Exp Brain Res (2015) 233:237252
DOI 10.1007/s00221-014-4107-6
RESEARCH ARTICLE
Interaction ofbrain areas ofvisual andvestibular simultaneous
activity withfMRI
HellenM.Della-Justina HumbertoR.Gamba
KaterinaLukasova MarianaP.Nucci-da-Silva
AndersonM.Winkler EdsonAmaroJr.
Received: 2 September 2013 / Accepted: 18 September 2014 / Published online: 10 October 2014
Springer-Verlag Berlin Heidelberg 2014
Abstract Static body equilibrium is an essential requi-
site for human daily life. It is known that visual and ves-
tibular systems must work together to support equilibrium.
However, the relationship between these two systems is not
fully understood. In this work, we present the results of a
study which identify the interaction of brain areas that are
involved with concurrent visual and vestibular inputs. The
visual and the vestibular systems were individually and
simultaneously stimulated, using flickering checkerboard
(without movement stimulus) and galvanic current, during
experiments of functional magnetic resonance imaging.
Twenty-four right-handed and non-symptomatic subjects
participated in this study. Single visual stimulation shows
positive blood-oxygen-level-dependent (BOLD) responses
(PBR) in the primary and associative visual cortices. Sin-
gle vestibular stimulation shows PBR in the parieto-insular
vestibular cortex, inferior parietal lobe, superior temporal
gyrus, precentral gyrus and lobules V and VI of the cer-
ebellar hemisphere. Simultaneous stimulation shows PBR
H.M.Della-Justina(*) H.R.Gamba
Graduate Program inElectrical andComputer Engineering,
Federal University ofTechnology-Parana, Av. Sete de Setembro,
3165, Curitiba, PR 80230-901, Brazil
e-mail: hellenjustina@gmail.com
K.Lukasova M.P.Nucci-da-Silva E.AmaroJr.
Department ofRadiology, University ofSao Paulo, Av. Dr. Enas
de Carvalho Aguiar, s/n, So Paulo, SP 05403-900, Brazil
A.M.Winkler
Oxford Centre forFunctional MRI ofthe Brain, University
ofOxford, OxfordOX3 9DU, UK
A.M.Winkler
Department ofPsychiatry, Yale University School ofMedicine,
300 George St, New Haven, CT 06511, USA
in the middle and inferior frontal gyri and in the precentral
gyrus. Vestibular- and somatosensory-related areas show
negative BOLD responses (NBR) during simultaneous
stimulation. NBR areas were also observed in the calcarine
gyrus, lingual gyrus, cuneus and precuneus during simulta-
neous and single visual stimulations. For static visual and
galvanic vestibular simultaneous stimulation, the reciprocal
inhibitory visualvestibular interaction pattern is observed
in our results. The experimental results revealed interac-
tions in frontal areas during concurrent visualvestibular
stimuli, which are affected by intermodal association areas
in occipital, parietal, and temporal lobes.
Keywords Visual system Vestibular system
Simultaneous stimulation Visualvestibular interaction
fMRI
Introduction
Static body equilibrium is an essential requisite for daily
life. It is known that equilibrium balance is controlled by
three sensory systems: the vestibular system, which acts by
tracking the position and the movements of the head; the
visual system, which gives a spatial estimate of the position
of objects relative to the body; and the proprioceptive sys-
tem, which monitors the relative position of different parts
of the body. The interactions among these three systems
prevent the imbalance sensation. The visual and vestibu-
lar systems can be individually or simultaneously studied,
using unimodal or bimodal stimulations with functional
magnetic resonance imaging (fMRI). A conventional static
visual task consists of a flickering checkerboard (Fox and
Raichle 1985; Singh et al. 2003). The vestibular stimulation
is accomplished using transcutaneous bipolar sinusoidal
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galvanic current, applied through a pair of electrodes that is
attached to the skin, one electrode over each mastoid pro-
cess (Wardman et al. 2003; Moore et al. 2006; MacDougall
et al. 2006; Aw et al. 2006). It is also possible to stimulate
the vestibular system with caloric stimulation in a neuro-
imaging study (Fasold et al. 2002; Dieterich et al. 2003a;
Marcelli et al. 2009). However, the main advantage of
using galvanic vestibular stimulation (GVS), compared to
the caloric stimulus, is that GVS acts on vestibular system
by firing the primary vestibular neurons, causing a quickly
stimulus response. Also, the GVS affects the otolithic affer-
ents (Watson et al. 1998) and the fiber of the semicircular
canals (Schneider et al. 2000) separately.
The earliest study that used GVS in a fMRI experiment
was published by Lobel et al. in 1998. In this study, it was
identified positive blood-oxygen-level-dependent (BOLD)
responses in the temporo-parietal junction, the central
gyrus, the intraparietal gyrus, and the frontal lobe. It was
also identified negative BOLD responses (NBR) in the
anterior part of the frontal lobe. Bense et al. (2001) used
galvanic vestibular stimulation and galvanic skin stimula-
tion to differentiate oculomotor vestibular and nociceptive
functions. They found areas with positive BOLD responses
(PBR) that are related with multisensory vestibular and
oculomotor functions. NBR in the bilateral visual cortex
was observed, and an inhibitory interaction pattern between
the visual and vestibular systems was perceived. A recipro-
cal inhibitory visualvestibular interaction pattern was first
described by Brandt et al. (1998), in a positron emission
tomography (PET) study, using a perception of self rota-
tion that was visually induced by a rotating stimulus. They
observed that during increase of regional cerebral blood
flow (rCBF) in the parietal and occipital cortices there was
a simultaneous decrease of rCBF in regional cerebral blood
flow in the bilateral posterior insula. As a functional con-
sequence, the decreased cortex activity protects the organ-
ism from conflicting inputs, i.e., the reduction response of
visual cortex during vestibular stimulation suppress visual
motion input and the signal reduction of vestibular cortex
during visual stimulation (constant velocity) should reduce
the vestibular systems sensitivity to head acceleration. The
theory proposed by Brandt and Dieterich (1999) for this
pattern is that a reciprocal inhibitory interaction mechanism
allows for a change in the dominant sensory system that is
dependent on the prevailing sensory mode. Other authors
also mentioned the reciprocal inhibitory pattern between
the visual and the vestibular systems during psychophysical
(Probst et al. 1985, 1986), electrophysiological (Probst and
Wist 1990), and neuroimage (Dieterich et al. 2003b; Naito
et al. 2003; Stephan et al. 2005) studies. More recently, a
transcranial magnetic stimulation (TMS) applied to visual
cortex demonstrated that vestibular activation modulates
human visual cortex excitability (Seemungal et al. 2013).
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Exp Brain Res (2015) 233:237252
Previous studies used visual and vestibular stimulations
in combination, Deutschlnder et al. (2002) conducted a
PET study with: (1) a small-field visual motion; (2) a ves-
tibular caloric irrigation stimulation; and (3) bimodal stim-
ulation, i.e., caloric and visual motion stimulation simulta-
neously. The vestibular stimulation led to decrease of rCBF
in visual cortex areas and increase of rCBF in temporo-
parietal vestibular areas, including posterior insula and
retroinsular regions, anterior cingulate gyrus, middle fron-
tal gyrus, and inferior parietal cortex. Visual stimulation
increased rCBF in bilateral occipital visual cortex areas and
in motion-sensitive area MT/V5, in the middle temporal
and middle occipital gyri. Visual stimulation was also asso-
ciated with decreased rCBF in the parieto-insular vestibular
cortex (PIVC), the superior and middle temporal gyri, the
inferior parietal lobule, the middle frontal gyrus, and the
anterior cingulate gyrus. Simultaneous vestibular and vis-
ual stimulation evoked activation in both sensory systems,
but with smaller and less statistical significance. The infe-
rior parietal lobule and the middle frontal gyrus activations
did not occur during bimodal stimulation.
Fink et al. (2003) conducted a fMRI study involving
visuospatial task and vestibular stimulation. The authors
manipulated the egocentric spatial reference frame by gal-
vanic vestibular stimulation and found that the right pos-
terior parietal and the right ventral premotor cortex are
responsible for an interaction between the neural mecha-
nisms that underlie allocentric visuospatial judgments and
vestibular stimulation. The vestibular stimulation showed
increased neural activity in the posterior insula and in the
superior temporal gyrus, including the PIVC. Line bisec-
tion judgments resulted in increased neural activity in bilat-
eral extrastriate cortex, parieto-occipital cortex, and poste-
rior parietal cortex, in the right premotor, dorsolateral, and
ventrolateral prefrontal cortex, and in the left cerebellum.
These results indicate an interaction between the visuos-
patial attentional systems and the vestibular processing
systems.
Although the cortical visualvestibular networks pro-
posed by Brandt et al. (1998) were confirmed by other stud-
ies, further investigations are needed. In previously studies,
the bimodal stimulation used was a visual stimulus that
exerted influence into the vestibular system (Deutschlnder
et al. 2002; Fink et al. 2003). This kind of stimulation
might intervene in the brain interaction processes that are
related to the visual and the vestibular systems. The visual
vestibular inhibitory reciprocal interaction occurs when
there are contradictory visual and vestibular inputs. Since
none of the earlier works included simultaneous static vis-
ual and vestibular stimuli, we question whether the brain
interactions areas will be the same when there is a purely,
but simultaneously, input for each system. Additionally, we
argue if it is possible to identify a specific region (i.e., a
Exp Brain Res (2015) 233:237252 239
critical node in the visual/auditory perception network) that
represents the interaction between static visual and vestibu-
lar inputs.
Also, the somatosensory processing was reported as
having strong integrative with the cortical vestibular system
(Lopez et al. 2008, 2012). Ferr et al. reported that cold
caloric vestibular stimulation (Ferr et al. 2011a, b) and
galvanic vestibular stimulation (Ferr et al. 2013) increase
tactile sensitivity on both hands. So, it is possible to con-
clude that the vestibular and somatosensory systems may
have a functional cross-modal perceptual interaction (Ferr
et al. 2011a, b; Bottini et al. 2013). Brain-damaged patients
also corroborates with these findings, the somatosensory
processing is affected by caloric vestibular stimulations
during left (Vallar 1998; Bottini et al. 2005) and right
neglect (Bottini et al. 2005; Ronchi et al. 2013). Corre-
spondingly, a recent electrophysiological study confirmed
that vestibular stimulation modulates somatosensory corti-
cal processing (Ferr et al. 2012).
In this paper, we present the results of a study which
identify the interaction of brain areas that are involved with
concurrent visual and vestibular inputs. The fMRI experi-
ments were performed with: a pure visual stimulus (flick-
ering checkerboard, without movement stimulus), where
there is no induction of vestibular responses; a pure vestib-
ular stimulus (galvanic vestibular stimulation); and a com-
bination of these two stimuli (simultaneous stimulation).
Materials andmethods
Participants
Twenty-four subjects were examined. The data from three
subjects were excluded; the stimuli were correlated with
head movements (above 1.0mm in translation and 1.0
in rotation). The data from 21 subjects were analyzed (12
men and 9 women, ages varying from 21 to 36years, mean
age 26.9years, SD 3.9years). The volunteers who partici-
pated in the study did not have history of neurological or
psychiatric disorders and did not make use of psychoac-
tive medications. All subjects had normal or corrected-to-
normal vision and normal vestibular functions (absence of
disorders regarding balance, vertigo or dizziness), by self-
reported. The absence of nystagmus during eye movement
was qualitatively assessed with eye tracking register during
rapid eye movements and visual pursue. The Edinburgh
Handedness Index (H.I) was computed for each subject
(Oldfield 1971). All the subjects were right-handed (mean
H.I =0.69, SD=0.24, median=0.67). This study was The vestibular stimulation was applied by a stimulator spe-
approved by Research Ethics Committee of University of
So Paulo, and informed consent was obtained from all
participants. Mental functioning was assessed with the
Mini International Neuropsychiatric Interview (M.I.N.I
Plus 5.0.0; Brazilian version by Amorim (2002)). The
neuropsychological tests focused on cognitive functions
such as attention (assessed by the continuous attention test
(Cambraia 2003)), trail making (Delis et al. 2001) and esti-
mated intelligence quotient (Matrix and vocabulary sub-
tests from WAIS-III battery (Wechsler 2004)). All subjects
were within the normal range, above 25%.
Experimental setup
Participants performed single visual (VIS), single vestibu-
lar (VES), simultaneous (DUAL) and baseline (BASE)
conditions in separate fMRI experiment blocks. One exper-
iment sequence consisted of a total of 17 blocks, includ-
ing 5 blocks of BASE, 4 blocks of VIS, 4 blocks of VES
and 4 blocks of DUAL conditions. In all fMRI experiment,
each block had duration of 21s, resulting in a total experi-
mental run-time of 6min and 12s. The stimulus sequence
was as follows: BASE, VIS, VES, DUAL, BASE, and was
repeated 4 times. A pseudo-randomized sequence was
selected to avoid that two consecutive blocks conditions
were the same and it was identical for all participants. To
ensure that tasks were performed correctly, eye position
was recorded using an Eye Tracker (HCET01 from Mag
Design & Engineering, USA) adapted for the MRI system.
The presentation used the software E-Prime 1.1 (Psychol-
ogy Software Tools, Inc., USA). To be familiarized with the
experiment stimuli sequence, in the same day and before
the fMRI experiment, the participants performed once the
same tasks but outside of the MRI scanner.
Visual stimulation
Checkerboard stimulation was used as visual stimulus dur-
ing the fMRI experiment. No component of spatial orien-
tation or movement was used, i.e., a pure visual task was
performed by the volunteers. The visual stimuli were back-
projected onto a screen (12.709.85) with the resulting
image visible to the participant inside the scanner through a
mirror mounted on the head coil. The visual stimulus con-
sisted of a black and white checkerboard (7.307.30,
33 squares) alternating every 62ms. The central square
was located in the center of the screen and contained a cen-
tral red cross (0.370.37) at which the subjects were
instructed to fixate their gaze.
Vestibular stimulation
cially built to be used inside the magnet room. The stimula-
tor consists of a signal generator, powered and controlled
by a computer that generates an optical signal for optical
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fiber transmission to a modulated electrical current source.
The current source produces an alternate current (AC) with
adjustable amplitude, frequency and waveform. Shield-
ing was made only for the circuits in the scanner proxim-
ity. During all the experiments, the subjects held a safety
switch button (handy switch) to interrupt the stimulus in
case of discomfort. A grounded shielded cable was used
to make the connection between the stimulator and the
safety button. The first 60cm of connection between the
handy switch and the electrodes was made with twisted
wires. To eliminate the currents induced in the wires by
the MRI RF pulses, three low-pass LC filters were placed
along the cable that made the connection between the
handy switch and the electrodes. The cutoff frequency of
the LC filters was in the order of 50Hz. After the third
LC filter, carbon wires were used to connect it to the elec-
trodes. Circular silicone (9cm2, in-house) electrodes were
bilaterally positioned over the mastoid processes of the
temporal bones (for more details about the equipment see
Della-Justina et al. 2014). Before placing the electrodes,
the skin was cleaned with alcohol, and a conductive gel,
soluble in water, was used to improve the conductance. A
sinusoidal electric current with a frequency of 1.0Hz was
applied in all volunteers. The amplitude of the current was
manually adjusted, according to the sensitivity of volun-
teers (on average 1.5mA). To determine the individual
stimulus amplitude, the sinusoidal galvanic stimulation was
slowly increased from 0mA until the subject first had a
clear perception of body movement, and then, the stimulus
amplitude was increased until the skin sensation started to
become unpleasant or painful. At that point, the stimulus
intensity was lowered to a level where the volunteer was
capable of perceiving an imbalance without a pain discom-
fort. Subjects were instructed to keep their eyes open and to
fixate their gaze in the center of a black screen.
Simultaneous stimulation
In the DUAL condition, the participants received both vis-
ual (VIS) and vestibular (VES) stimuli simultaneously. The
visual and the galvanic stimuli were identical to those that
were applied separately.
Baseline
In the baseline (BASE) condition, the participants were
instructed to fixate their gaze at a red cross in the center of
a black screen. No motor or cognitive tasks were required.
Scanning protocol
The fMRI experiments were performed in the Institute of
Radiology, University of So Paulo, using a Philips Intera
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Exp Brain Res (2015) 233:237252
Achieva 3.0 T scanner (Philips Healthcare, Andover,
MA, USA). Functional T*2-weighted echo-planar images
(EPI) sensitive to the BOLD contrast were acquired using
an 8-channel head coil and the following parameters:
TE/TR=30/3,000ms, flip angle=90, voxel size=3mm
isotropic, FOV=240mm, matrix size=8080, num-
ber of slices=41 (acquired sequentially), and slice
gap=0.5mm. A total of 124 EPI volumes were acquired.
The first five volumes were discarded to allow for signal
equilibrium effects. A structural, T1-weighted image was
also acquired for image registration, using the following
parameters: TE/TI/TR=3.2/900/7,000ms, flip angle=8,
180 slices, voxel size=1mm isotropic, FOV=240mm,
matrix size=240240, and sense 2.0.
Analysis
The fMRI data were submitted to statistical analysis using
the SPM8 software (Friston et al. 1995). Preprocess-
ing steps consisted of slice timing correction to the mid-
dle slice, motion correction through rigid-body realign-
ment to the mean functional image, co-registration to the
T1-weighted structural image, normalization (nonlinear
registration) to the MNI152 standard space and smoothing
with a Gaussian kernel with full width at half maximum
(FWHM) of 6mm. The signal was modeled as a boxcar
function convolved with a double-gamma hemodynamic
response function with a delay to peak of 6s (Glover
1999). Low-frequency drifts were removed with a tempo-
ral high-pass filter with a cutoff frequency of 0.0078Hz,
and errors were modeled using an auto-regressive model of
order 1 (AR(1)). The four epoch types (BASE, VIS, VES
and DUAL) were entered into a design matrix as separate
regressors. The statistical analysis was performed using the
general linear model as implemented in SPM8.
Statistical contrasts were used to identify both posi-
tive and negative BOLD responses, henceforth referred to
as PBR and NBR, respectively. Group analysis was per-
formed using a mixed effects model with one sample t
test as the summary statistic. The statistical maps for PBR
were obtained by the contrasts: VISBASE, VESBASE
and DUALBASE; and the statistical maps for NBR were
obtained by the contrasts: BASEVIS, BASEVES and
BASEDUAL. Quantitative comparisons between sin-
gle and DUAL conditions were performed using con-
trasts that directly compared VIS with DUAL conditions
(VISDUAL; DUALVIS) and VES with DUAL condi-
tions (VESDUAL; DUALVES). An interaction con-
trast, DUAL(VIS+VES), was also calculated to assess
the cortical areas that increased their signal in response
to the simultaneity of both stimuli. The resulting statisti-
cal maps for all the contrasts were corrected for multiple
comparisons. A voxel-wise control of the false discovery
Exp Brain Res (2015) 233:237252 241
rate (FDR) with a threshold of p<0.05 was used. To make
characterized BOLD response to pure stimulus conditions
(VISBASE and VESBASE), an individual percent signal
change analysis in four regions of interests (ROI) was done
using MarsBaR (Marseille bote rgion dintrt; Brett
et al. 2002). Functional ROIs were created by localizing the
clusters of interest in the statistical map for each contrast.
The right visual region had a total volume of 16,144 mm3
with a center of mass in x=+36, y=65, z=4; the publication by Eickhoff et al. (2005).
left visual region had a total volume of 1,872 mm3 with a
center of mass in x=27, y=85, z=7; the right ves-
tibular region had a total volume of 976 mm3 with a center
of mass in x=+39, y=0, z=13; and the left vestibular
region had a total volume of 144 mm3 with a center of mass
in x=37, y=+1, z=7.
Regions with statistically significant responses were
localized in the cytoarchitectonic probabilistic maps of the
human brain, using the SPM Anatomy Toolbox v1.8 (Eick-
hoff et al. 2005, 2006a, 2007). After locating the PBR and
NBR areas in the group data, a relative signal change anal-
ysis was conducted in ten anatomical areas (BA 18, BA 44,
Fig.1Statistical maps of PBR obtained for a sinlge visualVIS, b
single vestibularVES, and c simultaneousDUAL stimuli condi-
tions. The PBRs areas were marked through the group analysis by
comparing the BOLD response of each stimulus condition with the
BA 45, BA 6, IPC (PFcm), Insula (Ig2), Lobule VI (Hem),
hOC3v (V3v), hOC4 (V4), and hOC5 (V5)) for each con-
dition. The location and extension of the anatomical areas
were derived from the probabilistic cytoarchitectonic maps
embedded in the SPM Anatomy Toolbox. The relative sig-
nal change enables the analysis of differential activation
areas independent of the contrast definitions and statistical
thresholds. The analysis details are described in a previous
Results
Behavioral measures
Just after the experiment, all 21 subjects reported a clear
sensation of body movement only during the periods of
GVS. All they confirmed that during the single visual
and the resting periods this sensation completely disap-
pears. All subjects could also distinguish between the sin-
gle visual stimulation and the simultaneous stimulation.
base (BASE) condition. The statistical maps were superimposed onto
a standard brain template for anatomical interpretation. The results
are shown for a threshold of p<0.05, FDR corrected. Images are
shown in neurological convention
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Table1Coordinates (MNI152) and t values of regions showing PBR in visual and vestibular conditions
Brain region Visual stimulus Vestibular stimulus
x y z t score CA x y z t score CA

Frontal cortex
r Precentral gyrus +12 30 +70 3.28 BA 4a +44 +8 +42 5.18 BA 44
l Precentral gyrus 44 6 +36 5.26 BA 4p 44 +2 +38 3.93
r Inferior frontal gyrus +46 +14 +24 7.57 BA 44,45 +44 +24 +14 5.71 BA 45
l Inferior frontal gyrus 58 +20 +20 4.87 BA 44,45 38 +24 2 5.01
r Middle frontal gyrus +54 +26 +32 7.03 +54 +26 +32 4.44
l Middle frontal gyrus 44 +50 0 4.73 46 +26 +36 6.08 BA 45
r Dorsomedial prefrontal cortex +2 +34 +56 4.34
l Dorsomedial prefrontal cortex 0 +26 +54 4.20 BA 6
l Supplementary motor area 2 +24 +56 4.19 BA 6
Parietal cortex
r Precuneus +12 68 +42 4.15 SPL (7p)
r Supramarginal gyrus +48 38 +34 3.73 IPC (PFcm) +52 38 +36 5.54 IPC (PFcm)
l Supramarginal gyrus 44 36 +26 5.28 IPC (PFcm)
r Inferior parietal lobe +56 38 +50 4.97 IPC (PF) +52 46 +50 4.87 IPC (PFm)
l Inferior parietal lobe 40 52 +58 4.90 SPL (7PC) 46 46 +46 4.48 h (IP2)
l Superior parietal lobe 22 64 +50 4.40 SPL (7A)
r Rolandic operculum +54 28 +22 3.80 IPC (PFcm)
l Rolandic operculum 38 30 +20 5.01 OP1
r Paracentral lobe +6 38 +64 4.33 BA 4a
l Paracentral lobe 6 26 +64 4.32 BA 4a
r Angular gyrus +28 62 +44 4.44 SPL (7A)
Temporal cortex
r Superior temporal gyrus +54 38 +22 5.04 IPC (PFcm)
l Superior temporal gyrus 44 34 +16 4.82 IPC (PFcm)
r Middle temporal gyrus +46 60 +4 4.93 hOC5 (v5)
l Middle temporal gyrus 50 48 +18 3.65 IPC (PGa)
r Inferior temporal gyrus +44 66 4 6.98
Occipital cortex
r Lingual gyrus +24 82 8 7.82 hOC3v (v3v)
l Lingual gyrus 12 76 8 3.80 hOC3v (v3v)
r Fusiform gyrus +38 60 14 6.98 hOC4v (v4)
l Fusiform gyrus 36 70 14 7.82 hOC4v (v4) 40 62 14 4.12
r Superior occipital gyrus +28 66 +34 4.14 hLP1 +24 74 +36 3.97
r Middle occipital gyrus +24 90 +6 9.11 BA 18
l Middle occipital gyrus 22 90 +4 8.85 BA 17 42 68 +4 4.43 hoc5 (v5)
r Inferior occipital gyrus +36 74 10 4.40 hOC4v (v4)
l Inferior occipital gyrus 48 66 12 6.51
Cingulate cortex
r Anterior cingulate +10 +40 +24 3.72
l Anterior cingulate 2 2 +30 5.46
r Middle cingulate +2 10 +35 5.55 +4 8 +32 4.46
l Middle cingulate 6 6 +34 4.79
Insular cortex
r Insula +34 18 +6 3.57 Ig2 +42 +2 14 4.98
l Insula 38 0 6 3.47

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Table1continued
Brain region Visual stimulus Vestibular stimulus
x y z t score CA x y z t score CA
Subcortical and other structures
r Amygdala +24 2 12 4.83 Amyg (SF)
l Amygdala 26 6 12 6.41 Amyg (LB)
r Hippocampus +32 12 20 4.16 Hipp (CA) +34 18 8 4.20 Th-Visual
l Hippocampus 24 32 2 8.43 Hipp (Th-temporal)
r Putamen +34 16 +2 3.45 Ig2 +28 0 0 4.22
l Putamen 26 +6 +4 5.18
r Thalamus +14 32 +2 5.39 Th-temporal +10 14 +2 4.38 Th-Prefrontal
l Thalamus 8 30 0 6.03 Th-Visual/Prefrontal 12 14 +4 4.64 Th-Prefrontal
r Caudate Nucleus +14 2 +16 4.18
r Cerebellum +12 32 20 3.95 Lobules IIV (Hem) +20 40 20 3.67 Lobule V
l Cerebellum 34 74 24 4.85 Lobule VIIa Crus I (Hem) 24 62 28 4.90 Lobule VI (Hem)

Labeling of areas as provided in the cytoarchitectonicprobabilistic maps of the human brain (Eickhoff et al. 2005, 2006a, 2007)
CA cytoarchitectonic area

Some subjects reported that it was easier to focus in the
red cross during the simultaneous condition than dur-
ing the single vestibular condition. The vestibular sensa-
tions induced by GVS during the fMRI experiments were
similar in all subjects; they experienced oscillation of the
entire body at midsagittal axis. Somatosensory effects at
the electrode sites were present in all subjects and were
described mostly as a moderate heating sensation. None of
them reported nausea. The records of eye movements dur-
ing scanning demonstrated that all subjects executed cor-
rectly the instructions.
Positive responses related tosingle stimuli
We have identified characteristic BOLD response in pri-
mary visual and vestibular areas during the pure stimulus
condition. The BOLD response was above 10% for all the
subjects in right and left primary visual cortex during the
VISBASE contrast. The BOLD response was above 10 for
85% of subjects in right PIVC region and for 95% of sub-
jects in left PIVC region during VESBASE contrast. The
VISBASE contrast (Fig.1a; Table1) elicited a PBR in the
inferior occipital gyri, inferior temporal gyrus, hippocam-
pus, amygdala, paracentral lobe and superior parietal lobe.
The VESBASE contrast (Fig.1b; Table1) resulted in PBR
in the precuneus, superior and middle temporal gyri, hypo-
thalamus, supplementary motor area, central sulcus, dorso-
medial prefrontal cortex and caudate nucleus. In addition
to these areas, both conditions separately (VIS-BASE and
VES-BASE) elicited a PBR in the lingual gyrus, fusiform
gyrus, superior and middle occipital gyri, middle and infe-
rior frontal gyri, precentral gyrus, supramarginal gyrus,
insular cortex, inferior parietal lobe, anterior cingulate
gyrus, middle cingulate cortex, cerebellum, putamen and
thalamus.
Positive responses related tosimultaneous stimulation
The results of DUALBASE contrast (Fig.1c; Table2)
showed clusters of PBR in the major areas that are involved
with the visual and the vestibular systems: the lingual
gyrus, fusiform gyrus, superior and middle occipital gyri,
superior, middle and inferior frontal gyri, superior and infe-
rior temporal gyri, hypothalamus, precentral gyrus, supple-
mentary motor area, central sulcus, dorsomedial prefrontal
cortex, insular cortex, inferior parietal lobe, anterior cingu-
lated gyrus, middle cingulate cortex, putamen and caudate
nucleus.
Negative responses
The regions related to NBR are shown in Fig.2 and listed
in Table3. The BASEVIS contrast (Fig.2a) showed NBR
in the calcarine gyrus, lingual gyrus, cuneus, middle occip-
ital gyrus and precuneus. The BASEVES contrast did
not show any NBR regions. The BASEDUAL contrast
(Fig.2b) showed NBR in the calcarine gyrus, lingual gyrus,
cuneus, precuneus, superior parietal lobe, cerebellum and
postcentral gyrus.
Comparison betweensingle andsimultaneous stimulation
The VISDUAL and DUALVIS contrasts did not
show regions with significant differences when com-
pared with single visual and simultaneous conditions.
However, different regions had PBR when comparing

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244 Exp Brain Res (2015) 233:237252

the VESDUAL and DUALVES contrasts. The VES Table2Coordinates (MNI152) and t values of regions showing PBR
DUAL contrast showed significant voxels in the bilateral in the simultaneous condition
lingual gyrus, bilateral calcarine gyrus, right supramar- Brain region Simultaneous stimulation
ginal gyrus, right superior parietal lobule, right middle x y z t score CA
temporal gyrus, right insula, right cuneus, left superior,
middle and inferior frontal gyri, left precentral, left fusi- Frontal cortex
form gyri and bilateral thalamus (Fig.3a; Table4). The r Precentral gyrus +44 +8 +36 4.08 BA 44
DUALVES contrast showed significant voxels in the l Precentral gyrus 38 +4 +20 3.68 BA 44
bilateral middle occipital gyri, bilateral thalamus, the r Superior frontal gyrus +20 +52 +12 4.22
right lingual and inferior occipital gyri, and the left fusi- l Superior frontal gyrus 12 +44 +34 5.09
form gyrus and dorsomedial prefrontal cortex (Fig.3b; r Middle frontal gyrus +40 +30 +42 6.43
Table4). l Middle frontal gyrus 42 +24 +32 3.78 BA 45
The interaction contrast (DUAL(VIS+VES)) r Inferior frontal gyrus +52 +28 +33 5.22
showed areas with significant voxels in the bilateral mid- l Inferior frontal gyrus 44 +24 2 4.08 BA 45
dle occipital gyrus, bilateral anterior cingulate cortex, the r Dorsomedial prefrontal +12 +38 +44 3.94
right lingual and superior frontal gyri, and the left fusiform cortex
gyrus and dorsomedial prefrontal cortex (Fig.4; Table5). l Dorsomedial prefrontal 8 +30 +54 4.73
cortex
The relative BOLD signal change r Supplementary motor +12 22 +50 4.54 BA 6
area
l Supplementary motor 4 +24 +56 4.02 BA 6
The relative BOLD signal changes for the three stimuli area
are shown in Fig.5. The BOLD response was stronger in Parietal cortex
Lobule VI (Hem), hOC3v (V3v), hOC4 (V4), and hOC5 r Inferior parietal lobe +48 54 +52 3.85 IPO (PFcm)
(V5) areas. The BA 44, BA 45, BA 18, hOC3v (V3v), l Rolandic operculum 44 +4 +14 5.07 BA 44
hOC4 (V4) and hOC5 (V5) areas showed an increase in Temporal cortex
the BOLD signal during the VIS condition and a decrease
l Superior temporal gyrus 44 +4 10 4.85
in the BOLD signal during the VES condition. In contrast,
r Inferior temporal gyrus +44 64 4 5.41
the IPC (PFcm), Insula (Ig2) and Lobule VI (Hem) areas
Occipital cortex
showed an increase in the BOLD signal during the VES
r Lingual gyrus +24 82 8 8.23 hOC3v (v3v)
condition and a decrease in BOLD signal during the VIS
r Fusiform gyrus +32 80 4 8.13 hOC4v (v4)
condition. BA 6 did not show a significant change in the
l Fusiform gyrus 26 86 13 7.03 hOC4v (v4)
BOLD signal between the three different conditions. The
r Middle occipital gyrus +24 90 +6 9.60 BA 17
BOLD signal in the DUAL condition was generally in the
l Middle occipital gyrus 40 68 +2 5.12 hOC5 (v5)
same order as the BOLD signal in the VIS and VES condi-
r Inferior occipital gyrus +36 74 10 3.80 hOC4v (v4)
tions for BA 44 and BA 45 regions.
Cingulate cortex
r Anterior cingulate cortex +8 +46 +26 3.90
l Anterior cingulate cortex 8 +44 +12 3.84
Discussion r Middle cingulate cortex +16 18 +42 4.51
l Middle cingulate cortex 12 20 +40 3.58 SPL (5Ci)
In this study, galvanic vestibular stimulation and static Insular cortex
visual stimulus were applied separated and simultaneously r Insular +48 +10 10 4.39
during fMRI experiments. As we are interested in identify- l Insular 32 +2 +10 3.99
ing original regions involved with the interaction process Subcortical structures
during concurrent visual and vestibular input, we chose to l Hippocampus 20 34 +0 6.00 Th-Temporal
do a conservative statistical analysis to avoid false positives r Putamen +28 +16 +4 4.77
even knowing that this may eliminate signal in some acti- l Putamen 26 12 +10 3.98 Th-Motor
vated regions. We chose a FDR p value correction of 0.05 l Caudate nucleus 16 +10 +10 5.08
(Bennett et al. 2009). The results of our study showed that
during conflicting visualvestibular stimulus, the recipro- Labeling of areas as introduced in Table1
cal inhibitory visualvestibular interaction pattern is also CA cytoarchitectonic area

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Exp Brain Res (2015) 233:237252 245

Fig.2Statistical maps of NBR obtained for a sinlge visualVIS, b stimulus condition. The statistical maps were superimposed onto a
single vestibularVES, and c simultaneousDUAL stimuli condi- standard brain template for anatomical interpretation. The results are
tions. The NBRs areas were marked through the group analysis by shown for a threshold of p<0.05, FDR corrected. Images are shown
comparing the BOLD response of base (BASE) condition with each in neurological convention

Table3Coordinates (MNI152) and t values of NBR regions in visual and simultaneous conditions

Brain region Visual stimulus Simultaneous stimulus

x y z t score CA x y z t score CA

Parietal cortex
r Precuneus +22 44 +8 5.67 +24 44 +6 8.31
l Precuneus 14 54 +46 6.19 hlP3 14 48 +50 4.41 SPL (5m)
l Superior parietal lobe 20 50 +66 3.96 SPL (5l)
r Postcentral gyrus +26 46 +66 4.04 SPL (5l)
Occipital cortex
r Calcarine gyrus +26 70 +4 7.28 BA 17 +26 70 +4 7.83 BA 17
l Calcarine gyrus 10 82 +4 10.40 BA 17 10 82 +4 6.79 BA 17
r Lingual gyrus +10 72 +0 7.84 BA 18 +10 72 2 6.58 BA 18
l Lingual gyrus 12 72 6 9.00 hOC4v (v4) 12 78 +0 7.11 BA 18
r Cuneus +12 82 +20 7.68 BA 18 +14 82 +20 6.61 BA 18
l Cuneus 14 76 +26 5.98
r Middle occipital gyrus +52 70 +26 4.19 IPO (PGp)
l Middle occipital gyrus 38 78 +34 4.07 IPO (PGp)
Subcortical and other structures
l Cerebellum 26 54 28 5.10 Lobule VI (Hem)

Labeling of areas as introduced in Table 1

CA cytoarchitectonic area

present. Furthermore, our results show that there is an
interaction function in frontal areas during conflicting vis-
ualvestibular (DUAL) stimuli. This interactive functional-
ity is affected by intermodal associative areas in occipital,
parietal and temporal lobes.
Positive BOLD responses
From our experimental result analyses, it is possible to
infer that the middle and inferior frontal gyri, and the pre-
central gyrus are more related to the interaction process

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246
Fig.3Statistical maps obtained when comparing single and simul-
taneous conditions. a Brain areas that had stronger responses to VES
than to DUAL condition. b Brain areas that had stronger responses
to DUAL than to VES condition. The statistical maps were superim-
existed between the visual and vestibular. The PBR in the
precentral gyrus comprised the frontal eye field. The activa-
tion in the precentral gyrus region is related to eye move-
ments, such as saccades (Anderson et al. 1994; Della-Jus-
tina et al. 2008), smooth pursuit (Petit and Haxby 1999;
Della-Justina et al. 2008), optokinetic nystagmus (Dieterich
et al. 1998) and torsional eye deviation induced during ves-
tibular stimulation (Zink et al. 1998). The precentral gyrus
and the inferior and middle frontal gyri were also found to
be activated in earlier studies in which galvanic vestibu-
lar simulation was performed in healthy volunteers during
fMRI scans (Lobel et al. 1998; Bense et al. 2001; Stephan
et al. 2005). These areas send projections directly to the
vestibular nuclei, the PIVC and the 3aV region (Akbar-
ian et al. 1993, 1994), which, in addition to area 2v, form
a central circuit in the primates vestibular cortex (Guldin
et al. 1992).
The responses for the VES condition in the superior
parietal lobule and supramarginal gyrus were stronger than
in DUAL condition. These two regions are involved in the
interpretation of sensory information (Bremmer 2001) and
spatial orientation (Weeks et al. 1999). From Fig.5, it is
possible to suggest that, therefore, the activation of the
inferior parietal lobe is related more to the vestibular func-
tion than the visual function. Similar results were found by
Bense et al. (2001), Fink et al. (2003), and Stephan et al.
(2005) and, with caloric stimulation, by Bottini et al. (1994)
and Dieterich et al. (2003a). This area has connections to
the PIVC area and belongs to the internal vestibular circuit
described by (Guldin and Grsser 1996). It is interesting to
13
Exp Brain Res (2015) 233:237252
posed onto a standard brain template for anatomical interpretation.
The results are shown for a threshold of p<0.05, FDR corrected.
Images are shown in neurological convention
note that the precuneus has been mainly associated with the
visuospatial circuit, i.e., processing of self-movement or
egomotion (Smith et al. 2012). We found that this region
had a PBR during the single vestibular stimulation.
The temporal area that had stronger response to VES
condition, when compared to BASE condition, was the
middle temporal gyrus. This area is involved in visuomo-
tor processing, such as self-motion perceptions and moving
objects (Barton et al. 1996). It is interesting to note that dif-
ferently from Smith et al. (2012) that subdivided the mid-
dle temporal/medial superior temporal (MT/MST) area into
distinct functional regions and indicated that only the MST
region has vestibular connections, we found that MT region
have also vestibular connections.
As indicated in Tables1 and 2, when comparing VIS ver-
sus DUAL contrasts, we have not identified any region with
differential activity. This might be explained by the neural
activity saturation or BOLD response saturation induced
by the flickering checkerboard stimulation used in the sin-
gle visual stimulus. Another possibility is a vestibulo-ocular
reflex (VOR) suppression that is related to the BOLD activity
during the DUAL condition. An increase in the BOLD signal
in the VIS condition and a decrease in the BOLD signal dur-
ing the VES condition was observed in BA 18, hOC3v (V3v)
and hOC4 (V4) regions (Fig.5a). These results confirm the
involvement of the associative visual areas in processing
visual information. These regions are responsible for global
movement processing (Braddick et al. 2001).
The results also indicated that PBR in the insula was
more strongly related to the vestibula stimulus than the
Exp Brain Res (2015) 233:237252 247

Table4Coordinates (MNI152) and t values for purely vestibular versus simultaneous conditions
Brain region VES>DUAL DUAL>VES
x y z t score CA x y z t score CA

Frontal cortex
l Precentral gyrus 38 0 56 3.46 BA 6
l Superior frontal gyrus 26 4 60 3.03 BA 6
l Middle frontal gyrus 26 12 50 3.13 BA 6
l Inferior frontal gyrus 44 34 16 3.62 BA 45
l Dorsomedial prefrontal cortex 10 52 26 5.49
Parietal cortex
r Supramarginal gyrus 46 30 36 4.44 IPC (PFt)
r Superior parietal lobule 22 56 64 3.39 SPL (7A)
Temporal cortex
r Middle temporal gyrus 48 72 20 5.74 IPC (PGp)
Occipital cortex
l Calcarine gyrus 8 82 4 8.76 BA 17
r Calcarine gyrus 10 76 4 7.11 BA 17
l Lingual gyrus 10 78 0 9.42 BA 18
r Lingual gyrus 10 70 2 8.69 BA 18 24 82 8 7.34 hOC3v (v3v)
l Fusiform gyrus 28 48 8 7.48 26 82 18 7.84 hOC4v (v4)
r Cuneus 14 82 20 8.63 BA 18
r Middle occipital gyrus 26 88 4 9.57
l Middle occipital gyrus 22 90 4 9.27 BA 17
r Inferior occipital gyrus 40 68 16 5.06 hOC4v (Y4)
Insular cortex
r Insula 42 12 12 3.49 Ig2
Subcortical and other structures
r Thalamus 10 22 4 4.42 Th-Premotor 20 32 2 7.74 Th-Temporal
l Thalamus 16 10 12 3.38 Th-Prefrontal 22 32 0 6.17 Th-Temporal
l Cerebellum 10 68 10 8.15 Lobule VI (Hem)

Labeling of areas as introduced in Table1

CA Cytoarchitectonic area

Fig.4Brain areas that had

stronger responses to DUAL
than to VIS or VES conditions.
The statistical maps were super-
imposed onto a standard brain
template for anatomical inter-
pretation. The results are shown
for a threshold of p<0.05, FDR
corrected. Images are shown in
neurological convention

visual stimulus. Previous neuroimaging studies showed that
insula appears activated in response to purely vestibular
sensations (Marcelli et al. 2009; Eickhoff et al. 2006b; Die-
terich et al. 2003b; Fasold et al. 2002; Suzuki et al. 2001).
Furthermore, the area corresponding to PIVC is generally
considered to be the main vestibular cortex as the informa-
tion of other vestibular cortical areas converges into this
area (Guldin and Grsser 1998). Nevertheless, some stud-
ies suggest that although the insular cortex is activated by
vestibular stimulation, the insular may not be critically
involved in vestibular perception (Kahane et al. 2003; Baier
et al. 2013).
Our results showed that the left lobule VI responded
more strongly to the VES condition than to the DUAL

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248 Exp Brain Res (2015) 233:237252

condition. This left-lateralized pattern is in accordance with Table5Coordinates (MNI152) and t values for the interaction
a neuroimaging meta-analysis study of the human cerebel- contrast
lum. Stoodley and Schmahmann (2009) demonstrated that, Brain region DUAL>VIS+VES
in general, the cerebellar functions can be distinguished x y z t score CA
from sensorimotor processing, in the anterior lobe, from
cognitive/emotional processing, in the posterior lobe. Their Frontal cortex
work also showed that spatial tasks had a left-lateralized r Superior frontal gyrus +18 +16 +46 4.84 BA 6
pattern, reflecting crossed cerebrocerebellar projections. l Dorsomedial prefrontal 8 +54 +24 4.98
Moreover, human studies of the cerebellar function that cortex
combined pharmacological manipulation (Shaikh et al. Occipital cortex
2013) and brain imaging (Nigmatullina et al. 2013) demon- r Middle occipital gyrus +26 88 +4 10.13
strated that the cerebellum processes both vestibularocular l Middle occipital gyrus 22 90 +4 8.83 BA 17
reflex and perceptual vestibular function with a degree of r Lingual gyrus +24 82 8 7.52 hOC3v (v3v)
functional and neuroanatomical separation. l Fusiform gyrus 26 82 18 6.92 hOC4v (v4)
Cingulate cortex
Negative BOLD responses r Anteriorcingulate +14 +36 +14 4.65
l Anterior cingulate 8 +46 +4 5.00
The NBR, during visual stimulus, in the occipital cor-
Labeling of areas as introduced in Table1
tex were shown in previous studies (Shmuel et al. 2002;
CA cytoarchitectonic area
Bressler et al. 2007; Pasley et al. 2007; Bianciardi et al.
2011). In general, it is observed that the partial stimula-
tion of the visual field evokes PBR in retinotopic areas of Brain interactions betweenvisual andvestibular systems
the visual cortex while NBR are evoked around the non-
stimulated cortex (Harel et al. 2002; Shmuel et al. 2002, The results presented in Fig.4 and Table5 indicates that
2006). A visualvestibular cortical interaction process the main interaction between the visual and vestibular sys-
was described recently by Seemungal et al. (2013). In tems appears to occur in frontal regions. Specifically, dur-
this study, TMS was used to induce phosphenes to probe ing DUAL stimuli, the right superior frontal gyrus and the
cortical excitability, it was observed that during ves- left dorsomedial prefrontal cortex responded with stronger
tibular stimulation (caloric irrigation), the V5/MT excit- signals to the DUAL than VIS or VES conditions. Schu-
ability decreased and the early visual cortex excitability bert and Szameitat (2003) suggested that PBR of frontal
increased. The authors suggested that vestibular modula- regions, during simultaneous visual-auditory tasks, reflect
tion by the visual motion signals is related with the opti- an increase in neural activity that is associated with con-
mization of visual perception. During a simultaneous current responses induced by the stimulations. So, the pre-
visualvestibular stimulation, the visual stimulus reports frontal PBR effect showed in our work can be associated
no head motion and the vestibular stimulus reports head to the neural activity suppression for competing stimuli.
oscillation to the organs of the vestibular system. Thus, a Similar behavior was found in the dorsolateral prefrontal
NBR in the vestibular-related regions indicate that there is function when studying the working memory (Owen 1997;
a visual modulation over vestibular processing when there DEsposito et al. 1998). The superior frontal gyrus activa-
are visualvestibular conflicting signals. tion that was observed in our study may reflect an executive
Kleinschmidt et al. (2002) described that the activ- function which could be associated with the working mem-
ity levels remained identical in higher-order parieto- and ory network (Boisgueheneuc et al. 2006). For instance,
temporo-occipital areas, whereas decrease was observed choosing the appropriate action command to maintain the
in the motion-sensitive visual areas and in the PIVC dur- static body equilibrium.
ing a visual movement perception stimulus (constant From our results, the anterior cingulate cortex appeared
velocity roll-motion stimulation, with illusory perception to be more related to the visualvestibular integration
of self-motion). In our results, it was observed NBR in processes, as it showed stronger signal to the DUAL than
visual cortex and parietal lobe, which lead to the conclu- VIS or VES conditions. Now, it is interesting to note that
sion that the reciprocal inhibitory visualvestibular inter- in a previous fMRI study, Wall and Smith (2008) identi-
action pattern is also present during conflicting visual fied a cingulate area, the cingulate sulcus visual area
vestibular signals. Although NBR effect does not directly (CSv), which responded strongly to a single optic-flow
reflects an inhibitory mechanism, our results demonstrate stimulus and became unresponsive when the stimuli were
that the neural activity is reduced in this context (Logo- inconsistent with egomotion (movement of the whole
thetis 2002). body). They suggested that the visual cues to egomotion

13
Exp Brain Res (2015) 233:237252 249

Fig.5The relative BOLD

signal changes in the left (LH)
and the right (RH) hemisphere
for: a visualVIS, b vestibu-
larVES, and c simultaneous
(DUAL) stimuli. The ten
selected areas to determine the
relative BOLD signal changes
were labeled using the names
indicated in the probabilistic
cytoarchitectonic maps from
the SPM Anatomy Toolbox
(Eickhoff et al. 2005, 2006a,
2007). BA 18Lingual gyrus;
hOC3v(V3v)Lingual gyrus;
hOC4(V4)Fusiform gyrus;
hOC5(V5)Middle occipital
gyrus; IPC(PFcm)Supra-
marginal gyrus; Insula(Ig2)
Insula; Lobule VI(Hem)
Cerebellum; BA 44Inferior
frontal gyrus; BA 45Middle
frontal gyrus; BA 6Superior
frontal gyrus

are encoded by the CSv and the anterior portion of the
intraparietal sulcus. Cardin and Smith (2010) also dem-
onstrated the involvement of this cingulate area in the
egomotion encoding of visual signals. Naito et al. (2003)
suggested that the activation of the right anterior cingu-
late gyrus is involved with vestibular habituation during
caloric vestibular nystagmus, this cingulate region could
inhibit the VOR during the vestibular habituation. Thus,
our results corroborates with the fact that the cingulate
cortex is involved with the visualvestibular integration
processes.
The PBR effects in frontal regions, during DUAL con-
dition, could actually integrate the visual and vestibular
information to execute action commands. The multisensory
interplay affects not only multisensory convergence zones
but also brain areas that are traditionally considered to be
sensory specific (Driver and Noesselt 2008). This func-
tional integration of the visual and vestibular systems in
the frontal regions may be explained by the feed forward
convergence from lower-level, sensory-specific areas to
higher-order, heteromodal areas. Macaluso and Driver
(2005) suggested that the brain areas that are responsible
for processing different signals modalities can be influ-
enced by the stimulation and distribution of attention. Thus,
in the case of visual and vestibular systems, it appears that
the main interaction function occurs in the frontal areas
and may be influenced by occipital, parietal, and tempo-
ral regions. Indeed, in a recent review article, Lacquaniti

13

250
and coworkers formulated a hypothesis that the integration
of estimating the effects of gravity involves connections
between spatiotemporal visual areas and temporo-parietal-
insular regions, which are involved in multisensory integra-
tion (Lacquaniti et al. 2013).
Conclusion
In this study, galvanic vestibular stimulation and static
visual stimulus were applied separated and simultaneously
during fMRI experiments. The results of our study show
the encephalic regions involved with the static visual stim-
ulation, the vestibular stimulation, and the simultaneous
visualvestibular stimulation. Singular visual stimulation
showed positive BOLD responses in the primary and asso-
ciative visual cortex, while singular vestibular stimulation
led to positive BOLD responses in the PIVC, inferior pari-
etal lobe, superior temporal gyrus, precentral gyrus and in
lobules V and VI of the cerebellar hemisphere. The bimodal
stimulation resulted in additional positive BOLD responses
in the middle and inferior frontal gyri and in the precen-
tral gyrus. Vestibular and somatosensory systems regions
showed negative BOLD responses during the simultaneous
condition; single visual and simultaneous conditions also
led to a negative BOLD response in the visual cortex areas.
Our results not only experimentally confirmed the pres-
ence of a mechanism related to inhibitory visualvestibular
interaction pattern, as suggested by Brandt and cowork-
ers (Brandt et al. 1998), but also demonstrated that this
effect is more evident during static visual and galvanic
vestibular simultaneous stimulation as compared to single
stimulations.
The main contribution of our experimental results is that
the interaction between the visual and vestibular systems
occurs in frontal areas and this integration is influenced by
visual cortex areas and regions of the parietal and tempo-
ral lobes. The evidence of such interaction is only prelimi-
nary and will require further investigation to ensure how
the brain processes and combines the visual and vestibular
information. For instance, studies with an integrative com-
bination of different stimulation methods would be useful.
Acknowledgments The authors are grateful to CAPESCoorde-
nao de Aperfeioamente de Pessoal de Nvel Superiorand
CNPqConselho Nacional de Desenvolvimento Cientco e Tec-
nolgico, Brazilfor financial support.
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