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; [email protected]

ABSTRACT: In this review, the movement characteristics of the human masticatory system are discussed from a biomechani-
cal perspective. The discussion is based upon the three fundamental laws of mechanics applied to the various anatomical struc-
tures 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 acceler-
ate 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.

Introduction They are powerful tools for establishing causal relationships in

this field and have led to updates of or new formulations on var-
T here 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
ious insights into the function of the masticatory system.
Generally, monographs on jaw movement (Griffin and
field, the human masticatory system occupies a relatively small Malor, 1974; Brown, 1975; Goodson and Johansen, 1975) have
place. One of the reasons for this underexposure is probably its rel- been written from a more clinical perspective, and have provid-
ative complexity, which makes it more difficult to analyze than, for ed information mainly about the position, and positional
instance, the system of the shoulder, arm, hip, knee, or leg. changes, of the lower jaw. The dynamic aspects and their conse-
There are several reasons why masticatory dynamics are diffi- quences were rarely taken into account. Recent developments in
cult to analyze. First, the masticatory system consists of a large these areas have provided new insights. The present review is
number of muscles of various shapes and sizes, making it impos- based upon a selection of studies on jaw movement analysis,
sible to determine, unambiguously, how they might cooperate to from both clinical and basic perspectives. Its purpose is to estab-
lish an updated overview of the fundamentals of jaw movement,
perform a certain task (Wood, 1987; Lund, 1991; Hannam and
and it focuses on the contributions of muscles and the influence
McMillan, 1994). Second, they have a complex architecture
of passive constraints. It has been written from a biomechanical
(Schumacher, 1961; van Eijden et al., 1997), and their actions cannot
perspective and with an emphasis on masticatory dynamics.
be determined from their overall orientation only (van der Helm
Since relevant topics such as neural control and feedback as well
and Veenbaas, 1991). Third, the upper and lower jaws articulate
as properties of masticatory motor units have been reviewed rel-
through two very complexly shaped incongruent temporo- atively recently (Lund, 1991; Hannam and McMillan, 1994; van
mandibular joints (Werner et al., 1991; Wish-Baratz et al., 1996). Any Eijden and Turkawski, 2001), they will not be discussed here.
simplification of these joints based on concepts usually used for In section 1, the relevant anatomical properties of the
other joints (like hinges or balls-and-sockets) leads to considerable human masticatory system are reviewed briefly. They serve as
loss of functionality (van Loon et al., 1999). Furthermore, the artic- a basis for all aspects of jaw movement and its determinants.
ular surfaces are separated by a cartilaginous articular disc which Jaw movement is discussed in section 2, where its physics and
is able to move more or less freely between these surfaces (Bade et its properties are reviewed. Interactions among anatomical
al., 1994; Schmolke, 1994) and are influenced by, and affect, the structures and jaw movement properties are reviewed in sec-
movements of the jaw (Rees, 1954; Isberg and Westesson, 1998). tion 3. An attempt is made to analyze the causal relationships
Apart from the intrinsic complexity of the system, there are between the two components. Finally, relevant issues that have
several limitations to the collection of experimental data on not yet been resolved, but are assumed to be of critical impor-
masticatory function. For instance, some of the masticatory tance for jaw movement analysis, are discussed in section 4.
muscles run deep and are partially hidden behind bony struc-
tures, which prevents easy access for electromyographic (EMG) (1) The Human Masticatory System
measurements (Wood et al., 1986; Koole et al., 1990; Murray et The human masticatory system consists of a mandible which is
al., 1999a). Furthermore, many jaw movements are relatively able to move in relationship to the skull and is guided by two
small, posing stiff challenges to experimental systems designed temporomandibular joints through contractions of the mastica-
to record relevant properties adequately (Naeije et al., 1996). tory muscles. To establish the contribution of each individual
In summary, there are many factors that impede assessment structure to jaw movements, one must explore the construction
of the mutual contributions of the relevant active and passive of the joints and the muscular system as well as the mechanical
structures to jaw movements. Recently, the application of biome- consequences of this construction. The morphology of the
chanical models has provided an adequate experimental frame- human masticatory system will be summarized very briefly.
work to explore masticatory dynamics without several of the While there is a large quantity of literature in this area, the list
drawbacks that accompany experiments with human subjects. of relevant citations in this article is far from complete.

366 Crit Rev Oral Biol Med 13(4):366-376 (2002)

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 JOINTS the body of the mandible and is attached to the hyoid bone via a
Morphology fibrous loop which runs around its intermediate tendon. The lat-
eral pterygoid muscle completes the muscular system. It consists
The anatomy of the temporomandibular joint has been of a superior and inferior head running from the mandibular
described thoroughly (Rees, 1954). Mandibular movements neck in forward and medial directions. Since both heads are con-
are guided by its articular surfaces (Brown, 1975; Williams et sidered to have different actions, they cannot be regarded exclu-
al., 1995). These surfaces reside on the temporal bone of the
sively as elevator or depressor (Juniper, 1981).
skull, involving an articular eminence and a mandibular
The elevator muscles are heavily pennate (Hannam and
fossa, and on the roughly ovoid condylar head of the
McMillan, 1994; van Eijden et al., 1996, 1997). They have rela-
mandible. They are both irregularly shaped (Wish-Baratz et
al., 1996), covered with fibrocartilage, and inaccessible for tively large physiological cross-sectional areas and are suitable
direct measurements in vivo. for the generation of large forces. The fibers are short, which
The articular surfaces are separated by a cartilaginous artic- limits their capacity for active shortening during contraction.
ular disc with non-uniform thickness (Bade et al., 1994; The depressor muscles and the lateral pterygoid have more or
Schmolke, 1994). This disc is able to move together with the less parallel fibers and are therefore able to contract over a
mandibular condyle along the articular eminence while simul- longer distance with less force.
taneously rotating on the condyle (Rees, 1954). Disc movements Mechanical consequences
generally run smoothly with respect to the articular surfaces. The human masticatory system contains more muscles than are
The articular disc is connected superiorly to the temporal apparently necessary to accomplish its tasks. This seems to be
bone and inferiorly to the mandible by relatively loose fibrous unnecessary from a mechanical perspective, but it must be
structures. Together, these structures make up the articular cap- noted that there are also spatial requirements to the construc-
sule. It is reinforced laterally by the temporomandibular liga- tion of the muscular system. For instance, a muscular system
ment, and is the only capsular structure that runs directly that is mechanically optimal probably violates spatial require-
between the temporal bone and the mandible (Schmolke, 1994). ments with respect to the adjacent airway and alimentary tract.
Cadaver material reveals that the articular capsule is slack
The muscles can perform almost any task in various ways.
(Rees, 1954; Schumacher, 1983). There are two accessory liga-
Although the system is able to generate cyclic movements con-
ments: the sphenomandibular ligament, which runs medially
trolled by a central pattern generator (Lund, 1991; Ottenhoff et
from the mandibular ramus; and the stylomandibular liga-
al., 1993), its muscles cannot be lumped into a limited number
ment, which attaches to the mandibular angle from behind.
of alternating muscle groups. One of the reasons for this is that
Mechanical consequences they have to adapt constantly to the texture of the food between
The articular surfaces are highly incongruent, which means the teeth (Thexton, 1992). The system is mechanically redun-
that the shapes of the upper and lower surfaces differ consid- dant, which means that there is an infinite number of muscle
erably. This allows for a large amount of motion at the cost of contraction patterns which can cause the same movement.
a lessened joint stability and relatively small areas of joint con- It has been demonstrated that various masticatory mus-
tact. The articular disc is supposed to reduce joint incongruen- cles have the capacity to deploy regionally different portions
cy and increase joint stability by enlarging the contact area for different tasks (Møller, 1966; Wood, 1986; Miller, 1991;
(Williams et al., 1995). Blanksma and van Eijden, 1995; Blanksma et al., 1997; Murray
A second consequence of the incongruency of the joint, in et al., 1999b). Such functional heterogeneity, in combination
combination with the slackness of its capsule, is that the move- with a relatively large attachment area, may cause the direc-
ments in the joint are not restricted to rotations about more or tion of the line of action of such a muscle to vary as well (van
less fixed joint axes, as in classic joints (Andrews and Hay, der Helm and Veenbaas, 1991). While there is no a priori evi-
1983). The condyle and temporal bone can be regarded as two dence of co-activation between different muscle portions, the
separate bodies in space, usually held in appositional contact system is capable of fine-tuning, to a large extent, the orienta-
when the jaw moves. As a consequence, the mandible may be tion of the required muscle force (van Eijden et al., 1988) by
able to move with six degrees of freedom. Theoretically, it may selective activation of motor units (for reviews, see Hannam
rotate about an axis through, for instance, its incisor point. and McMillan, 1994; van Eijden and Turkawski, 2001). The rel-
Therefore, the motion of this point per se bears no relationship atively large extensive nature of some muscles may also cause
to condylar motion. This property is known as kinematic spatially distant fibers within a muscle to shorten to various
redundancy. Furthermore, if the incisor moves from one point degrees during mandibular movements (van Eijden and
to another its path is not necessarily defined a priori. It may Raadsheer, 1992; Hannam and McMillan, 1994; van Eijden et
choose to move along a straight path or along a detour. In prin- al., 1996, 1997). This may cause shifts in muscle lines of action
ciple, the number of possible paths is infinite. which are not caused by the nervous system.
The depressors are directly or indirectly attached to the
MUSCULAR SYSTEM hyoid bone. When this bone moves downward through action
Morphology of the infrahyoid muscles during wide jaw-opening (Muto and
From a classic anatomical perspective, the masticatory muscles Kanazawa, 1994), the jaw depressors are stretched, which, in
are divided into elevator and depressor groups. The elevator turn, lengthens their possibilities for active shortening. This
group consists of the masseter and temporalis muscles, which may help in obtaining wider jaw gapes.
are located more or less superficially, and the medial pterygoid (2) Jaw Movement Basics
muscle, which is located more deeply. The muscles of the depres-
sor group are located in the floor of the mouth. This group con- DEGREES OF FREEDOM FOR JAW MOVEMENT
sists (from superior to inferior) of the geniohyoid, the mylohy- In three-dimensional space, a body able to move freely may per-
oid, and digastric muscles. The geniohyoid and mylohyoid mus- form translations and/or rotations. This applies to the lower
cles connect the hyoid bone with the body of the mandible. The jaw, although the degrees of the various movements are limited.
digastric muscle connects the mastoid process of the skull with Translations can be performed along, and rotations about, three

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independent axes. The translation axes and the rotation axes are for plane motion (Wu et al., 1988; Lindauer et al., 1995; Chen,
not necessarily the same, but usually the three orthogonal axes 1998). It should be noted that the location and inclination of the
of some Cartesian systems aligned to anatomical planes are helical axis, and the amounts of rotation about and translation
used for this purpose. Translations can be described along axes along this axis, contain six independent variables according to
which are, for instance, anteroposterior or X, mediolateral or Y, the six degrees of freedom for movement.
and supero-inferior or Z. Rotations can be defined by terms Instantaneous helical axes thus provide a complete
such as azimuth (about the Z-axis), elevation (about the Y-axis), overview of jaw movement (Gallo et al., 1997, 2000; Koolstra and
and roll (about the X-axis) (Fig. 1) or yaw, pitch, and roll van Eijden, 1997b; Chen and Katona, 1999; Gal et al., 2000). The
(Baragar and Osborn, 1984). It must be noted that there are movement of teeth and condyles can be derived from them (Fig.
many other conventions about sets of axes which are applicable. 2). The relative contributions of rotations and translations of the
Independent of the applied set of axes, every movement can be mandibular condyle, for instance, can be determined from the
expressed by a unique combination of the six independent fun- distance to the axis. If, at a certain instant, the condylar move-
damental movements (which are known as the six degrees of ment is characterized primarily by a rotation, the helical axis will
freedom of movement). The lower jaw does not move freely but be situated close to the joint. If, in contrast, the translation com-
is guided by its joints. These structures, therefore, may reduce ponent is dominant, the helical axis will be located at a distant
the number of degrees of freedom (vide infra). Although transla- location. These differences were demonstrated for jaw-opening
tions and rotations relative to any of the three independent axes movements performed with different muscle recruitment pat-
remain possible, they are no longer independent. For instance, terns showing no clear visual differences from normal move-
if the joints should restrict one degree of freedom, the move- ments in terms of displacement of teeth and condyles, but great
ment is completely determined by the other five. differences in terms of helical axis positions (Koolstra and van
If the movement of a point is studied—for example, the Eijden, 1997b). This emphasizes that, from a clinical perspective
incisor point of the lower jaw or a point representing the as well, this approach can be relevant, for example, to an
condyle—it should be recognized that a point does not have an enhanced possibility of discrimination among different transla-
orientation. Rotations about axes through this point are thus tions of the same type of movement by quantification of the incli-
meaningless, and it should be recognized that movements nation and remoteness of the helical axis. Furthermore, it enables
accomplished by rotations about distant axes can also be per- on to discriminate between and among apparently similar
formed by translations. A point, therefore, can move with, at movements caused by different muscle contraction patterns.
most, three degrees of freedom. The consequence is that the PHYSICS OF JAW MOVEMENT: NEWTON’S LAWS
movement of any point on the jaw can be reconstructed from
the movement of the jaw, but not the other way around. The dynamics of a moving lower jaw are expressed by its posi-
It is not easy to recognize functional aspects of jaw move- tion, its velocity, and its acceleration. According to the six degrees
of freedom for movement, each of these three properties also con-
ment from a combination of translations and rotations with
sists of six independent variables. In a Cartesian system, the posi-
respect to pre-defined axes. An alternative way to describe a
tion is not defined only by the (X, Y, and Z) position of the center
movement is by a rotation about and a translation along a so-
of gravity with respect to the origin of this system, but also by the
called helical axis or screw axis (Kinzel et al., 1972). A move-
orientation (azimuth, elevation, and roll) of the jaw. The velocity
ment can be described by subsequent (six degrees of freedom)
and the acceleration also have three linear and three angular com-
small displacements. The orientation and location of the helical ponents. For each of the six components, velocity is the (time)
axis related to such a displacement (Fig. 2) provide information derivative of position and acceleration the derivative of velocity.
as to how it took place, while the moving body translates along Every moving body, including the lower jaw, obeys
and rotates about this axis. Generally, the helical axis is not sta- Newton’s laws. Movements are caused by forces acting on the
tionary and may itself undergo displacements during move- jaw. They may be active muscle forces and also passive (reac-
ment. Therefore, each instant of movement is connected to a tion) forces generated by joints, ligaments, and dental elements.
unique instantaneous helical axis. For sagittal movements, this The forces also have six components. Each linear force (Fx, Fy,
axis is directly analogous to the instantaneous center of rotation Fz) is accompanied by a moment (angular) or torque (Mazimuth,
Melevation, Mroll). The resultant forces and torques generate accel-
erations 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 con-
sists of three linear and three angular components. The three lin-
ear 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
Figure 1. Six degrees of freedom for jaw movement. Dashed lines: open-close movements than for latero-deviations. The accelera-
principal axes. a: (linear) accelerations. F: (linear) forces. m: mass. a: tions cause changes in (linear and angular) velocity, and the
angular accelerations. M: torques. I: moments of inertia. velocities cause changes in (linear and angular) jaw position.

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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 ball-
and-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 construc-
tion. While its articular capsule is relatively slack and its articu-
lating surfaces incongruent, the mandibular condyle is able to
move almost freely in the three-dimensional half-space bound-
ed superiorly by the articular surface of the temporal bone.
Articular contact is not necessarily maintained, although the Figure 2. Helical axis. (A) Rotations about and translations along a hypo-
distance along which the condyle is able to move perpendicular thetical helical axis during a non-midline jaw movement (dashed line). (B)
to the articular surfaces is relatively short. Furthermore, the Subsequent helical axes during jaw closure (after Gallo et al., 2000).
articular surfaces are not rigid. The articular cartilage and the
articular disc are deformable such that the distance between the ACTIVE ELEMENTS
bony surfaces will be proportional to the joint load (Huddleston
Slater et al., 1999). Consequently, the mandible is also able to Although passive structures in the masticatory system may act
move with six degrees of freedom. If the joints are assumed to as constraints for jaw movements and guide the mandible
maintain articular contact all the time, and the joint contact is along its path, active masticatory muscles are the prime
assumed to be rigid, a translation of the condyle in a direction movers in this system. Therefore, it can be expected that they
perpendicular to the articular surface of the temporal bone is will be the dominant determinants of jaw motion.
restricted, and the number of degrees of freedom for condylar Muscle lines of action
movement is reduced to five. Furthermore, if both joints are
assumed to be connected rigidly through the mandibular sym- While muscle lines of action differ considerably between mus-
physis, the rotation of the lower jaw about an antero-posterior cles, each contributes to masticatory movements in a unique
axis is restricted. In this (simplified) situation, it is able to move manner. Furthermore, the lines of action depend on the posi-
with four degrees of freedom (Schumacher, 1961). tion of the lower jaw with respect to the skull. This causes con-
tinuous changes in the interplay of muscle forces and torques.
INFLUENCE OF MUSCLES Investigators have estimated the lines of action of the mas-
The jaw moves through contractions of the masticatory mus- ticatory muscles, in vitro, by connecting the centers of the
cles. Each muscle contraction is associated with a force which attachment areas (centroids) on the skull and the mandible
is expressed by three independent variables: its magnitude, its (Baron and Debussy, 1979). However, functionally different
point of application, and its orientation. The latter two are muscle portions that share attachment areas cannot be discrim-
determined by the muscle’s line of action, defined by the inated with this approach. This drawback can be overcome by
geometry of the system. Each muscle can produce a translation measurement of the orientation of fiber bundles and incorpo-
of the lower jaw along its line of action, and a rotation about ration of the influence of tendinous sheets (van Eijden et al.,
an axis perpendicular to it and running through the jaw’s cen- 1997). Furthermore, this method facilitates the estimation of
ter of gravity (Stern, 1974; Koolstra and van Eijden, 1995), as fiber length as a function of jaw position (van Eijden et al., 1996;
illustrated in Fig. 3. The translation and rotation caused by a Koolstra and van Eijden, 1997b). The drawback of all in vitro
muscle are not independent, and they express only one degree methods is that results are not necessarily applicable to indi-
of freedom. Therefore, if such a muscle is activated homoge- vidual subjects. Therefore, muscle lines of action have been
neously, the nervous system is able to influence only one estimated in vivo by a determination of putative muscle attach-
degree of freedom through the magnitude of its force. If the ment points (Goto et al., 1995, 2001) or by fitting the long axis
muscle can be activated heterogeneously, and is represented through “Magnetic Resonance Imaging” sections (Koolstra et
by more than one independent line of action, it can influence al., 1990). Although this enables individual characteristics to be
more than one degree of freedom. Conversely, if separate mus- incorporated, the estimates of the lines of action remain coarse.
cles or muscle portions cannot be activated independently, Generally, the contribution of a muscle to jaw movements
then, together, they are able to influence only one degree of can be established by the direction of its line of action and the
freedom. A system of muscles, therefore, is represented by a position of this line with respect to the center of gravity of the
number of degrees of freedom equal to the number of inde- lower jaw. It accelerates the jaw in the direction of the line of
pendent lines of action. The masticatory system contains at action according to: a = F/m, where a is the linear acceleration
least 20 muscle portions which may be activated independent- vector, F the muscle force vector, and m the mass of the jaw. Also,
ly (Fig. 4). The number of degrees of freedom of the muscular an angular acceleration about the center of gravity occurs
system, therefore, is considerably larger than the (maximum) according to: a = M/I, where a is the angular acceleration vec-
six degrees of freedom of jaw movement. This causes a tor, M the muscle torque vector about the center of gravity, and
mechanical redundancy in the masticatory system. I the moment of inertia vector. The moment of inertia is depen-
dent on the related axis, whereas the mass is not (Koolstra and
(3) Determinants of Jaw Movement van Eijden, 1995). The actual movement, then, is determined by
Jaw movements caused by the masticatory muscles are guided the resultant instantaneous linear and angular accelerations ini-
by passive structures; thus, both passive and active elements tiated by the forces of all active and passive structures. For a sin-
generate forces and torques which accelerate the jaw (Fig. 5). gle muscle, the ratio between angular and linear acceleration is
Because of the large number of these forces, their changes dur- proportional to the length of the moment arm of the muscle with
ing jaw movements, and their strong interdependency, it is dif- respect to the center of gravity. Furthermore, it is proportional to
ficult to separate these influences. the ratio between the mass of the lower jaw and its moment of

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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.
inertia relative to the axis of the muscle torque. These combined Continuous lines: muscle lines of action. Cross-bar: muscle origin.
factors determine the effect of muscle contraction and, conse- Circle: muscle insertion. MAS_S: superficial masseter. MAS_P: deep
quently, the contribution of each single muscle to jaw movement. masseter. MPT: medial pterygoid. TEM_A: anterior temporalis. TEM_P:
It must be emphasized that these equations represent simplified posterior temporalis. LPT_S: superior lateral pterygoid. LPT_I: inferior
dynamics. To avoid excessive complexity, the terms relating to, lateral pterygoid. DIG: digastric. GEH: geniohyoid. MYH: mylohyoid.
for instance, inertial coupling, centripetal forces, and coriolis Dots: position of centers of right and left condyle and incisor point.
forces have been neglected (Nigg and Herzog, 1999).
In a sagittal plane analysis, the lines of action of most jaw- rotation remained close to the joint (Weijs et al., 1989).
closers are directed upward, and those of the jaw-openers, Muscle force is also dependent on the shortening velocity
downward and backward (Fig. 5). However, in both cases, each through the force-velocity relationship (Fig. 6). For jaw open-
line of action has a similarly directed moment with respect to close movements, where all jaw-opening or jaw-closing muscles
the sagittal axis through the center of gravity of the lower jaw. were activated simultaneously, it was demonstrated that the tra-
Jaw-closers and -openers are able to produce a similarly direct- jectory of movement is not very dependent on the speed of
ed torque about this axis which leads to an angular acceleration movement (Koolstra and van Eijden, 1997b). While this trajecto-
in the “negative elevation” (opening) direction. Consequently, ry depends on mutual muscle forces, the force-velocity relation-
almost every muscle pair that is activated symmetrically ship does not considerably affect the mutual ratio of instanta-
attempts, aside from its specific action, to perform an opening neous muscle forces. Consequently, the possibilities for force
rotation about the center of gravity. It is through this mecha- production are affected similarly in all contributing muscles
nism that both jaw-closers and -openers, despite their differ- (Koolstra and van Eijden, 1997a). In contrast, the force-velocity
ence in orientation, are able to maintain articular contact while relationship does appear to assist in deceleration of the lower
performing unloaded (symmetrical) jaw movements. jaw after a sudden disappearance of resistance during forceful
biting (Slager et al., 1997). When such an event occurs, the force
Muscle dynamics of the closing muscles can disappear instantly through the sud-
The optimum isometric force produced by a muscle Fopt is pro- denly large shortening velocity, which may be considered prof-
portional to its physiological cross-section S (in cm2) and its acti- itable when the teeth are near occlusion, and there is little time
vation A (in %), as denoted in the equation Fopt = 37 x S x A (Weijs to activate the jaw openers to decelerate the jaw.
and Hillen, 1985). Due to the dynamic muscle properties (Fig. 6),
the instantaneous force of a concentrically contracting muscle is Contribution of muscle action to jaw movements
less than Fopt, and an eccentrically contracting muscle may pro- Jaw movements are performed through co-contraction of various
duce an instantaneous force larger than Fopt. Due to these proper- muscles. Electromyographic (EMG) measurement of the mastica-
ties, the forces produced by masticatory muscles may change con- tory muscles during various jaw movement tasks (Carlsöö, 1952,
stantly during a movement, even though their activation levels 1956a,b; Møller, 1966; Wood, 1987), therefore, cannot be used to
remain constant (Koolstra and van Eijden, 1997a). The force- establish the individual contributions of the various muscles to a
length relationship quantifies the property that enables a muscle movement. For instance, it does not provide a means to decide
to produce a force when its sarcomeres are not shortened below, whether two or more active muscles assist each other to perform
or elongated beyond, certain lengths, and this property has been a certain movement or work against each other to stabilize the
demonstrated to be an important limiting factor for masticatory system. EMG measurement has been used to detect functional
muscle force development. For instance, the limited amount of heterogeneity in the activation of different muscle portions (e.g.,
maximum shortening of the fibers of the lateral pterygoid muscles Møller, 1966; Wood, 1986; Blanksma and van Eijden, 1990; Miller,
prevent protrusion of the jaw beyond its normal limits (Koolstra 1991; Blanksma et al., 1992; Murray et al., 1999b). For example, it
and van Eijden, 1996). Furthermore, maximum jaw-opening is has been shown that the temporalis muscle shows a gradual het-
limited by the maximum shortening of the jaw-openers, which is erogeneity in the antero-posterior direction, and that the masseter
counteracted by the passive forces of the elevators (Koolstra and muscle can be subdivided functionally into a superficial, an ante-
van Eijden, 1997b; Langenbach and Hannam, 1999). The latter rior deep, and a posterior deep portion.
effect would be even more dramatic if the instantaneous center of EMG registrations of masticatory muscles correlate muscle

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activity with jaw movements or static bite tasks. These correla-
tions, however, do not necessarily reflect causal relationships.
For causal relationships between muscle contraction and jaw
movement to be established, the influence of the passive con-
straints 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 mas-
ticatory 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 Figure 5. Forces acting on the lower jaw in the sagittal plane. Cross-
helical axis is relevant (Gal et al., 2000). hairs: center of gravity. Fclosers: mean force of the jaw-closing muscles.
A causal relationship between masticatory muscle contrac- Fopeners: mean force of jaw-opening muscles. Fjoint: joint force. Fbite:
tion and jaw movement can be demonstrated experimentally bite force. a: moment arm of the different forces.
only when muscles are activated independently. This is not an
option in a regular experimental setting, nor is it likely even erties of (Newtonian) rigid body dynamics were implemented,
with intensive training, due to the central organization of motor strengthening the reliability of its predictions. According to
control in the jaw muscles (Lund, 1991). An effective method, these rigid body dynamics, the temporomandibular ligaments
however, is registration of jaw movements evoked by electrical play an insignificant role during symmetrical jaw movements. It
stimulation of isolated muscles. This can be done with in- was demonstrated that activation of all jaw-closing muscles
dwelling electrodes (Zwijnenburg et al., 1996, 1999) or by elec- simultaneously leads to natural-looking jaw-closing move-
tromagnetic stimulation of selected portions of the motor cortex ments, including a condylar movement similar to that observed
(McMillan et al., 1998). The amplitude of such stimulation is nec- experimentally (Koolstra and van Eijden, 1995). Conversely,
essarily restricted, to protect the experimental subject. Thus, the activation of all jaw-opening muscles resulted in normal jaw-
evoked jaw movements are small and require a very sensitive opening movements (Koolstra and van Eijden, 1997b). It is pos-
jaw-tracking device to be recorded. When the masticatory mus- sible that the varying instantaneous center of rotation can be
cles are not fully relaxed, the other muscles may easily disturb used as a means for the assessment of different muscle recruit-
the measurements. Even if complete rest can be obtained (for ment patterns during apparently similar jaw movements.
instance, in the unconscious state), the passive muscle forces With the exception of the anterior temporalis and the super-
may be relatively large compared with the evoked force and ficial masseter, the principle that muscles, when activated unilat-
cannot be ignored. Consequently, direct muscle stimulation has erally, generate a translation along their line of action and a rota-
not yet convincingly demonstrated the contributions of individ- tion about the center of gravity has been confirmed. When their
ual masticatory muscles to jaw movements. torque about the vertical axis through the center of gravity is
Though direct experimental methods have failed to dis- considered, a contralateral latero-deviation would be expected,
close the full functional potential of the masticatory muscles, but instead, an ipsilateral latero-deviation occurs (Zwijnenburg
biomechanical modeling approaches can help to develop reli- et al., 1996, 1999). This apparent paradox can be resolved if one
able predictions on this subject. Such models are always simpli- takes into account that joint loads caused by muscle forces also
fications; care must be taken that all relevant properties be contribute to jaw movement (Fig. 7). Both the anterior temporalis
included, and correct assumptions regarding model behavior and superficial masseter tend to tilt the contralateral condyle
must be made. In one model, for instance, it has been predicted from the articular eminence because of a large ipsilateral joint
that the jaw-opening muscles have the tendency to dislocate the load. The ipsilateral joint reaction force results in an ipsilaterally
temporomandibular joint (Osborn, 1993), while in another, it directed joint torque about the vertical axis through the center of
has been suggested that they have the tendency to stabilize the gravity (Koolstra and van Eijden, 1999), which overcomes the
joint (Koolstra and van Eijden, 1997b). In the former model, muscle torque to cause an ipsilateral latero-deviation.
torques were computed not with respect to the center of gravi-
ty (Stern, 1974) but with respect to the joint, thereby neglecting PASSIVE STRUCTURES
the law of conservation of angular momentum (Nigg and Passive structures contribute to jaw motion because they have
Herzog, 1999). In contrast, in the latter model, all relevant prop- the ability to resist its movements along one or more degrees

13(4):366-376 (2002) Crit Rev Oral Biol Med 371

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 faces are compressed, and the larger the
magnitude of the reaction force.
Consequently, even if articular contact is
maintained during static and dynamic situa-
tions, some movement perpendicular to the
articular surfaces occurs. This is supported
indirectly by the finding that the condylar
path during an unloaded opening lies supe-
rior 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 jaw-
opening muscles (Koolstra and van Eijden,
1997b).
The movement range of the condyle is
not limited to any great extent by the articu-
lar surfaces of the skull. Only when the
joints are compressed in an upward or
obliquely backward direction by manipula-
tion can the fossa restrain jaw movements in
the region of the condyles. The articular sur-
Figure 6. Dynamic muscle properties. Total force is the sum of the forces produced by the faces do not restrict protrusive and medio-
sarcomeres (Fsarcomeres) The active force (Factive) is dependent on the activation through the lateral translations or rotations about any of
activation dynamics, the instantaneous sarcomere length, and contraction velocity. The the three axes. If the mandible performs a
parallel elastic force (Fpassive) is dependent on the instantaneous sarcomere length. latero-deviation, the contralateral condyle
has to move forward relative to the ipsilater-
of freedom. This resistance is expressed by the structure’s abil- al one, due to the interconnection of both condyles. During
ity to generate a reaction force and/or torque. Furthermore, this movement, it is forced to move downward along the artic-
the jaw itself contributes to its movements through its inertial ular eminence. Consequently, this movement includes not
properties. The ratio between linear and angular accelerations only an azimuth rotation, but also a roll rotation (Fig. 8). While
effected by a muscle is subtly dependent on the mass and these rotations are interdependent, they nevertheless imply
moments of inertia of the jaw, and all structures that are more only one degree of freedom.
or less rigidly attached to it. This attachment may include the Articular capsule and ligaments
part of the masticatory muscles attached to the mandible, the
tongue, skin, and other soft tissues. It has been demonstrated, The bony parts of the temporomandibular joint are connected
however, that the influence of these inertial properties on the by an articular capsule composed of relatively loose collage-
final movement is small (Koolstra and van Eijden, 1995). nous fiber bundles organized in a more or less parallel fashion
(Schmolke, 1994). On the lateral side, it is reinforced by a tem-
Articular surfaces poromandibular ligament. The fibers are able to withstand
When the temporomandibular joints are loaded, reaction forces some stretching. They deform according to their (non-linear)
are transferred to the mandibular condyles. The cartilaginous elastic properties (Woo et al., 1993) and, in doing so, generate
structures in the joint have very low friction, and their deforma- tensile forces. These forces may decelerate the attached
tion is fairly independent on loading direction (Beek et al., 2000), condyle when it moves away from the articular surface of the
so it is probable that these reaction forces are directed perpen- temporal bone. A similar mechanical function can be attrib-
dicular to the contacting articular surfaces. In a sagittal plane, uted to the accessory ligaments, though these are very thin
they are directed inferiorly and pass posteriorly to the center of (Williams et al., 1995) and their function is probably negligible.
gravity of the lower jaw. These forces, therefore, have the ten- For quite a while, the temporomandibular ligaments have
dency to lower the condyle and separate the articular surfaces been considered a dominant constraint for condylar movement
(Fig. 5). The joint reaction forces also apply torques with respect and, therefore, for jaw movement. This has been illustrated by
to the center of gravity of the lower jaw. While the line of action the construction of more or less fixed axes for mandibular rota-
of the reaction forces passes posteriorly to this center, the joint tions, especially in the final phase of jaw closure, and the idea
reaction torques lead to an angular acceleration about the sagit- has been applied widely in the areas of prosthetic dentistry
tal axis through the center of gravity, which is bound to produce (Ramfjord and Ash, 1966) and temporomandibular dysfunction
an elevation movement in the positive direction. Consequently, rehabilitation (Crawford, 1999). Presently, there is a consider-
the reaction forces attempt to perform a closing jaw rotation able amount of evidence that does not support the presence of a
about this axis (Koolstra and van Eijden, 1995). fixed hinge axis (Lindauer et al., 1995; Chen and Katona, 1999)
The cartilaginous temporomandibular joint disc and the in normal functional movements. Another dominant role for the
cartilage lining of the articular surfaces are deformable, with a temporomandibular ligaments has been proposed for symmet-
finite and non-linear elasticity (Beek et al., 2001). This elasticity rical jaw-opening movements. Here, it has been assumed that
causes the joint reaction force to be dependent on the deforma- the temporomandibular ligament is always taut and forces the
tion. The more both bony articular surfaces move toward each condyle to slide down the articular eminence (Osborn, 1993).
other, the more the cartilaginous structures between these sur- However, if the temporomandibular ligament is taut, it cannot

372 Crit Rev Oral Biol Med 13(4):366-376 (2002)

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allow for habitual latero-deviation movements unless there is a
mechanism for tightening the ligaments during midline move-
ments and for slackening them during non-midline movements.
It is very unlikely that such a mechanism exists. These consid-
erations 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 temporo-
mandibular ligaments were not present (Koolstra and van Figure 7. Schematic overview of the possible actions generated by force
Eijden, 1999; Koolstra et al., 2001). Due to the interconnection Fm of the superficial masseter or anterior temporalis muscle viewed in
of left and right condyles through the mandibular symphysis, the horizontal plane. (A) Possible rotations: Rg, about center of gravity;
the two ligaments are sufficient to limit both antero-posterior Rr, about a vertical axis behind the right joint for a latero-deviation to
and medio-lateral condylar movements, during protrusive the right; and Rl, about a vertical axis behind the left joint for a latero-
and retrusive mandibular movements, and during latero-devi- deviation to the left. (B) Influence of joint forces: am, moment arm of
ations, respectively. Furthermore, when the mandible reaches muscle force; Frj, right joint force; arj, moment arm of right joint force;
these positions, the contracting masticatory muscles have Fjl, left joint force; and ajl, moment arm of left joint force.
almost reached the length where they become insufficient for
force production (Koolstra and van Eijden, 1999). The forces slower than for soft foods. Notably, the muscles are able to gener-
applied to the ligaments will then probably not be large. ate larger forces when contracting slowly. The peak velocity that
follows the fracturing of hard, brittle food is much greater than
Teeth and food
that for soft food (Peyron et al., 1997). This peak velocity, however,
The direct influence of teeth and food on jaw movements is due is less than might be expected, possibly due to a decrease in mus-
to the reaction forces occurring when the upper and lower teeth cle force as a consequence of the force-velocity relationship (Slager
come into direct contact with each other or with a bolus of food et al., 1997). Also, the size of the food affects mandibular move-
in between. Through the interplay of muscle and joint forces, ments, since the mandible has to be opened wider for larger pieces
these reaction forces will be predominantly directed downward of food to be chewed. In the frontal plane, it has been observed
(Koolstra et al., 1988) and will be accompanied by an opening that subjects chewing hard food tended to perform larger lateral
torque with respect to the center of gravity (Fig. 5), thus causing excursions than when chewing soft food (Lewin, 1985).
a joint loading. There is also an indirect influence, since the cen- Impact loads on the dentition may have consequences for
tral nervous system is able to detect forces on the dental ele- the joints, since they transfer to the joints via the mandible. A
ments through mechanoreceptors in the periodontal ligament healthy periodontium partially absorbs impact loads, and thus,
(Lund, 1991). This system, therefore, is able to adapt muscle it may prevent peak loads on the joints. This property does not
activation as required by the presence of food. Furthermore, the exist if the dentition is connected with the mandible through an
nervous system is able to react even faster through reflexes. artificial implant. The mandible itself, however, is deformable
The reaction forces are due to the resistance to deformation (van Eijden, 2000), so it is possible that the transfer of impact
by the underlying structures. The dental elements are very hard, loads of the teeth to the joints may be reduced by its elasticity.
and should they come into contact with each other with some At present, there are no quantitative data on this subject.
velocity, even a small deformation could result in a very large
reaction force. Fortunately, the dentition is connected to the Muscles
mandible via a deformable, collagenous periodontium. When a When inactive, the masticatory muscles generate passive forces
tooth of the lower dentition comes into contact with its upper which are dependent on the instantaneous length of their sar-
neighbor with a certain velocity, it undergoes little deformation comeres (Epstein and Herzog, 1998). When the sarcomeres are
but is pushed into its socket. Due to the elasticity of the peri- at or below optimum length, estimated at 2.73 µm (van Ruijven
odontium, a reaction force occurs which becomes larger as the and Weijs, 1990), they are negligible, but increase exponential-
tooth is pushed further into its socket. This reaction force acts on ly if they are stretched beyond this length. Apart from these
the mandible and causes deceleration until the movement stops. passive forces, muscle stretch can, indirectly, cause reflexes,
If food is compressed between the teeth, the reaction force because it is detected by muscle receptors (Lund, 1991).
increases more slowly, while the bolus itself is deformable. The passive forces of the jaw-closing muscles are believed
The direct influence of the teeth on jaw movements is to decelerate the jaw at the end of jaw opening during mastica-
reflected by the superior portion of the Posselt envelope of tion (Ostry and Flanagan, 1989) and become significant when
incisal point motion, but the dentition can also have an indi- the jaw nears its maximum opening (Koolstra and van Eijden,
rect influence on jaw movements. It has been demonstrated 1997a). It has been proposed that they are a determinant of max-
that subjects with malocclusion have a more irregular chewing imum jaw opening (Langenbach and Hannam, 1999; Peck et al.,
pattern than normally found (Gibbs et al., 1971; Lewin, 1985). 2000; Koolstra et al., 2001). Mathematical models applied to the
It is not clear whether these aberrant patterns are due to tooth study of the passive forces of the masticatory muscles have been
contacts themselves or to external factors. unable to open the jaw more than about 3 cm, whereas an open-
During mastication, food is compressed and/or fractured ing of 6 cm is frequently observed in vivo (Posselt, 1962; Brown,
between the dentition to reduce the particle size and facilitate 1975). Therefore, the quantitative nature of these predictions is
swallowing. This compression and fracturing take place in the disputable. Due to the proposed exponential relationship
slow-closing phase of the masticatory cycle. It is apparent that the between the passive muscle forces and their sarcomere lengths,
movement in this phase will be dependent on the mechanical small errors in the constants that determine this relationship
properties of the food. For tough foods, the compression will be may lead to relatively large errors in the projected passive

13(4):366-376 (2002) Crit Rev Oral Biol Med 373

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 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 for-
mulate and validate or falsify testable hypotheses. In particu-
lar, the availability of dynamic biomechanical modeling meth-
ods 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 mus-
cles 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 knowl-
edge 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
Figure 8. Rotations about vertical and horizontal axes during a lat- been known, since the end of the 19th century, that such axes
ero-deviation movement. are non-existent during habitual jaw movements (Ulrich,
1896). The concept of a hinge axis may have been revived by
forces. As long as there are no accurate quantitative data on the the demonstration that rotary jaw movement near occlusion
relationship between sarcomere length and passive force of the can be accomplished through manipulation on cadaverous
human masticatory muscles, this issue remains uncertain. material (Rees, 1954). This concept has survived in clinical
INTERPLAY OF PASSIVE AND ACTIVE STRUCTURES practice, where, despite a lack of a scientific basis (Mohl et al.,
1990; Lindauer et al., 1995), it has been applied successfully for
The most dominant determinants for jaw movement are the a long period (Roth and Williams, 1996).
forces generated by active muscles. Passive forces may modu- While, for instance, with the use of dynamic biomechanical
late jaw movements, but become dominant as the jaw reaches models, hypotheses regarding muscle actions in the functioning
its movement boundaries. Axes of rotation of the jaw during masticatory system are being validated (Koolstra and van
free jaw movements are not primarily related to passive struc- Eijden, 1999), joint load predictions have not yet been verified
tures of the masticatory system, but are determined by muscle satisfactorily. This is a consequence of the fact that direct mea-
use. Nevertheless, in clinical practice, axes of rotation which surement of temporomandibular joint loading without disturb-
were assumed to be connected to the temporomandibular joint ing articular integrity has remained impossible. Unfortunately,
have been measured and applied successfully for the diagnosis this parameter is considered as a major influence on the devel-
and treatment planning of masticatory dysfunction (Roth and opment of wear and degeneration of the cartilaginous and bony
Williams, 1996). This indicates that conclusions drawn from structures of this joint. Insight on temporomandibular joint load-
studies which were performed to search for practical solutions ing is, therefore, still limited to model predictions, and the relia-
to a clinical problem cannot be automatically extrapolated to bility of these predictions is directly related to the assumptions
other problems in particular or to jaw movements in general. and parameters built in such models. These include joint mor-
The influence of the passive constraints appears to be more phology and the material properties of its deformable structures
dominant as jaw movement deviates from the midline. that contribute to load distribution. Furthermore, muscle ten-
Dynamic biomechanical analysis has demonstrated that the sions applied during joint loading are required. In each subject,
masticatory muscles are capable of maintaining the integrity of these parameters may be different, leading to the need for in vivo
the masticatory system, in most cases, without the need for an measurement. Progress has been made in reconstructing the rel-
articular capsule with ligaments to maintain articular apposi- evant muscle lines of action (Koolstra et al., 1990, 1992) and bony
tion (Koolstra and van Eijden, 1997b). In contrast, they appear parts of the joints (Krebs et al., 1994) in vivo, but, to date, no reli-
to play a role in reducing the medio-lateral movements of the able method is available to create reconstructions of the carti-
mandibular condyle during latero-deviation (Koolstra and van laginous tissues in the joints. For a complete overview of the
Eijden, 1999). If the joints are loaded asymmetrically, the influ- applied muscle tensions to be acquired, their physiological cross-
ence of their reaction forces on jaw movement has to be con- sections, architecture, and degree of activation must be estimat-
sidered. When a muscle is activated unilaterally, the condylar ed. Physiological cross-sections have been estimated from
reaction forces may produce a reverse movement compared anatomic cross-sections (Weijs and Hillen, 1984; Koolstra and
with the one expected from the muscle’s line of action (vide van Eijden, 1992) and muscle activation from EMG recordings
ante). In practice, however, the muscles contract as groups (Carlsöö, 1952, 1956a,b; Møller, 1966; Lehr et al., 1971; Widmalm
rather than in isolation. For both midline and non-midline jaw et al., 1988; Murray et al., 1999a,b), but it is questionable whether
movements, dynamic muscle properties should be taken into such methods can be applied routinely to all muscles involved.
account, since they limit the force-producing capacities of the Masticatory muscle architecture has been studied in vitro (van

374 Crit Rev Oral Biol Med 13(4):366-376 (2002)

Downloaded from http://cro.sagepub.com by on February 6, 2009
Eijden et al., 1997), but the influence of individual variations on sion and measured by the CPI instrument, and signs and symptoms
model predictions and the possibility of applying relevant cor- of temporomandibular dysfunction. Angle Orthod 69:103-116.
rections have not been established. Epstein M, Herzog W (1998). Theoretical models of skeletal muscle:
biological and mathematical considerations. Chichester: John
A start has been made on assessing the dynamic material
Wiley & Sons.
properties of the cartilaginous structures in the human tem- Gal JA, Gallo LM, Murray G, Klineberg IJ, Johnson CW, Palla S (2000).
poromandibular joint (Beek et al., 2001), but the nature and Screw axes and wrenches in the study of human jaw mechanics. In:
influence of individual variations are subject to speculation. Proceedings of a symposium commemorating the legacy, works,
Consequently, although much is known qualitatively, quantifi- and life of Sir Robert Stawell Ball. Cambridge, UK: University of
cation of joint forces that incorporate individual variation still Cambridge, Trinity College.
cannot be performed unambiguously. Gallo LM, Airoldi GB, Airoldi RL, Palla S (1997). Description of
The ultimate limiting factor for reliable masticatory function mandibular finite helical axis pathways in asymptomatic subjects.
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