THE ANATOMICAL RECORD 000:000–000 (2012)

The Arrangement of Fascicles in Whole

Muscle
BENJAMIN W. INFANTOLINO,1,2* THOMAS NEUBERGER,3,4
2
AND JOHN H. CHALLIS
1
Penn State Berks, Reading, Pennsylvania
2
Biomechanics Laboratory, Pennsylvania State University, Pennsylvania
3
Department of Bioengineering, Pennsylvania State University, Pennsylvania
4
The Huck Institutes of the Life Sciences, Pennsylvania State University, University Park, PA

ABSTRACT
The architecture of the muscle fascicles, here meaning their lengths
and their arrangement relative to one another, has important implica-
tions for the force a muscle can produce. Therefore, quantifying this
architectural arrangement and understanding the implications of the
architecture are important for understanding muscle function in vivo.
There were two purposes of this study: (1) to assess, via blunt dissection,
the number and the length of all the fascicles comprising the First Dorsal
Interosseous (FDI) muscle and (2) to visually identify, via magnetic reso-
nance imaging (MRI), the arrangement of the fascicles comprising the
FDI. Simple blunt dissection of all the fascicles comprising four FDI
muscles and their subsequent measurement demonstrated that the fas-
cicles comprising the whole muscle were not as long as the muscle belly
from which they were extracted. Muscle fascicles are surrounded by con-
nective tissue hence the paths of the fascicles in two whole FDI muscles
were identified via MRI by tracking the connective tissue surrounding
the fascicles. The fascicles had a spiral pattern along the length of each
muscle, within both muscles many of the fascicles were arranged in series
with other fascicles. These architectural features of the fascicles of the
FDI have important implications for the force–length and force–
velocity properties of the whole muscle. Anat Rec, 00:000–000,
2012. V C 2012 Wiley Periodicals, Inc.

Key words: first dorsal interosseous; muscle architecture;

fascicle

Two important characteristics of muscle are its force–
length and force–velocity properties. For a muscle fiber
both of these properties are in large part determined by
its length, which influences the range over which force
can be produced and the maximum velocity of shorten-
ing. For a whole muscle these properties depend on the
lengths of the fibers comprising the muscle and the
arrangement of the fibers relative to one another, and
relative to the muscle’s external tendon. The properties
of muscle fibers have received extensive study, for exam-
ple, the classic work of Gordon et al. (1966) on the force–
length properties, and that of A.V. Hill on the force–
velocity properties (Hill, 1938). The lengths and arrange-
ment of the fibers within whole muscle, and the
functional significance of this arrangement have
received less attention.
In 1878, August Froriep reported that the fibers com-
prising the belly of a parallel fibered muscle may not be
as long as the belly, providing evidence for complex ar-
chitectural arrangements of the fibers relative to one
another. There is evidence in cat muscle that the fibers
comprising a parallel fibered muscle are organized in
such a fashion that many of the fibers are shorter than
Additional Supporting Information may be found in the
online version of this article.
*Correspondence to: Benjamin W. Infantolino, Penn State
Berks, PO Box 7009, Tulpehocken Road, PA 19610. E-mail:
bwi100@psu.edu
Received 7 November 2011; Accepted 30 March 2012.
DOI 10.1002/ar.22484
Published online in Wiley Online Library
(wileyonlinelibrary.com).

V
C 2012 WILEY PERIODICALS, INC.
2 INFANTOLINO ET AL.
the muscle belly, the fibers do not always terminate at
the tendon, and some fibers appear to be arranged in se-
ries with other muscle fibers (Loeb et al., 1987). Fibers
terminating within a muscle belly can still transmit
force to the external tendon (Monti et al., 1999), and
fibers in series can transmit force between one another
(Trotter, 1993). In long human parallel fibered muscles,
the Sartorius and Gracilis, Heron and Richmond (1993)
found that these muscles were composed of relatively
short fibers arranged in series.
The architecture of the muscle fibers, here meaning
their lengths and their arrangement relative to one
another, has important implications for the force a mus-
cle can produce. Muscles fibers are bundled into
fascicles, and bundles of fascicles comprise the whole
muscle. Much recent focus has been on the behavior of
the fascicles in whole muscle, in part because current
technology permits their imaging (e.g., Rana and Wake-
ling, 2011; Sinha et al., 2011), and because the imaging
of fibers in whole muscle is not currently feasible. Fasci-
cle lengths are used as a surrogate for fiber length, and
while this assumption ignores some of the subtlety of
the fiber behavior it does give an indication of muscle
architecture and therefore the influence of architecture
on whole muscle function.
Current magnetic resonance imaging (MRI) data collec-
tion and processing methods have permitted the
identification of the fascicles within muscles. For example,
Kan et al. (2009) used diffusion tensor analysis of MRI
images to track the lines of action of the fascicles of the
quadriceps. Such an analysis provides information on the
line of action of the fascicles comprising a muscle, but
reported resolution values are not sufficient to discriminate
the orientation of one fascicle relative to another (Kan
et al., 2009; Sinha et al., 2011). In this current study, high-
resolution MRI was used to determine the location within a
whole muscle of the individual fascicles comprising the
muscle; such an approach is able to assess fascicle architec-
ture and if any of the fascicles are arranged in series.
The muscle of interest in this study was the First Dor-
sal Interosseous (FDI) muscle, a muscle solely responsible
for index finger abduction about the metacarpophalangeal
joint. This muscle has been studied in vivo as this is the
unusual instance of a single muscle being the cause of
motion about a joint. Previous in vivo studies have
exploited this feature of the FDI to investigate in vivo:
the influence of strength training on muscle properties
(e.g., Davies et al., 1985), tendon compliance (e.g., Cook
and McDonagh, 1996), and the order of recruitment of
motor units (e.g., Milner-Brown et al., 1973; Kornatz
et al., 2005). To compliment these in vivo studies there
have also been studies examining FDI muscle architec-
ture in cadavers (Brand et al., 1981; Jacobson et al.,
1992; Infantolino and Challis, 2010). It is feasible for a
muscle to have a much longer muscle belly than the
length of the fibers or fascicles comprising the muscle if
the muscle is pennated. Previous dissection of the fibers
and fascicles comprising cadaver FDIs by Infantolino and
Challis (2010) show that this muscle has a small amount
of pennation (10 ), and that even with this pennation
the fiber lengths are typically not sufficient to account for
the muscle belly length. These results indicate this mus-
cle may demonstrate complex architecture.
Two architectural features of the arrangement of mus-
cle fibers have been identified: the length of the fibers
(e.g., Loeb et al., 1987), and the arrangement of the
fibers relative to one another (e.g., Heron and Richmond,
1993). Although it is not feasible to measure these fea-
tures in whole muscle it is feasible to do so for muscle
fascicles (bundles of muscle fibers). Therefore, there
were two purposes of this study: (1) To assess, via blunt
dissection, the number and the length of all the fascicles
comprising the FDI muscle and (2) to visually identify,
via magnetic resonance imaging (MRI), the arrangement
of the fascicles comprising the FDI. Simple blunt dissec-
tion of all the fascicles comprising a muscle will provide
the lengths of the individual fascicles comprising a mus-
cle. The muscle fascicles are surrounded by connective
tissue which can be identified via MRI, therefore by
identifying the connective tissue surrounding the fas-
cicles it was feasible to examine the arrangement of the
fascicles within a muscle. It was hypothesized that the
fascicles comprising the FDI would demonstrate a com-
plex architecture indicated by a large range of fascicle
lengths, some shorter than the muscle belly, and that
some of the fascicles would be organized in a serial
fashion.
METHODS
The following sections describe the blunt dissection
procedure and measurement procedure for determining
the lengths of the fascicles comprising FDI muscles, and
MRI procedures and image analysis used for examining
the arrangement of the fascicles. All research with
cadaver materials was conducted following institution
approved ethical, safety, and biohazard standards.
Fascicle Dissection Procedure
Four FDI muscles were removed from two cadavers
using blunt dissection techniques described previously
(Infantolino and Challis, 2010). The cadaver characteris-
tics were: Cadaver 1: male, 92 years old, 178 cm in
height, cause of death – atherosclerotic heart disease;
Cadaver 2: female, 56 years old, 174 cm in height, cause
of death – hypothermia. The muscles had a normal
appearance, and had dimensions similar to those from
previous dissections (Infantolino and Challis, 2010).
Once the muscles were removed, they were soaked in a
20% nitric acid solution for approximately one half hour
to facilitate the dissection process by digesting some of
the connective tissue. Fascicles were dissected using
blunt dissection with the aid of a Cannon RE-350 video
visualizer (Cannon, New York). Each fascicle was placed
on a flat graduated surface and was imaged using the
video visualizer and the dissection was recorded with a
DVD recorder using the output of the video visualizer.
Still images of fascicles were taken from the dissection
videos. Subsequent analysis of the fascicle and fascicle
tendon dimensions were performed on the recorded fasci-
cle images.
All lengths were measured from the recorded images
using a custom MATLAB program (The MathWorks
Natick). The accuracy of these measurements was
assessed to be 60.5 mm based on the accuracy with
which known lengths could be made on a graticule slide.
Before acid digestion, the length of the whole muscle’s
externally visible tendon was measured, as was the
length of the muscle belly. After dissection, the fascicle
 FASCICLES IN MUSCLE
Fig. 1. Muscle fascicle image, from cadaver #2, on the graduated surface, Lfac is the fascicle length
and Lt is the fascicle tendon length. Grid spacing is 5 mm.
muscle belly length and the fascicle tendon lengths were
measured (Fig. 1). The fascicle tendons were portions of
the internal tendon that came free of the main aponeu-
rosis with gentle teasing with forceps.
MRI Procedure
Two FDI were removed from two additional cadavers.
The cadaver characteristics were: Cadaver 3: female, 76
years old, 152 cm in height, 67.4 kg in mass, cause of
death – end stage congestive heart failure; Cadaver 4:
male, 72 years old, 160 cm in height, 72.6 kg in mass,
cause of death – esophageal carcinoma. These muscles
also had a normal appearance, and had dimensions simi-
lar to those from previous dissections (Infantolino and
Challis, 2010). Each muscle was removed with the ten-
don intact using blunt dissection techniques.
The two FDIs were scanned using MRI to identify the
tubes of collagen, perimysium, within which the bundles
of muscle fibers comprising the fascicles ran. To reduce
the MRI scanning time the tissue was immersed in a
1.5% Magnevist (Bayer Health Care, Wayne, NJ) phos-
phor-buffered saline solution for 7 days. The achieved
short T1 (33 ms) and T2 (7 ms) times allowed for fast
imaging with a high contrast-to-noise ratio. To prevent
the tissue from drying out and to minimize magnetic
susceptibility artifacts during scanning the specimens
were surrounded by a flourinert liquid FC-43 (3M, St.
Paul, MN). All experiments were conducted using a ver-
tical wide bore 14.1 Tesla Varian MRI system (Varian,
Palo Alto, CA) with direct drive technology. A commer-
cially available millipede resonator (Varian) with an
inner diameter of 40 mm was used to acquire three-
dimensional spin echo images of the muscle tissue. A
standard imaging experiment with an isotropic resolu-
tion of 75 lm comprised a field of view of 45  20  20
mm3 and a matrix size of 600  268 (75% partial Fou-
rier: 201)  268. With 12 averages and a repetition time
of 75 ms (echo time 11.7 ms) the total scan time was
13.5 hr. MATLAB (The MathWorks, Natick, MA) was
used for post-processing. By zero-filling each direction by
a factor of two, the pixel resolution of the standard
imaging experiment was 37.5 lm isotropically.
SPIERS software was used to analyze the MRI images
(Sutton et al., 2001; Sutton et al., 2002). Connective tis-
sue was identified by assigning pixels, between user
defined intensity values, to a segment. Each segment
represents a different type of tissue. Intensity values
varied between 0 and 255. Each object of interest was
assigned to a mask by the operator. The boundary of the
3
muscle was assigned to a different mask by one trained
operator which allowed the entire muscle to be identified
without including any noise in the image outside of the
muscle. Noise was typically artifacts outside of the
image of the muscle. Two independent operators followed
the same procedure with the muscles and produced iden-
tical results. Three-dimensional objects were generated
by creating triangles from the pixels that were simulta-
neously assigned to a specific segment and mask (i.e.,
muscle tissue segment and muscle mask). This allowed
for rapid and accurate identification of objects as only
the pixels in the defined mask and connective tissue seg-
ment were considered to be part of that particular
connective tissue object. Pixels in the muscle mask that
were in the muscle tissue segment were not included in
the connective tissue object, thus reducing the time to
identify individual connective tissue objects.
The software identified all of the fascicles in a muscle.
Then the path of individual fascicles could be tracked
manually. The paths of 10 fascicles were tracked in each
muscle, with tracking continuing until the fascicle termi-
nated; if another fascicle was in series with the tracked
fascicle then that fascicle was also tracked. The collagen
tubes containing the fascicles were all tapered at both
ends. If a fascicle was assumed to be in series with
another fascicle the tapered end of one fascicle would be
in the same cross-sectional image with the tapered end
of another fascicle thereby demonstrating overlapping,
separate fascicles, not one longer fascicle. Both tapering
ends were in such close proximity, that they were clearly
different fascicles yet were capable of transmitting force
to each other (see Supporting Information video).
Evaluation of Hypothesis
To assess FDI fascicle architectural complexity, from
the dissection of the fascicles, the architecture of the
FDI was determined by visual inspection during the dis-
section process, and by the measurement of the length of
the muscle belly and length of the external tendon for
the whole muscle, and the lengths of the tendon and
fascicles comprising the muscle. From the MRI, the
arrangement of the fascicles, comprising each muscle,
relative to one another was quantified.
RESULTS
The results from the dissection are presented initially,
and then the imaging results.
4 INFANTOLINO ET AL.
Dissection Results
The number of fascicles in a muscle ranged from 61 to
101 fascicles per whole muscle. During dissection it was
observed that the fascicles did not always run from distal
to proximal tendon, that some fascicles were in series,
and that they were not the same length as the whole
muscle. Measurement of the lengths of the fascicles and
the tendon demonstrated that the fascicles were typically
much shorter than the muscle belly (Fig. 2). For example,
for muscle #1 the longest fascicle length was 40.0 mm
whereas the whole muscle belly length was 52.5 mm, the
shortest fascicle length was 8.0 mm. The Anderson–Dar-
ling test was used to formally test if the lengths of the
fascicles were normally distributed for each muscle,
which they were. Some of the fascicles had both proximal
and distal tendons, whereas others only had distinct dis-
tal or proximal tendons. Fascicles lacking tendon at one
Fig. 2. Fascicle lengths compared with whole muscle belly lengths
and fascicle tendon lengths compared with whole muscle tendon
lengths. The bars indicate the standard deviation of the respective
measure.
Fig. 3. Connective tissue in the FDI muscle from an oblique view for cadaver #4. Arrows indicate the
muscle line of action.
end, appeared during dissection to be attached to another
fascicle also lacking tendon at one end; these fascicles
were determined to be in series with one another.
MRI Results
In both FDI muscles, the connective tissue was
arranged in a spiraling fashion (Fig. 3). The connective
tissue made approximately a one quarter turn along the
length of the muscle (Fig. 4); see Supporting Information
video illustrating connective tissue alignment.
For only a portion of the 80 fascicles comprising the
FDI could the connective tissue surrounding each fascicle
be tracked. Of the 10 tracked fascicles in each FDI, these
fascicles did not extend from one end of the muscle to the
other as the connective tissue tubes did not run from one
end to the other of the muscle. These fascicles were
within the length ranges of the fascicles obtained during
dissection. In addition, the connective tissue tubes associ-
ated with the fascicles were seen to be arranged serially,
indicating that muscle fascicles were arranged in a serial
fashion (Fig. 5). Notice Sections V, VI, and VII, in Fig. 5,
are all arranged in series. Some sections of connective tis-
sue were long (III) whereas others were much shorter
(IV), which agreed with the observations of the dissected
fascicle length results. For cadaver #4 of the 10 tracked
fascicles only four terminated at the aponeurosis (Fig. 5
does not display two of these fascicles because they are
located on the other side of the aponeurosis). For cadaver
#3, three of the fascicles terminated at the aponeurosis,
two were in series with each other.
DISCUSSION
This study aimed to examine the complexity of the
architecture of fascicles of the FDI. Blunt dissection of
the muscle fascicles suggested some fascicles were
 FASCICLES IN MUSCLE 5

Fig. 4. Two MRI-derived images showing the connective tissue (teal) surround a fascicle spiraling
around the FDI aponeurosis (blue), for cadaver # 4.

Fig. 5. The central aponeurosis (large blue structure), and the fascicles tracked in the MR image from
their surrounding connective tissue, for Cadaver #4. Note the red (V), olive (VI), and African violet (VII) col-
ored connective tissue strands arranged in serial fashion in the center of the image.

arranged in series with others, and their lengths showed
a normal distribution. While the analysis of fascicle
lengths provided evidence of a lack of uniformity of fasci-
cle lengths the MRI analysis revealed a complex
arrangement of these fascicles. The fascicles spiraled
along the length of the FDI, with some of the fascicles
being arranged in series. These findings lend support to
the initial hypotheses that the fascicles comprising the
FDI would demonstrate a complex architecture. A com-
plex architecture was indicated by the large range of
fascicle lengths, by some of the fascicles being shorter
than the muscle belly, and by some of the fascicles being
organized in a serial fashion. The implications of these
findings will be discussed in the following two
paragraphs.
Dissection and measurement demonstrated that there
was variability in the lengths of the fascicles comprising
all analyzed FDI muscles, and these fascicles were not
as long as the muscle belly. If the fascicles were not as
long as the muscle belly this implies there must be some
overlapping structure of the fascicles to produce the
whole muscle. The operating range of the muscle belly
will therefore be a complex function of the lengths of its
fascicles and their arrangement relative to one another.
6 INFANTOLINO ET AL.
Indeed it is feasible that some of the fascicles, for a
given muscle belly length, will be operating on different
portions of the force–length curve. Clearly, the properties
of one fascicle cannot be used to infer the properties of all
the fascicles comprising a muscle, or of the whole muscle.
The MRI analysis of the FDI demonstrated that the
fascicles do not follow a straight line path in whole mus-
cle but have a spiral pattern, and some of the fascicles
are arranged in series with one another. This serial
arrangement has implications for both the force–length
and force–velocity properties of the whole muscle. By
having fascicles arranged in series they have the poten-
tial to function as one long fascicle, which would have a
greater range of lengths over which it can produce force,
and will have a greater maximum velocity of shortening
than either of the two component fascicles. These results
demonstrate that implying the properties of a whole
muscle from a limited sample of its fascicles can be mis-
leading if some of the fascicles are arranged in series.
In this study, muscle fascicles were used as a surro-
gate for the fibers, typical muscle fascicles contain
between 10 and 100 muscle fibers (Jones and Round,
1990). In reality the fiber architecture is probably more
complicated; hopefully improved technology will permit
the imaging of these structures in whole muscle to per-
mit further elucidation of architectural influences on
muscle function.
Although this study has been able to identify impor-
tant aspects of the architecture of a specific muscle, it
only provides a static snapshot of the muscle. In vivo
there will be interacting forces which will be dictated by
its dynamically changing architecture (Van Leeuwen
and Spoor, 1993). In vivo the changing architecture of a
muscle, the rat plantaris, has been measured but technical
restrictions mean only superficial fibers were monitored
(Savelberg et al., 2001). This study demonstrated helical
deformation of the muscle caused by transverse forces
between the fibers, presumably transferred via the interdi-
gitating connective tissues. Three-dimensional finite
element models (e.g., Yucesoy et al., 2002), offer the poten-
tial to examine the complex architectural interactions in
muscle with the static architecture from MRI providing
model parameters and initial conditions.
One limitation to this study was the sample size; four
FDI muscles were dissected and a further two were ana-
lyzed via MRI. These samples sizes are similar to those
in Loeb et al. (1987) and Heron and Richmond (1993),
and while it is not possible to generalize these results
the results do demonstrate important architectural
arrangements which can occur in the FDI muscle. The
muscles were obtained from older individuals, and so it
is feasible that the results may reflect architectural
changes that occur with aging. There is already evidence
that in old age human muscle cross-sectional area
decreases (Kent-Braun and Ng, 1999), muscle pennation
angle decreases (Kubo et al., 2003), and that there is a
reduction in the number of motor units within a muscle
(Galea, 1996). A useful extension of the current study
would be to analyze FDI muscles from a young sample,
and from additional older samples.
Another limitation to this study is the ability to gener-
alize the findings to other muscles in the body. A
previous dissection study in long human muscle, Heron
and Richmond (1993), has demonstrated fascicles seri-
ally arranged; other muscles should be examined to
elucidate their fascicle architecture. Currently, the MRI
techniques used in this study can only be used to inves-
tigate isolated cadaveric tissues as the bore of the MRI
is too small (40 mm) and the scan time too long (13.5 hr)
to be feasible to study live subjects. Identification of fas-
cicles paths was via the manual tracking of images from
MRI, thus meaning only a limited number could be
tracked. Improved tracking software may be able to
automatically identify and track the path of the fascicles
comprising a muscle, further enhancing our knowledge
of muscle structure and therefore function.
This study has offered insight into the complex archi-
tecture of the FDI muscle. Using blunt dissection, it was
found that individual fascicles are not representative of
the whole muscle. MRI demonstrated that some of the
fascicles were connected in series. These findings have
important implications for the function of the FDI, and
raise the question whether similar fascicle architectures
exist in other muscles.
ACKNOWLEDGEMENT
The authors acknowledge the contribution of Wade Shu-
maker for assistance with production of the video of the
muscle images and Brendan Casey and Michael Ellis for
their work with fascicle dissection.
LITERATURE CITED
Brand PW, Beach RB, Thompson DE. 1981. Relative tension and
potential excursion of muscles in the forearm and hand. J Hand
Surg 6:209–219.
Cook CS, McDonagh MJ. 1996. Measurement of muscle and tendon
stiffness in man. Eur J Appl Physiol O 72:380–382.
Davies CT, Dooley P, McDonagh MJ, White MJ. 1985. Adaptation of
mechanical properties of muscle to high force training in man.
J Physiol 365:277–284.
Froriep A. 1878. Ueber das sarcolemm und die muskelkerne. Archiv
fur Anatomie und Entwickelungsgeschichte 2:416–427.
Galea V. 1996. Changes in motor unit estimates with aging. J Clin
Neurophysiol 13:253–260.
Gordon AM, Huxley AF, Julian FJ. 1966. The variation in isometric
tension with sarcomere length in vertebrate muscle fibres.
J Physiol 184:170–192.
Heron MI, Richmond FJ. 1993. In-series fiber architecture in long
human muscles. Jf Morphol 216:35–45.
Hill AV. 1938. The heat of shortening and the dynamic constants of
muscle. Proc Roy Soc London 126:136–195.
Infantolino BW, Challis JH. 2010. Architectural properties of the
first dorsal interosseous. J Anat 216:463–469.
Jacobson MD, Raab R, Fazeli BM, Abrams RA, Botte MJ, Lieber
RL. 1992. Architectural design of the human intrinsic hand
muscles. J Hand Surg 17:804–809.
Jones DA, Round JM. 1990. Skeletal muscle in health and disease.
New York: St. Martin’s Press.
Kan JH, Heemskerk AM, Ding ZH, Gregory A, Menico G, Spindler
K, Damon BM. 2009. DTI-based muscle fiber tracking of the
quadriceps mechanism in lateral patellar dislocation. J Mag
Reson Imaging 29:663–670.
Kent-Braun JA, Ng AV. 1999. Specific strength and voluntary mus-
cle activation in young and elderly women and men. J Appl Phys-
iol 87:22–29.
Kornatz KW, Christou EA, Enoka RM. 2005. Practice reduces motor
unit discharge variability in a hand muscle and improves manual
dexterity in old adults. J Appl Physiol 98:2072–2080.
Kubo K, Kanehisa H, Azuma K, Ishizu M, Kuno SY, Okada M,
Fukunaga T. 2003. Muscle architectural characteristics in women
aged 20–79 years. Med Sci Sports Exer 35:39–44.
 FASCICLES IN MUSCLE
Loeb GE, Pratt CA, Chanaud CM, Richmond FJ. 1987. Distribution
and innervation of short, interdigitated muscle fibers in parallel-
fibered muscles of the cat hindlimb. J Morphol 191:1–15.
Milner-Brown HS, Stein RB, Yemm R. 1973. Changes in firing rate
of human motor units during linearly changing voluntary contrac-
tions. J Physiol 230:371–390.
Monti RJ, Roy RR, Hodgson JA, Edgerton VR. 1999. Transmission
of forces within mammalian skeletal muscles. J Biomech 32:
371–380.
Rana M, Wakeling JM. 2011. In vivo determination of 3D muscle
architecture of human muscle using free hand ultrasound. J Bio-
mech 44:2129–2135.
Savelberg HHCM, Willems PJB, Baan GC, Huijing PA. 2001. Defor-
mation and three-dimensional displacement of fibers in isometri-
cally contracting rat plantaris muscles. J Morphol 250:89–99.
Sinha U, Sinha S, Hodgson JA, Edgerton RV. 2011. Human soleus
muscle architecture at different ankle joint angles from magnetic
resonance diffusion tensor imaging. J Appl Physiol 110:807–819.
7
Sutton MD, Briggs DEG, Siveter DJ, Siveter DJ. 2001. Methodolo-
gies for the visualization and reconstruction of three-dimensional
fossils from the silurian herefordshire lagerst€ atte. Palaeontol
Electron 4:1–17.
Sutton MD, Briggs DEG, Siveter DJ, Siveter DJ, Orr PJ. 2002. The
arthropod Offacolus Aingi (Chelicerata) from the Silurian of Here-
fordshire, England: Computer based morphological reconstruc-
tions and phylogenetic affinities. Proc R Soc Lond Ser B-Biol Sci
269:1195–1203.
Trotter JA. 1993. Functional morphology of force transmission in
skeletal muscle. A brief review. Acta Anat 146:205–222.
Van Leeuwen JJL, Spoor CW. 1993. Modeling the pressure and
force equilibrium in unipennate muscles with in-line tendons.
Philos T Roy Soc B Biol Sci 342:321–333.
Yucesoy CA, Koopman B, Huijing PA, Grootenboer HJ. 2002. Three-
dimensional finite element modeling of skeletal muscle using a
two-domain approach: linked fiber-matrix mesh model. J Biomech
35:1253–1262.