Blazevich Et Al 2006 - Diferentes Locais de Medidas de Ultrassonografia

J. Anat. (2006) 209, pp289–310 doi: 10.1111/j.1469-7580.2006.00619.
Intra- and intermuscular variation in human quadriceps

Blackwell Publishing Ltd
femoris architecture assessed in vivo

Anthony J. Blazevich,1 Nicholas D. Gill2 and Shi Zhou3
1
Centre for Sports Medicine and Human Performance, School of Sport and Education, Brunel University, Uxbridge, UK
2
School of Sport and Exercise Science, Waikato Institute of Technology, Hamilton, New Zealand
3
School of Exercise Science and Sport Management, Southern Cross University, Lismore, Australia
Abstract
Despite the functional importance of the human quadriceps femoris in movements such as running, jumping, lifting
and climbing, and the known effects of muscle architecture on muscle function, no research has fully described the
complex architecture of this muscle group. We used ultrasound imaging techniques to measure muscle thickness,
fascicle angle and fascicle length at multiple regions of the four quadriceps muscles in vivo in 31 recreationally
active, but non-strength-trained adult men and women. Our analyses revealed a reasonable similarity in the super-
ficial quadriceps muscles, which is suggestive of functional similarity (at least during the uni-joint knee extension
task) given that they act via a common tendon. The deep vastus intermedius (VI) is architecturally dissimilar and
therefore probably serves a different function(s). Architecture varies significantly along the length of the superficial
muscles, which has implications for the accuracy of models that assume a constant intramuscular architecture. It might
also have consequences for the efficiency of intra- and intermuscular force transmission. Our results provide some
evidence that subjects with a given architecture of one superficial muscle, relative to the rest of the subject sample,
also have a similar architecture in other superficial muscles. However, this is not necessarily true for vastus lateralis
(VL), and was not the case for VI. Therefore, the relative architecture of one muscle cannot confidently be used to
estimate the relative architecture of another. To confirm this, we calculated a value of whole quadriceps architecture
by four different methods. Regardless of the method used, we found that the absolute or relative architecture of
one muscle could not be used as an indicator of whole quadriceps architecture, although vastus medialis, possibly
in concert with VL and the anterior portion of VI, could be used to provide a useful snapshot. Importantly, our estimates
of whole quadriceps architecture show a gender difference in whole quadriceps muscle thickness, and that muscle
thickness is positively correlated with fascicle angle whereas fascicle length is negatively, although weakly, correlated
with fascicle angle. These results are supportive of the validity of estimates of whole quadriceps architecture. These
data are interpreted with respect to their implications for neural control strategies, region-specific adaptations in
muscle size in response to training, and gender-dependent differences in the response to exercise training.
Key words fibre angle; fibre length; pennation; rectus; skeletal muscle; thigh; vastus.
Most notably, larger muscles typically have a large

Introduction
quantity of force-producing contractile material and
Gross muscle architecture, encompassing a muscle’s size are therefore able to develop high forces. However,
and the lengths and angles of its fibres, is a strong deter- the expression of this force is modulated by both fibre
minant of force production and movement performance. angle and fibre length (Woittiez et al. 1984; Lieber &
Fridén, 2000). Muscles with fibres attaching at large
Correspondence angles to the tendon or aponeurosis have a large physi-
Dr Anthony J. Blazevich, Centre for Sports Medicine and Human
ological cross-section for their volume, and therefore
Performance, School of Sport and Education, Brunel University,
Uxbridge UB8 3PH, UK. T: +44 1895266 465; F: +44 1895269 769; produce high relative forces (Powell et al. 1984). Because
E: anthony.blazevich@brunel.ac.uk fibres of pennate muscles rotate as they shorten (Muhl,
Accepted for publication 6 June 2006 1982), and consequently their fibre shortening distance
© 2006 The Authors

Journal compilation © 2006 Anatomical Society of Great Britain and Ireland
290 Human quadriceps femoris architecture, A. J. Blazevich et al.
is less than that of the tendon, fibres in pennate muscles might also require different patterns of force genera-
are also able to work at lower shortening speeds tion, and thus different force-generating capacities in
and have a greater opportunity to work near their opti- these muscles.
mum length. Muscles with long fibres, however, are This creates a problem with respect to optimum
capable of contracting with high shortening speeds muscle group design. Like the wrist muscle complex, the
and produce forces over longer length ranges, as both human quadriceps femoris comprises several muscles
of these characteristics are proportional to the number (including the mono-articular vastii and bi-articular
of serially arranged sarcomeres in the fibres. Also, rectus femoris). Variability in architecture between these
because sarcomeres of longer fibres shorten less for a muscles would ensure that a range of force requirements
given absolute fibre shortening, their ability to produce could be met. However, a comprehensive examination
force at high muscle shortening speeds is greater than of quadriceps femoris architecture has not been per-
that of short fibres. Given the significant effects of formed, so a detailed picture does not exist. Examina-
muscle architecture on force-generating ability, it makes tion of the intermuscular variability in architecture
sense for muscle architecture to vary across muscles and would allow a greater understanding of how synergists
muscle groups according to the muscles’ functional cooperate during force development. This is important
roles (Lieber et al. 1992; Zajac, 1992). for the construction of appropriate models of human
Although it would be ideal for a muscle, or muscle movement, and for the design of mechanized locomotor
group, to have an architecture that is suited specifically systems including prosthetic and robotic limbs. Despite
to the force requirements of the joint (or joints) it the importance of the human quadriceps in lifting and
serves, some joints perform numerous functions. At the locomotion, no research to our knowledge has compre-
wrist, for example, there is a requirement for high hensively described both its intra- and intermuscular
force (torque) generation when it is used during heavy architecture. It is also not known whether quadriceps
lifting, levering and pushing/pulling tasks, but it also architecture is uniform, or highly variable, between
functions through a large flexion–extension range of muscles or individuals. Because muscle architecture
motion, or at higher velocities in other tasks. This creates changes in response to physical loading (Kawakami
a difficulty in that an ideal wrist extensor would con- et al. 1995; Blazevich et al. 2003), differences in activity
tain fibres that are long, but attach at relatively large patterns between individuals should increase architec-
angles. The evolutionary solution to this problem is to tural heterogeneity.
have several muscles of varying architecture working in By way of comparison with a ‘theoretically optimum’
synergy so that muscles with a smaller volume can meet muscle group, the present research investigated
the force requirements (e.g. Jacobson et al. 1992; Lieber several hypotheses: (1) the architecture of each muscle
et al. 1992, 1997). within quadriceps femoris should be different, as mem-
The human quadriceps function primarily as knee bers of a synergistic group should have different force-
extensors, providing large forces during pushing/pull- production capabilities; (2) each muscle’s architecture
ing movements, and assisting in leg extension during should be relatively homogeneous along its length so
running and jumping. In comparison with other primates, that each muscle is optimally designed to fulfil its role;
humans walk (and run) with the knee joint relatively (3) the relative (i.e. ranked against others within the
extended (Winter et al. 1974; D’Août et al. 2002; Schmitt, subject sample) architecture of one muscle should be
2003), although during squatting and lifting the knee similar to the relative architecture of another muscle,
joint can become flexed acutely. Therefore, the qua- as humans have different genetic and physical activity
driceps must produce forces over large length ranges. profiles and each subject’s architecture should reflect
Moreover, during jumping, the knee extensor muscle that; and (4) given hypothesis 3, the relative (i.e. ranked)
stress reaches approximately 280 kN m−2 (Thorpe et al. architecture of individual muscles should be indicative
1998; bilateral countermovement jump) as the knee of whole quadriceps architecture, i.e. the measurement
extensor moment reaches approximately 280 Nm of one or more identified representative regions on the
(Bobbert et al. 1986). Therefore, human knee extensors muscles should allow estimation of a subject’s whole
must also be capable of producing relatively large quadriceps architecture. In order partially to validate
muscle forces, often at high shortening speeds. Addition- the estimation of whole quadriceps architecture, we
ally, the variability in human physical activity patterns compared men and women, and examined relationships
© 2006 The Authors

Human quadriceps femoris architecture, A. J. Blazevich et al. 291
between muscle thickness, fascicle angle and fascicle cadavers, and are similar to those measured using 3-D
length. Based on the published data, we expected men ultrasound unless the transducer is not orientated
to have a greater quadriceps muscle thickness than in the plane of the fascicles (error range 2.4–14.0%;
women (Abe et al. 1998; Chow et al. 2000; Kubo et al. Kurihara et al. 2005). Muscle thickness has also been
2003), for there to be a positive correlation between validated against magnetic resonance imaging scans in
muscle thickness and fascicle angle (Abe et al. 1998; various human muscles (< 2 mm; Juul-Kristensen et al.
Brechue & Abe, 2002; Kanehisa et al. 2003), and for 2000; Dupont et al. 2001).
there to be a weaker negative correlation between To obtain the sonographs, subjects lay supine with
fascicle angle and fascicle length (Calow & Alexander, their knees flexed to 45° with their legs supported and
1983; Kumagai et al. 2000; Kanehisa et al. 2003). We muscles relaxed. The knee bend was used to reduce
compared four methods of estimating whole quadriceps fascicle curvature and improve measurement reliability,
architecture to determine whether a ‘best method’ while still allowing significant interindividual variability
could be derived, with the methods differing in the to be seen. A water-soluble gel was applied to the
proportional contribution each muscle made to the probe to aid acoustic contact and remove the need to
whole quadriceps estimate. contact the skin, thus eliminating the deformation of
muscle that might occur if pressure was applied. Sono-
graphic scans were performed on the right leg with the
Materials and methods probe orientated parallel to the muscle fascicles and
perpendicular to the skin. The angle of the probe
Subjects
relative to the longitudinal axis of the thigh therefore
Sixteen women [age (mean ± SD) = 19.9 ± 3.1 years, varied between subjects, although approximate orien-
height = 170.0 ± 0.04 m, mass = 64.6 ± 7.9 kg] and 15 tations are shown in Fig. 1. Appropriate probe align-
men (age = 20.6 ± 2.6 years, height = 1.80 ± 0.09 m, mass ment was achieved when several fascicles could be
= 76.0 ± 13.0 kg) volunteered for the study. None of
the subjects had neuromuscular or inflammatory
diseases, nor had performed weight training in the
3 years prior to the study or performed intense physical
exercise more than three times a week. The subjects
refrained from intense physical exertion for at least
48 h prior to examination.
Muscle thickness and fascicle angle
In vivo muscle architecture was examined using two-

dimensional (2-D), B-mode ultrasonography (Acuson
Sequoia, Acuson Corporation, CA, USA) with a 3.8-cm
linear array probe (7.5 MHz). 2-D ultrasound is com-
monly used for this purpose (e.g. Kawakami et al. 1995;
Aagaard et al. 2001; Blazevich et al. 2003; Kanehisa Fig. 1 Quadriceps femoris scanning sites. Sonographs were
obtained at lengths equivalent to 5 –73% of thigh length
et al. 2003; Kubo et al. 2003) and involves direct measure-
measured from the superior border of the patella to the
ment of muscle thickness, fascicle angles and fascicle anterior superior iliac spine. Muscles were scanned at three
lengths from sonographs. The echoes reflected from sites as indicated in the diagram above. The anterior portion
the superficial and deep aponeuroses and the inter- of vastus intermedius (VIant, not shown) was scanned at the
same sites as RF, while the lateral portion (VIlat) was scanned
spaces among the fascicles clearly delineate these as per VL; thus, direct comparison can be made between sites
structures and allows architectural measurements to 1 and 2 of the anterior portion with sites 2 and 3 on the lateral
be made. Measures of fascicle angle (≤ 1.5°; Kawakami portion. VL: vastus lateralis, VM: vastus medialis, RF: rectus
femoris, VIant: vastus intermedius (anterior portion),
et al. 1993; Narici et al. 1996; Chleboun et al. 2001) and
VIlat: vastus intermedius (lateral portion). Solid bars show
fascicle length (≤ 1.5 mm; Kawakami et al. 1993) have the approximate transducer angles used to obtain images
been shown to be similar to those measured directly in parallel to the fascicles on VL, VM and RF (VI not shown).
© 2006 The Authors

calculated as the mean of the distances between super-

ficial and deep aponeuroses measured at the ends of
each 3.8-cm-wide sonograph. Each sonograph was digi-
tized twice on separate occasions and the mean values
recorded. Sonograph pairs were re-digitized separately
when muscle thickness differed by more than 2 mm or
fascicle angle differed by more than 1°.
Fascicle length
At most sites tested, the fascicles were too long to be

measured from origin to insertion. Hence, fascicle
length was estimated using Eq. (1):
Fascicle length = sin(γ + 90°) × MT/

Fig. 2 Sagittal plane ultrasound scan of vastus lateralis (VL)
sin(180° − (γ + 180° − θ)), (1)
and the lateral portion of vastus intermedius (VIlat). Landmarks
corresponding to aponeuroses (points 1– 4) and fascicles
(points 5–6) were digitized (see Methods), as in the image where MT is the muscle thickness, θ is the fascicle angle
above. Muscle thickness was calculated as the mean of the and γ is the angle between the superficial and deep
vertical distances between points 1–3 and 2– 4. Fascicle angle,
aponeuroses. Fascicle length was only estimated at the
measured from 3 to 4 mm above the deep aponeurosis
(dashed line) to mid-muscle was calculated as the angle mid-section of each muscle as: (1) such estimates assume
between points 2– 4 and 5 – 6. Aponeurosis angle was that no fascicle or aponeurosis curvature exists, which
calculated as the positive angle between points 3 – 4 and 1–2.
appeared to be true only at the mid-point of the
muscles, and (2) it is also assumed that muscle architecture
is relatively isotropic, which is not true toward the
traced without interruption across the image (i.e. they end of a muscle where it generally changes shape in
did not run out of the plane of the sonogram). Sono- response to space constraints. Owing to significant
graphs were obtained at proximal, middle and distal fascicle curvature in some subjects, the reliability of fas-
sites on vastus lateralis (VL), vastus medialis (VM), vastus cicle length estimates for VM was unacceptably low, so
intermedius (VI) and rectus femoris (RF). VI was scanned they are not presented. Errors of approximately 2–7%
simultaneously with both VL and RF to obtain sono- (Finni et al. 2001, 2003b) have been reported previously
graphs of both anterior (VIant) and lateral (VIlat) portions. with similar methods of estimating fascicle length when
Scans were taken at 15 muscle sites across the quadriceps the entire fascicle cannot be visualized.
femoris at proximal, middle and distal sites as described
in Fig. 1. An example sonograph is shown in Fig. 2.
Reliability of measures
The sonographs were imported into image digitizing
software (Peak Motus, Peak Technologies, CO, USA) Digitizing reliability was determined by calculation of
and landmarks corresponding to the muscle fascicles the intraclass correlation coefficient (ICC) and typical
and aponeuroses were digitized as shown in Fig. 3. Two errors based on the double digitization of 50 randomly
points on each selected fascicle were digitized, one selected images from each muscle site. ICCs were very
approximately 3 – 4 mm from the deep aponeurosis and high for muscle thickness and fascicle angle (0.948–
a second at 50% of the distance from the deep to 1.000), with little variation between muscles and muscle
superficial aponeurosis. Although there was little fas- sites. Typical errors between repeated measurements
cicle curvature at most sites, there was a tendency for for muscle thickness and fascicle angle ranged from
fascicles to curve slightly close to their insertion with 0.08 to 0.63 mm and 0.27 to 0.63°, respectively.
the deep aponeurosis, as predicted by Van Leeuwen & Reliability of the complete procedure, including the
Spoor (1992). Therefore, measuring from approxim- location of imaging sites, acquisition and digitization of
ately 3– 4 mm above the aponeurosis allowed accur- images, and calculations of architectural parameters, was
ate delineation of the fascicles. Muscle thickness was determined at each muscle’s midpoint. The procedure
© 2006 The Authors

Fig. 3 Intramuscular architecture of the

quadriceps femoris. Fascicle angles
[measured between the aponeurosis
(a) and the fascicle (f)] vary considerably
along the lengths of VL, VM and RF,
but are relatively constant in VI; the
magnitude of fascicle angle is
represented as depth of colour and
gradient of the representative fascicle
(dotted line). Example sonographs are
shown immediately below each muscle.
Muscle thickness is also relatively
variable in VL, VM and RF, but not in VI;
muscle thickness is drawn to scale. Distal
VM thickness was not measured in the
present study so an estimate has been
used to aid readability. RF is shown as a
shorter muscle than the vastii muscles;
however, muscle lengths are not drawn
to scale.
for measuring muscle thickness and fascicle angle was

Data analysis
also very reliable with ICCs of 0.882–0.970 and 0.899–
0.991 (typical errors of 0.74 – 0.97 mm and 0.24–1.22°), The intramuscular homogeneity of muscle architecture
respectively. The reliability of fascicle length estimates was examined in two ways. First, differences in the
was slightly less reliable with ICCs of 0.758 – 0.863 with absolute values of muscle thickness, fascicle angle and
typical errors of 10.2–19.4 mm. The slightly lower reli- fascicle length were compared between sites within each
ability is due to the small errors in the measurements muscle using paired t-tests with Bonferroni adjustment
of muscle thickness, fascicle angle and aponeurosis angle (α = 0.0167 for comparisons between three sites within
being multiplied during fascicle length estimation. each muscle); a reduction in type 1 error was considered
© 2006 The Authors

more important than a reduction in type 2 error. Second, pared with other subjects, a subject’s rank by magni-
relative variation at different muscle sites within each tude for an architectural parameter in one muscle was
muscle was examined using Pearson product moment the same as their rank for the parameter in another.
correlations to determine whether there were relation- In addition, subjects with above-median scores for an
ships between the muscle sites irrespective of their architectural parameter for one muscle were compared
absolute values (significant correlation = P < 0.05). against those with below-median architecture for their
Absolute differences between sites of different mus- architecture in other muscles using a multivariate
cles were not assessed because the positions at which analysis of variance (MANOVA) with repeated measures
the sonographs were taken from each muscle were not (α = 0.05). This test determined whether those that
identical. However, comparisons between muscles were were ‘above median’ for a parameter in one muscle
made in two ways. First, a dimensionless difference index were also above median in others, i.e. whether relative
(δ2−1) was calculated using a variation of the method architecture was similar across muscles.
proposed by Lieber et al. (1992): Finally, the mean z-score for each parameter for the
whole quadriceps femoris was calculated using the z-
2 2
([MT2 − MT1]/MT1) + ([θ2 − θ1]/ θ1) scores for each muscle. In order to examine relationships
δ 2−1 = , (2)
+ ([FL2 − FL1]/FL1)
2 between the architecture of each muscle with the whole
quadriceps femoris, we computed correlation statistics,
where MT is the muscle thickness, θ is the fascicle angle and used step-wise linear regression (inclusion criterion
and FL is the fascicle length measured at mid-muscle, P < 0.05; exclusion criterion P > 0.10) to predict whole
each representing muscles 1 and 2. MT and θ values quadriceps femoris architecture from that of individual
were estimated as the mean of measures taken at each muscles. This procedure assumes that each muscle’s
of the three sites (i.e. the value for each site was taken architecture contributes equally to the whole quadriceps
as representative of one-third of the muscle). For VM, femoris, which we have called the ‘equal ratio’ method.
muscle thickness was taken as the mean of proximal However, each muscle’s contribution to force pro-
and middle sites as it was not measured distally. However, duction is probably not the same, so we also ran the
the index was also computed assuming a thickness of analyses in three other ways. First, we apportioned each
30 mm at the distal site; there were no significant dif- muscle’s contribution according to its relative muscle
ferences in the values regardless of which method was volume (i.e. ‘volume’ method). Using MRI (3.0 T Mag-
used. Fascicle length values were taken as the fascicle netom, Siemens AG, Berlin, Germany), muscle volume
length measured at mid-muscle. The difference index was measured in a separate group of 11 women and 10
was determined for each muscle pair, with values of δ2−1 men whose physical characteristics matched as closely
< 0.3 representing architectural similarity, 0.3 < δ2−1 as possible the subjects used for the architecture analysis.
< 0.8 representing moderate similarity and δ2−1 > 0.8 Subjects lay supine with their legs straight while T2-
representing architectural dissimilarity. The index allows weighted axial scans using a standard body coil were
absolute comparisons of muscle architecture, i.e. inter- taken of the thigh from the inferior portion of the
muscular similarity. greater trochanter to the superior border of the patella
Second, an overall relative value for each architectural (TR: 4260 ms, TE: 95 ms, averages: 3, FoV: 200 × 200, slice
parameter was calculated for each muscle, and these thickness: 4 mm, slice separation: 14 mm, centre-to-centre
values were compared. To calculate an overall value, slice distance: 16 mm). Muscle cross-sectional area was
measurements at each muscle site were converted to z- measured for each slice using Scion Image for Windows
scores [(value – mean)/SD] and the values at each of the (free to download at http://www.scioncorp.com/frames/
three sites averaged, i.e. subjects were ranked based on fr_download_now.htm) and muscle volume was calcu-
their magnitudes of muscle thickness, fascicle angle lated by summing the product of cross-sectional area
and fascicle length. Each muscle therefore had an and slice thickness. The sum of the z-scores for each
overall value expressed as a mean z-score. Correlations muscle was taken as the whole quadriceps z-score after
between the z-scores of the muscles were computed to the values were multiplied by their proportion of the
determine whether subjects’ scores for one muscle muscle volume (i.e. 100% of the volume = 1). The pro-
were similar to their scores for another muscle. Thus, portional contributions of each muscle to the whole
from these analyses we could determine whether, com- quadriceps architecture are reported in Table 1.
© 2006 The Authors

Table 1 Muscle proportions used for estimation of whole Table 2 Muscle thickness and fascicle angle measured in
muscle architecture different muscles (means ± SD). For each muscle, data from
proximal (first number), middle (second number) and distal
Estimation method (third number) sites are presented in order of top to bottom
Equal Estimated Muscle thickness Fascicle

Muscle ratio* Volume† PCSA‡ contribution§ Muscle (mm) angle (°)
vastus lateralis 0.250 0.352 0.221 0.242 Vastus lateralis 21.5 ± 3.3 9.7 ± 3.9
vastus medialis 0.250 0.218 0.243 0.122 22.8 ± 3.2 13.2 ± 3.2a
rectus femoris 0.250 0.136 0.236 0.251 16.4 ± 4.3a,b 11.7 ± 3.8
vastus intermedius 0.250 0.294 0.300 0.396 Vastus medialis 20.9 ± 4.5 10.2 ± 3.1
26.8 ± 6.3a 14.0 ± 3.9a
*Whole muscle architecture estimated by assigning equal Not measured distally 28.5 ± 5.7a,b
weighting to each muscle.
Rectus femoris 26.9 ± 5.0 14.0 ± 4.3
†Muscle contribution apportioned by their contribution to whole
24.0 ± 4.8a 13.4 ± 3.6
quadriceps volume, which was determined by MRI imaging; data
20.1 ± 4.9a,b 10.1 ± 3.4a,b
collected on 21 subjects of similar age and physical stature.
‡Muscle contribution apportioned by their relative physiological Vastus intermedius 19.3 ± 4.6 6.0 ± 3.3
cross-sectional areas (PCSA); data from Narici et al. (1992). (anterior portion) 17.6 ± 4.1 7.1 ± 2.9
§Muscle contribution apportioned by their estimated contribution 17.0 ± 4.3a 6.4 ± 2.7
to total joint moment assessed by electrophysiological techniques; Vastus intermedius 6.8 ± 3.1 14.6 ± 2.2
data from Zhang et al. (2003). (lateral portion) 5.5 ± 2.4 15.9 ± 2.3
7.2 ± 3.2 14.7 ± 3.3
Significantly different from aproximal site and bmiddle site.

In a second analysis, we apportioned each muscle’s
contribution according to previously published estimates
of physiological cross-section (Narici et al. 1992), before (i.e. anterior vs. lateral portions); the anterior portion
summing z-scores to find an overall z-score (i.e. ‘PCSA’ was relatively uniform, whereas the lateral portion was
method). smaller at mid-thigh compared with proximal and distal
In a third analysis, we apportioned contributions regions. Fascicle angulation followed a slightly different
according to the estimated contribution to knee extensor pattern; whereas the fascicle angles of muscles generally
moment (i.e. ‘estimated contribution’ method) assessed decreased in the order VM > VIlat > VL > RF > VIant, the
electrophysiologically by Zhang et al. (2003). In their fascicle angulation usually decreased proximo-distally in
study, electrical stimulation was used to activate VL and RF, increased proximo-distally in VM, and remained
each quadriceps muscle independently while both the relatively constant in VI (anterior and lateral portions).
resultant force and EMG [indwelling (vastus intermedius) Thus, muscle thickness and fascicle angle varied consider-
or surface electrodes (vastii muscles)] were measured to ably along the lengths of some of the muscles (particularly
determine the activation–moment relationship. They then VM, VL and RF) but not others (particularly VI), so archi-
used the EMG collected during voluntary isometric tectural measurements taken at one part of the muscle
knee extensions to estimate the relative contributions seem not to be indicative of those taken at other sites.
of each muscle.
Homogeneity of within-muscle architecture

(relative differences)
Results
Proximal VL fascicle angle was significantly correlated
with the fascicle angles at middle (r = 0.48, P < 0.01)
(absolute differences)
and distal (r = 0.48, P < 0.01) sites, indicating that sub-
Muscle thickness and fascicle angle data are presented jects with larger fascicle angles at the proximal site also
in Table 2, and representations of these are shown had large angles at the other sites. The correlation
in Fig. 3. With respect to muscle thickness, VL and RF between the middle and distal sites, however, did not
decreased proximo-distally along the thigh, whereas reach significance (r = 0.24, P = 0.13). Muscle thickness
VM increased. Variations in VI thickness were different at each site was also correlated with the other sites
depending on the direction of sonograph acquisition (r > 0.5, P < 0.01), indicating a consistency in VL size. A
© 2006 The Authors

similar trend was noticed for VM and RF. In VM, there Table 3 Difference Index (δ2−1) quantifying the architectural
was a significant correlation between proximal fascicle difference between the quadriceps muscles
angle and that at the distal site (r = 0.41, P < 0.05),
VM RF VIant VIlat
although correlations with the middle site did not reach
significance. In addition, muscle thickness at proximal VL 0.36 0.20 0.66 0.76
and middle sites was significantly correlated (r = 0.57, VM – 0.08 0.46 0.55
RF – 0.70 0.77
P < 0.01); muscle thickness was not measured at the VIant – 1.97
distal site. In RF, there was a significant correlation VIlat –
between the fascicle angle at middle and distal sites
δ2−1 < 0.3 represents architectural similarity, 0.3 < δ2−1 < 0.8
(r = 0.38, P < 0.05), although correlations did not reach
represents moderate similarity, and δ2−1 > 0.8 represents
significance for the proximal site, whereas there were architectural dissimilarity. The three superficial quadriceps
significant correlations for muscle thickness at each of muscles were more similar to each other (δ2−1 < 0.36) than the
the three sites (0.56 < r < 0.74, P = 0.000 – 0.002). There- deeper vastus intermedius.
fore, for VL, VM and RF, subjects tended to have a

similar relative architecture along the length of the
muscles. These trends were similar when men and women between muscles. As detailed in Table 3, the index was
were analysed separately, although some of the cor- generally low for comparisons between the superficial
relations of similar magnitude did not reach statistical quadriceps muscles (δ2−1 < 0.36) but higher between
significance because of the smaller sample size. superficial and deep muscles (i.e. VI, δ2−1 = 0.46–0.77).
For VIant, there was a correlation between fascicle Thus, there was reasonable architectural similarity
angle at middle and distal sites (r = 0.47, P < 0.05), between the superficial quadriceps only. Furthermore,
although there was no correlation with the proximal the anterior and lateral portions of the VI were highly
site. Proximal muscle thickness was correlated with dissimilar in their architecture (δ2−1 = 1.97). So even
both the middle (r = 0.52, P < 0.01) and the distal sites within the same muscle, architectural dissimilarity was
(r = 0.38, P < 0.05), although the correlation between significant.
middle and distal sites did not reach significance z-scores were also calculated for each architectural
(r = 0.30, P = 0.10). Therefore, subjects who had larger parameter at each muscle site and an average z-score
fascicle angles at mid-muscle also had a larger fascicle was computed to provide a global parameter for the
angle distally, but not proximally, whereas subjects whole muscle; results of RM MANOVA and correlation
who had thicker muscles proximally were also thicker analyses are presented in Table 4. For VL, subjects with
in other regions. For VIlat, whereas there was a correla- a greater-than-median whole muscle fascicle angle or
tion between fascicle angle at proximal and middle fascicle length (fascicle length at mid-muscle was taken
sites (r = 0.71, P < 0.001), there was no correlation with as representative of the whole muscle) did not have
the distal site. There were no significant correlations statistically different fascicle angles or lengths in any
for muscle thickness between any of the sites, suggest- other muscle. Also, those with greater muscle thickness
ing that subjects with thicker muscles at one site did had a greater muscle thickness in VIlat (P < 0.05) but not
not necessarily have thicker muscles at another site. in any other muscle. These results were consistent with
Therefore, the consistency of muscle architecture in VI the correlation analysis, which showed no relationship
varied between anterior and lateral portions; subjects between the architecture of VL and other muscles, with
who had a specific muscle thickness and/or fascicle the single exception of VL thickness being correlated
angle at one site also had a similar relative architecture with VIlat thickness (r = 0.41, P < 0.05). Therefore,
at other sites in the anterior portion, but this was not subjects with a given magnitude of architecture in VL
the case for the lateral portion of the muscle. relative to the other subjects did not have the same
relative magnitude of architecture in the other muscles
in the quadriceps femoris.
Intermuscular variation in architecture
Nonetheless, subjects with larger VM fascicle angles
also had larger VL and RF fascicle angles (and there was
The difference index (δ2−1) calculated between muscles a trend toward larger angles in VIant, P = 0.09). There
allowed comparisons of overall muscle architecture was a strong relationship between VM and RF fascicle
© 2006 The Authors

© 2006 The Authors
Table 4 Relative relationships between muscle architectural variables for different muscles within the quadriceps femoris
VL VM RF VIant VIlat
± median FA TH FL FA TH FA TH FL FA TH FL FA TH FL
VL FA F = 52.8*** 4.3* 5.4* NS NS NS NS

r=1 0.55** −0.48** NS NS NS NS
TH 8.5** 34.8*** NS NS NS NS 6.3*
0.55** 1 NS NS NS NS 0.41*
FL NS NS 12.3** NS NS NS
−0.48** NS 1 NS NS NS
VM FA 5.6* 20.9*** NS 25.8*** NS NS
1 0.48** NS 0.38* NS
TH NS 24.8*** NS NS NS
0.48** 1 NS 0.46* NS
RF FA 6.6* 20.6*** 27.8*** NS NS NS NS
0.37* 0.93*** 1 NS NS NS NS
TH NS NS NS 54.4*** NS NS NS
NS NS NS 1 0.44* NS NS
FL NS NS 6.8* 17.6*** NS NS
NS NS 0.44* 1 NS NS

Vlant FA NS NS NS 43.9*** 5.5* NS NS
NS 0.38* NS 1 0.51** −0.40* NS
TH NS NS NS 4.9* 43.9*** NS NS
NS 0.46* NS 0.51* 1 NS NS
FL NS NS NS NS 17.0*** NS
NS NS −0.40* NS 1 NS
Vlant FA NS NS NS NS 25.8*** NS NS
NS NS NS NS 1 0.63*** NS
TH 10.3** NS NS NS NS 14.7** NS
0.41* NS NS NS 0.63*** 1 0.44*
FL NS NS NS NS NS 69.2***
NS NS NS NS 0.44* 1
After z-scores were computed for each variable for each subject, subjects with greater-than- or less-than-median scores were compared (left column) and correlations among the variables
were computed. For each relationship, the top number indicates the F-ratio and the bottom number the correlation (r). Asterisks indicate levels of significance: *P < 0.05, **P < 0.01,
***P < 0.001. FA: fascicle angle, TH: muscle thickness, FL: fascicle length.
Table 5 Correlations between individual muscle architecture scores and whole quadriceps femoris architecture estimated by four
methods (see Methods)
Equal ratio Volume PCSA Estimated contribution
Parameter FA TH FL FA TH FL FA TH FL FA TH FL
VL FA 0.61*** 0.73*** 0.59** 0.63***

TH 0.63*** 0.71*** 0.58** 0.59**
FL 0.70*** 0.81*** 0.64*** 0.61***
VM FA 0.91*** 0.83*** 0.91*** 0.85***
TH 0.75*** 0.75*** 0.76*** 0.61**
RF FA 0.91*** 0.55** 0.90*** 0.84***
TH 0.50** 0.40* 0.48** 0.58**
FL 0.59** NS 0.55** 0.49**
VIant FA 0.58** 0.64*** 0.62*** 0.70***
TH 0.60** 0.60** 0.66*** 0.71***
FL 0.60** 0.65*** 0.69*** 0.76***
VIant FA NS NS NS NS
TH 0.37* 0.37* NS 0.39*
FL NS NS NS NS
Generally, architecture in VIlat was poorly correlated with whole quadriceps femoris architecture. Asterisks indicate levels of significance:
*P < 0.05, **P < 0.01, ***P < 0.001. FA: fascicle angle, TH: muscle thickness, FL: fascicle length.
angles (r = 0.93, P < 0.001), suggesting that fascicle angle PCSA and (4) estimated contribution. Although there
in one muscle was highly predictive of the other. Despite was a strong intercorrelation between the methods
this, subjects with greater muscle thickness or fascicle for calculating whole quadriceps thickness (R2 > 0.95),
length in VM did not have greater magnitudes in other fascicle angle (R2 > 0.96) and fascicle length (R2 > 0.92),
muscles. These results were mirrored to some extent in the contributions of individual muscles toward the score
RF, where subjects with larger RF fascicle angles also had for whole quadriceps architecture varied (see Table 5).
larger VL and VM fascicle angles but subjects with greater In particular, apportioning muscle contribution relative
muscle thickness or fascicle lengths did not have greater to each muscle’s volume was different to the other
magnitudes in other muscles. Thus, although there was methods. For example, thickness of VL was more highly
a strong correlation between VM and RF fascicle angle, correlated with whole quadriceps thickness (r = 0.71,
there was no significant correlation with VL. P < 0.001) when whole quadriceps thickness was esti-
In VI, subjects with greater thickness of VIlat also had mated using the volume method than when muscle
a thicker VL (P < 0.01), but there were no other signifi- contributions were apportioned using equal ratio (r =
cant differences in either portion of VI. These results 0.63), PCSA (r = 0.58) or estimated contribution (r = 0.59),
were consistent with correlation statistics, which did as was VL fascicle length (r = 0.81, P < 0.001) when
not support a relationship between relative VI architec- compared with the equal ratio, PCSA and estimated
ture and that in other muscles. Therefore, although contribution methods (r = 0.70, 0.64 and 0.61, respec-
there was some constancy of fascicle angle between tively). Differences between the methods were also
VM, VL and RF, subjects with a certain relative magnitude highlighted by the fact that statistically significant
of muscle thickness or fascicle length of one quadriceps gender differences in whole quadriceps thickness were
muscle seemed not to have a similar relative magnitude detected when the equal ratio (P = 0.02), PCSA (P = 0.03)
of other muscles. and estimated contribution (P = 0.01) methods were
used, but not when contributions were apportioned
by muscle volume (P = 0.10). These data show that
Contribution of individual muscle architectures to
gender-related size differences were most clearly seen
whole quadriceps architecture
when quadriceps thickness was estimated by the esti-
Whole quadriceps architecture was estimated by four mated contribution method, although they could not
methods (see Methods): (1) equal ratio, (2) volume, (3) be detected using the volume method.
© 2006 The Authors

Table 6 Regression equations to predict

whole muscle architecture estimated by Method Parameter Equation
each of the four methods (see Methods)
Equal ratio FA VM × 0.347 + VL × 0.283 + VIant × 0.238 + RF × 0.407
TH VM × 0.528 + RF × 0.351 + VL × 0.379 + VIant × 0.312
FLa VL × 0.557 + RF × 0.513 + VIant × 0.513
Volume FA VM × 0.335 + VL × 0.441 + VIant × 0.310 + RF × 0.246
TH VM × 0.454 + VL × 0.527 + VIant × 0.362 + RF × 0.189
FLa VL × 0.699 + VIant × 0.538 + RF × 0.249
PCSA FA VM × 0.344 + VIant × 0.292 + VL × 0.255 + RF × 0.392
TH VM × 0.517 + RF × 0.334 + VIant × 0.378 + VL × 0.338
FLa VIant × 0.612 + VL × 0.489 + RF × 0.481
Estimated FA RF × 0.439 + VIant × 0.406 + VL × 0.295 + VM × 0.182
contribution TH VIant × 0.532 + VL × 0.396 + RF × 0.380 + VM × 0.278
FLa VIant × 0.680 + VL × 0.451 + RF × 0.431
Variables were added step-wise by greatest contribution to explained variance; equations

are presented in order of entry. Equation constants were less than 0.00008 in all cases and
are therefore not presented. Adjusted R2 = 1.000 and standard error of the estimate
< 0.0004 for all equations. aVM fascicle length was included in these analyses (see Methods).
FA: fascicle angle, TH: muscle thickness, FL: fascicle length.
equal ratio (r = 0.67), PCSA (r = 0.68), volume (r = 0.71)

Prediction of whole quadriceps architecture from
or estimated contribution (r = 0.65) methods were used
individual muscle architecture
(P < 0.001). In addition, fascicle angle and fascicle length
Regression equations showing the prediction of whole were negatively correlated (r = – 0.45, −0.48, −0.54 and
quadriceps architecture from individual muscles are −0.53 for the equal ratio, PCSA, volume and estimated
presented in Table 6. Variables were added step-wise contribution methods, respectively; P < 0.05). Thus, when
by greatest contribution to explained variance; equa- architecture at all muscles sites was combined to give
tions are presented in order of entry. For muscle thick- an average muscle architecture for the quadriceps
ness, VM was the best indicator of whole quadriceps femoris, those muscles with greater thickness tended
thickness, although, depending on the method of to have larger fascicle angles, whereas those with longer
estimating whole quadriceps thickness, VL, RF and VIlat fascicles had lesser fascicle angles.
were also reasonable indicators. VM fascicle angle was Significant correlations between muscle thickness
also the best predictor of whole quadriceps fascicle and fascicle angle were also seen for VL, VM, VIlat and
angle, except when the estimated contribution method VIant individually, but not RF (see Table 7a–e). However,
was used to estimate whole quadriceps fascicle angle; fascicle angle was only negatively correlated with
VL and VIant were also useful predictors. VL and VIant fascicle length for VL and VIant (fascicle length not
were the best indicators of whole quadriceps fascicle measured in VM). Thus, although there was a strong
length, although VM fascicle length was not used in indication that subjects with large quadriceps fascicle
this analysis (see Methods). Therefore, VM fascicle angle angles to have longer fibres, this was true only for the
and muscle thickness scores seemed to be the best VL and VIlat when muscles were analysed separately.
predictors of quadriceps femoris estimates, while VL and
VIant scores were reasonable predictors of all architec-
tural parameters. VIlat, however, was not a useful predictor Discussion
of quadriceps thickness, fascicle angle or fascicle length.
Intermuscular variation in quadriceps architecture
Validity of whole quadriceps estimates: interaction
In order to produce forces with broad magnitude, range
between muscle thickness, fascicle angle and fascicle
and velocity characteristics, it has been suggested that
length
muscles within synergistic groups tend to vary in their
In the whole quadriceps, muscle thickness was positively architecture (Lieber & Fridén, 2000). Our examination
correlated with fascicle angle, regardless of whether the suggests that this is only partly true for the human
© 2006 The Authors

Table 7 Correlations between muscle thickness, fascicle angle This intermuscular variation suggests that the super-
and fascicle length based on z-scores (see Methods) for each ficial and deep quadriceps muscles have different roles,
muscle (a–e)
and could be recruited uniquely based on their force-
FA TH FL generation properties. Such recruitment variability has
been previously shown. In the quadriceps femoris,
(a) vastus lateralis Zhang et al. (2003) found that the relative contribution
FA 1 0.55** −0.48**
TH 1 NS
of each muscle to the total isometric knee extensor
FL 1 moment changed as contraction force increased, such
(b) vastus medialis that the contribution of VI decreased whereas the con-
FA 1 0.48** tribution of VL, VM and RF increased (the increase in RF
TH 1
approached statistical significance, P = 0.06). Similar
FL
findings have been reported for the biceps brachii,
(c) rectus femoris
which also consists of multiple muscles attaching via a
FA 1 NS NS
TH 1 0.44* common tendon (Brown et al. 1993). Nonetheless, other
FL 1 studies employing surface EMG analyses have also shown
(d) vastus intermedius (anterior) muscle-specific changes in quadriceps muscle activation
FA 1 0.51** −0.40* as force levels are varied (Ebersole et al. 1999; Pincivero
TH 1 NS
FL 1
& Coelho, 2000; Rabita et al. 2000; Pincivero et al. 2002,
2004b; Cramer et al. 2004), knee joint angle, and there-
(e) vastus intermedius (lateral)
FA 1 0.63*** NS fore whole quadriceps femoris muscle length, is changed
TH 1 0.44* (Ebersole et al. 1999; Signorile et al. 1995), velocity is
FL 1
increased (Cramer et al. 2004) or contraction mode is
Asterisks indicate levels of significance: *P < 0.05, **P < 0.01, changed from concentric or isometric to eccentric
***P < 0.001. FA: fascicle angle, TH: muscle thickness, FL: fascicle (Narici et al. 1996; Pincivero et al. 2006). These findings
length. are not predicted by a model that assumes a similarity
in architecture of the superficial quadriceps muscles.
quadriceps muscles. The difference index calculated to However, these variations might be explained by
provide quantitative assessment of intermuscular vari- differences in intramuscular architecture, as discussed
ability revealed a close similarity between the mean below (‘Homogeneity of within-muscle architecture’).
architecture of the three superficial quadriceps muscles Movement-specific muscle activation strategies will have
(i.e. VL, VM and RF; δ2−1 < 0.36), despite RF being bi- to be examined further as a relationship between
articular and VL and VM being mono-articular. Given muscle architecture and recruitment has yet to be dem-
that muscles within the human quadriceps femoris act onstrated. Notwithstanding, there is some support for
via a common (patellar) tendon, this similarity in mean hypothesis 1, that the architecture of each muscle within
architecture is indicative of a similarity in their force- the quadriceps should be different, for the superficial
generating potential during uni-joint knee extension vs. deep quadriceps muscles; however, there seems to
(where RF functions as a mono-articular knee extensor), be some uniformity among the superficial muscles.
although their angle of attachment to the tendon The architectural differences between muscles would
would also influence their contribution to knee func- also affect the efficiency of force transfer to the tendon,
tion. Nonetheless, the deeper VI is architecturally dis- given that forces are transferred laterally between
similar to the superficial muscles, indicating that it has muscles (e.g. Huijing, 2003; Maas & Huijing, 2005). For
a different force-generating potential, and possibly a example, the shorter RF fascicles attaching at larger
different role during knee extension. A clear picture of angles (12.5°, 112.3 mm length at 45° knee angle) would
its role is difficult to ascribe because its architecture be expected to generate high relative peak forces over
varies significantly. The anterior portion is characterized shorter length ranges, which is commensurate with
by long fibres attaching at small angles to the aponeu- reports that it works quasi-isometrically during many
rosis, whereas the lateral portion has shorter fibres human movements (Gregoire et al. 1984; Ingen Schenau
attaching at larger angles (δ2−1 = 1.97); the medial por- et al. 1987; Jacobs & Ingen Schenau, 1992). This can be
tion was not examined. compared with the immediately adjacent, architecturally
© 2006 The Authors

Fig. 4 Effect of intermuscular fascicle length and angle differences on tendon/aponeurosis excursion. When rectus femoris (RF)
fascicles shorten (from FLRF1 to FLRF2, where FL is the fascicle length) and rotate (θRF1 to θRF2, where θ is the fascicle angle) in this
example, the aponeurosis or tendon displaces 10 mm. This represents a fascicle shortening of 9.7 mm, or 8.7%. If the vastus
intermedius (VI; anterior region) fascicle shortens by the same proportion, 14.0 mm, or 8.7%, and rotates, the aponeurosis or
tendon displaces 14.2 mm, which is a difference (Diff.) of 4.2 mm, or 42%. Thus, even when fascicle rotation is accounted for, the
differing architectures of RF and VI result in differential displacement of the tendon or aponeurosis. Either RF and VI must be
activated uniquely, or there will be a shear force in the tendon or aponeurosis. In this example, it was assumed that the muscle
thickness would remain constant given that RF and VI apply opposing forces of similar magnitude on the tendon/aponeurosis.
dissimilar VIant (6.5°, 162.1 mm at 45° knee angle), which scopy experiments has shown that the increase in O2
is suited to generating lower peak forces over greater utilization, and therefore energy consumption, by the
length ranges. Even when fibre rotation is accounted superficial quadriceps muscles with increasing force
for, which would allow RF fascicles to shorten less levels is similar between muscles (de Ruiter et al. 2005).
relative to the whole muscle shortening, there is still These data are consistent with the hypothesis that the
a significant force–length discrepancy between the unique activation of muscles optimizes efficiency under
muscles. As shown in Fig. 4, a similar relative shortening different loading conditions. Furthermore, strength
of RF and VI (anterior) fascicles would result in a different training of the quadriceps has not only been shown to
tendon or aponeurosis displacement (10.0 mm vs. 14.2 mm alter the proportional activation of the quadriceps
in the example). The discrepancy would result in a muscles (Pincivero et al. 2004a) but has been shown to
shear stress between their aponeuroses, and is likely to reduce the EMG measured from superficial quadriceps
affect force transfer efficiency. muscles at prescribed submaximal force levels (i.e
Alternatively, a lesser recruitment of VI as contraction decreased EMG–force ratio). This latter change is also
intensity increases (see Zhang et al. 2003) would allow indicative of a greater efficiency of force production
matching of muscle shortening. Because VIant contains (Narici et al. 1996; Rabita et al. 2000). The contention
longer fibres, and presumably a greater number of that unique muscle activation would be a useful opti-
sarcomeres in series, a similar absolute shortening of RF mization strategy also suggests that the EMG–force
and VI fascicles would result in a lesser shortening of VI ratios of individual muscles, and their adaptation with
sarcomeres. This would improve force production via movement training, would be specific to the person as
optimization of force–length and force–velocity relation- architecture varies considerably interindividually. This
ships and might allow more efficient force produc- contention has been given some credence by Rabita
tion. As discussed above, variations in the recruitment et al. (2000), who showed that alterations in the EMG–
of individual muscles has been demonstrated previously, force relationship varied significantly between muscles
but the hypothesis that the strategy of unique muscle and also between individuals. Thus, although direct
activation would improve force transfer efficiency has evidence that humans adopt a unique activation strategy
yet to be examined. Data from near-infrared spectro- in order to minimize shear forces and improve force
© 2006 The Authors

transfer efficiency in the human quadriceps is not stability (Van Leeuwen & Spoor, 1992), although such
available, published data are strongly supportive of the variation might further influence muscle function and
concept. force transfer between muscles. For the same relative
Importantly, many muscle models assume that archi- shortening of fascicles in different regions of the mus-
tectural variations have little impact on the active length cles, aponeurosis displacement will be different. Most
range of muscles working in synergistic groups (see Van planimetric models of muscle assume a constant fasci-
den Bogert et al. 1998), largely because it is supposed cle angle along the length of a muscle, which might not
that connective tissues would be overstretched /injured represent the muscles appropriately. More complex
if significant discrepancies existed. Based on our data, muscle models that allow fascicle angle and length
this assumption might need to be revised; significant parameters to vary (Woittiez et al. 1984; Otten, 1988;
intermuscular architectural differences might explain Savelberg & Schamhardt, 1995) will probably be more
some of the difficulties in accurately modelling com- accurate. To estimate the extent to which models might
plex muscle groups. Thus, the effects of architectural be affected when architecture is assumed to be constant,
dissimilarity in muscles functioning as a synergistic we constructed a planimetric model of a unipennate
group are complex, and more research is required to muscle containing three segments, or parallelepipeds
understand activation strategies used to optimize (Savelberg & Schamhardt, 1995), as shown in Fig. 5. It
muscle function. Regardless, the quadriceps femoris was assumed that each segment contributed to the
probably cannot be accurately modelled as a muscle tendon force in proportion to its estimated physio-
group that provides a force exactly equal to the sum of logical cross-section (muscle thickness2/fascicle length),
each of its constituents. and the total muscle force was equal to the sum of
forces provided by each segment (see Appendix A). The
model was run with three different inputs: (1) where
each segment was identical in its architecture, with mean
VL (middle) data being assumed to run throughout the
Although mean architecture measures were relatively muscle; (2) where each segment varied in its architecture,
consistent between muscles, except for VI, our data also with mean VL proximal, middle and distal data being
show that architecture does vary significantly within used as input for the three segments; and (3) where
them. For example, fascicle angle increased proximo- each segment varied, with VL (middle) data being used
distally in VM, decreased proximo-distally in RF and was as input for the middle segment but mean − 1 SD data
greater centrally in VL. Furthermore, fascicle angle was being used from distal and proximal VL as input for
significantly different in anterior and lateral regions segments 1 and 3 (see Fig. 5). The results of the model
of VI, although it changed little along the length of show little difference in the force–length curves when
the muscle. These results are consistent with previous the muscle is assumed to have a constant architecture
reports of intramuscular architectural variation (Scott compared with a varying architecture (Fig. 5A vs. Fig. 5B)
et al. 1993; Blazevich et al. 2003) and provide little sup- when the architecture varies within the orders of
port for our second hypothesis, that the intramuscular magnitude reported for VL in the present study. How-
architecture should be relatively homogeneous. They ever, when more significant architectural variation is
are also suggestive of the possibility that different input into the model (Fig. 5C), the force–velocity
regions of the muscles have different functional relationship changes far more dramatically. Muscles that
roles, and perhaps are activated uniquely during knee vary considerably in their architecture, such as VM, will
extension. Although there are no data describing probably not to be accurately modelled when intra-
region-specific variations in activation of the individual muscular architecture is assumed not to vary, so con-
quadriceps muscles, different regions of other muscles sideration must be given as to whether uniform muscle
such as the long head of biceps brachii are differentially models are appropriate for use.
activated as force patterns are altered (ter Haar Romeny The functional implications of intramuscular variations
et al. 1984). A similar examination of the quadriceps in architecture are unclear; however, we posit two
muscles might reveal region-specific activation patterns. hypotheses (additional to the requirement for mechanical
Intramuscular architectural variation is probably re- stability; Van Leeuwen & Spoor, 1992). First, the muscles
quired in pennate muscles in order to achieve mechanical might be intended to have both ‘force-generating’ and
© 2006 The Authors

Fig. 5 A planimetric muscle model was constructed, according to Appendix A. The muscle consisted of three segments which were
either identical in their architecture (non-varying architecture, A), varied according to normal data for the VL collected in the
present study (varying architecture, B), or varied more considerably in the proximal and distal segments than the measurements
in the present study (i.e. mean minus 1 SD) (varying architecture, C). Muscle thickness (MT) and fascicle angle (θ) parameters used
in the model are shown (parameters correspond to muscle optimum length). As shown in the graph of muscle force vs. muscle
length (D), modelling the muscle as three distinct segments makes a relatively small difference to the output of the model when
architecture varies only moderately (A vs. B). However, when the muscle’s architecture varies considerably (A vs. C), the ability
for a uniform muscle model to predict muscle force is significantly reduced.
© 2006 The Authors

‘force-transfer’ regions. The greater fascicle angle and also be influenced by the activity profiles of the subjects
subsequently larger physiological cross-section in one used. Thus, although the hypothesis that intramuscular
region might perform the function of generating high architectural variation should result in region-specific
muscle forces. A second, force-transfer, region might hypertrophic adaptations to exercise training has some
be designed to have fibres in line with the tendon to support, a direct link between hypertrophy and pre-
allow greater efficiency of force transfer. The use of a existing architecture has not yet been made.
transfer section consisting of muscle, rather than ten-
don, would permit greater control over muscle–tendon
stiffness, and subsequently over movement control.
The second hypothesis is that fascicles within muscles
tend to align with the joint’s centre of rotation, which Despite architecture varying considerably along the
would improve the efficiency of force transfer from length of three of the quadriceps quartet, there was a
muscle to bone. The present data cannot confirm this constant intersubject variability of regional variation in
hypothesis because the angle of the aponeurosis rela- VL, VM and RF. For example, subjects who had a greater
tive to the tendon was not measured, so pennation proximal fascicle angle in a given muscle, relative to the
angle could not be determined. However, our findings other subjects, also tended to have greater fascicle
that fascicle angle generally increased in VM proximo- angles at middle and distal sites in that muscle. This was
distally, but not in RF and VI, and that the lateral por- not necessarily the case for VI. In the anterior portion,
tion of VI had greater fascicle angles, tend to support muscle thickness between sites was highly correlated
the hypothesis, whereas the finding that fascicle angle (i.e. subjects with greater or lesser thickness at one region
decreased from middle to distal regions in VL is not also had greater or lesser thickness at another region),
supportive. but fascicle angle was only correlated between middle
Intramuscular architectural variation should also affect and distal regions. For the lateral portion, muscle thick-
the hypertrophic response to, and therefore the func- ness between regions was not correlated, and fascicle
tion of, different regions of the muscle. The variation angle was only correlated between proximal and middle
in fascicle angle (and length) along these muscles regions. Thus, while intersubject differences in one
would probably affect fascicle strain in response to an region of a superficial muscle were similar to those in
imposed stress (Maganaris & Paul, 2000; Muramatsu et al. other regions, and therefore in broad terms relative
2002; Finni et al. 2003a). Given the assumption that the architecture in one region is a reasonable indication of
hypertrophic response of muscle is somewhat related relative architecture in another, there was less evidence
to fibre strain (e.g. Goldspink, 1978; Vandenburgh & of this for VI. Because our second hypothesis was that
Kaufman, 1979; Gregory et al. 1986; Wong & Booth, muscle architecture should be uniform throughout a
1990; Hather et al. 1991; Goldspink et al. 1992, 1995), muscle, it stands also that we would expect a subject’s
one would predict region-specific muscle hypertrophy relative architecture (i.e. compared to the rest of the
in response to muscle loading. Published data are con- sample of subjects) at one region of a muscle to be
sistent with this hypothesis (Housh et al. 1992; Narici indicative of its relative architecture at another region
et al. 1996; Häkkinen et al. 2001). Interestingly, both in all muscles within the quadriceps femoris. Thus, our
Häkkinen et al. (2001) and Narici et al. (1996) reported hypothesis was proved to be incorrect.
greater proportional hypertrophy proximally in VM
and distally in VL; our data show that in these regions
Intermuscular variation in quadriceps architecture
the fibre angles of both muscles are relatively smaller.
Unfortunately, there are no data showing a direct link
between local muscle architecture and the hypertrophic Relative intermuscular variation was considerably greater
response. A direct conclusion is also made more diffi- than that within a muscle. For example, subjects who
cult in view of the findings of Housh et al. (1992) whose had a greater mean fascicle angle of VM also tended to
subjects increased at the mid-level of VM and VL only. have greater angles in VL and RF, and this was also true
Furthermore, there is a lack of agreement as to the for RF, where greater fascicle angles were indicative of
predominant sites of hypertrophy in VI and RF. It is pos- greater angles in VL and VM. However, this was not the
sible that adaptations are exercise-specific, and might case for VL and VI (both anterior and lateral regions),
© 2006 The Authors

where relative architecture was not predictive of archi- vidual muscles whose characteristics might be repre-
tecture in other muscles. Thus, although there is some sentative of the muscle group. An understanding of
indication that the architecture in one superficial muscle this would preclude the necessity to measure architec-
is reflective of that in another, this is not completely ture of all four muscles in order to characterize an indi-
true for VL and does not hold true for the deep VI. vidual’s whole quadriceps architecture. Unfortunately,
These data are not supportive of hypothesis 3, that the according to the regression analysis (Table 6), estima-
relative architecture of one muscle should be similar to tion of whole quadriceps architecture requires the
that of another. Therefore, to gain an overall impres- inclusion of data of up to four muscles: VL, VM, RF and
sion of whole quadriceps architecture a description of VIant. This is not supportive of hypothesis 4, that the
more than one muscle is required. relative architecture of individual muscles should be
indicative of relative whole quadriceps architecture,
i.e. compared with other subjects, a subject’s rank by
Contribution of individual muscle architectures to
magnitude for an architectural parameter in one muscle
whole quadriceps architecture
was not the same as their rank for the parameter in
We also sought a global measure of architecture for another. Most typically, VM architecture was the best
the quadriceps femoris. The low mean intermuscular predictor of quadriceps architecture, as evidenced by
architecture variability in superficial quadriceps muscles its primary inclusion in the estimation of fascicle angle
was suggestive that this might be possible, although and muscle thickness for almost all methods of estimat-
these muscles differed in a region-specific manner. We ing whole quadriceps thickness and fascicle angle (see
used four methods for estimating whole quadriceps Table 6). Because VM fascicle length measures were
architecture, including (1) allowing the mean architec- unreliable owing to the excessive curvature of the
ture of each muscle an equal weighting (equal ratio), fascicles in some subjects, they could not be used to
(2) weighting by relative muscle volumes (volume), (3) predict whole quadriceps fascicle length. In this instance,
weighting by relative muscle physiological cross-sections both VL and VIant were the primary predictors. In gen-
(PCSA) and (4) weighting by estimated contribution to eral, VL and VIant were most frequent secondary predic-
knee extensor torque as reported by Zhang et al. (2003; tors. VIlat was excluded in all instances and is therefore
estimated contribution). There were no significant dif- considered a poor estimate of whole quadriceps archi-
ferences in muscle thickness, fascicle angle or fascicle tecture. Thus, VM architecture was generally the best
length values found for the whole quadriceps between predictor of whole quadriceps architecture, although
the methods (R2 > 0.92), but there were some differ- the inclusion of at least VL and/or VIant is needed for
ences in the level of contribution of each of the muscles. more accurate estimates. Although these muscles might
Most notably, whereas the equal ratio, PCSA and esti- allow a ‘representative snapshot’ of the whole quadriceps
mated contribution methods gave statistically similar to be drawn, it is not known whether gross changes in
results, estimation by volume resulted in VL thickness quadriceps architecture that result from physical train-
and fascicle length being more highly correlated with ing or detraining can be quantified by measurement of
whole quadriceps thickness, and RF thickness being less only two of the quadriceps muscles. This needs to be
highly correlated with whole quadriceps thickness. Further- determined from studies specifically examining archi-
more, a statistically significant gender difference in tectural adaptation.
muscle thickness (men > women) was found using the
equal ratio, PCSA and estimated contribution methods,
Validity of whole quadriceps estimates: interaction
but not using the volume method. Given the above, we
between muscle thickness, fascicle angle and
suggest caution in weighting architectural values by
fascicle length
volume when estimating whole quadriceps architecture.
On the whole, our estimates of whole quadriceps
architecture appear to be a valid representation as they
Prediction of whole quadriceps architecture from
predict two important relationships that are normally
individual muscle architecture
found in individual muscles. First, a gender difference
An important benefit of estimating whole quadriceps in whole quadriceps thickness was obtained for all
architecture is that it allows the identification of indi- methods except when muscles were apportioned by
© 2006 The Authors

their relative volumes. This finding of larger muscles in for clinicians and therapists whose goal is to optimize
men than women is consistently reported in the litera- movement efficiency rapidly in patients after periods
ture (Abe et al. 1998; Kubo et al. 2003). It is interesting of disuse.
to note that fascicle angles and lengths did not differ A second indicator that our whole quadriceps archi-
significantly between sexes. While relatively few studies tecture estimates appear to provide a valid representa-
have reported sex variation, Chow et al. (2000) showed tion of the muscle group is that the data show a
differences in muscle size, fibre length and fibre angle significant positive correlation (r > 0.65, P < 0.001)
between men and women in the gastrocnemius and between muscle thickness and fascicle angle regardless
soleus muscles in non-athletic, younger subjects (Chow of the method used to estimate whole quadriceps
et al. 2000). Abe et al. (1998) observed that muscle architecture, and a weaker negative correlation between
thickness and fibre angle differences in triceps brachii, fascicle angle and fascicle length (r > 0.45, P < 0.05)
vastus lateralis and gastrocnemius muscles were also (see Fig. 6), and are thus in agreement with others (Abe
different between height- and weight-matched, et al. 1998; Kumagai et al. 2000; Brechue & Abe, 2002;
resistance-trained men and women. It is not clear whether Kanehisa et al. 2003). In general, the highest correla-
differences in bodyweight (not reported by Chow et al. tions were achieved using the ‘estimated contribution’
2000) or fat-free body mass (men > women; Abe et al. method of estimating whole quadriceps architecture,
1998), or muscle size (Abe et al. 1998; Chow et al. 2000), which might give some speculative validation to the
might at least partly explain these differences given results of Zhang et al. (2003). In some respects, these
that greater muscle size is generally associated with whole quadriceps data should be treated with some
either, or both, increased fascicle angles or lengths. caution because, for example, fascicle length was posi-
Nonetheless, other reports suggest there is no difference tively correlated with muscle thickness in RF, suggesting
between male and female trained sprinters (Abe et al. that important architectural characteristics of individual
2001), Olympic soccer, gymnastics and judo athletes muscles might be missed if only whole quadriceps values
(Ichinose et al. 1998). Given that there are sex-dependent are used for analysis.
differences in muscle force production and rates of
injury, more effort should perhaps be directed at
Conclusion
understanding sex-dependent architectural differences.
Our finding of a lack of sex difference in the quadri- To our knowledge these data are the first to describe
ceps also implies that muscle activation strategies, and the complex architecture of the quadriceps femoris in
their changes with exercise training, should also be normal humans. Although there is significant, region-
similar. This is consistent with published data showing specific variation in architecture of the superficial
that alterations in the EMG–force relationship of the quadriceps muscles that might affect force transfer
muscles are the same in men and women as movement efficiency and complicate muscle models, there is some
velocity (Cramer et al. 2004) and contraction mode generality with respect to relative intermuscular archi-
(Pincivero et al. 2006) are varied. However, it is not con- tecture of the superficial muscles. This is not true for
sistent with the finding that the proportional contribu- the deep VI. Our estimates of whole quadriceps archi-
tion of VL increased more in men, and the contribution tecture are important in that we were able to provide
of VM and RF decreased in women, as knee extension evidence that the architecture of single muscles cannot
contraction force increased (Pincivero et al. 2002). be used as an estimate for the whole quadriceps, but
Because muscle architectural measurements were not that examination of VM, along with possibly VL and
performed in these studies, it is not possible to ascribe VIant, can provide a snapshot. It also verified a sex-
their inconsistent findings to differences in muscle dependent size effect of muscles, and the assumed
architecture, although other factors such as differing relationships between muscle thickness, fascicle angle
exercise patterns between the men and women need and fascicle length that are often shown in single muscles.
also to be assessed. To our knowledge, no research has Assuming the architecture of individual muscles is
examined training-related changes in quadriceps indicative of their function, and given that the quadri-
activation between men and women. The idea that ceps muscles act via a common tendon during uni-joint
alterations in synergist activation patterns might be the knee extension, these data allow speculation of the
same in men and women has particular implications functional roles of individual muscles. This is important
© 2006 The Authors

Fig. 6 Relationships between whole

quadriceps muscle thickness and fascicle
angle (top graph: a), and fascicle angle
and fascicle length (bottom graph: b).
Whole quadriceps architecture in these
examples was calculated by the equal
ratio method (see Methods), which was
associated with the lowest correlations
between the variables when compared
with PCSA, volume and estimated
contribution methods. Muscle thickness
and fascicle angle were positively
correlated, while fascicle angle and
fascicle length were negatively
correlated. Magnitudes are presented as
z-scores.
for understanding human movement strategies, and be little sex difference in these adaptations as our data
assists in the development of muscle models and to do not show a sex difference in muscle architecture
inform the design of mechanized locomotor systems of the quadriceps in subjects with a common exercise
including prosthetic and robotic limbs. By examining background. Indeed, these hypotheses seem to be largely
architectural variations across the quadriceps and con- consistent with the published data; however, the
sidering the effects of muscle architecture on function, assumption that architecture reflects function has not
our data lead to the assumptions that: (1) changes in been empirically tested. Nor has it been determined
the relative activation of individual muscles should change whether the relationships reported here remain con-
as force, velocity movement range and contraction stant after periods of physical training or detraining.
mode are altered so that muscles with the most appro-
priate architectural design provide a greater contribu-
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© 2006 The Authors

Equations: 4. θnew = sin−1(MT/Lfasnew)
1. Lfasopt = MT/sin θopt where θnew is the new fascicle angle achieved after a
given fibre length change.
where Lfasopt is the fascicle (fibre) length at muscle opti-
mum length, MT is the muscle thickness and θopt is the
5. Fseg = cos θnew × Ffas
fascicle angle at optimum muscle length.
2. Lfasnew = √(c − (i × 33.75)2 + MT2) where Fseg is the force produced by each segment in the
three-segment model.
where Lfasnew is the new length of a fascicle after a
proportion of muscle shortening (i), where muscle 6. Ftotal = Fseg1 × a1 + Fseg2 × a2 + Fseg3 × a3
2
shortening has the value 1 to −1, and c = √( L fasopt + MT2).
33.75 refers to the possible muscle shortening distance where Ftotal is the total force of all three segments at
above and below optimum (33.75 mm), as described above. any given muscle length, Fseg1, Fseg2 and Fseg3 are the forces
produced by each of the three segments, and a1, a2 and
3. Ffas is the force produced by the fibre at a given fibre a3 are the relative contributions of each segment to
length, according to a parabolic force–length curve (over the total force based on their estimated physiological
the contraction range, Ffas ranged from approximately cross-sections (MT2/Lfasopt).
90 –100% of peak force).
© 2006 The Authors


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J. Anat. (2006) 209, pp289–310 doi: 10.1111/j.1469-7580.2006.00619.

Intra- and intermuscular variation in human quadriceps

femoris architecture assessed in vivo

Most notably, larger muscles typically have a large

© 2006 The Authors

© 2006 The Authors

Muscle thickness and fascicle angle

In vivo muscle architecture was examined using two-

© 2006 The Authors

calculated as the mean of the distances between super-

At most sites tested, the fascicles were too long to be

Fascicle length = sin(γ + 90°) × MT/

© 2006 The Authors

Fig. 3 Intramuscular architecture of the

for measuring muscle thickness and fascicle angle was

© 2006 The Authors

© 2006 The Authors

Equal Estimated Muscle thickness Fascicle

Significantly different from aproximal site and bmiddle site.

Homogeneity of within-muscle architecture

© 2006 The Authors

fore, for VL, VM and RF, subjects tended to have a

© 2006 The Authors

VL FA F = 52.8*** 4.3* 5.4* NS NS NS NS

Human quadriceps femoris architecture, A. J. Blazevich et al. 297

Equal ratio Volume PCSA Estimated contribution

VL FA 0.61*** 0.73*** 0.59** 0.63***

© 2006 The Authors

Table 6 Regression equations to predict

Variables were added step-wise by greatest contribution to explained variance; equations

equal ratio (r = 0.67), PCSA (r = 0.68), volume (r = 0.71)

© 2006 The Authors

© 2006 The Authors

© 2006 The Authors

© 2006 The Authors

© 2006 The Authors

© 2006 The Authors

© 2006 The Authors

© 2006 The Authors

Fig. 6 Relationships between whole

© 2006 The Authors

© 2006 The Authors

© 2006 The Authors

Equations: 4. θnew = sin−1(MT/Lfasnew)

© 2006 The Authors

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VL FA 0.61* 0.73* 0.59 0.63*