ArchsoralBid.Vol. 35,No. 4,pp.

311-318,1990 0003-9969/90
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EFFE.CTS OF CHEWING FREQUENCY AND BOLUS

HARDNESS ON HUMAN INCISOR TRAJECTORY AND
MASSETER MUSCLE ACTIVITY

B. BISHOP,’ 0. PLESH’ and W. D. MCCALL JR~

‘Department o,! Physiology, School of Medical Sciences, State University of New York at Buffalo,
*Department of Restorative Dentistry, School of Dentistry, University of California, San Francisco, CA
94143 and ‘Department of Oral Medicine, School of Dental Medicine, State University of New York at
Buffalo, Buffalo, NY 14214, U.S.A.

(Accepted 21 September 1989)

Summary-Nine adults with no orofacial dysfunctions were instructed to chew a standardized piece of
soft or hard gum on the right side in time with a metronome set at 46, 100 or 160 beats/min. Jaw
movements were recorded with a Myotronics kinesiograph and masseter electromyograms were detected
with surface elelztrodes. The chewing patterns on either gum were not significantly different in any of their
spatial or temporal aspects, in mean or peak opening or closing velocities, or in the timing or level of
activity in either masseter at any of the three chewing frequencies. These findings suggest that during
metronome-paced chewing the change in sensory feedback resulting from a change in gum hardness exerts
little or no effel:t on either the spatial or temporal aspects of masticatory motor output.

Key words: chewing pattern, spatial and temporal aspects of chewing, jaw velocities, metronome-paced
chewing, bolus texture, mast&tory motor c&o1

INTRODUCTION sensory feedback as revealed by kinematic and

electromyogram measures of masticatory motor out-
Mastication, like locomotion and respiration, is an put.
automatic, rhythmical movement originated by
central pattern generators within the central nervous
MATERIALS AND METHODS
system, without need for peripheral feedback (Dellow
and Lund, 1971; Kubo, Enomoto and Nakamura, Our methods were similar to those described in
1981; Luschei and Goldberg, 1981). Nonetheless, the detail by Plesh ef al. (1986, 1987, 1988). Six female
central nervous system integrates incoming periph- and 3 male adults, free of orofacial pain and with a
eral signals with centrally generated command signals full dentition, were studied while sitting in a dental
so that the rhythmical motor output is appropriate chair with the head supported. Protocols had full
for the load imposed by a bolus. For example, during approval of the human subjects review board of the
spontaneous chewing the hardness of a chewing gum School of Dental Medicine; all subjects gave in-
strongly influences the spatial and temporal aspects formed consent. The bolus was a tasteless gum base,
of an individual’s chewing pattern (Plesh, Bishop and supplied by the William Wrigley Jr Co., and was of
McCall, 1986) and the concomitant masseter muscle two detectably different consistencies, soft and hard.
activity (Plesh, Bishop and McCall, 1987, 1988). Each piece of gum was of a standard size
Thus, subjects chew hard gum more slowly than soft (10 x 18 x 2 mm) and weight (0.7 f 0.1 g).
gum and the slowing is due to an increase in the A subject was instructed to chew a piece of gum on
duration of the opening and occlusal phase; the the right side for a minute or two in time with a
duration of the closing phase is not significantly metronome. The metronome was set in random order
altered despite prolongation and enhancement of at 46, 100 or 160 beats/min. Each subject chewed each
masseter activity. Therefore, during spontaneous type of gum in succession, with the soft and hard
chewing, peripheral feedback, signalling the proper- order randomized.
ties of the bolus, strongly influences the output of the Movements of the mandibular incisor were regis-
central pattern generators even though it is not tered with a kinesiograph (Myotronics model
essential for the maintenance of rhythmic chewing. KSAR). Activity of the right or working masseter
We have now compared the spatial and temporal muscle (i.e. ipsilateral to the bolus) was detected with
aspects of the chewing pattern, the mean and peak surface electrodes and recorded on tape for subse-
opening and closing velocities, and the level and quent computer analysis. In some subjects the
duration of masseter activity when a subject chews electromyogram of the contralateral masseter was
either soft or hard gum in time with a metronome set recorded as well.
at 46, 100 and 160 beats/min. Our goals were to
Data analysis
determine whether a change in gum hardness affects
metronome-paced ch#:wing and whether a change in Tape-recorded data, played back to a polygraph,
chewing frequency induced by changing the revealed that the chewing pattern usually stabilized
metronome beat motdifies the central processing of within 3 or 4 cycles regardless of the metronome
311
312 B. BISHOP et al.
setting or the gum hardness. The subsequent 30
consecutive chewing cycles were analysed by com-
puter. The analogue signals of jaw movement from
the kinesiograph and the electromyographic activity
from the right masseter muscle were digitized at
1 kHz. The data were smoothed, displayed on an
Hewlett-Packard plotter (7475A), and compared to
their analogue counterpart to verify that smoothing
had not distorted the signals.
Spatial aspects of the chewing pattern were
assessed from the vertical excursions of the mandible.
Peak and mean opening and closing velocities were
determined from the peak and mean magnitudes of
the instantaneous velocity on the vertical channel.
Temporal aspects of the chewing pattern were deter-
mined from the digitized vertical signal after calibra-
tion (Myo-tronics Memorandum, 1978). The
beginning of each cycle (i.e. the start of opening) was
determined from the differentiated vertical signal by
identifying the first data point in the differentiated
waveform after which all subsequent points were
positive for at least 100 ms. The total cycle duration
was the time between the beginning of one cycle and
the beginning of the next. End of opening was defined
by the datum point which occurred when the vertical
distance reached a maximum. End of closing was
defined by the first point in the differentiated wave
which was not negative and which was preceded for
at least 100 ms by negative values. The duration of
the occlusal phase was defined as the difference
between the end of closing in one cycle and the onset
of opening in the next.
Raw electromyogram activity of the masseter was
assessed to obtain the onset latency and the burst
duration of muscle activity for each chewing cycle.
Onset latency was the time between the beginning of
the closing phase and the onset of masseter activity.
The integrated or averaged electromyogram was
obtained by rectifying, scaling and averaging the
digitized electromyogram. The mean rise times of the
bursts of masseter activity, determined from the
averaged signal, provided indices for rate coding and
rate of recruitment. The mean and peak amplitudes
of the averaged electromyograms provided indices for
the mean and maximum levels of muscle activity,
respectively
“-(A)
‘ii
E
E
E
15-
lo-
.
I(81
;/I
-/ I/ : ./ .
M%-ikTk-
0 5
Gape
Fig. 1. Mean (n = 30) vertical gapes for each of the 9
and hard gum in time with a metronome set at 46 (A), 100 (B) or 160 (C) beats/min. Lines of unit slope
are shown. Symbols denote individual subjects: No. 1 (open circle), No. 2 (filled circle), No. 3 (open
triangle), No. 4 (filled triangle), No. 5 (open square), No. 6 (filled square), No. 7 (open inverted triangle),
No. 8 (filled inverted triangle), No. 9 (open diamond).
Statistical analysis
Means and standard deviations for each variable
defined above were determined over the 30 cycles for
each subject. These averaged data were subjected to
paired or unpaired t-tests, and to two-way and
three-way within-subjects analyses of variance
(ANOVA) as appropriate.
RESULTS
EfSects of chewing frequency and gum hardness on
spatial aspects of the chewing pattern
Figure 1 shows the means of the peak vertical
gapes; differences between these means on either gum
were not statistically significant in any subject at any
of the chewing frequencies. In fact, most data points
fell closely about the lines of unit slope indicating that
gum hardness had little or no effect on gape.
Comparison of the range of vertical gapes in (A),
(B) and (C) of Fig. 1 suggested that an increase in
chewing frequency might be associated with a de-
crease in gape. This was confirmed by a two-way
ANOVA for repeated measures which showed that
(1) gape decreased significantly as chewing frequency
increased, (2) gape was not altered significantly by
gum hardness, and (3) the interaction between chew-
ing frequency and gum hardness was not significant.
Inspection of individual’s data suggested that the
motor strategy adopted to pace chewing with the
metronome beat varied. Some subjects maintained a
relatively stable vertical gape across all chewing
frequencies; others significantly changed gape when
pacing their mandibular movements. Examples of
these two strategies are shown in Fig. 2.
Subject No. 7 [Fig. 2(A)], when chewing at 46
beats/min, had a mean gape of 15.6 k 1.8 mm for soft
gum and 15.4 f 1.7 mm for hard gum. The means
when chewing at 100 (soft: 15.8 k 1.9; hard:
15.0 f 2.1) and 160 beats/min (soft: 17.4 & 1.6; hard:
15.6 f 1.7) were not significantly different from those
at 46 beats/min regardless of the texture of the gum.
Findings were qualitatively similar in 3 other subjects
(Nos 2, 3 and 4).
Subject No. 5 [Fig. 2(B)] had smaller mean vertical
gapes at every chewing frequency than subject No. 7
[Fig. l(A)]. In addition, means of peak gape were less
0 0 0 .0
l
10 20 0 20
on soft gum I mm1
subjects, indicated by symbol, while chewing soft
 Metronome-paced chewing patterns 313

18r(A) Figure 3 shows the mean total cycle duration for

15 each subject. The means in Fig. 3(A) range from 900
to 1460 ms; a subject chewing exactly in time with the
z 12
metronome (i.e. at 46 beats/min) would have had a
- 9 mean of 1304 ms. Although the mean cycle duration
$
8 6 while chewing soft gum was seldom identical to that
for hard gum in any subject, the differences between
3
the means for the 2 gums were not statistically
significant @ > 0.05) at any chewing frequency.
4.6 I$0 Figure 4 shows the group means of the total cycle
Metronome setting ( beots/min) duration. This duration, of course, had to decrease
when the chewing rate was increased; the decreases in
Gum type cycle duration were due to decreases in the durations
of each phase in the chewing cycle as shown in (B),
u Soft
(C) and (D) in Fig. 4. Note that a change in gum
m Hard hardness had no effect on any temporal aspect of the
chewing cycle.
Effect of chewing frequency andgum hardness on mean
and peak velocities of jaw excursions
Figure 5 shows the mean opening velocities and
mean closing velocities on hard or soft gum. With few
exceptions the data points cluster about the lines of
46 100 160
unit slope, suggesting that gum hardness had little or
Metronorre setting (beats/min)
no effect on either mean opening or closing velocity
regardless of the chewing frequency.
Fig. 2. Examples of two strategies used to keep chewing As might be predicted, in subjects who maintained
rhythm in time with the metronome. Mean peak vertical
gape across different chewing frequencies [i.e. subjects
gape of jaw excursions for subject No. 7 (A) and No. 5 (B)
when chewing soft gum (open bars) and hard sum (hatched No. 2 (filled circle), No. 3 (open triangle), No. 4 (filled
bars) at 3 different metronome settings (i.e. beats/min). triangle) and No. 7 (open inverted triangle)], both
mean opening and mean closing velocities tended to
increase with an increase in chewing frequency,
when this subject chewed at 100 (soft: 8.5 f 2.0; hard: whereas in subjects who reduced gape during fast
6.0 f 2.4) and at 160 beats/min (soft: 6.8 f 1.4; hard chewing [i.e. subjects No. 1 (open circle), No. 5 (open
3.5: + 1.2) than at 46 beats/min. Qualitatively similar square), No. 6 (filled square), No. 8 (filled inverted
data were obtained in 4 other subjects (Nos 1, 6, 8 triangle) and No. 9 (open diamond)], mean opening
and 9). and mean closing velocities either did not change or
tended to decrease with changes in chewing fre-
Eflects of chewing _/Yequency and gum hardness on quency.
temporal aspects of !he chewing pattern Gum hardness was without effect on the spatial or
As reported by Plesh et al. (1987), subjects varied temporal aspects of the chewing pattern when chew-
in their ability to chew in time with a metronome and ing frequency was metronome-paced; all velocity data
rarely, if ever, matched the duration of their chewing have been pooled in Fig. 6. Note that the data points
cycle exactly to that of the metronome beat. Even in for the opening velocities fall about a line whose slope
those subjects who followed the metronome quite is 10, whereas the points for closing velocities fall
closely, the duration of each phase in the cycle varied about a line whose slope is 5. In other words, during
considerably from cycle to cycle whether they chewed opening, the peak velocity is 10 times higher than the
soft or hard gum. mean opening velocity, whereas the peak closing

OX----
1.4 1.6 0.3 0.4 0.5 0.6

Total cycle duration on soft gum (s1

Fig. 3. Means (n = 30) for total cycle duration for each of the subjects while chewing the hard and soft
gums at 46 (A), 100 (B) and 160 (C) beats/min. The dashed line at 1.3 s in (A) indicates the cycle duration
if a subject had chewed exactly in time with the metronome. Symbols for subjects are as in Fig. 1.
314 B. BISHOP et al.

Gum type 600

(A)
0 Soft 2

-
46 100 160 46 100 160

600
2
E
P 450

‘?I
9
u 300
‘i;
s
i:,o 150

L
2

0
46 100 160

Metronome setting ( beats / min I

Fig. 4. Group means for the total cycle duration, and for the opening, closing and occlusal phases for
the 9 subjects while chewing soft (open bars) and hard (hatched bars) gum at the 3 metronome settings.

(A) (B) (Cl

.
.
Vm

:A 0
q
0s
.
I- I I
5 10 0 5 1” 5 10

Opening velocity on soft gum (cm/s I

E a- V
s 0
E
e 6-
s . 0
.
” - ‘A

kj\“4- 0 . vA
0
.
4 E
u
E” .
0
4 2- .
OV
”

I I I I
0 2 4 6 6 0 2 4 6 6

Closing velocity on soft gum 1cm/s )

Fig. 5. Upper panels: mean opening velocity while chewing hard gum (ordinate) versus mean opening
velocity on soft gum (abscissa) for the 9 subjects while chewing at 46 (A), 100 (B) and 160 (C) beats/min.
Lower panels: analogous plots for mean closing velocity. Symbols for subjects as in Fig. I. (Note
differences in scale on the axes between the upper and lower panels.)
 Metronome-paced chewing patterns 315

Effects of chewing frequency and gum hardness on

masseter activity
Burst durations. Figure 8 shows the mean burst
Shpe = 10 duration of the right and left masseter muscles for

1
x
P
one subject. At each metronome setting, the burst
a 200 duration of the non-working muscle (i.e. contra-
lateral to the bolus) was longer than that of the
working muscle. Other subjects showed a similar
I I I I pattern, but the differences between the group means
20 40 for burst duration of the 2 masseter muscles were not
Mean statistically significant at any chewing frequency.
Burst durations of both masseter muscles were
shortened in proportion to the shortening of the total
cycle duration when chewing was paced at 100 or 160
beats/min. In fact, masseter activity filled the same
proportion of each chewing cycle regardless of the
absolute total cycle duration, as pointed out by Plesh
et al. (1987). A change in gum hardness had no
significant effect on the masseters’ burst duration at
any of the chewing frequencies.
Onset latencies. The absolute value of onset latency
for the working masseter varied markedly among
I I I I subjects at every chewing frequency (Fig. 9). Mean
0 40 80 onset latencies on soft gum were not different from
Mean those on hard gum. With each increment in chewing
Fig. 6. Upper panel: peak versus mean opening velocities for frequency, onset latencies become significantly
all data. Lower panel: peak versus mean closing velocities shorter. In some subjects [e.g. No. 3 (open triangle),
for all data. No. 4 (filled triangle) and No. 6 (filled square)], when
chewing at 160 beats/min, the masseter became active
velocity exceeds the ‘mean closing velocity only five- prior to jaw closing. In all other cases masseter
fold. activity started after the start of jaw closing showing
As shown by the data in Fig. 7, mean closing that the masseter muscles do not initiate closing.
velocity was always higher than mean opening veloc- Masseter electromyogram activity. Figure 10 shows
ity, but peak closing and peak opening velocities were the mean peak amplitudes and the mean rise times of
not different. the integrated electromyograms from the working
masseter muscle for each subject. There was wide
inter-subject variability for each response. A two-
way, within-subject ANOVA revealed no significant
differences between group means for the 2 gums for
either the peak amplitudes or the rise times. Thus,
neither a change in chewing frequency nor a change
in gum hardness caused a predictable change in the
level or timing of recruitment in the masseter muscles.

600
0 soft gum
r
;; 500 m Non-workmg
t
z Mean dosing velocity (mm /s )

46 100 l&O
Metronome setting
g
!?! Oo% 2bo 3bo 4bo 5do
(beats/min)
Peak dosing velocity (mm 1s) Fig. 8. Mean burst durations of masseter activity in the
Fig. 7. Upper panel: mean opening versus mean closing working (open bars) and non-working (solid bars) muscles
velocity. Lower panel: peak opening versus peak closing of subject No. 1 when chewing the soft and hard gums in
velocity. Lines of identity are shown. time with the 3 metronome settings.
316 B. BISHOP et al.

‘O” (A) . 3oo (Cl

‘-E- VI 225
: E 600 c F

0 200 400 600 800 -50 50 150 250 350

onset latency on soft gum (Ins1

Fig. 9. Mean onset latencies for the right (i.e. working) masseter muscle while chewing hard and soft gums
at the 3 metronome settings. Symbols as in Fig. 1.

DISCUSSION These findings are in marked contrast to those

from earlier studies in which the subject’s attention
We have shown that when chewing is in time with was distracted from chewing by having him view a
a metronome, a change in the hardness of the gum video tape while chewing the soft and hard gums
being chewed has no effect on any of the spatial or (Plesh et al., 1987, 1988). Under these conditions
temporal aspects of the chewing cycle. The mean when a subject chewed at his automatic or preferred
gapes (Fig. l), total cycle duration (Fig. 3) and the rate, the change in gum hardness significantly
duration of phases within the chewing cycle (Fig. 4) modified the chewing pattern. For example, the open-
were not significantly different for either gum at any ing and occlusal phases, and hence the total cycle,
metronome setting. Similarly, the mean and peak were significantly longer when chewing hard gum,
velocities during opening and closing were not whereas the duration of the closing phase was not
modified by the change in gum hardness (Fig. 5). different on the 2 gums. The change in gum hardness
Furthermore, the modification of peripheral feedback did not significantly modify the vertical gape even
caused by changing gum consistency resulted in no though burst duration and level of masseter activity
significant change in the mean burst durations (Fig. were significantly greater on the hard gum (Plesh et
8), onset latencies (Fig. 9) or level or rate of recruit- al., 1986).
ment of masseter activity (Fig. 10). Thus, an alter- The fact that a change in gum hardness exerts
ation in peripheral feedback had little effect on either different effects on the chewing pattern, depending on
chewing pattern or masseter activity when the whether the movement is automatic or metronome-
chewing rate was dictated by the metronome. paced, raises some interesting, but as yet unanswered

Peak amplitude on soft gum (AU)

)_I 3,6yy2~;i;r:/, ‘, j_. , I;

0 100 200 300 0 100 200 300 0 100 200 300

Rise tome on soft gum 1 msl

Fig. 10. Upper panel: mean peak amplitudes of the integrated electromyograms (in arbitary units, AU)
recorded from the right masseter muscle during chewing hard gum versus those while chewing soft gum
on the right side in time with the metronome. Lower panel: analogous plots for mean rise times of the
integrated electromyograms. Symbols as in Fig. I.
 Metronome-paced chewing patterns
questions. Metronome-paced chewing imposes a con-
straint on the duration of jaw movements; automatic
chewing does not. Does one neuronal network serve
both tasks? If so, the peripheral influences must be
‘gated’ during metronome-paced chewing. Perhaps
the same neural mechanisms are used differently in
the 2 tasks.
During automatic chewing the rhythmicity and
mandibular trajectory are controlled by central pat-
tern generators in the brain stem and pontomedullary
region of the medial reticular formation (Chandler
and Goldberg, 1982; Enomoto, Katakura and Naka-
mura, 1983; Katoh er al., 1982; Kubo et al., 1981;
Nakamura and Kubo, 1978; Nozaki, Iriki and Naka-
mura, 1986). Whether metronome-paced chewing
and automatic chewing use the central pattern gener-
ators in identical ways remains to be determined.
During metronome-paced chewing the subject is
required to control his jaw movements at rates below,
close to and above the set point used for automatic
chewing. The descending motor command for this
task could exert its control over the masticatory
effecters by way of the central pattern generators, so
ignoring intra-oral feedback, or possibly the central
pattern generators are by-passed with the motor
commands being conducted directly to the final com-
mon pathway (i.e. the trigeminal motoneurones in the
nucleus of V).
Rhythmical jaw movements can be induced by
mechanical or electrical stimuli applied to the lips of
the decerebrate rat pup (Thexton and Griffiths, 1979),
by electrical stimulation of the masticatory area of
the cerebral cortex in various animals (Chandler,
Goldberg and Lambert, 1985; Lund and Dellow,
1971; Lund, Rossignol and Murakami, 1981; Naka-
mura et al., 1976; Nozaki et al., 1986; Sumi, 1969,
1977), or by non-rhythmical repetitive stimulation of
the pontine pyramid.al tract in the isolated brain stem
of the guinea pig (Nozaki et al., 1986) or decerebrate
rabbit (Lund and Dellow, 1971). During cortical
stimulation in the cat (Kubo ez al., 1981; Nakamuro
and Kubo, 1978) or guinea pig, jaw-closer moto-
neurones undergo alternating excitatory and in-
hibitory phases, whereas jaw-opener motoneurones
undergo only rhythmical excitatory phases (Chandler
and Goldberg, 1982). Transection of the brain stem
at the pontobulbar junction abolishes rhythmical
masticatory activity in the masseter motoneurones
(Enomoto et al., 1983), whereas cortical excitatory
effects on masseter motoneurones persist (Enomoto
et al., 1983; Katoh et al., 1982). These observations
suggest that the cortex has access to the masseter
motoneurones both directly and by way of the central
pattern generators.
Lund et al. (1984.) have demonstrated that at least
three different forms of mastication (incisal gnawing,
vertical mastication, and the ruminatory type) are
represented in separate zones of the rabbit sensori-
motor cortex. These masticatory forms, first defined
by Bremer in 1923, are assumed to be generated by
brain-stem circuits in a way analogous to that
produced by spinal circuits for scratching, walking
and galloping. Thus, brain-stem neural circuitry sub-
serving the masticatory rhythm generators appears to
be endowed with considerable versatility depending
upon the cortical zone from which they receive
317
activation. The fact that the nervous system can
activate the same muscles using different components
of the central motor system shows that different
motor systems converge on the same motor pools
(Chez, 1985). Perhaps direct corticotrigeminal path-
ways from the motor cortex converge on premotor
neurones that may or may not be components of the
masticatory pattern generators in order to exert
control over voluntary jaw movements. No informa-
tion is at present available on this point.
Nearly a century ago, Hughlings Jackson proposed
different hierarchical levels of motor control (Jack-
son, 1931). The highest levels involve the prefrontal
cortex, the middle levels involve the pre- and post-
central (i.e. the primary sensorimotor) cortex, and
the lowest levels mediating automatic control involve
the brain stem, the spinal cord or both. Lund et al.
(1984) have speculated that “alterations in the level
of cortical activity appear to be capable of changing
specific components of the chewing pattern”.
Although high-level cortical influences may play a
role in automatic chewing, they are not essential, as
shown by rhythmical jaw movements in decerebrate
preparations (Thexton, Griffiths and McGarrick,
1980) and in ancephalic infants (Monnier and Willi,
1947).
In our experiments, with each resetting of the
metronome, subjects had to change their chewing rate
to keep jaw movements paced to the auditory signal.
This limitation imposed by movement rate made it
unpredictable which way a subject would alter their
jaw movement to satisfy the constraint. Some
subjects retained gape and altered velocity, others
reduced gape while maintaining velocity (see exam-
ples in Fig. 2). Regardless of the strategy selected, the
duration of every phase in the chewing cycle was
altered with an alteration in chewing frequency and
the magnitudes of these alterations in phase durations
were independent of the gum’s hardness. We assume
that this reflects a cerebral resetting of the central
motor command.
Our findings do not negate the importance of
peripheral feedback during metronome-paced
chewing. In fact, we have shown previously that
cycle-to-cycle variability in all chewing parameters is
significantly less during metronome-paced chewing
than during automatic chewing (Plesh et al., 1988).
This suggests that peripheral feedback serves the
important function of resisting fluctuations in
chewing performance (Goodwin and Luschei, 1974;
Thexton, Hiiemae and Crompton, 1980; Lund and
Olsson, 1983).
Our data thus show that an alteration in peripheral
feedback has little or no effect on metronome-paced
chewing. The exact neural networks involved in such
chewing remain to be defined. They are no doubt also
involved in other voluntarily controlled jaw move-
ments including speaking, shouting, singing and
laughing. If this assumption is correct, then it appears
that the neural circuitry controlling voluntary
mandibular movements is less affected by peripheral
feedback than that involved in the control of auto-
matic chewing.
Acknowledgement-This study was supported by NIDR
Grant DE-06717.
318 B. BISHOPet al.
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