Sports Med 2006; 36 (12): 1003-1017

LEADING ARTICLE 0112-1642/06/0012-1003/$39.95/0

 2006 Adis Data Information BV. All rights reserved.

Effects of Physical Training and

Detraining, Immobilisation, Growth
and Aging on Human
Fascicle Geometry
Anthony J. Blazevich
Centre for Sports Medicine and Human Performance, Brunel University, Uxbridge,
Middlesex, UK

Abstract In addition to its size and the extent of its neural activation, a muscle’s
geometry (the angles and lengths of its fibres or fascicles) strongly influences its
force production characteristics. As with many other tissues within the body,
muscle displays significant plasticity in its geometry. This review summarises
geometric differences between various athlete populations and describes research
examining the plasticity of muscle geometry with physical training, immobilisa-
tion/detraining, growth and aging. Typically, heavy resistance training in young
adults has been shown to cause significant increases in fascicle angle of vastus
lateralis and triceps brachii as measured by ultrasonography, while high-speed/
plyometrics training in the absence of weight training has been associated with
increases in fascicle length and a reduction in angles of vastus lateralis fascicles.
These changes indicate that differences in geometry between various athletic
populations might be at least partly attributable to their differing training regimes.
Despite some inter-muscular differences, detraining/unloading is associated with
decreases in fascicle angle, although little change was shown in muscles such as
vastus lateralis and triceps brachii in studies examining the effects of prolonged
bed rest. No research has examined the effects of other interventions such as
endurance or chronic stretching training. Few data exist describing geometric
adaptation during growth and maturation, although increases in gastrocnemius
fascicle angle and length seem to occur until maturation in late adolescence.
Although some evidence suggests that a decrease in both fascicle angle and length
accompanies the normal aging process, there is a paucity of data examining the
issue; heavy weight training might attenuate the decline, at least in fascicle length.
A significant research effort is required to more fully understand geometric
adaptation in response to physical training, immobilisation/detraining, growth and
aging.
1004
Historically, changes in human fascicle geome-
try1 have been impossible to examine since the
excision and subsequent study of muscle could only
be performed in cadavers. More recently, imaging
techniques such as magnetic resonance (MRI)[1] and
ultrasound[2,3] imaging (figure 1) have allowed the in
vivo examination of geometric, i.e. fascicle length
and angle, changes.[2,4-8] Results of such research
have predominately shown geometry to be highly
changeable. An understanding of how different
forms of muscle loading, or exercise training, affects
geometry would allow us to bring about deliberate
and specific changes that would influence the mus-
cles’ force-generating properties. There are, howev-
er, a limited number of studies that have examined
the plasticity of fascicle geometry, or that have
compared populations of individuals who have per-
formed different exercise training. The present re-
view summarises research that quantifies changes in
fascicle geometry with exercise and detraining, and
development and aging. A greater understanding of
fascicle geometric adaptation could inform both
physical performance and rehabilitative training
programmes, as well as allowing the development of
muscle models that predict physical performance
θ PCSA
b 
β
α  ACSA
lf = ∆a − b a 
Muscle length
Fig. 1. Muscle architectural parameters include: fibre length = dis-
tance between ends of a fibre (a to b); pennation angle (θ) =
fascicle angle (relative to the aponeurosis [α]) minus the aponeuro-
sis angle (relative to the tendon [β]); muscle length; and anatomical
(ACSA) or physiological (PCSA) cross-sectional area. The PCSA
can be calculated as (V/t) • sinθ for a simple, uni-pennate muscle,
where V is the muscle volume, t is the muscle thickness from one
aponeurosis to the other, and θ is the pennation angle. In more
complex muscles, PCSA is calculated as V/ lf • cosθ where lf is the
mean fibre/fascicle length.
1 The term ‘fascicle geometry’ as used here describes the angulation and length of muscle fascicles. The broader term
‘muscle architecture’ will be reserved for the description of the whole muscle structure including fascicle geometry,
muscle length and muscle volume (or physiological cross-sectional area).
 2006 Adis Data Information BV. All rights reserved.
Blazevich
changes after training and detraining interventions
and/or aging and development.
1. Geometry of Human Muscles
Many human muscles contain fascicles that do
not run directly from origin to insertion, but are
angled and attach to the muscles’ aponeuroses (fig-
ure 2). These ‘pennate’ muscles can have complex
fascicular arrangements that differ markedly be-
tween muscles. Their fascicle arrangement is proba-
bly a greater determinant of general muscle function
than other aspects such as fibre type.[9] As a general
rule, muscles with large fascicle attachment angles
and correspondingly short fascicles are suited to
contractions involving high forces[10] and/or pro-
duce forces over a relatively short range of mo-
tion.[11] Examples include the large proximal mus-
cles of the legs (e.g. vastus lateralis; see table I),
smaller muscles such as those involved in mastica-
tion,[10] and those involved in postural control such
as soleus and tibialis posterior. Large fascicle angles
are also seen in muscles that attach to long tendons,
such as gastrocnemius lateralis, which produce suf-
ficient force to remain relatively isometric during
the propulsive phase of stretch-shorten contractions
in walking,[12-14] running[12,14] and jumping with[15,16]
and without[12,14,17] countermovement, and allow op-
timum elongation and recoil of the tendon.[12] Mus-
cles that commonly participate in movements re-
quiring significant length change or a high shorten-
ing velocity tend to be characterised by long
fascicles attaching at relatively small angles to the
tendon.[9,18] Examples include adductor magnus and
longus, whose longer fascicles attaching at smaller
angles allows force generation over a large range of
motion at the hip during flexion/adduction tasks,
and the topologically similar long head of biceps
femoris, which is highly active during sprint run-
ning.[19-21]
Sports Med 2006; 36 (12)
Plasticity of Fascicle Geometry 1005

2. Effects of Geometry on
Force Production θ θ θ

Fascicle
2.1 Fibre Length force

Muscle-specific geometric differences can be ra-

tionalised by considering the effects of different
fascicle arrangements on force production. Muscles
containing long fascicles would produce forces over
large length ranges and at high shortening speeds
because they have a large number of simultaneously
contracting, serially arranged sarcomeres. Moreo-
ver, since the shortening speed of each sarcomere in
a fibre or fascicle would be slower for a given speed
of whole-fibre shortening when there are more
sarcomeres in series, sarcomere force would not
decrease as rapidly as fibre-shortening speeds in-
crease, according to the force-velocity relationship.
Therefore, at high shortening velocities, longer fas-
cicles are capable of generating greater force. None-
theless, the increase in sarcomere number would
increase the energy cost of force production, since
force output is not improved with the increase in
energy-consuming sarcomeres. Indeed, it is theoreti-
cally possible that energy consumption per sarcom-
ere is increased due to the absorption of energy by
neighbouring sarcomeres in protein structures such
as titin,[23-26] cross-bridges,[27,28] and actin and myo-
sin filaments,[29-31] as well as by the re-arrangement
of the z-band lattice,[32] which has been shown to
extend by approximately 20% in tetanised fibres.[33]
Thus, muscle containing longer fascicles produce
force over long length ranges and at high shortening
speeds, but the relative energy cost of force produc-
tion is high.
2.2 Fibre Angle
While increases in fascicle angulation, or penna-
tion, reduces the proportion of fibre force directed
along the tendon (tendon force = sum of fibre forces
× cos[fibre angle]), this effect is minimal when
fascicle angle is moderate (e.g. <25°), as shown in
figure 2. However, angulation probably improves
force generation in three main ways. First, for the
same muscle volume, a muscle with larger fascicle
Tendon/
aponeurosis
m m m
θ = 5° θ = 25° θ = 45°
Tendon force (% muscle force)
100 θ = 5° θ = 25°
80 θ = 45°
60
40
20
0
5 15 25 35 45 55
Fascicle angle (°)
Fig. 2. The effect of fascicle angle on the quantity of force directed
along the tendon axis. As fascicle angle (θ) increases, the propor-
tion of fibre force directed along the tendon decreases (tendon
force = sum of fibre forces × cos[fibre angle]), where the fibre force
is represented by the arrow attached to the tendon/aponeurosis.
The tendon is attached to a mass (m) representing the inertia of the
system on which the muscle-tendon complex does work. The effect
of fascicle angulation on the proportion of force directed along the
tendon is minimal when fascicle angle is moderate (e.g. <25°), but
increases non-linearly as fascicle angle increases, as shown in the
graph.
angulation will have a greater physiological cross-
section and thus a greater force-generating capaci-
ty.[34] Essentially, this fascicular geometry allows a
greater amount of contractile tissue to attach to a
given area of tendon or aponeurosis.[7] Second, fas-
cicle angulation probably increases force by al-
lowing fibres to operate closer to their optimum
length. Fibres in pennate muscles rotate as they
shorten[35] so tendon excursion is greater than the
shortening distance of the individual fibres. Accord-
ing to the length-tension relationship, there will be
an optimum sarcomere length at which fibres pro-
duce their greatest active force. Since optimum
sarcomere length seems to occur at lengths where

 2006 Adis Data Information BV. All rights reserved. Sports Med 2006; 36 (12)
1006
Table I. Examples of fascicle angles and lengths in human muscles (average of two cadavers)[22]
Force characteristics Action
High Force generation
Prolonged (time) Postural control
Rapid/high Stretch-shorten
High velocity/large range of motion Long-range force
Eccentric/large range of motion Long-range force
the highest forces are required (rather than at the
muscle’s resting length for example; Herring et
al.[36] [human]; Lutz and Rome[37,38] [frog]), fibres
that shorten less for a given tendon excursion are
likely to stay closer to their optimum for force
generation. Third, since the time-dependent shorten-
ing distance of a fibre decreases during a contrac-
tion, the shortening speed will also decrease (v = d/t,
where v = velocity of shortening, d = fibre displace-
ment and t = time) and force increases as per the
force-velocity relationship for muscle. So in pennate
muscle, a more forceful contraction might be possi-
ble from exploitation of both the force-velocity and
length-force relationships in addition to the greater
quantity of contractile material that can attach to the
tendon or aponeurosis. It is clear then that having
large fibre angles would be an important adaptation
to allow high force production.
3. Plasticity of Human Muscle Geometry
Tissues within the human body have a great
ability to remodel in response to stress (e.g. Wolff’s
Law for bone). At the gross level of muscle, in-
creased loading is commonly associated with an
early (<6 weeks) change in their neural activation
during heavy lifting[3,38-44] (for a review of neural
mechanisms see Enoka[45]). However, subsequent
periods of loading result in an increase in muscle
size with more contractile tissue being available for
force generation.[3,39,42,46,47] After prolonged periods
of training, when the rate of hypertrophy has
reached a plateau, further small strength increases
are most likely to occur by neural mechanisms in
response to variations in training intensity or vol-
ume.[48] While this sequence of adaptation is well
 2006 Adis Data Information BV. All rights reserved.
Blazevich
Muscle Fascicle angle Fascicle length
(°) (mm)
Vastus lateralis 13 80.3
Soleus 32 30.3
Tibialis posterior 19 28.6
Gastrocnemius lateralis 17.5 60.6
Adductor magnus 4 113.0
Adductor longus 3.5 94.2
Biceps femoris (long head) 7 72.6
accepted, it fails to take into account changes in
fascicle geometry as a possible contributing factor to
strength adaptation. Since fascicle geometry signifi-
cantly impacts on force generation, loading-depen-
dent geometric changes would influence force pro-
duction and ultimately influence movement capaci-
ty. There are two ways in which information
regarding changes in fascicle geometry can be
gained, one is to examine differences between popu-
lations who perform different training regimes, and
another is to measure its changes in response to
training.
3.1 Measuring Fascicle Geometry in Humans
MRI has been scarcely used to measure fascicle
geometry in humans. The striation patterns resulting
from inter-fascicular fat and connective tissue ele-
ments can be visualised using MRI scanning since
their signal (brightness in the image) differs from
the muscle tissue because of variations in water
content of the tissues. The reconstruction of separate
two-dimensional (2-D) images, with the striation
patterns visually marked, into three dimensions
(3-D) allows the accurate determination of fascicle
geometry. Scott et al.[1] found good agreement be-
tween MRI and dissection estimates of fibre geome-
try in the quadriceps of human cadavers, while
Gaudy et al.[49,50] reported the muscle architectural
arrangements of masseter, temporalis and lateral
pterygoid, with little variance between cadaver dis-
section and MRI measurements. Lam et al.[51] vali-
dated the measurement of fibre angles of the masse-
ter by MRI in a single human cadaver. To date, no
studies have examined fascicle geometry in athletes,
or determined longitudinal changes in fascicle ge-
Sports Med 2006; 36 (12)
Plasticity of Fascicle Geometry
ometry using MRI, perhaps because of the difficulty
of manually tracing fascicles and then reconstruct-
ing 3-D images.
More recently, other MRI-based techniques have
been developed. Diffusion tensor imaging relies on
the diffusion of water in the muscle being con-
strained by membranes and other cellular constitu-
ents, so in muscles the diffusion occurs largely along
the plane of the fibres. The diffusion tensor can be
calculated from the measurement of the direction of
the diffusion.[52,53] This technique is more efficient
than traditional methods of MRI determination be-
cause the fibres can be tracked rapidly by a fibre-
tracking algorithm[54,55] and the fibre orientations
can be quickly reconstructed. While the technique
has been used to determine fascicle geometry in
animal muscles,[53,56-59] it has yet to be used to
quantify human fascicle geometry adaptations. In
addition to the diffusion tensor technique, magnetic
resonance elastography has recently been shown to
provide information with regard to fascicle geome-
try in humans.[60] After low frequency (<1000Hz)
oscillations are induced in the muscle by a pneumat-
ic or mechanical driver, MRI is used to measure the
spread of shear waves travelling through the muscle.
The spread of waves is constrained by the stiffness
of muscle-based tissues, where lateral (with respect
to the muscle fibres) transmission is affected by
parallel layers of connective tissue but longitudinal
transmission is relatively unimpeded along the fib-
res. Again, this technique has not yet been used to
quantify muscle fascicle geometric changes longitu-
dinally; however, both diffusion tensor imaging and
elastography show great promise as fascicle imag-
ing techniques.
The most common method of measuring fascicle
geometry is via the acquisition of 2-D longitudinal
images of the muscle using ultrasound[2,4-7,61,62] (fig-
ure 3). When the ultrasound transducer (probe) is
oriented in the plane of the fascicles, the fascicle
angles and lengths can be accurately measured.
Measures of fascicle angle (≤1.5°)[2,63,64] and fasci-
cle length (≤1.5mm)[2] have been shown to be simi-
lar to those measured directly in cadavers. When the
transducer is not oriented in the plane of the fasci-
 2006 Adis Data Information BV. All rights reserved.
1007
cles, larger errors in fascicle length estimates have
been observed when compared with 3-D ultrasound
images (2.4–14.0%).[65] Fascicle length is also diffi-
cult to accurately measure when the width of the
probe (often 3.8 or 6cm) is insufficient to capture an
entire fascicle. In these instances, linear approxima-
tions are calculated from muscle thickness and fasci-
cle angle measures (length = muscle thickness/sin ×
fascicle angle). These do not account for fibre curva-
ture that is present in some hypertrophied or con-
tracting muscles and errors range from approximate-
ly 0% (non-contracted gastrocnemius) to 6.6% (con-
tracted gastrocnemius) according to Muramatsu et
al.,[66] or 2.4% in the non-contracted tibialis anterior
muscle.[67] Fascicle length can also be estimated by
measuring the visible portion directly and then esti-
mating the non-visible portion. Errors of 2–7% have
been reported for this method.[68,69] Regardless, ul-
trasound imaging techniques allow valid and relia-
ble measures of fascicle geometry in vivo, and have
been commonly used to examine population differ-
ences and longitudinal adaptations.
3.2 Population Differences in
Fascicle Geometry
A muscle’s phenotype is at least partly reflective
of its long-term activity patterns. Although a per-
son’s genetic predisposition might play a role, it is
a b
Skin/fat layer
Vastus
lateralis
Vastus
intermedius
Bone
Fig. 3. Images of human muscles can be obtained using ultrasound
imaging. In this longitudinal section of the thigh, the skin/fat layer,
femur (bone), and vastus lateralis and vastus intermedius muscles
are clearly visible. The aponeurosis (apo) and line of fascicles (fas)
are also visible, from which fascicle angle can be determined, and
fascicle length can be estimated using standard trigonometric pro-
cedures.[5-7,68,69] Physical training can alter a muscle’s fascicle ge-
ometry. Here, a period of heavy strength training has resulted in an
increase in fascicle angle concomitant with the increase in muscle
thickness (a vs b).
Sports Med 2006; 36 (12)
1008
expected that the physical work capacity of athletes,
who have consistently trained with specific move-
ment patterns for a period of many years, would
vary according to the adaptations elicited by their
training. Indeed, significant, training-specific differ-
ences in work capacity have been shown among
groups of athletes with varied training histo-
ries.[70-76] Thus, one method of considering training-
dependent fascicle geometric adaptations is to com-
pare populations with different physical activity his-
tories.
A significant quantity of research has examined
differences in fascicle geometry between different
athlete populations. For example, both vastus later-
alis and gastrocnemius in highly trained male sprint-
ers (100m in <11.0 seconds) were found to contain
longer fascicles attached at smaller angles than in
lesser-trained sprinters (100m in >11.0 seconds[77]).
In females, sprinters have been shown to have small-
er fascicle angles in vastus lateralis than non-trained
controls (with no differences in gastrocnemius) and
longer fascicles in vastus lateralis and gastrocnemi-
us lateralis, which were significantly correlated with
100m best time (r = –0.51 and r = –0.44, respective-
ly[78]). A comparison of male and female sprinters
revealed no differences in absolute or relative (to leg
length) fascicle length.[78] When the fascicle geome-
try of sprinters was compared with well trained
endurance runners, differences were also seen.[79]
While endurance runners (13.5–14.5 minutes for
5000m) had a lesser muscle thickness in the vastus
lateralis and gastrocnemius muscles than sprinters
(10.0–10.9 seconds for 100m), their fascicles were
also shorter and their fascicle angles greater. Com-
pared with non-active subjects, young endurance
runners had larger fascicle angles (13.7%) but a
similar fascicle length in gastrocnemius medialis;
there were no differences in fascicle geometry in the
vastus lateralis.[61] At least for the gastrocnemius,
muscles in endurance runners appear to be adapted
to produce forces with minimal metabolic cost (i.e.
short fibres or fascicles require less energy for a
given level of force production because they have
fewer sarcomeres in series), whereas in sprinters
they are adapted for high-speed force development
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Blazevich
relatively regardless of cost. These data would sug-
gest that there are significant differences in the
fascicle geometry of well trained sprinters compared
with lesser-trained sprinters, endurance runners or
control subjects.
Unique geometry is also seen in weight-trained
individuals. As expected, muscle thickness mea-
sured at 13 sites on muscles including triceps
brachii, vastus lateralis and gastrocnemius lateralis
of powerlifters was significantly correlated with
squat, bench-press and deadlift performance (r =
0.63–0.91[80]). However, fascicle length of the vas-
tus lateralis was also positively correlated with squat
lift (r = 0.50) and deadlift (r = 0.54) strength, and
triceps fascicle length was correlated with bench-
press strength (r = 0.52). While muscle thickness
and fascicle angle were positively correlated (triceps
brachii r = 0.64, p < 0.01; gastrocnemius r = 0.48, p
< 0.05), triceps brachii fascicle angle was negatively
correlated with bench-press performance (relative to
fat-free mass, r = –0.45) and gastrocnemius fascicle
angle was negatively correlated with deadlift per-
formance (relative to fat-free mass, r = –0.56). In
another study, bodybuilders were shown to have
greater muscle thickness and fascicle angles than
non-weight-trained controls.[2] There is also some
evidence that muscle size, fascicle angle and fasci-
cle length are simultaneously greater in athletes who
require a large body mass and physical strength as
well as fast movement velocities (e.g. Sumo wres-
tlers[81]), than they are in the normal population.
3.3 Training Effects on Fascicle Geometry
While the assessment of population differences
provides an insight into possible activity-dependent
adaptations in fascicle geometry, a more satisfactory
method of examining the plasticity of fascicle geom-
etry is to directly measure the longitudinal adapta-
tions.
3.3.1 Resistance Training
Early research was suggestive of there being little
change in geometry in response to resistance train-
ing. Rutherford and Jones[8] showed no change in
vastus lateralis or intermedius fascicle angles after 3
months of quadriceps strength training in seven men
Sports Med 2006; 36 (12)
Plasticity of Fascicle Geometry 1009

Table II. Muscle architecture changes after strength training in humansa

Study
Rutherford and Jones[8]
Kawakami et al.[7]
Aagaard et al.[4]
Blazevich and Giorgi[5]
Blazevich et al.[84]
a Significant geometric change seems possible in previously untrained subjects, but these seem more limited in highly trained
athletes.
Method
3mo weight training (previously
untrained)
16wk elbow extensor training
(previously untrained)
14wk leg extensor training
(previously untrained)
12wk bench-press and triceps
extension training (well trained)
12wk bench-press and triceps
extension training + testosterone
injections (well trained)
5wk isokinetic leg extension
(previously untrained)
Muscles
Vastus lateralis, vastus
intermedius
Triceps brachii (lateral head)
Vastus lateralis
Triceps brachii (lateral head)
Triceps brachii (lateral head)
Vastus lateralis, vastus
intermedius, vastus medialis
Change
No change in fascicle angle or length
29.1% increase in fascicle angle, no
changes in fascicle length (0.9%)
33.8% (2.7°) increase in fascicle angle
No change in fascicle angle
39.5% (3.2°) increase in fascicle angle
No change in fascicle angle or length

and five women. There was also no relationship
between fascicle angle and force-generating capaci-
ty, although the authors reported a moderately high
coefficient of variation (13.5%), making small
changes (<2°) in fascicle angle difficult to detect.
Further research showed that significant changes in
fascicle geometry occurred in response to training
(see example in figure 3 and a summary in table II).
Kawakami et al.[7] reported significant increases in
triceps brachii physiological cross-sectional area
(PCSA) [+33.3%] and fascicle angle (+29.1%, 4.8°)
with no changes in fascicle length (–0.9%) after 16
weeks of elbow extensor training in five men. Aag-
aard et al.[4] found that fascicle angle (+33.8%, 2.7°;
vastus lateralis) increased significantly after 14
weeks of leg extensor training in 11 men simultane-
ously with anatomical cross-sectional area (+9.7%;
quadriceps total area) and muscle fibre cross-sec-
tional area (+12.9%; vastus lateralis). More recently,
Gondin et al.[82] reported fascicle angle increases in
vastus lateralis (14%) after 8 weeks of static electro-
myostimulation training of the knee extensors that
resulted in an increase of 27% in maximum contrac-
tion force and a 6% increase in quadriceps cross
section. Also, Alegre et al.[83] reported increases in
fascicle length in vastus lateralis after 13 weeks of
strength training when the concentric phase was
performed with maximum speed. These data indi-
cate that fascicle length is also adaptable, although it
is presently unclear whether the training velocity or
range of motion most influenced the changes (sec-
tion 5). Thus, recent research has shown a signifi-
cant adaptability of fascicle geometry in response to
longer-term resistance training.
Other research has examined changes in well
trained, athletic populations. A small change in tri-
ceps brachii muscle thickness (+13.6%) after a
12-week period of heavy resistance training in pre-
viously weight-trained men (n = 5) was shown to
occur without fascicle geometry or strength
changes.[5] However, in the same study, another
group (n = 4) showed a significant strength increase
(15.8%, bench-press) accompanied by increases in
fascicle angle (39.5%, 3.2°) after receiving once-
weekly intramuscular testosterone enanthate injec-
tions (3.5 mg/kg). Little change in geometry or
strength appears to occur, therefore, in well trained
lifters who continue to perform similar training,
although the administration of anabolic steroids
might allow greater strength increases accompanied
by significant geometric adaptation. In another
study,[6] however, two resistance training groups
comprising competitive athletes (n = 7 and n = 8)
with at least 1 year of resistance training experience,
showed increases in vastus lateralis fascicle angle
after 5 weeks of heavy (first session in the week) and
explosive (second session) squat lift training that
were significantly greater than a group that omitted
its strength training, but performed only sprint/jump
training (n = 8; no change). In the same study, vastus

 2006 Adis Data Information BV. All rights reserved. Sports Med 2006; 36 (12)
1010
lateralis fascicle length increased only for the sprint/
jump-trained group. Muscle thickness of both vastus
lateralis and rectus femoris increased in all training
groups also, although there were no differences in
performance (sprint, jump and strength tests) be-
tween the groups after training. Similar to previous
studies on untrained subjects,[7,83] these results show
that changes in fascicle geometry occur in response
to resistance training in well trained subjects. The
results may have differed from previous studies
where little change was noticed in trained subjects
because the training programme differed sufficient-
ly from that which the athletes had previously per-
formed. These data were the first to show short-term
changes in geometry in response to training, and
also show that increases in fascicle length and de-
creases in fascicle angle can occur after high-veloc-
ity movement training in humans; it is not known,
however, whether other factors such as the training
range of motion influence the architectural adapta-
tion.
Given that rapid (5-week) changes in geometry
have been shown to occur in athletes, one might
speculate that rapid adaptation is a mechanism by
which early strength increases occur in previously
untrained subjects. However, recent data[84] showed
no changes in quadriceps muscle size or geometry in
previously untrained men (n = 7) and women (n = 8)
despite significant increases in concentric and ec-
centric isokinetic strength of the quadriceps muscle
group after 5 weeks of isokinetic knee extensor
training. Also, Gondin et al.[82] did not find signifi-
cant increases in vastus lateralis fascicle angle after
4 weeks of electromyostimulation training, despite
there being significant increases after 8 weeks. It is
therefore likely that changes in fascicle geometry
are not a significant factor influencing early strength
increases in previously untrained subjects, although
whether rapid adaptations occur after training using
other modes (e.g. isotonic/isoinertial) is not
known.[7]
Geometric changes are probably not a factor in-
volved in strength increases that have been reported
in the untrained contralateral limb in response to
unilateral training.[85-87] Kawakami et al.[7] reported
 2006 Adis Data Information BV. All rights reserved.
Blazevich
no changes in muscle size and geometry of elbow
extensors of an untrained limb in five men after 16
weeks of training, although no strength changes
were found either. Blazevich et al.[84] have reported
that significant increases in contralateral knee exten-
sor strength after 5 weeks of isokinetic training
occurred without a change in geometry. Thus, al-
though few data are available, there is no current
evidence to suggest that fascicle geometric changes
are a factor affecting short-term strength increases in
a contralateral limb.
3.3.2 Endurance Training
No research has examined changes in geometry
after long-term endurance training. It is therefore
not known whether endurance training can elicit
geometric adaptations, or how different modes of
training (e.g. cycling, running, lifting) might affect
them. It makes intuitive sense that if a muscle could
generate force more efficiently, then muscle endur-
ance would improve. Given that muscles with large
pennation and shorter fascicles would generate force
with less metabolic cost, such an adaptation might
be expected with endurance training. Certainly, effi-
ciency during stretch-shortening contractions could
be expected to increase following such a geometric
adaptation. This reasoning is consistent with the
finding that vastus lateralis and gastrocnemius fasci-
cles were shorter and fascicle angles greater in a
group of endurance runners compared with sprint-
ers.[79] However, it could also be argued that longer
fibres would benefit muscles that are commonly
recruited during endurance tasks where forces are
produced over large ranges of motion, since such
fibre geometry is uniquely adapted to perform such
work. Also, intramuscular pressure (IMP) is greatest
in muscles with large fibre angles;[88,89] therefore,
occlusion of blood flow during muscle contraction
would be greater. This hypothesis is consistent with
the decreases in blood volume and oxygen satura-
tion seen during exercise (1Hz plantar flexion) in the
distal region of gastrocnemius medialis, where fas-
cicle angle and length are greatest.[90] During loco-
motion, IMP increases with movement speed[91] in
line with the known positive relationship between
IMP and muscle force,[92-95] so the limitation of
Sports Med 2006; 36 (12)
Plasticity of Fascicle Geometry
Table III. Muscle architecture changes after detraining/unloading in humansa
Study Method
Abe et al.[103] 20d bed rest
Kawakami et al.[98] 20d bed rest
Narici and Cerretelli[62] Disuse/injury
Bleakney and Maffulli[102] Disuse/injury
a Some data are suggestive of decreases in fascicle angle and length in line with muscle atrophy. Differences between studies might
be related to study duration or the length at which muscles were held during unloading.
blood flow at increased speeds of locomotion might
be expected to impact on muscle endurance. Of
course, during most forms of locomotion, muscle
contractions are phasic and blood flow will not be
significantly affected when the duty cycle is below a
threshold limit (e.g. <40%),[93] so the effect of ge-
ometry on endurance performance is probably com-
plex. It would seem likely that fibre type is a more
important muscle-based determinant of muscle en-
durance while fascicle geometry affects more the
length-tension and force-velocity characteristics of a
muscle.
3.3.3 Detraining/Unloading
Numerous studies have investigated the effect of
detraining/unloading on muscle size and strength.
For example, significant reductions in strength and
muscle size follow prolonged bed rest[96-98] along
with decreases in fibre size[99] and changes in fibre
type (slow- to fast-twitch transformation[100]). How-
ever, few studies have measured fascicle geometry
after a period of detraining/unloading, or have not
reported changes in geometry when it has been
measured for the determination of PCSA. In one
study, Kawakami et al.[98] found no changes in vas-
tus lateralis, plantarflexors or triceps brachii fascicle
lengths or angles in subjects (n = 5) after 20 days of
bed rest despite significant reductions in PCSA (ta-
ble III); a similar finding was reported in four sub-
jects in a later publication.[97] These results are simi-
lar to those reported in rat studies where immobilisa-
tion of gastrocnemius and soleus muscles at
shortened lengths was not associated with changes
in fibre angles.[101] However, Narici and Cerretel-
 2006 Adis Data Information BV. All rights reserved.
1011
Muscles Change
Gastrocnemius medialis, No change in vastus lateralis or triceps brachii,
vastus lateralis, triceps brachii significant decrease in gastrocnemius (5.5%)
measured standing, but not lying (4.7% decrease)
Vastus lateralis, plantarflexors, No change in fascicle angle or length
triceps brachii
Gastrocnemius lateralis 16.4% decrease in fascicle angle, 12.7% decrease in
fascicle length
Vastus lateralis 4.8° decrease in fascicle angle, 7.5mm decrease in
fascicle length
li[62] found that disuse atrophy of gastrocnemius
lateralis (23.1% reduction in anatomical cross-sec-
tional area) in an injured leg was associated with a
significant decrease in fascicle angle and length
(compared with uninjured leg; time period of injury
not reported). Also, although measurements were
taken at varying times after injury, Bleakney and
Maffulli[102] found significant decreases in fibre an-
gle and length of vastus lateralis in legs of subjects
recovering from tibia/femur fracture, compared with
the non-injured limb (time after fracture, mean = 7.6
months, range = 14 days to 2 years, 7 months). Abe
et al.[103] reported mixed results in subjects who
underwent 20 days of bed rest where there was a
decrease in gastrocnemius medialis fascicle angle
measured during standing (5.5%) and a non-signifi-
cant reduction during lying (4.7%), but no changes
in triceps brachii or vastus lateralis muscles. There-
fore, while there is some discrepancy in the litera-
ture, there is good evidence that fascicle geometry
changes can occur in response to unloading/de-
training in humans.
The reasons for the disparate results in de-
training/unloading studies are not clear; however,
the interaction effects of pre-unloading fascicle ge-
ometry and the muscle length during immobilisation
might be two factors affecting the magnitude of
geometric change during detraining. Studies on cat,
mouse, rat and rabbit muscles have shown signifi-
cant decreases in fibre length after periods of un-
loading when muscles were immobilised at lengths
shorter than normal rest length.[94,104-108] These ef-
fects have been reduced or reversed when muscles
Sports Med 2006; 36 (12)
1012
were immobilised in anatomical[109] or a lengthened
position.[93,106,108,109] Immobilisation in a lengthened
position has also been shown to counteract the slow-
to fast-twitch transformation that often accompanies
immobilisation[110] and reduces the loss of whole
muscle mass.[111] These data on animal muscles
show that the length of immobilisation has an im-
pact on muscle and fibre characteristics, although
this has not been fully examined in humans. In
particular, given that the fibre angle of a shortened
pennate muscle is greater than in a lengthened mus-
cle, one might speculate that immobilisation of mus-
cles in a shortened position might maintain the ge-
ometry of highly pennate, short-fibred muscles. Fi-
nally, given that immobilisation in lengthened
positions,[106] or the application of moderate stretch-
ing (30 min/day),[107] has been shown to prevent
fibre shortening during disuse, it is also pertinent to
determine whether such interventions might prevent
the fibre shortening that has sometimes been shown
to accompany disuse in humans.[62] Certainly the
addition of small amounts of stretching/mobilisa-
tion[99] during detraining has been shown to arrest
muscle atrophy in the human soleus.
3.3.4 Chronic Stretching
No research has examined changes in fascicle
geometry after acute or chronic stretching of mus-
cles in humans. Evidence from animal studies indi-
cates that chronic stretching might have a significant
effect on fibre length. Immobilisation of muscle or
muscle fibres in a lengthened position for periods of
days to weeks has resulted in increases in the num-
ber of serially arranged sarcomeres and overall fibre
length in animal muscles.[106,108,110] Interestingly,
moderate stretching performed for only 30 min/day
was enough to reduce or reverse the fibre shortening
that accompanied short-length immobilisation in rat
muscle.[107] It is not known whether flexibility train-
ing in humans results in an increase in fibre/fascicle
length. It is also not clear what effect, if any, chronic
stretching would have on fascicle angle.
 2006 Adis Data Information BV. All rights reserved.
Blazevich
4. Fascicle Geometry through Growth
and Aging
Fascicle geometric changes have yet to be de-
scribed in most human muscles through develop-
ment from infant to adult. Binzoni et al.[112] mea-
sured medial gastrocnemius fascicle angle in 134
subjects (87 men and 47 women) in the age range of
0–70 years. Very small fascicle angles at birth in-
creased through childhood and adolescence to reach
their maxima in early adulthood. A comparison of
bodyweight and height, and tibia length indicated a
strong relationship between fascicle angle and phys-
ical size, possibly indicating that increases in fasci-
cle angle of the gastrocnemius were related to the
loads imposed on it. No research has examined other
muscles, making it difficult to predict fascicle geo-
metric change with development, particularly in
non-weight-bearing muscles.
More data are available with regard to fascicle
geometry with aging from early adult to old age.
Aging is associated with a loss of muscle size, or
sarcopenia,[113-117] resulting from a loss of fibres/
motor units[118] and a significant reduction in fibre
area, particularly of the fast-twitch fibres.[116,119-121]
These morphological changes in muscle have been
implicated, along with possible reductions in muscle
activation,[122,123] in the significant loss of muscle
force[114,115,119,124-126] and power[114,115,126,127] seen in
aged individuals. Changes in fascicle geometry are
therefore likely given the positive relationship be-
tween fascicle angle and muscle size. Data collected
on 229 women from the ages of 20–79 years indicat-
ed that pennation of vastus lateralis (r = –0.50, p <
0.001), but not medial gastrocnemius or triceps
brachii, might be reduced with age, although there
was no evidence for a change in fascicle length
(relative to limb length).[128] The lesser pennation of
vastus lateralis in elderly compared with young, but
lack of differences in gastrocnemius or triceps
brachii, or in the length of fascicles in any of these
muscles, has also been reported for men.[129] In a
smaller (n21–32y = 19, n60–69y = 30), but well
matched (anthropometrically) sample, Karamanidis
and Arampatzis[61] found no differences in gas-
trocnemius medialis or vastus lateralis fascicle
Sports Med 2006; 36 (12)
Plasticity of Fascicle Geometry
lengths or angles, regardless of whether the subjects
were habitual endurance runners or not. Nonethe-
less, other data have shown significant differences
in gastrocnemius pennation (young [27–42 years] >
13.2% [elderly = 70–81 years]) and fascicle length
(young >10.2%) in subjects matched for height,
mass and physical activity.[117] These data suggest
that the aging process is accompanied by a signifi-
cant change in fascicle geometry that cannot neces-
sarily be explained by physical activity levels. Fur-
thermore, Morse et al.[130] showed that decreases in
muscle volume of soleus and medial and lateral
gastrocnemius were associated with decreases in
both fascicle angle and length. These decreases were
similar amongst the three muscles indicating that
these muscles, which fulfil different functional
roles, are affected similarly during aging. Thus,
while it seems that fascicle geometry changes from
adult to old age, more research is required to sub-
stantiate these early findings and further examine
inter-muscular differences.
The significant improvements in both muscle
strength[131-133] and power[127] after extended periods
of resistance exercise in older individuals are associ-
ated with increases in cross-sectional area or volume
of muscles.[127-130] This increase in cross-sectional
area is often largely explained by increases in areas
of both type I and II muscle fibres.[131,134] Few
researchers have examined the effects of changing
geometry on muscle size, or indeed the effects of
geometry on force development. One study[133] has
shown that an increase in muscle volume was attrib-
uted partly to an increase in fascicle length of the
vastus lateralis. This change largely accounted for
the increase in muscle PCSA, and suggests that
significant changes in geometry might occur with
training in aged subjects. In this case, the change
toward longer fascicle lengths is at odds with the
increases in fascicle angle (with no change or a
slight reduction in fascicle length) seen after training
in younger subjects. This might reflect an adaptation
in less mobile older subjects to a greater movement
range of motion used in exercise training compared
with their normal daily activities. More research is
required to determine how the changes in fascicle
 2006 Adis Data Information BV. All rights reserved.
1013
geometry shown with aging are influenced by exer-
cise interventions.
5. Conclusions and Future Directions
Human fascicle geometry is highly adaptable.
Heavy weight training in physically active individu-
als is often associated with increases in fascicle
angle (with some evidence for fascicle shortening)
with these changes amplified by anabolic steroid
use. In the elderly, the prominent adaptation seems
to be an increase in fascicle length, although it is not
clear how geometry changes in response to longer-
term training. Both detraining and aging are associ-
ated with decreases in fascicle angle and/or de-
creases in fascicle length, although there appears to
be some inter-muscular differences. These changes
have important consequences for force generation
since increases in fascicle angle allow higher muscle
forces, while increases in fascicle length allow
higher shortening speeds and for forces to be gener-
ated over larger length ranges.
While there are significant data showing the plas-
ticity of fascicle geometry, more research is required
to fully understand the effects of physical training.
Not least, no research has examined the effects of
endurance or chronic stretching training. Further-
more, in order to more fully understand how fascicle
geometry responds to heavy loading (e.g. resistance
training), a number of research lines need to be
explored. For example, despite a well established
link between fascicle geometry and function, and
significant evidence of geometric changes in re-
sponse to resistance training, relatively little re-
search has examined geometric changes in response
to athletic training interventions. Only one research
study[6] has examined changes in geometry in re-
sponse to training in athletic subjects who perform
mixed training regimens, and only one study[128] has
examined changes in older individuals, so the ef-
fects of training programme manipulations in ath-
letes and the aged remain relatively unstudied. Also,
no research has specifically investigated the effects
of training mode, contraction type or range of mo-
tion. It is unclear whether, for example, adaptations
to isokinetic training are different to isotonic/
Sports Med 2006; 36 (12)
1014
isoinertial training. Despite changes in sarcomere
number being reported after periods of eccentric
training in rats (downhill running),[135,136] which
might suggest that overall fibre/fascicle length also
increases with such training, no research has ex-
amined differences between the effects of concen-
tric, eccentric or isometric training programmes on
geometric adaptation. Also, while range of motion-
specific strength adaptations cannot be fully ex-
plained by neural mechanisms,[87] no research has
examined the effects of training range of motion. A
greater understanding of these parameters could
have significant benefits for injury/illness rehabilita-
tion in that maintenance of muscle strength during
relative immobilisation might accelerate physical
recovery. No less, a more complete understanding
might also inform athletic training practices.
Acknowledgements
Special thanks to Prof. Craig Sharp for his insightful
comments and discussion during the preparation of this arti-
cle. This research was partly funded by an American Society
of Biomechanics research grant (ASB-406015). The author
has no conflicts of interest that are directly related to the
content in this article.
References
1. Scott SJ, Engstrom CM, Loeb GE. Morphometry of human
thigh muscles: determination of fascicle architecture by mag-
netic resonance imaging. J Anat 1993; 182: 249-57
2. Kawakami Y, Abe T, Fukunaga T. Muscle-fiber pennation
angles are greater in hypertrophied than in normal muscles. J
Appl Physiol 1993; 74: 2740-4
3. Narici MV, Hoppeler H, Kayser B, et al. Human quadriceps
cross-sectional area, torque and neural activation during 6 mo
strength training. Acta Physiol Scand 1996; 157: 175-86
4. Aagaard P, Andersen JL, Dyhre-Poulsen P, et al. A mechanism
for increased contractile strength of human pennate muscle in
response to strength training: changes in muscle architecture. J
Physiol 2001; 534: 613-23
5. Blazevich AJ, Giorgi A. Effect of testosterone administration
and weight training on muscle architecture. Med Sci Sports
Exerc 2001; 33: 1688-93
6. Blazevich AJ, Gill ND, Bronks R, et al. Training-specific mus-
cle architecture adaptation after 5-wk concurrent training in
athletes. Med Sci Sports Exerc 2003; 35: 2013-22
7. Kawakami Y, Abe T, Kuno S-Y, et al. Training-induced
changes in muscle architecture and specific tension. Eur J Appl
Physiol 1995; 72: 37-43
8. Rutherford OM, Jones DA. Measurement of fibre pennation
using ultrasound in the human quadriceps in vivo. Eur J Appl
Physiol 1992; 65: 433-7
 2006 Adis Data Information BV. All rights reserved.
Blazevich
9. Burkholder TJ, Fingado B, Baron S, et al. Relationship between
muscle fiber types and sizes and muscle architectural proper-
ties in the mouse hindlimb. J Morphol 1994; 221: 177-90
10. Van Eijden TMGJ, Korfage JAM, Brugman P, et al. Architec-
ture of the human jaw-closing and jaw-opening muscles. Anat
Rec 1997; 248: 464-74
11. Lieber RL, Fridén J. Functional and clinical significance of
skeletal muscle architecture. Muscle Nerve 2000; 23: 1647-66
12. Hof AL. Muscle mechanics and neuromuscular control. J Bi-
omech 2003; 36: 1031-8
13. Hof AL, Geelen BA, Van den Berg J. Calf muscle moment,
work and efficiency in level walking: role of series elasticity. J
Biomech 1983, 37
14. Hof AL, Van Zandwijk JP, Bobbert MF. Mechanics of human
triceps surae muscle in walking, running and jumping. Acta
Physiol Scand 2002; 174: 17-30
15. Kawakami Y, Muraoka T, Ito S, et al. In-vivo muscle-fibre
behaviour during counter-movement exercise in humans
reveals significant role of tendon elasticity. J Physiol 2002;
540: 635-46
16. Kubo K, Kanehisa H, Takeshita D, et al. In vivo dynamics of
human medial gastrocnemius muscle-tendon complex during
strength-shortening cycle exercise. Acta Physiol Scand 2000;
170: 127-35
17. Kurokawa S, Fukunaga T, Fukashiro S. Behavior of fascicles
and tendinous structures of human gastrocnemius during verti-
cal jumping. J Appl Physiol 2001; 90: 1349-58
18. Lieber RL, Jacobson MD, Fazelle BM, et al. Architecture of
selected muscles of the arm and forearm: anatomy and impli-
cations for tendon transfer. J Hand Surg 1992; 17A: 787-98
19. Elliott BC, Blanksby BA. The synchronization of muscle activi-
ty and body segment movements during a running cycle. Med
Sci Sports 1979; 11: 322-7
20. Wiemann K, Tidow G. Relative activity of hip and knee exten-
sors in sprinting: implications for training. N Studies Athl
1995; 10: 29-49
21. Wood GA. Optimal performance criteria and limiting factors in
sprint running. N Studies Athl 1986; 2: 55-63
22. Friederich JA, Brand RA. Muscle fiber architecture in the
human lower limb. J Biomech 1990; 23: 91-5
23. Keller TCS. Molecular bungees. Nature 1997; 387: 233-5
24. Politou AS, Thomas DJ, Pastore A. The folding and stability of
titin immunoglobulin-like modules, with implications for the
mechanism of elasticity. Biophys J 1995; 96: 2601-10
25. Tskhovrebova L, Trinick J, Sleep JA, et al. Elasticity and
unfolding of single molecules of the giant muscle protein titin.
Nature 1997; 387: 308-12
26. Tskhovrebova L, Trinick J. Extensibility in the titin molecule
and its relation to muscle elasticity. Adv Exp Med Biol 2000;
481: 163-78
27. Linari M, Dobbie I, Reconditi M, et al. The stiffness of skeletal
muscle in isometric contraction and rigor: the fraction of
myosin heads bound to actin. Biophys J 1998; 74: 2459-73
28. Tawada K, Kimura M. Stiffness of glycerinated rabbit psoas
fibers in the rigor state: filament-overlap relation. Biophys J
1984; 45: 593-602
29. Huxley HE, Stewart A, Sosa H, et al. X-ray diffraction measure-
ments of the extensibility of actin and myosin filaments in
contracting muscle. Biophys J 1994; 67: 2411-21
30. Martyn DA, Chase PB, Regnier M, et al. A simple model with
myofilament compliance predicts activation-dependent cross-
bridge kinetics in skinned skeletal fibers. Biophys J 2002; 83:
3425-34
Sports Med 2006; 36 (12)
Plasticity of Fascicle Geometry
31. Wakabayashi K, Sugimoto Y, Tanaka H, et al. X-ray diffraction
evidence for the extensibility of actin and myosin filaments
during contraction. Biophys J 1997; 67: 2422-35
32. Schroeter P. P, Bretaudiere JP, Sass RL, et al. Three-dimension-
al structure of the Z band in a normal mammalian skeletal
muscle. J Cell Biol 1996; 133: 571-83
33. Goldstein MA, Michael LH, Schroeter JP, et al. The Z-band
lattice in skeletal muscle before, during and after tetanic con-
traction. J Muscle Res Cell Motil 1986; 7: 527-36
34. Fukunaga T, Miyatani M, Tachi M, et al. Muscle volume is a
major determinant of joint torque in humans. Acta Physiol
Scand 2001; 172: 249-55
35. Muhl ZF. Active length-tension relation and the effect of muscle
pinnation of fiber lengthening. J Morphol 1982; 173: 285-92
36. Herring SW, Grimm AF, Grimm BR. Regulation of sarcomere
number in skeletal muscle: a comparison of hypotheses. Mus-
cle Nerve 1984; 7: 161-73
37. Lutz GJ, Rome LC. Muscle function during jumping in frogs I:
sarcomere length change, EMG pattern, and jumping perform-
ance. Am J Physiol 1996; 271: C563-70
38. Lutz GJ, Rome LC. Muscle function during jumping in frogs II:
mechanical properties of muscle – implications for system
design. Am J Physiol 1996; 271: C571-8
39. Häkkinen K, Komi PV. Effect of explosive type strength train-
ing on electromyographic and force production characteristics
of leg extensor muscles during concentric and various stretch-
shortening cycle exercises. Scand J Sports Sci 1985; 7: 65-76
40. Häkkinen K, Komi PV. Electromyographic changes during
strength training and detraining. Med Sci Sports Exerc 1983;
15: 455-60
41. Hortobágyi T, Barrier J, Beard D, et al. Greater initial adapta-
tions to submaximal muscle lengthening than maximal short-
ening. J Appl Physiol 1996; 81: 1677-82
42. Moritani T, DeVries HA. Neural factors versus hypertrophy in
the time course of muscle strength gain. Am J Phys Med 1979;
58: 115-30
43. Patten C, Kamen G, Rowland D, et al. Rapid adaptations of
motor unit firing rate during the initial phase of strength
development. Med Sci Sports Exerc 1995; 27: S6
44. Ploutz LL, Tesch PA, Biro RL, et al. Effect of resistance training
on muscle use during exercise. J Appl Physiol 1994; 76:
1675-81
45. Enoka RM. Neural adaptations with chronic physical activity. J
Biomech 1997; 30: 447-55
46. Chilibeck PD, Calder AW, Sale DG, et al. A comparison of
strength and muscle mass increases during resistance training
in young women. Eur J Appl Physiol 1998; 77: 170-5
47. Hickson RC, Hidaka K, Foster C, et al. Successive time courses
of strength development and steroid hormone responses to
heavy-resistance training. J Appl Physiol 1994; 76: 663-70
48. Häkkinen K, Komi PV, Alén M, et al. EMG, muscle fibre and
force production characteristics during a 1 year training period
in elite weight-lifters. Eur J Appl Physiol 1987; 56: 419-27
49. Gaudy JF, Zouaoui A, Bravetti P, et al. Functional anatomy of
the human temporal muscle. Surg Radiol Anat 2001; 23:
389-98
50. Gaudy JF, Zouaoui A, Bravetti P, et al. Functional organization
of the human masseter muscle. Surg Radiol Anat 2000; 22:
181-90
51. Lam EW, Hannam AG, Christiansen EL. Estimation of tendon-
plane orientation within human masseter muscle from recon-
structed magnetic resonance images. Arch Oral Biol 1991; 36:
845-53
 2006 Adis Data Information BV. All rights reserved.
1015
52. Basser PJ, Mattiello J, LeBihan D. MR diffusion tensor spec-
troscopy and imaging. Biophys J 1994; 66: 259-67
53. van Donkelaar CC, Kretzers LJ, Bovendeerd PH, et al. Diffu-
sion tensor imaging in biomechanical studies of skeletal mus-
cle function. J Anat 1999; 194: 79-88
54. Mori S, Crain BJ, Chacko VP, et al. Three-dimensional tracking
of axonal projections in the brain by magnetic resonance
imaging. Ann Neurol 1999; 45: 265-9
55. Mori S, van Zijl PC. Fiber tracking: principles and strategies – a
technical review. NMR Biomed 2002; 15: 468-80
56. Heemskerk AM, Strijkers GJ, Vilanova A, et al. Determination
of mouse skeletal muscle architecture using three-dimensional
diffusion tensor imaging. Mag Res Med 2005; 53: 1333-40
57. van Doorn A, Bovendeerd PH, Nicolay K, et al. Determination
of muscle fibre orientation using diffusion-weighted MRI. Eur
J Morphol 1996; 34: 5-10
58. Damon BM, Ding Z, Anderson AW, et al. Validation of diffu-
sion tensor MRI-based muscle fiber tracking. Mag Res Med
2002; 48: 97-104
59. Cleveland GG, Chang DC, Hazlewood CF, et al. Nuclear mag-
netic resonance measurement of the intracellular water. Bi-
ophys J 1976; 16: 1043-53
60. Bensamoun SF, Ringleb SI, Littrell L, et al. Determination of
thigh muscle stiffness using magnetic resonance elastography.
J Mag Res Imag 2006; 23: 242-7
61. Karamanidis K, Arampatzis A. Mechanical and morphological
properties of human quadriceps femoris and triceps surae
muscle-tendon unit in relation to aging and running. J Biomech
2006; 39: 406-17
62. Narici M, Cerretelli P. Changes in human muscle architecture in
disuse-atrophy evaluated by ultrasound imaging. J Gravit
Physiol 1998; 5: 73-4
63. Chleboun GS, France AR, Crill MT, et al. In vivo measurement
of fascicle length and pennation angle of the human biceps
femoris muscle. Cells Tiss Org 2001; 169: 401-9
64. Narici MV, Binzoni T, Hiltbrand E, et al. Changes in human
gastrocnemius architecture with joint angle, at rest and with
isometric contraction, evaluated in vivo. J Physiol 1996; 496:
287-97
65. Kurihara T, Oda T, Chino K, et al. Use of three-dimensional
ultrasonography for the analysis of the fascicle length of
human gastrocnemius muscle during contractions. Int J Sport
Health Sci 2005; 3: 226-34
66. Muramatsu T, Muraoka T, Kawakami Y, et al. In vivo determi-
nation of fascicle curvature in contracting human skeletal
muscles. J Appl Physiol 2002; 92: 129-34
67. Reeves ND, Narici MV. Behavior of human muscle fascicles
during shortening and lengthening contractions in vivo. J Appl
Physiol 2003; 95: 1090-6
68. Finni T, Ikegawa S, Komi PV. Concentric force enhancement
during human movement. Acta Physiol Scand 2001; 173:
369-77
69. Finni T, Ikegawa S, Lepola V, et al. Comparison of force-
velocity relationships of vastus lateralis muscle in isokinetic
and in stretch-shortening cycle exercises. Acta Physiol Scand
2003; 177: 483-91
70. Izquerdo M, Ibañez J, Häkkinen K, et al. Maximal strength and
power, muscle mass, endurance and serum hormones in
weightlifters and road cyclists. J Sport Sci 2004; 22: 465-78
71. Izquierdo M, Häkkinen K, Gonzalez-Badillo JJ, et al. Effects of
long-term training specificity on maximal strength and power
of the upper and lower extremities in athletes from different
sports. Eur J Appl Physiol 2002; 87: 264-71
Sports Med 2006; 36 (12)
1016
72. Coyle EF. Improved muscular efficiency displayed as Tour de
France champion matures. J Appl Physiol 2005; 98: 2191-6
73. Alexander MJL. The relationship between muscle strength and
sprint kinematics in elite sprinters. Can J Sport Sci 1989; 14:
148-57
74. Taylor NAS, Cotter JD, Stanley SN, et al. Functional torque-
velocity and power-velocity characteristics of elite athletes.
Eur J Appl Physiol 1991; 62: 116-21
75. Blazevich AJ, Jenkins DG. Physical performance differences
between weight-trained sprinters and weight trainers. J Sci
Med Sport 1998; 1: 12-21
76. Sleivert GG, Backus RD, Wenger HA. Neuromuscular differ-
ences between volleyball players, middle distance runners and
untrained controls. Int J Sport Med 1995; 16: 390-8
77. Kumagai K, Abe T, Brechue WF, et al. Sprint performance is
related to muscle fascicle length in male 100-m sprinters. J
Appl Physiol 2000; 88: 811-6
78. Abe T, Fukashiro S, Harada Y, et al. Relationship between
sprint performance and muscle fascicle length in female sprint-
ers. J Physiol Anthropol Appl Human Sci 2001; 20: 141-7
79. Abe T, Kumagai K, Brechue WF. Fascicle length of leg muscles
is greater in sprinters than distance runners. Med Sci Sports
Exerc 2000; 32: 1125-9
80. Brechue WF, Abe T. The role of FFM accumulation and skeletal
muscle architecture in powerlifting performance. Eur J Appl
Physiol 2002; 86: 327-36
81. Kearns CF, Abe T, Brechue WF. Muscle enlargement in sumo
wrestlers includes increased muscle fascicle length. Eur J Appl
Physiol 2000; 83: 289-96
82. Gondin J, Guette M, Ballay Y, et al. Electromyostimulation
training effects on neural drive and muscle architecture. Med
Sci Sports Exerc 2005; 37: 1291-9
83. Alegre LM, Jimenez F, Gonzalo-Orden JM, et al. Effects of
dynamic resistance training on fascicle length and isometric
strength. J Sports Sci 2006; 24: 501-8
84. Blazevich AJ, Gill ND, Deans N, et al. Lack of human muscle
architectural adaptation after short-term strength training.
Muscle Nerve. Epub 2006 Oct 12
85. Häkkinen K, Kallinen M, Linnamo V, et al. Neuromuscular
adaptations during bilateral versus unilateral strength training
in middle-aged and elderly men and women. Acta Physiol
Scand 1996; 158: 77-88
86. Tanaguchi Y. Laterality specificity in resistance training: the
effect of bilateral and unilateral training. Eur J Appl Physiol
1997; 75: 144-50
87. Weir JP, Housh DJ, Housh TJ, et al. The effect of unilateral
concentric weight training and detraining on joint angle speci-
ficity, cross-training, and the bilateral deficit. J Orthop Sports
Phys Ther 1997; 25: 264-70
88. Jarvholm U, Palmerud G, Karlsson D, et al. Intramuscular
pressure and electromyography in four shoulder muscles. J
Orthop Res 1991; 9: 609-19
89. Sejersted OM, Hagens AR. Intramuscular pressures for moni-
toring different tasks and muscle conditions. Adv Expl Med
Biol 1995; 384: 339-50
90. Miura H, McCully K, Nioka S, et al. Relationship between
muscle architectural features and oxygenation status deter-
mined by near infrared device. Eur J Appl Physiol 2004; 91:
273-8
91. Ballard RE, Watenpaugh DE, Breit GA, et al. Leg intramuscular
pressures during locomotion in humans. J Appl Physiol 1998;
84: 1976-81
 2006 Adis Data Information BV. All rights reserved.
Blazevich
92. Aratow M, Ballard RE, Crenshaw AG, et al. Intramuscular
pressure and electromyography as indexes of force during
isokinetic exercise. J Appl Physiol 1993; 74: 2634-40
93. Degens H, Salmons S, Jarvis JC. Intramuscular pressure, force
and blood flow in rabbit tibialis anterior muscles during single
and repetitive contractions. Eur J Appl Physiol Occup Physiol
1998; 78: 13-9
94. Jarvinen MJ, Einola SA, Virtanen EO. Effect of the position of
immobilization upon the tensile properties of the rat gas-
trocnemius muscle. Arch Phys Med Rehab 1992; 73: 253-7
95. Sejersted OM, Hagens AR, Kardel KR, et al. Intramuscular fluid
pressure during isometric contraction of human skeletal mus-
cle. J Appl Physiol 1984; 56: 287-95
96. Akima H, Kuno S, Suzuki Y, et al. Effects of 20 days of bed rest
on physiological cross-sectional area of human thigh and leg
muscles evaluated by magnetic resonance imaging. J Gravit
Physiol 1997; 4: S15-21
97. Kawakami Y, Akima H, Kubo K, et al. Changes in muscle size,
architecture, and neural activation after 20 days of bed rest
with and without resistance exercise. Eur J Appl Physiol 2001;
84: 7-12
98. Kawakami Y, Muraoka Y, Kubo K, et al. Changes in muscle
size and architecture following 20 days of bed rest. J Gravit
Physiol 2000; 7: 53-60
99. Yamashita-Goto K, Okuyama R, Honda M, et al. Maximal and
submaximal forces of slow fibers in human soleus after bed
rest. J Appl Physiol 2001; 91: 417-24
100. Andersen JL, Gruschy-Knudsen T, Sandri C, et al. Bed rest
increases the amount of mismatched fibers in human skeletal
muscle. J Appl Physiol 1999; 86: 455-60
101. Heslinga JW, Huijing PA. Effects of short length immobiliza-
tion of medial gastrocnemius muscle of growing young adult
rats. Eur J Morphol 1992; 30: 257-73
102. Bleakney R, Maffulli N. Ultrasound changes to intramuscular
architecture of the quadriceps following intramedullary nail-
ing. J Sports Med Phys Fit 2002; 42: 120-5
103. Abe T, Kawakami Y, Suzuki Y, et al. Effects of 20 days bed rest
on muscle morphology. J Gravit Physiol 1997; 4: S10-4
104. Baker JH, Matsumoto DE. Adaptation of skeletal muscle to
immobilization in a shortened position. Muscle Nerve 1988;
11: 231-44
105. Hayat A, Tardieu C, Tabary JC, et al. Effects of denervation on
the reduction of sarcomere number in cat soleus muscle immo-
bilized in shortened position during seven days. J Physiol
(Paris) 1978; 74: 563-7
106. Tabary JC, Tabary C, Tardieu C, et al. Physiological and struc-
tural changes in the cat’s soleus muscle due to immobilization
at different lengths by plaster casts. J Physiol 1972; 224:
231-44
107. Williams PE. Use of intermittent stretch in the prevention of
serial sarcomere loss in immobilised muscle. Ann Rheum Dis
1990; 49: 316-7
108. Williams PE, Goldspink G. Changes in sarcomere length and
physiological properties in immobilized muscle. J Anat 1978;
127: 459-68
109. Nicks DK, Beneke WM, Key RM, et al. Muscle fibre size and
number following immobilisation atrophy. J Anat 1989; 163:
1-5
110. Leterme D, Cordonnier C, Mounier Y, et al. Influence of chron-
ic stretching upon rat soleus muscle during non-weight-bear-
ing conditions. Eur J Physiol 1994; 429: 274-9
Sports Med 2006; 36 (12)
Plasticity of Fascicle Geometry
111. Herbert RD, Balnave RJ. The effect of position of immobilisa-
tion on resting length, resting stiffness, and weight of the
soleus muscle of the rabbit. J Orthop Res 1993; 11: 358-6
112. Binzoni T, Bianchi S, Hanquinet S, et al. Human gastrocnemius
medialis pennation angle as a function of age: from newborn to
the elderly. J Physiol Anthropol Appl Hum Sci 2001; 20: 293-8
113. Häkkinen K, Pakarinen A. Muscle strength and serum hormones
in middle-aged and elderly men and women. Acta Physiol
Scand 1993; 148: 199-207
114. Izquierdo M, Ibañez J, Gorostiaga E, et al. Maximal strength
and power characteristics in isometric and dynamic actions of
the upper and lower extremities in middle-aged and older men.
Acta Physiol Scand 1999; 167: 57-68
115. Izquerdo M, Häkkinen K, Antón A, et al. Maximal strength and
power, endurance performance, and serum hormones in mid-
dle-aged and elderly men. Med Sci Sports Exerc 2001; 33:
1577-87
116. Klitgaard H, Mantoni M, Schiaffino S, et al. Function, morphol-
ogy and protein expression of ageing skeletal muscle: a cross-
sectional study of elderly men with different training back-
grounds. Acta Physiol Scand 1990; 140: 41-54
117. Narici MV, Maganaris CN, Reeves ND, et al. Effect of aging on
human muscle architecture. J Appl Physiol 2003; 95: 2229-34
118. Lexell J, Taylor CC, Sjostrom M. What is the cause of the aging
atrophy? Total number, size and proportion of different fiber
types studies in whole vastus lateralis muscle from 15- to
83-year-old men. J Neurol Sci 1988; 84: 275-94
119. Hortobágyi T, Zheng D, Weidner M, et al. The influence of
aging on muscle strength and muscle fiber characteristics with
special reference to eccentric strength. J Gerontol A: Biol Sci
Med Sci 1995; 50: B399-406
120. Larsson L, Grimby G, Karlsson J. Muscle strength and speed of
movement in relation to age and muscle morphology. J Appl
Physiol 1979; 46: 451-6
121. Poggi P, Marchettic C, Scelsi R. Automatic morphometric anal-
ysis of skeletal muscle fibers in the aging man. Anat Rec 1987;
217: 30-4
122. Häkkinen K, Alen M, Kallinen M, et al. Neuromuscular adapta-
tion during prolonged strength training, detraining and re-
strength-training in middle-aged and elderly people. Eur J
Appl Physiol 2000; 83: 51-62
123. Häkkinen K, Kallinen M, Izquierdo M, et al. Changes in ago-
nist-antagonist EMG, muscle CSA, and force during strength
training in middle-aged and older people. J Appl Physiol 1998;
84: 1341-9
124. Knight CA, Kamen G. Adaptations in muscular activation of the
knee extensor muscles with strength training in young and
older adults. J Electromyogr Kinesiol 2001; 11: 405-12
 2006 Adis Data Information BV. All rights reserved.
1017
125. Roos MR, Rice CL, Connelly DM, et al. Quadriceps muscle
strength, contractile properties, and motor unit firing rates in
young and old men. Muscle Nerve 1999; 22: 1094-103
126. Valour D, Ochala J, Ballay Y, et al. The influence of ageing on
the force-velocity-power characteristics of human elbow flex-
or muscles. Exp Gerontol 2003; 38: 387-95
127. Newton RU, Häkkinen K, Häkkinen A, et al. Mixed-methods
resistance training increases power and strength of young and
older men. Med Sci Sports Exerc 1999; 34: 1367-75
128. Kubo K, Kanehisa H, Azuma K, et al. Muscle architectural
characteristics in women aged 20-79 years. Med Sci Sports
Exerc 2003; 35: 39-44
129. Kubo K, Kanehisa H, Azuma K, et al. Muscle architectural
characteristics in young and elderly men and women. Int J
Sports Med 2003; 24: 125-30
130. Morse CI, Thom JM, Birch KM, et al. Changes in triceps surae
muscle architecture with sarcopenia. Acta Physiol Scand 2005;
183: 291-8
131. Häkkinen K, Kraemer WJ, Pakarinen A, et al. Effects of heavy
resistance/power training on maximal strength, muscle mor-
phology, and hormonal response patterns in 60-75-year-old
men and women. Can J Appl Physiol 2002; 27: 213-31
132. Izquerdo M, Häkkinen K, Ibañez J, et al. Effects of strength
training on submaximal and maximal endurance performance
capacity in middle-aged and older men. J Strength Cond Res
2003; 17: 129-39
133. Reeves ND, Narici MV, Maganaris CN. Effect of resistance
training on skeletal muscle-specific force in elderly humans. J
Appl Physiol 2004; 96: 885-92
134. Trappe S, Williamson D, Godard M, et al. Effect of resistance
training on single muscle fiber contractile function in older
men. J Appl Physiol 2000; 89: 143-52
135. Lynn R, Morgan DL. Decline running produced more sarcomer-
es in rat vastus intermedius muscle fibers than does incline
running. J Appl Physiol 1994; 77: 1439-44
136. Butterfield TA, Leonard TR, Herzog W. Differential serial
sarcomere number adaptations in knee extensor muscles of rats
is contraction type dependent. J Appl Physiol 2005; 99: 1352-8
Correspondence and offprints: Dr Anthony J. Blazevich,
Centre for Sports Medicine and Human Performance,
Brunel University, Uxbridge, UB8 3PH, UK.
E-mail: anthony.blazevich@brunel.ac.uk
Sports Med 2006; 36 (12)