Vous êtes sur la page 1sur 15

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 Blazevich

Historically, changes in human fascicle geome- changes after training and detraining interventions
try1 have been impossible to examine since the and/or aging and development.
excision and subsequent study of muscle could only
be performed in cadavers. More recently, imaging
techniques such as magnetic resonance (MRI)[1] and 1. Geometry of Human Muscles
ultrasound[2,3] imaging (figure 1) have allowed the in
vivo examination of geometric, i.e. fascicle length Many human muscles contain fascicles that do
and angle, changes.[2,4-8] Results of such research not run directly from origin to insertion, but are
have predominately shown geometry to be highly angled and attach to the muscles’ aponeuroses (fig-
changeable. An understanding of how different ure 2). These ‘pennate’ muscles can have complex
forms of muscle loading, or exercise training, affects fascicular arrangements that differ markedly be-
geometry would allow us to bring about deliberate tween muscles. Their fascicle arrangement is proba-
and specific changes that would influence the mus- bly a greater determinant of general muscle function
cles’ force-generating properties. There are, howev- than other aspects such as fibre type.[9] As a general
er, a limited number of studies that have examined rule, muscles with large fascicle attachment angles
the plasticity of fascicle geometry, or that have and correspondingly short fascicles are suited to
compared populations of individuals who have per- contractions involving high forces[10] and/or pro-
formed different exercise training. The present re- duce forces over a relatively short range of mo-
view summarises research that quantifies changes in tion.[11] Examples include the large proximal mus-
fascicle geometry with exercise and detraining, and cles of the legs (e.g. vastus lateralis; see table I),
development and aging. A greater understanding of smaller muscles such as those involved in mastica-
fascicle geometric adaptation could inform both tion,[10] and those involved in postural control such
physical performance and rehabilitative training as soleus and tibialis posterior. Large fascicle angles
programmes, as well as allowing the development of are also seen in muscles that attach to long tendons,
muscle models that predict physical performance such as gastrocnemius lateralis, which produce suf-
ficient force to remain relatively isometric during
the propulsive phase of stretch-shorten contractions

θ PCSA
b  in walking,[12-14] running[12,14] and jumping with[15,16]
and without[12,14,17] countermovement, and allow op-
β
α  ACSA
timum elongation and recoil of the tendon.[12] Mus-

lf = ∆a − b a  cles that commonly participate in movements re-


quiring significant length change or a high shorten-
Muscle length
ing velocity tend to be characterised by long
Fig. 1. Muscle architectural parameters include: fibre length = dis- fascicles attaching at relatively small angles to the
tance between ends of a fibre (a to b); pennation angle (θ) = tendon.[9,18] Examples include adductor magnus and
fascicle angle (relative to the aponeurosis [α]) minus the aponeuro- longus, whose longer fascicles attaching at smaller
sis angle (relative to the tendon [β]); muscle length; and anatomical
(ACSA) or physiological (PCSA) cross-sectional area. The PCSA angles allows force generation over a large range of
can be calculated as (V/t) • sinθ for a simple, uni-pennate muscle, motion at the hip during flexion/adduction tasks,
where V is the muscle volume, t is the muscle thickness from one
and the topologically similar long head of biceps
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 femoris, which is highly active during sprint run-
mean fibre/fascicle length. ning.[19-21]

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. 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-


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

 2006 Adis Data Information BV. All rights reserved. Sports Med 2006; 36 (12)
1006 Blazevich

Table I. Examples of fascicle angles and lengths in human muscles (average of two cadavers)[22]
Force characteristics Action Muscle Fascicle angle Fascicle length
(°) (mm)
High Force generation Vastus lateralis 13 80.3
Prolonged (time) Postural control Soleus 32 30.3
Tibialis posterior 19 28.6
Rapid/high Stretch-shorten Gastrocnemius lateralis 17.5 60.6
High velocity/large range of motion Long-range force Adductor magnus 4 113.0
Adductor longus 3.5 94.2
Eccentric/large range of motion Long-range force Biceps femoris (long head) 7 72.6

the highest forces are required (rather than at the accepted, it fails to take into account changes in
muscle’s resting length for example; Herring et fascicle geometry as a possible contributing factor to
al.[36] [human]; Lutz and Rome[37,38] [frog]), fibres strength adaptation. Since fascicle geometry signifi-
that shorten less for a given tendon excursion are cantly impacts on force generation, loading-depen-
likely to stay closer to their optimum for force dent geometric changes would influence force pro-
generation. Third, since the time-dependent shorten- duction and ultimately influence movement capaci-
ing distance of a fibre decreases during a contrac- ty. There are two ways in which information
tion, the shortening speed will also decrease (v = d/t, regarding changes in fascicle geometry can be
where v = velocity of shortening, d = fibre displace- gained, one is to examine differences between popu-
ment and t = time) and force increases as per the lations who perform different training regimes, and
force-velocity relationship for muscle. So in pennate another is to measure its changes in response to
muscle, a more forceful contraction might be possi- training.
ble from exploitation of both the force-velocity and
length-force relationships in addition to the greater 3.1 Measuring Fascicle Geometry in Humans
quantity of contractile material that can attach to the
tendon or aponeurosis. It is clear then that having MRI has been scarcely used to measure fascicle
large fibre angles would be an important adaptation geometry in humans. The striation patterns resulting
to allow high force production. from inter-fascicular fat and connective tissue ele-
ments can be visualised using MRI scanning since
3. Plasticity of Human Muscle Geometry their signal (brightness in the image) differs from
the muscle tissue because of variations in water
Tissues within the human body have a great content of the tissues. The reconstruction of separate
ability to remodel in response to stress (e.g. Wolff’s two-dimensional (2-D) images, with the striation
Law for bone). At the gross level of muscle, in- patterns visually marked, into three dimensions
creased loading is commonly associated with an (3-D) allows the accurate determination of fascicle
early (<6 weeks) change in their neural activation geometry. Scott et al.[1] found good agreement be-
during heavy lifting[3,38-44] (for a review of neural tween MRI and dissection estimates of fibre geome-
mechanisms see Enoka[45]). However, subsequent try in the quadriceps of human cadavers, while
periods of loading result in an increase in muscle Gaudy et al.[49,50] reported the muscle architectural
size with more contractile tissue being available for arrangements of masseter, temporalis and lateral
force generation.[3,39,42,46,47] After prolonged periods pterygoid, with little variance between cadaver dis-
of training, when the rate of hypertrophy has section and MRI measurements. Lam et al.[51] vali-
reached a plateau, further small strength increases dated the measurement of fibre angles of the masse-
are most likely to occur by neural mechanisms in ter by MRI in a single human cadaver. To date, no
response to variations in training intensity or vol- studies have examined fascicle geometry in athletes,
ume.[48] While this sequence of adaptation is well or determined longitudinal changes in fascicle ge-

 2006 Adis Data Information BV. All rights reserved. Sports Med 2006; 36 (12)
Plasticity of Fascicle Geometry 1007

ometry using MRI, perhaps because of the difficulty cles, larger errors in fascicle length estimates have
of manually tracing fascicles and then reconstruct- been observed when compared with 3-D ultrasound
ing 3-D images. images (2.4–14.0%).[65] Fascicle length is also diffi-
cult to accurately measure when the width of the
More recently, other MRI-based techniques have
probe (often 3.8 or 6cm) is insufficient to capture an
been developed. Diffusion tensor imaging relies on
entire fascicle. In these instances, linear approxima-
the diffusion of water in the muscle being con-
tions are calculated from muscle thickness and fasci-
strained by membranes and other cellular constitu-
cle angle measures (length = muscle thickness/sin ×
ents, so in muscles the diffusion occurs largely along
fascicle angle). These do not account for fibre curva-
the plane of the fibres. The diffusion tensor can be
ture that is present in some hypertrophied or con-
calculated from the measurement of the direction of
tracting muscles and errors range from approximate-
the diffusion.[52,53] This technique is more efficient
ly 0% (non-contracted gastrocnemius) to 6.6% (con-
than traditional methods of MRI determination be-
tracted gastrocnemius) according to Muramatsu et
cause the fibres can be tracked rapidly by a fibre-
al.,[66] or 2.4% in the non-contracted tibialis anterior
tracking algorithm[54,55] and the fibre orientations
muscle.[67] Fascicle length can also be estimated by
can be quickly reconstructed. While the technique
measuring the visible portion directly and then esti-
has been used to determine fascicle geometry in
mating the non-visible portion. Errors of 2–7% have
animal muscles,[53,56-59] it has yet to be used to
been reported for this method.[68,69] Regardless, ul-
quantify human fascicle geometry adaptations. In
trasound imaging techniques allow valid and relia-
addition to the diffusion tensor technique, magnetic
ble measures of fascicle geometry in vivo, and have
resonance elastography has recently been shown to
been commonly used to examine population differ-
provide information with regard to fascicle geome-
ences and longitudinal adaptations.
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 3.2 Population Differences in
spread of shear waves travelling through the muscle. Fascicle Geometry
The spread of waves is constrained by the stiffness
of muscle-based tissues, where lateral (with respect A muscle’s phenotype is at least partly reflective
to the muscle fibres) transmission is affected by of its long-term activity patterns. Although a per-
parallel layers of connective tissue but longitudinal son’s genetic predisposition might play a role, it is
transmission is relatively unimpeded along the fib- a b
res. Again, this technique has not yet been used to Skin/fat layer
quantify muscle fascicle geometric changes longitu-
dinally; however, both diffusion tensor imaging and Vastus
lateralis
elastography show great promise as fascicle imag-
ing techniques.
Vastus
The most common method of measuring fascicle intermedius
geometry is via the acquisition of 2-D longitudinal Bone
images of the muscle using ultrasound[2,4-7,61,62] (fig- Fig. 3. Images of human muscles can be obtained using ultrasound
ure 3). When the ultrasound transducer (probe) is imaging. In this longitudinal section of the thigh, the skin/fat layer,
femur (bone), and vastus lateralis and vastus intermedius muscles
oriented in the plane of the fascicles, the fascicle are clearly visible. The aponeurosis (apo) and line of fascicles (fas)
angles and lengths can be accurately measured. are also visible, from which fascicle angle can be determined, and
Measures of fascicle angle (≤1.5°)[2,63,64] and fasci- fascicle length can be estimated using standard trigonometric pro-
cedures.[5-7,68,69] Physical training can alter a muscle’s fascicle ge-
cle length (≤1.5mm)[2] have been shown to be simi- ometry. Here, a period of heavy strength training has resulted in an
lar to those measured directly in cadavers. When the increase in fascicle angle concomitant with the increase in muscle
transducer is not oriented in the plane of the fasci- thickness (a vs b).

 2006 Adis Data Information BV. All rights reserved. Sports Med 2006; 36 (12)
1008 Blazevich

expected that the physical work capacity of athletes, relatively regardless of cost. These data would sug-
who have consistently trained with specific move- gest that there are significant differences in the
ment patterns for a period of many years, would fascicle geometry of well trained sprinters compared
vary according to the adaptations elicited by their with lesser-trained sprinters, endurance runners or
training. Indeed, significant, training-specific differ- control subjects.
ences in work capacity have been shown among Unique geometry is also seen in weight-trained
groups of athletes with varied training histo- individuals. As expected, muscle thickness mea-
ries.[70-76] Thus, one method of considering training- sured at 13 sites on muscles including triceps
dependent fascicle geometric adaptations is to com- brachii, vastus lateralis and gastrocnemius lateralis
pare populations with different physical activity his- of powerlifters was significantly correlated with
tories. squat, bench-press and deadlift performance (r =
0.63–0.91[80]). However, fascicle length of the vas-
A significant quantity of research has examined
tus lateralis was also positively correlated with squat
differences in fascicle geometry between different
lift (r = 0.50) and deadlift (r = 0.54) strength, and
athlete populations. For example, both vastus later-
triceps fascicle length was correlated with bench-
alis and gastrocnemius in highly trained male sprint-
press strength (r = 0.52). While muscle thickness
ers (100m in <11.0 seconds) were found to contain
and fascicle angle were positively correlated (triceps
longer fascicles attached at smaller angles than in
brachii r = 0.64, p < 0.01; gastrocnemius r = 0.48, p
lesser-trained sprinters (100m in >11.0 seconds[77]).
< 0.05), triceps brachii fascicle angle was negatively
In females, sprinters have been shown to have small-
correlated with bench-press performance (relative to
er fascicle angles in vastus lateralis than non-trained
fat-free mass, r = –0.45) and gastrocnemius fascicle
controls (with no differences in gastrocnemius) and
angle was negatively correlated with deadlift per-
longer fascicles in vastus lateralis and gastrocnemi- formance (relative to fat-free mass, r = –0.56). In
us lateralis, which were significantly correlated with another study, bodybuilders were shown to have
100m best time (r = –0.51 and r = –0.44, respective- greater muscle thickness and fascicle angles than
ly[78]). A comparison of male and female sprinters non-weight-trained controls.[2] There is also some
revealed no differences in absolute or relative (to leg evidence that muscle size, fascicle angle and fasci-
length) fascicle length.[78] When the fascicle geome- cle length are simultaneously greater in athletes who
try of sprinters was compared with well trained require a large body mass and physical strength as
endurance runners, differences were also seen.[79] well as fast movement velocities (e.g. Sumo wres-
While endurance runners (13.5–14.5 minutes for tlers[81]), than they are in the normal population.
5000m) had a lesser muscle thickness in the vastus
lateralis and gastrocnemius muscles than sprinters 3.3 Training Effects on Fascicle Geometry
(10.0–10.9 seconds for 100m), their fascicles were
also shorter and their fascicle angles greater. Com- While the assessment of population differences
pared with non-active subjects, young endurance provides an insight into possible activity-dependent
runners had larger fascicle angles (13.7%) but a adaptations in fascicle geometry, a more satisfactory
similar fascicle length in gastrocnemius medialis; method of examining the plasticity of fascicle geom-
there were no differences in fascicle geometry in the etry is to directly measure the longitudinal adapta-
vastus lateralis.[61] At least for the gastrocnemius, tions.
muscles in endurance runners appear to be adapted 3.3.1 Resistance Training
to produce forces with minimal metabolic cost (i.e. Early research was suggestive of there being little
short fibres or fascicles require less energy for a change in geometry in response to resistance train-
given level of force production because they have ing. Rutherford and Jones[8] showed no change in
fewer sarcomeres in series), whereas in sprinters vastus lateralis or intermedius fascicle angles after 3
they are adapted for high-speed force development months of quadriceps strength training in seven men

 2006 Adis Data Information BV. All rights reserved. Sports Med 2006; 36 (12)
Plasticity of Fascicle Geometry 1009

Table II. Muscle architecture changes after strength training in humansa


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

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

 2006 Adis Data Information BV. All rights reserved. Sports Med 2006; 36 (12)
1010 Blazevich

lateralis fascicle length increased only for the sprint/ no changes in muscle size and geometry of elbow
jump-trained group. Muscle thickness of both vastus extensors of an untrained limb in five men after 16
lateralis and rectus femoris increased in all training weeks of training, although no strength changes
groups also, although there were no differences in were found either. Blazevich et al.[84] have reported
performance (sprint, jump and strength tests) be- that significant increases in contralateral knee exten-
tween the groups after training. Similar to previous sor strength after 5 weeks of isokinetic training
studies on untrained subjects,[7,83] these results show occurred without a change in geometry. Thus, al-
that changes in fascicle geometry occur in response though few data are available, there is no current
to resistance training in well trained subjects. The evidence to suggest that fascicle geometric changes
results may have differed from previous studies are a factor affecting short-term strength increases in
where little change was noticed in trained subjects a contralateral limb.
because the training programme differed sufficient-
ly from that which the athletes had previously per- 3.3.2 Endurance Training
formed. These data were the first to show short-term No research has examined changes in geometry
changes in geometry in response to training, and after long-term endurance training. It is therefore
also show that increases in fascicle length and de- not known whether endurance training can elicit
creases in fascicle angle can occur after high-veloc- geometric adaptations, or how different modes of
ity movement training in humans; it is not known, training (e.g. cycling, running, lifting) might affect
however, whether other factors such as the training them. It makes intuitive sense that if a muscle could
range of motion influence the architectural adapta- generate force more efficiently, then muscle endur-
tion. ance would improve. Given that muscles with large
pennation and shorter fascicles would generate force
Given that rapid (5-week) changes in geometry
with less metabolic cost, such an adaptation might
have been shown to occur in athletes, one might
be expected with endurance training. Certainly, effi-
speculate that rapid adaptation is a mechanism by
ciency during stretch-shortening contractions could
which early strength increases occur in previously
be expected to increase following such a geometric
untrained subjects. However, recent data[84] showed
adaptation. This reasoning is consistent with the
no changes in quadriceps muscle size or geometry in
finding that vastus lateralis and gastrocnemius fasci-
previously untrained men (n = 7) and women (n = 8)
cles were shorter and fascicle angles greater in a
despite significant increases in concentric and ec-
group of endurance runners compared with sprint-
centric isokinetic strength of the quadriceps muscle ers.[79] However, it could also be argued that longer
group after 5 weeks of isokinetic knee extensor fibres would benefit muscles that are commonly
training. Also, Gondin et al.[82] did not find signifi- recruited during endurance tasks where forces are
cant increases in vastus lateralis fascicle angle after produced over large ranges of motion, since such
4 weeks of electromyostimulation training, despite fibre geometry is uniquely adapted to perform such
there being significant increases after 8 weeks. It is work. Also, intramuscular pressure (IMP) is greatest
therefore likely that changes in fascicle geometry in muscles with large fibre angles;[88,89] therefore,
are not a significant factor influencing early strength occlusion of blood flow during muscle contraction
increases in previously untrained subjects, although would be greater. This hypothesis is consistent with
whether rapid adaptations occur after training using the decreases in blood volume and oxygen satura-
other modes (e.g. isotonic/isoinertial) is not tion seen during exercise (1Hz plantar flexion) in the
known.[7] distal region of gastrocnemius medialis, where fas-
Geometric changes are probably not a factor in- cicle angle and length are greatest.[90] During loco-
volved in strength increases that have been reported motion, IMP increases with movement speed[91] in
in the untrained contralateral limb in response to line with the known positive relationship between
unilateral training.[85-87] Kawakami et al.[7] reported IMP and muscle force,[92-95] so the limitation of

 2006 Adis Data Information BV. All rights reserved. Sports Med 2006; 36 (12)
Plasticity of Fascicle Geometry 1011

Table III. Muscle architecture changes after detraining/unloading in humansa


Study Method Muscles Change
Abe et al.[103] 20d bed rest 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)
Kawakami et al.[98] 20d bed rest Vastus lateralis, plantarflexors, No change in fascicle angle or length
triceps brachii
Narici and Cerretelli[62] Disuse/injury Gastrocnemius lateralis 16.4% decrease in fascicle angle, 12.7% decrease in
fascicle length
Bleakney and Maffulli[102] Disuse/injury Vastus lateralis 4.8° decrease in fascicle angle, 7.5mm decrease in
fascicle length
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 li[62] found that disuse atrophy of gastrocnemius
be expected to impact on muscle endurance. Of lateralis (23.1% reduction in anatomical cross-sec-
course, during most forms of locomotion, muscle tional area) in an injured leg was associated with a
contractions are phasic and blood flow will not be significant decrease in fascicle angle and length
significantly affected when the duty cycle is below a (compared with uninjured leg; time period of injury
threshold limit (e.g. <40%),[93] so the effect of ge- not reported). Also, although measurements were
ometry on endurance performance is probably com- taken at varying times after injury, Bleakney and
plex. It would seem likely that fibre type is a more Maffulli[102] found significant decreases in fibre an-
important muscle-based determinant of muscle en- gle and length of vastus lateralis in legs of subjects
durance while fascicle geometry affects more the recovering from tibia/femur fracture, compared with
length-tension and force-velocity characteristics of a the non-injured limb (time after fracture, mean = 7.6
muscle. months, range = 14 days to 2 years, 7 months). Abe
et al.[103] reported mixed results in subjects who
3.3.3 Detraining/Unloading
underwent 20 days of bed rest where there was a
Numerous studies have investigated the effect of decrease in gastrocnemius medialis fascicle angle
detraining/unloading on muscle size and strength.
measured during standing (5.5%) and a non-signifi-
For example, significant reductions in strength and
cant reduction during lying (4.7%), but no changes
muscle size follow prolonged bed rest[96-98] along
in triceps brachii or vastus lateralis muscles. There-
with decreases in fibre size[99] and changes in fibre
fore, while there is some discrepancy in the litera-
type (slow- to fast-twitch transformation[100]). How-
ture, there is good evidence that fascicle geometry
ever, few studies have measured fascicle geometry
changes can occur in response to unloading/de-
after a period of detraining/unloading, or have not
training in humans.
reported changes in geometry when it has been
measured for the determination of PCSA. In one The reasons for the disparate results in de-
study, Kawakami et al.[98] found no changes in vas- training/unloading studies are not clear; however,
tus lateralis, plantarflexors or triceps brachii fascicle the interaction effects of pre-unloading fascicle ge-
lengths or angles in subjects (n = 5) after 20 days of ometry and the muscle length during immobilisation
bed rest despite significant reductions in PCSA (ta- might be two factors affecting the magnitude of
ble III); a similar finding was reported in four sub- geometric change during detraining. Studies on cat,
jects in a later publication.[97] These results are simi- mouse, rat and rabbit muscles have shown signifi-
lar to those reported in rat studies where immobilisa- cant decreases in fibre length after periods of un-
tion of gastrocnemius and soleus muscles at loading when muscles were immobilised at lengths
shortened lengths was not associated with changes shorter than normal rest length.[94,104-108] These ef-
in fibre angles.[101] However, Narici and Cerretel- fects have been reduced or reversed when muscles

 2006 Adis Data Information BV. All rights reserved. Sports Med 2006; 36 (12)
1012 Blazevich

were immobilised in anatomical[109] or a lengthened 4. Fascicle Geometry through Growth


position.[93,106,108,109] Immobilisation in a lengthened and Aging
position has also been shown to counteract the slow-
to fast-twitch transformation that often accompanies Fascicle geometric changes have yet to be de-
scribed in most human muscles through develop-
immobilisation[110] and reduces the loss of whole
ment from infant to adult. Binzoni et al.[112] mea-
muscle mass.[111] These data on animal muscles
sured medial gastrocnemius fascicle angle in 134
show that the length of immobilisation has an im- subjects (87 men and 47 women) in the age range of
pact on muscle and fibre characteristics, although 0–70 years. Very small fascicle angles at birth in-
this has not been fully examined in humans. In creased through childhood and adolescence to reach
particular, given that the fibre angle of a shortened their maxima in early adulthood. A comparison of
pennate muscle is greater than in a lengthened mus- bodyweight and height, and tibia length indicated a
cle, one might speculate that immobilisation of mus- strong relationship between fascicle angle and phys-
cles in a shortened position might maintain the ge- ical size, possibly indicating that increases in fasci-
ometry of highly pennate, short-fibred muscles. Fi- cle angle of the gastrocnemius were related to the
loads imposed on it. No research has examined other
nally, given that immobilisation in lengthened
muscles, making it difficult to predict fascicle geo-
positions,[106] or the application of moderate stretch-
metric change with development, particularly in
ing (30 min/day),[107] has been shown to prevent non-weight-bearing muscles.
fibre shortening during disuse, it is also pertinent to More data are available with regard to fascicle
determine whether such interventions might prevent geometry with aging from early adult to old age.
the fibre shortening that has sometimes been shown Aging is associated with a loss of muscle size, or
to accompany disuse in humans.[62] Certainly the sarcopenia,[113-117] resulting from a loss of fibres/
addition of small amounts of stretching/mobilisa- motor units[118] and a significant reduction in fibre
tion[99] during detraining has been shown to arrest area, particularly of the fast-twitch fibres.[116,119-121]
muscle atrophy in the human soleus. These morphological changes in muscle have been
implicated, along with possible reductions in muscle
3.3.4 Chronic Stretching activation,[122,123] in the significant loss of muscle
force[114,115,119,124-126] and power[114,115,126,127] seen in
No research has examined changes in fascicle aged individuals. Changes in fascicle geometry are
geometry after acute or chronic stretching of mus- therefore likely given the positive relationship be-
cles in humans. Evidence from animal studies indi- tween fascicle angle and muscle size. Data collected
cates that chronic stretching might have a significant on 229 women from the ages of 20–79 years indicat-
effect on fibre length. Immobilisation of muscle or ed that pennation of vastus lateralis (r = –0.50, p <
muscle fibres in a lengthened position for periods of 0.001), but not medial gastrocnemius or triceps
days to weeks has resulted in increases in the num- brachii, might be reduced with age, although there
ber of serially arranged sarcomeres and overall fibre was no evidence for a change in fascicle length
length in animal muscles.[106,108,110] Interestingly, (relative to limb length).[128] The lesser pennation of
vastus lateralis in elderly compared with young, but
moderate stretching performed for only 30 min/day
lack of differences in gastrocnemius or triceps
was enough to reduce or reverse the fibre shortening
brachii, or in the length of fascicles in any of these
that accompanied short-length immobilisation in rat muscles, has also been reported for men.[129] In a
muscle.[107] It is not known whether flexibility train- smaller (n21–32y = 19, n60–69y = 30), but well
ing in humans results in an increase in fibre/fascicle matched (anthropometrically) sample, Karamanidis
length. It is also not clear what effect, if any, chronic and Arampatzis[61] found no differences in gas-
stretching would have on fascicle angle. trocnemius medialis or vastus lateralis fascicle

 2006 Adis Data Information BV. All rights reserved. Sports Med 2006; 36 (12)
Plasticity of Fascicle Geometry 1013

lengths or angles, regardless of whether the subjects geometry shown with aging are influenced by exer-
were habitual endurance runners or not. Nonethe- cise interventions.
less, other data have shown significant differences
in gastrocnemius pennation (young [27–42 years] > 5. Conclusions and Future Directions
13.2% [elderly = 70–81 years]) and fascicle length
(young >10.2%) in subjects matched for height, Human fascicle geometry is highly adaptable.
mass and physical activity.[117] These data suggest Heavy weight training in physically active individu-
that the aging process is accompanied by a signifi- als is often associated with increases in fascicle
cant change in fascicle geometry that cannot neces- angle (with some evidence for fascicle shortening)
sarily be explained by physical activity levels. Fur- with these changes amplified by anabolic steroid
thermore, Morse et al.[130] showed that decreases in use. In the elderly, the prominent adaptation seems
muscle volume of soleus and medial and lateral to be an increase in fascicle length, although it is not
gastrocnemius were associated with decreases in clear how geometry changes in response to longer-
both fascicle angle and length. These decreases were term training. Both detraining and aging are associ-
similar amongst the three muscles indicating that ated with decreases in fascicle angle and/or de-
these muscles, which fulfil different functional creases in fascicle length, although there appears to
roles, are affected similarly during aging. Thus, be some inter-muscular differences. These changes
while it seems that fascicle geometry changes from have important consequences for force generation
adult to old age, more research is required to sub- since increases in fascicle angle allow higher muscle
stantiate these early findings and further examine forces, while increases in fascicle length allow
inter-muscular differences. higher shortening speeds and for forces to be gener-
ated over larger length ranges.
The significant improvements in both muscle While there are significant data showing the plas-
strength[131-133] and power[127] after extended periods ticity of fascicle geometry, more research is required
of resistance exercise in older individuals are associ- to fully understand the effects of physical training.
ated with increases in cross-sectional area or volume Not least, no research has examined the effects of
of muscles.[127-130] This increase in cross-sectional endurance or chronic stretching training. Further-
area is often largely explained by increases in areas more, in order to more fully understand how fascicle
of both type I and II muscle fibres.[131,134] Few geometry responds to heavy loading (e.g. resistance
researchers have examined the effects of changing training), a number of research lines need to be
geometry on muscle size, or indeed the effects of explored. For example, despite a well established
geometry on force development. One study[133] has link between fascicle geometry and function, and
shown that an increase in muscle volume was attrib- significant evidence of geometric changes in re-
uted partly to an increase in fascicle length of the sponse to resistance training, relatively little re-
vastus lateralis. This change largely accounted for search has examined geometric changes in response
the increase in muscle PCSA, and suggests that to athletic training interventions. Only one research
significant changes in geometry might occur with study[6] has examined changes in geometry in re-
training in aged subjects. In this case, the change sponse to training in athletic subjects who perform
toward longer fascicle lengths is at odds with the mixed training regimens, and only one study[128] has
increases in fascicle angle (with no change or a examined changes in older individuals, so the ef-
slight reduction in fascicle length) seen after training fects of training programme manipulations in ath-
in younger subjects. This might reflect an adaptation letes and the aged remain relatively unstudied. Also,
in less mobile older subjects to a greater movement no research has specifically investigated the effects
range of motion used in exercise training compared of training mode, contraction type or range of mo-
with their normal daily activities. More research is tion. It is unclear whether, for example, adaptations
required to determine how the changes in fascicle to isokinetic training are different to isotonic/

 2006 Adis Data Information BV. All rights reserved. Sports Med 2006; 36 (12)
1014 Blazevich

isoinertial training. Despite changes in sarcomere 9. Burkholder TJ, Fingado B, Baron S, et al. Relationship between
muscle fiber types and sizes and muscle architectural proper-
number being reported after periods of eccentric ties in the mouse hindlimb. J Morphol 1994; 221: 177-90
training in rats (downhill running),[135,136] which 10. Van Eijden TMGJ, Korfage JAM, Brugman P, et al. Architec-
ture of the human jaw-closing and jaw-opening muscles. Anat
might suggest that overall fibre/fascicle length also Rec 1997; 248: 464-74
increases with such training, no research has ex- 11. Lieber RL, Fridén J. Functional and clinical significance of
amined differences between the effects of concen- skeletal muscle architecture. Muscle Nerve 2000; 23: 1647-66
12. Hof AL. Muscle mechanics and neuromuscular control. J Bi-
tric, eccentric or isometric training programmes on omech 2003; 36: 1031-8
geometric adaptation. Also, while range of motion- 13. Hof AL, Geelen BA, Van den Berg J. Calf muscle moment,
specific strength adaptations cannot be fully ex- work and efficiency in level walking: role of series elasticity. J
Biomech 1983, 37
plained by neural mechanisms,[87] no research has 14. Hof AL, Van Zandwijk JP, Bobbert MF. Mechanics of human
examined the effects of training range of motion. A triceps surae muscle in walking, running and jumping. Acta
Physiol Scand 2002; 174: 17-30
greater understanding of these parameters could
15. Kawakami Y, Muraoka T, Ito S, et al. In-vivo muscle-fibre
have significant benefits for injury/illness rehabilita- behaviour during counter-movement exercise in humans
tion in that maintenance of muscle strength during reveals significant role of tendon elasticity. J Physiol 2002;
540: 635-46
relative immobilisation might accelerate physical 16. Kubo K, Kanehisa H, Takeshita D, et al. In vivo dynamics of
recovery. No less, a more complete understanding human medial gastrocnemius muscle-tendon complex during
might also inform athletic training practices. 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-
Acknowledgements cal jumping. J Appl Physiol 2001; 90: 1349-58
18. Lieber RL, Jacobson MD, Fazelle BM, et al. Architecture of
Special thanks to Prof. Craig Sharp for his insightful selected muscles of the arm and forearm: anatomy and impli-
comments and discussion during the preparation of this arti- cations for tendon transfer. J Hand Surg 1992; 17A: 787-98
cle. This research was partly funded by an American Society 19. Elliott BC, Blanksby BA. The synchronization of muscle activi-
ty and body segment movements during a running cycle. Med
of Biomechanics research grant (ASB-406015). The author Sci Sports 1979; 11: 322-7
has no conflicts of interest that are directly related to the 20. Wiemann K, Tidow G. Relative activity of hip and knee exten-
content in this article. sors in sprinting: implications for training. N Studies Athl
1995; 10: 29-49
21. Wood GA. Optimal performance criteria and limiting factors in
References sprint running. N Studies Athl 1986; 2: 55-63
1. Scott SJ, Engstrom CM, Loeb GE. Morphometry of human 22. Friederich JA, Brand RA. Muscle fiber architecture in the
thigh muscles: determination of fascicle architecture by mag- human lower limb. J Biomech 1990; 23: 91-5
netic resonance imaging. J Anat 1993; 182: 249-57 23. Keller TCS. Molecular bungees. Nature 1997; 387: 233-5
2. Kawakami Y, Abe T, Fukunaga T. Muscle-fiber pennation 24. Politou AS, Thomas DJ, Pastore A. The folding and stability of
angles are greater in hypertrophied than in normal muscles. J titin immunoglobulin-like modules, with implications for the
Appl Physiol 1993; 74: 2740-4 mechanism of elasticity. Biophys J 1995; 96: 2601-10
3. Narici MV, Hoppeler H, Kayser B, et al. Human quadriceps 25. Tskhovrebova L, Trinick J, Sleep JA, et al. Elasticity and
cross-sectional area, torque and neural activation during 6 mo unfolding of single molecules of the giant muscle protein titin.
strength training. Acta Physiol Scand 1996; 157: 175-86 Nature 1997; 387: 308-12
4. Aagaard P, Andersen JL, Dyhre-Poulsen P, et al. A mechanism 26. Tskhovrebova L, Trinick J. Extensibility in the titin molecule
for increased contractile strength of human pennate muscle in and its relation to muscle elasticity. Adv Exp Med Biol 2000;
response to strength training: changes in muscle architecture. J 481: 163-78
Physiol 2001; 534: 613-23 27. Linari M, Dobbie I, Reconditi M, et al. The stiffness of skeletal
5. Blazevich AJ, Giorgi A. Effect of testosterone administration muscle in isometric contraction and rigor: the fraction of
and weight training on muscle architecture. Med Sci Sports myosin heads bound to actin. Biophys J 1998; 74: 2459-73
Exerc 2001; 33: 1688-93 28. Tawada K, Kimura M. Stiffness of glycerinated rabbit psoas
6. Blazevich AJ, Gill ND, Bronks R, et al. Training-specific mus- fibers in the rigor state: filament-overlap relation. Biophys J
cle architecture adaptation after 5-wk concurrent training in 1984; 45: 593-602
athletes. Med Sci Sports Exerc 2003; 35: 2013-22 29. Huxley HE, Stewart A, Sosa H, et al. X-ray diffraction measure-
7. Kawakami Y, Abe T, Kuno S-Y, et al. Training-induced ments of the extensibility of actin and myosin filaments in
changes in muscle architecture and specific tension. Eur J Appl contracting muscle. Biophys J 1994; 67: 2411-21
Physiol 1995; 72: 37-43 30. Martyn DA, Chase PB, Regnier M, et al. A simple model with
8. Rutherford OM, Jones DA. Measurement of fibre pennation myofilament compliance predicts activation-dependent cross-
using ultrasound in the human quadriceps in vivo. Eur J Appl bridge kinetics in skinned skeletal fibers. Biophys J 2002; 83:
Physiol 1992; 65: 433-7 3425-34

 2006 Adis Data Information BV. All rights reserved. Sports Med 2006; 36 (12)
Plasticity of Fascicle Geometry 1015

31. Wakabayashi K, Sugimoto Y, Tanaka H, et al. X-ray diffraction 52. Basser PJ, Mattiello J, LeBihan D. MR diffusion tensor spec-
evidence for the extensibility of actin and myosin filaments troscopy and imaging. Biophys J 1994; 66: 259-67
during contraction. Biophys J 1997; 67: 2422-35 53. van Donkelaar CC, Kretzers LJ, Bovendeerd PH, et al. Diffu-
32. Schroeter P. P, Bretaudiere JP, Sass RL, et al. Three-dimension- sion tensor imaging in biomechanical studies of skeletal mus-
al structure of the Z band in a normal mammalian skeletal cle function. J Anat 1999; 194: 79-88
muscle. J Cell Biol 1996; 133: 571-83 54. Mori S, Crain BJ, Chacko VP, et al. Three-dimensional tracking
33. Goldstein MA, Michael LH, Schroeter JP, et al. The Z-band of axonal projections in the brain by magnetic resonance
lattice in skeletal muscle before, during and after tetanic con- imaging. Ann Neurol 1999; 45: 265-9
traction. J Muscle Res Cell Motil 1986; 7: 527-36 55. Mori S, van Zijl PC. Fiber tracking: principles and strategies – a
34. Fukunaga T, Miyatani M, Tachi M, et al. Muscle volume is a technical review. NMR Biomed 2002; 15: 468-80
major determinant of joint torque in humans. Acta Physiol 56. Heemskerk AM, Strijkers GJ, Vilanova A, et al. Determination
Scand 2001; 172: 249-55 of mouse skeletal muscle architecture using three-dimensional
35. Muhl ZF. Active length-tension relation and the effect of muscle diffusion tensor imaging. Mag Res Med 2005; 53: 1333-40
pinnation of fiber lengthening. J Morphol 1982; 173: 285-92 57. van Doorn A, Bovendeerd PH, Nicolay K, et al. Determination
36. Herring SW, Grimm AF, Grimm BR. Regulation of sarcomere of muscle fibre orientation using diffusion-weighted MRI. Eur
number in skeletal muscle: a comparison of hypotheses. Mus- J Morphol 1996; 34: 5-10
cle Nerve 1984; 7: 161-73 58. Damon BM, Ding Z, Anderson AW, et al. Validation of diffu-
37. Lutz GJ, Rome LC. Muscle function during jumping in frogs I: sion tensor MRI-based muscle fiber tracking. Mag Res Med
sarcomere length change, EMG pattern, and jumping perform- 2002; 48: 97-104
ance. Am J Physiol 1996; 271: C563-70 59. Cleveland GG, Chang DC, Hazlewood CF, et al. Nuclear mag-
38. Lutz GJ, Rome LC. Muscle function during jumping in frogs II: netic resonance measurement of the intracellular water. Bi-
mechanical properties of muscle – implications for system ophys J 1976; 16: 1043-53
design. Am J Physiol 1996; 271: C571-8 60. Bensamoun SF, Ringleb SI, Littrell L, et al. Determination of
39. Häkkinen K, Komi PV. Effect of explosive type strength train- thigh muscle stiffness using magnetic resonance elastography.
ing on electromyographic and force production characteristics J Mag Res Imag 2006; 23: 242-7
of leg extensor muscles during concentric and various stretch- 61. Karamanidis K, Arampatzis A. Mechanical and morphological
shortening cycle exercises. Scand J Sports Sci 1985; 7: 65-76 properties of human quadriceps femoris and triceps surae
40. Häkkinen K, Komi PV. Electromyographic changes during muscle-tendon unit in relation to aging and running. J Biomech
strength training and detraining. Med Sci Sports Exerc 1983; 2006; 39: 406-17
15: 455-60 62. Narici M, Cerretelli P. Changes in human muscle architecture in
41. Hortobágyi T, Barrier J, Beard D, et al. Greater initial adapta- disuse-atrophy evaluated by ultrasound imaging. J Gravit
tions to submaximal muscle lengthening than maximal short- Physiol 1998; 5: 73-4
ening. J Appl Physiol 1996; 81: 1677-82 63. Chleboun GS, France AR, Crill MT, et al. In vivo measurement
42. Moritani T, DeVries HA. Neural factors versus hypertrophy in of fascicle length and pennation angle of the human biceps
the time course of muscle strength gain. Am J Phys Med 1979; femoris muscle. Cells Tiss Org 2001; 169: 401-9
58: 115-30 64. Narici MV, Binzoni T, Hiltbrand E, et al. Changes in human
43. Patten C, Kamen G, Rowland D, et al. Rapid adaptations of gastrocnemius architecture with joint angle, at rest and with
motor unit firing rate during the initial phase of strength isometric contraction, evaluated in vivo. J Physiol 1996; 496:
development. Med Sci Sports Exerc 1995; 27: S6 287-97
44. Ploutz LL, Tesch PA, Biro RL, et al. Effect of resistance training 65. Kurihara T, Oda T, Chino K, et al. Use of three-dimensional
on muscle use during exercise. J Appl Physiol 1994; 76: ultrasonography for the analysis of the fascicle length of
1675-81 human gastrocnemius muscle during contractions. Int J Sport
45. Enoka RM. Neural adaptations with chronic physical activity. J Health Sci 2005; 3: 226-34
Biomech 1997; 30: 447-55 66. Muramatsu T, Muraoka T, Kawakami Y, et al. In vivo determi-
46. Chilibeck PD, Calder AW, Sale DG, et al. A comparison of nation of fascicle curvature in contracting human skeletal
strength and muscle mass increases during resistance training muscles. J Appl Physiol 2002; 92: 129-34
in young women. Eur J Appl Physiol 1998; 77: 170-5 67. Reeves ND, Narici MV. Behavior of human muscle fascicles
47. Hickson RC, Hidaka K, Foster C, et al. Successive time courses during shortening and lengthening contractions in vivo. J Appl
of strength development and steroid hormone responses to Physiol 2003; 95: 1090-6
heavy-resistance training. J Appl Physiol 1994; 76: 663-70 68. Finni T, Ikegawa S, Komi PV. Concentric force enhancement
48. Häkkinen K, Komi PV, Alén M, et al. EMG, muscle fibre and during human movement. Acta Physiol Scand 2001; 173:
force production characteristics during a 1 year training period 369-77
in elite weight-lifters. Eur J Appl Physiol 1987; 56: 419-27 69. Finni T, Ikegawa S, Lepola V, et al. Comparison of force-
49. Gaudy JF, Zouaoui A, Bravetti P, et al. Functional anatomy of velocity relationships of vastus lateralis muscle in isokinetic
the human temporal muscle. Surg Radiol Anat 2001; 23: and in stretch-shortening cycle exercises. Acta Physiol Scand
389-98 2003; 177: 483-91
50. Gaudy JF, Zouaoui A, Bravetti P, et al. Functional organization 70. Izquerdo M, Ibañez J, Häkkinen K, et al. Maximal strength and
of the human masseter muscle. Surg Radiol Anat 2000; 22: power, muscle mass, endurance and serum hormones in
181-90 weightlifters and road cyclists. J Sport Sci 2004; 22: 465-78
51. Lam EW, Hannam AG, Christiansen EL. Estimation of tendon- 71. Izquierdo M, Häkkinen K, Gonzalez-Badillo JJ, et al. Effects of
plane orientation within human masseter muscle from recon- long-term training specificity on maximal strength and power
structed magnetic resonance images. Arch Oral Biol 1991; 36: of the upper and lower extremities in athletes from different
845-53 sports. Eur J Appl Physiol 2002; 87: 264-71

 2006 Adis Data Information BV. All rights reserved. Sports Med 2006; 36 (12)
1016 Blazevich

72. Coyle EF. Improved muscular efficiency displayed as Tour de 92. Aratow M, Ballard RE, Crenshaw AG, et al. Intramuscular
France champion matures. J Appl Physiol 2005; 98: 2191-6 pressure and electromyography as indexes of force during
73. Alexander MJL. The relationship between muscle strength and isokinetic exercise. J Appl Physiol 1993; 74: 2634-40
sprint kinematics in elite sprinters. Can J Sport Sci 1989; 14: 93. Degens H, Salmons S, Jarvis JC. Intramuscular pressure, force
148-57 and blood flow in rabbit tibialis anterior muscles during single
74. Taylor NAS, Cotter JD, Stanley SN, et al. Functional torque- and repetitive contractions. Eur J Appl Physiol Occup Physiol
velocity and power-velocity characteristics of elite athletes. 1998; 78: 13-9
Eur J Appl Physiol 1991; 62: 116-21 94. Jarvinen MJ, Einola SA, Virtanen EO. Effect of the position of
75. Blazevich AJ, Jenkins DG. Physical performance differences immobilization upon the tensile properties of the rat gas-
between weight-trained sprinters and weight trainers. J Sci trocnemius muscle. Arch Phys Med Rehab 1992; 73: 253-7
Med Sport 1998; 1: 12-21 95. Sejersted OM, Hagens AR, Kardel KR, et al. Intramuscular fluid
76. Sleivert GG, Backus RD, Wenger HA. Neuromuscular differ- pressure during isometric contraction of human skeletal mus-
ences between volleyball players, middle distance runners and cle. J Appl Physiol 1984; 56: 287-95
untrained controls. Int J Sport Med 1995; 16: 390-8 96. Akima H, Kuno S, Suzuki Y, et al. Effects of 20 days of bed rest
77. Kumagai K, Abe T, Brechue WF, et al. Sprint performance is on physiological cross-sectional area of human thigh and leg
related to muscle fascicle length in male 100-m sprinters. J muscles evaluated by magnetic resonance imaging. J Gravit
Appl Physiol 2000; 88: 811-6 Physiol 1997; 4: S15-21
78. Abe T, Fukashiro S, Harada Y, et al. Relationship between 97. Kawakami Y, Akima H, Kubo K, et al. Changes in muscle size,
sprint performance and muscle fascicle length in female sprint- architecture, and neural activation after 20 days of bed rest
ers. J Physiol Anthropol Appl Human Sci 2001; 20: 141-7 with and without resistance exercise. Eur J Appl Physiol 2001;
79. Abe T, Kumagai K, Brechue WF. Fascicle length of leg muscles 84: 7-12
is greater in sprinters than distance runners. Med Sci Sports 98. Kawakami Y, Muraoka Y, Kubo K, et al. Changes in muscle
Exerc 2000; 32: 1125-9 size and architecture following 20 days of bed rest. J Gravit
80. Brechue WF, Abe T. The role of FFM accumulation and skeletal Physiol 2000; 7: 53-60
muscle architecture in powerlifting performance. Eur J Appl 99. Yamashita-Goto K, Okuyama R, Honda M, et al. Maximal and
Physiol 2002; 86: 327-36 submaximal forces of slow fibers in human soleus after bed
81. Kearns CF, Abe T, Brechue WF. Muscle enlargement in sumo rest. J Appl Physiol 2001; 91: 417-24
wrestlers includes increased muscle fascicle length. Eur J Appl 100. Andersen JL, Gruschy-Knudsen T, Sandri C, et al. Bed rest
Physiol 2000; 83: 289-96 increases the amount of mismatched fibers in human skeletal
82. Gondin J, Guette M, Ballay Y, et al. Electromyostimulation muscle. J Appl Physiol 1999; 86: 455-60
training effects on neural drive and muscle architecture. Med
101. Heslinga JW, Huijing PA. Effects of short length immobiliza-
Sci Sports Exerc 2005; 37: 1291-9
tion of medial gastrocnemius muscle of growing young adult
83. Alegre LM, Jimenez F, Gonzalo-Orden JM, et al. Effects of rats. Eur J Morphol 1992; 30: 257-73
dynamic resistance training on fascicle length and isometric
102. Bleakney R, Maffulli N. Ultrasound changes to intramuscular
strength. J Sports Sci 2006; 24: 501-8
architecture of the quadriceps following intramedullary nail-
84. Blazevich AJ, Gill ND, Deans N, et al. Lack of human muscle ing. J Sports Med Phys Fit 2002; 42: 120-5
architectural adaptation after short-term strength training.
Muscle Nerve. Epub 2006 Oct 12 103. Abe T, Kawakami Y, Suzuki Y, et al. Effects of 20 days bed rest
on muscle morphology. J Gravit Physiol 1997; 4: S10-4
85. Häkkinen K, Kallinen M, Linnamo V, et al. Neuromuscular
adaptations during bilateral versus unilateral strength training 104. Baker JH, Matsumoto DE. Adaptation of skeletal muscle to
in middle-aged and elderly men and women. Acta Physiol immobilization in a shortened position. Muscle Nerve 1988;
Scand 1996; 158: 77-88 11: 231-44
86. Tanaguchi Y. Laterality specificity in resistance training: the 105. Hayat A, Tardieu C, Tabary JC, et al. Effects of denervation on
effect of bilateral and unilateral training. Eur J Appl Physiol the reduction of sarcomere number in cat soleus muscle immo-
1997; 75: 144-50 bilized in shortened position during seven days. J Physiol
(Paris) 1978; 74: 563-7
87. Weir JP, Housh DJ, Housh TJ, et al. The effect of unilateral
concentric weight training and detraining on joint angle speci- 106. Tabary JC, Tabary C, Tardieu C, et al. Physiological and struc-
ficity, cross-training, and the bilateral deficit. J Orthop Sports tural changes in the cat’s soleus muscle due to immobilization
Phys Ther 1997; 25: 264-70 at different lengths by plaster casts. J Physiol 1972; 224:
88. Jarvholm U, Palmerud G, Karlsson D, et al. Intramuscular 231-44
pressure and electromyography in four shoulder muscles. J 107. Williams PE. Use of intermittent stretch in the prevention of
Orthop Res 1991; 9: 609-19 serial sarcomere loss in immobilised muscle. Ann Rheum Dis
89. Sejersted OM, Hagens AR. Intramuscular pressures for moni- 1990; 49: 316-7
toring different tasks and muscle conditions. Adv Expl Med 108. Williams PE, Goldspink G. Changes in sarcomere length and
Biol 1995; 384: 339-50 physiological properties in immobilized muscle. J Anat 1978;
90. Miura H, McCully K, Nioka S, et al. Relationship between 127: 459-68
muscle architectural features and oxygenation status deter- 109. Nicks DK, Beneke WM, Key RM, et al. Muscle fibre size and
mined by near infrared device. Eur J Appl Physiol 2004; 91: number following immobilisation atrophy. J Anat 1989; 163:
273-8 1-5
91. Ballard RE, Watenpaugh DE, Breit GA, et al. Leg intramuscular 110. Leterme D, Cordonnier C, Mounier Y, et al. Influence of chron-
pressures during locomotion in humans. J Appl Physiol 1998; ic stretching upon rat soleus muscle during non-weight-bear-
84: 1976-81 ing conditions. Eur J Physiol 1994; 429: 274-9

 2006 Adis Data Information BV. All rights reserved. Sports Med 2006; 36 (12)
Plasticity of Fascicle Geometry 1017

111. Herbert RD, Balnave RJ. The effect of position of immobilisa- 125. Roos MR, Rice CL, Connelly DM, et al. Quadriceps muscle
tion on resting length, resting stiffness, and weight of the strength, contractile properties, and motor unit firing rates in
soleus muscle of the rabbit. J Orthop Res 1993; 11: 358-6
young and old men. Muscle Nerve 1999; 22: 1094-103
112. Binzoni T, Bianchi S, Hanquinet S, et al. Human gastrocnemius 126. Valour D, Ochala J, Ballay Y, et al. The influence of ageing on
medialis pennation angle as a function of age: from newborn to
the force-velocity-power characteristics of human elbow flex-
the elderly. J Physiol Anthropol Appl Hum Sci 2001; 20: 293-8
or muscles. Exp Gerontol 2003; 38: 387-95
113. Häkkinen K, Pakarinen A. Muscle strength and serum hormones
127. Newton RU, Häkkinen K, Häkkinen A, et al. Mixed-methods
in middle-aged and elderly men and women. Acta Physiol
Scand 1993; 148: 199-207 resistance training increases power and strength of young and
older men. Med Sci Sports Exerc 1999; 34: 1367-75
114. Izquierdo M, Ibañez J, Gorostiaga E, et al. Maximal strength
and power characteristics in isometric and dynamic actions of 128. Kubo K, Kanehisa H, Azuma K, et al. Muscle architectural
the upper and lower extremities in middle-aged and older men. characteristics in women aged 20-79 years. Med Sci Sports
Acta Physiol Scand 1999; 167: 57-68 Exerc 2003; 35: 39-44
115. Izquerdo M, Häkkinen K, Antón A, et al. Maximal strength and 129. Kubo K, Kanehisa H, Azuma K, et al. Muscle architectural
power, endurance performance, and serum hormones in mid- characteristics in young and elderly men and women. Int J
dle-aged and elderly men. Med Sci Sports Exerc 2001; 33: Sports Med 2003; 24: 125-30
1577-87
130. Morse CI, Thom JM, Birch KM, et al. Changes in triceps surae
116. Klitgaard H, Mantoni M, Schiaffino S, et al. Function, morphol-
muscle architecture with sarcopenia. Acta Physiol Scand 2005;
ogy and protein expression of ageing skeletal muscle: a cross-
sectional study of elderly men with different training back- 183: 291-8
grounds. Acta Physiol Scand 1990; 140: 41-54 131. Häkkinen K, Kraemer WJ, Pakarinen A, et al. Effects of heavy
117. Narici MV, Maganaris CN, Reeves ND, et al. Effect of aging on resistance/power training on maximal strength, muscle mor-
human muscle architecture. J Appl Physiol 2003; 95: 2229-34 phology, and hormonal response patterns in 60-75-year-old
118. Lexell J, Taylor CC, Sjostrom M. What is the cause of the aging men and women. Can J Appl Physiol 2002; 27: 213-31
atrophy? Total number, size and proportion of different fiber 132. Izquerdo M, Häkkinen K, Ibañez J, et al. Effects of strength
types studies in whole vastus lateralis muscle from 15- to training on submaximal and maximal endurance performance
83-year-old men. J Neurol Sci 1988; 84: 275-94 capacity in middle-aged and older men. J Strength Cond Res
119. Hortobágyi T, Zheng D, Weidner M, et al. The influence of 2003; 17: 129-39
aging on muscle strength and muscle fiber characteristics with
133. Reeves ND, Narici MV, Maganaris CN. Effect of resistance
special reference to eccentric strength. J Gerontol A: Biol Sci
Med Sci 1995; 50: B399-406 training on skeletal muscle-specific force in elderly humans. J
Appl Physiol 2004; 96: 885-92
120. Larsson L, Grimby G, Karlsson J. Muscle strength and speed of
movement in relation to age and muscle morphology. J Appl 134. Trappe S, Williamson D, Godard M, et al. Effect of resistance
Physiol 1979; 46: 451-6 training on single muscle fiber contractile function in older
men. J Appl Physiol 2000; 89: 143-52
121. Poggi P, Marchettic C, Scelsi R. Automatic morphometric anal-
ysis of skeletal muscle fibers in the aging man. Anat Rec 1987; 135. Lynn R, Morgan DL. Decline running produced more sarcomer-
217: 30-4 es in rat vastus intermedius muscle fibers than does incline
122. Häkkinen K, Alen M, Kallinen M, et al. Neuromuscular adapta- running. J Appl Physiol 1994; 77: 1439-44
tion during prolonged strength training, detraining and re- 136. Butterfield TA, Leonard TR, Herzog W. Differential serial
strength-training in middle-aged and elderly people. Eur J sarcomere number adaptations in knee extensor muscles of rats
Appl Physiol 2000; 83: 51-62
is contraction type dependent. J Appl Physiol 2005; 99: 1352-8
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; Correspondence and offprints: Dr Anthony J. Blazevich,
84: 1341-9
Centre for Sports Medicine and Human Performance,
124. Knight CA, Kamen G. Adaptations in muscular activation of the
knee extensor muscles with strength training in young and Brunel University, Uxbridge, UB8 3PH, UK.
older adults. J Electromyogr Kinesiol 2001; 11: 405-12 E-mail: anthony.blazevich@brunel.ac.uk

 2006 Adis Data Information BV. All rights reserved. Sports Med 2006; 36 (12)