DR.

KENJI FUEKI (Orcid ID : 0000-0002-5885-2447)

Article type : Original Article

Accepted Article
Adaptive change in chewing-related brain activity while wearing a palatal plate: An fMRI study

1 1 2 2 1
Yuka Inamochi , Kenji Fueki , Nobuo Usui , Masato Taira , and Noriyuki Wakabayashi

1
Removable Partial Prosthodontics, Department of Masticatory Function Rehabilitation, Graduate

School of Medical and Dental Sciences, Tokyo Medical and Dental University, Tokyo, Japan.
2
Department of Cognitive Neurobiology, The Center for Brain Integration Research, Graduate School

of Medical and Dental Sciences, Tokyo Medical and Dental University, Tokyo, Japan.

Correspondence to: Kenji Fueki

Removable Partial Prosthodontics, Graduate School of Medical and Dental Sciences, Tokyo Medical

and Dental University, 1-5-45 Yushima, Bunkyo-ku, Tokyo, Japan.

Phone +81-3-5803-5514 FAX +81-3-5803-5514

kunfu.rpro@tmd.ac.jp

Article category: Original research

Running head: Adaptive change in chewing with palatal coverage

Conflict of interest: none

Key words: adaptation, mastication, functional magnetic resonance imaging, motor learning, palatal

plate, putamen
This article has been accepted for publication and undergone full peer review but has not been through
the copyediting, typesetting, pagination and proofreading process, which may lead to differences
between this version and the Version of Record. Please cite this article as doi: 10.1111/joor.12541

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 Abstract
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Prosthodontic treatment success depends on patients’ ability to adapt to an altered oral environment

containing removable prostheses. We investigated adaptive chewing-related brain activity changes in

response to a new oral environment. Twenty-eight fully dentate subjects (mean age: 28.6 years) wore

experimental denture-base palatal plates (3-mm thick), for 7 days. We measured food mixing ability and

cycle time, and assessed brain activity by functional magnetic resonance imaging during chewing at

pre-insertion (Day 0), and immediately (Day 1), 3 days (Day 3), and 7 days (Day 7) after insertion. Food

mixing ability significantly decreased and cycle time increased on Day 1 as compared to Day 0 (P <

0.001), and tended to recover to Day 0 level by Day 7. Brain activation in the right face primary

sensorimotor cortex and putamen significantly decreased on Day 1 as compared to Day 0 (P < 0.001),

and recovered to Day 0 level by Day 7. Brain activation in the left face primary sensorimotor cortex,

putamen, anterior cingulate gyrus (ACG), and right posterior medial frontal cortex (pMFC) significantly

decreased on Day 1 as compared to Day 0 (P < 0.001), and did not recover by Day 7. Thus, oral

environment changes involving palate covering affected chewing and induced adaptive brain activity

changes in the face primary sensorimotor cortex and putamen, possibly associated with motor learning.

Since ACG and pMFC activity remained unrecovered by 7 days after plate insertion, automatization of

chewing while wearing a palatal plate may require longer adaptation periods.

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 Introduction
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Prosthodontic treatment success depends on patients’ ability to adapt to an altered oral environment

that contains removable prostheses (1). Covering the palate with removable dentures is essentially

uncomfortable for patients and they have difficulty habituating to using such dentures (2), although

maxillary dentures often cover the palate to obtain adequate denture retention and stability (3).

Behavioural studies have shown that palatal coverage impaired masticatory performance (3, 4) and

bolus formation (4), and that adaptive changes occurred in 1 week. These findings suggested that

adaptation to palatal coverage during chewing is an important factor in adaptation to removable denture

use; however, the underlying mechanism remains unclear.

Some studies have indicated that changes in the oral environment could induce cortical adaptive

changes. For example, tooth extraction induced neuroplastic changes in the face primary motor cortex

in rats (5). In humans, differential neuronal activity involved in chewing and clenching occurred in

patients with implant-supported fixed dentures, implant-supported overdentures, and complete

dentures (6, 7). The dental arch length of dentures elicited different patterns of brain activity in the

middle frontal gyrus (8). Furthermore, the insertion of new dentures induced changes in precentral and

postcentral gyrus activation (1). Thus, cortical adaptive changes in chewing might also occur during

adaptation to newly inserted appliances.

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 Mastication is a rhythmic motor behaviour generated by central pattern generator (9). Additionally,
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the central nervous system coordinates these motor programs by sensory feedback from the peripheral

nerves (9). Several areas, such as the primary sensorimotor areas, supplementary motor area,

prefrontal cortex, insula, thalamus, basal ganglia, anterior cingulate cortex, and cerebellum are

activated during gum-chewing (10, 11). Therefore, we hypothesized that adaptive changes in the

activation of brain regions associated with chewing behaviour would occur in response to chewing with

palatal coverage. Such coverage may mask the input from the palate and prevent the tongue from

moving as usual. The decrease in these orofacial afferent inputs during mastication may induce

decreased activation of the relevant cortical areas, and such activation would recover during the

adaptation period. Thus, this study investigated adaptive changes in chewing-related brain activity in

response to palatal coverage.

Materials and methods

Subjects

Experience of denture use affects adaptability to new dentures (12). Therefore, to control the denture

experience among subjects as a confounder when investigating adaptive change in brain activity to a

new oral condition, we recruited fully dentate subjects, who had no experience of removable dentures.

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 A previous study indicated that more than 20 subjects are needed to obtain sufficient statistical power
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to detect a small effect in an fMRI neuroimaging study (13). We also referred to the sample size in fMRI

studies on human chewing (n = 17 [10], n = 29 [11]). Based on these studies, we decided to include 28

subjects in this study after considering drop-out due to artefacts on fMRI images caused by head

movement during chewing.

Right-handed subjects, who had individual normal occlusion and had no decayed teeth or teeth

under treatment, were recruited from among the students and clinical staff of Tokyo Medical and Dental

University. Subjects who had past experience of using removable dentures, who were currently

undergoing orthodontic therapy, who had temporomandibular disorder symptoms or psychiatric or

neurological diseases, who had undergone glossectomy or had tongue diseases, or who had oro-facial

pain were excluded. Written informed consent was obtained from each subject. This study was

approved by the Ethics Committee of Tokyo Medical and Dental University (Approval No. 1219).

Experimental palatal plate

A resin plate made of pour-type denture-base acrylic resin (PROCAST DSP, GC, Tokyo, Japan) was

used for full-coverage of the palatal area; plate thickness was 3.0 mm, as for conventional complete

dentures. Wrought wire clasps (0.9-mm diameter) made of cobaltchrome alloy were placed on

bilateral second molars to retain the palatal plates. The wire was adjusted to have no contact with the

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 opposing teeth during occlusion, prior to plate insertion. In a preliminary experiment, we confirmed that
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these wires created no artefact on brain images. Subjects were asked to wear the palatal plates for 1

week, except when sleeping.

Behavioural experiment

We used a colour-changeable Masticatory Performance Evaluating Gum XYLITOL (Lotte, Tokyo,

Japan) as the test food for evaluating food mixing ability (14). Subjects were seated on a chair in a

relaxed position and chewed the gum for 60 strokes, naturally, and at their own pace. The colour

measurement of the chewed gum and the calculation of ⊿E were performed as described previously

(15); a higher ⊿E score indicated greater food mixing ability. We recorded the chewing time during the

chewing test and calculated cycle time (CT) (s). Measurements were repeated three times at 1-minute

intervals. These experiments were performed at pre-insertion (Day 0), immediately post-insertion (Day

1), and on 3 days (Day 3) and 7 days (Day 7) post-insertion. We calculated the individual data of ⊿E

and CT for each experimental day.

functional magnetic resonance imaging (fMRI)

Subjects lay supine on the scanner table with their head immobilized with foam pads and straps around

the forehead. The task paradigm, which was a blocked design, began with 18 s of rest followed by six

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 sets of alternating 18-s tasks and 18-s rests. Visual stimuli were projected on a small screen attached
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to an MRI head coil by a DLA-10 projector (JVC, Yokohama, Japan) and a CF-T4GWKAXP personal

computer (Panasonic, Osaka, Japan). The stimulus presentation was controlled by E-Prime 2.0

software (Psychology Software Tools, Pittsburgh, USA). During the rest period, a white fixation cross

was displayed on the centre of the black screen for 18 s, and subjects were instructed to relax the jaw

and focus on the fixation point without chewing. After the rest period, the fixation cross disappeared and

a yellow character indicating “chewing” was shown on the screen for 18 s. Subjects chewed the

odourless, tasteless gum (GC, Tokyo, Japan) naturally, at their own pace, during the task period. The

volume of the gum was similar to that used for the behavioural experiment. The measurements were

performed on same days as behavioural experiment.

Magnetic resonance images were acquired using a MAGNETOM Spectra 3T scanner (Siemens,

Erlangen, Germany) with a T2*-weighted gradient-echo echo-planar imaging (EPI) sequence (192-mm

field of view; matrix: 64 × 64; 34 axial slices [3-mm thickness, 0.75-mm gap]; 2000-ms repetition time;

30-ms echo time; 77° flip angle; 3 mm × 3 mm × 3 mm voxel size). The initial nine volumes of each

session were dummy scans, to allow for magnetisation stabilisation. High-resolution T1-weighted

images were obtained as anatomical references (250-mm field of view; 192 sagittal slices [1-mm

thickness]; 1900-ms repetition time; 2.42-ms echo time; 1 mm × 1 mm × 1 mm voxel size).

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 Image analyses were performed using Statistical Parametric Mapping software SPM12
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(Wellcome Trust Centre for Neuroimaging, http://www.fil.ion.ucl.ac.uk/spm/software/spm12/). Prior to

statistical analysis, all images were realigned to the first volume to correct for head motion and the

mean of the realigned images was generated. The middle time-point slice acquired at 1000 ms in each

scan was chosen as the reference slice for the slice-acquisition timing correction. The T1-weighted

image was coregistered to the mean EPI image and transformed to the standard Montreal Neurological

Institute (MNI) space. Functional data were then normalised using the same transformation parameters

and data were spatially smoothed with an 8-mm full-width at half-maximum. The low noise frequencies

were removed by high-pass filtering (128-s cut-off). Voxel-based statistical analysis was performed for

each subject on each experiment day, using a general linear model with predictors for each task block,

convolved with a canonical haemodynamic response function and six nuisance variables for head

motion. A contrast image representing brain activation during the chewing task relative to baseline was

generated through each individual analysis.

All individual contrast images for each experiment day were also analysed as a group, using a

second-level random-effects procedure, using within-subject one-way analysis of variance on each

contrast. We computed statistical activation maps for the chewing task relative to baseline. The

threshold was set as P < 0.001 familywise error (FWE)-correction.

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 To compare brain activation between experimental days, subtraction analysis was performed
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between the statistical activation maps of each day. The voxel-level threshold was set at P < 0.001,

uncorrected for multiple comparisons, and the cluster level was set at P < 0.05 uncorrected for multiple

comparisons. Brain regions were anatomically defined and labelled according to a Tarairach atlas (16).

The beta-values from regions-of-interest (ROIs) in the brain were extracted using the Mars Bar software

(http://marsbar.sourceforge.net/).

Statistical analysis

We compared behavioural data and beta-values from ROIs in the brain across time-points, using a

linear mixed model analysis with post-hoc Bonferroni corrections for multiple comparisons. Significance

was set at P < 0.05. SPSS Statistics Version 20 (IBM, Tokyo, Japan) was used for statistical analysis.

Results

Twenty-eight subjects (15 male, 13 female; mean age ± standard deviation: 28.6 ± 2.5 years) were

enrolled in the study and completed all measurements. One subject exhibited head movement

exceeding 1 voxel in fMRI measurement. The motion artefact was observed on the image, and the

subject was thus excluded from further analyses of brain activity.

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 Food mixing ability and cycle time
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⊿E decreased and CT increased on Day 1 over that on Day 0, but both values tended to recover to the

Day 0 level by Day 7 (Fig. 1). The linear mixed model analysis found a significant time effect on ⊿E

and CT (P < 0.001) (Table S1). Post-hoc multiple comparisons of ⊿E found significant differences

between Day 0 and Day 1, Day 1 and Day 7, and Day 0 and Day 3 ( P < 0.05). Significant differences in

CT were also found between Day 0 and Day 1, Day 1 and Day 3, and Day 1 and Day 7 (P < 0.001).

Brain activity

Fig. 2 shows significant activity during gum-chewing on each experimental day. The bilateral

sensorimotor cortex, cerebellum, putamen, thalamus, superior frontal gyrus, and cuneus were the

regions commonly significantly activated at all time-points (P < 0.001, FWE-corrected).

We directory compared the activation on Day 0, with the highest ⊿E, with that on Day 1, with the

lowest ⊿E (Table 1, Fig. 3). Voxels in the right superior frontal gyrus, anterior cingulate gyrus (ACG),

bilateral precentral gyrus, and bilateral putamen, which may be involved in mastication, and the left

posterior cingulate gyrus, were significantly more activated on Day 0 than on Day 1 (P < 0.001). No

voxels were significantly activated on Days 1 and 7 as compared to Day 0, while voxels in the white

matter revealed significant activation on Day 3 as compared to Day 0 (P < 0.001).

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 To evaluate the activation changes in these six regions during the first week post-insertion, we
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compared activation on Day 0 with that on Days 3 and 7. Voxels in the right superior frontal gyrus,

bilateral putamen, and right precentral gyrus showed more significant activation on Day 0 than on Day

3 (P < 0.001). Furthermore, there was significantly more activation in the right superior frontal gyrus on

Day 0 than on Day 7 (P < 0.001).

We selected six regions that showed significantly more activation on Day 0 than on Day 1 for ROI

analysis. All ROIs were defined in MNI space as peak coordinates of activity and the beta-values from

these ROIs were extracted for each time point. The linear mixed model analysis found a significant time

effect on the beta-values of all six ROIs (P < 0.05) (Fig. 4, Table S1). Post-hoc analysis showed that the

beta-values of all six ROIs on Day 3 were significantly lower than those on Day 0, and were also

significantly lower on Day 7 than on Day 0, except for the left putamen (P < 0.05).

The beta-values of the right primary sensorimotor cortex on Days 3 and 7 and of the right

putamen on Day 7 were significantly higher than on Day 1 (P < 0.05). The beta-values of the left

primary sensorimotor cortex, left putamen, ACG, and pMFC on Days 3 and 7 were greater than on Day

1, but the values for Days 3 and 7 were not significantly different from that on Day 1 (P > 0.05).

Discussion

Here, the brain activation in the primary sensorimotor cortex, putamen, ACG, and pMFC during

chewing decreased immediately after palatal plate insertion and the activation in the primary

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 sensorimotor cortex and putamen tended to increase by 7 days after insertion. ⊿E decreased and CT
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increased immediately after plate insertion, indicating chewing impairment, and recovered to

pre-insertion level by 7 days post-insertion. Thus, the new oral environment induced by covering the

palate with a plate affected the changes of brain activity during chewing and chewing behaviour within 7

days.

The brain areas activated during gum-chewing on each experimental day (Fig. 2) were similar to

those previously identified (10, 11). The voxels of the primary sensorimotor cortex activated in this

study were found in the oral region, which previously appeared to be associated with a chewing-specific

process (8). The decrease in brain activity in the primary sensorimotor cortex on Day 1 and the

tendency to increase on Day 7 may be related to the masticatory performance reduction and CT

increase on Day 1, and recovery on Day 7, respectively. This may be explained as follows: on Day 1,

orofacial afferent input during mastication may have decreased, due to the reduction of the input from

the palate and prevention of tongue movement, and activation of the cortical areas that rely on such

sensory input decreased. Therefore, a reduction of coordination by cortical neurons during mastication

could have resulted in a decrease in food mixing ability and an increase in CT. It has previously been

reported that sensory disturbance of oral tissues in stroke patients was associated with impaired

masticatory function (17), and that reduced perception of oral structures caused by local anaesthesia

led to a decrease in food comminution and mixing ability (18). These findings suggest that reduced

sensory input from the palate may be responsible for masticatory impairment on Day 1. On the other

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 hand, by Day 7, learning of a new pattern of the tongue movement could have induced a change in
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brain activity in the primary sensorimotor cortex, resulting in the recovery of food mixing ability and CT.

The tongue plays an important role in bolus formation and transportation during mastication, and there

is reportedly a relationship between tongue motor skills and masticatory performance (19). However,

we could not assess how the palatal coverage changed the tongue movement. Previous studies have

measured tongue pressure using a palatal plate with pressure sensors (20), or assessed tongue

movements during mastication using videofluorography (21), and ultrasonography (22). We were

unable to use such approaches, because of the burden of physical strength or examination time it

would place on the subjects. Instead, the change in CT might in part indicate the motion parameters

during chewing. Further studies are needed to evaluate tongue movement more simply.

A previous study had shown that ⊿E did not recover to the Day 0 level by Day 7 (4), which was

not consistent with our findings. It is possible that differences in the thickness of the palatal plate, which

was 1.5 mm in the previous study, could be related to this inconsistency. However, further studies are

needed to clarify how wearing palatal plates of different thicknesses may influence chewing behaviour.

Motor adaptation is associated with profound changes in brain activation patterns over time (23).

Findings obtained from neuroimaging studies suggest that cortico-thalamic-cerebellar and

cortico-thalamic-striatal systems contribute to skill acquisition and long-term learning (24, 25). We

expected that the activation in the cerebellum would also differ between each experimental day, but in

this study, the activation on the cerebellum did not change statistically significantly, and only the

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 putamen showed significant changes. A previous study had shown that there is a general shift between
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the cortico-cerebellar to the cortico-striatal system with increased practice and motor learning (24). The

shift from the cerebellar cortex to the deep nuclei appears to take place within one session of practice

(26), whereas the shift from associative to sensorimotor regions within the striatum takes place over a

matter of days (27). The peak coordinate extracted in ROIs in this study was located in the

sensorimotor (caudal) region in the putamen. From the change in brain activation in the putamen, we

could detect late-stage learning in a new oral environment during 7 days post-insertion. We assumed

that we were unable to detect fast learning on Day 1 because of the block design of the fMRI.

The activation of the ACG and the pMFC decreased immediately after insertion of the plate.

These areas are presumably involved in the monitoring and control of complex movements (28, 29).

More specifically, monitoring for errors and signalling the need for adjustments are associated with the

pMFC (29). Experimental evidence from previous studies has demonstrated associative/premotor (AP)

and sensorimotor networks (SN) that operate within cortico-cerebellar and cortico-striatal systems. AP

areas contribute to early-stage performance and SN areas support performance in later practice stages

(24). The AP areas include the dorsolateral prefrontal cortex, rostal premotor areas, the inferior parietal

cortex, cerebellar cortex, and the dentate nucleus, which seem to be associated with the ACG and

pMFC (24, 28, 29). Our results suggested that the change in activation in AP areas appeared to occur

within one session on Day 1, similar to the findings for the cerebellum, whereas the shift from AP to SN

areas took place over a matter of days. Additionally, previous studies have shown that activation in the

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 prefrontal cortex and ACG increased when subjects were instructed to attend to a previously learnt or
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automatized sequence (28). Increase in the activation in AP areas during chewing with the palatal plate

may have occurred after more than 7 days.

There were some limitations in this study. Subjects were instructed to chew the gum naturally

during fMRI measurement, in a supine position. This was an unnatural position that may have affected

brain activation. This study suggested that similar adaptive changes in chewing-related brain activity

may also occur with removable dentures. However, it is unknown whether adaptation to removable

dentures could be changed by alteration of occlusion or of denture design. Moreover, the period of

adaptation to removable dentures varies from 23 weeks, to 30 days, to 3 months (1, 12, 30). Therefore,

the adaptation period to removable dentures may be longer for elderly subjects. We focused on the

adaptive changes during chewing in this study; however, the adaptation to removable dentures may

also be related to other factors, such as oral sensory function, speech, and swallowing. Further

consideration is required to clarify the adaptation in elderly individuals who require dentures from the

point of view of these varied factors, and to investigate individual differences in adaptation to new oral

environments.

Conclusion

In this study, we investigated the adaptive changes in brain activity during mastication with palatal

coverage. The results suggest that the recovery of activation in the face primary sensorimotor cortex

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 and putamen after 1 week may be associated with motor learning of gum-chewing in a new oral
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environment.

Acknowledgements

All MR images were obtained using the MRI scanner of the Department of Oral and Maxillofacial

Radiology, and the authors would like to thank Drs. Tohru Kurabayashi and Norio Yoshino for allowing

us to use the MRI scanner. This study was carried out without funding. The authors have no conflicts of

interests to declare.

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 Table 1: Significant clusters and their main activation peaks in the comparison between pre-insertion
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(Day 0) and post-insertion (Day 1, Day 3, and Day 7) are listed.

MNI coordinates Talairach coordinates

Cluster Peak Peak
(mm) (mm)

voxel voxel
Contrast Region Side size x y z x y z
T Z

Day 0 >
Superior frontal gyrus R 523 4.69 4.39 16 36 46 16 37 41
Day1

Anterior Cingulate R 176 4.60 4.31 10 18 24 10 19 21

Precentral gyrus R 1439 6.67 5.92 60 -4 24 59 -3 22

Putamen L 338 4.86 4.53 -28 -6 -8 -28 -6 -6

Putamen R 331 6.18 5.56 28 -6 -4 28 -6 -3

Precentral gyrus L 896 5.46 5.01 -46 -14 38 -46 -12 36

Posterior Cingulate L 643 4.75 4.44 -6 -60 6 -6 -58 8

Day 0 >
Superior frontal gyrus L 119 3.91 3.72 -22 54 14 -22 53 10
Day 3

Superior frontal gyrus R 485 5.27 4.86 20 54 4 20 52 1

Superior frontal gyrus R 324 4.72 4.41 14 38 44 14 39 39

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 Superior frontal gyrus L 175 4.36 4.11 -20 28 54 -20 30 48
Accepted Article
Putamen L 108 4.23 4.00 -28 -4 -10 -28 -4 -8

Putamen R 516 5.15 4.76 26 -8 -4 26 -8 -3

Precentral gyrus R 192 4.29 4.05 48 -10 18 48 -9 17

Posterior Cingulate L 708 4.71 4.40 -12 -56 14 -12 -54 16

Middle temporal gyrus L 206 4.45 4.19 -58 -16 -14 -57 -16 -11

Day 0 >
Superior frontal gyrus R 188 4.36 4.11 12 38 44 12 39 39
Day 7

Day 1 >
No significant activation
Day 0

Day 3 >
White matter L 177 4.00 3.80 -48 -32 32 -48 -30 31
Day 0

Day 7 >
No significant activation
Day 0

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 Figure Legends
Accepted Article
Fig. 1:

Change in ⊿E and cycle time (Means and SDs) (n = 28).

Fig. 2:

Activated voxels during the gum-chewing task projected onto the surface of the three-dimensional

brain (n = 27). Day 0: pre-insertion; Day 1: immediately post-insertion; Day 3: 3 days

post-insertion; Day 7: 7 days post-insertion. Left, right, and each sagittal section image: left and

right hemispheres and section images of X = 0. Cu: cuneus; sfg: superior frontal gyrus; p:

putamen; t: thalamus; C: cerebellum; smc: sensorimotor cortex.

Fig. 3:

Brain regions in the axial view significantly activated in chewing at pre-insertion of the palatal

plate (Day 0), as compared to post-insertion (Day 1, Day 3, and Day 7) (n = 27).

p: putamen; ACG: anterior cingulate gyrus; PRCG: precentral gyrus; sfg: superior frontal gyrus.

Comparison of “Day 0 > Day1, Day 0 > Day 3, Day 0 > Day 7”. Each coordinate indicates the

peak of the activated regions in Day 0 > Day 1.

Fig. 4:

Change in the beta-value in the peak coordinate in Day 0 > Day 1 in the ROI (Means and SDs) (n

= 27).

* P < 0.05, ** P < 0.01, *** P < 0.005, **** P < 0.001.

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Accepted Article

This article is protected by copyright. All rights reserved.

Accepted Article

This article is protected by copyright. All rights reserved.