ, Gill ND, Bronks R, Newton RU. Training-spe-
cific muscle architecture adaptation after 5-wk concurrent
training in athletes. Med Sci Sports Exerc 2003;35:2013–2022.
6. Blazevich AJ, Gill ND, Zhou S. Intra- and inter-muscular vari-
ation in human quadriceps femoris architecture assessed in
vivo. J Anat 2006;209:289 –310.
7. Carson JA, Nettleton D, Reecy JM. Differential gene expres-
sion in the rat soleus muscle during early work overload-
induced hypertrophy. FASEB J 2002;16:207–209.
8. Chen Y-W, Hubal MJ, Hoffman EP, Thompson PD, Clarkson
PM. Molecular responses of human muscle to eccentric exer-
cise. J Appl Physiol 2003;95:2485–2494.
9. Chesley A, MacDougall JD, Tarnopolsky MA, Atkinson SA,
Smith K. Changes in human muscle protein synthesis after
resistance exercise. J Appl Physiol 1992;73:1383–1388.
10. Eklund B, Kaijser L, Knutsson E. Blood flow in resting (con-
tralateral) arm and leg during isometric contraction. J Physiol
(Lond) 1974;240:111–123.
11. Finni T, Ikegawa S, Komi PV. Concentric force enhancement
during human movement. Acta Physiol Scand 2001;173:369 –
377.
12. Finni T, Ikegawa S, Lepola V, Komi PV. Comparison of force-
velocity relationships of vastus lateralis muscle in isokinetic
and in stretch-shortening cycle exercises. Acta Physiol Scand
2003;177:483– 491.
13. Heslinga JW, Huijing PS. Muscle length-force characteristics
in relation to muscle architecture: a bilateral study of gastroc-
nemius medialis muscles of unilaterally immobilized rats. Eur
J Appl Physiol 1993;66:289 –298.
14. Heslinga JW, Rozendal RH, Huijing PA. Unilateral immobili-
zation affects contralateral rat gastrocnemius muscle architec-
ture. Acta Anat 1992;143:231–235.
15. Ikeda S, Yoshida A, Matayoshi S, Horinouchi K, Tanaka N.
Induction of myogenin messenger ribonucleic acid in rat
skeletal muscle after 1 hour of passive repetitive stretching.
Arch Phys Med Rehabil 2004;85:166 –167.
16. Kawakami Y, Abe T, Fukunaga T. Muscle-fiber pennation
angles are greater in hypertrophied than in normal muscles.
J Appl Physiol 1993;74:2740 –2744.
17. Kawakami Y, Abe T, Kuno S-Y, Fukunaga T. Training-induced
changes in muscle architecture and specific tension. Eur
J Appl Physiol 1995;72:37– 43.
18. Kim PL, Staron RS, Phillips SM. Fasted-state skeletal muscle
protein synthesis after resistance exercise is altered with train-
ing. J Physiol (Lond) 2005;568:283–290.
19. Lacerte M, deLateur BJ, Alquist AD, Questad KA. Concentric
versus combined concentric-eccentric isokinetic training pro-
grams: effect on peak torque of human quadriceps femoris
muscle. Arch Phys Med Rehabil 1992;73:1059 –1062.
20. Leterme D, Cordonnier C, Mounier Y, Falempin M. Influence
of chronic stretching upon rat soleus muscle during non-
weight-bearing conditions. Eur J Appl Physiol 1994;429:274 –
279.
86 Lack of Muscle Adaptation
21. Lynn R, Talbot JA, Morgan DL. Differences in rat skeletal
muscles after incline and decline running. J Appl Physiol
1998;85:98 –104.
22. MacDougall JD, Gibala MJ, Tarnopolsky MA, MacDonald JR,
Interisano SA, Yarasheski KE. The time course for elevated
muscle protein synthesis following heavy resistance exercise.
Can J Appl Physiol 1995;20:480 – 486.
23. Moritani T, DeVries HA. Neural factors versus hypertrophy in
the time course of muscle strength gain. Am J Phys Med
1979;58:115–130.
24. Munn J, Herbert RD, Gandevia SC. Contralateral effects of
unilateral resistance training: a meta-analysis. J Appl Physiol
2004;96:1861–1866.
25. Narici MV, Hoppeler H, Kayser B, Landoni L, Claassen H,
Gavardi C, et al. Human quadriceps cross-sectional area,
torque and neural activation during 6 mo strength training.
Acta Physiol Scand 1996;157:175–186.
26. Narici MV, Roi GS, Landoni L, Minetti AE, Cerretelli P.
Changes in forces, cross-sectional areas and neural activation
during strength training and detraining of the human quad-
riceps. Eur J Appl Physiol 1989;59:310 –319.
27. Peters D, Barash IA, Burdi M, Yuan PS, Mathew L, Fridén J, et
al. Asynchronous functional, cellular and transcriptional
changes after a bout of eccentric exercise in the rat. J Physiol
(Lond) 2003;553:947–957.
28. Shah SB, Peters D, Jordan KA, Milner DJ, Fridén J, Capet-
anaki Y, et al. Sarcomere number regulation maintained after
immobilization in desmin-null mouse skeletal muscle. J Exp
Biol 2001;204:1703–1710.
29. Tabary JC, Tabary C, Tardieu C, Tardieu G, Goldspink G.
Physiological and structural changes in the cat’s soleus muscle
due to immobilization at different lengths by plaster casts.
J Physiol (Lond) 1972;224:231–244.
30. Tesch PA, Ekberg A, Lindquist DM, Trieschmann JT. Muscle
hypertrophy following 5-week resistance training using a non-
gravity-dependent exercise system. Acta Physiol Scand 2004;
180:89 –98.
31. Trappe TA, Raue U, Tesch PA. Human soleus muscle protein
synthesis following resistance exercise. Acta Physiol Scand
2004;182:189 –196.
32. Williams PE. Use of intermittent stretch in the prevention of
serial sarcomere loss in immobilised muscle. Ann Rheum Dis
1990;49:316 –317.
33. Williams PE, Goldspink G. Changes in sarcomere length and
physiological properties in immobilized muscle. J Anat 1978;
127:459 – 468.
34. Yasuda Y, Miyamura M. Cross transfer effects of muscular
training on blood flow in the ipsilateral and contralateral
forearms. Eur J Appl Physiol 1983;51:321–329.
35. Zhou S. Chronic neural adaptations to unilateral exercise:
mechanisms of cross-education. Exerc Sport Sci Rev 2000;28:
177–184.
MUSCLE & NERVE January 2007