Neuroscience Letters 360 (2004) 137–140

www.elsevier.com/locate/neulet

A fronto-parietal network for chewing of gum: a study on human subjects

with functional magnetic resonance imaging
Toshihisa Takadaa,*, Tamaki Miyamotob
a
Department of Oral Functional Science, Graduate School of Dental Medicine, Hokkaido University, Kita-13, Nishi-7, Kita-Ku, Sapporo, 006-8586, Japan
b
Department of Physiology, Graduate School of Medicine, Hokkaido University, Sapporo, Japan
Received 3 December 2003; received in revised form 10 February 2004; accepted 23 February 2004

Abstract
The purpose of this study is to investigate human brain activity during mastication using fMRI. Twelve right-handed normal subjects
performed two tasks: chewing of gum at their own pace, and imitating the movements of chewing gum. In order to reveal which areas of the
brain are more strongly activated while chewing gum, we performed the conjunction analyses of gum chewing minus sham chewing with
gum chewing minus rest. The common activity in the orofacial sensorimotor and premotor cortex was subtracted out since it was common to
both tasks, but there were some differences in activity in some prefrontal and posterior parietal cortex areas. Our results suggest that a fronto-
parietal network for mastication exists and may contribute to higher cognitive information processing.
q 2004 Elsevier Ireland Ltd. All rights reserved.
Keywords: Functional magnetic resonance imaging; Mastication; Manipulation; Prefrontal cortex; Fronto-parietal network

Can chewing of gum improve brain performance? It is difficult
to answer this question. The masticatory movements activate
the brain; therefore, it has been generally said that the learning
effect is improved or dementia is prevented. Recently,
Wilkinson et al. discovered that chewing gum can improve
the memory of the person chewing it [14]. They suggest that
chewing of gum can improve episodic memory (involving the
learning, storage and retrieval of information) and working
memory (where information is held ‘on line’). These findings
have raised the possibility of a differential involvement of the
cortical regions related to human memory in the act of
mastication. When humans think about eating, higher-order
mental activities must be absolutely essential. For instance,
while eating we always have to recognize the features of foods
within the mouth and manipulate them adequately during
mastication. However, previous neuroimaging studies have
not shown that the cortical regions other than primary and
nonprimary motor areas were activated during mastication [7].
This result does not reinforce the idea that mastication needs
higher-order mental activities. Several neuroimaging findings
* Corresponding author. 9-22, 2-jo, 1-tyoume, Satozuka, Kiyota-Ku,
Sapporo, 004-0802, Japan. Tel.: þ81-11-889-5057; fax: þ 81-11-889-
5057.
E-mail address: takada@den.hokudai.ac.jp (T. Takada).
0304-3940/03/$ - see front matter q 2004 Elsevier Ireland Ltd. All rights reserved.
doi:10.1016/j.neulet.2004.02.052
provide the strongest evidence available to date from human
studies that the prefrontal cortex plays a role in working
memory function [5]. If ‘mastication’ results in increased
activity in the prefrontal cortex, this evidence may give a very
significant scientific basis to the general idea that ‘mastication’
improves brain performance. However, no one has demon-
strated a correlation between the activity in the prefrontal
cortex and chewing of gum previously. Moreover, little is
known about whether brain activity related to mastication and
mere jaw movements are different.
In the present experiment, 12 right-handed participants
between the ages of 20 and 28 years were tested. Six women
and six men participated in this study. None had any history
of neurological or psychiatric disorders. Informed consent
was obtained from each subject prior to the experiment. The
Committee of Medical Ethics, Hokkaido University,
approved the protocol of this study. We employed two
tasks: (1) a self-paced gum chewing task; (2) a sham
chewing task. In the gum chewing task, all subjects were
instructed to chew a piece of gum base (Lotte Co., Ltd.,
Tokyo, Japan) at their own, but constant, pace. During the
sham chewing task, the subjects were instructed to mimic
chewing movements, but they did not actually have a piece
of gum in their mouths. Both task sessions (gum chewing
task and sham chewing task) were designed in a block
138 T. Takada, T. Miyamoto / Neuroscience Letters 360 (2004) 137–140
manner (four rest and four task blocks, respectively, for 28
and 224 s for a single task session). All subjects participated
in both sessions in random order. Each trial began with the
rest block, followed by the task block (gum chewing or
sham chewing) at a given auditory signal. During the
experiment, the room lights were dimmed and the subjects’
eyes were closed. Brain imaging data were acquired on a GE
Signa 1.5 T whole-body MRI scanner using a head coil. For
functional imaging, we used a gradient-echo echo-planar
imaging sequence (GRE-EPI) with the following par-
ameters: FOV 24 £ 24 cm; matrix 64 £ 64; slice thickness
5 mm; slice gap 2.5 mm; TR 4 s; TE 40 ms. The spatial
resolution was 3.75 £ 3.75 £ 5 mm. In addition, 12 slices
were obtained from a single session. Sessions that showed
brain motion exceeding 0.375 mm were repeated to avoid
fictional activation due to pixel misalignment. After image
construction, fMRI time series data consisting of consecu-
tive EPI images for each slice were analyzed using SPM99
software (Wellcome Department of Cognitive Neurology,
London, UK). T1 anatomical images were coregistered to
the mean of the functional scans and aligned to the SPM T1
template in the Talairach space. The calculated nonlinear
transformation was applied to all functional images for
spatial normalization. Finally, the functional images were
smoothed with a 6 mm full width at half maximum
(FWHM) kernel, Gaussian filter. Contrasts were calculated
by fixed-SOA from the protocol as epochs and by
convolving them with the hemodynamic response function
(HRF) to specify the appropriate design matrix. Condition
and subject effects were estimated according to the general
liner model at each voxel in brain space. To minimize
effects of physiological noise, a high pass filter of 80 s was
applied within the design matrix. Moreover, a low pass filter
of Gaussian type (4 s) was also used. Specific effects were
tested by applying appropriate linear contrasts to the
parameter estimates for each condition, resulting in a
statistic for each and every voxel, while group data were
analyzed using a random effects model. The resulting
correlates were transformed into a Z score map. The
significant Z value was threshold at z . 3:09 (corresponding
to P , 0:001 at each voxel level). Only clusters . 10
activated voxels were reported. Activated brain structures
were identified by transforming the MNI coordinate into the
brain atlas of Talairach and Tournoux. In this study, we
undertook the following comparisons of the main exper-
iment: gum chewing task minus sham chewing task with
gum chewing task minus rest. The purpose of this
comparison is to reveal which areas of the brain are
relatively more strongly activated while chewing gum.
We found that the mean ^ SD frequency of the jaw
movements during the gum chewing task was 1.15 ^ 0.11
Hz, while the mean frequency was faster than that noted
during the sham chewing task (1.24 Hz).
The gum chewing task significantly activated the primary
sensorimotor cortex extending down into the upper bank of
the operculum and insula, the dorsal and ventral premotor
areas (PMA) bilaterally, the supplementary motor area
(SMA), the left inferior frontal cortex, the middle frontal
gyrus bilaterally, the intraparietal cortex bilaterally and the
superior parietal lobe (SPL) bilaterally. However, the SMA,
the inferior frontal cortex, and the middle frontal cortex
activation did not reach statistical significance (P , 0:05,
corrected), although an uncorrected significance level of
P , 0:001 was observed. The same areas in the primary
sensorimotor areas bilaterally, the dorsal and ventral PMA,
and the SMA were also significantly activated during the
sham chewing task (P , 0:005, corrected), however in the
inferior and middle frontal cortex no increase in activity was
observed (P , 0:001, uncorrected).
Then, to reveal which areas of the brain were relatively
more strongly activated while chewing gum, we performed
the conjunction analyses of gum chewing minus sham
chewing with gum chewing minus rest. This conjunction
analysis was applied by masking, whereby the second
subtraction was tested only in pixels that reached signifi-
cance (P , 0:001) in the first subtraction. This analysis
revealed significant activation in the middle frontal cortex
bilaterally (x ¼ 50, y ¼ 42, z ¼ 24 in Talairach coordinates
with the peak score of Z ¼ 5:78, x ¼ 244, y ¼ 58, z ¼ 14
with Z ¼ 5:62), the left inferior cortex (x ¼ 252, y ¼ 46,
z ¼ 214 with Z ¼ 7:34), the inferior parietal lobule
(x ¼ 42, y ¼ 266, z ¼ 50 with Z ¼ 3:83, x ¼ 240,
y ¼ 268, z ¼ 40 with Z ¼ 3:98) and the right superior
parietal lobule (x ¼ 18, y ¼ 280, z ¼ 50 with Z ¼ 4:84)
(Fig. 1).
Specific activations were obtained in the dorsolateral
prefrontal cortex, ventral prefrontal cortex, and parietal
cortex during the gum chewing task. In the present study
using conjunction analysis, the common activity in the
orofacial sensorimotor and premotor cortex was subtracted
out since it was common to both tasks. Consequently, we
hypothesize that the parieto-frontal network plays an
essential role in ‘mastication’.
Areas in the middle frontal gyrus and inferior frontal
gyrus were activated during the gum chewing task in this
study, but were not activated when the subjects merely
moved their jaw, tang, and oral mucosa rhythmically (sham
chewing task). Therefore, the role of these areas must be
involved in whether the gum is actually in the oral cavity.
The functional subdivision of the prefrontal cortex has
been an issue of much debate recently. A review by Fletcher
and Henson [5] more generally states the involvement of the
ventrolateral prefrontal area in processes such as updating
and maintenance of information. In contrast, processes such
as information manipulation activate the dorsolateral
prefrontal cortex [4]. Applying these concepts to our data,
we would like to argue that prefrontal activation during
mastication can be related to the manipulation of somato-
sensory information and the retention of stimulus related
information. Our results may extend recent models on the
prefrontal cortex and its contribution to higher cognitive
information processing as established in the verbal [2,6],
 T. Takada, T. Miyamoto / Neuroscience Letters 360 (2004) 137–140
Fig. 1. Surface projections of color-coded statistical parametric maps (SPMs) superimposed onto MNI standard brain. Activation revealed by gum chewing
minus sham chewing with gum chewing minus rest. Left and right sides represent the left and right hemispheres, respectively. The center represents an
overhead view. The threshold is set at P , 0:05, corrected.
visual [3], and visuospatial [11] domain by demonstrating
their relevance and applicability for the somatosensory
domain related to automaticity such as mastication.
However, further studies are needed to investigate the
precise role of these prefrontal areas. In particular, studies
that manipulate the degree of memory load required are
needed.
Additionally, an area in the inferior parietal lobule
showed activation. This result is consistent with the results
reported in a previous study indicating that the activation of
the inferior parietal lobule is related to tactile object
identification [1]. Furthermore, focal lesions in the inferior
parietal lobule are known to produce a selective deficit in
shape perception and representation with preserved lower
somatosensory functions, spatial abilities, and tactile
exploration behavior [9]. In addition, coactivation of the
ventral PMA and the inferior parietal lobule was observed.
Classically, the ventral sector of the frontal cortex was
thought of as an area controlling oro-laryngeal movements.
However, a series of recent studies showed that this view
only partially describes the function of the ventral PMA [8,
12]. These data suggest that, in addition to the control of
oro-laryngeal movements, the ventral PMA is of importance
for motor tasks with high motor execution demands [15].
Therefore, we suggest that coactivation of the parietal and
the premotor cortex indicates that these areas are part of the
cortical network involved in tactile information processing
during the mastication.
Finally, the SMA was activated during the sham chewing
task. The focus of SMA activation was probably located in
the pre-SMA [10,13]. However, the SMA showed slight
increases in activity (P , 0:001, uncorrected) during the
gum chewing task. Our findings may indicate that activity in
the cerebral cortex is different between mastication with
food and rhythmical jaw movements. However, the
contribution of the pre-SMA in mastication requires further
research.
139
In contrast to the present study, a previous neuroimaging
study gave negative results for the existence of a fronto-
parietal network subserving the masticatory movement [7].
We have demonstrated that the coactivation of the
prefrontal and the parietal cortex was observed with
conjunction analysis. As for these findings, we suggest
that our results may extend recent models on the prefrontal
cortex and its contribution to higher cognitive information
processing. Presumably, the tactile information from the
oral cavity, related to a piece of gum, may become a trigger
to recollect the ‘mastication’. However, it is unclear what
the mechanism underlying oral stereognosis and recognition
of the food bolus is, so further investigation is required.
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