First published online August 25, 2003
Mechanical properties of rat soleus aponeurosis and tendon during variable recruitment in situ
Ryan J. Monti1,
Roland R. Roy2,*,
Hui Zhong2 and
V. R. Edgerton1,2
1 Department of Physiological Science, University of California Los Angeles,
Los Angeles, CA 90095-1761, USA
2 Brain Research Institute, University of California Los Angeles, Los
Angeles, CA 90095-1761, USA

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Fig. 1. (A) Captured video frame showing the typical spacing of the metal particles
along the length of the tendon and aponeurosis. The circled particle is
implanted in the tibia for use as a reference point to compensate for any limb
movement. (B) Representation of the extent of the aponeurosis as a reference
for the actual placement of the particles.
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Fig. 2. (A) The same captured video frame shown in
Fig. 1, indicating the level of
the histological section shown in B. The section is stained with hemotoxylin
and eosin and illustrates the placement and encapsulation of the metal
particle. Note that it is completely surrounded by connective tissue that is
physically continuous with the aponeurosis.
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Fig. 3. Representative force (A) and strain (B) patterns for a single particle pair
during a tetanic contraction. The mean force and strain from a series of video
frames (shaded region; 100 ms) were used to construct the force-strain
curves for every particle pair studied.
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Fig. 4. A representative force-strain curve for a single pair of particles (middle
aponeurosis; Lo). Each point (x) is from a maximal tetanic
contraction via a different combination of ventral root filament
bundles. At the lowest recruitment levels used in this study (15-20% of
Po), the strains were typically more than 50% of the
strains attained at Po. This implies the existence of a
highly compliant toe region. The straight line is a tangent to the
force-strain curve at 75% of Po, and its slope is the
stiffness of the tissue at that point. Po for this muscle
was 1.58 N. Thus, it can be seen from the relationship between this tangent
line and the force-strain curve that the stiffness of the tissue did not vary
much above 30% of Po (0.5 N for this muscle).
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Fig. 5. The relationship between peak strain (strain at Po) and
muscle-tendon unit (MTU) length for each of the five segments defined in this
study: DA, dorsal aponeurosis; MA, middle aponeurosis; PA, proximal
aponeurosis; PT/DA, proximal tendon/distal aponeurosis; TEND, tendon. Strains
in the tendon were significantly (P<0.05) less than those in all
regions of the aponeurosis and did not vary significantly with increasing MTU
length. By contrast, peak strains in the aponeurosis tended to increase with
lengthening of the MTU. Values are means ± 1 S.D.
(N=4).
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Fig. 6. The relationship between stiffness (N/% strain) and muscle-tendon unit
(MTU) length for each of the five segments defined in this study; for
definitions of abbreviations, see Fig.
5. Tendon stiffness was greater than that of all regions of the
aponeurosis and did not vary significantly with increasing MTU length. The
stiffness of the aponeurosis was relatively constant across lengths but was
significantly (P<0.05) higher at Lo+2 mm than
at any other length. Values are means ± 1 S.D.
(N=4).
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Fig. 7. Variations in stiffness among the five segments studied at
Po or 0.25 of Po. Stiffness declines
from the tendon to the aponeurosis, and the tendon was significantly
(P<0.05; indicated on graph) stiffer than all three regions that
did not include any overlap between tendon and aponeurosis. In addition, while
the general pattern holds for contractions at 0.25 of Po,
the tissue is much more compliant. TEND, tendon; PTDA, proximal tendon/distal
aponeurosis; DA, dorsal aponeurosis; MA, middle aponeurosis; PA, proximal
aponeurosis.
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Fig. 8. Hypothetical relationships between muscle fiber and connective tissue
recruitment. (A) All connective tissue is loaded at every recruitment level,
leading to a linear increase in connective tissue stress with an increasing
number of muscle fibers recruited. (B) The amount of connective tissue is a
constant function of recruitment, leading to a constant stress in the
connective tissue. (C) The amount of connective tissue recruited is
disproportionately high at low forces, with an increase as force rises. This
leads to a non-linear increase in connective tissue stress with recruitment.
For details, see text.
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© The Company of Biologists Ltd 2003