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Critical Reviews in Oral Biology & Medicine
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DOI: 10.1177/154411130201300406
CROBM 2002 13: 366
J.H. Koolstra
Dynamics of the Human Masticatory System
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366 Crit Rev Oral Biol Med 13(4):366-376 (2002)
Introduction
There is a large body of literature describing the dynamics of the
human musculoskeletal system (for a review, see, for instance,
Nigg and Herzog, 1999). In this large and mostly well-developed
field, the human masticatory system occupies a relatively small
place. One of the reasons for this underexposure is probably its relative
complexity, which makes it more difficult to analyze than, for
instance, the system of the shoulder, arm, hip, knee, or leg.
There are several reasons why masticatory dynamics are difficult
to analyze. First, the masticatory system consists of a large
number of muscles of various shapes and sizes, making it impossible
to determine, unambiguously, how they might cooperate to
perform a certain task (Wood, 1987; Lund, 1991; Hannam and
McMillan, 1994). Second, they have a complex architecture
(Schumacher, 1961; van Eijden et al., 1997), and their actions cannot
be determined from their overall orientation only (van der Helm
and Veenbaas, 1991). Third, the upper and lower jaws articulate
through two very complexly shaped incongruent temporomandibular
joints (Werner et al., 1991; Wish-Baratz et al., 1996). Any
simplification of these joints based on concepts usually used for
other joints (like hinges or balls-and-sockets) leads to considerable
loss of functionality (van Loon et al., 1999). Furthermore, the articular
surfaces are separated by a cartilaginous articular disc which
is able to move more or less freely between these surfaces (Bade et
al., 1994; Schmolke, 1994) and are influenced by, and affect, the
movements of the jaw (Rees, 1954; Isberg and Westesson, 1998).
Apart from the intrinsic complexity of the system, there are
several limitations to the collection of experimental data on
masticatory function. For instance, some of the masticatory
muscles run deep and are partially hidden behind bony structures,
which prevents easy access for electromyographic (EMG)
measurements (Wood et al., 1986; Koole et al., 1990; Murray et
al., 1999a). Furthermore, many jaw movements are relatively
small, posing stiff challenges to experimental systems designed
to record relevant properties adequately (Naeije et al., 1996).
In summary, there are many factors that impede assessment
of the mutual contributions of the relevant active and passive
structures to jaw movements. Recently, the application of biomechanical
models has provided an adequate experimental framework
to explore masticatory dynamics without several of the
drawbacks that accompany experiments with human subjects.
They are powerful tools for establishing causal relationships in
this field and have led to updates of or new formulations on various
insights into the function of the masticatory system.
Generally, monographs on jaw movement (Griffin and
Malor, 1974; Brown, 1975; Goodson and Johansen, 1975) have
been written from a more clinical perspective, and have provided
information mainly about the position, and positional
changes, of the lower jaw. The dynamic aspects and their consequences
were rarely taken into account. Recent developments in
these areas have provided new insights. The present review is
based upon a selection of studies on jaw movement analysis,
from both clinical and basic perspectives. Its purpose is to establish
an updated overview of the fundamentals of jaw movement,
and it focuses on the contributions of muscles and the influence
of passive constraints. It has been written from a biomechanical
perspective and with an emphasis on masticatory dynamics.
Since relevant topics such as neural control and feedback as well
as properties of masticatory motor units have been reviewed relatively
recently (Lund, 1991; Hannam and McMillan, 1994; van
Eijden and Turkawski, 2001), they will not be discussed here.
In section 1, the relevant anatomical properties of the
human masticatory system are reviewed briefly. They serve as
a basis for all aspects of jaw movement and its determinants.
Jaw movement is discussed in section 2, where its physics and
its properties are reviewed. Interactions among anatomical
structures and jaw movement properties are reviewed in section
3. An attempt is made to analyze the causal relationships
between the two components. Finally, relevant issues that have
not yet been resolved, but are assumed to be of critical importance
for jaw movement analysis, are discussed in section 4.
(1) The Human Masticatory System
The human masticatory system consists of a mandible which is
able to move in relationship to the skull and is guided by two
temporomandibular joints through contractions of the masticatory
muscles. To establish the contribution of each individual
structure to jaw movements, one must explore the construction
of the joints and the muscular system as well as the mechanical
consequences of this construction. The morphology of the
human masticatory system will be summarized very briefly.
While there is a large quantity of literature in this area, the list
of relevant citations in this article is far from complete.
DYNAMICS OF THE HUMAN MASTICATORY SYSTEM
J.H. Koolstra
Department of Functional Anatomy, Academic Centre for Dentistry
Amsterdam, Meibergdreef 15, 1105 AZ Amsterdam, the Netherlands;
j.h.koolstra@amc.uva.nl
ABSTRACT: In this review, the movement characteristics of the human
masticatory system are discussed from a biomechanical
perspective. The discussion is based upon the three fundamental laws of
mechanics applied to the various anatomical structures
that are part of the masticatory system. An analysis of the forces and
torques applied to the mandible by muscles, joints,
articular capsules, and teeth is used to assess the determinants of jaw
movement. The principle of relating the interplay of forces
to the center of gravity of the lower jaw, in contrast to a hinge axis near
its joints, is introduced. It is evident that the muscles
are the dominant determinants of jaw movement. The contributions of the
individual muscles to jaw movements can be derived
from the orientation of their lines of action with respect to the center of
gravity of the lower jaw. They cause the jaw to accelerate
with six degrees of freedom. The ratio between linear and angular
accelerations is subtly dependent on the mass and
moments of inertia of the jaw, and the structures that are more or less
rigidly attached to it. The effects of articular forces must
be taken into account, especially if the joints are loaded asymmetrically.
The muscles not only move the jaw but also maintain
articular stability during midline movements. Passive structures, such as
the ligaments, become dominant only when the jaw
reaches its movement boundaries. These ligaments are assumed to
prevent joint dislocation during non-midline movements.
Key words. Jaw movement, masticatory muscles, biomechanics.
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JOINTS
Morphology
The anatomy of the temporomandibular joint has been
described thoroughly (Rees, 1954). Mandibular movements
are guided by its articular surfaces (Brown, 1975; Williams et
al., 1995). These surfaces reside on the temporal bone of the
skull, involving an articular eminence and a mandibular
fossa, and on the roughly ovoid condylar head of the
mandible. They are both irregularly shaped (Wish-Baratz et
al., 1996), covered with fibrocartilage, and inaccessible for
direct measurements in vivo.
The articular surfaces are separated by a cartilaginous articular
disc with non-uniform thickness (Bade et al., 1994;
Schmolke, 1994). This disc is able to move together with the
mandibular condyle along the articular eminence while simultaneously
rotating on the condyle (Rees, 1954). Disc movements
generally run smoothly with respect to the articular surfaces.
The articular disc is connected superiorly to the temporal
bone and inferiorly to the mandible by relatively loose fibrous
structures. Together, these structures make up the articular capsule.
It is reinforced laterally by the temporomandibular ligament,
and is the only capsular structure that runs directly
between the temporal bone and the mandible (Schmolke, 1994).
Cadaver material reveals that the articular capsule is slack
(Rees, 1954; Schumacher, 1983). There are two accessory ligaments:
the sphenomandibular ligament, which runs medially
from the mandibular ramus; and the stylomandibular ligament,
which attaches to the mandibular angle from behind.
Mechanical consequences
The articular surfaces are highly incongruent, which means
that the shapes of the upper and lower surfaces differ considerably.
This allows for a large amount of motion at the cost of
a lessened joint stability and relatively small areas of joint contact.
The articular disc is supposed to reduce joint incongruency
and increase joint stability by enlarging the contact area
(Williams et al., 1995).
A second consequence of the incongruency of the joint, in
combination with the slackness of its capsule, is that the movements
in the joint are not restricted to rotations about more or
less fixed joint axes, as in classic joints (Andrews and Hay,
1983). The condyle and temporal bone can be regarded as two
separate bodies in space, usually held in appositional contact
when the jaw moves. As a consequence, the mandible may be
able to move with six degrees of freedom. Theoretically, it may
rotate about an axis through, for instance, its incisor point.
Therefore, the motion of this point per se bears no relationship
to condylar motion. This property is known as kinematic
redundancy. Furthermore, if the incisor moves from one point
to another its path is not necessarily defined a priori. It may
choose to move along a straight path or along a detour. In principle,
the number of possible paths is infinite.
MUSCULAR SYSTEM
Morphology
From a classic anatomical perspective, the masticatory muscles
are divided into elevator and depressor groups. The elevator
group consists of the masseter and temporalis muscles, which
are located more or less superficially, and the medial pterygoid
muscle, which is located more deeply. The muscles of the depressor
group are located in the floor of the mouth. This group consists
(from superior to inferior) of the geniohyoid, the mylohyoid,
and digastric muscles. The geniohyoid and mylohyoid muscles
connect the hyoid bone with the body of the mandible. The
digastric muscle connects the mastoid process of the skull with
the body of the mandible and is attached to the hyoid bone via a
fibrous loop which runs around its intermediate tendon. The lateral
pterygoid muscle completes the muscular system. It consists
of a superior and inferior head running from the mandibular
neck in forward and medial directions. Since both heads are considered
to have different actions, they cannot be regarded exclusively
as elevator or depressor (Juniper, 1981).
The elevator muscles are heavily pennate (Hannam and
McMillan, 1994; van Eijden et al., 1996, 1997). They have relatively
large physiological cross-sectional areas and are suitable
for the generation of large forces. The fibers are short, which
limits their capacity for active shortening during contraction.
The depressor muscles and the lateral pterygoid have more or
less parallel fibers and are therefore able to contract over a
longer distance with less force.
Mechanical consequences
The human masticatory system contains more muscles than are
apparently necessary to accomplish its tasks. This seems to be
unnecessary from a mechanical perspective, but it must be
noted that there are also spatial requirements to the construction
of the muscular system. For instance, a muscular system
that is mechanically optimal probably violates spatial requirements
with respect to the adjacent airway and alimentary tract.
The muscles can perform almost any task in various ways.
Although the system is able to generate cyclic movements controlled
by a central pattern generator (Lund, 1991; Ottenhoff et
al., 1993), its muscles cannot be lumped into a limited number
of alternating muscle groups. One of the reasons for this is that
they have to adapt constantly to the texture of the food between
the teeth (Thexton, 1992). The system is mechanically redundant,
which means that there is an infinite number of muscle
contraction patterns which can cause the same movement.
It has been demonstrated that various masticatory muscles
have the capacity to deploy regionally different portions
for different tasks (Møller, 1966; Wood, 1986; Miller, 1991;
Blanksma and van Eijden, 1995; Blanksma et al., 1997; Murray
et al., 1999b). Such functional heterogeneity, in combination
with a relatively large attachment area, may cause the direction
of the line of action of such a muscle to vary as well (van
der Helm and Veenbaas, 1991). While there is no a priori evidence
of co-activation between different muscle portions, the
system is capable of fine-tuning, to a large extent, the orientation
of the required muscle force (van Eijden et al., 1988) by
selective activation of motor units (for reviews, see Hannam
and McMillan, 1994; van Eijden and Turkawski, 2001). The relatively
large extensive nature of some muscles may also cause
spatially distant fibers within a muscle to shorten to various
degrees during mandibular movements (van Eijden and
Raadsheer, 1992; Hannam and McMillan, 1994; van Eijden et
al., 1996, 1997). This may cause shifts in muscle lines of action
which are not caused by the nervous system.
The depressors are directly or indirectly attached to the
hyoid bone. When this bone moves downward through action
of the infrahyoid muscles during wide jaw-opening (Muto and
Kanazawa, 1994), the jaw depressors are stretched, which, in
turn, lengthens their possibilities for active shortening. This
may help in obtaining wider jaw gapes.
(2) Jaw Movement Basics
DEGREES OF FREEDOM FOR JAW MOVEMENT
In three-dimensional space, a body able to move freely may perform
translations and/or rotations. This applies to the lower
jaw, although the degrees of the various movements are limited.
Translations can be performed along, and rotations about, three
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independent axes. The translation axes and the rotation axes are
not necessarily the same, but usually the three orthogonal axes
of some Cartesian systems aligned to anatomical planes are
used for this purpose. Translations can be described along axes
which are, for instance, anteroposterior or X, mediolateral or Y,
and supero-inferior or Z. Rotations can be defined by terms
such as azimuth (about the Z-axis), elevation (about the Y-axis),
and roll (about the X-axis) (Fig. 1) or yaw, pitch, and roll
(Baragar and Osborn, 1984). It must be noted that there are
many other conventions about sets of axes which are applicable.
Independent of the applied set of axes, every movement can be
expressed by a unique combination of the six independent fundamental
movements (which are known as the six degrees of
freedom of movement). The lower jaw does not move freely but
is guided by its joints. These structures, therefore, may reduce
the number of degrees of freedom (vide infra). Although translations
and rotations relative to any of the three independent axes
remain possible, they are no longer independent. For instance,
if the joints should restrict one degree of freedom, the movement
is completely determined by the other five.
If the movement of a point is studied—for example, the
incisor point of the lower jaw or a point representing the
condyle—it should be recognized that a point does not have an
orientation. Rotations about axes through this point are thus
meaningless, and it should be recognized that movements
accomplished by rotations about distant axes can also be performed
by translations. A point, therefore, can move with, at
most, three degrees of freedom. The consequence is that the
movement of any point on the jaw can be reconstructed from
the movement of the jaw, but not the other way around.
It is not easy to recognize functional aspects of jaw movement
from a combination of translations and rotations with
respect to pre-defined axes. An alternative way to describe a
movement is by a rotation about and a translation along a socalled
helical axis or screw axis (Kinzel et al., 1972). A movement
can be described by subsequent (six degrees of freedom)
small displacements. The orientation and location of the helical
axis related to such a displacement (Fig. 2) provide information
as to how it took place, while the moving body translates along
and rotates about this axis. Generally, the helical axis is not stationary
and may itself undergo displacements during movement.
Therefore, each instant of movement is connected to a
unique instantaneous helical axis. For sagittal movements, this
axis is directly analogous to the instantaneous center of rotation
for plane motion (Wu et al., 1988; Lindauer et al., 1995; Chen,
1998). It should be noted that the location and inclination of the
helical axis, and the amounts of rotation about and translation
along this axis, contain six independent variables according to
the six degrees of freedom for movement.
Instantaneous helical axes thus provide a complete
overview of jaw movement (Gallo et al., 1997, 2000; Koolstra and
van Eijden, 1997b; Chen and Katona, 1999; Gal et al., 2000). The
movement of teeth and condyles can be derived from them (Fig.
2). The relative contributions of rotations and translations of the
mandibular condyle, for instance, can be determined from the
distance to the axis. If, at a certain instant, the condylar movement
is characterized primarily by a rotation, the helical axis will
be situated close to the joint. If, in contrast, the translation component
is dominant, the helical axis will be located at a distant
location. These differences were demonstrated for jaw-opening
movements performed with different muscle recruitment patterns
showing no clear visual differences from normal movements
in terms of displacement of teeth and condyles, but great
differences in terms of helical axis positions (Koolstra and van
Eijden, 1997b). This emphasizes that, from a clinical perspective
as well, this approach can be relevant, for example, to an
enhanced possibility of discrimination among different translations
of the same type of movement by quantification of the inclination
and remoteness of the helical axis. Furthermore, it enables
on to discriminate between and among apparently similar
movements caused by different muscle contraction patterns.
PHYSICS OF JAW MOVEMENT: NEWTON’S LAWS
The dynamics of a moving lower jaw are expressed by its position,
its velocity, and its acceleration. According to the six degrees
of freedom for movement, each of these three properties also consists
of six independent variables. In a Cartesian system, the position
is not defined only by the (X, Y, and Z) position of the center
of gravity with respect to the origin of this system, but also by the
orientation (azimuth, elevation, and roll) of the jaw. The velocity
and the acceleration also have three linear and three angular
components.
For each of the six components, velocity is the (time)
derivative of position and acceleration the derivative of velocity.
Every moving body, including the lower jaw, obeys
Newton’s laws. Movements are caused by forces acting on the
jaw. They may be active muscle forces and also passive (reaction)
forces generated by joints, ligaments, and dental elements.
The forces also have six components. Each linear force (Fx, Fy,
Fz) is accompanied by a moment (angular) or torque (Mazimuth,
Melevation, Mroll). The resultant forces and torques generate
accelerations
according to Newton’s second law (acceleration equals
force divided by mass) (Fig. 1). This accounts for each degree of
freedom, emphasizing the fact that the mass of the jaw also consists
of three linear and three angular components. The three linear
components of the mass of the lower jaw are equal to the
common mass. The three angular masses (moments of inertia)
are dependent on the distribution of mass about the axis under
consideration and therefore on the shape of the lower jaw and
adhering structures. The moment of inertia about an axis is
defined as the sum of the mass of each particle multiplied by the
distance between this particle and the relevant axis to the power
of two (Nigg, 1999). For a lower jaw of about 0.44 kg, the
moments of inertia have been estimated as 8.6 kg.cm2, 2.9
kg.cm2, and 6.1 kg.cm2 for Iazimuth (about the z-axis), Ielevation
(about the y-axis), and Iroll (about the x-axis), respectively
(Koolstra and van Eijden, 1995). This means that it requires
about three times less muscle torque to accelerate the jaw for
open-close movements than for latero-deviations. The accelerations
cause changes in (linear and angular) velocity, and the
velocities cause changes in (linear and angular) jaw position.
Figure 1. Six degrees of freedom for jaw movement. Dashed lines:
principal axes. a: (linear) accelerations. F: (linear) forces. m: mass. a:
angular accelerations. M: torques. I: moments of inertia.
368 Crit Rev Oral Biol Med 13(4):366-376 (2002)
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INFLUENCE OF JOINTS
The degrees of freedom of articulating bones are often reduced
due to the construction of the connecting joint. An ideal balland-
socket joint, for instance, does not allow for translations.
Therefore, such a joint allows for movements with a maximum
of three degrees of freedom. In contrast, the degrees of freedom
in the temporomandibular joint are not reduced by its construction.
While its articular capsule is relatively slack and its articulating
surfaces incongruent, the mandibular condyle is able to
move almost freely in the three-dimensional half-space bounded
superiorly by the articular surface of the temporal bone.
Articular contact is not necessarily maintained, although the
distance along which the condyle is able to move perpendicular
to the articular surfaces is relatively short. Furthermore, the
articular surfaces are not rigid. The articular cartilage and the
articular disc are deformable such that the distance between the
bony surfaces will be proportional to the joint load (Huddleston
Slater et al., 1999). Consequently, the mandible is also able to
move with six degrees of freedom. If the joints are assumed to
maintain articular contact all the time, and the joint contact is
assumed to be rigid, a translation of the condyle in a direction
perpendicular to the articular surface of the temporal bone is
restricted, and the number of degrees of freedom for condylar
movement is reduced to five. Furthermore, if both joints are
assumed to be connected rigidly through the mandibular symphysis,
the rotation of the lower jaw about an antero-posterior
axis is restricted. In this (simplified) situation, it is able to move
with four degrees of freedom (Schumacher, 1961).
INFLUENCE OF MUSCLES
The jaw moves through contractions of the masticatory muscles.
Each muscle contraction is associated with a force which
is expressed by three independent variables: its magnitude, its
point of application, and its orientation. The latter two are
determined by the muscle’s line of action, defined by the
geometry of the system. Each muscle can produce a translation
of the lower jaw along its line of action, and a rotation about
an axis perpendicular to it and running through the jaw’s center
of gravity (Stern, 1974; Koolstra and van Eijden, 1995), as
illustrated in Fig. 3. The translation and rotation caused by a
muscle are not independent, and they express only one degree
of freedom. Therefore, if such a muscle is activated homogeneously,
the nervous system is able to influence only one
degree of freedom through the magnitude of its force. If the
muscle can be activated heterogeneously, and is represented
by more than one independent line of action, it can influence
more than one degree of freedom. Conversely, if separate muscles
or muscle portions cannot be activated independently,
then, together, they are able to influence only one degree of
freedom. A system of muscles, therefore, is represented by a
number of degrees of freedom equal to the number of independent
lines of action. The masticatory system contains at
least 20 muscle portions which may be activated independently
(Fig. 4). The number of degrees of freedom of the muscular
system, therefore, is considerably larger than the (maximum)
six degrees of freedom of jaw movement. This causes a
mechanical redundancy in the masticatory system.
(3) Determinants of Jaw Movement
Jaw movements caused by the masticatory muscles are guided
by passive structures; thus, both passive and active elements
generate forces and torques which accelerate the jaw (Fig. 5).
Because of the large number of these forces, their changes during
jaw movements, and their strong interdependency, it is difficult
to separate these influences.
ACTIVE ELEMENTS
Although passive structures in the masticatory system may act
as constraints for jaw movements and guide the mandible
along its path, active masticatory muscles are the prime
movers in this system. Therefore, it can be expected that they
will be the dominant determinants of jaw motion.
Muscle lines of action
While muscle lines of action differ considerably between muscles,
each contributes to masticatory movements in a unique
manner. Furthermore, the lines of action depend on the position
of the lower jaw with respect to the skull. This causes continuous
changes in the interplay of muscle forces and torques.
Investigators have estimated the lines of action of the masticatory
muscles, in vitro, by connecting the centers of the
attachment areas (centroids) on the skull and the mandible
(Baron and Debussy, 1979). However, functionally different
muscle portions that share attachment areas cannot be discriminated
with this approach. This drawback can be overcome by
measurement of the orientation of fiber bundles and incorporation
of the influence of tendinous sheets (van Eijden et al.,
1997). Furthermore, this method facilitates the estimation of
fiber length as a function of jaw position (van Eijden et al., 1996;
Koolstra and van Eijden, 1997b). The drawback of all in vitro
methods is that results are not necessarily applicable to individual
subjects. Therefore, muscle lines of action have been
estimated in vivo by a determination of putative muscle attachment
points (Goto et al., 1995, 2001) or by fitting the long axis
through “Magnetic Resonance Imaging” sections (Koolstra et
al., 1990). Although this enables individual characteristics to be
incorporated, the estimates of the lines of action remain coarse.
Generally, the contribution of a muscle to jaw movements
can be established by the direction of its line of action and the
position of this line with respect to the center of gravity of the
lower jaw. It accelerates the jaw in the direction of the line of
action according to: a = F/m, where a is the linear acceleration
vector, F the muscle force vector, and m the mass of the jaw. Also,
an angular acceleration about the center of gravity occurs
according to: a = M/I, where a is the angular acceleration vector,
M the muscle torque vector about the center of gravity, and
I the moment of inertia vector. The moment of inertia is dependent
on the related axis, whereas the mass is not (Koolstra and
van Eijden, 1995). The actual movement, then, is determined by
the resultant instantaneous linear and angular accelerations initiated
by the forces of all active and passive structures. For a single
muscle, the ratio between angular and linear acceleration is
proportional to the length of the moment arm of the muscle with
respect to the center of gravity. Furthermore, it is proportional to
the ratio between the mass of the lower jaw and its moment of
13(4):366-376 (2002) Crit Rev Oral Biol Med 369
Figure 2. Helical axis. (A) Rotations about and translations along a
hypothetical
helical axis during a non-midline jaw movement (dashed line). (B)
Subsequent helical axes during jaw closure (after Gallo et al., 2000).
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inertia relative to the axis of the muscle torque. These combined
factors determine the effect of muscle contraction and, consequently,
the contribution of each single muscle to jaw movement.
It must be emphasized that these equations represent simplified
dynamics. To avoid excessive complexity, the terms relating to,
for instance, inertial coupling, centripetal forces, and coriolis
forces have been neglected (Nigg and Herzog, 1999).
In a sagittal plane analysis, the lines of action of most jawclosers
are directed upward, and those of the jaw-openers,
downward and backward (Fig. 5). However, in both cases, each
line of action has a similarly directed moment with respect to
the sagittal axis through the center of gravity of the lower jaw.
Jaw-closers and -openers are able to produce a similarly directed
torque about this axis which leads to an angular acceleration
in the “negative elevation” (opening) direction. Consequently,
almost every muscle pair that is activated symmetrically
attempts, aside from its specific action, to perform an opening
rotation about the center of gravity. It is through this mechanism
that both jaw-closers and -openers, despite their difference
in orientation, are able to maintain articular contact while
performing unloaded (symmetrical) jaw movements.
Muscle dynamics
The optimum isometric force produced by a muscle Fopt is proportional
to its physiological cross-section S (in cm2) and its activation
A (in %), as denoted in the equation Fopt = 37 x S x A (Weijs
and Hillen, 1985). Due to the dynamic muscle properties (Fig. 6),
the instantaneous force of a concentrically contracting muscle is
less than Fopt, and an eccentrically contracting muscle may produce
an instantaneous force larger than Fopt. Due to these properties,
the forces produced by masticatory muscles may change constantly
during a movement, even though their activation levels
remain constant (Koolstra and van Eijden, 1997a). The forcelength
relationship quantifies the property that enables a muscle
to produce a force when its sarcomeres are not shortened below,
or elongated beyond, certain lengths, and this property has been
demonstrated to be an important limiting factor for masticatory
muscle force development. For instance, the limited amount of
maximum shortening of the fibers of the lateral pterygoid muscles
prevent protrusion of the jaw beyond its normal limits (Koolstra
and van Eijden, 1996). Furthermore, maximum jaw-opening is
limited by the maximum shortening of the jaw-openers, which is
counteracted by the passive forces of the elevators (Koolstra and
van Eijden, 1997b; Langenbach and Hannam, 1999). The latter
effect would be even more dramatic if the instantaneous center of
rotation remained close to the joint (Weijs et al., 1989).
Muscle force is also dependent on the shortening velocity
through the force-velocity relationship (Fig. 6). For jaw openclose
movements, where all jaw-opening or jaw-closing muscles
were activated simultaneously, it was demonstrated that the trajectory
of movement is not very dependent on the speed of
movement (Koolstra and van Eijden, 1997b). While this trajectory
depends on mutual muscle forces, the force-velocity relationship
does not considerably affect the mutual ratio of instantaneous
muscle forces. Consequently, the possibilities for force
production are affected similarly in all contributing muscles
(Koolstra and van Eijden, 1997a). In contrast, the force-velocity
relationship does appear to assist in deceleration of the lower
jaw after a sudden disappearance of resistance during forceful
biting (Slager et al., 1997). When such an event occurs, the force
of the closing muscles can disappear instantly through the suddenly
large shortening velocity, which may be considered profitable
when the teeth are near occlusion, and there is little time
to activate the jaw openers to decelerate the jaw.
Contribution of muscle action to jaw movements
Jaw movements are performed through co-contraction of various
muscles. Electromyographic (EMG) measurement of the masticatory
muscles during various jaw movement tasks (Carlsöö, 1952,
1956a,b; Møller, 1966; Wood, 1987), therefore, cannot be used to
establish the individual contributions of the various muscles to a
movement. For instance, it does not provide a means to decide
whether two or more active muscles assist each other to perform
a certain movement or work against each other to stabilize the
system. EMG measurement has been used to detect functional
heterogeneity in the activation of different muscle portions (e.g.,
Møller, 1966; Wood, 1986; Blanksma and van Eijden, 1990; Miller,
1991; Blanksma et al., 1992; Murray et al., 1999b). For example, it
has been shown that the temporalis muscle shows a gradual
heterogeneity
in the antero-posterior direction, and that the masseter
muscle can be subdivided functionally into a superficial, an anterior
deep, and a posterior deep portion.
EMG registrations of masticatory muscles correlate muscle
370 Crit Rev Oral Biol Med 13(4):366-376 (2002)
Figure 3. Force and torque generated by a muscle (arrow) with
respect to the center of gravity of the lower jaw.
Figure 4. Overview of the masticatory system. Ventro-lateral view.
Continuous lines: muscle lines of action. Cross-bar: muscle origin.
Circle: muscle insertion. MAS_S: superficial masseter. MAS_P: deep
masseter. MPT: medial pterygoid. TEM_A: anterior temporalis. TEM_P:
posterior temporalis. LPT_S: superior lateral pterygoid. LPT_I: inferior
lateral pterygoid. DIG: digastric. GEH: geniohyoid. MYH: mylohyoid.
Dots: position of centers of right and left condyle and incisor point.
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31, 2010
activity with jaw movements or static bite tasks. These correlations,
however, do not necessarily reflect causal relationships.
For causal relationships between muscle contraction and jaw
movement to be established, the influence of the passive constraints
must also be taken into account. Generally, this influence
can be simplified to a screw displacement axis defined by the
movement constraints in the joint (Andrews and Hay, 1983). The
function of a muscle can then be defined by its moment with
respect to this pre-determined axis. This concept was adopted by
Grant (1973) and applied to the instantaneous center of rotation
during jaw movements. Unfortunately, the method applies to
joints influenced by only one degree of freedom by any muscle.
Since the temporomandibular joint allows for movements with
at least four degrees of freedom, this kind of analysis is irrelevant
both for static situations (when the influence of the joint force is
omitted) and for dynamic situations (when torques are
expressed with respect to the center of gravity; Stern, 1974). In
contrast, masticatory muscle function is not dependent on the
location of the center of rotation. Conversely, the location of the
instantaneous center of rotation (or instantaneous helical axis for
non-midline movements) is dependent on the actions of the masticatory
muscles. For instance, it has been demonstrated that
jaw-open movements can be performed with different muscle
recruitment patterns. Despite the relatively similar appearance
of these movements, the trajectories of the instantaneous centers
of rotation are very different (Koolstra and van Eijden, 1997b). If
one wishes to estimate the work done by a muscle force during
a movement, then its position with respect to the instantaneous
helical axis is relevant (Gal et al., 2000).
A causal relationship between masticatory muscle contraction
and jaw movement can be demonstrated experimentally
only when muscles are activated independently. This is not an
option in a regular experimental setting, nor is it likely even
with intensive training, due to the central organization of motor
control in the jaw muscles (Lund, 1991). An effective method,
however, is registration of jaw movements evoked by electrical
stimulation of isolated muscles. This can be done with indwelling
electrodes (Zwijnenburg et al., 1996, 1999) or by electromagnetic
stimulation of selected portions of the motor cortex
(McMillan et al., 1998). The amplitude of such stimulation is necessarily
restricted, to protect the experimental subject. Thus, the
evoked jaw movements are small and require a very sensitive
jaw-tracking device to be recorded. When the masticatory muscles
are not fully relaxed, the other muscles may easily disturb
the measurements. Even if complete rest can be obtained (for
instance, in the unconscious state), the passive muscle forces
may be relatively large compared with the evoked force and
cannot be ignored. Consequently, direct muscle stimulation has
not yet convincingly demonstrated the contributions of individual
masticatory muscles to jaw movements.
Though direct experimental methods have failed to disclose
the full functional potential of the masticatory muscles,
biomechanical modeling approaches can help to develop reliable
predictions on this subject. Such models are always simplifications;
care must be taken that all relevant properties be
included, and correct assumptions regarding model behavior
must be made. In one model, for instance, it has been predicted
that the jaw-opening muscles have the tendency to dislocate the
temporomandibular joint (Osborn, 1993), while in another, it
has been suggested that they have the tendency to stabilize the
joint (Koolstra and van Eijden, 1997b). In the former model,
torques were computed not with respect to the center of gravity
(Stern, 1974) but with respect to the joint, thereby neglecting
the law of conservation of angular momentum (Nigg and
Herzog, 1999). In contrast, in the latter model, all relevant properties
of (Newtonian) rigid body dynamics were implemented,
strengthening the reliability of its predictions. According to
these rigid body dynamics, the temporomandibular ligaments
play an insignificant role during symmetrical jaw movements. It
was demonstrated that activation of all jaw-closing muscles
simultaneously leads to natural-looking jaw-closing movements,
including a condylar movement similar to that observed
experimentally (Koolstra and van Eijden, 1995). Conversely,
activation of all jaw-opening muscles resulted in normal jawopening
movements (Koolstra and van Eijden, 1997b). It is possible
that the varying instantaneous center of rotation can be
used as a means for the assessment of different muscle recruitment
patterns during apparently similar jaw movements.
With the exception of the anterior temporalis and the superficial
masseter, the principle that muscles, when activated unilaterally,
generate a translation along their line of action and a rotation
about the center of gravity has been confirmed. When their
torque about the vertical axis through the center of gravity is
considered, a contralateral latero-deviation would be expected,
but instead, an ipsilateral latero-deviation occurs (Zwijnenburg
et al., 1996, 1999). This apparent paradox can be resolved if one
takes into account that joint loads caused by muscle forces also
contribute to jaw movement (Fig. 7). Both the anterior temporalis
and superficial masseter tend to tilt the contralateral condyle
from the articular eminence because of a large ipsilateral joint
load. The ipsilateral joint reaction force results in an ipsilaterally
directed joint torque about the vertical axis through the center of
gravity (Koolstra and van Eijden, 1999), which overcomes the
muscle torque to cause an ipsilateral latero-deviation.
PASSIVE STRUCTURES
Passive structures contribute to jaw motion because they have
the ability to resist its movements along one or more degrees
13(4):366-376 (2002) Crit Rev Oral Biol Med 371
Figure 5. Forces acting on the lower jaw in the sagittal plane. Crosshairs:
center of gravity. Fclosers: mean force of the jaw-closing muscles.
Fopeners: mean force of jaw-opening muscles. Fjoint: joint force. Fbite:
bite force. a: moment arm of the different forces.
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31, 2010
of freedom. This resistance is expressed by the structure’s ability
to generate a reaction force and/or torque. Furthermore,
the jaw itself contributes to its movements through its inertial
properties. The ratio between linear and angular accelerations
effected by a muscle is subtly dependent on the mass and
moments of inertia of the jaw, and all structures that are more
or less rigidly attached to it. This attachment may include the
part of the masticatory muscles attached to the mandible, the
tongue, skin, and other soft tissues. It has been demonstrated,
however, that the influence of these inertial properties on the
final movement is small (Koolstra and van Eijden, 1995).
Articular surfaces
When the temporomandibular joints are loaded, reaction forces
are transferred to the mandibular condyles. The cartilaginous
structures in the joint have very low friction, and their deformation
is fairly independent on loading direction (Beek et al., 2000),
so it is probable that these reaction forces are directed perpendicular
to the contacting articular surfaces. In a sagittal plane,
they are directed inferiorly and pass posteriorly to the center of
gravity of the lower jaw. These forces, therefore, have the tendency
to lower the condyle and separate the articular surfaces
(Fig. 5). The joint reaction forces also apply torques with respect
to the center of gravity of the lower jaw. While the line of action
of the reaction forces passes posteriorly to this center, the joint
reaction torques lead to an angular acceleration about the sagittal
axis through the center of gravity, which is bound to produce
an elevation movement in the positive direction. Consequently,
the reaction forces attempt to perform a closing jaw rotation
about this axis (Koolstra and van Eijden, 1995).
The cartilaginous temporomandibular joint disc and the
cartilage lining of the articular surfaces are deformable, with a
finite and non-linear elasticity (Beek et al., 2001). This elasticity
causes the joint reaction force to be dependent on the deformation.
The more both bony articular surfaces move toward each
other, the more the cartilaginous structures between these surfaces
are compressed, and the larger the
magnitude of the reaction force.
Consequently, even if articular contact is
maintained during static and dynamic situations,
some movement perpendicular to the
articular surfaces occurs. This is supported
indirectly by the finding that the condylar
path during an unloaded opening lies superior
to the path during an unloaded closing
movement (Yatabe et al., 1997; Huddleston
Slater et al., 1999). The consequence of this
observation is that joint loading during
(unloaded) jaw opening is most likely larger
than that during unloaded jaw closing.
Furthermore, it supports the suggestion that
the joints are loaded by the torque of the jawopening
muscles (Koolstra and van Eijden,
1997b).
The movement range of the condyle is
not limited to any great extent by the articular
surfaces of the skull. Only when the
joints are compressed in an upward or
obliquely backward direction by manipulation
can the fossa restrain jaw movements in
the region of the condyles. The articular surfaces
do not restrict protrusive and mediolateral
translations or rotations about any of
the three axes. If the mandible performs a
latero-deviation, the contralateral condyle
has to move forward relative to the ipsilateral
one, due to the interconnection of both condyles. During
this movement, it is forced to move downward along the articular
eminence. Consequently, this movement includes not
only an azimuth rotation, but also a roll rotation (Fig. 8). While
these rotations are interdependent, they nevertheless imply
only one degree of freedom.
Articular capsule and ligaments
The bony parts of the temporomandibular joint are connected
by an articular capsule composed of relatively loose collagenous
fiber bundles organized in a more or less parallel fashion
(Schmolke, 1994). On the lateral side, it is reinforced by a
temporomandibular
ligament. The fibers are able to withstand
some stretching. They deform according to their (non-linear)
elastic properties (Woo et al., 1993) and, in doing so, generate
tensile forces. These forces may decelerate the attached
condyle when it moves away from the articular surface of the
temporal bone. A similar mechanical function can be attributed
to the accessory ligaments, though these are very thin
(Williams et al., 1995) and their function is probably negligible.
For quite a while, the temporomandibular ligaments have
been considered a dominant constraint for condylar movement
and, therefore, for jaw movement. This has been illustrated by
the construction of more or less fixed axes for mandibular rotations,
especially in the final phase of jaw closure, and the idea
has been applied widely in the areas of prosthetic dentistry
(Ramfjord and Ash, 1966) and temporomandibular dysfunction
rehabilitation (Crawford, 1999). Presently, there is a considerable
amount of evidence that does not support the presence of a
fixed hinge axis (Lindauer et al., 1995; Chen and Katona, 1999)
in normal functional movements. Another dominant role for the
temporomandibular ligaments has been proposed for symmetrical
jaw-opening movements. Here, it has been assumed that
the temporomandibular ligament is always taut and forces the
condyle to slide down the articular eminence (Osborn, 1993).
However, if the temporomandibular ligament is taut, it cannot
372 Crit Rev Oral Biol Med 13(4):366-376 (2002)
Figure 6. Dynamic muscle properties. Total force is the sum of the forces
produced by the
sarcomeres (Fsarcomeres) The active force (Factive) is dependent on the
activation through the
activation dynamics, the instantaneous sarcomere length, and contraction
velocity. The
parallel elastic force (Fpassive) is dependent on the instantaneous
sarcomere length.
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13(4):366-376 (2002) Crit Rev Oral Biol Med 373
allow for habitual latero-deviation movements unless there is a
mechanism for tightening the ligaments during midline movements
and for slackening them during non-midline movements.
It is very unlikely that such a mechanism exists. These considerations
favor a less dominant role for the temporomandibular
ligaments in controlling habitual jaw movements.
The temporomandibular ligament may play a dominant
role in preventing the mandibular condyle from moving
beyond the limits of the articular surface of the temporal bone.
It has been shown that the masticatory muscles could pull the
condyles a few mm beyond these limits if the temporomandibular
ligaments were not present (Koolstra and van
Eijden, 1999; Koolstra et al., 2001). Due to the interconnection
of left and right condyles through the mandibular symphysis,
the two ligaments are sufficient to limit both antero-posterior
and medio-lateral condylar movements, during protrusive
and retrusive mandibular movements, and during latero-deviations,
respectively. Furthermore, when the mandible reaches
these positions, the contracting masticatory muscles have
almost reached the length where they become insufficient for
force production (Koolstra and van Eijden, 1999). The forces
applied to the ligaments will then probably not be large.
Teeth and food
The direct influence of teeth and food on jaw movements is due
to the reaction forces occurring when the upper and lower teeth
come into direct contact with each other or with a bolus of food
in between. Through the interplay of muscle and joint forces,
these reaction forces will be predominantly directed downward
(Koolstra et al., 1988) and will be accompanied by an opening
torque with respect to the center of gravity (Fig. 5), thus causing
a joint loading. There is also an indirect influence, since the central
nervous system is able to detect forces on the dental elements
through mechanoreceptors in the periodontal ligament
(Lund, 1991). This system, therefore, is able to adapt muscle
activation as required by the presence of food. Furthermore, the
nervous system is able to react even faster through reflexes.
The reaction forces are due to the resistance to deformation
by the underlying structures. The dental elements are very hard,
and should they come into contact with each other with some
velocity, even a small deformation could result in a very large
reaction force. Fortunately, the dentition is connected to the
mandible via a deformable, collagenous periodontium. When a
tooth of the lower dentition comes into contact with its upper
neighbor with a certain velocity, it undergoes little deformation
but is pushed into its socket. Due to the elasticity of the periodontium,
a reaction force occurs which becomes larger as the
tooth is pushed further into its socket. This reaction force acts on
the mandible and causes deceleration until the movement stops.
If food is compressed between the teeth, the reaction force
increases more slowly, while the bolus itself is deformable.
The direct influence of the teeth on jaw movements is
reflected by the superior portion of the Posselt envelope of
incisal point motion, but the dentition can also have an indirect
influence on jaw movements. It has been demonstrated
that subjects with malocclusion have a more irregular chewing
pattern than normally found (Gibbs et al., 1971; Lewin, 1985).
It is not clear whether these aberrant patterns are due to tooth
contacts themselves or to external factors.
During mastication, food is compressed and/or fractured
between the dentition to reduce the particle size and facilitate
swallowing. This compression and fracturing take place in the
slow-closing phase of the masticatory cycle. It is apparent that the
movement in this phase will be dependent on the mechanical
properties of the food. For tough foods, the compression will be
slower than for soft foods. Notably, the muscles are able to generate
larger forces when contracting slowly. The peak velocity that
follows the fracturing of hard, brittle food is much greater than
that for soft food (Peyron et al., 1997). This peak velocity, however,
is less than might be expected, possibly due to a decrease in muscle
force as a consequence of the force-velocity relationship (Slager
et al., 1997). Also, the size of the food affects mandibular movements,
since the mandible has to be opened wider for larger pieces
of food to be chewed. In the frontal plane, it has been observed
that subjects chewing hard food tended to perform larger lateral
excursions than when chewing soft food (Lewin, 1985).
Impact loads on the dentition may have consequences for
the joints, since they transfer to the joints via the mandible. A
healthy periodontium partially absorbs impact loads, and thus,
it may prevent peak loads on the joints. This property does not
exist if the dentition is connected with the mandible through an
artificial implant. The mandible itself, however, is deformable
(van Eijden, 2000), so it is possible that the transfer of impact
loads of the teeth to the joints may be reduced by its elasticity.
At present, there are no quantitative data on this subject.
Muscles
When inactive, the masticatory muscles generate passive forces
which are dependent on the instantaneous length of their sarcomeres
(Epstein and Herzog, 1998). When the sarcomeres are
at or below optimum length, estimated at 2.73 μm (van Ruijven
and Weijs, 1990), they are negligible, but increase exponentially
if they are stretched beyond this length. Apart from these
passive forces, muscle stretch can, indirectly, cause reflexes,
because it is detected by muscle receptors (Lund, 1991).
The passive forces of the jaw-closing muscles are believed
to decelerate the jaw at the end of jaw opening during mastication
(Ostry and Flanagan, 1989) and become significant when
the jaw nears its maximum opening (Koolstra and van Eijden,
1997a). It has been proposed that they are a determinant of maximum
jaw opening (Langenbach and Hannam, 1999; Peck et al.,
2000; Koolstra et al., 2001). Mathematical models applied to the
study of the passive forces of the masticatory muscles have been
unable to open the jaw more than about 3 cm, whereas an opening
of 6 cm is frequently observed in vivo (Posselt, 1962; Brown,
1975). Therefore, the quantitative nature of these predictions is
disputable. Due to the proposed exponential relationship
between the passive muscle forces and their sarcomere lengths,
small errors in the constants that determine this relationship
may lead to relatively large errors in the projected passive
Figure 7. Schematic overview of the possible actions generated by force
Fm of the superficial masseter or anterior temporalis muscle viewed in
the horizontal plane. (A) Possible rotations: Rg, about center of gravity;
Rr, about a vertical axis behind the right joint for a latero-deviation to
the right; and Rl, about a vertical axis behind the left joint for a
laterodeviation
to the left. (B) Influence of joint forces: am, moment arm of
muscle force; Frj, right joint force; arj, moment arm of right joint force;
Fjl, left joint force; and ajl, moment arm of left joint force.
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31, 2010
374 Crit Rev Oral Biol Med 13(4):366-376 (2002)
forces. As long as there are no accurate quantitative data on the
relationship between sarcomere length and passive force of the
human masticatory muscles, this issue remains uncertain.
INTERPLAY OF PASSIVE AND ACTIVE STRUCTURES
The most dominant determinants for jaw movement are the
forces generated by active muscles. Passive forces may modulate
jaw movements, but become dominant as the jaw reaches
its movement boundaries. Axes of rotation of the jaw during
free jaw movements are not primarily related to passive structures
of the masticatory system, but are determined by muscle
use. Nevertheless, in clinical practice, axes of rotation which
were assumed to be connected to the temporomandibular joint
have been measured and applied successfully for the diagnosis
and treatment planning of masticatory dysfunction (Roth and
Williams, 1996). This indicates that conclusions drawn from
studies which were performed to search for practical solutions
to a clinical problem cannot be automatically extrapolated to
other problems in particular or to jaw movements in general.
The influence of the passive constraints appears to be more
dominant as jaw movement deviates from the midline.
Dynamic biomechanical analysis has demonstrated that the
masticatory muscles are capable of maintaining the integrity of
the masticatory system, in most cases, without the need for an
articular capsule with ligaments to maintain articular apposition
(Koolstra and van Eijden, 1997b). In contrast, they appear
to play a role in reducing the medio-lateral movements of the
mandibular condyle during latero-deviation (Koolstra and van
Eijden, 1999). If the joints are loaded asymmetrically, the influence
of their reaction forces on jaw movement has to be considered.
When a muscle is activated unilaterally, the condylar
reaction forces may produce a reverse movement compared
with the one expected from the muscle’s line of action (vide
ante). In practice, however, the muscles contract as groups
rather than in isolation. For both midline and non-midline jaw
movements, dynamic muscle properties should be taken into
account, since they limit the force-producing capacities of the
muscles, thereby restricting jaw movement possibilities.
(4) Final Remarks
Jaw movement analysis has evolved from early observation
(Ulrich, 1896; Bennett, 1908) to experiments designed to formulate
and validate or falsify testable hypotheses. In particular,
the availability of dynamic biomechanical modeling methods
has accelerated our understanding of jaw movements and
the masticatory system (Koolstra and van Eijden, 1995, 1997b,
1999; Hannam et al., 1997; Langenbach and Hannam, 1999;
Peck et al., 2000). It is now possible to predict the actions of the
different muscles in this complex system by applying
Newton’s laws. It has become clear that the masticatory muscles
not only control jaw movements, but also maintain the
physical integrity of the masticatory system. However, during
relatively large medio-lateral excursions, the muscles may fail
to keep the articular components in apposition, at which time
the articular ligaments may be presumed to perform this role.
Although these developments have improved our knowledge
of the working of the human masticatory system, the
persistence of some outdated theories is striking. A prominent
example is related to the articulation (Gysi, 1910), where there
is a need to find a simple, reliable method for describing jaw
movements near dental occlusion in anatomically different
patients. Often, researchers and clinicians have attempted to
use hinge axes to describe this movement. However, it has
been known, since the end of the 19th century, that such axes
are non-existent during habitual jaw movements (Ulrich,
1896). The concept of a hinge axis may have been revived by
the demonstration that rotary jaw movement near occlusion
can be accomplished through manipulation on cadaverous
material (Rees, 1954). This concept has survived in clinical
practice, where, despite a lack of a scientific basis (Mohl et al.,
1990; Lindauer et al., 1995), it has been applied successfully for
a long period (Roth and Williams, 1996).
While, for instance, with the use of dynamic biomechanical
models, hypotheses regarding muscle actions in the functioning
masticatory system are being validated (Koolstra and van
Eijden, 1999), joint load predictions have not yet been verified
satisfactorily. This is a consequence of the fact that direct measurement
of temporomandibular joint loading without disturbing
articular integrity has remained impossible. Unfortunately,
this parameter is considered as a major influence on the development
of wear and degeneration of the cartilaginous and bony
structures of this joint. Insight on temporomandibular joint loading
is, therefore, still limited to model predictions, and the reliability
of these predictions is directly related to the assumptions
and parameters built in such models. These include joint morphology
and the material properties of its deformable structures
that contribute to load distribution. Furthermore, muscle tensions
applied during joint loading are required. In each subject,
these parameters may be different, leading to the need for in vivo
measurement. Progress has been made in reconstructing the relevant
muscle lines of action (Koolstra et al., 1990, 1992) and bony
parts of the joints (Krebs et al., 1994) in vivo, but, to date, no reliable
method is available to create reconstructions of the cartilaginous
tissues in the joints. For a complete overview of the
applied muscle tensions to be acquired, their physiological crosssections,
architecture, and degree of activation must be estimated.
Physiological cross-sections have been estimated from
anatomic cross-sections (Weijs and Hillen, 1984; Koolstra and
van Eijden, 1992) and muscle activation from EMG recordings
(Carlsöö, 1952, 1956a,b; Møller, 1966; Lehr et al., 1971; Widmalm
et al., 1988; Murray et al., 1999a,b), but it is questionable whether
such methods can be applied routinely to all muscles involved.
Masticatory muscle architecture has been studied in vitro (van
Figure 8. Rotations about vertical and horizontal axes during a latero-
deviation movement.
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13(4):366-376 (2002) Crit Rev Oral Biol Med 375
Eijden et al., 1997), but the influence of individual variations on
model predictions and the possibility of applying relevant corrections
have not been established.
A start has been made on assessing the dynamic material
properties of the cartilaginous structures in the human
temporomandibular
joint (Beek et al., 2001), but the nature and
influence of individual variations are subject to speculation.
Consequently, although much is known qualitatively, quantification
of joint forces that incorporate individual variation still
cannot be performed unambiguously.
The ultimate limiting factor for reliable masticatory function
analysis incorporating biological variation is the lack of knowledge
about masticatory muscle recruitment patterns. The
mechanical redundancy of the masticatory system prevents their
unambiguous prediction (Koolstra and van Eijden, 2001). The
forces generated by the active muscles are the most dominant
determinants of jaw movement and joint loading. The search for
a rational way to predict muscle recruitment patterns remains a
dominant challenge in the field of jaw movement analysis.
Acknowledgments
This research was supported by the Interuniversity Research School of
Dentistry, through the Academic Center of Dentistry Amsterdam. I am
grateful
to Prof. T.M.G.J. van Eijden for his critical suggestions.
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