Dental Research
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Introduction
The force a muscle can produce depends on the length and
shortening velocity of the sarcomeres. The length changes to
which the sarcomeres are subjected during joint motion are
closely related to muscle architecture (Gans, 1982; Otten,
1988). Parameters of architecture-such as muscle fiber
position relative to the axis of rotation, pennation angle,
fiber length, sarcomere length, tendon length, and
physiological cross-sectional area-differ between muscles
(Wickiewicz et al., 1983; Friederich and Brand, 1990). In
addition, these parameters can vary within anatomically
complex muscles. For example, the anteriormost and
posteriormost regions of the human masseter have different
fiber and moment arm lengths (Van Eijden and Raadsheer,
1992). Hence, it is likely that the effect of joint motion upon
sarcomere length-and, as a consequence, upon the
magnitude of force and the velocity of shortening-differs
between and within muscles.
The human pterygoid muscles have a complex
architectural design. The lateral pterygoid is considered as
being fan-shaped with relatively long muscle fibers and the
medial pterygoid as being multipennate with short fibers
(Schumacher, 1961; Williams et al., 1989). In addition, the
lateral pterygoid is composed of two separate heads, which
are fused in front of the temporomandibular joint. Such a
division is less clear in the medial pterygoid, although its
anterior and posterior muscle fibers have distinct origins.
The action lines of the muscle heads differ in orientation
and position, suggesting a variety of functions in selective
activation (Van Eijden et al., 1988). The superior head of the
lateral pterygoid is believed to be activated during jaw
closing, whereas the inferior head is activated during
opening (Juniper, 1981; Wood et al., 1986). The medial
pterygoid is considered one of the main elevators of the
mandible.
The aim of the present study was to determine
quantitatively the structural characteristics of the pterygoid
muscles and to examine possible functional consequences.
1489
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1490
Age
F
F
F
M
F
M
F
M
56
69
56
89
55
82
83
74
12
10
8
10
5
11
8
5
Sarcomere lengths
From each cadaver, 16 samples (length, 5 mm; diameter, 2 mm),
eight per muscle and four per muscle head, were obtained. They
were taken halfway from each muscle belly, by means of a
sharp scalpel. The long axes of the samples were parallel to the
fiber direction of the muscle head and at equal craniocaudal
(inferior head, lateral pterygoid), mediolateral (superior head,
lateral pterygoid), and anteroposterior (medial pterygoid)
distances from each other. The samples were dehydrated in
alcohol and embedded in hydroxyethyl methacrylate
(Technovit). Three groups of four longitudinal sections (2 pm)
from each sample were stained with phosphotungstic acidhematein (PTAH); the distance between the groups was 200 pm
(i.e., 100 sections of 2 pm thickness). Fifteen randomly selected
rows, five per group, of ten sarcomeres were measured at a
magnification of xlOOO, by means of a Zeiss microscope with a
micrometer eyepiece (Weijs and Van der Wielen-Drent, 1982).
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Model simulations
For each muscle portion, average sarcomere length, fiber length,
and the three-dimensional attachment coordinates were
measured for the closed-jaw position. For all other mandibular
positions, sarcomere length and concomitant active muscle
force were calculated from the changed distance between origin
and insertion with respect to the closed-jaw position by means
of a three-dimensional rigid-body model (Weijs et al., 1987). The
muscle tendons were modeled as stiff structures. The active
length-force relationship that was incorporated into the model
was a third-degree equation (Van Ruijven and Weijs, 1990) and
was based on experimental data from rabbit digastric muscle
(Muhl et al., 1978). Optimum sarcomere length was assumed to
be 2.73 jpm, and active force ranged between 1.89 pm and 4.00
pm. Since the three-dimensional displacements of the insertions
of the muscle portions are not known for normal activities (such
as chewing), open\close excursions were chosen as rotations of
the jaw (Van Eijden and Raadsheer, 1992). The following
rotations, in 10 increments, were simulated: (1) about a
transverse axis situated 2.0 cm below the temporomandibular
joint, i.e., about halfway up the ramus of the mandible through
1491
Sarcomere Length
Sarcomere Length
3.00
2.80
2.60
2.40
2.20
2.00
ant
post
Fiber Length
Fiber Length
E~
f
-i
post
ant
Physiological Cross-section
Physiological Cross-section
E
'a
inf
sup
Lateral Pterygoid
'ezez,,,ant
Medial
,
post
Pterygoid
Statistics
For survey of differences in sarcomere length within each
muscle, one-way analysis of variance (ANOVA) was used.
Paired Student t tests were used for comparison between
different muscle heads. In all tests, the level of significance was
set to p < 0.05.
Results
Sarcomere lengths
In the eight regions of the lateral pterygoid, the average
sarcomere length for all subjects ranged from 2.68 to 2.90
jm. The difference in sarcomere length among the eight
regions of the muscle was not significant. Average
sarcomere lengths in the regions of the medial pterygoid
muscle ranged between 2.43 and 2.60 pm and did not differ
significantly from each other. Fig. 1 shows grand means of
sarcomere length of the muscle heads. For each subject,
average value of the 4 samples per muscle head was
calculated (n = 60, 4 samples x 15 fibers); the standard
deviation bars are a measure of interindividual variability.
Sarcomeres in the lateral pterygoid (inferior head, 2.83 pm;
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1492
Sarcomere Length
Mean
SD
Lateral pterygoid
inferior head
superior head
8.0%
7.1%
2.2%
1.9%
3.4%
3.0%
Medial pterygoid
anterior head
7.3%
0.8%
10.2%
5.6%
7.2%
1.2%
posterior head
13.4%
5.6%
a
Mean SD of the coefficient of variation of the eight muscles.
Mean
SDb
Lateral pterygoid
inferior head
length
sagittal angle
frontal angle
transverse angle
superior head
length
sagittal angle
frontal angle
transverse angle
32.6 mm
120.70
28.40
47.40
3.1 mm
11.70
8.40
7.30
31.3 mm
76.40
16.70
41.70
2.9 mm
7.80
11.50
8.00
Medial pterygoid
anterior head
length
sagittal angle
frontal angle
transverse angle
posterior head
length
sagittal angle
frontal angle
37.9 mm
33.80
31.30
42.10
6.0 mm
6.30
5.80
9.50
44.4 mm
5.5 mm
8.20
8.40
4.20
39.20
42.80
48.80
transverse angle
See text for definition of angles.
b Mean SD of the eight muscles.
a
Model simulations
During jaw opening, the distance between origin and
insertion became shorter in the lateral pterygoid and longer
in the medial pterygoid. This is illustrated in the top panels
of Fig. 2, where sarcomere length positions on the
normalized active length-force curve are shown for the
closed and open jaw, after a rotation about the ramus axis.
Note that the lateral pterygoid is mainly on the ascending
limb of the curve and the medial pterygoid mainly on the
descending limb. The amounts of sarcomere shortening in
the inferior and superior heads of the lateral pterygoid were
0.70 pm and 0.87 pm, respectively. In the anterior and
posterior heads of the medial pterygoid, the amounts of
sarcomere lengthening were 1.57 pm and 0.88 pm,
respectively. Differences in sarcomere excursion between
the heads of the muscles were primarily the result of
different moment-arm lengths relative to the axis of rotation.
In the middle and bottom panels of Fig. 2, the relationship
between jaw angle and active force is compared between
open/close rotation around the ramus axis and the joint
axis. The model predicted that, in the case of jaw opening
around the axis situated within the ramus, active force
decreased in both pterygoid muscles. Change of force was
relatively small in the inferior head of the lateral pterygoid
and in the posterior head of the medial pterygoid. When the
axis was located in the temporomandibular joint, the
moment-arm length of the medial pterygoids became
relatively large, while that of the lateral pterygoids was
negligible. Consequently, sarcomere excursions in the
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1493
Medial Pterygoid
Lateral Pterygoid
--O interior head
0.80
040 open
pen
0L0.40
0.200
0.20
clos
Xn
0.60
o.oo
1.50
2.00
Discussion
2.50
3.00
3.50
4.00
.,50
4.50
2.00
inferior head
1.00
2.50
3.00
3.50
4.00
4.50
A posterior head
cis
1.00
0.60
close
z2S
1.00 -
superior head
anterior head
posterior head
00
1)
.------------
o.so
ab0.60
LL
0.60/
0.40.
U-
0.20
0.40.\"
\
0 20
o.oo
"
o.oo
20
25
5
t0o
t5
open angle (c) ondylar axis
0.80
superior head
inferior head
(-
1.00
0.80
LL
L0. 40
0.20
0.00
o.oo
15
25
1o
20
open angle () ramus axis
.\
0.40
0.20
--posterior head
tD0.60
."
anterior head
30
:'
e 0.60
5
10o
25
t5
20
open angle () condylar axis
30
30
.
o
10
15
20
25
30
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1494
Acknowledgments
We are grateful to W.A. Weijs for his constructive criticism,
L. van Ruijven for help with various technical problems, and
H. Korfage for help with histological processing of the
muscle samples. Institutional funds supported this project.
References
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1495
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