MUSCLE ARCHITECTURE ADAPTATIONS TO KNEE EXTENSOR

ECCENTRIC TRAINING: RECTUS FEMORIS VS. VASTUS LATERALIS

BRUNO MANFREDINI BARONI, MSc,1 JEAM MARCEL GEREMIA, MSc,1 RODRIGO RODRIGUES, PE,1

RODRIGO DE AZEVEDO FRANKE, PE,1 KIROS KARAMANIDIS, PhD,2,3 and MARCO AURELIO VAZ, PhD1
1
Exercise Research Laboratory, School of Physical Education, Federal University of Rio Grande do Sul, Porto Alegre, Brazil
2
Institute of Biomechanics and Orthopaedics, German Sport University of Cologne, Cologne, Germany
3
Faculty of Mathematics and Technology, RheinAhrCampus Remagen, Koblenz University of Applied Sciences, Koblenz, Germany
Accepted 9 January 2013

ABSTRACT: Introduction: Changes in muscle architecture
induced by eccentric knee extensor training remain unclear, as
well the adaptive responses of synergistic knee extensor
muscles with different geometrical designs. Methods: Ultraso-
nography images were taken from rectus femoris (RF) and
vastus lateralis (VL) of 20 male volunteers before and after a
non-training control period of 4 weeks, and additional evalua-
tions were performed after 4, 8, and 12 weeks of isokinetic
eccentric training. Results: RF and VL had significant changes
in muscle architecture within the first 4 training weeks, and the
adaptive response throughout the intervention was similar. Mus-
cle thickness increased by around 7–10%, fascicle length
increased 17–19%, and pennation angle was unchanged. Con-
clusions: Increased muscle thickness due to eccentric training
was related to increased fascicle length and not to pennation
angle changes. Although RF and VL have a different fascicular
geometry, they had similar morphological adaptations to eccen-
tric training.
Muscle Nerve 48: 498–506, 2013
The effects of eccentric resistance training on the
neuromuscular system have been evaluated in sev-
eral studies involving healthy subjects,1–12 conven-
tional resistance training practitioners (performed
with concentric and eccentric muscle actions),13–15
and athletes of different sports and competition
levels.16–18 To counteract the deleterious effects of
aging,19,20 this kind of exercise has been proposed
for the elderly population21–23 and has also been
used widely in rehabilitation programs of musculo-
skeletal injuries.24,25 Probably due to their clinical
importance for the locomotor system, knee exten-
sors are the most frequently assessed muscle
group, and significant increases in strength and
muscle mass due to eccentric quadriceps training
have been reported.1–4,6,7,10–12,22,23
The hypertrophic response has been commonly
assessed through measurement of muscle anatomic
cross-sectional area (ACSA) in resistance training
Abbreviations: ACSA, anatomical cross-sectional area; ANOVA, analysis
of variance; PCSA, physiological cross-sectional area; RF, rectus femoris;
VI, vastus intermedius; VL, vastus lateralis; VM, vastus medialis
Key words: eccentric exercise; fascicular geometry; muscle plasticity;
quadriceps femoris; ultrasonography
Correspondence to: B.M. Baroni; e-mail: bmbaroni@yahoo.com.br
This study was supported by the Coordenac¸a ~o de Aperfeiçoamento de
Pessoal de Nıvel Superior (CAPES Brazil), Financiadora de Estudos e Pro-
jetos (FINEP Brazil), and Conselho Nacional de Pesquisa (CNPq Brazil).
C 2013 Wiley Periodicals, Inc.
V allow for comparisons between VM and VL in an
Published online 30 January 2013 in Wiley Online Library
(wileyonlinelibrary.com). DOI 10.1002/mus.23785
studies.26,27 However, ACSA of pennate muscles,
such as most locomotor muscles (e.g., quadriceps
femoris), do not represent the perpendicular
cross-sectional area of all muscle fibers, that is,
physiological cross-sectional area (PCSA). The lat-
ter is considered to be the architectural parameter
directly proportional to the maximum tension gen-
erated by the muscle.28 Because PCSA is estimated
from muscle volume, fiber pennation angle, and
fiber length, changes in fascicular geometry have
significant effects on functionality of pennate
muscles.28 Therefore, fascicular geometric adapta-
tions of knee extensor muscles have been investi-
gated in several studies involving conventional
resistance training,22,23,29–33 but only a few eccen-
tric training studies.6,22,23
Eccentric training programs have shown signifi-
cant increases in muscle thickness after 10 training
weeks in young subjects6 and 14–16 training weeks
in elderly subjects.22,23 Some evidence has sug-
gested that the pennation angle increases,6
whereas others suggested that there are no
changes22,23 in response to eccentric training. Con-
troversial results can also be found in relation to
fascicle length adaptations.6,22,23 Moreover, the
time-course of muscle architecture adaptations
remains unclear, as the study by Blazevich et al.6
seems to be the only one to have shown muscle
architecture changes after a short period of quadri-
ceps femoris eccentric training (i.e., 5 weeks). In
other words, at present there is no consensus
about how and when the changes in fascicular ge-
ometry occur in a knee extensor eccentric training
program.
Another open question concerns the fact that
eccentric training studies have focused attention on
vastus lateralis (VL) muscle,6,22,23 usually assuming
that changes in VL architecture are representative
of the whole quadriceps muscle. However, different
adaptations in fascicular pennation angle after
eccentric training have been shown between VL
and vastus medialis (VM).6 The complex architec-
ture of VM led to an unacceptably low reliability in
fascicle length measurements,6,31,34 and did not
earlier study.6 However, evidence supports a

498 Adaptations to Eccentric Training MUSCLE & NERVE October 2013

FIGURE 1. Experimental design of the study: a control period of 4 weeks and a training period of 12 weeks. There were 5 evaluation
sessions (indicated by the arrows) throughout the study.
different magnitude of sarcomerogenesis (in-series
addition of sarcomeres within a muscle fiber)
between vastus intermedius (VI) and VL of rats sub-
jected to chronic eccentric exercise.35 Therefore,
the idea of specific adaptive responses to eccentric
training on fascicle geometry in synergistic knee ex-
tensor muscles seems reasonable.
The singular adaptation of each knee extensor
muscle to eccentric training may be related to its
fascicular geometry and specific response to eccen-
tric exercise. The most common acute response to
repeated eccentric contractions is called exercise-
induced muscle damage,36,37 which is an activation
mechanism of the repair process that leads to addi-
tion of sarcomeres aligned in-parallel38 and in-se-
ries.39 A higher number of in-parallel sarcomeres
leads to an increase in fiber diameter, possibly
leading to an increase in fiber pennation angle40,41
and resulting in a higher PCSA. The addition of
sarcomeres aligned in-series determines increased
fiber length.28,41 Assuming that muscles with larger
pennation angles present stronger fibers due to a
higher number of in-parallel sarcomeres, they are
less susceptible to damage. At the same time,
muscles with large fiber diameters and pennation
angles usually have small fascicle lengths.28,42 Short
fibers are more susceptible to muscle damage
because they work closer to the descending limb
of the force–length relationship and undergo a
great magnitude of strain during eccentric
actions.39
Ultrasonography scans have shown variability of
pennation angle and fascicle length values between
different sites of the quadriceps femoris muscle,34
demonstrating the complex fascicular geometry of
the knee extensor muscles in humans. Data from
human cadavers have shown that VL has longer
muscle fibers and higher pennation angles and
PCSA than rectus femoris (RF),42 thus reducing its
susceptibility to exercise-induced muscle damage.
On the other hand, RF is the only biarticular knee
extensor muscle, and therefore it has a lower prob-
ability of reaching the maximal physiological
length (descending limb of force–length curve)
during open kinetic chain exercises performed in
a sitting position (e.g., isokinetic dynamometers or
Adaptations to Eccentric Training
weight-lifting machines) due to hip flexion.
Because specific behavior from RF and VL muscle
architecture during this kind of exercise was not
assessed in previous human studies, it remains
unclear how it affects the adaptive response of
these muscles to eccentric training.
This study was motivated by: (1) conflicting
results related to pennation angle and fascicle
length increments; (2) the lack of evidence regard-
ing the time-course of fascicular geometry changes;
and (3) the lack of studies comparing the adaptive
response to eccentric training from synergistic
knee extensor muscles in humans. Therefore, our
aim was to assess and compare the effect of a 12-
week eccentric training program on the fascicular
geometry of 2 synergistic knee extensor muscles
with different architectural characteristics: RF
and VL.
METHODS
Experimental Design. A longitudinal clinical trial
was designed to verify morphological adaptations
of RF and VL during a 12-week knee extensor
eccentric training program. Volunteers were eval-
uated 5 times during the 4-month study to assess
architectural parameters of RF and VL (i.e., muscle
thickness, pennation angle, and fascicle length)
through ultrasonography images. A 4-week interval
period was observed between consecutive evalua-
tions. In the first 4 weeks of the study, the interval
period between the first and second evaluations
(called baseline and pre-training, respectively), vol-
unteers were subjected to a non-training control
period. After the second evaluation, subjects initi-
ated the 12-week eccentric training program. Addi-
tional evaluations were performed after 4 (post-4),
8 (post-8), and 12 weeks (post-12) of training (see
study design in Fig. 1). The study was conducted
according to the provisions of the Declaration of
Helsinki and was approved previously by the insti-
tutional ethics in research committee.
Subjects. Twenty male university students (age
24.05 6 3.73 years, height 1.75 6 0.06 m, weight
73.95 6 6.99 kg) completed the full study schedule.
All were physically active men who were not
MUSCLE & NERVE October 2013 499
enrolled in any kind of lower limb systematic resist-
Table 1. Eccentric training program.
ance training program for a period of 6 months
Mesocycle Week Frequency Sets Repetitions Volume
prior to their initial participation in the study. Vol-
unteers were carefully informed about the study 1 1 1 3 10 30
design and procedures during an interview session 2 2 3 10 30
3 2 3 10 30
when eligible subjects were selected. Exclusion cri- 4 2 3 10 30
teria included: (1) history of lower limb musculo- 2 5 1 4 10 40
skeletal disorders that could be a contraindication 6 2 4 10 40
for maximal tests or that could impair perform- 7 2 4 10 40
ance during training and tests (e.g., patellar tendi- 8 2 4 10 40
3 9 1 5 10 50
nitis, knee surgery, ruptured but not operated 10 2 5 10 50
knee ligaments, recent muscle strains or joint 11 2 5 10 50
sprains); (2) respiratory or cardiovascular diseases 12 2 5 10 50
considered a risk or a limiting factor for maximal
exercise; and (3) users of nutritional supplements
or anabolic steroids. J software (National Institutes of Health, Bethesda,
Maryland). Muscle thickness was considered to be
Muscle Architecture Assessment. A B-mode ultraso-
the distance between deep and superficial apo-
nography system (SSD-4000; Aloka Inc., Tokyo, neuroses and was calculated through the mean
Japan) with a linear-array probe (60 mm, 7.5 MHZ; value of 5 parallel lines drawn at right angles
Aloka) was used to determine muscle thickness, between the superficial and deep aponeuroses
pennation angle, and fascicle length of RF and VL along each ultrasonography image. The best fasci-
muscles. Great care was taken to determine the cle in each image was used for pennation angle
specific sites where the images were collected. Ana- and fascicle length analysis. Pennation angle was
tomical reference points (i.e., border of patella), calculated as the angle between the muscle fascicle
markings on the skin (i.e., freckles and scars), and and the deep aponeurosis, whereas fascicle length
the specific ultrasonography scanning sites were was measured as the length of the fascicular path
mapped on a malleable plastic sheet to ensure that between the 2 aponeuroses. When fascicle length
repeated scans were taken from the same was greater than the probe surface, the fascicle
site.6,23,30,31 All measurements taken during the line was extrapolated and calculated through a
study were performed by the same investigator trigonometric function. This mathematical proce-
with extensive experience in ultrasonography. Sub- dure has been used in several studies involving fas-
jects were evaluated in the supine position with cicle length evaluations of the vastus
the legs fully extended and the muscles relaxed.6 lateralis,6,22,23,29–31,34,43 and the estimated coeffi-
Subjects rested for 10 minutes before ultrasonogra- cient of variation of this fascicle length determina-
phy evaluations, and volunteers were instructed to tion is 4%.22,30
not engage in any vigorous physical activity for 48
hours before the tests. Eccentric Training. A 12-week knee extensor
Scans were taken at the midpoint between the eccentric training intervention was performed on
greater trochanter and the lateral condyle of the an isokinetic dynamometer (Biodex System 3; Bio-
femur. The ultrasonography probe was covered dex Medical Systems, Shirley, New York). The
with water-soluble transmission gel and oriented training program comprised 2 sessions per week
parallel to the muscle fascicles and perpendicular with a mimimum interval of 72 hours between ses-
to the skin over RF and VL. The transducer orien- sions. In accordance with the progression princi-
tation relative to the longitudinal axis of the thigh ple, the training program was divided into 3
was different between subjects due to their individ- mesocycles, and the training session volume (num-
ual anatomical characteristics. Probe alignment was ber of sets per session 3 number of repetitions
considered appropriate when several fascicles per set) was increased in each mesocycle (Table
could be easily delineated without interruption 1). Therefore, each training session comprised a 5-
across the image. minute warm up exercise on a cycloergometer and
Three ultrasonography scans were taken at 3–5 sets of 10 maximal eccentric contractions with
each site, and muscle architecture parameters a 1-minute rest period between sets. Subjects per-
from RF and VL were analyzed by the same investi- formed only 1 training session in the first training
gator, who was blinded to the identity of the par- week to allow for progressive introduction to the
ticipants and time-point at which each training regimen and to avoid having participants
ultrasonography image was obtained. Ultrasonogra- undertake an exercise session in the presence of
phy images were digitized and analyzed with Image clinical symptoms of exercise-induced muscle
500 Adaptations to Eccentric Training MUSCLE & NERVE October 2013
damage.36,37 Moreover, evaluation weeks through-
out the training program (weeks 5 and 9 of train-
ing) coincided with the exchange of mesocycles
and consequent increment of training volume, and
therefore only 1 training session was performed on
these weeks.
Eccentric contractions were performed accord-
ing to the protocol described by Baroni et al.44
Subjects were positioned in the dynamometer
chair with 85 of hip flexion and 90 of knee flex-
ion (0 5 full extension for hip and knee). Before
each eccentric contraction, the limb was extended
passively to 30 of knee flexion. Subjects were
encouraged to perform a maximal contraction of
the knee extensor as soon as the dynamometer
arm reached this position. In response to the sub-
ject’s extensor torque, the dynamometer drove the
segment to 90 of knee flexion (range of
motion 5 60 ) at an angular velocity of 60 /sec-
ond. Therefore, each eccentric contraction lasted
1 second, followed by 1-second rest during which
the limb was extended passively.
To avoid muscular imbalance between right
and left lower limbs due to training adaptations,
all volunteers trained both lower limbs. However,
some subjects had joint pain in 1 leg during the
training sessions, and others had musculoskeletal
injury in 1 leg during their daily activities (unre-
lated to the study intervention). In these cases,
only the uninjured segment from these subjects
performed the 12-week training program and was
considered for analysis. As a result, the statistical
analysis included 20 volunteers (15 with the left
leg and 5 with the right leg, or 16 non-dominant
and 4 dominant legs). Eighteen volunteers com-
pleted the 21 training sessions, and 2 subjects
missed 1 session each due to personal reasons,
leading our sample to an adherence level of
99.5%.
Statistical Analysis. A mean value obtained from
the measurements made in 3 ultrasonography
images from RF and VL for muscle thickness, pen-
nation angle, and fascicle length was considered
for statistical analysis. When these 3 values had a
dispersion >10%, the more discrepant value was
excluded, and only the 2 closest values were con-
sidered for analysis.
Values of muscle thickness, pennation angle,
and fascicle length from all volunteers at baseline
and pre-training evaluations were used to verify the
test–retest reliability of our measures through an
intraclass correlation coefficient (ICC).
The absolute values of muscle thickness, penna-
tion angle, and fascicle length from both muscles
from each of the 5 time-points (baseline, pre-training,
post-4, post-8, and post-12) were compared through a
Adaptations to Eccentric Training
repeated-measures analysis of variance (ANOVA).
The Mauchly test was performed, and the Green-
house–Geisser correction factor was applied when the
assumption of sphericity was violated. If a significant
time effect was determined, a least-significant-differ-
ence (LSD) post-hoc test was used to identify the differ-
ences between the time-points within each muscle.
The relative changes in relation to the pre-
training values were calculated for each parameter.
A 2-way repeated-measures ANOVA was used to
verify the interaction between muscle (RF and VL)
and time of assessment (baseline, pre-training,
post-4, post-8, and post-12). The Mauchly test and
Greenhouse–Geisser correction factors were
applied. If a significant interaction between muscle
and time-point was observed, a 1-way ANOVA was
used within each time-point to compare the RF
and VL percent change, and a repeated-measures
ANOVA followed by an LSD post-hoc test was used
within each muscle to compare the time-points. If
only time effect was observed, an LSD post-hoc test
was used to identify the differences between
time-points.
All statistical analyses were performed with an
alpha level of P  0.05. Results are presented in
the tables as mean 6 standard deviation and in the
figures as mean 6 standard error.
RESULTS
High scores for test–retest reliability between
baseline and pre-training evaluations were obtained
for all measures: muscle thickness (RF 5 0.96,
VL 5 0.91); pennation angle (RF 5 0.95, VL 5 0.90);
and fascicle length (RF 5 0.95, VL 5 0.91). More-
over, there were no significant differences
(P > 0.05) in any variable between these 2 time-
points (Table 2).
Concerning the absolute values, there was a
statistically significant time effect on muscle thick-
ness (RF: P < 0.01; VL: P < 0.01) and fascicle
length (RF: P < 0.01; VL: P < 0.01), but not for
pennation angle (RF: P 5 0.70; VL: P 5 0.54).
There was a significant increase in muscle thick-
ness in both muscles from pre-training to post-4
(P < 0.01), which remained constant until the end
of the training program (no significant differences
between post-4 and post-8 or post-12; P > 0.05).
Fascicle length of both muscles showed a signifi-
cant increment after 4 training weeks (RF:
P 5 0.02; VL: P 5 0.05) and between time-points
post-4 and post-8 (P < 0.01 for RF and VL), without
significant changes in post-12 compared with post-
4 or post-8 (P > 0.05 for both muscles).
Figure 2 shows the percent change (normalized
to the pre-training values) of each muscle architec-
tural parameter for all time-points analyzed. The
statistical analysis showed no significant interaction
MUSCLE & NERVE October 2013 501
Table 2. Muscle thickness (MT), pennation angle (PA), and fascicle length (FL) from rectus femoris (RF) and vastus lateralis (VL) at various
evaluation times of the study.
Baseline Pre-training Post-4 Post-8 Post-12
* *
RF MT (cm) 2.01 6 0.34 2.01 6 0.35 2.15 6 0.35 2.20 6 0.35 2.19 6 0.33*
PA ( ) 8.48 6 1.85 8.70 6 1.87 8.55 6 1.43 8.42 6 1.48 8.67 6 1.35
FL (cm) 13.49 6 2.48 13.63 6 2.29 14.31 6 1.97* 15.32 6 1.86*† 15.61 6 1.51*†
VL MT (cm) 2.61 6 0.24 2.59 6 0.23 2.72 6 0.23* 2.75 6 0.25* 2.77 6 0.29*
PA ( ) 17.18 6 2.54 17.28 6 2.50 17.79 6 2.50 17.02 6 2.78 17.15 6 2.58
FL (cm) 8.10 6 1.11 8.22 6 1.32 8.54 6 1.21* 9.51 6 1.50*† 9.67 6 1.75*†

*Different from baseline and pre-training (P < 0.05).

†
Different from post-4 (P < 0.05).

between muscle and time-point for muscle thick-

ness (P 5 0.07), pennation angle (P 5 0.68), or fas-
cicle length (P 5 0.73), demonstrating that RF and
VL had similar relative changes throughout the
training program. Muscle effect was not observed
in the 3 muscle architectural parameters (P 5 0.23,
P 5 0.77, and P 5 0.76; respectively). A time effect
was observed for muscle thickness (P < 0.01) and
fascicle length (P < 0.01), but not for pennation
angle (P 5 0.38). Muscle thickness and fascicle
length increased from pre-training to post-4
(P < 0.01) and from post-4 to post-8 (P < 0.01), but
not from post-8 to post-12 (P 5 0.90 and P 5 0.09,
respectively).
Figure 3 illustrates the changes in RF and VL
muscle architecture of a representative subject
through ultrasonography images for pre-training
and post-12 evaluations. Note the remarkable in-
crement of muscle thickness for RF and VL after
12 weeks of eccentric training.

DISCUSSION
In this study we have assessed and compared
the fascicular geometric adaptations between RF
and VL muscles throughout an eccentric training
program in humans. An animal model study could
be the optimal approach, because it allows for a
more precise evaluation of sarcomeres aligned in-
parallel and in-series per muscle fiber. However,
the specific adaptations of synergistic muscles to
eccentric training in animals have already been
reported,35 and the assessment of these adapta-
tions in humans is considered to be the novelty of
this study. Therefore, the main findings of this
study are that: (1) knee extensor eccentric training
increased muscle thickness and fascicle length of
RF and VL but produced no adaptive measurable
changes in pennation angles; (2) a 4-week eccen-
tric training period was enough to promote signifi-
cant muscle architectural adaptations; and (3) FIGURE 2. Relative changes in muscle thickness (MT), penna-
despite the differences between the muscle archi- tion angle (PA), and fascicle length (FL) of rectus femoris (RF)
and vastus lateralis (VL) throughout the study. Note that there
tecture of RF and VL, there were no differences in was no difference between muscles throughout the study. *Dif-
fascicular geometric adaptations between these syn- ferent from baseline and pre-training (P < 0.05); #different from
ergistic knee extensor muscles. post-4 (P < 0.05).

502 Adaptations to Eccentric Training MUSCLE & NERVE October 2013

FIGURE 3. Ultrasonography images from a representative subject showing rectus femoris (RF) and vastus lateralis (VL) muscles
before (pre-training) and after (post-12) the eccentric training program.

This experimental design eliminated the need
for a non-trained control group, because all volun-
teers underwent a control period immediately
before the eccentric training program. The high
ICC scores (0.90–0.96) and the absence of signifi-
cant differences between baseline and pre-training
evaluations confirmed the test–retest reliability of
the measurement procedures, strengthening the
trust in the results observed through the training
period.
The increment in muscle thickness is an
expected adaptive response to resistance train-
ing,41,45 and it can be attributed to increased pen-
nation angle, increased fascicle length, or both.
Adaptations to Eccentric Training
Our initial hypothesis was that both pennation
angle and fascicle length of RF and VL would
increase due to the in-parallel and in-series sarco-
mere addition as a response to eccentric exercise–
induced muscle damage.38,39 The simultaneous
increase of pennation angle and fascicle length
due to eccentric training has been reported for
VL,6 although no changes in these variables after
an eccentric training program have been reported
previously.23 The findings of our study are in
agreement with Reeves et al.,22 who suggested that
the increment in muscle thickness due to eccentric
training is attributed to increased fascicle length
but not to pennation angle changes.
MUSCLE & NERVE October 2013 503
 Because previous studies showed that eccentric
training increased muscle fiber area,2,5,14 we
expected an increased pennation angle after
eccentric training as a response to the hyper-
trophic process.40,41 However, studies involving
conventional and eccentric resistance training of
the knee extensors have reported conflicting
results regarding pennation angle changes.
Although some studies have described increments
in VL pennation angle after conventional resist-
ance training,22,29,30,33 others showed no changes
in this parameter.23,31 Blazevich et al.6 reported
increases in VL but not in VM pennation angle
due to eccentric training, whereas findings from
other eccentric training studies22,23 suggested no
changes in VL pennation angle. It is hard to deter-
mine the reason for these differing findings with
respect to pennation angle adaptation after differ-
ent kinds of resistance training, but a possible ex-
planation may be related to the measurement
error of the ultrasonography analysis technique.
Whereas eccentric training and conventional resist-
ance training seem to increase VL pennation angle
by around 0.1–5.1 ,6,22,23,29–31,33 ultrasonographic
analyses have shown a typical error of 0.24–
1.22 .31,34 Therefore, the adaptive response of pen-
nation angle is usually within or close to the mea-
surement error of the analysis technique, which
could explain why some studies described incre-
ments in hypertrophy markers (e.g., muscle thick-
ness, ACSA, volume) without changes in pennation
angle.
Although studies with 14 weeks of conventional
resistance training have shown increments in VL
fascicle length,22,30 most showed no changes in
this parameter after 5,29,31 9,33 or even 1623 train-
ing weeks. On the other hand, Seynnes et al.32
used a flywheel ergometer with a stronger eccen-
tric component than conventional weight-lifting
machines for knee extensor exercise and found
increases in VL fascicle length after only 10 days of
training. The effect of eccentric contractions on
fascicular length is supported by studies showing
increases in VL with 5,6 10,6 and 1422 weeks of
eccentric training, as well as increases in fascicle
length of biceps femoris9 and gastrocnemius medi-
alis8 after 7 and 8 training weeks, respectively. In a
similar way, studies in animal models have demon-
strated sarcomerogenesis induced by eccentric
exercise.35,46 This adaptation has been proposed as
a protective mechanism against exercise-induced
muscle damage in a subsequent eccentric exercise
session,39 commonly associated with the repeated
bout effect (for a review, see McHugh47).
Muscles with longer fibers would produce
forces over a greater length range and at high
shortening velocities, because they have a large
504 Adaptations to Eccentric Training
number of simultaneously contracting, serially
arranged sarcomeres.28,45 The shift of optimal
length in direction to longer fiber lengths17 is of
interest for athletes aiming to avoid muscle inju-
ries,18 whereas the increased shortening velocity of
longer fibers14 improves athletic performance
through movements with greater power.16,21 Kuma-
gai et al.,43 for example, demonstrated that profes-
sional runners with a best 100-meter sprint time of
between 10.0 and 10.9 s have greater fascicle
lengths than slower runners (11.0–11.7 s), and fas-
cicle length was correlated significantly with the
athletes’ performance. In addition, the decrease in
fiber length associated with sarcopenia19,20 seems
to be responsible for almost half of the muscle
shortening velocity difference between young and
elderly subjects48; this implies that the deleterious
aging effect could be counteracted with eccentric
training.21,22 Therefore, our findings suggest that
these and other possible benefits related to fascicle
length increment could be reached with relatively
short periods of knee extensor eccentric training.
Whereas VL has longer muscle fibers and
higher pennation angles and PCSA,42 RF is a biar-
ticular knee extensor muscle that does not reach
its maximal physiological length during open ki-
netic chain exercises performed in a sitting posi-
tion. In other words, each muscle has favorable
and unfavorable factors regarding onset of exer-
cise-induced muscle damage, which should be
taken into account together in order to under-
stand the responses of RF and VL to eccentric
training. Therefore, the similar adaptations in fas-
cicular geometry between muscles found in this
study should be related to the specific response of
RF and VL to the type of exercise performed dur-
ing the training sessions.
Guilhem et al.49 demonstrated that RF and VL
have similar activation patterns throughout the
range of motion during maximal isokinetic eccen-
tric contractions. Both muscles showed a progres-
sive activation increase from 30 to 65 of knee
flexion, a plateau region between 65 and 75
(near to 80–90% of electromyographic activity
observed in maximal isometric contractions for
each muscle), and an activation fall from 75 to
90 of knee flexion. This means that both muscles
were recruited to generate force at a similar level
during the range of motion of the isokinetic knee
extensor eccentric exercise used during the train-
ing sessions. Therefore, the similar activation
might be an important factor that could explain
why neither RF nor VL had a larger adaptive
response after eccentric training.
Although VL and other vasti muscles have lon-
ger muscle fibers than RF,42 studies50,51 with run-
ners showed that both mono- and biarticular knee
MUSCLE & NERVE October 2013
extensor muscles do not reach the most critical
lengths of the force–length relationship52 during
contractions performed in the sitting position up
to 90 of knee flexion. This means that the eccen-
tric training program performed in our study
(30 –90 of knee flexion) probably represents an
exercise that does not reach the critical lengths for
muscle damage.53 If RF and VL were exercised in
a similar part of the force–length relationship (i.e.,
ascending limb, plateau region, and possibly initial
part of the descending limb), then these muscles
may have undergone similar levels of exercise-
induced muscle damage after the training sessions,
which could explain the similar muscle architec-
ture adaptations observed in our study.
Although clinical programs of eccentric training
or rehabilitation are designed to avoid the undesir-
able symptoms of muscle damage (mainly delayed-
onset muscle soreness), our eccentric training pro-
gram was designed to induce muscle damage and
potentiate the adaptations to eccentric exercise.
Although other mechanisms are related to muscular
adaptation to resistance training, the regeneration
process subsequent to exercise-induced muscle
damage has been considered to be the main factor
explaining why eccentric training leads to a higher
hypertrophic response54 and larger in-series sarco-
mere addition35,46 compared with concentric train-
ing. These aspects justify why the rationale for this
study and the explanation for our findings are
based in the effects of eccentric exercise–induced
muscle damage.
It is of interest to note that knee extensor exer-
cises performed in open and closed kinetic chain
showed different acute responses55 as well as open
kinetic chain exercises performed in isotonic and
isokinetic conditions.49 In other words, other types
of knee extensor exercises would likely produce
adaptations distinct from those observed in our
study. Therefore, grounded in previous evidence
that eccentric training adaptations have demon-
strated important functional implications for pre-
vention and rehabilitation of musculoskeletal
injuries, athletic performance, and elderly func-
tionality, it is imperative that future studies con-
centrate their attention on determining the
contribution of each training variable (i.e., type of
exercise, range of motion, angular velocity, train-
ing volume, etc.) to the muscle adaptive response.
In conclusion, these results show that increased
VL and RF muscle thickness after knee extensor
eccentric training is related to increments in fasci-
cle length and not to any measurable adaptive
changes in pennation angle. Knee extensor muscle
architectural adaptations were identified after only
4 weeks of eccentric training. Although RF and VL
differ significantly in their geometrical design,
Adaptations to Eccentric Training
knee extensor eccentric training performed in an
isokinetic device did not provoke an inhomogene-
ous adaptation between these muscles throughout
the 12-week eccentric exercise training program in
our study.
The authors thank all the study volunteers for their participation
in this project.
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