Behavioural Brain Research 228 (2012) 254–260

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Behavioural Brain Research

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Research report

Increased occlusal vertical dimension induces cortical plasticity in the rat face
primary motor cortex
C. Kato ∗ , K. Fujita, S. Kokai, T. Ishida, M. Shibata, S. Naito, T. Yabushita, T. Ono
Department of Orofacial Development and Function, Orthodontic Science, Tokyo Medical and Dental University, 1-5-45 Yushima, Bunkyo-ku, Tokyo 113-8549, Japan

a r t i c l e i n f o a b s t r a c t

Article history: Previous studies have demonstrated that functional plasticity in the primary motor cortex (M1) is related
Received 24 September 2011 to motor-skill learning and changes in the environment. Increased occlusal vertical dimension (iOVD) may
Received in revised form modulate mastication, such as in the masticatory cycle, and the firing properties of jaw-muscle spindles.
10 November 2011
However, little is known about the changes in motor representation within the face primary motor cortex
Accepted 11 November 2011
(face-M1) after iOVD. The purpose of the present study was to determine the effect of iOVD on the face-
Available online 22 November 2011
M1 using intracortical microstimulation (ICMS). In an iOVD group, the maxillary molars were built-up by
2 mm with acrylic. The electromyographic (EMG) activities from the left (LAD) and right (RAD) anterior
Keywords:
Plasticity
digastric (AD), masseter and genioglossus (GG) muscles elicited by ICMS within the right face-M1 were
Intracortical microstimulation recorded 1, 2 and 8 weeks after iOVD. IOVD was associated with a significant increase in the number of
Orofacial sites within the face-M1 from which ICMS evoked LAD and/or GG EMG activities, as well as a lateral shift
Jaw muscle in the center of gravity of the RAD and LAD muscles at 1 and 2 weeks, but not at 8 weeks. These findings
Occlusal vertical dimension suggest that a time-dependent neuroplastic change within the rat face-M1 occurs in association with
Rat iOVD. This may be related to the animal’s ability to adapt to a change in the oral environment.
Crown Copyright © 2011 Published by Elsevier B.V. All rights reserved.

1. Introduction microstimulation (ICMS) [14]. ICMS can identify M1 neuroplastic-

Various parts of the body are precisely encoded into the primary
motor cortex (M1) of the cerebrum [1]. Representations of parts of
the body that are capable of fine motor control (e.g., fingers and
face) occupy a greater amount of the M1 area than those that exhibit
less-precise motor control (e.g., the trunk) [2]. The orofacial region
occupies 68% of the M1 area of the rat [3], which is greater than
the actual proportion. Since orofacial movements are controlled
by feedback from the periphery [4–7] and the M1 receives input
indirectly from the orofacial region [8,9], the exaggerated orofacial
motor representation in rats and humans indicates the importance
of feedback from the orofacial region in these species. Previous
studies using lesions or reversible cold-block of the M1 have shown
that the duration of some masticatory-related electromyographic
(EMG) activities [10] is affected, the amount of mouth-opening in
mastication is decreased [11], and the performance in a tongue-
protrusion task is impaired [12].
Previous studies have demonstrated that plastic changes occur
in the M1 in association with motor-skill learning and changes in
the environment [4,11–18]. One of the methods used to measure
such changes involves the mapping of motor cortical represen-
tations of muscle movements through the use of intracortical
∗ Corresponding author. Tel.: +81 3 5803 5963; fax: +81 3 5803 5963.
E-mail address: c.kato.orts@tmd.ac.jp (C. Kato).
ity induced by motor-skill learning or peripheral manipulations.
The M1 shows a use-dependent increase in movement represen-
tations [15,16], and these increased motor representations endure
long after the manipulations [17,18]. Meanwhile, peripheral nerve
lesions result in a decrease in motor representation and an increase
in association with nerve recovery [19]. Moreover, limb immobi-
lization in humans with a surgical cast has been shown to affect
M1 neuroplasticity [20,21].
In the orofacial region of the M1 (face-M1), neuronal activity
increases in relation to the training of tongue movements [22,23].
Whisker motor representation decreases following facial nerve
damage and increases with age [24]. Dental extraction is associ-
ated with a significant increase in the number of sites within the
face-M1 from which ICMS evokes EMG activity in jaw-opening [i.e.,
anterior digastrics (AD)] and tongue-protruding [i.e., genioglossus
(GG)] muscles [25]. Likewise, lingual nerve transaction is associ-
ated with neuroplastic changes in tongue motor representations in
the face-M1 [26].
The orofacial region responds to environmental changes, which
can include tooth [25] and dental pulp [27] extraction that
are accompanied by sensory disruption, and changes in the
occlusal vertical dimension (OVD) that are accompanied by sen-
sory changes. OVD is defined as the vertical dimension of the face
when the teeth and/or dentures are in maximum contact in centric
occlusion [28]. A decreased OVD due to the loss or severe abra-
sion/attrition of the posterior teeth can be restored by treatment

0166-4328/$ – see front matter. Crown Copyright © 2011 Published by Elsevier B.V. All rights reserved.
doi:10.1016/j.bbr.2011.11.013
 C. Kato et al. / Behavioural Brain Research 228 (2012) 254–260 255

Instruments,Tokyo, Japan) were used to record EMG activities from the left (LAD)
and right (RAD) AD and jaw-closing (masseter) muscles as well as the left GG muscle.
A craniotomy was performed to expose the right cortical hemispheres. To avoid
thick surface blood vessels, a fine glass-insulated tungsten microelectrode (250 ␮m-
diameter shaft with an 8◦ tapered tip; A-M Systems, Inc., Carlsborg, WA, USA) was
inserted transdurally into the exposed hemispheres in a standardized systematic
series of microelectrode penetrations. Penetrations had a horizontal spatial resolu-
tion of 0.5 mm, which coincided with the estimated extent of ICMS current spread
of less than 0.5 mm at an intensity of 60 ␮A [3,36]. Monophasic, cathodal, constant-
current stimulation trains at 333 Hz (i.e., 33.2-ms trains comprising 12 pulses of
0.2-ms and 2.8-ms interpulse intervals) were delivered through a stimulus isolator.
At each ICMS site, five trains were delivered at 1 Hz with a suprathreshold ICMS
intensity of 60 ␮A.

2.4. Histological analysis

Fig. 1. A schematic illustration of an animal in the increased occlusal vertical dimen-
sion (iOVD) group. The vertical dimension between the maxillary and mandibular
molars was increased by 2.0 mm with the build-up of acrylic on the maxillary molars.
The rats were sacrificed and fixed by the transcardial perfusion of 10% buffered
formalin. Nissl staining of coronal sections (100 ␮m) was used for subsequent veri-
fication of the three-dimensional positions of positive ICMS-penetrations and sites.
Sites located outside the cortical grey matter were excluded from the data analy-
sis, and cytoarchitectonic features were used to delineate the borders between the
granular and agranular cortex [37].
2.5. Data acquisition and analysis

[29,30]. In orthodontic treatment, anterior cross-bite patients can EMG activity was amplified using a high-input impedance differential amplifier
benefit from an increase in the OVD [31]. On the other hand, ante- (DAM 80, World Precision Instruments, Sarasota, FL, USA; ×1000 gain, 300 Hz and
rior open-bite patients can benefit from a decrease in the OVD 3 kHz for low- and high-pass filters, respectively) followed by full-wave rectifica-
[32]. In either situation, the change in the OVD should be limited tion and integration. All data were captured and analyzed on a computer with a CED
1401 interface and Spike2 software for Windows, version 5.21 (Cambridge Elec-
to within a physiological range [33]. The mechanism of the adap-
tronic Design, Cambridge, UK). An ICMS site was defined and counted as a positive
tation in response to a change in the OVD is believed to involve
denaturation of the muscle spindle and temporomandibular joint
mechanoreceptors in the short term, and adaptation in the long
term [27,34,35]. However, it is unclear whether a change in the
peripheral environment (i.e., an increase in OVD) is associated with
reorganization of the central nervous system. Thus, we tested the
null hypothesis that an increase in OVD (iOVD) is not associated
with a plastic change in the motor cortex.

2. Materials and methods

2.1. Animals

In this study, 28 13-week-old male Wistar albino rats were used. They were
housed in groups of 2–3 in plastic cages in an animal colony under a 12 h–12 h
light–dark cycle, with lights off at 19:00 pm. All experimental procedures described
here were approved by the Animal Welfare Committee and performed in accor-
dance with the Animal Care Standards of Tokyo Medical and Dental University
(#0100204C, 0110197A, 0110337A).

2.2. Surgical preparation

The rats were randomly divided into control (n = 7) and iOVD (n = 21) groups. The
animals were lightly anesthetized with thiamylal sodium (Isozol® , Yoshitomi Phar-
maceutical, Osaka, Japan; 60 mg/kg i.p.). In the iOVD group, the dimension between
the maxillary and mandibular molars was increased by 2.0 mm with the build-up
of acrylic on the bilateral maxillary molars (Fig. 1). To minimize the reduction in
the molar height due to abrasive movement of the mandible, the occlusal surfaces
of the lower molars were coated with acrylic fluid. Lateral radiographic images of
the skull of all rats were taken using a soft X-ray system (SRO-M50, Sofron, Tokyo,
Japan). The rats were then returned to their cages and allowed to recover from anes-
thesia. Intracortical microstimulation mapping was performed in iOVD animals after
1 (iOVD1 w ), 2 (iOVD2 w ) and 8 (iOVD8 w ) weeks (n = 7 each). ICMS was performed in
the control animals at 17-week old. The body weights of rats in both the control and
iOVD groups were monitored during the entire experimental period for assessment
of the general health status.

2.3. ICMS and EMG recordings

Before ICMS, lateral radiographic images of the skull of all rats were taken again
and we confirmed that the amount of iOVD did not change. The rats were kept
under a stable level of general anesthesia with ketamine–HCl (Ketalor® , Daiichi
Sankyo Propharma, Tokyo, Japan; 25–50 mg/kg/h, i.v.) throughout the ICMS experi-
ments. Additional anesthetic agent was injected when mild spontaneous whisking
or mild pinch-withdrawal and eyelid reflexes were observed. A stereotaxic appara-
tus (models SN-2 and SM-15 M, Narishige Scientific Instruments, Tokyo, Japan) was
used to place the animals in a prone position. Bipolar electrodes (100 ␮m diameter,
single-stranded, Teflon-insulated stainless-steel wires; type E-2, Narishige Scientific
Fig. 2. A representative electromyographic (EMG) recording of the responses
to intracortical microstimulation (ICMS) taken from the anterior digastric and
genioglossus muscles of a rat. The latency was defined as the time when the
ICMS-evoked EMG response exceeded 2 standard deviations of the baseline activ-
ity. Abbreviations: LAD, left anterior digastric muscle; RAD, right anterior digastric
muscle; GG, genioglossus muscle.
256 C. Kato et al. / Behavioural Brain Research 228 (2012) 254–260

Fig. 3. Representative motor maps of the left (LAD) and right (RAD) digastric and genioglossus (GG) muscles in a rat from the increased occlusal vertical dimension group at
2 weeks (iOVD2 w ; right) and a rat from the control group (control; left). The sites at which intracortical microstimulation (ICMS) was effective for eliciting electromyographic
(EMG) responses in the LAD, RAD and GG muscles are depicted by orange, green and blue dots, respectively. Black dots represent sites where ICMS did not evoke EMG activity
in the LAD, RAD, or GG muscles. Abbreviation: AP, anteroposterior.

ICMS site if at least three of five ICMS trains evoked an EMG response in a muscle
with an onset latency of maximally 40 ms and a peak activity that exceeded the
mean value in the initial 10 ms of the EMG response plus 2SDs (Fig. 2).

2.6. Center of gravity (CoG)

The CoG of positive ICMS sites, which defines the three-dimensional [medio-
lateral (ML)/superoinferior (SI)/anteroposterior (AP)] center position of the motor
maps weighted relative to the extent of the muscle motor representations,
was determined as described previously [38]. The following equation was used:
X = ai Xi /ai , where ai is the number of positive ICMS sites at a cortical ML coor-
dinate, Xi . In a similar manner, the SI coordinates Yi and the AP coordinate Zi were
determined.

2.7. Statistical analyses

Data from all groups and subgroups were compared using repeated measures
ANOVA, followed by the Bonferroni/Dunn post hoc test (5% significance level).
Statview for Windows, version 5.0 (SAS Institute, Cary, NC, USA), was used for the
statistical analysis.

3. Results
There was no significant difference in mean body weight
between the control and iOVD groups throughout the experiment.
Within the mapped area in the control and iOVD groups, ICMS
evoked EMG activities in the AD and/or GG muscles from an exten-
sive area within the right face-M1; the positive ICMS sites were
located along the entire depth of layers V and VI (Fig. 3). Within
the mapped area, both the AD and GG muscles showed extensive
motor representations within the face-M1, but there were very
few positive ICMS sites for the masseter muscle. Since the num-
ber of positive ICMS sites for the masseter muscle was negligible,
the masseter muscle was excluded from further analysis.
There were no significant differences in the number of inde-
pendent positive ICMS sites among muscles in the control group.
The LAD muscle had a significantly more independent positive site
than the RAD muscle in the right hemisphere, although there were
Fig. 4. The number of independent sites within the face-M1 at which intracortical
microstimulation (ICMS) effectively elicited electromyographic (EMG) responses in
the left (LAD) or right anterior digastric (RAD) or genioglossus (GG) muscles in the
control (n = 7) and iOVD (1, 2 and 8 W) groups (n = 7 each). Significant differences in
number of sites at different times were labeled with asterisks (p < 0.05). Bars indicate
the standard deviations of the mean.
no statistically significant differences between the LAD and GG
muscles in the right hemisphere in the control and iOVD groups.
There were significant differences between the control and both
the iOVD1 w and iOVD2 w groups with regard to the number of inde-
pendent positive ICMS sites in the LAD and GG muscles, but not the
RAD muscle (Fig. 4). On the other hand, there were no significant
differences between the control and iOVD8 w groups with regard to
the number of positive ICMS sites in the LAD and GG muscles.
 C. Kato et al. / Behavioural Brain Research 228 (2012) 254–260 257

Table 1
Onset latency of electromyographic (EMG) activities in the left (LAD) and right
anterior digastric (RAD) and genioglossus (GG) muscles evoked by intracortical
microstimulation (ICMS).

Variables LAD RAD GG

Mean SD Mean SD Mean SD

a
Control 15. 3 4.1 22.3 5.6 19.6 5.4
iOVD1 w 15.4a 5.6 24.1 2.7 20.2 6.1
iOVD2 w 15.4a 5.4 23.8 2.7 21.0 5.7
iOVD8 w 12.7a 4.1 20.6 3.2 17.7 7.4

Means and standard deviations of the mean (SD) are shown in ms. Within each of
the iOVD groups, the LAD muscle showed a significantly shorter onset latency than
the RAD muscle (control: p = 0.015, iOVD1 w : p < 0.0001, iOVD2 w : p = 0.0015, iOVD8 w :
p < 0.0001). Within each of the iOVD groups, the LAD muscle showed a significantly
shorter onset latency than the GG muscle (control: p = 0.013, iOVD1 w : p = 0.0053,
iOVD2 w : p = 0.015, iOVD8 w : p < 0.0001).
a
Indicates a significant difference between the LAD and RAD/GG muscles in the
control and iOVD1 w , iOVD2 w and iOVD8 w groups.

Fig. 5. The number of overlapping sites within the face-M1 at which intracortical
microstimulation (ICMS) effectively elicited electromyographic (EMG) responses in
two or more of the left (LAD) and right anterior digastric (RAD) and genioglossus
(GG) muscles, in the control (n = 7) and iOVD (1, 2 and 8 W) groups (n = 7 each). Sig-
nificant differences in number of sites at different times were labeled with asterisks
(p < 0.05). Bars indicate the standard deviations of the mean.
The number of sites at which ICMS evoked concurrent EMG
activity in the LAD and GG muscles (i.e., overlapping positive ICMS
sites) was significantly increased in both the iOVD1 w and iOVD2 w
groups (Fig. 5). However, the number of overlapping positive ICMS
sites for other pairs or triads did not show any significant differ-
ence between the iOVD groups and the control group, except for
the RAD, LAD and GG muscles in the iOVD1 w group and control
group.
The LAD, RAD and GG muscles did not show any significant dif-
ferences in onset latency between the control and iOVD groups.
Within each of the iOVD subgroups, the LAD muscle showed
significantly shorter onset latencies than the RAD muscle (con-
trol: p = 0.015, iOVD1 w : p < 0.0001, iOVD2 w : p = 0.0015, iOVD8 w :
p < 0.0001; see Table 1). Within each of the iOVD groups, the LAD
muscle showed a significantly shorter onset latency than the GG
muscle (control: p = 0.013, iOVD1 w : p = 0.0053, iOVD2 w : p = 0.015,
iOVD8 w : p < 0.0001).
With regard to the temporal displacement of the CoG, there was
a significant difference in the CoG between the LAD and GG mus-
cles in the iOVD1 w and iOVD2 w groups compared with the iOVD8 w
group and the control group. (Fig. 6)

Fig. 6. Centers of gravity weighted against the number of effective intracortical microstimulation (ICMS) sites for the right (B) and left anterior digastric (C) and genioglossus
(D) muscles in three-dimensional space. The directions of the three axes are indicated on the rat brain (modified from Paxinos G, Watson C. The Rat Brain. sixth ed. The
Netherlands: Amsterdam; 2007.) (A). Abbreviations: AP, anteroposterior; ML, mediolateral; SI, superoinferior.
258 C. Kato et al. / Behavioural Brain Research 228 (2012) 254–260
4. Discussion
In this study, the null hypothesis was rejected in the
iOVD1 w and iOVD2 w groups. However, it was supported in the
iOVD8 w group. These findings suggest that a time-dependent
neuroplastic change occurs in the face-M1 in association with
iOVD.
4.1. Methodological considerations
In this study, we selected an animal model with an increase in
OVD to investigate the effects of the passive restriction of move-
ment without any damage to the muscles or nerves. We increased
OVD, and this was estimated to be within 30% of the rat’s maxi-
mum mouth-opening. This is equivalent to the freeway space in
humans [39,40]. The freeway space is defined as the interocclusal
separation between the maxillary and mandibular teeth when the
mandible is in the physiological rest position; the position when the
condylar head is nearly in the centric position in the glenoid fossa.
It has been reported that an increase in the OVD within the range of
the freeway space satisfactorily adapted and remained stable over
a two-year observation period [40]. In our study, there was no sig-
nificant difference in body weight between the control and iOVD
groups after 8 weeks; iOVD had no effect on the general condition.
This suggests that weight differences between iOVD and control
groups did not influence the oral condition or the face-M1. Indeed,
dentists often increase OVD within the range of the freeway space
during the course of treatment [35,41], which is accompanied by
changes in EMG activities during mastication [42].
We used ICMS in the right hemisphere and recordings of evoked
EMG activity in the present study. The LAD muscle had signifi-
cantly more positive sites than the RAD muscle in both the iOVD
and control groups. Contralateral dominance of the motor cortex
has been demonstrated in humans [43], monkeys [6,12,44,45] and
rats [3,45]. The masseter muscle had very few positive sites, and
this finding is consistent with previous studies in rats [24] and
monkeys [6,7,24]. In contrast, a previous study on cortical map-
ping of the human masticatory muscle using transcranial magnetic
stimulation found that the masseter muscle was represented [46].
4.2. Changes in the face-M1 responsible for the AD and GG
muscles
IOVD was associated with a significant increase in the num-
ber of positive ICMS sites, and with a shift in the CoG of the LAD
and GG muscles within the face-M1. It should be pointed out that
any modification (i.e., increase or decrease) of cortical maps, occur-
ring without a shift in the CoG of positive ICMS sites, cannot truly
distinguish the plastic reorganization of the cortical motor out-
put from increased excitability of the corticospinal system. In fact,
when cortical stimuli are delivered to the region surrounding the
motor-representation area, they may be equally able to induce acti-
vation of the muscles simply because the corticospinal system is
more excitable [47]. Therefore, any enlargement of the motor area
extension can be considered to reflect cortical rearrangement only
if spinal excitability is not modified [48]. Thus, we speculate that
there was no significant change from the face-M1 to the trigeminal
motor nucleus that was analogous to the corticospinal system for
limbs, while there were changes in motor representations based
on a shift in the CoG and a greater number of positive ICMS sites
within the face-M1.
Skill training has been associated with cortical map expansion
and synaptogenesis [15,16,25,49]. Strength and endurance train-
ing induce angiogenesis in the motor cortex [50,51], but not motor
map reorganization or an increase in the number of synapses. In
the iOVD1 w and iOVD2 w groups, the number of positive ICMS sites
for both the AD and GG muscles was significantly greater than that
in the control group at the same recording timing, which suggests
the occurrence of cortical map expansion associated with synap-
togenesis. Therefore, iOVD is more analogous to skill training than
strength or endurance training, since iOVD may have changed the
oral environment and rats acquired a new skill.
The orofacial area has rich sensory innervations, and M1 receives
these inputs indirectly. Somatosensory inputs provide peripheral
feedback that is used to control orofacial motor functions [4–7].
IOVD appeared to modulate afferent inputs, and in turn affected
efferent output. Increased EMG activity of AD muscles throughout
the day has been attributed to iOVD [42,52,53]. Moreover, subjects
with anterior open-bite showed more GG EMG activity than those
without anterior open-bite [54]. These findings in previous studies
are consistent with the increased AD and GG EMG activities at 1 or
2 weeks after iOVD.
In the iOVD8 w group, there were significantly fewer positive
ICMS sites for the AD and GG muscles than in the iOVD1 w and
iOVD2 w groups. Moreover, there were no significant differences
between the iOVD8 w and control groups. This means that there
were no significant long-term changes associated with iOVD in the
face-M1. Playing with string and reading Braille result in long-term
changes in cortical motor representations [17,18]. It has been sug-
gested that, under these conditions, changes in muscle activities
modify the state of the M1 [55]. Learning is a voluntary activity,
but orofacial movements (e.g., tongue protrusion and jaw-opening)
are semi-automatic and peripheral feedback plays an important
role [56,57]. According to our study, the mechanism of the change
in neural activity observed 8 weeks after iOVD can be explained
by the assumption that recovery in peripheral structures causes
reconstruction of the central nervous system, and efferent output
then adapts to this change in the CNS.
4.3. Overlapping positive ICMS sites for the AD and GG muscles
ICMS simultaneously evoked more AD and GG EMG activities
in the iOVD1 w and iOVD2 w groups. Such overlapping of cortical
motor representations, along with the extensive representations
of the AD and GG muscles, may be important for the coordina-
tion of dynamic orofacial movements that involve several muscles
and may also serve as the substrate that underlies neuroplastic
mechanisms that are manifest in the reorganization of motor rep-
resentations in the face-M1 [58,59]. The overlap of jaw and tongue
motor representations has previously been reported after 1 week
of extraction of the incisor in the rat [25]. This indicates that the
face-M1 adjust in response to a new environment in the iOVD1 w
and iOVD2 w . In a study in the cat, sites at which multiple responses
could be evoked decreased as the cat grew and the brain map was
segmented [60]. Therefore, the temporal decrease in the overlap of
motor representations suggests both an increase in accuracy and
a reduction in the diversity of movement. We speculate that the
simultaneous decrease in the ICMS-evoked AD and GG EMG activ-
ities in the iOVD8 w indicates a decreased demand for coordinated
motor control.
4.4. Latency of ICMS-evoked EMG activity
A significant contralateral dominance was reflected in the dif-
ference in the latency of ICMS-evoked jaw responses in the control
and iOVD groups, which is attributable to the difference in the
pathways to the RAD and LAD muscles from the right hemisphere.
These findings are consistent with findings in previous studies in
the rat AD muscle [25,61]. Also, there was no significant change
in the latency with iOVD. This indicates that iOVD had no signif-
icant effects on the fastest corticofugal projections to trigeminal
motor nucleus premotoneuronal structures [62]. A previous study
 C. Kato et al. / Behavioural Brain Research 228 (2012) 254–260
found that motor nerve conduction velocity decreases depending
on dysfunction [63]. Therefore, findings in our study suggest that
peripheral nerve lesion by iOVD had no significant influence on the
latencies of the ICMS-evoked EMG activities in the GG, LAD and
RAD muscles.
4.5. Peripheral and central interactions in association with iOVD
M1 plays a crucial role when an animal adapts to an altered
state for the limbs. For example, M1 is involved in adaptation to
microgravity [13] and immobilization of the lower limbs [20,25].
A previous study reported that there were no significant differ-
ences in the jaw and tongue motor representations between two
groups fed a soft- and hard-diet after 2–3 weeks [61]. Although
the experimental set-up in that study [61] was similar to that in
ours, neuroplastic changes following iOVD were seen only in our
study. There are at least three possible explanations for this discrep-
ancy. First, the previous study [61] overlooked the cortical change
that probably occurred after 3 weeks. Second, the cortical change
occurred much earlier and recovered to the original state before
recording. Third, differences in diet consistency and iOVD may have
diverse effects on peripheral organs. The diet consistency influ-
ences the masseter muscle [64,65] and temporomandibular joint
[66] during the growth period, but not in adulthood. This suggests
that diet consistency has less of an effect on the orofacial region
than iOVD.
The influence of iOVD has been reported with regard to
changes in peripheral inputs; a short-term decrease in the fir-
ing activity of the muscle spindle and temporomandibular joint
mechanoreceptors and long-term adaptation [34,35]. During vol-
untary movement, the input from the muscle spindle is reflected
in M1 [67], indicating that information regarding muscle length
is important. In the present study, ICMS and recording were per-
formed 8 weeks after iOVD. Interestingly, the muscle spindle
has been reported to adapt to OVD in 8 weeks [35]. Our novel
findings suggest that iOVD may be associated with significant neu-
roplastic changes that reflect adaptation of the central nervous
system.
5. Conclusions
The present findings suggest that a significant time-dependent
neuroplastic change within the rat face-M1 occurs in association
with iOVD. This may be related to the animal’s ability to adapt to a
change in the oral environment.
Acknowledgments
This study was supported by Grants-in-Aid for Scientific
Research (21792061; 21792062, 2009–2010) from the Japan Soci-
ety for the Promotion of Science. Part of this study was presented
at the 70th Annual Meeting of the Japanese Orthodontic Society,
Nagoya City, October 17–20, 2011.
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