Ambiguous Figures - What Happens in The Brain When Perception Changes But Not The Stimulus

REVIEW ARTICLE
HUMAN NEUROSCIENCE published: 22 March 2012

doi: 10.3389/fnhum.2012.00051
Ambiguous figures – what happens in the brain when

perception changes but not the stimulus
Jürgen Kornmeier 1,2 * and Michael Bach 2
1
Institute for Frontier Areas of Psychology and Mental Health, Freiburg, Germany
2
University Eye-Hospital, Freiburg, Germany
Edited by: During observation of ambiguous figures our perception reverses spontaneously although
Theofanis Panagiotaropoulos, Max
the visual information stays unchanged. Research on this phenomenon so far suffered
Planck Institute for Biological
Cybernetics, Germany from the difficulty to determine the instant of the endogenous reversals with sufficient
Reviewed by: temporal precision. A novel experimental paradigm with discontinuous stimulus presenta-
Theofanis Panagiotaropoulos, Max tion improved on previous temporal estimates of the reversal event by a factor of three.
Planck Institute for Biological It revealed that disambiguation of ambiguous visual information takes roughly 50 ms or
Cybernetics, Germany
two loops of recurrent neural activity. Further, the decision about the perceptual outcome
Kevin Whittingstall, Université de
Sherbrooke, Canada has taken place at least 340 ms before the observer is able to indicate the consciously
*Correspondence: perceived reversal manually. We provide a short review about physiological studies on mul-
Jürgen Kornmeier , Institute for tistable perception with a focus on electrophysiological data. We further present a new
Frontier Areas of Psychology and perspective on multistable perception that can easily integrate previous apparently con-
Mental Health, Wilhelmstraße 3a,
tradicting explanatory approaches. Finally we propose possible extensions toward other
79098 Freiburg i. Br., Germany.
e-mail: [email protected] research fields where ambiguous figure perception may be useful as an investigative tool.
Keywords: ambiguous figures, multistable perception, Necker cube, old/young woman, EEG/ERP, event-related
potentials, reversal positivity, reversal negativity
INTRODUCTION Koch, 1998; Blake and Logothetis, 2002; Dehaene and Changeux,
Normally we experience our visual world as stable and unambigu- 2011).
ous – it seems to be as we see it. Numerous optical illusions (Bach, We will here review the findings from physiological studies on
1997), however, demonstrate that the information provided via ambiguous figures with a specific focus on EEG studies. Based on
our eyes is restricted, thus incomplete and often ambiguous. Our the results available so far we will propose a new perspective on the
perceptual system needs to disambiguate and interpret it in order phenomenon, which easily integrates approaches that appeared to
to construct stable unambiguous percepts that allow us to success- be contradictory so far.
fully act in our environment. Extreme cases are ambiguous figures, Scientific studies of perceptual instability phenomena have
like the classical Necker cube (Figure 1A, Necker, 1832) or Borings been carried out for nearly 200 years and explanations so far fol-
Old/Young woman (Figure 1C, Boring, 1930), designed to render lowed two main traditions, namely the bottom-up (or sensory)
two (or even more) perceptual interpretations about equally prob- and the top-down (or cognitive) explanatory approaches.
able (indicated in Figures 1B,D). Another example is binocular The bottom-up approach assumes that perceptual reversals
rivalry, where the observer’s two eyes see different images (Blake, result from cycles of passive adaptation, recovery, and mutual inhi-
1989, 2001). In all of these cases the brain states corresponding bition of competing neural units or channels in early visual areas
to the two interpretations become unstable and spontaneous per- (e.g., Köhler, 1940; Toppino and Long, 1987). There is plenty of
ceptual reversals can occur although the external stimulus stays experimental evidence for this approach and some key findings
unchanged. are listed below:
Ambiguous figures can be found in any textbook about cogni- Several studies demonstrated local (retinotopic) adaptation
tive sciences and neuroscience. The perspective to experimentally effects (e.g., Howard and Durham, 1961; Long et al., 1992; Long
separate perceptual interpretation – which changes periodically – and Olszweski, 1999), effects of stimulus features (e.g., Washburn
from the earliest steps of visual processing – which should stay et al., 1931; Ammons and Ammons, 1963), and of presentation
unchanged, like the visual input – has attracted scientists from mode (e.g., Orbach et al., 1963; Kornmeier et al., 2007) on the
various disciplines in the field. It is believed that understand- initial percept and the reversal dynamics of ambiguous figures.
ing the mechanisms underlying the perceptual reversal or finding The top-down approach, in apparent contrast, assumes per-
the location(s) of the switch between sensory and perceptual ceptual reversals as the result from active high-level/cognitive
processing may help to understand how the activity pattern of processes like attention, expectation, decision-making, and learn-
sensory receptors is translated into a complex representation of ing (Gregory, 1974; Rock et al., 1994b; Leopold and Logo-
the perceptual world (perceptual organization, e.g., Pomerantz thetis, 1999). Some key findings supporting this approach are:
and Kubovy, 1981), how this representation is realized neurally (1) Although the bottom-up approach implies regular oscilla-
(object representation) and how it gets conscious (e.g., Crick and tions between the two interpretations, the durations of successive
Frontiers in Human Neuroscience www.frontiersin.org March 2012 | Volume 6 | Article 51 | 1

Kornmeier and Bach Ambiguous figures
reversals as time reference to analyze the data in a certain interval

backward in time. Typical control intervals in manual response
studies were periods of stable percept (e.g., Basar-Eroglu et al.,
1996) and spontaneous EEG intervals during fixation of a fixa-
tion target (e.g., Strüber et al., 2001). Spontaneous key presses not
related to perceptual reversals were considered as control for EEG
components related to motor preparation and execution (Basar-
Eroglu et al., 1993). This paradigm has been applied in a number
of studies and the results are summarized in the following:
P300-like parietal positivity

A positive event-related potential (ERP) with a right parietal maxi-
mum (P4 electrode,“parietal positivity”) occurred ≈250 ms before
key press with reversals of motion direction of the stroboscopic
alternative motion (SAM) stimulus (Schiller, 1933; Basar-Eroglu
et al., 1993; Strüber and Herrmann, 2002) and with orientation
FIGURE 1 | Examples of ambiguous figures and their disambiguated reversals of the Necker cube (Strüber et al., 2001; Mathes et al.,
variants. (A) The Necker cube (Necker, 1832) and (B) disambiguated 2006). This parietal positivity was interpreted as a variant of the
versions with depth cues and a lightning model. (C) Outline version of well-known cognitive P300 ERP component (e.g., Pritchard, 1981;
Borings Old/Young woman figure (Boring, 1930) adapted from Gale and
Verleger et al., 2005) and thus taken as evidence for top-down
Findlay (1983) and (D) disambiguated old (left) and young woman figure
(right). processes. Strüber and Herrmann (2002) compared MEG activity
during endogenous motion reversals of the SAM with exogenously
induced motion reversals of disambiguated SAM variants (the two
intervals of transiently stable percepts (“dwell times”) are unpre- dots moved concurrently either horizontally or vertically in the
dictable and show characteristics of a stochastic process (e.g., same direction). They found a reversal-related parietal positivity
fit to gamma distribution, Borsellino et al., 1972; Murata et al., both for endogenous as well as for exogenously induced motion
2003; Brascamp et al., 2005). According to Leopold and Logothetis reversals. Given reaction times from their unambiguous stimulus
(1999) this is a signature of high-level exploratory behavior. (2) conditions of about 550 ms, the authors assumed that their parietal
Perceptual reversals are very rare or even absent, when observers do positivity occurs subsequent to the motion reversal and reflects its
not know that alternative interpretational possibilities exist (e.g., conscious recognition.
Girgus et al., 1977; Rock et al., 1994a), a finding hard to recon-
cile with fully automatic adaptation processes. Further, the rate of Alpha-band power decrease
reversals can be modulated volitionally, although they cannot be İşoğlu-Alkaç (2000) used the Necker cube in their study and
prevented totally (e.g., Liebert and Burk, 1985; Strüber and Stadler, reported a decrease of alpha-band power in the time range of
1999). the parietal positivity compared to alpha activity during a time
While most of the authors so far exclusively favored either range immediately before. The authors argue that because alpha-
a bottom-up or a top-down explanation, more and more stud- power decrease occurs with active brain states while alpha-power
ies indicate that both classes of factors play important roles and increase with passive states of decreased cortical excitability, alpha
must be integrated in an explanation of the phenomenon (e.g., power (8–16 Hz) decrease together with the P300-like positiv-
Hochberg and Peterson, 1987; Blake and Logothetis, 2002; Tong ity indicate active cognitive processes during perceptual reversals
et al., 2006; Kornmeier et al., 2009; Bartels and Logothetis, 2010). (İşoğlu-Alkaç et al., 2000). Strüber and Herrmann (2002) found
a relatively constant value of alpha oscillation (10 Hz) until it
EEG STUDIES AND THE TIME REFERENCE PROBLEM suddenly drops between 300 and 200 ms before button press
The EEG allows millisecond resolution of neural processes under- for exogenously induced SAM-motion reversal, and a continu-
lying perceptual reversals and thus may provide the necessary ous decrease in alpha activity within 1000 ms before key press for
temporal precision to resolve the debate about whether low-level endogenous reversals. They interpreted the latter as evidence for a
bottom-up or high-level/cognitive processes are at work. However slow bottom-up destabilization of the active perceptual interpre-
a precise time reference is necessary for EEG analysis but difficult to tation compared to a sudden exogenously driven destabilization.
establish because of the endogenous character of the reversal event. İşoğlu-Alkaç and Strüber (2006) recently found that alpha-power
Two major strategies have evolved to circumvent this problem, decrease is restricted to the lower alpha bands (6–10 Hz) whereas
both with advantages and disadvantages. In the following we will the upper alpha band (10–12 Hz) was unaffected.
discuss the relevant studies and relate their results to each other.
Gamma-band power increase
THE “MANUAL RESPONSE PARADIGM” – USING MANUAL RESPONSE Basar-Eroglu et al. (1996) found (non-phase-locked) EEG gamma
AS TIME REFERENCE band (30–50 Hz) enhancement within 1000 ms before key press
The principal idea is to present an ambiguous stimulus contin- related to SAM reversals compared to perceptual stability (dur-
uously and use participants’ manual indications of perceptual ing a time interval after key press) most prominent at the right

frontal electrode position (F4). The results were interpreted as

evidence for cognitive destabilization processes underlying per-
ceptual reversals. Strüber et al. (2000) found higher SAM reversal-
related gamma activity (28–48 Hz) for participants with high
reversal rates (reversals per time unit) compared to participants
with low reversal rates. The authors interpret their results as
further support for the involvement of attentional top-down pro-
cessing during endogenous motion reversals. Strüber et al. (2001)
confirmed these results for the Necker cube.
In Mathes et al.’s (2006) study participants had to volitionally
either speed up or slow down the reversal rate or keep a passive atti-
tude toward a Necker cube stimulus. The authors reported higher
amplitude of the P300-like positivity and higher gamma activity
(28–48 Hz) during the slow down condition than during the speed
up condition.
Summary
Taking the above studies together, gamma power increases and
alpha-power decreases in a time interval 1000 ms before partici-
pants’ manual indication of perceptual reversals of an ambiguous
figure. Between 500 and 250 ms before key press a parietal positiv-
ity occurs which is interpreted as indicating conscious recognition
of a perceptual reversal. The reversal event itself then must have
occurred earlier.
PROBLEMS OF THE MANUAL RESPONSE PARADIGM

EEG results from the manual response paradigm suggest that
endogenous perceptual reversals take place between 1000 and
550 ms before observers’ manual response. However, the available
data can easily be interpreted either in bottom-up or top-down
direction.
Further, as the authors emphasize, manual reaction times
strongly vary both intra-individually and inter-individually
(Strüber et al., 2000; Strüber and Herrmann, 2002). Recently we
estimated the amount of reaction time variance related to orienta-
tion reversals of a Necker cube (Kornmeier and Bach, 2004b) in the
following way: using depth cues, we created two unambiguous and FIGURE 2 | Estimating manual reaction time variance. Participants
thus perceptually stable variants of the Necker cube (Figure 1B). observed unambiguous cubes (similar to Figure 1B) that randomly reversed
(16 times per minute on the average) and indicated manually each
These appeared in random order, and participants indicated per-
perceived orientation reversal. (A) ERP traces (±SEM) from the occipital
ceived orientation reversals with a key press. Median reaction time location of one participant with the exogenous reversal instance as time
from five subjects was 616 ms with an interquartile range from 530 reference for averaging (vertical line) are seen. Sharply defined early ERP
to 733 ms. ERP example traces (±SEM) from the occipital location components result; the reaction time is widely scattered (200 ms
(Oz) of one participant are seen in Figure 2A: When the EEG is interquartile range; dashed line indicates median reaction time). (B) The
same EEG data averaged backward with reaction times as time references
averaged with respect to the exogenous stimulus reversal (continu-
(vertical line). The early visual ERP components are largely obliterated. Only
ous vertical line), sharply defined ERP components result (dashed the late positivity is roughly similar to (A). The Onset Paradigm (Figure 4)
vertical line indicates the median reaction time). Figure 2B: When avoided this problem.
the same EEG is “backward averaged” with respect to the par-
ticipant’s response, the early visual ERP components are largely
obliterated. It is thus likely, and indeed we will show that endoge- in a non-monotonic manner mainly as a function of ISI. Figure 3
nous reversals are accompanied by ERP structures earlier than the depicts their data together with data from our own lab (Kornmeier
parietal positivity that can be indentified only with a more precise et al., 2002).
time reference. As can be seen, for ISIs shorter than about 400 ms reversal
rates increase monotonically up to more than twice the rate dur-
THE ONSET PARADIGM: STIMULUS ONSET AS TIME REFERENCE ing continuous observation. After 400 ms a monotonic decrease
The onset paradigm goes back to the work of the Orbach group of reversal rates can be observed. Reversals can even be brought
(e.g., Orbach et al., 1963; Orbach et al., 1966). They presented a to stand still with ISIs in the range of seconds (Leopold et al.,
Necker cube discontinuously, varied presentation time and inter- 2002; Maier et al., 2003). Most relevant for EEG studies is that
stimulus interval (ISI) and found that reversal rates are modulated reversals occurred typically near stimulus onset if the presentation

interruptions, Figure 3, left part), and another responsible for the

decreasing part (perceptual reversals with long ISIs, Figure 3 right
part). O’Donnell et al.’s choice of 3300 ms ISI places their reversals
to the second type and thus raises doubts about whether they are
a good model for the continuous case (see also below).
(3) A perceptual reversal during continuous observation of an
ambiguous figure involves an implicit comparison of the current
with the previous (different) percept, thus working memory may
play a role (e.g., Pearson and Brascamp, 2008). In O’Donnell et
al.’s (1988) study participants were instructed to report their per-
cept of each presentation. Comparisons across presentations were
not necessary. This task together with the relatively long ISIs may
have allowed separate percepts of cube stimuli at each stimulus
presentation instead of perceptual reversal experiences.
We adopted O’Donnell et al. (1988)’s paradigm but modified it
FIGURE 3 | Reversal rate as a function of ISI. Reversal rates with in the following ways (Figure 4):
continuous and discontinuous presentations of ambiguous Necker stimuli
from different studies. Reversal rates behave non-monotonically, increasing
with ISI up to 400 ms. Thereafter they decrease with further increasing ISIs.
(a) Optimizing presentation time and ISI: We shortened our ISI
(Fig from Kornmeier et al., 2007, modified). to 400 ms to be as close to the continuous viewing condition as
possible while concurrently enabling manual responses within
the ISI. Presentation time was set to 800 ms, long enough
time was short enough (Orbach et al., 1966). Stimulus onset thus to allow the full development of a P300-like positivity and
promised to be a more precise time reference of the reversal event short enough to keep the probability of additional perceptual
than reaction times. reversals later during the presentation interval low.
O’Donnell et al. (1988) were the first study to use stimulus (b) Comparison task across presentations: Our participants com-
onset as time reference for averaging EEG data. In two condi- pared in a go/nogo task the perceived front-back orienta-
tions participants viewed the ambiguous Necker cube and (appar- tion of the current Necker lattice (a combination of nine
ently) unambiguous cube variants for 700 ms followed by ISIs of Necker cubes, Figure 4) with that of the preceding one in
3300 ms. After each stimulus offset they indicated their previous two experimental conditions. In one condition they pressed
percept by pressing one of two possible keys. In the case of the a key whenever the currently perceived orientation differed
ambiguous Necker cube, perceptual reversals between two suc- from the preceding one (“reversal condition”). The second,
cessive stimulus presentations were purely endogenous, whereas control condition was identical except that the task was the
in the case of the unambiguous cube variants perceptual rever- opposite: participants indicated when they perceived orien-
sals were regarded as exogenously induced by stimulus alterna- tation of the stimulus as remaining the same (“stability con-
tions. The authors compared reversal trials with non-reversal dition”). All manual responses were executed in the ISI after
trials and found a late parietal/frontal positivity for both exoge- the relevant percept in order to keep neural activity related
nously induced and endogenous perceptual reversals. In the case to motor-preparation and -execution as far away as possible
of exogenously induced reversals this positivity was larger and from the critical time interval of the reversal. Any ERP sig-
occurred earlier (around 550 ms after stimulus onset) compared nature related to lower-level stimulus processing, afterimages,
to endogenous reversals (around 650 ms after onset). or motor preparation should be identical in the two condi-
Identifying O’Donnell et al.’s (1988) positivity with the positions and should disappear after subtraction of the related
tivity found with the Manual Response Paradigm and accepting ERP traces from each other.
Strüber and Herrmann’s (2002) hypothesis about the positivity (c) Like O’Donnell et al. (1988) we exogenously induced orienta-
as an index of conscious realization of the reversal, O’Donnell et tion reversals of unambiguous stimulus variants in a second
al.’s data restrict the reversal event to a time window of 650 ms experiment. Our unambiguous cube variants contained depth
between stimulus onset and the late positivity. Further, the similar cues, like shading, central perspective, and a lighting model
results of O’Donnell et al.’s (1988) illusory and physical reversal (Figure 1B, Woo et al., 1998).
conditions suggest similar processes underlying endogenous and
exogenously induced perceptual reversals. Four types of ERP traces were calculated, related to (1) endoge-
O’Donnell et al.’s (1988) data provide a sizable improvement nous perceptual reversals and (2) endogenous perceptual stability
on temporal precision, but some limitations have to be faced: (1) of ambiguous Necker stimuli, (3) exogenously induced percep-
Their “unambiguous” cube variants are not really unambiguous. tual reversals, and (4) exogenously induced perceptual stability of
They can easily be perceived in two different orientations as either disambiguated stimulus variants.
inside or outside corners of a box. (2) The non-monotonic charac-
ter of the reversal rate as a function of ISI (Figure 3) suggests two THE PROCESSING CHAIN OF PERCEPTUAL REVERSALS
separate processes, one responsible for the increasing part of the According to our null hypothesis, the EEG traces from the reversal
function (reversals during continuous observation and with short and stability conditions should not differ, because both conditions

FIGURE 4 | Onset paradigm. Participants viewed in different experiments perspective reversal [reversal conditions in (A,C)] or perceived stability
either unambiguous lattices (A,B) or ambiguous Necker lattices (C,D) and [stability conditions in (B,D)] across two successive stimulus presentations by
compared the 3D perspective of successively presented stimuli. In separate a key press in the ISI following the respective perceptual event. Each key
experimental conditions they indicated in a go/nogo task either a perceived press extended the current ISI from 400 to 1000 ms.
contain identical stimuli, a comparison task, a mental decision, electrode positions are indicated schematically in Figures 5B and
and aspects of response preparation. Further, the amount of 9B. At a first glance ERP traces (Figure 5A) are very similar across
required attention to execute the task should be equal in the two stimuli and conditions. Especially the P100 components as the
conditions. earliest visual responses after stimulus onset are roughly identi-
The difference traces (“dERP,” reversal condition minus sta- cal (Figure 6). The subsequent components match in latency but
bility condition) should thus be flat and any residuum would be differ in amplitude. These differences are better seen in the dif-
related to the perceptual reversal aspect. Indeed we did find a ference traces (dERPs, Figure 5C) and their temporal relations
series of reversal related modulations in lower (dERP) as well as are schematically presented in Figure 8. Likewise modulations at
higher frequencies. Figure 5A presents the ERP traces from per- higher frequency are very similar across averaged conditions and
ceptual reversal trials (interrupted lines) and perceptual stability experiments, and are depicted in the time–frequency charts in
trials (continuous lines) on a schematic head (data from Korn- Figure 9A. Most conspicuous is an initial power enhancement in
meier and Bach, 2004b). Figure 9 presents the related (induced) the alpha to beta range immediately after onset that quickly shifts
modulations at higher frequencies. Like in Figure 5, for each to the theta and lower alpha range where it sustains. Further, a sus-
electrode position a pair of graphs depicts data from the unam- tained beta deactivation starts about 100 ms after stimulus onset
biguous lattice variants on bottom on a gray background and data and spreads to the whole beta and upper alpha band. Differences in
from the ambiguous lattices on top on a white background. The higher frequency power between conditions and experiments are

FIGURE 5 | Grand mean ERPs (16 participants, baseline −60 to traces) and across experiments (ambiguous and unambiguous stimuli).
+40 ms). For each electrode position a pair of graphs contain data from This is especially true for the early visual ERP components at the occipital
the unambiguous lattice variants in red on bottom on a gray background electrode position). (C) Difference traces (reversal minus stability).
and the data from the ambiguous lattices in blue on top on a white Differences ERPs (dERPs) are very similar between ambiguous and
background. Dashed vertical lines indicate stimulus onset, electrode unambiguous stimuli with two exceptions: (1) All dERPs found with
positions are indicated schematically in (B). (A) Raw ERPs before unambiguous stimuli occur earlier and (2) the early occipital Reversal
subtraction. The global shapes of the ERP traces at each electrode are very Positivity (RP) is restricted to endogenous perceptual reversals of
similar across conditions (reversal, dotted traces, and stability, continuous ambiguous Necker lattices. Data from Kornmeier and Bach (2004b).
better seen in the difference time–frequency charts in Figure 9C In the following we will discuss the individual dERP compo-
and their temporal relation schematically in Figure 10. nents and related modulations at higher frequencies separately:

ratio and thus the number of EEG trials per subject that enter the
ERP calculation. This was between 100 and 120 per condition in
our studies. Britz et al. (2009) reported about 52 trials per condi-
tion and participant on average. In Pitts et al.’s (2007, 2008) studies
the number of trials was down to 30 and below. Their absolute
number of trials (not reported) may have been large enough to let
a positive deflection appear but too low to render it significant1
(Luck, 2005; Pitts et al., 2007).
In studies using the Manual Response Paradigm the RP was
probably obliterated due to reaction time variability. This would
imply that reaction times are considerably less precise compared to
stimulus onset as time reference. We estimated the precision of the
Onset Paradigm in the following way: The earliest ERP response
after stimulus onset, the P100, occurred in our data 80 ms after
onset with a peak width of ±20 ms and was regarded as a low-level
ERP and fully synchronized to stimulus onset. In comparison, the
RP is the earliest reversal-related dERP component has a peak
width of about ±35 ms. Let us now assume that it results from a
convolution of the variance producing the P100-width with the
variance producing the Reversal Positivity. Assuming a Gaussian
peak shape, the unknown width of the temporal variance would
FIGURE 6 | Grand mean ERP traces from the occipital electrode
be equal to:
position. As can be seen, the early visual ERP components are very similar
across conditions (reversal, dotted traces and stability, continuous traces)
and experiments (Necker lattices, blue, and unambiguous variants, red). Temporal variance of the reversal instance = 352 — 202 ≈ 30
Data from Kornmeier and Bach (2004b).
(based on additivity of variances)

Thus endogenous perceptual reversals were synchronized with
SIGNATURES SPECIFIC FOR ENDOGENOUS REVERSALS OF stimulus onset with a precision of about ±30 ms, an improvement
AMBIGUOUS FIGURES of more than a factor of three over the manual response reference
Reversal positivity (Figure 7).
The reversal positivity (RP) occurs around 130 ms after stimulus
onset, is most prominent at occipital electrode positions and Early alpha modulations
restricted to endogenous reversals of the ambiguous stimuli. So A left-hemispheric power reduction in the alpha band (8–15 Hz)
far it has been reported by studies using the Onset Paradigm ranging from the occipital to frontopolar electrode (Component
with Necker lattices (Kornmeier and Bach, 2005, 2006; Korn- A5 in Figures 9 and 10) shows some similarities to the RP and may
meier et al., 2007; Britz et al., 2009), with the classical Necker cube thus be related: Both (1) start around 130 ms, (2) are restricted
(Kornmeier et al., 2011), with Borings (1930) Old/Young Woman to endogenous reversals, and (3) have an opposite sign as the
(Figure 1C, Kornmeier and Bach, 2004a) and recently even with subsequent components.
binocular rivalry stimuli (Britz and Pitts, 2011). Pitts et al. (2007)
found modulations in the P1 ERP component during reversals of Interpretation of the RP: decision conflict during interpretation
the Vase/Face stimulus and Schroeder’s staircase stimulus fitting of ambiguous information. In summary, the RP shows three
with polarity and time window of the RP. Britz and Pitts recently interesting features:
discussed the RP as a modulation of the P1 component and local-
ized it in “primary visual areas” (Britz and Pitts, 2011). Recent (1) It is restricted to endogenous reversals.
evidence from our lab suggests, however, that P1 and RP are dif- (2) It occurs relatively “early” during visual processing, which is
ferent components (Kornmeier et al., 2011) suggesting that the reflected in its latency and its spatial distribution at occipi-
P1 effect in Pitts et al. (2007) may be a superimposed RP. Pitts tal electrodes. For discontinuous stimulus presentation, early
et al. (2008) and Intaite et al. (2010) did not find a significant RP is well defined as occurring with short latency after stimulus
with perceived reversals of the Necker lattice. In the latter study onset. But what would early mean for perceptual reversals
a strongly modified variant of the Onset Paradigm (e.g., reversals during continuous observation? Here early and late can be
from unambiguous to ambiguous lattices) may be the reason. Pitts alternatively defined as temporal distance from reaction time.
et al. (2009) did not analyze the data in the RP time window. Reaction times can be estimated with ambiguous figures using
A possible explanation for the remaining inhomogeneity of the Onset Paradigm and instructing participants to respond
results may be the following: The RP is a small component typi-
cally with amplitudes around or even below 1 μV (e.g., Kornmeier 1 The multiple testing problem is inherent to the analysis of EEG data. Depending
and Bach, 2005; Kornmeier and Bach, 2006; Britz et al., 2009). A on the number of tests, corrections for alpha-error-inflation can strongly reduce
critical parameter for its statistical validity is the signal-to-noise test-sensitivity, especially in cases of small signals like with the Reversal Positivity.

even with binocular rivalry stimuli). This indicates gener-

ality across low-level stimulus features and across stimulus
types. Further, the RP differs from the spatiotemporal close
P1 (Kornmeier et al., 2011).
Currently we regard the RP as an indicator of the visual sys-

tem’s “detection” of ambiguity, or the related decision conflict at
a certain processing level during perceptual organization (Pomer-
antz and Kubovy, 1981) where the ambiguity gets evident. For the
Necker cube the identification of line orientations and their bind-
ing together would be low-level steps and perhaps related to the
C1 and P1 ERP components (discussed in Kornmeier et al., 2011).
Ambiguity occurs subsequently during the interpretation of acute
and obtuse angles as orthogonal and the allocation of depth values
to the different square planes. Here a perceptual decision has to be
made and the RP may reflect the related conflict or its detection.
Two questions arise immediately from this speculation:
(1) In the case of the Necker cube the stimulus information is

always ambiguous. Why should there be a difference in deal-
ing with this ambiguity in reversal and stability trials (remind
that the RP is a dERP, resulting from the subtraction of stabil-
FIGURE 7 | Estimation of the temporal precision with the Onset ity ERP traces from reversal ERP traces)? A possible answer to
Paradigm. Continuous trace: Grand mean ERP trace from the occipital this question may be the following: All sensory information
electrode in the reversal condition in the P100 time window. Dotted trace:
Grand Mean dERP trace from the occipital electrode position in the time
is incomplete and to some degree ambiguous. Our perceptual
window of the RP. The dERP trace is shifted on the time scale in order to system uses (amongst other strategies) past perceptual expe-
align the RP with the P100 and thus to simplify peak-with comparison. Data riences (on various time scales) to disambiguate and interpret
from Kornmeier and Bach (2005). this information. Several adaptation and priming studies sup-
port this view (e.g., Long et al., 1992; Woerner et al., 2009).
The above mentioned decision conflict need not occur if the
immediately after perceived reversals. Substituting the rever- perceptual system is already strongly biased in favor of one
sal instance with stimulus onset and assuming that reversals (e.g., the previous) percept at the moment the ambiguous
occur only after stimulus onset, the upper limit of reaction stimulus appears. This may be the case in our stability tri-
time to endogenous reversals is in the range of 600 ms (Korn- als. We have estimated the disambiguation time in the case
meier and Bach, 2006). Thus the RP occurs roughly 470 ms of a decision conflict with the Necker lattice as 40–70 ms
before the manual indication of a perceived reversal (Korn- (Kornmeier and Bach, 2006, see also below). If our above
meier and Bach, 2006). In perceptual processing times 470 ms speculation is correct, then reaction times should be delayed
is very long and thus the RP can also be labelled as early from by this amount in the reversal trials, where a conflict arises
this point of view. and disambiguation is necessary, compared to the stability
(3) The RP occurs with different types of ambiguous stimuli trials without conflict. This is indeed what we recently found
(Necker cube, Necker lattice, Old/Young woman stimulus and (Kornmeier et al., 2011).
FIGURE 8 | Schematic time scale of endogenous and exogenous component before subtraction. Negative values on the time axis reflect
dERP components. dERP components are indicated by rectangles. The the time period of the ISI before stimulus onset. Data from Kornmeier
vertical dashed line indicates the P100 as earliest occipital ERP and Bach (2004b, 2006).

FIGURE 9 | Grand mean EEG time–frequency charts. For each charts (reversal minus stability). Entries in (B) (e.g., A1, U2, . . .) indicate
electrode position a pair of time–frequency charts contain data from the the position of significant deflections from zero in the time–frequency
unambiguous lattice variants (bottom, grey background) and the data from plane. A, ambiguous; U, unambiguous; numbers indicate the temporal
the ambiguous lattices (above, white background). Dashed vertical lines order of effects; black/white colouring of the numbers and letters is only
mark the stimulus onset. (A) Raw time–frequency charts. Grand mean for saliency. Most remarkable is the gamma band modulation before onset
time–frequency charts were obtained by averaging data across of the ambiguous Necker stimulus (deflection A1 at electrodes P4 and C4
participants and conditions. Electrode positions are indicated and Cz; and deflection A2 at electrode Oz). No such modulation occurs
schematically in (B). (C) Differences of grand mean EEG time–frequency with the unambiguous stimulus variants. (from Ehm et al., 2011, Figure 2).
(2) Disambiguation of incoming information as described above accumulating evidence that perception is discontinuous and
makes sense if the stimulus reappears periodically after a that our impression of perceptual continuity is an illusion
blank screen interval and a percept has to be created. But (e.g., Dubois and Vanrullen, 2011). According to this view
how can this be translated to the continuous case? There is some neural instances would “perceive” a discontinuously

FIGURE 10 | Schematic display of significant components from the (negative) excursions appear above (below) the time axes. Top: Necker
time–frequency analyses. The black dots within the schematic heads mark stimulus; bottom: unambiguous stimulus. Gray backgrounds highlight
electrode positions. The closed polygons surrounding electrode positions analogies between ambiguous and unambiguous stimuli. Their temporal
mark significant test results, their color indicates the corresponding frequency difference is consistent with dERP results. Pre-onset gamma modulations (A1
range. Head positions on the time axes indicate peak time. Positive and and A2) are restricted to the Necker stimuli.
presented stimulus similarly to a continuously presented alpha activity is discussed in the contexts of change from a resting
stimulus (given a certain frequency). Some (potentially higher state to excitation/activation, attention and top-down modulation
level) updating instance may periodically re-evaluate and re- of perceptual processing, execution of cognitive tasks or temporal
interpret the visual input in order to notice environmental segmentation of perception and consciousness (for a review, see
changes. Thus, periodic decisions and decision conflicts may Palva and Palva, 2007). In the current experiment the alpha reduc-
also arise during continuous observation of an ambiguous tion may indicate recurrent activity between occipital and frontal
figure. Such a concept had already been proposed in top- areas to resolve the ambiguity and the related decision conflict
down explanation approaches (e.g., Vickers, 1972; Leopold indicated by the RP. It may thus reflect a disambiguation time of
and Logothetis, 1999) and also in a recent Necker-Zeno Model roughly 60 ms in the case of the Necker lattice. Interestingly, Foxe
for Bistable Perception (Atmanspacher et al., 2004, 2008). and Simpson (2002) indicated that in humans visual information
Accordingly, the RP should not only occur with the Onset Par- needs only about 30 ms from striate to prefrontal cortex. Two loops
adigm but also with spontaneous reversals of a continuously of recurrent activity thus seem to be sufficient to disambiguate the
presentated ambiguous figure. And it should be detectable Necker lattice.
if a sufficiently precise time reference were available with
continuous stimulus presentation. Additional experiments SIGNATURES OCCURRING WITH BOTH EXOGENOUSLY INDUCED AND
with ambiguous figures from other categories (e.g., appar- ENDOGENOUS REVERSALS
ent motion, figure-background stimuli) need to further test Reversal negativity
the generality of the RP. The reversal negativity (RN) is the first dERP component found
with both endogenous reversals of the Necker lattices and exoge-
Interpretation of the alpha-power reduction: disambiguation nously induced reversals of the unambiguous lattice variants.
time. The left-hemispheric reduction of alpha-power starts at the It occurs roughly 220 ms after stimulus onset with exogenously
latency of the RP (130 ms), it extends from posterior to anterior induced reversals and about 40 ms later at 260 ms with endogenous
sites and lasts for about 60 ms (Figures 9 and 10). Modulation of reversals (Kornmeier and Bach, 2006; Intaite et al., 2010; Figure 8).

This component has been reported in all studies using the onset studies did not discern between Frontopolar and Parietal Posi-
paradigm (Kornmeier and Bach, 2004b, 2005, 2006; Kornmeier tivities and regarded them as one and the same component with
et al., 2007, 2011; Pitts et al., 2007, 2008, 2009; Britz et al., 2009; a parietal to frontopolar extent (e.g., Britz et al., 2009; Britz and
Intaite et al., 2010; Britz and Pitts, 2011). Source analysis revealed Pitts, 2011). Pitts et al. (2008) reported a frontal positive compo-
locations in the lateral occipital and inferior temporal areas (Pitts nent at around 300 ms and interpreted it as a “selection positivity,”
et al., 2009; Britz and Pitts, 2011). i.e., a sign-inverted frontal counterpart of the occipital/parietal
There are, however, some differences concerning the temporal Selection Negativity (Anllo-Vento and Hillyard, 1996).
extent and the spatial distribution of the RN. These differences
may be traced back to differences in the experimental paradigm Interpretation of the frontopolar positivity. Recently we found in
concerning the time window of participants’ responses. In some our data strong evidence in support of two separate positive com-
studies the participants were allowed to respond immediately after ponents: With our go–nogo variant of the Onset Paradigm we were
a perceived reversal, i.e., already during the stimulus presentation able to differ between manually indicated reversal events (rever-
period. In those cases the negative dERP component was both tem- sal percepts in the reversal condition, i.e., go trials) and reversal
porally and spatially extended (e.g., Pitts et al., 2008; Intaite et al., events without indication (reversal percepts in the stability con-
2010; Kornmeier et al., 2011) compared to the case when responses ditions, i.e., nogo trials) – and with the same logic corresponding
were only allowed in the ISI following a perceived reversal. In the stability events. The dERPs from the go trials showed a decrease
latter case the negative dERP component was shorter and restricted of a positivity at 400 ms and a increase of a positivity at 300 ms
to occipital/parietal locations (e.g., Kornmeier and Bach, 2004b; from parietal to frontopolar electrode positions in the case of the
Kornmeier and Bach, 2005). Kornmeier et al. (2007) demonstrated unambiguous lattices (Figure 11C: from bright to dark colors).
that in the cases of immediate responses (already in the stimulus This is similarly indicated but less pronounced with the ambigu-
presentation interval) the occipital/parietal RN is superimposed ous lattices with the above mentioned latency shifts (Figure 11A).
by a residual Bereitschaftspotential, which occurs later (at about In contrast, the dERPs related to the nogo trials showed a sim-
300 ms after onset) and which has a distribution from parietal to ple amplitude reduction from parietal to frontopolar electrodes
frontal positions. Elbert et al. (1985) have previously shown influ- without any difference in latency (Figures 11B,D). Thus, the Fron-
ence of psychological variables on the Bereitschaftspotential in the topolar Positivity is absent in the nogo trials. This observation
context of perceptual reversals of the Necker cube. decouples the Frontopolar Positivity both from the Parietal Pos-
itivity and the RN and may indicate a role of working memory
Interpretation of the RN. So far the functional role of the RN related to the delayed response in the subsequent ISIs of the go tri-
is unclear. It’s role as an indicator of top-down influence of the als – no delayed response and thus no related memory is necessary
reversal process (e.g., Pitts et al., 2008) and its potential identity in the nogo trials. This interpretation suggests that some response-
with the spatiotemporally very similar Selection Negativity (Anllo- related brain instance must already know about the perceptual
Vento and Hillyard, 1996) are in discussion (Kornmeier and Bach, outcome at 300 ms (unambiguous lattices) or 340 ms (ambiguous
2004b; Pitts et al., 2008; Intaite et al., 2010). A specific role for the lattices). More research is necessary to test this interpretation of
RN in endogenous perceptual reversals can be ruled out by the the Frontopolar Positivity.
fact that it also occurs when perceptual reversals are exogenously
driven by a physical stimulus change (Kornmeier and Bach, 2006). Interpretation of the parietal positivity. So far all electrophysio-
Interestingly, the RN’s spatial location seems to be related to the logical studies on ambiguous figures report a slow positivity with
stimulus type. It is most prominent at occipital/parietal locations parietal to frontal dominance. With the Onset Paradigm such a
with cube type stimuli whereas it seems to be more dominant positivity occurs between 400 and 500 ms after stimulus onset and
at temporal locations in the case of Boring’s old/young woman between 100 and 150 ms before the manual response (e.g., Korn-
(Kornmeier and Bach, 2004a). meier and Bach, 2006; Pitts et al., 2009). With the Manual Response
Paradigm a slow positivity peaks roughly 250 ms before the man-
Frontopolar and parietal positivities ual response. Assuming that both paradigms identified the same
The Frontopolar Positivity follows the RN. It is most prominent at component, the mismatch between latencies of about 100 ms may
the frontopolar electrode and occurs 300 ms after stimulus onset result from (1) different time references for averaging (stimulus
in the case of exogenously induced reversals, and 40 ms later at onset or reaction instances) together with a temporal coupling
340 ms in the case of endogenous reversals (Figures 5C and 8). No asymmetry of this component to the two time references and (2) a
source analysis of this component has been done so far. merging of the Frontopolar with the Parietal Positivity in the case
The Parietal Positivity follows the Frontopolar Positivity. It is of backward averaging from manual responses.
maximal at parietal electrodes and occurs 400 ms after stimulus Several authors identified the Parietal Positivity with the well-
onset with exogenously induced reversals and about 70 ms later at known P3b component, which typically occurs in oddball par-
470 ms in the case of endogenous reversals. Most of the ERP stud- adigms (Picton, 1992), and indeed Verleger et al. (2005) demon-
ies using the Onset Paradigm reported a reversal-related parietal strated the persistence of the classical P3b across onset and reaction
positivity with very similar spatiotemporal patterns (e.g., Korn- time as time references for averaging. The interpretations of the
meier and Bach, 2004b; Pitts et al., 2008; Britz et al., 2009). It was Parietal Positivity range from (1) indication of attentional and
recently localized in the bilateral superior and middle temporal cognitive processes during a perceptual reversal (O’Donnell et al.,
as well as left inferior frontal areas (Britz and Pitts, 2011). Some 1988) or (2) following it (İşoğlu-Alkac et al., 2000), (3) closure

FIGURE 11 | Parietal and frontopolar positivities (dERPs) along the however, is absent in the no–go trials. Instead, the amplitude of the Parietal
midline electrodes. (A,C) Temporally and spatially separated Parietal and Positivity decreases from parietal to frontopolar positions, while its latency
Frontopolar Positivities occur in the go trials. (B,D) The Frontopolar Positivity, stays unchanged (Data from Kornmeier and Bach, 2004b).
of the switching process (Strüber et al., 2001), or (4) cogni- a temporal delay. A similar pattern of results can be observed
tive/conscious recognition or appraisal of the reversal (Strüber in the beta and gamma frequency range. In the case of exoge-
and Herrmann, 2002; Kornmeier and Bach, 2006; Britz and nously induced reversals a left-central increase in gamma activ-
Pitts, 2011). The following consideration supports the cognitive ity at about 150 ms (40–65 Hz, Figures 9 and 10, component
appraisal/visual awareness hypothesis from above: The minimal A6) followed by a bilateral central increase in beta activity (14–
reaction time with the simplest auditory task (faster than visual) is 26 Hz, Figures 9 and 10, component A7) at around 260 ms can
in the range of 150 ms (Debecker and Desmedt, 1970). This gives be observed. Correspondingly, in the case of endogenous rever-
an upper limit for a pure motor execution after the conscious sals a left-central gamma increase starts at about 250 ms and
response decision has taken place. It fits well with our average thus 100 ms later than the exogenous counterpart (Figures 9
temporal distance between the Parietal Positivity and the manual and 10, component U2). An increase in beta activity follows
reaction and thus may place the moment of conscious experi- at around 320 ms (Figures 9 and 10, component U3). Signif-
ence of the perceptual change (and the decision to indicate this) icance for the latter, however, is restricted to the right-central
roughly at the latency of the Parietal Positivity. Further support electrode position. Although the pattern of results agrees less well
comes from several ERP studies on ERP correlates of conscious- than the dERP components, which might partly be due to the
ness. There, a P3b occurs if perceptual processing of a stimulus lower temporal resolution of time–frequency analyses compared
reaches consciousness, whereas this component is absent, when to ERPs, component A6 might be associated with component
the stimulus stays subliminal (Dehaene and Changeux, 2011, for a U2 and the same may apply to components A7 and U3 (indi-
recent review). cated by a gray shading in Figure 10) and the temporal delay
between these components has the same sign as with the dERP
Beta and gamma modulations common to both endogenous and components.
exogenously induced reversals Several studies using the Manual Response Paradigm reported
All dERP components found with exogenously induced rever- enhanced frontal gamma activity within 1000 ms before the man-
sals seem to occur also with endogenous reversals, however with ual response (e.g., Basar-Eroglu et al., 1996; Strüber et al., 2000).

The gamma enhancement found with the Onset-Paradigm lies was considered as a kind of perceptual decision about the rep-
within the above 1000-ms time interval and may reflect the resentation of an ambiguous stimulus appearing (and then dis-
same neural processes. The spatial differences between gamma appearing) on a blank screen. The underlying mechanisms were
enhancements and the missing beta effect with the Manual assumed to differ from those of percept switch mechanisms,
Response Paradigm may be explained by different control con- i.e., reversals during continuous observation of an ambiguous
ditions. figure.
According to Noest et al. (2007), all findings with the Onset
Interpretation of the beta and gamma modulations. The Paradigm belong to the latter reversal type. It is not entirely
gamma-band modulation at 250 ms is the earliest signature at clear, however, how and when exactly (at which ISI) the transition
higher frequencies in the case of endogenous reversals with a between percept choice and percept switch takes place. Based on
corresponding component from exogenously induced reversals. the following theoretical and empirical arguments, including the
Interestingly, its start matches well the latency of the RN, the earli- data presented above, we argue that reversals during interrupted
est dERP component common to both types of reversal. According stimulation with short ISIs (e.g., <400 ms) may still belong to the
to Kornmeier and Bach (2006) these findings indicate that at least percept switch events and that the above-presented results from
250 ms after stimulus onset, the “decision” about the perceptual the Onset Paradigm may also apply to the continuous case.
outcome has taken place. Thus all signatures at 250 ms and later
must be of secondary order and subsequent to the reversal process, 1. The empirical data show a non-monotonic modulation of
and the underlying processes must be very similar if not identical reversal rates mainly as a function of ISIs with a maximum
for the two types of stimuli. Their detailed functional roles have around an ISI of 300 ms (Orbach et al., 1963, 1966; Kornmeier
to be discovered in future experiments. et al., 2002), an asymptotic decrease toward zero reversals for
IS IT VALID TO ANALYZE COMPONENTS FROM DIFFERENCE TRACES? longer ISIs (e.g., Orbach et al., 1963; Orbach et al., 1966; Korn-
Amplitude modulations in an ERP difference trace can result from meier et al., 2002; Leopold et al., 2002; Maier et al., 2003),
a modulation in strength of components before subtraction or they and a smooth monotonic decrease of reversal rates toward the
are simple artifacts from differential latency shifts of the under- continuous case for shorter ISIs (Orbach et al., 1963, 1966;
lying components in the two conditions. A combination of both Kornmeier et al., 2002, 2007; see also Figure 3). This non-
is also possible. Figures 5A and 6 show roughly equal latencies of monotonic behavior of reversal rates provides evidence for the
our raw ERP traces (before subtraction) across reversal and sta- existence of two different processes and suggests an estimate
bility conditions for each of the two stimulus types. Analyzing the of roughly 400 ms for the transition point. For shorter ISIs the
dERPs thus seems to be valid. underlying processes may be similar to the continuous case to
Any EEG deflection, however, is a spatial summation of an which the observed reversal rates converge (switch events). For
unknown number of differing generators. A component of a ISIs above 400 ms, perceptual reversals may be simply separate
difference trace can thus result from a complex and differential percepts (choice events). The recently proposed Necker-Zeno
interplay of different underlying generators in the two condi- model of bistable perception (Atmanspacher et al., 2004, 2008)
tions. Britz and Pitts (2011) thus additionally compared current lends additional, theoretical support to this view. It success-
density maps and source images of raw ERPs and of dERPs. Fur- fully models the non-monotonic behavior of reversal rates with
ther they identified clusters of brain microstates (Lehmann and discontinuous stimulus presentations and predicts a simple
Skrandies, 1980; Michel et al., 2004) with temporally stable global relation between three basic time scales in cognitive neuro-
field power (spatial SD of the potential field) and compared them science. ISIs of 300 ms and longer can be identified with one
between the reversal, stability and difference traces. In summary of these time scales, while for ISIs smaller than 300 ms this is
they found converging evidence that all the reversal-related sig- not the case. A most recent summary of the results obtained
natures reported above result from strength-based modulations with the Necker-Zeno Model can be found in Atmanspacher
of stimulus-related neural generators. This qualifies the stability and Filk (2010).
condition as a valid control and further validates the analysis of 2. The RP, as a dERP component specifically related to endoge-
the signatures from the difference traces. nous reversals, has been reported so far in several studies with
discontinuous presentation of ambiguous figures using small
ARE PERCEPTUAL REVERSALS DURING DISCONTINUOUS STIMULUS ISIs: from 14 to 390 ms in Kornmeier et al. (2007), and even
PRESENTATION A GOOD MODEL FOR THE CONTINUOUS CASE? 600 ms in Britz et al. (2009)2 . It is invariant in amplitude and
The gain in temporal resolution of the endogenous reversal
process, allowing a high temporal resolution of ERP compo- 2 Psychophysical data collected with the Necker cube indicate a transition point
nents, comes at the cost of the periodical interruption of the between switch reversals and choice reversals at ISIs between 300 and 400 ms
stimulus presentation. Noest et al. (2007) recently discussed exten- (Orbach et al., 1963). Kornmeier et al. (2007) presented a more complex Necker
sively whether the basic mechanisms of reversals induced by these lattice, composed of nine simple cubes, and found a rising reversal rate together
interruptions resemble or differ from the reversal mechanism with a unchanged RP in a ISI-range between 10 and 400 ms. Britz et al. (2009), pre-
during continuous observation. They distinguished between “per- senting a lattice composed of four Necker cubes, identified the RP even with an ISI
of 600 ms, which indicates a discrepancy between a physiological and a psychophys-
cept switch” events during continuous presentation and “percept ical estimate of the transition point. Results from Kornmeier et al. (2002) however
choice” events during discontinuous presentation of an ambigu- indicate that the transition point in the case of more complex Necker lattices are at
ous figure and present a model for the latter. A percept choice larger ISIs compared to the simple Necker cubes.

latency within an ISI-range at least up to 400 ms (Kornmeier Table 1 provides an overview of all here discussed EEG
et al., 2007). Interestingly, O’Donnell et al. (1988), presenting signatures found with perceptual reversals of ambiguous and
their Necker cubes discontinuously with ISIs as long as 3300 ms, unambiguous figures, their latencies, their locations and the stim-
did not find a RP. We would agree with Noest et al. (2007) that, uli used to induce them. Interestingly, Lumer et al. (1998) reported
given such long ISIs, they observed perceptual choices rather frontal and parietal fMRI activation during perceptual reversals
than perceptual switches. of binocular rivalry stimuli. Further, Sterzer and Kleinschmidt
3. A typical eye-blink roughly lasts 200 ms, occurs every 4 s on (2007) found overlapping parietal and frontal brain structures
average (e.g., Caffier et al., 2003) and interrupts continuous with higher fMRI BOLD activity during endogenous reversals
stimulus presentation. Thus even the continuous case (percep- of the SAM stimulus and exogenously induced reversals of a
tual switch event) is full of short perceptual interruptions. disambiguated SAM version compared to related control con-
ditions. This is in line with several fMRI studies report about
In summary, it seems appropriate to systematically distinguish a parietal–frontal network being activated during spontaneous
between percept choice and percept switch events. However, both perceptual reversals of ambiguous figures and binocular rivalry
empirical and theoretical arguments indicate that the mechanisms stimuli (Sterzer et al., 2009 for a review). Due to the much
underlying perceptual reversals during discontinuous stimulus coarser temporal resolution of BOLD responses it is difficult to
presentations with short ISIs are very similar – if not identical – as discern between primary processes directly related to the rever-
during continuous observation. We thus expect that the RP as the sal event and secondary processes after the perceptual decision
earliest ERP correlate of perceptual reversals during discontinu- has taken place. Keeping this in mind, it may well be possi-
ous presentation of ambiguous figures would also be identified ble that the parietal and frontal fMRI activity may be related to
in the continuous case if the necessary temporal resolution were the RN and the Parietal and Frontopolar Positivities, discussed
available. The exact transition point between percept switch and above. According to our reasoning at least the latter three sig-
percept choice events may depend on the type of ambiguous natures are of secondary nature. A recent review discussing cor-
stimulus used. relates of conscious perception indicates that perception related
neural activity after 200 ms reflects the “ignition” of a large-scale
SUMMARY AND PREDICTIONS prefronto-parietal network, necessary for conscious perception
The Onset Paradigm successfully synchronizes endogenous per- and that the P3b ERP component is a relevant part of the related
ceptual reversals of ambiguous figures with stimulus onset neural activity (Dehaene and Changeux, 2011). The Parietal and
with a precision of about ±30 ms. It provides a series of Frontopolar Positivies are broad and temporally overlapping ERP
EEG signatures related to endogenous reversals which is very occurring in the same time range as reported for the P3b. They
similar to a series of EEG signatures related to exogenously are thus good candidates to reflect the recurrent activity of this
induced reversals of unambiguous stimulus variants with three network.
exceptions:
(1) An occipital RP 130 ms after stimulus onset and (2) a
FOCUSING THE BOTTOM-UP VS. TOP-DOWN CONTROVERSY
left-hemispheric occipital to frontopolar decrease in alpha-band
WITH EEG
activity, starting at the same time and lasting for about 60 ms, are
The present results suggests as optimal starting point to probe the
restricted to endogenous reversals of ambiguous figures. (3) All
influence of bottom-up and top-down factors on the EEG in a
subsequent signatures are delayed with endogenous compared to
critical time window (temporal ROI) between stimulus onset and
exogenously induced reversals.
the RN at 250 ms.
The smallest delay lasts roughly 40 ms and is visible in the
earliest component occurring with both types of reversals, the
occipital/parietal RN, starting at 260 ms with endogenous reversals EEG CORRELATES OF TOP-DOWN INFLUENCE
and at 220 ms with exogenously induced reversals. One of the strongest top-down factor influencing the reversal
We speculate as follows: The RP is a marker of a decision conflict dynamics of ambiguous figures is volitional control: Although per-
arising with ambiguity at a certain stimulus-specific perceptual ceptual reversals cannot be totally prevented, the rate of perceptual
processing step. The concurrent alpha-band decrease may reflect reversals can be doubled with the instruction to alternate the per-
two loops of recurrent activity in order to resolve the ambiguity cept as fast as possible, and it can be halved by the instruction to
within at most 60 ms. prevent reversals (e.g., Liebert and Burk, 1985; van Ee et al., 2005;
The similarity of the subsequent EEG components indicates Kornmeier et al., 2009). To our knowledge only three EEG stud-
that after a certain step perceptual processing of endogenous ies on volitional control about ambiguous figures perception have
reversals and exogenously induced reversals are very similar if been conducted so far, two of them already published: Mathes et al.
not identical. The earliest and shortest temporal delay between (2006) used the Manual Response Paradigm with the Necker cube
EEG components to endogenous and exogenously induced rever- and found increase in both a slow positive component (labeled as
sals is in the same time range as the duration of the alpha- increase in delta band activity and probably identical to our Pari-
band decrease and thus provides a good estimation of 40–60 ms etal Positivity) and in gamma-band power when participants tried
endogenous disambiguation time (Kornmeier and Bach, 2006). to volitionally prevent reversals compared to trials when they tried
Disambiguation is completed at the latest 250 ms after stimulus to volitionally induce them. While the slow positivity is clearly
onset. outside our ROI before 250 ms, it is not possible to decide this

Table 1 | EEG correlates of endogenous and exogenously induced (parentheses) perceptual reversals and their temporal occurrences with
reference to stimulus onset (onset paradigm) or to reaction times (manual response paradigm).
Peak latency (ms) Location Stimuli
Stimulus onset = 0 Reaction time = 0
Reversal positivity 1301–6 −470 Occipital electrodes1–3 Necker cube2 , Necker lattice1, 3 , old/young
(RP) woman4 , vase/face stimulus5 , Schroeders
Primary visual areas6 staircase5 , Binocular rivalry stimuli6
Alpha-power decrease −400 to +600 −1000 to 08, 10, 20 Parietally distributed8 Necker lattice7 , Necker cube10, 19 , SAM8
(≈10 Hz) (−1400– +600) (−2000–0)8
130–2007 −470 to −400 Left-hemispheric, from occipital
to frontopolar electrodes7
Reversal negativity 2601–6, 11–14 −340 (−380) Occipital and parietal Necker lattice1, 3, 7, 12–14 , Necker cube9 ,
(RN) (220)1, 11, 12, 14 electrodes1, 11–13 face/vase5 , old/young woman4 , Schröder
Lateral occipital and inferior staircase5 , Binocular rivalry stimuli6
temporal areas6, 9
Late (incl. parietal and 3401, 4, 11–13 −260 (−300) Frontopolar electrode1, 4, 12, 13 Necker cube9, 16–18 , Necker lattice7 , old/young
frontopolar) positivity (300)1, 11, 12 woman4 , SAM8, 15 , Binocular rivalry stimuli6
4701, 3, 4, 11–15 −130 (−200) Parietal electrodes1, 4, 12–14 ,
(400)1, 11, 12, 15 inferior temporal, and superior
parietal regions9
350 −2508, 16–18 Right parietal electrodes15–18, 20
Beta power increase 340 (180)7 −260 (−420) Parietal and central electrodes7 Necker lattice7
(14–26 Hz)
Gamma power −400 to +600 −1000 to Right frontal electrode20 SAM20, 21 , Necker cube16, 17 , Necker lattice7
increase (≈30–70 Hz) –016, 17, 20, 21
−2007 −800 Right parietal/central electrodes7
300 (150)7 −300 (−450) Left-central electrodes7
Global field power −503 −650 Right inferior parietal lobe3, 22 Necker lattice3 , SAM24 , Binocular rivalry
effects −30024 −900 stimuli22
Bold indicates raw values, regular type indicates values are translated by a 600-ms reaction time2, 8, 12 .
1
Kornmeier and Bach (2005), 2 Kornmeier et al. (2011), 3 Britz et al. (2009), 4 Kornmeier and Bach (2004a), 5 Pitts et al. (2007), 6 Britz and Pitts (2011), 7 Ehm et al. (2011),
8
Strüber and Herrmann (2002), 9 Pitts et al. (2009), 10
İşoğlu-Alkaç (2000), 11 Kornmeier et al. (2001), 12
Kornmeier and Bach (2004b), 13
Kornmeier et al. (2007), 14
Pitts
15 16 17 18 19 20
et al. (2008), Intaite et al. (2010), Basar-Eroglu et al. (1993), Strüber et al. (2001), Mathes et al. (2006), O’Donnell et al. (1988), İşoğlu-Alkaç and Struber (2006),
21
Basar-Eroglu et al. (1996), 22 Strüber et al. (2000), 23 Britz et al. (2010), 24 Muller et al. (2005).
for the gamma modulation, occurring in a time window 500 ms EEG CORRELATES OF BOTTOM-UP INFLUENCE
before the late positivity peaks. Presentation mode is regarded as a bottom-up factor (Long and
Pitts et al. (2008) used the Onset Paradigm with the Necker Toppino, 2004) strongly modulating the reversal dynamics of both,
lattice and found an increased RN with a perceptual reversal when ambiguous figures and binocular rivalry stimuli (e.g., Orbach et al.,
participants were instructed to reverse as often as possible com- 1963; Kornmeier et al., 2002; Leopold et al., 2002; Maier et al., 2003;
pared to a passive viewing condition without volitional effort. No Kanai et al., 2005, Noest et al., 2007; see Figure 3). The reversal rates
effects were found in the ROI before 250 ms. of different ambiguous figures and also binocular rivalry stimuli
In an EEG study from our lab with a slightly different variant can more than double or even brought to stand still as a function
of the Onset Paradigm (so far only published as diploma thesis, of ISI with discontinuous stimulus presentation (Orbach et al.,
Hein, 2006) we found central and left temporal ERP components 1963; Kornmeier et al., 2002; Leopold et al., 2002; Maier et al.,
370 ms after stimulus onset, which differed between an instruction 2003; Klink et al., 2008). In an EEG study with the Necker lattice
to reverse condition and a passive condition as well as between we looked for modulations in the first 250 ms after onset as a func-
the instruction to reverse condition and an instruction to prevent tion of ISI. All EEG modulations we found, however, occurred at
reversals condition. Again no effects were found in the ROI before and after 250 ms, no modulation within the critical ISI could be
250 ms. observed (Kornmeier et al., 2007).
In summary, participants could successfully modulate their
reversal rates if instructed to do so, but no EEG effects related to SUMMARY
volitional control were found in the critical time window between Neither the top-down factors nor the bottom-up factors tested
stimulus onset and the RN at 250 ms. so far, significantly modulated EEG data within the critical time

window between stimulus onset and the occurrence of the RN at attractor, the related neural assembly is “activated” and the per-
250 ms. This raises some doubts about the expected importance cept is (consciously) perceived (e.g., Kornmeier et al., 2004). Let
of this time period for the perceptual reversal process and about us look at two extreme cases: If the visual stimulus is completely
our above interpretations of the EEG literature on multistable unambiguous we have a single unique and deep attractor and
perception. However, maybe our interpretations are correct but thus a stable conscious experience of the represented object. In
our predictions concerning expected bottom-up and top-down the case of an ambiguous figure, two (or even more) represen-
modulations were wrong. tations fit with the visual information and thus two (or more)
perhaps shallower attractors are in close vicinity with a rela-
PROPOSAL OF AN INTEGRATIVE THEORY tively low barrier in between. For the ambiguous Necker cube,
In the following, we present a new perspective that can nicely the two 3D interpretations would correspond to the two attractors
integrate our and previous bottom-up and top-down results. It (Figure 12). A perceptual reversal occurs if the system escapes from
assumes two separate processes underlying spontaneous percep- one attractor and jumps into the alternative one. The main ques-
tual reversals, destabilization and restabilization/disambiguation, tion addressed by the research of the last 200 years on ambiguous
working on different time scales (seconds and milliseconds). figures could be translated to “what is the mechanism under-
The following thoughts are based on accumulating evidence lying the spontaneous state change from one attractor to the
from psychophysical, EEG, and fMRI studies. All the so far used other?”
methods have advantages and limitations and none of the tech- We here regard it as conceptually fruitful to subdivide the
niques allow a complete view on the processes at work during reversal event into two steps:
spontaneous perceptual alternations. It may thus well be pos-
sible that highly relevant neural activity is restricted to a small DESTABILIZATION
number of neurons and neither detectable with EEG nor with The observation of an ambiguous figure typically leads to a
fMRI methods. The absence of detectable signatures in a spe- transiently stable percept, which destabilizes over time. This
cific paradigm can thus mean that no activity is present or that destabilization is relatively slow and takes on average 5–6 s if a
such activity is not measurable with the methods at hand. Hav- Necker cube is passively observed (Orbach et al., 1963; Kornmeier
ing this in mind, we regard the following as a working hypothesis, et al., 2009). However, destabilization times (also known as stabil-
allowing us to develop predictions that may be tested in future ity durations or dwell times) are highly variable within and across
experiments. participants (e.g., Borsellino et al., 1972) and also differ between
The terminology from non-linear dynamics provides a help- different types of ambiguous figures (e.g., Strüber and Stadler,
ful framework to describe the mechanisms underlying sponta- 1999); most of the bottom-up and top-down literature on ambigu-
neous perceptual reversals of ambiguous figures (Haken, 1983; ous figures describes the variables that alter destabilization time.
Atmanspacher, 1992; Kruse et al., 1996; Kornmeier et al., 2004; Often, passive adaptation of low-level neurons is discussed as the
Braun and Mattia, 2010). Brain states are regarded as points in mechanism underlying a slow destabilization (e.g., Orbach et al.,
a hypothetical state space. Representations of objects, e.g., a 3D 1963; Toppino and Long, 1987). This can be modeled as a slow
cube, are modeled as attractors and their depth as a measure of depth reduction of the attractor in which the state is located. Spon-
the current representation’s stability. Physiologically, the form of taneous fluctuations (e.g., synaptic fluctuations, variations in neu-
the attractor may depend on the connectivity matrix of neurons rotransmitter concentrations, etc.) or exogenously induced neural
within a relevant assembly but also on the quality of the visual fluctuations (e.g., by blinks and/or eye movements, visual tran-
input. If the state of the perceptual system is located within an sients, like short interruptions (Orbach et al., 1963) of stimulus
FIGURE 12 | Destabilization and disambiguation of ambiguous visual factors can accelerate and decelerate this process. After transition of a state
information. With prolonged observation of the ambiguous lattice stimulus a of maximal instability, fast (40–60 ms) disambiguation takes place, leading to a
transiently stable percept gets destabilized slowly, symbolized as a depth different stable percept. The brain state is now located in the alternative,
reduction of the brain states current attractor. Both bottom-up and top-down deeper attractor.

presentation or light flashes (Kanai et al., 2005)) may also be rele- a more stable state. The early RP dERP component and the Beta
vant factors for a perceptual reversal and even more, the flatter the band deactivation may be related to the fast disambiguation of
current attractor is. At least a part of the data from the Orbach ambiguous visual input, taking place immediately after stimulus
group may thus be explained by an interaction between tran- onset (within the Onset Paradigm) and being time-locked to it
sients (fluctuations) and adaptation (influencing attractor depth). (±30 ms). Destabilization, on the other hand, may start imme-
Moreno-Bote et al. (2007) recently provided a neurally plausible diately after the new percept has been established and is much
attractor model that assumes weak adaptation and noise under- slower. It can take from seconds to minutes and its dynamic can
lying perceptual alternations. This model explains nicely several be changed in different ways and perhaps at different time points.
empirical findings like gamma distributed dwell times and more. Any EEG correlate of destabilization should thus occur before the
Effects of volitional control on reversal rates fit within the onset of the stimulus that is perceived as reversed and it should
attractor model in the following way: The instruction to voli- indicate an upcoming reversal.
tionally control the percept, typically given at the begin of an
experimental block, may cause an a priori “top-down” change of An EEG correlate of perceptual destabilization
depth of specific attractors in the sense of a global change of base- Indeed, we recently found an increase in lower gamma-band
line activity or of a certain threshold, and thus prolong or shorten activity (26–40 Hz) at the right-hemispheric central and parietal
the destabilization time and increase or decrease the influence of electrodes roughly 200 ms before onset of a Necker lattice that is
fluctuations on the transition probability. perceived as reversed compared to a lattice that is perceived as sta-
In summary a neural representation can be modeled as a brain ble (Figures 9 and 10, component U1) together with an occipital
state, located in an attractor within a state space. The depth of the decrease of higher gamma-band activity (40–65 Hz; Figures 9 and
attractor depends on the quality of the visual input (amongst oth- 10, component U2). No such gamma modulations were observed
ers). The more ambiguous the input is, the shallower is the attrac- with exogenously induced reversals of unambiguous lattice vari-
tor, the more sensitive the representation is to spontaneous fluctu- ants (Ehm et al., 2008, 2011). Britz et al. (2009) also used the
ations and the more probable is a reversal between representations Necker lattice with the Onset Paradigm. They compared dom-
(attractors). It is reasonable to assume that in the case of ambigu- inant potential maps with temporally stable global field power
ous visual information the attractor, which is currently occupied, (spatial SD of the potential field) and current source distributions
is initially shallow and slowly flattens over time. Different bottom- for reversal and stability trials in a 50-ms pre-stimulus interval and
up (e.g., mode of stimulus presentation, Kanai et al., 2005) as well also found significant differences in the right inferior parietal lobe.
as top-down factors (like volitional control, Kornmeier et al., 2009) No unambiguous stimulus variants were used in their experiment.
may be able to alter attractor depth and/or increase the noise level A series of related studies underscore the importance of the
(Moreno-Bote et al., 2007) and thus influence the reversal dynam- above findings: Basar-Eroglu et al. (1996) reported a right ante-
ics, even simultaneously in an additive manner (Kornmeier et al., rior gamma power increase within 1000 ms before participants
2009). Bottom-up and top-down explanations are thus no longer indicated an endogenous perceptual motion reversal of the SAM
mutually exclusive with this conception. stimulus. We assume that their gamma modulation contains
both, the pre-stimulus and post-stimulus gamma modulations
RESTABILIZATION/DISAMBIGUATION indicated in our data (Ehm et al., 2011). Roeber et al. (2008)
Each reversal from one stable percept to another passes through a recently reported a right-hemispheric ERP correlate of percep-
point of maximal instability when the perceptual state is on top of tual reversals of binocular rivalrous sine wave gratings. Nakatani
the barrier between the two related attractors (Figure 12). and van Leeuwen (2006) found EEG gamma-band synchroniza-
The sensory information we receive is inherently incomplete tion between right-hemispheric parietal and right-hemispheric
and ambiguous. We have to disambiguate and interpret it in order frontal electrode positions 800–600 ms before the manual indi-
to perceive it. Our perceptual system is optimized to disambiguate cation of a Necker cube reversal. VanRullen et al. (2006) found
and interpret the visual information as fast as possible (e.g., we higher gamma activity at right-hemispheric central locations with
immediately perceive faces or objects in the formation of clouds illusory motion direction reversals of the Wagon-Wheel Illusion
in the sky or in the formation of country rocks). This indicates compared to real motion reversals. Lumer et al. (1998) reported
that due to evolutionary reasons our perceptual system tries to selective right-hemispheric BOLD (fMRI) activation during per-
keep the inevitably instable brain states in between attractors as ceptual transitions of binocular rivalry stimuli, but no such activity
short as possible and thus to achieve a fast perceptual interpreta- with exogenous transitions of unambiguous stimulus variants.
tion of whatever sensory information is available in order to be Sterzer and Kleinschmidt (2007) found increased fMRI response in
able to react immediately. the right inferior frontal cortex with endogenous motion reversals
These considerations apply to ambiguous-figure perceptions of the SAM stimulus compared to exogenously induced reversals of
in the following way: Leaving of an attractor (destabilization) and unambiguous SAM variants. Similarly, Ilg et al. (2008) found pos-
arriving at another attractor (disambiguation/restabilization) are terior right-hemispheric fMRI activity with spontaneous motion
different processes, working on different time scales (minutes and direction reversals of the spinning wheel illusion (Wertheimer,
seconds vs. milliseconds): Given a brain state of maximal insta- 1912), but no such activity with exogenously induced reversals.
bility either at the onset of an ambiguous figure or as a result of Müller et al. (2005) used the onset of the SAM stimulus imme-
the above described slow destabilization process during prolonged diately before a button press as the time reference for reversals
observation, our perceptual system tries to find as fast as possible of motion direction. They found changes in EEG activity about

300 ms before the reversal-related SAM flashes, i.e., temporally a continuously observed Necker cube. Again, a sufficiently precise
close to our pre-onset gamma modulation. Meenan and Miller time reference for a reversal in the continuous case is necessary to
(1994) reported about difficulties of patients with lesions in right test this.
frontal areas to recognize more than one interpretation of several
ambiguous figures. TWO (OF SEVERAL) MAJOR OPEN QUESTIONS IN THE
Further evidence for an important role of right-hemispheric CONTEXT OF MULTISTABLE PERCEPTION
areas comes from recent studies with transcranial magnetic stim- WHAT DO NEURAL REPRESENTATIONS LOOK LIKE – AND WHY ARE
ulation (TMS). It was shown, that TMS stimulation of the parietal THEY SOMETIMES UNSTABLE?
lobe can modulate dwell times of ambiguous structure-from- This is essentially the question about perceptual states and their
motion stimuli (Kanai et al., 2010, with bilateral stimulation) destabilization over time with ambiguous visual input. One of
and binocular rivalrous moving gratings (Carmel et al., 2010; the basic assumptions in the multistable perception literature
Kanai et al., 2011, with right-hemispheric stimulation). Stimu- is that each conscious interpretation of an ambiguous figure
lation of the posterior parts of the parietal lobe increased dwell is based on the activity of a distinct neural assembly (e.g.,
times whereas stimulation of anterior parts decreased dwell times Blake and Logothetis, 2002, for a review). Thus one central
(Kanai et al., 2011). Zaretskaya et al. (2010) found during percep- goal of the physiological studies of multistable perception was
tual reversals of a rivalrous face/house stimulus a stronger BOLD to find brain areas where neural activity fluctuates in correla-
response in the right intraparietal sulcus in nine participants and tion with the perceptual experience, in contrast to areas with
stronger BOLD response in the left intraparietal sulcus in 6 partic- unchanged neural activity, reflecting the unchanged visual input.
ipants. TMS stimulation of the right intraparietal sulcus increased Especially the border between sensory and percept-related activ-
dwell times. ity was assumed to play a key role for the perceptual reversal
In summary, there is converging evidence that right- process and consciousness in general (e.g., Blake and Logothetis,
hemispheric brain areas play an important role during sponta- 2002).
neous perceptual reversals across stimulus types and categories Logothetis et al. (e.g., Logothetis, 1998; Leopold and Logothetis,
(ambiguous figures and binocular rivalry stimuli) and experi- 1999) recorded from primate single cells in different brain areas,
mental paradigms. More precisely, some studies, providing precise while the animals reported their percepts of binocular rivalry stim-
temporal information, indicate that this activity precedes percep- uli. They found that the number of neurons with a percept-related
tual reversals and may thus have predictive character, although this firing pattern increased from V1 (striate cortex, 20%) over V2, V4,
has to be demonstrated on a single trial level. Further, the reversal MT, medial superior temporal sulcus (MST, 80), the inferotempo-
dynamics can be altered by right-hemispheric TMS stimulation. ral cortex (IT, 80%), and the superior temporal sulcus (STS, 80%;
The spatial variability of the right-hemispheric signature across Logothetis, 1998; Leopold and Logothetis, 1999). A large number
studies is so far unexplained. of V1 neurons remained active whether the stimulus was per-
So far we can only speculate about the function of this ceived or suppressed. Percept-related firing patterns were neither
right-hemispheric pre-onset activity. A common feature across found in monocular V1 neurons (e.g., Leopold and Logothetis,
all stimulus types, stimulus categories, and paradigms is that 1999) nor in LGN neurons (Lehky and Maunsell, 1996). Several
perception changes spontaneously while the visual information results from fMRI studies are in line with these single cell findings,
stays unchanged. As a working hypothesis we suggest that the indicating the importance of higher cortical areas for object rep-
pre-onset gamma modulation indicates a transient brain state of resentation. Tong et al. (1998) presented binocular rivalry stimuli
maximal instability at the transition from one stable brain state containing the picture of a house and of a face and found recipro-
to another or its recognition by some unconscious and so far cal modulations in BOLD activity in the parahippocampal place
unknown neural instance. Such an instable brain state in between area and the fusiform face area highly correlating with partici-
two adjacent attractors is a necessary pre-condition for a percep- pants’ reports of seeing a face or a house. Interestingly, this level
tual reversal to occur and thus for a decision conflict preceding a of modulation was comparable to that with binocular house and
reversal. Moments of instable brain states should thus be inherent face perceptions (non-rivalrous conditions). Similarly, Andrews
in all types of perceptual reversal and related right-hemispheric et al. (2002) presented Rubin’s ambiguous Face/Vase stimulus and
modulations – perhaps in the EEG gamma band – should be found slightly increased BOLD activity in the fusiform face area
observable. A sufficiently precise time reference for a reversal in during the perception of the faces compared to trials with house
the continuous case is necessary to test this percepts. Recently Watanabe et al. (2011) found in V1 strong fMRI
During continuous viewing of an ambiguous figure, state insta- BOLD effects of attention but none of dominance vs. suppression
bility (or its recognition) and the subsequent decision conflict intervals of binocular rivalry stimuli.
during (re)interpretation of the ambiguous visual information Other studies point to the importance of lower-level areas for
may be in close temporal vicinity. The short interruptions in the object representations: Dodd et al. (2001) reported correlation
Onset Paradigm may act as a “temporal magnifier,” slowing down between single cell activity in primate area MT and the per-
the reversal process and thus temporally separating the two steps. ceived motion direction of an ambiguous structure-from-motion
According to these considerations a second working hypothesis, stimulus. Further, fMRI studies provided evidence for percept-
may thus be that the right-hemispheric central gamma modu- related activity in V1 (e.g., Tong and Engel, 2001) and in the LGN
lation, which is pre-onset with the Onset Paradigm, may occur (Haynes et al., 2005) during humans’ observation of binocular
immediately before the occipital RP during a perceptual reversal of rivalry stimuli.

Today there are potential explanations for the inconsistency this process is located at lower visual or at high-level cognitive
between single cell and fMRI findings: fMRI BOLD activity is processing units (e.g., Long and Toppino, 2004).
(like EEG activity) more related to local field potentials than to We suggest the reversal process can be subdivided into two
neural spiking activity (Logothetis et al., 2001). Thus percept- separate steps, which need to be understood:
related modulation in early visual areas, as shown with fMRI, (1) How the current brain state gets out of its attractor and on
should be reflected in local field potentials rather than in spike top of the barrier in between this one and the alternative attrac-
modulation. This has been demonstrated recently in primates tor. (2) How the “decision” takes place about whether the instable
(Wilke et al., 2006; Maier et al., 2008). Further, information may be brain state returns to the old or the alternative attractor or perhaps
coded by temporal synchronization of neural activity (i.e., tempo- whether it approaches another, so far less probable attractor.
ral coding), rather than amplitude modulations. Fries et al. (1997) Both steps depend on several factors like the energy distri-
presented binocular rivalry stimuli to cats and found precept- bution of the state space at this specific moment and thus the
related synchronization of neural gamma-band oscillations in V1. depth of the currently activated and the alternate attractors and
Like surface EEG, the MEG is believed to reflect synchronous spik- the amount of energy fluctuations within the system (endogenous
ing activity of a large number of cortical neurons. MEG studies on and/or exogenously introduced neural noise or background activ-
binocular rivalry showed correlations between activity over a wide ity). Recent empirical evidence indicates that these factors depend
range of sensors (from occipital to frontal lobes) and observers at least in part on the perceptual history (on different time scales;
dominance and suppression reports, indicating the involvement of e.g., Hesselmann et al., 2008; Pastukhov and Braun, 2008). Influ-
the entire cortex in conscious object representation (e.g., Tononi ence may be exerted at different steps and levels of complexity
et al., 1998; Dehaene and Changeux, 2011). during visual perception (Blake and Logothetis, 2002; Kornmeier
In summary, there is no isolated cortical area selectively cor- and Bach, 2006; Sterzer and Rees, 2008; Bartels and Logothetis,
relating with the participant’s current percept. The locus of the 2010), which is in line with the distributed object representation
attractor and its specific neural realization is so far not under- account discussed above. Nakatani and van Leeuwen (2006), e.g.,
stood (e.g., Moreno-Bote et al., 2007). Rather, object representa- provided evidence for different types of reversal of a Necker cube
tions seem to result from a complex, probably reciprocal interplay within and between participants, reflected by different patterns
between early visual and several higher brain areas across the cor- of synchrony in EEG oscillations. Blake and Logothetis (2002)
tex. Further, most of the physiological correlates are transient in summarized related evidence from the binocular rivalry literature
nature and so far it is unclear how the maintenance of a stable and Bartels and Logothetis (2010) found recently with binocu-
conscious percept is neuronally realized (Ehm et al., 2011). This lar rivalry stimuli, that perceptual reversals can be triggered at
however, has to be understood in order to understand the destabi- different levels of complexity during the perceptual process.
lization of a given representation in the case of ambiguous figures In summary, there is probably no unique neural switch area or
and its susceptibility to bottom-up and top-down factors. There “unique gate to conscious perception” within the brain that gov-
is some experimental and theoretical evidence that neural activity erns perceptual reversals. However, the EEG findings discussed
is somewhat weaker with ambiguous or rivalrous stimuli com- above indeed indicate some generality aspects across types of
pared to disambiguated variants (e.g., Leopold and Logothetis, reversals, types of stimuli (e.g., Necker cube or old/young woman)
1999; Kornmeier and Bach, 2006; Moreno-Bote et al., 2007; Pitts and even across categories (ambiguous figures and binocular
et al., 2010). This may indicate that the activated attractors are in rivalry stimuli). First, each endogenous perceptual reversal comes
general flatter and thus more susceptible to any type of fluctuation with a transient state of instability at the barrier between the
the more ambiguous the visual information is. In a recent study two alternative attractors (Figure 11). The generality of the pre-
we analyzed this systematically with ambiguous figures from dif- reversal right-hemispheric activity (e.g., Britz et al., 2009; Britz
ferent categories: Slight low-level figural changes of an ambiguous et al., 2010; Ehm et al., 2011) may classify this modulation as a
figure disambiguate it and produce more sustained stable per- good candidate reflecting this instable brain state or its detection.
cepts. We found dramatic ERP differences between ambiguous What this would mean in terms of neural processing, however, is
figures and their disambiguated variants, independent of the rever- so far unclear. More research has to be done in order to determine
sal dynamics and across different types of figures (Necker cube, more attributes of this right-hemispheric activity modulation in
Old/Young Woman and SAM stimulus, as latest – unpublished- detail.
results from our lap indicate). This difference might (directly Second, a general finding across stimulus types (Necker cube,
or indirectly) reflect the difference in depth of the respective Old/Young woman) and categories (ambiguous figures and binoc-
attractors (Kornmeier and Bach, 2009). ular rivalry stimuli) is the early occipital RP. It may be possible that
after some central instance (perhaps general across types of rever-
WHO DECIDES ON THE PERCEPTUAL RESULT IF THE VISUAL INPUT IS sals) has notified an instable perceptual brain state (reflected in
AMBIGUOUS? the right-hemispheric activity modulation, described above), the
This is essentially the question about perceptual transitions. A “decision” about the perceptual outcome needs one (or two) top-
large number of psychophysical studies about ambiguous figures down reactivation(s) of early visual areas for reinterpretation of
aimed to find or understand “the one mechanism” that underlies the available visual information. It may also be possible that the
spontaneous perceptual reversals – they looked for the deciding surface EEG can only detect the strong activity of the “low-level
instance or mechanism. The major difference between bottom-up receiver” (i.e., the RP) but not the potentially weaker activity of the
and top-down explanations thus pertains to the question whether “higher-level sender,” wherever in the brain it may be located. This

is highly speculative and further experiments are necessary, e.g., scribed brain regions but involve almost the entire cortex. A
to describe the attributes of the RP in more detail. A step in this clear border between sensory and perceptual processing is thus
direction has been done recently by demonstrating the RP’s inde- difficult to find and may depend on experimental and stimulus
pendence of stimulus size (Kornmeier et al., 2011) and stimulus details.
complexity (cubes or lattices Kornmeier and Bach, 2003). The available evidence further suggests that, in contrary to
previous expectations, no circumscribed neural unit exists that
SUMMARY AND CONCLUSION decides about the perceptual outcome. Rather, perceptual rever-
Applying the Onset Paradigm to investigate spontaneous percep- sals can be induced at different locations and levels of complexity
tual reversals of both ambiguous figures and binocular rivalry along the processing of visual information.
stimuli sizably improved the temporal resolution of the underly- Recent evidence from our lab suggests the existence of a central
ing processes. Since its first application several independent lines evaluation instance that estimates the reliability of the percep-
of evidence encouraged us to propose that spontaneous perceptual tual outcome, given a certain quality of visual information. The
reversals are governed by two independent processes working on outcome of this evaluation thus may indirectly reflect the depth
different time scales: (1) The transiently stable perceptual interpre- of the activated attractor (e.g., Kornmeier and Bach, 2009; Has-
tation of the ambiguous information destabilizes over time until sberg, 2010). Understanding the mechanisms of such reliability
the perceptual system reaches a state of maximal instability. This estimation may be highly relevant for a better understanding of
destabilization is slow in the order of seconds to minutes and sub- psychiatric perception disorders and ambiguous figure may be an
ject to multiple endogenous and exogenous influences. The result interesting tool for this.
of this destabilization, a perceptual brain state of maximal insta- Ambiguous figure perception is an ideal model to investigate
bility seems to be correlated with a right-hemispheric modulation changes of brain states between already existing attractors in other
in EEG activity which occurs pre-onset in the case of discontinu- domains. Already the Gestalt Psychologists regarded this phenom-
ous stimulus presentation. (2) Due to evolutionary pressure our enon as interesting for the understanding of insight phenomena
brain is optimized to keep unavoidably instable brain states as brief (Knoblich and Öllinger, 2006), where probably a transition from
as possible. Electrophysiological evidence indicates that the transi- an existing attractor to an instantaneous newly built attractor
tion from maximal instability to a (potentially altered) stable brain takes place. Most interesting in this respect may be the tran-
state is very fast, in the order of 40–60 ms, and not susceptible to sient states of maximal instability on top of the barrier between
endogenous or exogenous manipulations. EEG data further indi- two attractors (“acategorial states,” Feil and Atmanspacher, 2010).
cates that perceptual processes within 350 ms before the manual Thus ambiguous figure perception may also be an interesting tool
indication of a perceptual reversal are post-decision and thus of for future research in insight processes. The gain in temporal
secondary nature with respect to the reversal process. The (uncon- resolution attained with the Onset Paradigm provided specific
scious) decision about the perceptual outcome thus seems to be electrophysiological marker of critical processing steps underly-
rather early. ing spontaneous perceptual reversals of ambiguous figures. The
Numerous psychophysical and physiological evidence indi- speculations above about their functional role encourage precise
cates that neural representations are not restricted to circum- experimental hypotheses that may be tested in future experiments.
REFERENCES Atmanspacher, H., and Filk, T. (2010). Blake, R. (1989). A neural theory of Britz, J., Landis, T., and Michel, C. M.
Ammons, C. H., and Ammons, R. A proposed test of temporal nonlo- binocular rivalry. Psychol. Rev. 96, (2009). Right parietal brain activ-
B. (1963). Perspective reversal as cality in bistable perception. J. Math. 145–167. ity precedes perceptual alternation
affected by physical characteristics of Psychol. 54, 314–321. Blake, R. (2001). A primer on binocular of bistable stimuli. Cereb. Cortex 19,
Necker cube drawings. Proc. Mon- Atmanspacher, H., Filk, T., and Römer, rivalry, including current controver- 55–65.
tana Acad. Soc. 23, 287–302. H. (2004). Quantum Zeno features sies. Brain Mind 2, 5–38. Britz, J., and Pitts, M. A. (2011).
Andrews, T. J., Schluppeck, D., Homfray, of bistable perception. Biol. Cybern. Blake, R., and Logothetis, N. K. (2002). Perceptual reversals during binoc-
D., Matthews, P., and Blakemore, 90, 33–40. Visual competition. Nat. Rev. Neu- ular rivalry: ERP components
C. (2002). Activity in the fusiform Bach, M. (1997). Some Visual Phenom- rosci. 3, 13–21. and their concomitant source
gyrus predicts conscious perception ena and Optical Illusions. Available Boring, E. G. (1930). A new ambiguous differences. Psychophysiology 48,
of Rubin’s vase-face illusion. Neu- at: http://www.michaelbach.de/ot/ figure. Am. J. Psychol. 42, 444–445. 1490–1499.
roimage 17, 890–901. Bartels, A., and Logothetis, N. K. (2010). Borsellino, A., De Marco, A., Allazetta, Britz, J., Pitts, M. A., and Michel, C. M.
Anllo-Vento, L., and Hillyard, S. A. Binocular rivalry: a time dependence A., Rinesi, S., and Bartolini, B. (2010). Right parietal brain activ-
(1996). Selective attention to the of eye and stimulus contributions. J. (1972). Reversal time distribu- ity precedes perceptual alternation
color and direction of moving stim- Vis. 10, 3. tion in the perception of visual during binocular rivalry. Hum. Brain
uli: electrophysiological correlates of Basar-Eroglu, C., Struber, D., Kruse, P., ambiguous stimuli. Kybernetik 10, Mapp. 32, 1432–1442.
hierarchical feature selection. Per- Basar, E., and Stadler, M. (1996). 139–144. Caffier, P. P., Erdmann, U., and
cept. Psychophys. 58, 191–206. Frontal gamma-band enhance- Brascamp, J. W., van Ee, R., Pestman, W. Ullsperger, P. (2003). Experimental
Atmanspacher, H. (1992). Categoreal ment during multistable visual R., and van den Berg, A. V. (2005). evaluation of eye-blink parameters
and acategoreal representation of perception. Int. J. Psychophysiol. 24, Distributions of alternation rates in as a drowsiness measure. Eur. J. Appl.
knowledge. Cogn. Syst. 3, 259–288. 113–125. various forms of bistable perception. Physiol. 89, 319–325.
Atmanspacher, H., Bach, M., Filk, T., Basar-Eroglu, C., Strüber, D., Stadler, J. Vis. 5, 287–298. Carmel, D., Walsh, V., Lavie, N., and
Kornmeier, J., and Römer, H. (2008). M., and Kruse, E. (1993). Multistable Braun, J., and Mattia, M. (2010). Attrac- Rees, G. (2010). Right parietal TMS
Cognitive time scales in a Necker- visual perception induces a slow pos- tors and noise: twin drivers of deci- shortens dominance durations in
Zeno model for bistable perception. itive EEG wave. Int. J. Neurosci. 73, sions and multistability. Neuroimage binocular rivalry. Curr. Biol. 20,
Open Cybern. Syst. J. 2, 234–251. 139–151. 52, 740–751. R799–R800.

Crick, F., and Koch, C. (1998). Con- Gregory, R. (1974). “Choosing a par- long-term EEG recordings: sub- Kornmeier, J., Bach, M., and
sciousness and neuroscience. Cereb. adigm for perception,” in Hand- bands of alpha activity. Int. J. Psy- Atmanspacher, H. (2004). Cor-
Cortex 8, 97–107. book of Perception: Vol. 1. Historical chophysiol. 59, 179–189. relates of perspective instabilities
Debecker, J., and Desmedt, J. E. (1970). and Philosophical Roots of Perception, Kanai, R., Bahrami, B., and Rees, in visually evoked potentials.
Maximum capacity for sequential eds C. Carterette and M. Fried- G. (2010). Human parietal cortex Int. J. Bifurcat. Chaos 14,
one-bit auditory decisions. J. Exp. man (New York: Academic Press), structure predicts individual differ- 727–736.
Psychol. 83, 366–372. 255–283. ences in perceptual rivalry. Curr. Kornmeier, J., Ehm, W., Bigalke, H.,
Dehaene, S., and Changeux, J. P. Haken, H. (1983). Synergetics: An Biol. 20, 1626–1630. and Bach, M. (2007). Discontin-
(2011). Experimental and theoreti- Introduction; Nonequilibrium Phase Kanai, R., Carmel, D., Bahrami, B., uous presentation of ambiguous
cal approaches to conscious process- Transitions and Self-Organization and Rees, G. (2011). Structural figures: how interstimulus-interval
ing. Neuron 70, 200–227. in Physics, Chemistry, and Biology. and functional fractionation of durations affect reversal dynam-
Dodd, J. V., Krug, K., Cumming, B. Berlin: Springer. right superior parietal cortex in ics and ERPs. Psychophysiology 44,
G., and Parker, A. J. (2001). Per- Hassberg, T. (2010). Aufmerksamkeits- bistable perception. Curr. Biol. 21, 552–560.
ceptually bistable three-dimensional und Adaptationseffekte bei R106–R107. Kornmeier, J., Hein, C. M., and
figures evoke high choice probabili- instabiler Wahrnehmung. Med- Kanai, R., Moradi, F., Shimojo, S., and Bach, M. (2009). Multistable
ties in cortical area MT. J. Neurosci. ical Dissertation, University Verstraten, F. A. (2005). Perceptual perception: when bottom-up and
21, 4809–4821. of Freiburg, 79. Available at: alternation induced by visual tran- top-down coincide. Brain Cogn. 69,
Dubois, J., and Vanrullen, R. (2011). http://www.freidok.uni-freiburg.de/ sients. Perception 34, 803–822. 138–147.
Visual trials: do the doors of volltexte/7875/ Klink, P. C., van Ee, R., Nijs, M. M., Kornmeier, J., Heinrich, S. P.,
perception open periodically? PLoS Haynes, J. D., Deichmann, R., and Rees, Brouwer, G. J., Noest, A. J., and van Atmanspacher, H., and Bach,
Biol. 9, e1001056. doi:10.1371/jour- G. (2005). Eye-specific effects of Wezel, R. J. (2008). Early interactions M. (2001). The reversing “Necker
nal.pbio.1001056 binocular rivalry in the human lat- between neuronal adaptation and Wall” – a new paradigm with
Ehm, W., Bach, M., and Kornmeier, eral geniculate nucleus. Nature 438, voluntary control determine percep- reversal entrainment reveals an early
J. (2008). What role for gamma 496–499. tual choices in bistable vision. J. Vis. EEG correlate. Invest. Ophthalmol.
oscillations in the perception of Hein, C. (2006). Cognitive Influence on 8, 11–18. Vis. Sci. 42, 409.
ambiguous figures? Perception 37, Perceptual Reversals – ERP Correlates Knoblich, G., and Öllinger, M. (2006). Kornmeier, J., Heinrich, S. P., and
116. to the Disambiguation of the Necker “Einsicht und Umstrukturierung Bach, M. (2002). “Necker-Würfel:
Ehm, W., Bach, M., and Kornmeier, Cube. Diploma Thesis, 89. beim Problemlösen. Enzyklopädie Neuronale Sättigung und “top-
J. (2011). Ambiguous figures Hesselmann, G., Kell, C. A., Eger, E., der Psychologie,” in Enzyklopädie down” Einflüsse,” in Beiträge zur 5.
and binding: EEG frequency and Kleinschmidt, A. (2008). Spon- der Psychologie, Denken und Prob- Tübinger Wahrnehmungskonferenz,
modulations during multistable taneous local variations in ongoing lemlösen, ed. J. Funke (Göttingen: 69.
perception. Psychophysiology 48, neural activity bias perceptual deci- Hogrefe), C/II/8, 782. Kornmeier, J., Pfaffle, M., and Bach,
547–558. sions. Proc. Natl. Acad. Sci. U.S.A. Köhler, W. (1940). Dynamics in Psychol- M. (2011). Necker cube: stimulus-
Elbert, T., Hommel, J., and Lutzen- 105, 10984–10989. ogy. New York: Liverlight. related (low-level) and percept-
berger, W. (1985). The perception Hochberg, J., and Peterson, M. Kornmeier, J., and Bach, M. (2003). related (high-level) EEG signatures
of Necker cube reversal interacts (1987). Piecemeal organization and “Zwei Varianten des Necker- early in occipital cortex. J. Vis. 11,
with the Bereitschaftspotential. Int. cognitive components in object Würfels: Frühe, okzipitale lateral- 1–11.
J. Psychophysiol. 3, 5–12. perception. J. Exp. Psychol. Gen. 116, isierte ERP-Korrelate des Kruse, P., Carmesin, H. O., Pahlke,
Feil, D., and Atmanspacher, H. 370–380. Wahrnehmungswechsels,” in L., Struber, D., and Stadler, M.
(2010). Acategorial states in a Howard, I. P., and Durham, U. (1961). Beiträge zur 6. Tübinger Wahrneh- (1996). Continuous phase transi-
representational theory of mental An investigation of a satiation mungskonferenz, 157. tions in the perception of multistable
processes. J. Conscious. Stud. 17, process in the reversible perspective Kornmeier, J., and Bach, M. (2004a). visual patterns. Biol. Cybern. 75,
72–101. of revolving skeletal shapes. Q. J. “More evidence for early visual pro- 321–330.
Foxe, J. J., and Simpson, G. V. (2002). Exp. Psychol. 13, 19–33. cessing in perceptual disambigua- Lehky, S. R., and Maunsell, J. H. (1996).
Flow of activation from V1 to frontal Ilg, R., Wohlschlager, A. M., Buraza- tion of ambiguous figures,” in Pro- No binocular rivalry in the LGN of
cortex in humans. A framework for nis, S., Woller, A., Nunnemann, ceeding of the 7th Tübinger Perception alert macaque monkeys. Vision Res.
defining “early” visual processing. S., and Muhlau, M. (2008). Conference, Tübingen, 105. 36, 1225–1234.
Exp. Brain Res. 142, 139–150. Neural correlates of spontaneous Kornmeier, J., and Bach, M. (2004b). Lehmann, D., and Skrandies, W.
Fries, P., Roelfsema, P. R., Engel, A. K., percept switches in ambigu- Early neural activity in Necker-cube (1980). Reference-free iden-
Konig, P., and Singer, W. (1997). Syn- ous stimuli: an event-related reversal: evidence for low-level tification of components of
chronization of oscillatory responses functional magnetic resonance processing of a gestalt phe- checkerboard-evoked multichannel
in visual cortex correlates with imaging study. Eur. J. Neurosci. 28, nomenon. Psychophysiology 41, potential fields. Electroen-
perception in interocular rivalry. 2325–2332. 1–8. cephalogr. Clin. Neurophysiol. 48,
Proc. Natl. Acad. Sci. U.S.A. 94, Intaite, M., Koivisto, M., Ruksenas, Kornmeier, J., and Bach, M. (2005). 609–621.
12699–12704. O., and Revonsuo, A. (2010). The Necker cube – an ambigu- Leopold, D. A., and Logothetis, N.
Gale, A. G., and Findlay, J. M. (1983). Reversal negativity and bistable ous figure disambiguated in early K. (1999). Multistable phenom-
“Eye movement patterns in view- stimuli: attention, awareness, or visual processing. Vision Res. 45, ena: changing views in perception.
ing ambiguous figures,” in Eye something else? Brain Cogn. 74, 955–960. Trends Cogn. Sci. (Regul. Ed.) 3,
Movements and Psychological Func- 24–34. Kornmeier, J., and Bach, M. (2006). 254–264.
tions: International Views, eds R. İşoğlu-Alkaç, U., Basar-Eroglu, C., Ade- Bistable perception – along the Leopold, D. A., Wilke, M., Maier, A.,
Groner, C. Menz, D. F. Fischer, and moglu, A., Demiralp, T., Miener, M., processing chain from ambigu- and Logothetis, N. K. (2002). Sta-
R. A. Monty (Hillsdale: Erlbaum), and Stadler, M. (2000). Alpha activ- ous visual input to a stable per- ble perception of visually ambigu-
145–168. ity decreases during the perception cept. Int. J. Psychophysiol. 62, ous patterns. Nat. Neurosci. 5,
Girgus, J. J., Rock, I., and Egatz, R. of Necker cube reversals: an appli- 345–349. 605–609.
(1977). The effect of knowledge cation of wavelet transform. Biol. Kornmeier, J., and Bach, M. (2009). Liebert, R. M., and Burk, B. (1985).
of reversibility on the reversibility Cybern. 82, 313–320. Object perception: when our brain Voluntary control of reversible
of ambiguous figures. Percept. Psy- İşoğlu-Alkaç, Ü., and Strüber, D. is impressed but we do not notice it. figures. Percept. Mot. Skills 61,
chophys. 22, 550–556. (2006). Necker cube reversals during J. Vis. 9, 1–10. 1307–1310.

Logothetis, N. K. (1998). Single units state in illusory multistable motion Pitts, M. A., Martinez, A., Stalmas- on the reversal rate of differ-
and conscious vision. Philos. Trans. perception. J. Neural Transm. 112, ter, C., Nerger, J. L., and Hill- ent categories of reversible figures.
R. Soc. Lond. B Biol. Sci. 353, 565–576. yard, S. A. (2009). Neural gener- Perception 28, 1185–1196.
1801–1818. Murata, T., Matsui, N., Miyauchi, S., ators of ERPs linked with Necker Tong, F., and Engel, S. A. (2001).
Logothetis, N. K., Pauls, J., Augath, Kakita, Y., and Yanagida, T. (2003). cube reversals. Psychophysiology 46, Interocular rivalry revealed
M., Trinath, T., and Oeltermann, A. Discrete stochastic process underly- 694–702. in the human cortical blind-
(2001). Neurophysiological investi- ing perceptual rivalry. Neuroreport Pitts, M. A., Nerger, J. L., and Davis, spot representation. Nature 411,
gation of the basis of the fMRI signal. 14, 1347–1352. T. J. R. (2007). Electrophysiological 195–199.
Nature 412, 150–157. Nakatani, H., and van Leeuwen, correlates of perceptual reversals for Tong, F., Meng, M., and Blake, R. (2006).
Long, G. M., and Olszweski, A. D. C. (2006). Transient synchrony three different types of multistable Neural bases of binocular rivalry.
(1999). To reverse or not to reverse: of distant brain areas and images. J. Vis. 7, 1–14. Trends Cogn. Sci. (Regul. Ed.) 10,
when is an ambiguous figure not perceptual switching in ambigu- Pomerantz, J., and Kubovy, M. (1981). 502–511.
ambiguous? Am. J. Psychol. 112, ous figures. Biol. Cybern. 94, “Perceptual organization: an Tong, F., Nakayama, K., Vaughan, J. T.,
41–71. 445–457. overview,” in Perceptual Orga- and Kanwisher, N. (1998). Binocu-
Long, G. M., and Toppino, T. C. Necker, L. A. (1832). Observations on nization, eds M. Kubovy and J. lar rivalry and visual awareness in
(2004). Enduring interest in percep- some remarkable optical phaenom- Pomerantz (Hillsdale: Erlbaum), human extrastriate cortex. Neuron
tual ambiguity: alternating views of ena seen in Switzerland; and on an 423–456. 21, 753–759.
reversible figures. Psychol. Bull. 130, optical phaenomenon which occurs Pritchard, W. S. (1981). Psychophys- Tononi, G., Srinivasan, R., Russell, D.
748–768. on viewing a Figure of a crystal or iology of P300. Psychol. Bull. 89, P., and Edelman, G. M. (1998).
Long, G. M., Toppino, T. C., and geometrical solid. Philos. Mag. J. Sci. 506–540. Investigating neural correlates of
Mondin, G. W. (1992). Prime 1, 329–337. Rock, I., Gopnik, A., and Hall, S. conscious perception by frequency-
time: fatigue and set effects Noest, A. J., van Ee, R., Nijs, M. (1994a). Do young children reverse tagged neuromagnetic responses.
in the perception of reversible M., and van Wezel, R. J. (2007). ambiguous figures? Perception 23, Proc. Natl. Acad. Sci. U.S.A. 95,
figures. Percept. Psychophys. 52, Percept-choice sequences driven by 635–644. 3198–3203.
609–616. interrupted ambiguous stimuli: a Rock, I., Hall, S., and Davis, J. Toppino, T. C., and Long, G. M.
Luck, S. (2005). An Introduction to low-level neural model. J. Vis. (1994b). Why do ambiguous figures (1987). Selective adaptation with
the Event-Related Potential Tech- 7, 10. reverse? Acta Psychol. (Amst.) 87, reversible figures: don’t change that
nique. Cambridge, MA, USA: MIT O’Donnell, B. F., Hendler, T., and 33–59. channel. Percept. Psychophys. 42,
Press. Squires, N. K. (1988). Visual evoked Roeber, U., Widmann, A., Trujillo- 37–48.
Lumer, E. D., Friston, K. J., and Rees, potentials to illusory reversals of the Barreto, N. J., Herrmann, C. S., van Ee, R., van Dam, L. C., and
G. (1998). Neural correlates of per- Necker cube. Psychophysiology 25, O’Shea, R. P., and Schröger, E. Brouwer, G. J. (2005). Voluntary
ceptual rivalry in the human brain. 137–143. (2008). Early correlates of visual control and the dynamics of per-
Science 280, 1930–1934. Orbach, J., Ehrlich, D., and Heath, awareness in the human brain: time ceptual bi-stability. Vision Res. 45,
Maier, A., Wilke, M., Aura, C., Zhu, C., H. (1963). Reversibility of the and place from event-related brain 41–55.
Ye, F. Q., and Leopold, D. A. (2008). Necker cube: I. An examination of potentials. J. Vis. 8, 1–12. VanRullen, R., Reddy, L., and Koch,
Divergence of fMRI and neural sig- the concept of “satiation of ori- Schiller, P. V. (1933). Stroboskopische C. (2006). The continuous wagon
nals in V1 during perceptual sup- entation”. Percept. Mot. Skills 17, Alternativversuche. Psychol. Forsch. wheel illusion is associated with
pression in the awake monkey. Nat. 439–458. 17, 179–214. changes in electroencephalogram
Neurosci. 11, 1193–1200. Orbach, J., Zucker, E., and Olson, R. Sterzer, P., and Kleinschmidt, A. (2007). power at approximately 13 Hz. J.
Maier, A., Wilke, M., Logothetis, N. (1966). Reversibility of the Necker A neural basis for inference in per- Neurosci. 26, 502–507.
K., and Leopold, D. A. (2003). cube: VII: reversal rate as a func- ceptual ambiguity. Proc. Natl. Acad. Verleger, R., Jaskowski, P., and Wascher,
Perception of temporally interleaved tion of figure-on and figure-off Sci. U.S.A. 104, 323–328. E. (2005). Evidence for an integra-
ambiguous patterns. Curr. Biol. 13, durations. Percept. Mot. Skills 22, Sterzer, P., Kleinschmidt, A., and Rees, tive role of P3b in linking reaction
1076–1085. 615–618. G. (2009). The neural bases of mul- to perception. J. Psychophysiol. 19,
Mathes, B., Struber, D., Stadler, M. Palva, S., and Palva, J. M. (2007). tistable perception. Trends Cogn. Sci. 165–181.
A., and Basar-Eroglu, C. (2006). New vistas for alpha-frequency band (Regul. Ed.) 13, 310–318. Vickers, D. (1972). A cyclic decision
Voluntary control of Necker cube oscillations. Trends Neurosci. 30, Sterzer, P., and Rees, G. (2008). A neural model of perceptual alternation.
reversals modulates the EEG delta- 150–158. basis for percept stabilization in Perception 1, 31–48.
and gamma-band response. Neu- Pastukhov, A., and Braun, J. (2008). A binocular rivalry. J. Cogn. Neurosci. Washburn, M. F., Mallat, H., and Naylor,
rosci. Lett. 402, 145–149. short-term memory of multi-stable 20, 389–399. A. (1931). The influence of the size of
Meenan, J. P., and Miller, L. A. (1994). perception. J. Vis. 8, 1–14. Strüber, D., Basar-Eroglu, C., Hoff, E., an outline cube on the fluctuations
Perceptual flexibility after frontal or Pearson, J., and Brascamp, J. (2008). and Stadler, M. (2000). Reversal- of its perspective. Am. J. Psychol. 43,
temporal lobectomy. Neuropsycholo- Sensory memory for ambiguous rate dependent differences in the 484–489.
gia 32, 1145–1149. vision. Trends Cogn. Sci. (Regul. Ed.) EEG gamma-band during multi- Watanabe, M., Cheng, K., Murayama, Y.,
Michel, C. M., Murray, M. M., Lantz, 12, 334–341. stable visual perception. Int. J. Psy- Ueno, K., Asamizuya, T., Tanaka, K.,
G., Gonzalez, S., Spinelli, L., and Picton, T. W. (1992). The P300 chophysiol. 38, 243–252. and Logothetis, N. (2011). Attention
Grave de Peralta, R. (2004). EEG wave of the human event-related Strüber, D., Basar-Eroglu, C., Miener, but not awareness modulates the
source imaging. Clin. Neurophysiol. potential. J. Clin. Neurophysiol. 9, M., and Stadler, M. (2001). EEG BOLD signal in the human V1 dur-
115, 2195–2222. 456–479. gamma-band response during the ing binocular suppression. Science
Moreno-Bote, R., Rinzel, J., and Rubin, Pitts, M. A., Gavin, W. J., and Nerger, J. perception of Necker cube reversals. 334, 829–831.
N. (2007). Noise-induced alterna- L. (2008). Early top-down influences Vis. Cogn. 8, 609–621. Wertheimer, M. (1912). Experi-
tions in an attractor network model on bistable perception revealed by Strüber, D., and Herrmann, C. S. (2002). mentelle Studien über das Sehen
of perceptual bistability. J. Neuro- event-related potentials. Brain Cogn. MEG alpha activity decrease reflects von Bewegungen. Z. Psychol. 61,
physiol. 98, 1125–1139. 67, 11–24. destabilization of multistable per- 161–265.
Müller, T. J., Koenig, T., Wackermann, Pitts, M. A., Martinez, A., and Hill- cepts. Brain Res. Cogn. Brain Res. 14, Wilke, M., Logothetis, N. K., and
J., Kalus, P., Fallgatter, A., Strik, yard, S. A. (2010). When and where 370–382. Leopold, D. A. (2006). Local field
W., and Lehmann, D. (2005). Sub- is binocular rivalry resolved in the Strüber, D., and Stadler, M. (1999). potential reflects perceptual sup-
second changes of global brain visual cortex? J. Vis. 10, 1–11. Differences in top-down influences pression in monkey visual cortex.

Proc. Natl. Acad. Sci. U.S.A. 103, Zaretskaya, N., Thielscher, A., Logo- commercial or financial relationships perception changes but not the stimulus.
17507–17512. thetis, N. K., and Bartels, A. that could be construed as a potential Front. Hum. Neurosci. 6:51. doi:
Woerner, R., Bach, M., and Kornmeier, (2010). Disrupting parietal func- conflict of interest. 10.3389/fnhum.2012.00051
J. (2009). Ambiguous figures: differ- tion prolongs dominance durations Copyright © 2012 Kornmeier and Bach.
ent levels of perceptual adaptation. in binocular rivalry. Curr. Biol. 20, Received: 29 October 2011; paper pending This is an open-access article distributed
Perception 38, 138. 2106–2111. published: 27 November 2011; accepted: under the terms of the Creative Commons
Woo, M., Neider, J., and Davis, T. (1998). 26 February 2012; published online: 22 Attribution Non Commercial License,
OpenGL Programming Guide. The March 2012. which permits non-commercial use, dis-
Official Guide to Learning OpenGL, Conflict of Interest Statement: The Citation: Kornmeier J and Bach tribution, and reproduction in other
Version 1.1. Reading, MA: Addison- authors declare that the research was M (2012) Ambiguous figures – forums, provided the original authors and
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HUMAN NEUROSCIENCE published: 22 March 2012

Ambiguous ﬁgures – what happens in the brain when

Frontiers in Human Neuroscience www.frontiersin.org March 2012 | Volume 6 | Article 51 | 1

reversals as time reference to analyze the data in a certain interval

P300-like parietal positivity

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frontal electrode position (F4). The results were interpreted as

PROBLEMS OF THE MANUAL RESPONSE PARADIGM

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interruptions, Figure 3, left part), and another responsible for the

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(based on additivity of variances)

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even with binocular rivalry stimuli). This indicates gener-

Currently we regard the RP as an indicator of the visual sys-

(1) In the case of the Necker cube the stimulus information is

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Peak latency (ms) Location Stimuli

Stimulus onset = 0 Reaction time = 0

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