Beruflich Dokumente
Kultur Dokumente
Muscle length
l
f
= a b
PCSA
Fig. 1. Muscle architectural parameters include: fibre length = dis-
tance between ends of a fibre (a to b); pennation angle () =
fascicle angle (relative to the aponeurosis []) minus the aponeuro-
sis angle (relative to the tendon []); muscle length; and anatomical
(ACSA) or physiological (PCSA) cross-sectional area. The PCSA
can be calculated as (V/t) sin for a simple, uni-pennate muscle,
where V is the muscle volume, t is the muscle thickness from one
aponeurosis to the other, and is the pennation angle. In more
complex muscles, PCSA is calculated as V/ lf cos 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. Sports Med 2006; 36 (12)
Plasticity of Fascicle Geometry 1005
2. Effects of Geometry on
Force Production
2.1 Fibre Length
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
100
m
Tendon/
aponeurosis
Fascicle
force
= 5
T
e
n
d
o
n
f
o
r
c
e
(
%
m
u
s
c
l
e
f
o
r
c
e
)
80
= 5
= 25
= 45
60
40
20
0
5 15
Fascicle angle ()
25 35 45 55
m
= 25
m
= 45
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 (13.514.5 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
(10.010.9 seconds for 100m), their fascicles were
While the assessment of population differences
also shorter and their fascicle angles greater. Com-
provides an insight into possible activity-dependent
pared with non-active subjects, young endurance
adaptations in fascicle geometry, a more satisfactory
runners had larger fascicle angles (13.7%) but a
method of examining the plasticity of fascicle geom-
similar fascicle length in gastrocnemius medialis;
etry is to directly measure the longitudinal adapta-
there were no differences in fascicle geometry in the
tions.
vastus lateralis.
[61]
At least for the gastrocnemius,
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 humans
a
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-
tion 5). Thus, recent research has shown a signifi- between fascicle angle and force-generating capaci-
cant adaptability of fascicle geometry in response to ty, although the authors reported a moderately high
longer-term resistance training. coefficient of variation (13.5%), making small
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
training (n = 8; no change). In the same study, vastus is presently unclear whether the training velocity or
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
extensors of an untrained limb in five men after 16 jump-trained group. Muscle thickness of both vastus
weeks of training, although no strength changes lateralis and rectus femoris increased in all training
were found either. Blazevich et al.
[84]
have reported groups also, although there were no differences in
that significant increases in contralateral knee exten- performance (sprint, jump and strength tests) be-
sor strength after 5 weeks of isokinetic training
tween the groups after training. Similar to previous
occurred without a change in geometry. Thus, al-
studies on untrained subjects,
[7,83]
these results show
though few data are available, there is no current
that changes in fascicle geometry occur in response
evidence to suggest that fascicle geometric changes
to resistance training in well trained subjects. The
are a factor affecting short-term strength increases in
results may have differed from previous studies
a contralateral limb.
where little change was noticed in trained subjects
because the training programme differed sufficient-
3.3.2 Endurance Training
ly from that which the athletes had previously per-
No research has examined changes in geometry
formed. These data were the first to show short-term
after long-term endurance training. It is therefore
changes in geometry in response to training, and
not known whether endurance training can elicit
also show that increases in fascicle length and de-
geometric adaptations, or how different modes of
creases in fascicle angle can occur after high-veloc-
training (e.g. cycling, running, lifting) might affect
ity movement training in humans; it is not known,
them. It makes intuitive sense that if a muscle could
however, whether other factors such as the training
generate force more efficiently, then muscle endur-
range of motion influence the architectural adapta-
ance would improve. Given that muscles with large
tion.
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 humans
a
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
The reasons for the disparate results in de-
measured for the determination of PCSA. In one
training/unloading studies are not clear; however,
study, Kawakami et al.
[98]
found no changes in vas-
the interaction effects of pre-unloading fascicle ge-
tus lateralis, plantarflexors or triceps brachii fascicle
ometry and the muscle length during immobilisation
lengths or angles in subjects (n = 5) after 20 days of
might be two factors affecting the magnitude of
bed rest despite significant reductions in PCSA (ta-
geometric change during detraining. Studies on cat,
ble III); a similar finding was reported in four sub-
mouse, rat and rabbit muscles have shown signifi- jects in a later publication.
[97]
These results are simi-
cant decreases in fibre length after periods of un- lar to those reported in rat studies where immobilisa-
loading when muscles were immobilised at lengths tion of gastrocnemius and soleus muscles at
shorter than normal rest length.
[94,104-108]
These ef- shortened lengths was not associated with changes
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
and Aging
position.
[93,106,108,109]
Immobilisation in a lengthened
position has also been shown to counteract the slow-
Fascicle geometric changes have yet to be de-
to fast-twitch transformation that often accompanies
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
070 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
bodyweight and height, and tibia length indicated a pennate muscle is greater than in a lengthened mus-
strong relationship between fascicle angle and phys-
cle, one might speculate that immobilisation of mus-
ical size, possibly indicating that increases in fasci-
cles in a shortened position might maintain the ge-
cle angle of the gastrocnemius were related to the
ometry of highly pennate, short-fibred muscles. Fi-
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/
motor units
[118]
and a significant reduction in fibre
addition of small amounts of stretching/mobilisa-
area, particularly of the fast-twitch fibres.
[116,119-121]
tion
[99]
during detraining has been shown to arrest
These morphological changes in muscle have been
muscle atrophy in the human soleus.
implicated, along with possible reductions in muscle
activation,
[122,123]
in the significant loss of muscle
3.3.4 Chronic Stretching
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 2079 years indicat-
effect on fibre length. Immobilisation of muscle or ed that pennation of vastus lateralis (r =
0.50, p <
0.001), but not medial gastrocnemius or triceps
muscle fibres in a lengthened position for periods of
brachii, might be reduced with age, although there
days to weeks has resulted in increases in the num-
was no evidence for a change in fascicle length
ber of serially arranged sarcomeres and overall fibre
(relative to limb length).
[128]
The lesser pennation of
length in animal muscles.
[106,108,110]
Interestingly,
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 (n
2132y
= 19, n
6069y
= 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-
cise interventions. were habitual endurance runners or not. Nonethe-
less, other data have shown significant differences
in gastrocnemius pennation (young [2742 years] > 5. Conclusions and Future Directions
13.2% [elderly = 7081 years]) and fascicle length
Human fascicle geometry is highly adaptable.
(young >10.2%) in subjects matched for height,
Heavy weight training in physically active individu-
mass and physical activity.
[117]
These data suggest
als is often associated with increases in fascicle
that the aging process is accompanied by a signifi-
angle (with some evidence for fascicle shortening)
cant change in fascicle geometry that cannot neces-
with these changes amplified by anabolic steroid
sarily be explained by physical activity levels. Fur-
use. In the elderly, the prominent adaptation seems
thermore, Morse et al.
[130]
showed that decreases in
to be an increase in fascicle length, although it is not
muscle volume of soleus and medial and lateral
clear how geometry changes in response to longer-
gastrocnemius were associated with decreases in
term training. Both detraining and aging are associ-
both fascicle angle and length. These decreases were
ated with decreases in fascicle angle and/or de-
similar amongst the three muscles indicating that
creases in fascicle length, although there appears to
these muscles, which fulfil different functional
be some inter-muscular differences. These changes
roles, are affected similarly during aging. Thus,
have important consequences for force generation
while it seems that fascicle geometry changes from
since increases in fascicle angle allow higher muscle
adult to old age, more research is required to sub-
forces, while increases in fascicle length allow
stantiate these early findings and further examine
higher shortening speeds and for forces to be gener-
inter-muscular differences.
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
9. Burkholder TJ, Fingado B, Baron S, et al. Relationship between
isoinertial training. Despite changes in sarcomere
muscle fiber types and sizes and muscle architectural proper-
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10. Van Eijden TMGJ, Korfage JAM, Brugman P, et al. Architec-
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[135,136]
which
ture of the human jaw-closing and jaw-opening muscles. Anat
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skeletal muscle architecture. Muscle Nerve 2000; 23: 1647-66
amined differences between the effects of concen-
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omech 2003; 36: 1031-8
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of Biomechanics research grant (ASB-406015). The author
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has no conflicts of interest that are directly related to the
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content in this article.
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