Braz J Oral Sci . April/June 2002 - Vol.
1 - Number 1
Andries van der Bilt 1
1
Department of Oral-Maxilofacial Surgery,
Prosthodontics and Special Dental Care.
University Medical Center Utrecht
Correspondence to:
A. Van der Bilt,
University Medical Center,
PO Box 85.060,
3508 AB, Utrecht, The Netherlands
Telephone: +31-30-2533540
Telefax: +31-30-2535537
e-mail: a.vanderbilt@med.uu.nl
7
Human oral function: a review
Abstract
Chewing is the first step in the process of digestion and is meant to
prepare the food for swallowing and further processing in the
digestive system. During chewing, the food bolus or food
particles are reduced in size, saliva is produced to moisten the
food and flavors are released. Taste and texture of the food are
perceived and have their influence on the chewing process. There
are several factors determining the chewing result. The teeth are
important in the masticatory system. They form the occlusal area
where the food particles are fragmented. This fragmentation
depends on the total occlusal area and thus on the number of
teeth. Another important factor in mastication is the bite force.
The bite force depends on muscle volume, jaw muscle activity,
and the coordination between the various chewing muscles. Also
the movement of the jaw, and thus the neuromuscular control of
chewing, plays an important role in the fragmentation of the food.
Another aspect of chewing is how well the tongue and cheeks
manipulate the food particles between the teeth. Finally, the
production of sufficient saliva is indispensable for good chewing.
Large differences in oral function exist among various groups of
subjects, such as dentate subjects, partial and complete denture
wearers, and subjects with implant-retained overdentures. Both
maximum bite force and masticatory performance are
significantly reduced, when dentures replace natural teeth. Oral
function also is impaired in orthognathic patients, both before
and after surgery, and in patients with a neuromuscular disease,
myasthenia gravis. The muscle activity, needed to overcome the
resistance of the food during mastication, consists of two
contributions: an anticipating and a sensory-induced component.
The anticipating muscle activity is generated only if food
resistance is expected. The sensory-induced muscle activity
immediately starts after food contact. About 85% of the muscle
activity needed to crush the food is sensory induced, which
indicates that jaw muscle activity is mainly of sensory origin.
Key Words:
Mastication, chewing efficiency, food comminution, particle size
distribution, saliva, swallowing, neuromuscular control
Braz J Oral Sci 1(1): 7-18
Introduction
Chewing, talking, laughing, smiling, and yawning are
important functions of the masticatory system. Often,
patients are confronted with problems in performing
these functions. This may be due to a malfunction of the
jaw muscles, jaws and/or jaw joints or the neural system.
The research on the masticatory system deals with the
general question of how (neuro-) physiological and
mechanical properties of the jaw muscles, jaws, and jaw
joints affect the development of normal and abnormal
form and function of the masticatory system. In this way
a better insight can be obtained into the etiological
factors involved in the development of disorders of the
masticatory system and an improvement can be gained
in the diagnostics and the treatment modalities.
Masticatory function has been extensively investigated
in both animal and human. In this review, the focus will
be on human oral function.
Masticatory Function
Subjective masticatory function (defined as masticatory
ability) has been studied by interviewing subjects as to
their own assessment of that function, e.g.1, 6 . Masticatory
ability appeared to be closely related to the number of
teeth. Impairment of chewing ability occurs when less
than 20 well-distributed teeth are present1.
Objective masticatory function (defined as masticatory
performance) has often been measured by determining
an individual's capacity to grind or pulverize a test food.
Several studies have shown that masticatory function is
reduced in people who have lost postcanine teeth, e.g.2,7-12
and in those who wear removable dentures, e.g.13-15 .
Implant-supported prostheses improve the oral
function and satisfaction in edentulous patients, e.g.16-20 .
Methods to Determine Masticatory Function
The rate of breakdown of food depends on many
anatomical and physiological variables. Study of the
comminution of solid food is of interest in revealing the
relative importance of these variables to masticatory
performance. A wide variety of methods has been used
to analyze chewing performance: for example,
measuring sugar loss from chewing gum21, measuring
color change in chewing gum22, 23, a colorimetric method
to determine the release of dye when chewing raw
carrots24, photometric methods to quantify changes in
color25-27, and optical scanning of chewed particles28,29.
However, in the majority of the studies the degree of
Human oral function: a review
breakdown of the food has been determined by sieving,
e.g.11, 19, 30-33 . Both natural foods, such as peanuts, almonds,
carrots and synthetic materials have been used as test
materials in experiments determining the masticatory
performance. A natural test food has the advantage that it
is normally consumed, so that subjects are accustomed to
it. However, the consistency of the food may vary due to
seasonal and geographical influences. To avoid these
variations in consistency, artificial food is a good
alternative, which has often been used, e.g.15, 19, 31, 33 .
Sieving and analysis methods
A wide variety of sieving methods has been applied in
studies on chewing. Some authors have used test sieving
with only one sieve, e.g.8,30,34-37 . In these studies the
masticatory performance was defined by the weight
percentage of masticated food that would pass a sieve
with a fixed aperture. Sieving methods, which use more
than one sieve, give more detailed information on the
distribution of particle sizes in the chewed food, e.g. 7, 12, 15, 19,
31-33
. Recently, the single and multiple sieve methods were
compared38. It was concluded, that the multiple sieve
method yields better results than the single sieve method.
Thus, the multiple sieve method should be preferred.
However, the larger amount of data obtained with the
multiple sieve method complicates the description of the
particle size distribution. Several mathematical equations
have been proposed, which express the masticatory
efficiency in terms of the weights retained on the various
sieves, e.g. 7, 31. In other studies, the cumulative weight of
the comminuted food particles was represented in a plot
showing the cumulative weight percentage undersize as
a function of the sieve aperture or particle size, e.g.30, 32, 33 .
The cumulative weight percentage undersize for a certain
sieve aperture is defined as the percentage of the particles
by weight that can pass that sieve. An example of data
points plotted in this way can be seen in Fig. 1.
The figure shows the results of a chewing experiment in
which a dentate subject with an average masticatory
function chewed 10, 20, and 55 times, respectively on an
artificial test food. The data points are curve-fitted with a
distribution function39. A logarithmic scale was chosen for
the sieve apertures, because of the wide range in sieve
apertures (0.25-8.0 mm), thus avoiding congestion of data
points in the region of small sieve apertures. As may be
expected, the data points of the experiments with
different number of chewing cycles shift towards smaller
particle size upon an increasing number of chewing
8
Braz J Oral Sci 1(1): 7-18
100
%
80
60 N: 55 20 10
40
20
0 selection and breakage of particles in each chewing cycle,
0.1 sieve diameter 1 10 mm
Fig. 1 - Cumulative weight-fraction undersize as a function of the
logarithm of the sieve aperture for a dentate subject after 10, 20, and 55
chewing strokes. Drawn lines are best fits through the data points
according to the Rosin-Rammler distribution function. The median
particle sizes for the 3 particle size distributions are indicated by the
vertical lines.
cycles. The test food consisted of 8 cubes with an edge
size of 8 mm of a dental impression material (Optosil®,
www.bayer.com)15. The cumulative weight distribution
of particle sizes may be characterized by the median
particle size, X50, which can be obtained from the graph
by determining the theoretical sieve aperture through
which 50% of the weight can pass. The number of
chewing cycles needed to halve the initial particle size,
denoted as N1/2, is a good measure of chewing
efficiency40. The median particle size decreases as a
function of the number of chewing strokes, N, according
to the relation X50 = c N-d. This relation describes the rate
of food breakdown and therefore the variables c and d
characterize the masticatory efficiency of the subjects40.
Models Describing Comminution
Matrix method for calculating particle size distributions
An approach in which the breakdown of a material is
considered as the composite result of two processes,
selection and breakage41, has been successfully applied
to the analysis of food comminution32, 40, 42, 43. Selection was
defined as the chance that a food particle is placed
between the teeth and comminuted or at least damaged
during a chewing cycle. Factors related to selection are
for instance movement of jaw, tongue and cheeks, the
total occlusal area of the molar teeth, tooth shape,
particle shape, particle size and the total amount of food
in the mouth. If a food particle has been selected, it will
be fractured between the teeth into fragments of variable
number and size. This is called the breakage process.
Breakage depends on factors such as tooth shape,
fracture characteristics of the food and the intensity and
coordination of jaw-muscle activity, which is generating
the bite force. Experiments on comminution of natural
9
Human oral function: a review
food42 and artificial food43 showed that the selection and
breakage processes could be adequately described by
rather simple functions. The selection chance was found
to increase as a power function of the particle size,
whereas the weight fractions of the fragments of a
selected particle could be adequately described by a
cumulative distribution function44,45. Due to the repeated
the size distribution of the particles of the test food will
change. Relatively large particles will gradually
disappear, while small particles will become more
abundant. The way in which selection and breakage
influence the process of food comminution can be
determined by means of a mathematical model based on
matrix algebra40. This model enables description of the
changes in particle size distribution during
comminution. The matrix also describes how particles of
different sizes are comminuted. To calculate this matrix,
the particle size distributions in various phases of
chewing must be known and the breakage of particles
must be determined in a calibration experiment. The
results from the matrix model were verified
experimentally by a double-labeling method43.
An analytic probability density for particle size in mastication
In the beginning of the chewing process, when many
unbroken food particles are still present in the mixture,
empirical functions, describing the distribution of
particle sizes, fail to give a good description. To solve this
problem, mathematical formulae were derived to
characterize the distribution of chewed food particles by
size as a function of the number of chewing cycles46, 47. The
reduction of food particle sizes was again considered to
be the composite result of a selection and a breakage
process. The probability density Pn+1(x) of finding a
particle of size x after n+1 chewing cycles was computed
from Pn(x) by selecting a proportion of particles of size y
from Pn to be converted to particles of size x < y by a
breakage function. Measures of central tendency,
average, median, and most probable size were obtained
as a function of the number of chewing cycles47. The
measures of central tendency characterize the degree of
food comminution during the chewing process and can
be used to quantify chewing performance. The
comminution of food is described in terms of the selection
and breakage functions in a convenient, efficient analytic
way, valid for all phases of the chewing process.
Braz J Oral Sci 1(1): 7-18
A selection model to estimate the interaction between food
particles and molar teeth
In the selection process, every food particle has a chance
of being placed between the antagonistic post-canine
teeth and being subjected to subsequent breakage. The
selection chance, being the ratio between the number of
selected and offered particles, has been mathematically
described as a function of the number of particles
offered, in terms of the number of breakage sites
available on the teeth and particle affinity, i.e. the
fraction of breakage sites occupied by one particle48. It
was demonstrated that the number of selected food
particles of a single size asymptotically approaches the
total number of breakage sites available for that size,
when the number of particles offered increases. From
this relationship the critical particle number can be
determined. The critical particle number indicates the
number of particles by which the breakage sites become
saturated.
Factors Influencing Masticatory Function
Many factors are known to influence masticatory
performance, such as loss and restoration of postcanine
teeth2, 7, 8, 10-12, 49, 50, bite force19, 37, 51, 52, age and gender3, 8, 19,
sensory feedback53, occlusal contact area54, and oral
motor function55. Only one factor at a time was studied in
the majority of these studies. The main factors
influencing masticatory performance were determined
in only a few of these studies10, 37, 54. Number of functional
tooth units and bite force were confirmed as the key
determinants of masticatory performance37.
Occlusal factors
Many people have deficiencies in masticatory
performance because of loss of teeth, malocclusion, or
periodontal disease. However, in spite of their handicap,
most people manage to eat successfully even though
they are unable to comminute their food perfectly before
swallowing. In order to improve the masticatory
function, missing teeth are often replaced by fixed or
removable prosthodontic appliances. Indeed, a clear
relationship exists between dental state and masticatory
performance as determined with a chewing test, e.g.2, 7-12,
37, 49, 54, 56, 57
. A considerable variation in the masticatory
performance is found, which may be related to many
different factors such as the total occlusal surface, the
occlusal contact area, the number of teeth present, the
number of occluding pairs of teeth, tooth shape, the
Human oral function: a review
preferred side and the action of the soft tissues. In a study
on the influence of occlusal factors on the masticatory
performance in 32 young dentate subjects, it was found
that the rate of comminution was most highly correlated
with the occlusal area of the postcanine teeth9. An even
more important factor controlling the masticatory
performance of people with natural teeth proved to be the
amount of occlusal contact area of molar and premolar
teeth, which is on average one fifth of the total occlusal
surface57. A multiple regression analysis was used to
predict the masticatory performance from occlusal
factors10. Their findings revealed a significant correlation
between masticatory performance and the state of
occlusion. However, the multiple regression analysis
showed that only 48% of the masticatory performance
score could be predicted from the occlusal factors, which
indicates that other factors may also affect masticatory
performance. The number of postcanine teeth appeared
to be less important than the number of occlusal contacts
of these teeth7, 12, 57. In these studies, the number of
occluding posterior teeth was expressed in occlusal units.
An occluding molar pair was counted as two occlusal
units, whereas a premolar pair was counted as one
occlusal unit2. The number of occlusal units per side was
also determined as the distribution of occlusal units is
known to influence chewing performance, e.g.2, 8. The
number of occlusal units appeared to be a good predictor
of chewing performance7, 11, 12, 37.
Maximum bite force
Significant correlations are reported between masticatory
performance and maximum bite force for subjects with
natural dentition, shortened arch as well as complete
arch19, 37, 51, 52. This indicates that a higher bite force leads to a
better fragmentation of the food. However, in some
groups of edentate subjects, such a correlation did not
exist19. In a group of subjects with implant-retained
overdentures maximum bite force and masticatory
performance were not related. This was caused by large
variations in both maximum bite force and masticatory
efficiency.
Sensory feedback and manipulation of food
The selection process of human mastication is normally
treated as a mechanico-statistical function rather than a
specific motor response driven by sensory input58.
However, the selection of food depends on the
manipulation of the food by the tongue and cheeks to sort
10
Braz J Oral Sci 1(1): 7-18
out coarse particles and bring them on to the occlusal
surfaces of the teeth for further reduction53. A chewing
stroke will be ineffective for particle reduction when
there is an insufficient quantity of large particles
between the occlusal surfaces. It has been shown, on a
quantitative basis, that local anesthesia impairs chewing
efficiency53. Ten persons with intact dentitions chewed
on standard quantities of peanuts on their preferred
chewing side. An average of 40 chewing strokes was
required after unilateral anesthesia to achieve almost the
same performance achieved with 20 strokes before
anesthesia. Thus, peripheral sensory impairment affects
masticatory performance in dentate persons. Also
tongue motor skill may play an important role in the
manipulation of the food. Tongue motor skill was
examined by an ultrasound system in 3 groups of
subjects: 30 healthy adult dentates, 10 elderly dentates,
and 20 complete denture wearers55. The tongue motor
skill of the elderly dentates and complete denture
wearers were statistically lower than those of the adult
dentates. Furthermore, the tongue motor skill and
masticatory performance were significantly correlated.
In a recent study, an index of oral manipulative skill was
introduced by measuring the fastest possible movement
to bite on a small rubber ball (diameter of 8mm)
repetitively between the right and left posterior teeth59
The cycle time of the movement was significantly
correlated with the masticatory performance as
obtained from chewing peanuts. Masticatory
performance appeared not to be correlated with oral
stereognostic ability in a group of 71 dentate subjects
and 64 denture wearers60Apparently, the ability to
recognize the edible test forms (made from raw carrots)
in the mouth is not related with the chewing result in
both dentate subjects and denture wearers.
Age
In a comprehensive study of oral function in humans3
the masticatory performance was determined for a
group of 814 subjects between 25 and 75 years of age. The
investigation was limited to persons with natural
dentition or with missing dentition replaced by a fixed
prosthesis. According to this study, masticatory
performance does not alter significantly with age in
persons who have a complete or almost complete
dentition. However, in these persons, there is a
significant increase with age in the number of strokes
used to prepare the test food for swallowing. Although
11
Human oral function: a review
maximum bite force appeared to be significantly larger in
a group of young dentate subjects as compared to old
dentate subjects, no significant differences in masticatory
efficiency were observed19 . Apparently, the lower
maximum bite force of the older subjects was still high
enough for chewing the test food.
Saliva
Saliva participates in digestive functions by contributing
to food breakdown, by dissolving and releasing food taste
and odor chemicals, and by lubricating the food bolus for
swallowing61 . Reduction of particle size, reduction of
resistance against food deformation and the formation of
a coherent bolus that can be swallowed are the main goals
of the chewing process. Saliva moistens the fragmented
food particles during chewing, so that the food can be
swallowed62, 63 . However, in a group of 177 dentate
subjects only a low, but significant correlation (r = 0.19)
was found between the flow rate of saliva and masticatory
performance as obtained with an artificial food
(unpublished observations). Salivary glands and the
saliva they produce also play a major role in the health of
the oral cavity and the proximal portion of the
gastrointestinal tract64.
Food texture and taste
Food texture has a large influence on the chewing process,
e.g.61, 65, 66. A clear relationship between muscular activity
and food properties has been reported67, 68. The number of
chewing cycles preceding the first swallow depends on
the texture of the food62, 65. A firm and dry food, as for
instance nuts, requires many chewing cycles to fragment
the particles and to wet them with saliva before they can
be swallowed62. Although it may be expected that the taste
of food also has much influence on the chewing process, it
is disappointing to notice that experiments and thus data
on this question are scarce69. It is suggested that an
influence of taste on the chewing process runs via its
relation with saliva flow rate. Flavor release from food
may be related to the surface of the fragmented food
particles and thus it will be related to masticatory
performance70.
Swallowing threshold
The swallowing threshold is influenced by the
masticatory performance. Subjects with a reduced
chewing performance, due to an inadequate dentition,
Braz J Oral Sci 1(1): 7-18
needed more chewing cycles to prepare the food for
swallowing than those with a good performance7, 11, 71.
Furthermore, they swallowed larger food particles than
those with good performance8, 11, 56, 72. Thus, subjects with
an inadequate dentition compensate for their reduced
chewing performance by chewing for a longer period of
time and by swallowing larger food particles. The
initiation of swallowing, which is voluntary, has been
thought to depend on separate thresholds for food
particle size and for particle lubrication73. However,
instead of this duality, it has also been suggested that
swallowing is initiated when it is sensed that a batch of
food particles is binding together under viscous forces
so as to form a bolus63. The swallowing process has been
studied directly with videofluorography74. Food
movements during complete feeding sequences on soft
and hard foods, coated with barium sulfate, were
investigated with this technique. The movements of
tongue, jaw, hyoid, and food were recorded, so that the
various stages of intra-oral transport, including
swallowing, could be quantified.
Oral function in various groups of individuals
Dentate subjects
In most studies on the relation between dental state and
masticatory function, subjects with a poor dental state
were compared with subjects who have a complete
dentition. However, there have been few studies that
directly determine the influence on masticatory function
of prosthodontic treatment. The objective masticatory
function improved immediately after prosthodontic
treatment56. Thereafter, a further gradual incre1ase in
masticatory efficiency occurred until after a month the
masticatory function was optimal. The masticatory
performance of partially edentulous patients has been
studied before and after completion of prosthetic
restoration12. The results of the patient group were
compared with data obtained for a control group of
subjects having a full dentition. In this way it was
quantified to what extent the masticatory function could
be restored to normal levels. The total number of
occluding postcanine teeth increased as a result of the
prosthodontic treatment, yielding a significantly
improved objective masticatory function. The average
masticatory performance was found to approach the
level of a control group of subjects with a complete
dentition, if all occlusal units of the longest posterior
Human oral function: a review
side were replaced. Subjects with an incomplete dentition
tended to chew predominantly at the side of the longest
posterior arch. Rehabilitation of post-canine teeth
restores some of the objective masticatory function and
leads to an increased appreciation of the masticatory
function, although no correlation between the change in
objective and subjective masticatory function was found.
Complete denture wearers
The ability of denture wearers to break down test food is
very poor as compared with that of young adults with
natural dentitions, e.g.13-15, 75, 76. Food pulverization
experiments have shown that complete denture wearers
have a much lower chewing efficiency than persons with
natural dentitions. Complete denture wearers needed on
average 4 times,19, 30, 77 , 6 times,13and even 8 times78 the
number of chewing strokes of dentate persons to achieve
the same degree of pulverization. It was found that the
difference in chewing efficiency between dentates and
denture wearers depends on the consistency of the food15.
The denture wearers needed on average 3 to 5 times the
number of chewing cycles for a soft, respectively hard
food. One of the factors leading to the decrease in chewing
performance is the reduced bite force that denture
wearers can develop due to a lack of retention and
stability of the denture. The bite force of these subjects
obtained with maximum clenching (unilateral) ranges
from 77 to 135 N14, 19, 75, 79, 80, whereas the average maximum
bite force (unilateral) for dentate persons varies from 330
to 522 N14, 19, 80-83. The maximum bite forces of denture
wearers may be even lower than the forces needed to
penetrate natural foods, such as boiled meat (80 N), raw
carrot (118 N), and rye bread (167 N)79. Thus, denture
wearers might have difficulties in biting and incising such
foods. As a consequence, full denture wearers select only
a few food particles at a time, so the total force needed to
crush the food is limited15. We may conclude that
edentulous persons are handicapped in masticatory
function and even clinically satisfactory complete
dentures are poor substitutes for natural teeth.
Implant-retained overdentures
The objective oral function significantly improves when
the mandibular denture is supported by oral implants.
The maximum bite force of subjects with a mandibular
denture supported by implants is 60 to 200% higher than
that of subjects with a conventional denture20, 84-87.
12
Braz J Oral Sci 1(1): 7-18
However, the average bite force after treatment was still
only two thirds of the value obtained for dentate
subjects 2 0 . The masticatory performance also
significantly improved after implant treatment16, 77. The
number of chewing cycles needed to halve the initial size
of the food decreased from 51 to 2877. The improvement
of oral function after implant treatment does not seem to
depend on the degree of implant support16, 87. In these
studies, it was shown that bite force and masticatory
performance did not differ among subjects with mainly
implant-borne overdentures on a transmandibular
implant and subjects with mucosa-borne overdentures
on two implants.
The attachment type of implant-retained mandibular
overdentures may influence the retention and the
stability of the denture, and thus the oral function. In a
within-subject crossover clinical trial the influence of
retention and stability of the denture on the maximum
bite force and the corresponding EMG, and on the
masticatory performance was studied20, 77. Eighteen
subjects received two permucosal implants and three
suprastructure modalities: magnet, bar-clip, or ball
attachment. The 3 suprastructures were worn
successively by all 18 subjects, so a within-subject
comparison of the oral function obtained with the 3
attachment types could be made. No significant
differences in maximum bite force, muscle activity, and
masticatory performance were found among the 3
attachment types20, 77.
The subjective oral function of subjects with an implant-
retained overdenture has been recently reviewed88. Both
chewing ability and satisfaction improved as a result of
implant treatment89, 90.
Orthognathic patients
A reduced chewing performance has been reported for
patients with a mandibular prognathism, who were
scheduled for orthognathic surgery91-95. After surgical
correction, chewing performance improved
significantly in three out of five of these studies91, 93, 95.
Patients with a hypoplastic mandible also showed a
reduced chewing performance before treatment96. In this
study the patients needed about twice the number of
chewing cycles as the controls to halve the initial size of
the food particles. The chance to select a food particle for
breakage as well as the degree of breakage was
significantly smaller for the patients than for the
13
Human oral function: a review
controls. These findings suggest that the reduced
chewing performance of pre-orthognathic surgery
patients is due to an impairment of both selection and
breakage of food particles.
Neuromuscular disease: Myasthenia Gravis
Fluctuating weakness of voluntary muscles and
abnormal fatigability on exertion are the most prominent
characteristics of myasthenia gravis, a neuromuscular
disease in which the neuromuscular transmission is
impaired by an autoimmune reaction to the postsynaptic
acetylcholine receptors. All skeletal muscles may be
involved. Bulbar myasthenia gravis refers to weakness in
jaw muscles. Myasthenia gravis patients with bulbar
involvement may suffer from difficulties in swallowing
and chewing, changed facial expression, and dysarthria.
A group of 20 bulbar myasthenia gravis patients has been
compared with a group of 20 healthy controls, matched
for age, gender, and dental status97. Masticatory muscle
strength, chewing performance, and tongue force were
quantified in patients with bulbar myasthenia gravis and
compared with that of the healthy subjects. The
maximum bite force was significantly lower for the
myasthenia gravis patients than for the controls: 264 ± 40
N versus 487 ± 48 N98. The chewing performance was
impaired in the patient group99. The median particle size
after 15 chewing cycles was 5.6 ± 2.0 and 3.2 ± 1.4 mm for
the patients and controls, respectively. The force of the
tongue, which plays an important role in the swallowing
process, was measured with the use of a tongue force
transducer100. The results indicated that maximum tongue
force in the patient group was significantly lower in all
directions than the tongue force of the controls101.
Neuromuscular control of chewing
The movement of the jaw, and thus the neuromuscular
control of chewing, also plays an important role in the
comminution of the food. Chewing requires muscle
activity to make the movements of the jaw and to exert
forces in order to cut or grind the food. A relatively low
level of muscle activity is observed in the surface EMG of
the closing muscles of subjects making pseudo-chewing
movements without food. More muscle activity is
generated if the closing movement is counteracted by
food resistance58, 102. Apparently, a small part of the muscle
activity observed during chewing is needed just for the
basic rhythmic movements of the jaw, and additional
Braz J Oral Sci 1(1): 7-18 Human oral function: a review

muscle activity is required to overcome the resistance of pressoreceptors by deafferentiation. It was concluded
the food. The total amount of EMG activity has been that periodontal pressoreceptors, and muscle spindles,
shown to depend on the texture of the food: more EMG provide positive feedback to the jaw-closing muscles
activity is observed for harder foods66-68. during mastication.

Central pattern generator Simulated chewing experiments

The brain stem has been shown to be an essential part of The neuromuscular control of chewing in humans has
the central nervous system that is necessary for been studied in our laboratory, e.g.113-115 . In these studies
mastication, because decerebrate animals and animals food resistance was simulated by a computer controlled
without a cerebellum or spinal cord can still chew103-107. external load, acting on the mandible in a downward
The basic rhythmic activity of the jaw-opening and jaw- direction during closing (Fig. 2).
closing muscles is probably evoked by a central pattern
generator located in the brain stem58. Cortically evoked
rhythmic trigeminal activity remained present in
animals after elimination of sensory feedback from
peripheral receptors103, 108. This shows that neither muscle
spindle afferents nor periodontal afferents are essential
to the basic rhythmic activity patterns of mastication.
Cortical stimulation of the anaesthetized rabbit induced
rhythmical mandibular movements in the awake
animal109. The central pattern generator may be switched
on by activity of higher centers or by intraoral stimuli104,
110
.

Peripheral feedback
Comparison of the movements and the activity patterns Fig. 2 - Schematic drawing of the experimental setup. In the experiment, an
external force simulates food resistance.
in the motor nerves evoked by cortical stimulation of the
paralyzed animal with those of natural chewing before
Sequences of cycles with a load were unexpectedly
paralysis, has demonstrated the important role of
alternated with sequences of cycles without a load. Jaw
sensory feedback in mastication107. During cortical
movement, and EMG of the masseter, temporalis, and
stimulation, the central pattern generator produces
digastric muscles were recorded. It was demonstrated
stereotyped open-close cycles, whereas during natural
that the additional muscle activity, needed to counteract
chewing the movement trajectories of the consecutive
the external load, consists of two components: an
chewing cycles vary considerably107. Moreover, the
anticipating component starting before the onset of the
activity of the jaw-closer -motoneurons is much smaller
food simulating load and a peripherally induced
in fictive mastication than during natural chewing. This
component starting after the onset of the load. The
suggests that to adequately fulfil the motor tasks of the
anticipating component is generated only if a
mandible during chewing, the central nervous system
counteracting load is expected. The onset of the
requires information about the position and velocity of
anticipating muscle activity occurs immediately after the
the mandible, about the forces acting on the mandible
moment that the jaw starts closing. Peripherally induced
and on the teeth, and about the length and contraction
muscle activity is generated on average 23 ms after the
velocity of the muscles involved. An increase of the
onset of the load. About 85% of the muscle activity needed
amplitude and the duration of the activity of the jaw
to overcome the external load is peripherally induced,
closing muscles of the rabbit was observed, when
which indicates that the muscle activity is mainly of
cortically induced rhythmic open-close movements
sensory origin. However, when the movement rate of
were obstructed by a steel ball or a foam strip between
chewing was doubled (fast chewing with 120 cycles per
antagonistic teeth 109, 111, 112. This effect was reduced after
minute), the contribution of peripherally induced muscle
elimination of feedback from the periodontal
14
Braz J Oral Sci 1(1): 7-18
activity decreased to only 40%. Therefore, as jaw
movement speed increased, emphasis in the control of
the muscle activity shifted from sensory induced
(closed-loop) to feed forward (open-loop) control116.
Muscles spindles are primarily responsible for the
peripherally induced muscle activity as was
demonstrated in an experiment on anesthetized
rabbits112. The masticatory system is mainly closed-loop
controlled. A reason for this may be the fact that the
relatively large forces needed for food fragmentation
must be controlled under uncertain conditions. First, no
optical feedback is available in the chewing process.
Furthermore, food resistance may vary largely from
cycle to cycle. Thus, immediate muscle response is
necessary to maintain a constant chewing rhythm.
Force-velocity properties of the jaw-closing muscles
play a major role in the situation that the food resistance
suddenly disappears117. In that case reflex activity is too
slow to limit the jaw velocity at impact. The force-
velocity properties of the muscles provide a quick
mechanism for dealing with unexpected closing
movements and so avoid damage to the dental elements.
An experiment with rhythmic arm movements,
comparable to the rhythmic jaw movements described
above, showed that arm and jaw muscles respond
differently to loading118. In the arm muscles, there was
little reflex activity, but a large anticipatory response.
This indicates that reflexes do not play an important role
in these rhythmic arm movements. This emphasizes that
the mainly reflexly induced control of the jaw closing
muscles is a unique phenomenon.
Acknowledgements
This work was supported by the University Medical
Center Utrecht and the Netherlands Institute for Dental
Sciences. I am grateful to Dr. F.A. Fontijn-Tekamp for
critical reading of the manuscript.
References
1. Agerberg G, Carlsson GE. Chewing ability in relation to
dental and general health. Analyses of data obtained from
a questionnaire. Acta Odontol Scand 1981; 39:147-153.
2. Käyser AF. Shortened dental arches and oral function. J
Oral Rehabil 1981; 8:457-462.
3. Wayler AH, Chauncey HH. Impact of complete dentures
and impaired natural dentition on masticatory
performance and food choice in healthy aging men. J
Prosthet Dent 1983; 49:427-433.
4. Chauncey HH, Muench ME, Kapur KK, Wayler AH. The
effect of the loss of teeth on diet and nutrition. Int Dent J
15
Human oral function: a review
1984; 34:98-104.
5. Oosterhaven SP, Westert GP, Schaub RH, van der Bilt A.
Social and psychologic implications of missing teeth for
chewing ability. Community Dent Oral Epidemiol 1988;
16:79-82.
6. Witter DJ, Elteren P, Käyser AF, Rossum GM. Oral comfort
in shortened dental arches. J Oral Rehabil 1990; 17:137-143.
7. Helkimo E, Carlsson GE, Helkimo M. Chewing efficiency
and state of dentition. A methodologic study. Acta Odontol
Scand 1978; 36:33-41.
8. Feldman RS, Kapur KK, Alman JE, Chauncey HH. Aging
and mastication: changes in performance and in the
swallowing threshold with natural dentition. J Am Geriatr
Soc 1980; 28:97-103.
9. Luke DA, Lucas PW. Chewing efficiency in relation to
occlusal and other variations in the natural human
dentition. Br Dent J 1985; 159:401-403.
10. Omar SM, McEwen JD, Ogston SA. A test for occlusal
function. Brit J Orthod 1987; 14:85-90.
11. van der Bilt A, Olthoff LW, Bosman F, Oosterhaven SP. The
effect of missing postcanine teeth on chewing performance
in man. Arch Oral Biol 1993; 38:423-429.
12. Van der Bilt A, Olthoff LW, Bosman F, Oosterhaven SP.
Chewing performance before and after rehabilitation of
postcanine teeth in man. J Dent Res 1994; 73:1677-
1683.
13. Kapur KK, Soman SD. Masticatory performance and
efficiency in denture wearers. J Prosthet Dent 1964;
14:687-693.
14. Helkimo E, Carlsson GE, Helkimo M. Bite force and state
of dentition. Acta Odontol Scand 1977; 35:297-303.
15. Slagter AP, Bosman F, van der Bilt A. Comminution of two
artificial test foods by dentate and edentulous subjects. J
Oral Rehabil 1993; 20:159-176.
16. Geertman ME, Slagter AP, van Waas MAJ, Kalk W.
Comminution of food with mandibular implant-retained
overdentures. J Dent Res 1994; 73:1858-1864.
17. Kapur KK, Garrett NR, Hamada MO, Roumanas ED,
Freymiller E, Han T, Diener RM, Levin S, Ida R. A
randomized clinicla trial comparing the efficacy of
mandibular implant-supported overdentures and
conventional dentures in diabetic patients. Part I:
Methodology and clinical outcomes. J Prosthet Dent 1998;
79:555-569.
18. Tang L, Lund JP, Taché R, Clokie CML, Feine JS. A within-
subject comparison of mandibular long-bar and hybrid
implant-supported prostheses: Evaluation of masticatory
function. J Dent Res 1999; 78:1544-1553.
19. Fontijn-Tekamp FA, Slagter AP, van der Bilt A, van 't Hof
MA, Witter DJ, Kalk W, Jansen JA. Biting and chewing with
mandibular implant-retained overdentures compared
with other states of artificial and natural dentition. J Dent
Res 2000; 79:1519-1524.
20. van Kampen FMC, van der Bilt A, Cune MS, Bosman F. The
influence of various attachment types in mandibular
implant-retained overdentures on maximum bite force
and EMG. J Dent Res 2002; 81:170-173.
21. Heath MR. The effect of maximum biting force and bone
loss upon masticatory function and dietary selection of the
elderly. Int Dent J 1982; 32:345-356.
Braz J Oral Sci 1(1): 7-18
22. Hayakawa I, Watanabe I, Hirano S, Nagao M, Seki T. A
simple method for evaluating masticatory performance
using a color- changeable chewing gum. Int J Prosthodont
1998; 11:173-176.
23. Prinz JF. Quantitative evaluation of the effect of bolus size
and number of chewing strokes on the intra-oral mixing of
a two-colour chewing gum. J Oral Rehabil 1999; 26:243-
247.
24. Käyser AF, Hoeven v. Colorimetric determination of the
masticatory performance. J Oral Rehabil 1977; 4:145-148.
25. Masuda G, Fujiyama N, Koga I, Fukagai T, Masuda M,
Ohtani T, Suzuki S. The new method of measuring
masticatory performance using spectrophtometer with
ATP granules. 1st Evaluation of measuring method and
materials. J Jpn Stomatol Soc 1981; 30:103-110.
26. Gunne H-SJ. Masticatory efficiency. A new method for
determination of the breakdown of masticated test
material. Acta Odontol Scand 1983; 41:271-276.
27. Nakasima A, Higashi K, Ichinose M. A new, simple and
accurate method for evaluating masticatory ability. J Oral
Rehabil 1989; 16:373-380.
28. Shi CS, Ouyang G, Guo TW. Comparison of food particle
distribution masticated by subjects wearing complet
dentures and with natural teeth. J Oral Rehabil 1990;
17:611-615.
29. van der Bilt A, van der Glas HW, Mowlana F, Heath MR. A
comparison between sieving and optical scanning for the
determination of particle size distributions obtained by
mastication in man. Arch Oral Biol 1993; 38:159-162.
30. Manly RS, Braley LC. Mastication performance and
efficiency. J Dent Res 1950; 29:448-462.
31. Edlund J, Lamm CJ. Masticatory efficiency. J Oral Rehabil
1980; 7:123-130.
32. Lucas PW, Luke DA.Methods for analysing the
breakdown of food in human mastication. Arch Oral Biol
1983; 28:813-819.
33. Olthoff LW, van der Bilt A, Bosman F, Kleizen HH.
Distribution of particle sizes in food comminuted by
human mastication. Arch Oral Biol 1984; 29:899-903.
34. Demers M, Bourdages J, Brodeur JM, Benigeri M.
Indicators of masticatory performance among elderly
complete denture wearers. J Prosthet Dent 1996; 75:188-
193.
35. Kapur KK, Garrett NR, Dent RJ, Hasse AL. A randomized
clinical trialof two basic removable denture designs. Part
II: Comparisons of masticatory scores. J Prosthet Dent
1997; 78:15-21.
36. Garrett NR, Kapur KK, Hamada MO, Roumanas ED,
Freymiller E, Han T, Diener RM, Levin S, Chen T. A
randomized clinical trial comparing the eficiacy of
mandibular implant-supported overdentures and
conventional dentures in diabetic patients. Part II.
Comparisons of masticatory performance. J Prosthet Dent
1998; 79:632-640.
37. Hatch JP, Shinkai RSA, Sakai S, Rugh JD, Paunovich ED.
Determinants of masticatory performance in dentate
adults. Arch Oral Biol. 2000; 46:641-648.
38. van der Bilt A, Fontijn-Tekamp FA. Comparison of single
and multiple sieve methods for the determination of
masticatory performance. J Dent Res 2002; 81:A-455.
Human oral function: a review
39. Rosin P, Rammler E. Gesetzmassigkeiten in der
Kornzusammensetzung des Zementes. Zement 1933;
31:427-433.
40. van der Bilt A, Olthoff LW, van der Glas HW, van der
Weelen K, Bosman F. A mathematical description of the
comminution of food during mastication in man. Arch Oral
Biol 1987; 32:579-586.
41. Epstein B. The mathematical description of certain
breakage mechanisms leading to the logarithmico-normal
distribution. J Franklin Inst 1947; 244:471-477.
42. Lucas PW, Luke DA. Computer simulation of the
breakdown of carrot particles during human mastication.
Arch Oral Biol 1983; 28:821-826.
43. van der Glas HW, van der Bilt A, Olthoff LW, Bosman F.
Measurement of selection chances and breakage functions
during chewing in man. J Dent Res 1987; 66:1547-1550.
44. Gaudin AM, Meloy TP. Model and a comminution
distribution equation for single fracture. AIME Trans 1962;
223:40-43.
45. Austin LG. A review; introduction to the mathematical
description of grinding as a rate process. Powder
Technology 1971; 5:1-17.
46. Voon FT, Lucas PW, Chew KL, Luke DA. A simulation
approach to understanding the masticatory process. J
Theor Biol 1986; 119:251-262.
47. Baragar FA, van der Bilt A, van der Glas HW. An analytic
probability density for particle size in human mastication. J
Theor Biology 1996; 181:169-178.
48. van der Glas HW, van der Bilt A, Bosman F. A selection
model to estimate the interaction between food particles
and the post-canine teeth in human mastication. J Theor
Biol 1992; 155:103-120.
49. Carlsson GE. Masticatory efficiency: the effect of age, the
loss of teeth and prosthetic rehabilitation. Int Dent J 1984;
34:93-97.
50. Akeel R, Nilner M, Nilner K. Masticatory efficiency in
individuals with natural dentition. Swed Dent J 1992;
16:191-198.
51. Wilding RC. The association between chewing efficiency
and occlusal contact area in man. Arch Oral Biol 1993;
38:589-596.
52. Boretti G, Bickel M, Geering AH. A review of masticatory
ability and efficiency. J Prosthet Dent 1995; 74:400-403.
53. Kapur KK, Garrett NR, Fischer E. Effects of anaesthesia of
human oral structures on masticatory performance and
food particle size distribution. Arch Oral Biol 1990; 35:397-
403.
54. Julien KC, Buschang PH, Throckmorton GS, Dechow PC.
Normal masticatory performance in young adults and
children. Arch Oral Biol 1996; 41:69-75.
55. Koshino H, Hirai T, Ishijima T, Ikeda Y. Tongue motor skills
and masticatory performance in adult dentates, elderly
dentates, and complete denture wearers. J Prosthet Dent
1997; 77:147-152.
56. Abel LF, Manly RS. Masticatory function of partial denture
patients among navy personel. J Prosth Dent 1953; 3:382-
392.
57. Yurkstas AA. The masticatory act. A Review. J Prosthet
Dent 1965; 15:248-260.
58. Thexton AJ. Mastication and swallowing: an overview. Brit
16
Braz J Oral Sci 1(1): 7-18
Dent J 1992; 173:197-206.
59. Suzuki T, Okamoto Y, Hayakawa I. Evaluation of an index
of oral manipulative skill. J Dent Res 2002; 81:A-158.
60. Garrett NR, Kapur KK, Jochen DG. Oral stereognostic
ability and masticatory performance in denture wearers.
Int J Prosthodont 1994; 7:567-573.
61. Christensen CM. Food texture perception. Adv Food Res
1984; 29:159-199.
62. Prinz JF, Lucas PW. Swallow thresholds in human
mastication. Arch Oral Biol 1995; 40:401-403.
63. Prinz J, Lucas P. An optimization model for mastication
and swallowing in mammals. Proc R Soc Lond B
1997; 264:1715-1721.
64. Mandel ID. The functions of saliva. J Dent Res 1987;
66:623-627.
65. Hiiemae KM, Heath MR, Heath G, Kazazoglu E, Murray J,
Sapper D, Hamblett K. Natural bites, food consistency
and feeding behaviour in man. Arch Oral Biol 1996;
41:175-189.
66. Mioche L, Bourdiol P, Martin J-F, Noël Y. Variations in
human masseter and temporalis muscle activity related to
food texture during free and side-imposed mastication.
Arch Oral Biol 1999; 44:1005-1012.
67. Mathevon E, Mioche L, Brown WE, Culioli J. Texture
analysis of beef cooked at various temperatures by
mechanical measurements, sensory assessments and
electromyography. J Texture Stud 1995; 26:175-192.
68. Agrawal KR, Lucas PW, Bruce IC, Prinz JF. Food
properties that influence neuromuscular activity during
human mastication. J Dent Res 1998; 77:1931-1938.
69. Bosman F, van der Bilt A, van der Glas HW. The taste of
chewing. In: Schieberle P, ed. Food science and technology
Cost 96. 1998. p.113-120.
70. Brown WE, Langley KR, Martin A, MacFie HH.
Characterisation of patterns of chewing behaviour in
human subjects and their influence on texture perception.
J Texture Stud 1994; 25:455-468.
71. Lucas PW, Luke DA. Is food particle size a criterion for the
initiation of swallowing? J Oral Rehabil 1986; 13:127-136.
72. Yurkstas A. Compensation for inadequate mastication.
Brit Dent J 1951; 91:261-262.
73. Hutchings JB, Lillford PJ. The perception of food texture-
the philosophy of the breakdown path. J Texture Stud
1988; 19:103-115.
74. Hiiemae KM, Palmer JB. Food transport and bolus
formation during complete feeding sequences on
foods of different initial consistency. Dysphagia
1999; 14:31-42.
75. Haraldson T, Karlsson U, Carlsson GE. Bite force and oral
function in complete denture wearers. J Oral Rehabil 1979;
6:41-48.
76. Slagter AP, Olthoff LW, Steen WA, Bosman F.
Comminution of food by complete-denture wearers. J
Dent Res 1992; 71:380-386.
77. van Kampen FMC, van der Bilt A, Cune MS, Bosman F.
Mandibular implant-overdenture treatment and oral
function. J Dent Res 2002; 81:A-60.
78. Yurkstas AA, Emerson WH. Decreased masticatory
function in denture patients. J Prosthet Dent 1964; 14:931-
934.
17
Human oral function: a review
79. Eerikäinen E, Könönen M. Forces required by complete
dentures for penetrating food in simulated function. J
Oral Rehabil 1987; 14:607-613.
80. Slagter AP, Bosman F, van der Glas HW, van der Bilt A.
Human jaw elevator muscle activity and food
comminution in the dentate and edentulous state. Arch
Oral Biol 1993; 38:195-205.
81. Bakke M, Michler L, Han K, Möller E. Clinical significance
of isometric bite force versus electrical activity in temporal
and masseter muscles. Scand J Dent Res 1989; 97:539-551.
82. Bakke M, Holm B, Jensen BL, Michler L, Möller E.
Unilateral, isometric bite force in 8-68-year old women and
men related to occlusal factors. Scand J Dent Res 1990;
98:149-158.
83. van der Bilt A, Abbink JH, Fontijn-Tekamp FA, Bosman F.
Maximal bite force and EMG during bilateral and
unilateral clenching. J Oral Rehabil 2002; in press.
84. Lindquist LW, Carlsson GE. Long-term effects on chewing
with mandibular fixed prostheses on osseo-integrated
implants. Acta Odontol Scand 1985; 43:39-45.
85. Haraldson T, Jemt T, Stålblad P, Lekholm U. Oral function
in subjects with overdentures supported by
osseointegrated implants. Scand J Dent Res 1988; 96:235-
242.
86. Carlsson GE, Lindquist LW. Ten-year longitudinal study of
masticatory function in edentulous patients treated with
fixed complete dentures on osseointegrated implants. Int J
Prosthodont 1994; 7:448-453.
87. Fontijn-Tekamp FA, Slagter AP, van 't Hof MA, Geertman
ME, Kalk W. Bite forces with mandibular implant-retained
overdentures. J Dent Res 1998; 77:1832-1839.
88. Locker D. Patient-based assessment of the outcomes of
implant therapy: A review of the literature. Int J
Prosthodont 1998; 11:453-461.
89. de Grandmont P, Feine JS, Tach‚ R, Boudrias P, Donohue
WB, Tanguay R, Lund JP. Within-subject comparisons of
implant-supported mandibular prostheses: psychometric
evaluation. J Dent Res 1994; 73:1096-1104.
90. Geertman ME, Boerrigter EM, van 't Hof MA, van Waas
MAJ, van mandibular overdentures versus complete
dentures - chewing ability. Community Dent Oral
Epidemiol 1996; 24:79-84.
91. Shiratsuchi Y, Kouno K, Tashiro H. Evaluation of
masticatory function following orthognathic
surgical correction of mandibular prognathism. J Cranio
Maxill Surg 1991; 19:299-303.
92. Kobayashi T, Honma K, Hanada K. Masticatory function
in patients with mandibular prognatism before and after
orthognathic surgery. J Oral Maxil Surg 1993; 51:997-1001.
93. Kikuta T, Hara I, Seto T, Yoshioka I, Nakashima T,
Yasumitsu C. Evaluation of masticatory function after
sagittal split ramus osteotomy for patients with
mandibular prognathism. Int J Adult Orthod
Orthognath Surg 1994; 9:9-17.
94. Zarrinkelk HM, Throckmorton GS, Ellis E, Sinn DP. A
longitudinal study of changes in masticatory performance
of patients undergoing orthognatic surgery. J Oral Maxil
Surg 1995; 53:777-782.
95. Kobayashi T, Honma K, Shingaki S, Nakajima T. Changes
in masticatory function after orthognathic treatment in
Braz J Oral Sci 1(1): 7-18 Human oral function: a review

patients with mandibular prognathism. Brit J Maxil Surg Inoue T, Masuda Y, Saito O, Mizuno J, Kato T.
2001; 39:260-265. Autoregulation of masticatory force in the anesthetized
96. van den Braber W, van der Glas HW, van der Bilt A, rabbit. In: Morimoto T, Matsuya T, Takada K, eds. Brain and
Bosman F. Chewing efficiency of pre-orthognathic oral functions. Amsterdam: Elsevier Science B.V.; 1995.
surgery patients: selection and breakage of food p.115-124.
particles. Eur J Oral Sci 2001; 109:306-311. 113. Ottenhoff FM, van der Bilt A, van der Glas HW, Bosman F.
97. Weijnen FG, van der Bilt A, Wokke JHJ, Wassenberg Peripherally induced and anticipated elevator muscle
MWM, Oudenaarde I. Oral functions of patients with activity during simulated chewing in humans. J
myasthenia gravis. In: Richman P, ed. Myasthenia Gravis Neurophysiol 1992; 67:75-83.
and related diseases. New York: The New York Academy 114. van der Bilt A, Ottenhoff FM, van der Glas HW, Bosman F,
of Sciences; 1998. p.773-776. Abbink JH, Weijnen FG. Neuromuscular control of
98. Weijnen FG, van der Bilt A, Wokke JHJ, Kuks JBM, van der simulated chewing in humans. In: Morimoto T, Matsuya T,
Glas HW, Bosman F. Maximal bite force and surface EMG Takada K, eds. Brain and oral functions.Amsterdam:
in patients with myasthenia gravis. Muscle & Nerve 2000; Elsevier Science B.V.; 1995. p.125-140.
23:1694-1699. 115. Abbink JH, van der Bilt A, Bosman F, van der Glas HW. A
99. Weijnen FG, van der Bilt A, Kuks JBM, van der Glas HW, comparison of jaw-opener and jaw-closing muscle activity
Oudenaarde I, Bosman F. Masticatory performance in in humans to overcome an external force counteracting jaw
patients with Myasthenia Gravis. Arch Oral Biol 2001; in movement. Exp Brain Res 1997; 118:269-278.
press. 116. Abbink JH, van der Bilt A, Bosman F, van der Glas HW.
100. Robinovitch SN, Hershler C, Romilly DP. A tongue force Speed-dependent control of cyclic open-close movements
measurement system for the assesment of oral-phase of the human jaw with an external force counteracting
swallowing disorders. Arch Phys Med Rehabil 1991; closing. J Dent Res 1999; 78:878-886.
72:38-42. 117. Slager GEC, Otten E, van Eijden TMGJ, van Willigen JD.
101. Weijnen FG, Kuks JBM, van der Bilt A, van der Glas HW, Mathematical model of the human jaw system simulating
Wassenberg MWM, Bosman F. Tongue force in patients static biting and movements after unloading. J
with myasthenia gravis. Acta Neurol Scand 2000; 102:303- Neurophysiol 1997; 78:3222-3233.
308. 118. Abbink JH, van der Bilt A, Bosman F, van der Glas HW,
102. van der Glas HW, Olthoff LW, van der Bilt A, Bosman F. Erkelens CJ, Klaassen MFH. Comparison of external-load
Control of elevator muscle activity during chewing in compensation during rhythmic arm movements and
man. Soc Neurosci Abstr 1987; 13:11. rhythmic jaw movements in humans. J Neurophysiol 1999;
103. Dellow PG, Lund JP. Evidence for central timing of 82:1209-1217.
rhythmical mastication. J Physiol 1971; 215:1-13.
104. Lund JP. Evidence for a central neural pattern generator
regulating the chewing cycle. In: Anderson DJ, Matthews
B, eds. Mastication. Bristol: John Wright and Sons; 1976.
p.204-212.
105. Nozaki S, Iriki A, Nakamura Y. Localization of central
rhythm generator involved in cortically induced
rhythmical masticatory jaw-opening movement in the
guinea-pig. J Neurophysiol 1986; 55:806-825.
106. Goldberg LJ, Chandler SH. Central mechanisms of
rhythmic trigeminal activity. In: Taylor A,ed.
Neurophysiologyof the jaws and teeth. Hong
Kong: The Macmillan Press ltd; 1990. p.268-293.
107. Lund JP. Mastication and its control by the brain stem.
Critical Reviews in Oral Biology and Medicine 1991; 2:33-
64.
108. Goodwin GM, Luschei ES. Effects of destroying spindle
afferents from jaw muscles on mastication in monkeys. J
Neurophysiol 1974; 37:967-981.
109. Morimoto T, Inoue T, Masuda Y, Nagashima T. Sensory
components facilitating jaw-closing muscle activities in
the rabbit. Exp Brain Res 1989; 76:424-440.
110. Thexton AJ. Some aspects of neurophysiology of dental
interest. I. Theories of oral function. J Dent 1973; 2:49-54.
111. Lavigne G, Kim JS, Valiquette C, Lund JP. Evidence that
periodontal pressoreceptors provide positive feedback to
jaw closing muscles during mastication. J Neurophysiol
1987; 58:342-358.
112. Morimoto T, Nakamura O, Ogata K, Liu ZJ, Matsuo R,

18