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Brain Research Bulletin 75 (2008) 610618

Research report

The changing face of perceptual rivalry

Trung T. Ngo a,d,e, , Guang B. Liu a,b , Andrew J. Tilley c ,
John D. Pettigrew a , Steven M. Miller d,e,f

Vision Touch and Hearing Research Centre, School of Biomedical Sciences, University of Queensland, Brisbane, Queensland 4072, Australia
b Department of Biological and Physical Sciences, University of Southern Queensland, Toowoomba, Queensland 4350, Australia
c School of Psychology, University of Queensland, Brisbane, Queensland 4072, Australia
d Cauleld Pain Management and Research Centre, Cauleld General Medical Centre, Cauleld, Melbourne, Victoria 3162, Australia
e Brain Stimulation Laboratory, Alfred Psychiatry Research Centre, The Alfred Hospital, Prahran, Melbourne, Victoria 3004, Australia
f Department of Psychological Medicine, Monash University, Clayton, Melbourne, Victoria 3800, Australia
Received 20 March 2007; received in revised form 3 September 2007; accepted 17 October 2007
Available online 20 November 2007

Functional brain-imaging studies of houseface binocular rivalry and Rubins vasefaces illusion have consistently reported face perceptiondependent activity in the right fusiform gyrus. Here we use Rubins illusion and report that activation of the left hemisphere by caloric vestibular
stimulation increases the predominance of the faces percept in a substantial number of test subjects. While partially supporting the brain-imaging
lateralization reports, our findings also challenge these studies by suggesting that neural mechanisms of Rubins illusion cannot be limited to
extrastriate perception-dependent processing. In accordance with our previously proposed interhemispheric switch model, the present findings
support the notion that perceptual rivalry engages high-level cortical structures that mediate unihemispheric attentional selection.
2007 Elsevier Inc. All rights reserved.
Keywords: Ambiguous figures; Rubins figure-ground illusion; Fusiform gyrus; Face processing; Caloric vestibular stimulation; Interhemispheric switching

1. Introduction
When presented with a two-dimensional ambiguous figure
that has two perceptual possibilities, the brain deals with the
conflict by alternating perception between each interpretation,
every few seconds. The most well-known of such stimuli are
the perspective-reversing Necker cube and the figure-ground
alternating Rubins vasefaces illusion (Fig. 1A and B, respectively). Another means of achieving perceptual rivalry is to
simultaneously present different images such as orthogonal gratings, one to each eye, in corresponding retinal locations. During
such binocular rivalry (Fig. 1C), the brain alternately samples
each eyes presented image, also every few seconds. For over a
hundred years, investigators have been interested in the psychophysical features of ambiguous figure rivalry (AFR) and
binocular rivalry (BR). More recently, there has been grow-

Corresponding author. Tel.: +61 3 9076 6834; fax: +61 3 9076 6675.
E-mail addresses: (T.T. Ngo), (S.M. Miller).
0361-9230/$ see front matter 2007 Elsevier Inc. All rights reserved.

ing interest in their underlying neural mechanisms and indeed,

whether they share mechanisms in common.
In the case of BR, the past 15 years has seen the prevailing view of its mechanism undergo revision from low-level,
bottom-up models [4] to high-level, stimulus-representation
interpretations [52], finally developing into an amalgam view
[8]. This amalgam view considers that BR involves a series of
processes implemented by neural mechanisms at different stages
of visual processing. Despite generally supporting this view, Lee
and Blake continue to assert that BR is fundamentally a lowlevel phenomenon that can engage high-level processes under
the right conditions [45]. We, on the other hand, take the opposite stance and consider that the phenomenon is fundamentally
one that occurs via high-level, involuntary attentional selection
mechanisms that can at times engage low-level processes [67].
Given the perpetual swing of the explanatory pendulum for BR
[5], it is not surprising that this dichotomy has been mirrored
in explanations of AFR, by opposing bottom-up and top-down
views that were recently developed into a hybrid model [54].
Along with asserting a high-level emphasis, we earlier proposed a specific high-level mechanism of BR and AFR

T.T. Ngo et al. / Brain Research Bulletin 75 (2008) 610618


Fig. 1. Types of perceptual rivalry and the CVS technique. (A) The Necker cube induces perspective reversals. (B) Rubins vasefaces illusion switches between
figure and ground. (C) Binocular rivalry dichoptically presented orthogonal gratings induce alternating perception of each image. (D) Irrigation of iced water into
the external ear canal (in this case right ear) activates contralateral (in this case left-hemisphere) structures involved in attentional processing (TPA, temporo-parietal
areas, and ACC, anterior cingulate cortex; insular cortex activation not indicated).

the interhemispheric switch (IHS) hypothesis [60,63,76]. This

model suggested that the independent attentional resources of
each hemisphere select one but not the other image/perspective
during perceptual rivalry, and that the perceptual alternations
are mediated by a process of alternating hemispheric activation
(or alternating unihemispheric attentional selection; [60]). We
obtained evidence in support of the IHS model by demonstrating
that unilateral hemispheric activation of attentional structures
with caloric vestibular stimulation (CVS) significantly changed
predominance (i.e., the time spent perceiving one percept relative to its rival) during BR and Necker cube AFR [63]. In these
experiments, left- but not right-hemisphere activation caused the
significant predominance change (see Section 4). Furthermore,
we found that single-pulse transcranial magnetic stimulation
(TMS), applied to the left hemisphere, caused phase-specific
perceptual disruption during BR, thus also supporting the IHS
model. The CVS and TMS data challenge explanations of perceptual rivalry that rely on neural competition occurring locally
(at any level) and that assume synchrony between the hemispheres.
CVS has traditionally been used as a neurodiagnostic tool
[23,81], however more recently it has been applied in a wide
range of cognitive, behavioural and clinical contexts (reviewed
in [64]). Our claim that CVS activates attentional structures is
based on the known functions of brain regions activated by CVS
and on the known effects of CVS in disorders of attention. Thus,
brain-imaging studies of unilateral CVS demonstrate contralateral activation in temporo-parietal cortex, insular cortex and
anterior cingulate cortex [3,1113,20,33,40,95,99], and these
regions have been strongly implicated in attentional processing
(e.g., [14,18,80]). Lesions in any of these regions (usually on the
right side) can result in unilateral attentional neglect [39,48,49]
and remarkably, left-ear/right-hemisphere CVS can temporarily
ameliorate this deficit [86,91]. The effect of CVS on rivalry predominance, as we have demonstrated with four different rivalry
stimuli [63,67], therefore supports three conclusions: (i) rivalry
is mediated by interhemispheric switching (given CVS causes
unilateral hemisphere activation; Fig. 1D), (ii) rivalry involves
mechanisms of (involuntary) attention (given CVS activates cor-

tical regions involved in such functions), and (iii) rivalry engages

high-level mechanisms (given CVS induces high-level cortical
The debate over whether BR occurs at a high or low level
in the visual pathway has been informed by psychophysical
and electrophysiological studies and these have been extensively reviewed elsewhere [4,5,8,52,67,89]. Brain-imaging data
also bear on this debate (see Section 4). Of particular interest among the brain-imaging studies of BR and AFR were two
that specifically addressed face perception during rivalry. Tong
et al. [90] used functional magnetic resonance imaging (fMRI)
to investigate BR between houses and faces, utilizing the face
response-selectivity of the fusiform gyrus (fusiform face area)
in the right hemisphere [37,38,79]. They found right-lateralized
face perception-dependent activation in this region. A recent BR
experiment using affective face stimuli has reported data consistent with this result [101]. The study by Tong et al. [90] also
found perception-dependent activation for house images in bilateral parahippocampal gyri (regions that display non-lateralized
response-selectivity for places and inanimate objects; [21]). On
the background of the study by Tong et al. [90], Andrews et
al. [2] investigated Rubins illusion with fMRI and found that
activity in the right fusiform gyrus was statistically predictive
of face perception (though parahippocampal gyri activation was
not statistically predictive of vase perception). Right-lateralized
fusiform gyrus activation has also recently been demonstrated
during face perception with ambiguous Mooney face images
[1]. These findings suggest that perception-dependent extrastriate activity is either fundamental to resolving what is perceived
during rivalry or is the outcome of other fundamental resolution
In the present experiment, we were particularly interested in
subjecting Rubins illusion to the CVS technique given the brainimaging data described above. In addition to CVS inducing
unilateral temporo-parietal, insular and anterior cingulate activation, it has also been reported that the technique induces bilateral
fusiform gyri deactivation [11]. Thus, even if one rejected the
IHS model of rivalry, it could be argued that CVS applied during
viewing of Rubins illusion should, based on CVS deactivation


T.T. Ngo et al. / Brain Research Bulletin 75 (2008) 610618

of fusiform gyri, nevertheless have an effect on predominance

of faces and vase percepts. Specifically, a shift in favour of the
vase image should occur. Alternatively, if one accepts the IHS
model, a predominance change is also expected, independent of
any effect of fusiform gyri deactivation, but it is not immediately
apparent in which direction this should be. Our previous Necker
cube AFR data [63] suggested that the significant predominance
shifts due to CVS could occur in either direction, implying an
arbitrary selection of perspective-to-hemisphere. On the other
hand, the BR data from that study suggested some influence of
higher-order lateralizing factors on determining which image
was selected by which hemisphere. In a separate paper in preparation, we report on CVS experiments during BR designed
specifically to address this issue of percept-to-hemisphere selection, showing that despite some small influence of higher-order
lateralization factors on the selection of a particular percept by a
particular hemisphere, it remains the case that either hemisphere
can select either percept. It is the parsing of percepts, one to each
hemisphere, irrespective of direction, that is the critical feature
of the IHS model.
Therefore, if during viewing of Rubins illusion a predominance shift is found following right-ear CVS (i.e., following
left-hemisphere high-level activation and bilateral fusiform gyri
deactivation), and the direction of that shift is clearly in favour of
vase perception, the cause of such an effect could either be due
to (i) fusiform gyri deactivation (irrespective of the IHS model),
or (ii) activation of high-level unihemispheric attentional selection mechanisms (in accordance with the IHS model) with the
higher-order lateralizing influence of face perception in the right
hemisphere (leaving the vase percept to the left hemisphere). If,
on the other hand, a predominance shift is found and the direction of that shift is clearly in favour of the faces percept, support
for such a finding could not find basis in bilateral fusiform gyri
deactivation, nor would such a finding, on an IHS interpretation,
be consistent with our previous CVS rivalry studies regarding
the direction of predominance shifts. Finally, if a predominance
shift is found and the direction of shifts is arbitrary or only
minimally biased (i.e., towards faces in some subjects and vase
in others, in an equal or slightly biased fashion), then the IHS

model with an arbitrary or only minimally biased percept-tohemisphere selection process would be the most parsimonious
explanation for such a finding. We report the latter to be the case.
2. Materials and methods
2.1. Subjects
Volunteers were recruited from an on-campus employment website and
advertisements and were paid a small remuneration for their participation.
Thirty-two right-handed male subjects (School of Psychology handedness
questionnaire) aged 1828 years had normal or corrected-to-normal Snellen
visual acuity in both eyes. Exclusion criteria included any psychiatric history,
epilepsy or other neurological disorder, ear disease, vestibular dysfunction or
other significant medical history such as cardiac disease. Subjects were nave
to the experimental hypothesis and written, informed consent was obtained
prior to each experimental session, according to a protocol approved by the
University of Queenslands Medical Research Ethics Committee and in keeping with the National Statement on Ethical Conduct in Research Involving
Humans (1999) issued by the National Health and Medical Research Council of

2.2. Recording AFR

Subjects viewed Rubins illusion (identical to Fig. 1B) which subtended
7.4 (height) 7.7 (width) of visual angle, presented on a matt white surface
100 cm from the subject at eye level (fixation at centre) in a quiet, well lit room.
Participants sat upright and were instructed to record their passive alternating
perceptions (i.e., no preferential response to any percept) using three response
keys: one for the faces percept, one for the vase percept (percept key designations and hand used were counterbalanced across all subjects), and the third
for undecided or indeterminate percepts or if the subjects fixation deviated.
The third response option was not included in analyses.
Subjects underwent one half-hour recording schedule consisting of three
6-min blocks interspersed with 5-min rest periods at the start of all three experimental sessions (Fig. 2). Each block of recording consisted of three 100-s trials
separated by 60-s breaks. In the initial control sessions, blocks 3 and 4 were
separated by a non-intervention 5-min rest period for all subjects, while the
subsequent sessions counterbalanced left-ear (right-hemisphere) and right-ear
(left-hemisphere) CVS across all subjects. In these latter sessions, the postCVS recording schedule was the same as the half-hour pre-CVS schedule
(and similarly for the control sessions). Both the PC-compatible online rivalry
recording and offline analysis programs were generated with MATLAB software
(The MathWorks, Inc., Natick, MA, USA). All within-group statistical analyses
employed two-tailed Wilcoxon signed-ranks test ( = 0.05).

Fig. 2. Protocol for recording AFR. Subjects recorded AFR according to this protocol for all three experimental sessions of approximately 1 h each (on separate days).
An initial control session was followed by counterbalanced left-ear CVS (i.e., right-hemisphere activation) and right-ear CVS (left-hemisphere activation) sessions
across all subjects. Monaural stimulation was delivered around 1 min after the end of baseline AFR recording. The duration until the resumption of post-stimulation
AFR recording was about 5 min. To assess for a CVS effect, the absolute magnitude of change in predominance (i.e., |log predominance|) between blocks 2 and
3 (random variation) was compared with that between blocks 3 and 4 (random variation and CVS effect).

T.T. Ngo et al. / Brain Research Bulletin 75 (2008) 610618


2.3. CVS procedure

A medical officer screened all volunteers for any signs of ear disease or cerumen impaction before participation. An explanation of the CVS procedure was
repeated to subjects, having earlier been given via an information sheet. They
were instructed to report the expected signs of stimulation onset (see below).
Participants were reclined with their mid-sagittal plane vertical and head positioned 30 from the horizontal plane, thus placing the lateral semicircular canal
in the vertical plane. Cold (iced) water was slowly instilled using a 50-ml syringe
with a short piece of silastic tubing attached and carefully placed in the external ear canal close to the tympanic membrane without touching it (Fig. 1D).
Irrigation stopped once the subject reported vertigo following about 2030 ml,
usually just after nystagmus (slow-phase direction ipsilateral to the stimulated
ear) was detected by the experimenter. The refluent water was collected in a
plastic container placed on the subjects shoulder. After reports that vertigo had
ceased and there was no further observable nystagmus, participants sat upright
for the resumption of data collection.

3. Results
The predominance ratio for each recording block was calculated as the total time in seconds perceiving one image divided
by the total time in seconds perceiving the other image, after
exclusion of the third response option. The ratios were then
log-transformed to account for the disproportionate numerical
representation in predominance (i.e., >1 for one percept cf. <1
for the other). The first (training) block was excluded before
analysis. To assess the effect of CVS on predominance, the absolute magnitude of change in predominance (i.e., irrespective of
the direction of change), or |log predominance|, between two
pre-CVS blocks of AFR (blocks 2 and 3) was compared to that
between the blocks immediately before and after CVS (blocks 3
and 4). Thus, a significantly greater predominance change across
Blocks 34 (experimental effect plus random variation; Fig. 2A)
than across Blocks 23 (random variation alone; Fig. 2A) would
indicate an effect from the CVS.
Left-hemisphere activation by right-ear CVS significantly
changed predominance during viewing of Rubins illusion
(Fig. 3; Mean |log predominance| for Blocks 23 and 34,
respectively = 0.099 and 0.197, W = 238, p = <0.05). Righthemisphere activation by left-ear CVS did not induce a
significant shift in the predominance of faces and vase percepts
(Mean |log predominance| for Blocks 23 and 34, respectively = 0.102 and 0.177, W = 176, p = 0.10). Similarly, the no
stimulation condition did not reveal significant predominance
changes (Mean |log predominance| for Blocks 23 and 34,
respectively = 0.127 and 0.099, W = 176, p = 0.10).
To assess the direction of predominance shifts in the significant left-hemisphere stimulation condition, inspection of
subjects individual data was performed on the 22 observers
who had larger predominance changes after CVS than between
the two pre-stimulation blocks (Fig. 3A; inverted triangles
above zero). Nine had post-CVS predominance shifts towards
the faces percept and thirteen had post-CVS predominance
shifts in favour of the vase percept. There was no significant
difference found between subjects favouring the faces following CVS and those favouring the vase following CVS, with
respect to the magnitude of the predominance changes (Mean
 |log predominance| = 0.284 and 0.114, respectively; two-

Fig. 3. Left-hemisphere activation significantly changes predominance of

Rubins illusion. (A) The points above zero depict individuals with larger predominance shifts after CVS than in baseline viewing (22 subjects in total), while
those below zero indicate larger baseline shifts in predominance than that following CVS. For each experimental condition, the data points were arranged
in descending order of magnitude, thus individual subjects do not necessarily
correspond between conditions. The points were calculated by subtracting the
absolute magnitude of change in the log-transformed ratio of predominance (i.e.,
|log predominance|) for blocks 2 and 3 from that of blocks 3 and 4 in each
subject. These pairs of values are shown for the significant left-hemisphere stimulation condition (B) wherein each pair corresponds to an individual subjects
data arranged according to descending order of the Block 34 magnitudes.

tailed MannWhitney U = 40, p = 0.23). However, the former did

tend to display larger post-CVS shifts than the latter (see Fig. 4).
To assess the effect of CVS on the distribution of perceptual intervals, gamma function distributions were calculated
separately for Block 3 (immediately pre-CVS) and Block 4
(immediately post-CVS) of the left-hemisphere stimulation
condition in the same 22 subjects using f(x) = r / (r)xr1
exp(x) where (r) = (r 1)! (following [53]). As depicted
in Fig. 5, the fit of the gamma curve before and after CVS
shows that the distribution of perceptual intervals during
Rubins illusion is not affected by significant CVS-induced
changes in predominance. Finally, analysis of the number of
space bar responses before and after CVS (left-hemisphere
activation) revealed no significant difference across all 32
subjects (Mean for blocks 3 and 4, respectively = 6.56 and
6.00, W = 99, p = 0.18) or across the 22 subjects (Mean for


T.T. Ngo et al. / Brain Research Bulletin 75 (2008) 610618

Fig. 5. CVS and the distribution of Rubins illusion perceptual intervals. The
normalised frequency histograms both before and after CVS (left-hemisphere
stimulation) are well-described by the gamma curve given the coefficient of
determination (R2 ) values are 0.95. (A) The fit to the pre-CVS distribution of
perceptual intervals in Rubins illusion is consistent with evidence in the literature on AFR (see Section 4). (B) The fit to the post-CVS distribution shows that
despite the demonstrated significant CVS-induced changes in predominance,
the distribution of intervals is not affected by the CVS intervention. This finding
is consistent with a similar lack of CVS effect on the gamma function curves for
BR with orthogonal oblique gratings in our earlier studies (unpublished observations from the data in [63]). Thus, these findings, in addition to the CVS
predominance findings, provide further evidence for the view that AFR and
BR share some degree of common neural mechanism (see Section 4). The following gamma function parameters are reported: N = total number of intervals,
n=number of subjects, = mean interval duration (seconds) across all subjects,
= standard deviation of all intervals, R2 = coefficient of determination. and r
are parameters which specify the values that generate a gamma curve of best fit
to the normalised distribution of interval durations.

Fig. 4. Individual subjects perceptual predominance for each block of their

left-hemisphere stimulation condition. Shown are data for the 22 subjects who
had larger post-CVS than pre-CVS predominance changes, corresponding to the
same 22 subjects above the zero line in Fig. 3A. The log-transformed predominance ratio is calculated for each block of AFR responses. The dotted and full
lines indicate subjects who showed post-CVS predominance shifts towards the
faces percept and vase percept, respectively. Each subjects plot is arranged corresponding to their descending order of  |log predominance| values depicted
in Fig. 3A (left-hemisphere activation condition). For ease of comparison, the
plotted lines for all subjects are aligned such that from Block 3 to Block 4,
the slope runs upward. Hence, those subjects in whom this slope actually runs
downward have had their entire plots inverted for the purposes of comparison.
The subjects with inverted plots are indicated by an asterisk. Each subjects data
points are plotted on the same ordinate scale of logged predominance values but
these plots have been stacked rather than superimposed, also for ease of comparison. As such, the ordinate axis is not numerically labeled but a log-predominance
scale bar is given.

blocks 3 and 4, respectively = 6.05 and 5.14, W = 60, p = 0.16)

who showed larger post-CVS than pre-CVS predominance
4. Discussion
The finding of a significant CVS effect on perceptual predominance of Rubins illusion is consistent with our previous CVS
studies of horizontal/vertical BR, orthogonal oblique BR, AFR
with the Necker cube [63], and rivalling coherent images with
DazCaneja stimuli [67]. This finding demonstrates that like
BR and perspective-reversing AFR according to our previous
work, figure-ground AFR is also mediated by interhemispheric
switching, with the involvement of high-level attentional selection mechanisms. In all these experiments, including the current
study, predominance is significantly affected by left- but not

T.T. Ngo et al. / Brain Research Bulletin 75 (2008) 610618

right-hemisphere CVS, a finding we interpret on the basis

of reported hemispheric asymmetries of spatial representation
[30,60] or reported hemispheric asymmetries during BR transitions [55,63]. Despite attempts to falsify the IHS hypothesis
(see [7274] and our response in [67]), no convincing alternative explanations for our consistent CVS findings have yet to be
Furthermore, the present finding of CVS-induced face preference in a substantial number of subjects (i.e., 40% of those
subjects who showed a greater CVS-induced predominance
change than random variation) challenges the brain-imaging
data of Tong et al. [90] and Andrews et al. [2]. While we acknowledge that the slight bias in direction of CVS-induced shifts
favouring vase over faces perception (60% cf. 40%, respectively)
does provide partial support for the face-lateralization finding in
those imaging studies, we consider the increase in predominance
of faces in 40% of subjects to be of far more interest. How could
such a finding be reconciled with the brain-imaging data? As
referred to in Section 1, we suggest the IHS model provides the
most parsimonious explanation.
According to the imaging studies of BR and AFR involving face percepts, and the imaging studies of CVS showing
contralateral activation (temporo-parietal, anterior cingulate and
insular cortex) and bilateral fusiform gyri deactivation, the findings of the present study are best explained by an IHS model
of rivalry with an arbitrary or only minimally biased percept-tohemisphere selection process. The possibility that CVS-induced
bilateral deactivation of fusiform gyri is relevant to the present
results is not excluded by our data but can be considered
unlikely because this interpretation does not predict that a substantial number of subjects would spend more time perceiving
faces following CVS. Similarly, an IHS model interpretation
with a strong higher-order influence of percept-to-hemisphere
selection (i.e., with right-lateralization of the faces percept and
consequent left-lateralization of the vase percept), also does not
account well for the finding of a substantial number of subjects
perceiving more of the faces percept following left-hemisphere
Thus, it seems that in 40% of subjects, the left hemisphere
selects the faces image despite the common finding of rightlateralization for face processing and the demonstration of
right-lateralized, face perception-dependent activity in fusiform
gyrus with Rubins illusion. This interpretation does not doubt
the existence of right-lateralized face processing during viewing
of Rubins illusion. Rather, it argues that other high-level resolution processes must also exist during AFR (as during BR), in
addition to the brain-imaging documented extrastriate activity.
These other processes appear to involve attentional selection of
the faces percept by the left hemisphere in a substantial number
of cases.
It could alternatively be argued that the present findings are
somehow explained by qualifications to the notion of rightlateralized extrastriate face processing, including for example,
the fact that bilateral fusiform gyri activation was found during face perception in one of the four subjects in Tong et al.s
[90] study, in another BR experiment using face stimuli ([55];
see also [35]), and in one study with ambiguous Mooney face


images ([58] cf. [1]). However, it is left-lateralized fusiform

gyrus activity, rather than bilateral activity, that would need to
be demonstrated to support any such appeal. That is, this line
of reasoning would require the demonstration of left-lateralized
fusiform gyrus activation during face processing in a substantial number of subjects scanned. No studies have reported such
activation patterns.
Eye-movement explanations also do not provide an adequate account of the present findings because any effect of
residual nystagmus from CVS would equally affect faces and
vase perception and thus cannot explain predominance shifts
in either direction. Such interpretations were ruled out in our
previous work by demonstrating the same CVS findings when
BR involved horizontal/vertical gratings and orthogonal oblique
gratings [63]. Earlier studies showing that BR and AFR could
occur with afterimages [16,26,43,57] or when the stimuli were
retinally-stabilised [7,82,87] similarly cast doubt on significant
effects of eye movements, though recent work has found that saccades are linked with perceptual transitions in both rivalry types
[19,92]. We consider the effect of eye movements an unlikely
account of our present findings and suggest instead that the
data argue for a resolution mechanism separate to extrastriate
processing which (i) occurs at a high-level, (ii) engages mechanisms of (involuntary) attention, and (iii) is consistent with
the well-documented unilateral activation induced by CVS. The
mechanism that accommodates all these criteria is interhemispheric switching.
However, significant questions remain. Other brain-imaging
studies on BR in humans have shown low-level perceptiondependent activity (in primary visual cortex and lateral
geniculate nucleus; [28,29,44,46,47,78,88,104]). Such studies
though, are not supported by low-level single-unit recordings in macaque monkeys that demonstrate predominantly
presentation-dependent rather than perception-dependent activity at this level [50]. A recent local field potential study of
generalized flash suppression in macaque monkeys may shed
some light on this apparent discrepancy [102], although the
investigators acknowledge the need for further studies on the
relationship between microelectrode neural signals and the fMRI
blood oxygenation level-dependent signal. Nonetheless, the IHS
model has yet to be assessed with brain-imaging techniques. To
do so would require each subjects data be analysed separately
and likewise for each direction of perceptual switch. Such experiments should also consider high-level brain regions involved
in attentional selection as regions of interest, including cortical
areas activated by CVS. Also warranted are further studies on the
relationship between mechanisms of attention and mechanisms
of visual consciousness during BR and AFR (see [42]), however
such investigations are not without substantial methodological
challenges [60,61,67].
Our present and previous demonstrations of significant CVSinduced predominance effects on perceptual rivalry support
the intuitions of early investigators such as Wheatstone [100]
and Helmholtz [31] who likened BR and AFR to processes
of (involuntary) attention. Voluntary and involuntary attention
mechanisms during rivalry have recently been the subject of substantial investigation (e.g., [9,17,27,36,47,59,65,68,75,9294];


T.T. Ngo et al. / Brain Research Bulletin 75 (2008) 610618

see also [67]). Such studies however, commonly fail to acknowledge the support provided by our series of CVS experiments for
a role of (involuntary) attention mechanisms in rivalry [63,67].
These CVS studies, including the present experiment, not only
support an involuntary attention interpretation of rivalry, but also
add weight to the view that BR and AFR share some degree of
common neural mechanism.
The notion of a common neural mechanism(s) for AFR
and BR is supported by psychophysical similarities reported
for the two rivalry types. Although it has been known for a
long time that BR and AFR are not subject to complete voluntary control [16,31,100], early comparisons suggested that
the influence of voluntary attention on the former is less than
that for the latter [25,57,98]. Recent work has demonstrated
support for [59,93,94], but also questioned, this contention
[27,59,94]. A survey of the respective literatures and of studies directly comparing AFR and BR suggest more similarities
than differences between the two types of perceptual rivalry.
The similarities include, for example: (i) their temporal dynamics [7,10,24,53,66] cf. [15]); (ii) detection thresholds for test
probes presented to the suppressed image/unperceived percept
[6,52,96,103]; (iii) effects on predominance during intermittent
stimulus presentation [51,56,6971,85]; (iv) intra-individual
consistency in response parameters such as alternation rate and
voluntary modulation of percept dominance [93]; (v) effects on
rate when concurrently engaging in an attentional or cognitive
task [75,83,84,97]; (vi) brain regions activated during perceptual
transitions [34,41,55]; (vii) brain regions activated during perceptual states [2,90,101]; (viii) effects of CVS on predominance
[63,67, present study] and lack of CVS effects on the gamma
function distribution (present study and unpublished observations from the data in [63]); and (ix) slow rivalry rate in bipolar
disorder but not in schizophrenia [22,32,62,77]. Our present and
previous CVS findings also support the view that the mechanism in common between BR and AFR is interhemispheric
In conclusion, we have demonstrated a significant effect
of left-hemisphere CVS on perceptual predominance during
viewing of Rubins illusion, with a substantial number of subjects favouring faces-perception following the intervention. This
finding suggests that neural mechanisms of AFR with Rubins
illusion are not limited to the right-sided perception-dependent
extrastriate activity shown in brain-imaging studies, and must
also involve other resolution processes. The high-level unilateral
activation of attentional structures induced by CVS suggests that
these other resolution processes are high-level unihemispheric
attentional selection mechanisms that alternate in accordance
with perceptual alternations (i.e., interhemispheric switching).
The present study brings to five the total number of experiments
in which CVS has been shown to induce a significant change
in predominance during perceptual rivalry. Our work cumulatively represents well over a hundred subjects reporting their
rivalry perceptions and the findings consistently support the IHS
model, one of the few tangible and readily testable models of
rivalry in the current literature. We encourage attempts at falsification of the model and have elsewhere [67] detailed strategies to
this end.

Conicts of interest
None declared.
We thank two anonymous reviewers for helpful comments
on an earlier version of the manuscript, and all experimental
volunteers for their participation.
[1] T.J. Andrews, D. Schluppeck, Neural responses to Mooney images reveal
a modular representation of faces in human visual cortex, Neuroimage 21
(2004) 9198.
[2] T.J. Andrews, D. Schluppeck, D. Homfray, P. Matthews, C. Blakemore,
Activity in the fusiform gyrus predicts conscious perception of Rubins
vaseface illusion, Neuroimage 17 (2002) 890901.
[3] S. Bense, P. Bartenstein, S. Lutz, T. Stephan, M. Schwaiger, T. Brandt,
M. Dieterich, Three determinants of vestibular hemispheric dominance
during caloric stimulation. A positron emission tomography study, Ann.
N. Y. Acad. Sci. 1004 (2003) 440445.
[4] R. Blake, A neural theory of binocular rivalry, Psychol. Rev. 96 (1989)
[5] R. Blake, A primer on binocular rivalry, including current controversies,
Brain Mind 2 (2001) 538.
[6] R. Blake, J. Camisa, On the inhibitory nature of binocular rivalry
suppression, J. Exp. Psychol. Human Percept. Perform. 5 (1979)
[7] R. Blake, R. Fox, C. McIntyre, Stochastic properties of stabilizedimage binocular rivalry alternations, J. Exp. Psychol. 88 (1971)
[8] R. Blake, N.K. Logothetis, Visual competition, Nat. Rev. Neurosci. 3
(2002) 1321.
[9] Y.S. Bonneh, M. Pavlovskaya, H. Ring, N. Soroker, Abnormal binocular
rivalry in unilateral neglect: evidence for a non-spatial mechanism of
extinction, Neuroreport 15 (2004) 473477.
[10] A. Borsellino, A. de Marco, A. Allazetta, S. Rinesi, B. Bartolini, Reversal
time distribution in the perception of visual ambiguous stimuli, Kybernetik 10 (1972) 139144.
[11] G. Bottini, H.-O. Karnath, G. Vallar, R. Sterzi, C.D. Frith, R.S.J.
Frackowiak, E. Paulesu, Cerebral representations for egocentric space.
Functionalanatomical evidence from caloric vestibular stimulation and
neck vibration, Brain 124 (2001) 11821196.
[12] G. Bottini, E. Paulesu, R. Sterzi, E. Warburton, R.J.S. Wise, G. Vallar,
R.S.J. Frackowiak, C.D. Frith, Modulation of conscious experience by
peripheral sensory stimuli, Nature 376 (1995) 778781.
[13] G. Bottini, R. Sterzi, E. Paulesu, G. Vallar, S.F. Cappa, F. Erminio, R.E.
Passingham, C.D. Frith, R.S.J. Frackowiak, Identification of the central
vestibular projections in man: a positron emission tomography activation
study, Exp. Brain Res. 99 (1994) 164169.
[14] M.M. Botvinick, J.D. Cohen, C.S. Carter, Conflict monitoring and anterior cingulate cortex: an update, Trends Cogn. Sci. 8 (2004) 539546.
[15] J.W. Brascamp, R. van Ee, W.R. Pestman, A.V. van den Berg, Distributions of alternation rates in various forms of bistable perception, J. Vis. 5
(2005) 287298.
[16] B.B. Breese, On inhibition, Psychol. Monogr. 3 (1899) 165.
[17] S.C. Chong, R. Blake, Exogenous attention and endogenous attention
influence initial dominance in binocular rivalry, Vision Res. 46 (2006)
[18] M. Corbetta, G.L. Shulman, Control of goal-directed and stimulus-driven
attention in the brain, Nat. Rev. Neurosci. 3 (2002) 201215.
[19] W. Einhauser, K.A.C. Martin, P. Konig, Are switches in perception of
the Necker cube related to eye position? Eur. J. Neurosci. 20 (2004)

T.T. Ngo et al. / Brain Research Bulletin 75 (2008) 610618

[20] M. Emri, M. Kisely, Z. Lengyel, L. Balkay, T. Marian, L. Miko, E.
Toth, Cortical projection of peripheral
Berenyi, I. Sziklai, L. Tron, A.
vestibular signalling, J. Neurophysiol. 89 (2003) 26392646.
[21] R. Epstein, N. Kanwisher, A cortical representation of the local visual
environment, Nature 392 (1998) 598601.
[22] J.H. Ewen, The psychological estimation of the effects of certain drugs
upon the syntonic and schizophrenic psychoses. With a brief enquiry
into a physiological basis of temperament, J. Mental Sci. 77 (1931)
[23] T.D. Fife, R.J. Tusa, J.M. Furman, D.S. Zee, E. Frohman, R.W. Baloh,
T. Hain, J. Goebel, J. Demer, L. Eviatar, Assessment: vestibular testing
techniques in adults and children. Report of the therapeutics and technology assessment subcommittee of the American Academy of Neurology,
Neurology 55 (2000) 14311441.
[24] R. Fox, J. Herrmann, Stochastic properties of binocular rivalry alternations, Percept. Psychophys. 2 (1967) 432436.
[25] R.W. George, The significance of the fluctuations experienced in observing ambiguous figures and in binocular rivalry, J. Gen. Psychol. 15 (1936)
[26] R. Gregory, The Intelligent Eye, McGraw Hill, New York, 1970.
[27] S. Hancock, T.J. Andrews, The role of voluntary and involuntary attention
in selecting perceptual dominance during binocular rivalry, Perception 36
(2007) 288298.
[28] J.-D. Haynes, R. Deichmann, G. Rees, Eye-specific effects of binocular rivalry in the human lateral geniculate nucleus, Nature 438 (2005)
[29] J.-D. Haynes, G. Rees, Predicting the stream of consciousness from activity in human visual cortex, Curr. Biol. 15 (2005) 13011307.
[30] K.M. Heilman, T. Van Den Abell, Right hemisphere dominance for attention: the mechanism underlying hemispheric asymmetries of inattention
(neglect), Neurology 30 (1980) 327330.
[31] H.L.F. von Helmholtz, Handbuch der Physiologischen Optik, Voss,
Leipzig, 1867. A. Gullstrand, J. von Kries, W. Nagel (Eds.), third German
ed., Voss, Hamburg, 1910. J.P.C. Southall (Ed.), Helmholtzs Treatise on
Physiological Optics, Translated from the Third German Edition, Dover,
New York, 1962.
[32] J. Hunt, J.P. Guilford, Fluctuation of an ambiguous figure in dementia
praecox and in manic-depressive patients, J. Abnormal Soc. Psychol. 27
(1933) 443452.
[33] I. Indovina, V. Maffei, G. Bosco, M. Zago, E. Macaluso, F. Lacquaniti,
Representation of visual gravitational motion in the human vestibular
cortex, Science 308 (2005) 416419.
[34] T. Inui, S. Tanaka, T. Okada, S. Nishizawa, M. Katayama, J. Konishi,
Neural substrates for depth perception of the Necker cube; a functional
magnetic resonance imaging study in human subjects, Neurosci. Lett. 282
(2000) 145148.
[35] Y. Jiang, S. He, Cortical responses to invisible faces: dissociating
subsystems for facial-information processing, Curr. Biol. 16 (2006)
[36] A.P. Kamphuisen, R.J.A. van Wezel, R. van Ee, Inter-ocular transfer of
stimulus cueing in dominance selection at the onset of binocular rivalry,
Vision Res. 47 (2007) 11421144.
[37] N. Kanwisher, J. McDermott, M.M. Chun, The fusiform face area: a
module in human extrastriate cortex specialized for face perception, J.
Neurosci. 17 (1997) 43024311.
[38] N. Kanwisher, G. Yovel, The fusiform face area: a cortical region specialized for the perception of faces, Philos. Trans. R. Soc. Lond. B Biol.
Sci. 361 (2006) 21092128.
[39] H.-O. Karnath, M.F. Berger, W. Kuker, C. Rorden, The anatomy of spatial
neglect based on voxelwise statistical analysis: a study of 140 patients,
Cereb. Cortex 14 (2004) 11641172.
[40] M. Kisely, M. Emri, Z. Lengyel, B. Kalvin, G. Horvath, L. Tron, L.
Toth, Changes in brain activation caused by caloric
Miko, I. Sziklai, A.
stimulation in the case of cochleovestibular denervation PET study,
Nucl. Med. Commun. 23 (2002) 967973.
[41] A. Kleinschmidt, C. Buchel, S. Zeki, R.S.J. Frackowiak, Human brain
activity during spontaneously reversing perception of ambiguous pictures,
Proc. R. Soc. Lond. B Biol. Sci. 265 (1998) 24272433.


[42] C. Koch, N. Tsuchiya, Attention and consciousness: two distinct brain

processes, Trends Cogn. Sci. 11 (2007) 1622.
[43] L.C. Lack, The role of accommodation in the control of binocular rivalry,
Percept. Psychophys. 10 (1971) 3842.
[44] S.-H. Lee, R. Blake, V1 activity is reduced during binocular rivalry, J.
Vis. 2 (2002) 618626.
[45] S.-H. Lee, R. Blake, A fresh look at interocular grouping during binocular
rivalry, Vision Res. 44 (2004) 983991.
[46] S.-H. Lee, R. Blake, D.J. Heeger, Traveling waves of activity in primary
visual cortex during binocular rivalry, Nat. Neurosci. 8 (2005) 2223.
[47] S.-H. Lee, R. Blake, D.J. Heeger, Hierarchy of cortical responses underlying binocular rivalry, Nat. Neurosci. 10 (2007) 10481054.
[48] F.S. Leibovitch, S.E. Black, C.B. Caldwell, P.L. Ebert, L.E. Ehrlich,
J.P. Szalai, Brainbehavior correlations in hemispatial neglect using
CT and SPECT: the Sunnybrook stroke study, Neurology 50 (1998)
[49] F.S. Leibovitch, S.E. Black, C.B. Caldwell, A.R. McIntosh, L.E. Ehrlich,
J.P. Szalai, Brain SPECT imaging and left hemispatial neglect covaried
using partial least squares: the Sunnybrook stroke study, Hum. Brain
Mapp. 7 (1999) 244253.
[50] D.A. Leopold, N.K. Logothetis, Activity changes in early visual cortex
reflect monkeys percepts during binocular rivalry, Nature 379 (1996)
[51] D.A. Leopold, M. Wilke, A. Maier, N.K. Logothetis, Stable perception
of visually ambiguous patterns, Nat. Neurosci. 5 (2002) 605609.
[52] N.K. Logothetis, Single units and conscious vision, Philos. Trans. R. Soc.
Lond. B Biol. Sci. 353 (1998) 18011818.
[53] N.K. Logothetis, D.A. Leopold, D.L. Sheinberg, What is rivalling during
binocular rivalry? Nature 380 (1996) 621624.
[54] G.M. Long, T.C. Toppino, Enduring interest in perceptual ambiguity: alternating views of reversible figures, Psychol. Bull. 130 (2004)
[55] E.D. Lumer, K.J. Friston, G. Rees, Neural correlates of perceptual rivalry
in the human brain, Science 280 (1998) 19301934.
[56] A. Maier, M. Wilke, N.K. Logothetis, D.A. Leopold, Perception of temporally interleaved ambiguous patterns, Curr. Biol. 13 (2003) 10761085.
[57] W. McDougall, III. The physiological factors of the attention-process
(III.), Mind 12 (1903) 473488.
[58] T.J. McKeeff, F. Tong, The timing of perceptual decisions for ambiguous
face stimuli in the human ventral visual cortex, Cereb. Cortex 17 (2007)
[59] M. Meng, F. Tong, Can attention selectively bias bistable perception?
Differences between binocular rivalry and ambiguous figures, J. Vis. 4
(2004) 539551.
[60] S.M. Miller, Binocular rivalry and the cerebral hemispheres: with a note
on the correlates and constitution of visual consciousness, Brain Mind 2
(2001) 119149.
[61] S.M. Miller, On the correlation/constitution distinction problem (and
other hard problems) in the scientific study of consciousness, Acta Neuropsychiatr. 19 (2007) 159176.
[62] S.M. Miller, B.D. Gynther, K.R. Heslop, G.B. Liu, P.B. Mitchell, T.T.
Ngo, J.D. Pettigrew, L.B. Geffen, Slow binocular rivalry in bipolar disorder, Psychol. Med. 33 (2003) 683692.
[63] S.M. Miller, G.B. Liu, T.T. Ngo, G. Hooper, S. Riek, R.G. Carson, J.D.
Pettigrew, Interhemispheric switching mediates perceptual rivalry, Curr.
Biol. 10 (2000) 383392.
[64] S.M. Miller, T.T. Ngo, Studies of caloric vestibular stimulation: implications for the cognitive neurosciences, the clinical neurosciences and
neurophilosophy, Acta Neuropsychiatr. 19 (2007) 183203.
[65] J.F. Mitchell, G.R. Stoner, J.H. Reynolds, Object-based attention determines dominance in binocular rivalry, Nature 429 (2004) 410413.
[66] T. Murata, N. Matsui, S. Miyauchi, Y. Kakita, T. Yanagida, Discrete
stochastic process underlying perceptual rivalry, Neuroreport 14 (2003)
[67] T.T. Ngo, G.B. Liu, A.J. Tilley, J.D. Pettigrew, S.M. Miller, Caloric
vestibular stimulation reveals discrete neural mechanisms for coherence
rivalry and eye rivalry: a meta-rivalry model, Vision Res. 47 (2007)


T.T. Ngo et al. / Brain Research Bulletin 75 (2008) 610618

[68] T.L. Ooi, Z.J. He, Surface representation and attention modulation mechanisms in binocular rivalry, in: D. Alais, R. Blake (Eds.), Binocular Rivalry,
MIT Press, Cambridge, MA, 2005, pp. 117135.
[69] J. Orbach, D. Ehrlich, H.A. Heath, Reversibility of the Necker cube: I.
An examination of the concept of satiation of orientation, Percept. Mot.
Skills 17 (1963) 439458.
[70] J. Orbach, D. Ehrlich, E. Vainstein, Reversibility of the Necker cube: III.
Effects of interpolation on reversal rate of the cube presented intermittently, Percept. Mot. Skills 17 (1963) 571582.
[71] J. Orbach, E. Zucker, R. Olson, Reversibility of the Necker cube: VII.
Reversal rate as a function of figure-on and figure-off durations, Percept.
Mot. Skills 22 (1966) 615618.
[72] R.P. OShea, P.M. Corballis, Binocular rivalry between complex stimuli
in split-brain observers, Brain Mind 2 (2001) 151160.
[73] R.P. OShea, P.M. Corballis, Binocular rivalry in split-brain observers, J.
Vis. 3 (2003) 610615.
[74] R.P. OShea, P.M. Corballis, Visual grouping on binocular rivalry in a
split-brain observer, Vision Res. 45 (2005) 247261.
[75] C.L.E. Paffen, D. Alais, F.A.J. Verstraten, Attention speeds binocular
rivalry, Psychol. Sci. 17 (2006) 752756.
[76] J.D. Pettigrew, Searching for the switch: neural bases for perceptual
rivalry alternations, Brain Mind 2 (2001) 85118.
[77] J.D. Pettigrew, S.M. Miller, A sticky interhemispheric switch in bipolar
disorder? Proc. R. Soc. Lond. B Biol. Sci. 265 (1998) 21412148.
[78] A. Polonsky, R. Blake, J. Braun, D.J. Heeger, Neuronal activity in human
primary visual cortex correlates with perception during binocular rivalry,
Nat. Neurosci. 3 (2000) 11531159.
[79] M.T. Posamentier, H. Abdi, Processing faces and facial expressions, Neuropsychol. Rev. 13 (2003) 113143.
[80] M.I. Posner, M.K. Rothbart, Attention, self-regulation and consciousness,
Philos. Trans. R. Soc. Lond. B Biol. Sci. 353 (1998) 19151927.
[81] Practice parameters for determining brain death in adults (Summary,
Statement), The quality standards subcommittee of the American
academy of neurology, Neurology 45 (1995) 10121014.
[82] R.M. Pritchard, Visual illusions viewed as stabilized retinal images,
Quart. J. Exp. Psychol. 10 (1958) 7781.
[83] D. Reisberg, General mental resources and perceptual judgments, J. Exp.
Psychol. Human Percept. Perform. 9 (1983) 966979.
[84] D. Reisberg, M. OShaughnessy, Diverting subjects concentration slows
figural reversal, Perception 13 (1984) 461468.
[85] J. Ross, A. Ma-Wyatt, Saccades actively maintain perceptual continuity,
Nat. Neurosci. 7 (2004) 6569.
[86] Y. Rossetti, G. Rode, Reducing spatial neglect by visual and other sensory
manipulations: noncognitive (physiological) routes to the rehabilitation
of a cognitive disorder, in: H.-O. Karnath, D. Milner, G. Vallar (Eds.),
The Cognitive and Neural Bases of Spatial Neglect, Oxford University
Press, Oxford, 2002, pp. 375396.
[87] M.A. Scotto, G.A. Oliva, M.T. Tuccio, Eye movements and reversal rates
of ambiguous patterns, Percept. Mot. Skills 70 (1990) 10591073.
[88] F. Tong, S. Engel, Interocular rivalry revealed in the human cortical blindspot representation, Nature 411 (2001) 195199.

[89] F. Tong, M. Meng, R. Blake, Neural bases of binocular rivalry, Trends

Cogn. Sci. 10 (2006) 502511.
[90] F. Tong, K. Nakayama, J.T. Vaughan, N. Kanwisher, Binocular rivalry
and visual awareness in human extrastriate cortex, Neuron 21 (1998)
[91] G. Vallar, C. Guariglia, M.L. Rusconi, Modulation of the neglect syndrome by sensory stimulation, in: P. Their, H.-O. Karnath (Eds.), Parietal
Lobe Contributions to Orientation in 3D Space, Springer-Verlag, Berlin,
1997, pp. 555578.
[92] L.C.J. van Dam, R. van Ee, The role of saccades in exerting voluntary control in perceptual and binocular rivalry, Vision Res. 46 (2006)
[93] R. van Ee, A.J. Noest, J.W. Brascamp, A.V. van den Berg, Attentional
control over either of the two competing percepts of ambiguous stimuli
revealed by a two-parameter analysis: means do not make the difference,
Vision Res. 46 (2006) 31293141.
[94] R. van Ee, L.C.J. van Dam, G.J. Brouwer, Voluntary control and the
dynamics of perceptual bi-stability, Vision Res. 45 (2005) 4155.
[95] E. Vitte, C. Derosier, Y. Caritu, A. Berthoz, D. Hasboun, S. Soulie,
Activation of the hippocampal formation by vestibular stimulation: a
functional magnetic resonance imaging study, Exp. Brain Res. 112 (1996)
[96] R. Wales, R. Fox, Increment detection thresholds during binocular rivalry
suppression, Percept. Psychophys. 8 (1970) 9094.
[97] B. Wallace, F.A. Priebe, Hypnotic susceptibility, interference and alternation frequency to the Necker cube illusion, J. Gen. Psychol. 112 (1985)
[98] M.F. Washburn, A. Gillette, Motor factors in voluntary control of cube
perspective fluctuations and retinal rivalry fluctuations, Am. J. Psychol.
45 (1933) 315319.
[99] R. Wenzel, P. Bartenstein, M. Dieterich, A. Danek, A. Weindl, S.
Minoshima, S. Ziegler, M. Schwaiger, Th. Brandt, Deactivation of human
visual cortex during involuntary ocular oscillations. A PET activation
study, Brain 119 (1996) 101110.
[100] C. Wheatstone, The Bakerian Lecture.Contributions to the physiology
of vision.Part the second. On some remarkable, and hitherto unobserved, phenomena of binocular vision (continued), Philos. Trans. R. Soc.
Lond 142 (1852) 117.
[101] M.A. Williams, A.P. Morris, F. McGlone, D.F. Abbott, J.B. Mattingley, Amygdala responses to fearful and happy facial expressions
under conditions of binocular suppression, J. Neurosci. 24 (2004)
[102] M. Wilke, N.K. Logothetis, D.A. Leopold, Local field potential reflects
perceptual suppression in monkey visual cortex, Proc. Natl. Acad. Sci.
U.S.A. 103 (2006) 1750717512.
[103] E. Wong, N. Weisstein, A new perceptual context-superiority effect: line
segments are more visible against a figure than against a ground, Science
218 (1982) 587589.
[104] K. Wunderlich, K.A. Schneider, S. Kastner, Neural correlates of binocular
rivalry in the human lateral geniculate nucleus, Nat. Neurosci. 8 (2005)