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brain research 1520 (2013) 107115

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Research Report

The fate of object memory traces under change


detection and change blindness
Niko A. Buscha,b,n
a

Institute of Medical Psychology, CharitUniversittsmedizin, Berlin, Germany


Berlin School of Mind and Brain, Humboldt Universitt zu, Berlin, Germany

art i cle i nfo

ab st rac t

Article history:

Observers often fail to detect substantial changes in a visual scene. This so-called change

Accepted 8 May 2013

blindness is often taken as evidence that visual representations are sparse and volatile.

Available online 15 May 2013

This notion rests on the assumption that the failure to detect a change implies that

Keywords:

representations of the changing objects are lost all together. However, recent evidence

EEG

suggests that under change blindness, object memory representations may be formed and

Long-term memory (LTM)

stored, but not retrieved. This study investigated the fate of object memory representations

Short-term memory (STM)

when changes go unnoticed. Participants were presented with scenes consisting of real

Attention

world objects, one of which changed on each trial, while recording event-related potentials

Inattentional blindness

(ERPs). Participants were rst asked to localize where the change had occurred. In an

Awareness

additional recognition task, participants then discriminated old objects, either from the

Consciousness

pre-change or the post-change scene, from entirely new objects. Neural traces of object
memories were studied by comparing ERPs for old and novel objects. Participants
performed poorly in the detection task and often failed to recognize objects from the
scene, especially pre-change objects. However, a robust old/novel effect was observed in
the ERP, even when participants were change blind and did not recognize the old object.
This implicit memory trace was found both for pre-change and post-change objects. These
ndings suggest that object memories are stored even under change blindness. Thus,
visual representations may not be as sparse and volatile as previously thought. Rather,
change blindness may point to a failure to retrieve and use these representations for
change detection.
& 2013 Elsevier B.V. All rights reserved.

1.

Introduction

Subjective visual experience suggests that we have access to


a rich and stable representation of a visual scene. However,
this intuition has been challenged by results obtained with
the change blindness paradigm (see Rensink, 2002; Simons

and Rensink, 2005, for reviews). This line of research has


demonstrated that observers are often unable to detect large
changes in a scene when the change occurs simultaneously
with a brief visual disruption, be it a saccade (McConkie and
Currie, 1996; Hayhoe et al., 1998), an eye blink (O'Regan et al.,
2000), a icker (Rensink et al., 1997), or a distracting stimulus

n
Correspondence address: Charit Universittsmedizin, Institut fr Medizinische Psychologie, Luisenstr. 57, 10117 Berlin, Germany.
Fax: +49 30 450 529 990.
E-mail address: niko.busch@charite.de

0006-8993/$ - see front matter & 2013 Elsevier B.V. All rights reserved.
http://dx.doi.org/10.1016/j.brainres.2013.05.014

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brain research 1520 (2013) 107115

(O'Regan et al., 1999). Under these conditions, a change cannot


be detected directly by detecting the motion or contrast
transient. Instead, change detection depends on preserving
and comparing object representations of pre-change and postchange objects. Change blindness obviously points to a limit in
our ability to represent, process, and maintain visual scenes.
But which of the numerous perceptual processes involved in
change detection is subject to this limit?
In order to detect a change, an observer has to process and
represent visual information through a number of stages.
First, the pre-change display must be encoded, and the
representation of the pre-change display must be maintained
in short-term or long-term memory, or both. Second, the
post-change display must be encoded and maintained. Third,
pre-change and post-change information must be compared,
based on either information currently active in short-term
memory or on information retrieved from long-term memory. Finally, the observer needs to make a decision as to
whether a change has or has not occurred.
Several authors have argued that change blindness results
from a limitation at the encoding stage (O'Regan, 1992;
Blackmore et al., 1995). Proponents of this view argue that the
capacity to represent visual information is strongly limited and
suggest that we represent no more than the semantic and
structural gist of a scene plus a small portion of the scene's
details, which are currently attended. A strong interpretation of
the change blindness phenomenon is that our every-day
intuition is wrongvisual representations may be, in fact, more
sparse than we believe. An alternative account argues that
change blindness can result from a failure to form a stable
representation in short-term or long-term memory. Thus, as
long as the scene is in view, visual representations may be rich,
in line with our phenomenology of a rich experience. However,
these representations are volatile and are easily overwritten
once the original scene disappears and the modied scene is
presented (Becker and Pashler, 2002; Landman et al., 2003; Beck
and Levin, 2003). Another cause for change blindness has been
demonstrated by Mitroff et al. (2004) who found that sometimes
observers are able to recognize both pre-change and postchange objects in a subsequent memory test even when they
were blind to the changes made to these objects, indicating that
change blindness can result as well from a failure to compare
existing representations of pre- and post-change information.
Recently, a number of behavioral studies have investigated the
role of long-term memory in change detection. Performance on
memory tasks, that require recognition of the changing objects,
is often better than change detection performance, suggesting
that object memory traces for the changing object are formed,
but are not used in the change detection task (Varakin and

Levin, 2006; Hollingworth, 2005; Hollingworth and Henderson,


2002; Beck et al., 2007). Other authors have demonstrated that
cues presented after the change has occured improve change
detection performance by facilitating retrieval of object information from long-term memory (Hollingworth, 2003; Beck and
van Lamsweerde, 2011). In sum, these studies have found that
change blindness can as well result from a failure to retrieve
object representations from long-term memory, in spite of
existing memory representations.
The present study investigated the fate of object representations in long-term memory under change blindness. To
my knowledge, this is the rst study combining a change
detection paradigm with a subsequent recognition memory
task that investigates neural markers of memory representations under change blindness. Participants were presented
with scenes consisting of familiar, nameable objects, one of
which changed on each trial, while recording event-related
potentials (ERPs). Participants were asked to detect changes
and report their location. In an additional recognition task,
participants were to discriminate old objects, either from the
pre-change or the post-change scene, from entirely new
objects. The analysis focused on the ERP-difference between
old and new items presented in the recognition task. If
change blindness was only due to a capacity limit during
encoding or maintenance, or due to overwriting, the visual
system should not be able to recognize the changing objects
as old under change blindness, and no ERP old/new effect
should ensue. If, however, change blindness can result from a
failure to retrieve object representations from memory during
detection, it might be possible to observe a memory effect (i.e.
old/new ERP differences) in the subsequent recognition task
even when the change detection task had failed. Indeed, I
found a robust ERP difference between old and novel objects
in spite of change blindness, indicating that traces of object
memories persisted even when changes to these objects were
not noticed. Importantly, this effect was found even when
both localization and recognition failed. This nding suggests
that in addition to limited capacity for encoding and overwriting, change blindness can also occur when the visual
system fails to retrieve object representations from memory.

2.

Results

2.1.

Behavioral results

Participants correctly localized the change on 48% of all trials


(see Table 1).

Table 1 Behavioral performance in the change localization task and the old/new recognition task. The table gives average
proportion correct (and standard error of the mean) relative to the number of pre-change and post-change trials.
Pre-change

Rec. correct
Rec. incorrect

Post-change

Loc. correct

Loc. incorrect

Loc. correct

Loc. incorrect

0.30 (0.02)
0.17 (0.01)

0.28 (0.02)
0.24 (0.01)

0.46 (0.03)
0.03 (0.01)

0.30 (0.02)
0.21 (0.02)

brain research 1520 (2013) 107115

109

Performance in the old/new recognition task was better when


the old item was a post-change item compared to pre-change
items F1;19 146; po0:001. On an average, old/new recognition was better on trials on which the change was also correctly
detected F1;19 624; po0:001. However, a signicant interaction F1;19 89; po0:0001 between probe-type (pre-change/postchange)  localization (correct/incorrect) indicated that the effect
of detection on recognition performance was less pronounced
for pre-change objects (t19 6:4; po0:001; two-tailed paired
samples t test) compared to post-change objects (t19
22:8; po0:001; Table 1). These results are displayed in Fig. 2.
The next analysis tested if participants were able to
recognize old probe objects better than chance on trials when
the change was not correctly detected. Indeed, recognition
performance was low, but above chance (i.e. signicantly
greater than 0.5) both for pre-change probe objects
(t19 7:9; po0:001; one-sample t test) and for post-change
objects t19 7:3; po0:001.

to 1200 ms. The analysis of the global dissimilarity index


revealed that the topographies of the early and late old/novel
effect were signicantly different po0:001.

2.2.

Fig. 2 Behavioral results: the gure shows proportion correct


(averaged across participants) in the recognition task, in which
participants had to discriminate old objects (either pre-change
or post-change) from novel objects. Error bars show standard
error of the mean. Participants were better at recognizing old
objects on trials when they also detected where the change
had happened. Additionally, they were better at recognizing
post-change objects. Although recognition performance was
poor on trials without correct change detection, recognition
was still signicantly above chance.

EEG results

In order to search for memory traces under change blindness,


I rst identied electrodes and time points at which the old/
new effect was most pronounced when subjects correctly
localized the change and recognized the old probe object
(Fig. 3). Under these conditions, ERPs were generally more
negative for novel probe objects than for old probe objects.
This effect was most pronounced at central electrodes from
200 to 700 ms and at right fronto-central electrodes from 800

Fig. 1 Schematic time course of a trial sequence. Each trial started with a display consisting of 9 nameable, familiar objects. On
each trial, one of these objects changed after a brief interruption of the display. Participants rst had to detect the change by
clicking on the location where the change had occurred. Following the detection response, two single objects were presented,
one of which was a novel object that had never been presented before in this experiment. The other object was either the prechange or the post-change object from the same trial. Participants reported which of the two objects had been presented before
on this trial. The ERP analysis focused on the difference between old and novel probe objects.

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brain research 1520 (2013) 107115

Fig. 3 Event related potentials (ERPs) for old and novel probes (averaged across participants) on trials with correct detection and
recognition. Left panel: difference between old and novel probe objects tested with a two-tailed t test. The old/novel effect was
tested at each sampling point and electrode, and the resulting t-values are color-coded. T-values corresponding to a p40:01 are
masked in grey. Each horizontal line represents one electrode, and electrodes are sorted according to their anteriorposterior
position. This analysis revealed temporally and spatially widespread old/new effects, starting at around 200 ms at frontal
electrodes, followed by a more parietal distribution from 500 to 700 ms, and a fronto-central distribution from 800 ms. Based on
this analysis, two spatio-temporal regions of interest were selected for further analyses of the old/novel effect. Middle panel: ERP
time courses averaged across electrodes within the central region of interest (see white markers in the topography), used for
testing the old/novel effect between 200 and 700 ms (grey shaded area). The topography shows the difference between old and
novel probe objects. Right panel: ERPs and difference topography for the old/novel effect between 800 and 1200 ms at right
fronto-central electrodes.

Subsequently, a three-factor ANOVA with the factors probe


novelty (old probe vs. novel probe), detection (correct vs.
incorrect) and recognition (correct vs. incorrect) was applied
to investigate whether old/novel effects in these two time
windows were present even when both change detection and
probe recognition failed.

2.2.1.

ERPs 200700 ms

For the early time window at central electrodes (Fig. 4, top


panel), this analysis conrmed that ERP amplitudes were
signicantly more negative for novel than for old probe
objects (probe novelty: F1;19 10:60; po0:001). The magnitude
of the old/novel effect was reduced when the change was not
detected (probe novelty  detection: F1;19 30:90; po0:001)
and when the old probe object was not recognized (probe
novelty  recognition: F1;19 20:80; po0:001). These interactions are not particularly surprising given that the selection
of time points and electrodes was based on the subset of
trials with successful change detection and probe recognition. The crucial question was whether this selection of
electrodes and time points would yield an old/novel effect
for other conditions, too. Thus, follow-up tests (two-tailed t
tests) were conducted that tested the old/novel effect for each
condition separately. These tests revealed that the old/novel
effect was present on trials with correct detection but no
recognition t19 2:58; p 0:018, on trials without detection but correct recognition t19 2:92; p 0:009, and even
on trials when both detection and recognition were incorrect

t19 3:11; p 0:006. In other words, even when the behavioral response indicated that the participant was not able to
discriminate an old from a novel probe object, the neural
response to old probe objects was nevertheless affected by
the prior presentation of this object.
A second ANOVA with factors probe novelty (old/novel) and
probe type (pre-change/post-change) compared the old/novel
effect under incorrect detection and recognition between prechange and post-change probes. This analysis conrmed the
main effect of probe novelty (F1;19 17:02; p 0:001; no interaction). Follow up tests conrmed that the old/novel difference was signicant for both pre-change probes (t19 3:74;
p 0:001; two-tailed t test) and post-change probes (t19
2:708; p 0:014).

2.2.2.

ERPs 8001200 ms

At longer latencies and frontocentral electrodes (Fig. 4, bottom panel), the polarity of the old/novel effect depended on
the recognition of the probe object, as indicated by a probe
novelty  recognition interaction F1;19 30:97; po0:001: when
recognition of the probe object was correct, ERPs for novel
probe objects were more negative-going than ERPs for old
objects. The opposite pattern was found when recognition
was incorrect, ERPs for novel probe objects were more
positive than ERPs for old objects. Moreover, the old/novel
effect was strongest when both detection and recognition
were correct, as indicated by a probe novelty  detection 
recognition interaction F1;19 4:68; p 0:043. Post hoc test

brain research 1520 (2013) 107115

111

Fig. 4 Event related potentials (ERPs) for old and novel probes averaged across participants. Upper panel: ERPs at central
channels (for channel locations see Fig. 3). The grey shaded area shows the old/novel effect in the time range from 200 to
700 ms. Statistically signicant old/novel effects were found in all conditions, even when detection and recognition had failed.
Bottom panel: ERPs at right fronto-central channels. The grey shaded area shows the old/novel effect in the time range from 800
to 1200 ms. The polarity of the late old/new effect depends on the success of recognition: ERPs are more positive for the object
which participants decided to report as old.

revealed that the old/novel effect was signicant when both


detection and recognition were correct (t19 6:15; po0:001;
two-tailed t test), when only recognition was correct t19
2:85; p 0:010, when only detection was correct t19
2:99; p 0:007, and even when neither detection nor recognition was correct t19 2:46; p 0:023. In other words, the
polarity of the ERP effect in this time range did not depend on
the novelty of the object per se, but on the participants'
decision: ERPs were more positive for the object which participants decided to report as old.
A further analysis tested this decision-related effect under
incorrect localization and recognition separately for prechange and post-change probes. An old/novel effect (i.e. more
negative ERP for novel items) was found for pre-change
probes (t19 2:35; p 0:30; two-tailed t test) as well as for
post-change probes (t19 2:55; p 0:019).

3.

Discussion

Change blindness is often taken as an evidence that visual


representations are sparse or fragile. A tacit assumption in
parts of the change blindness literature has been that the
explicit change detection is a direct reection of object
memory representations: if a change is detected, information
about the object had been stored in visual memory; if the
change goes undetected, then no object memory had been
stored. To test this assumption, the present study investigated the fate of object memory representations under
change blindness: when observers fail to detect that objects
change, do they store information about these objects in
memory despite their change blindness? Participants viewed
scenes consisting of nameable objects, one of which changed

to another object on each trial (see Fig. 1). Participants had to


detect where the change happened. Additionally, after the
detection task, one old object from the scene was presented
along with a novel object, and participants had to discriminate old from novel probe objects. To study memory for the
changing objects, behavioral performance in the old/novel
recognition task as well as event-related potentials (ERPs)
evoked by old and novel probes were analyzed. The results
argue against a direct correspondence of the success of
change detection and the storage of object memories.
Two main ndings stand out. First, performance in the
old/novel recognition task was strongly impaired when the
change was not detected. However, even under change
blindness, recognition performance was well above chance
(Fig. 2). This indicates that change blindness does not preclude the formation of memory representations of the changing objects, or their retrieval in a recognition memory task,
even though these object memories are not automatically
used for change detection.
Second, the analysis of ERPs in the present report extends
previous behavioral studies in an important aspect. Previous
studies have tested participants' explicit retrieval using the
behavioral reports in old/new recognition tests (e.g.
Hollingworth and Henderson, 2002; Varakin and Levin, 2006;
Beck et al., 2007; Beck and van Lamsweerde, 2011). The
analysis of event-related potentials in the present study
additionally allowed for testing neural markers of implicit
memories. This analysis revealed that even when participants could not consciously recognize objects from the
preceding scene as old, ERPs nonetheless showed a reliable
difference between old and novel items (Fig. 4), indicating an
implicit memory trace when explicit retrieval had failed.
Importantly, implicit memories for objects were found even

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brain research 1520 (2013) 107115

when participants were not aware of changes to these


objects. These implicit memory traces demonstrate that
change blindness can result not only from a failure to use
explicit object memories in the change detection task but also
from a failure to consciously retrieve the memory representation all together. It is possible that the information about a
changing object that is available under change blindness only
supports formation of this weak kind of memory. While this
indicates that the visual representation under change blindness is indeed impoverished compared to representations
under conscious change detection, the existence of even this
impoverished representation is not predicted by sparseness
and overwriting accounts of change blindness. In sum, the
ndings have important implications for our understanding
of how visual information processing is compromised under
change blindness: change blindness does not necessarily
point to a limit in our capacity to represent and maintain
perceptual information (O'Regan, 1992; Blackmore et al., 1995;
Becker et al., 2000; Becker and Pashler, 2002; Beck and Levin,
2003; Landman et al., 2003; Mitroff et al., 2004). Instead, visual
representations may be richer and more stable than previously suggested, in line with previous behavioral studies
demonstrating that change blindness can result from a postperceptual failure to retrieve or compare object memory
representations (Hollingworth and Henderson, 2002;
Hollingworth, 2005; Varakin and Levin, 2006; Beck et al., 2007).
This conclusion does not preclude other causes of change
blindness. In particular, another reason why change blindness can occur is that information about the pre-change
scene is overwritten by the post-change scene (Becker and
Pashler, 2002; Landman et al., 2003; Beck and Levin, 2003). For
example, Landman et al. (2003) investigated the availability of
object information by presenting partial-report cues a
centrally presented arrow that pointed to the location of
one item in the scene and asked participants to judge
whether the cued item had changed. A partial-report cue can
facilitate change detection because it strongly limits the
number of items that need to be retained and compared,
but only if a representation of the item is still available by the
time the cue is presented. The authors found that cues during
the blank interval between pre-change and post-change
scene strongly improved change detection. By contrast, cues
presented after the onset of the post-change scene did not
improve change detection. This nding indicates that the
representations of pre-change objects were available after the
offset of pre-change scenes, but were overwritten by the
onset of the post-change scene. In line with this nding,
Mitroff et al. (2004) investigated memory for pre-change and
post-change objects, and found that participants were signicantly better at recognizing post-change objects. Consistent with these studies, performance in the old/novel
recognition task was signicantly better for post-change
items (see Fig. 1). The old/novel ERP effect also tended to be
larger when the old item was a post-change item; however,
this interaction did not reach statistical signicance. Thus,
overwriting of pre-change information is a strong determinant for the change detection task and possibly for the
storage of longer-lasting object memories.
The present study focused on the old/novel ERP effect as
an electrophysiological correlate of object memory

representations during change detection, and the question


at hand was whether any such effect would show under
change blindness. ERP studies on recognition memory have
identied a number of old/new ERP effects and have related
them to different memory sub-systems (see reviews in Paller,
2004; Rugg and Curran, 2007; Mecklinger, 2000). While the
present study was not specically designed for such a
purpose, it may be worth speculating on the nature of the
observed old/new ERP effects. Correctly classied items generally elicit more positive-going ERPs than new items. Specically, an early mid-frontal old/new effect around 300500 ms
has been reported as a correlate of item familiarity. Old/new
effects in this time range are largely insensitive to depth-ofstudy manipulations and do not require conscious recognition of the item as old (Rugg et al., 1998). Others have reported
effects of implicit perceptual priming at similar latencies
when stimulus encoding was made difcult so as to promote
priming in the absence of conscious recognition (Paller et al.,
2003). Thus, there is a controversy as to whether the early
mid-frontal old/new effect reects familiarity or a form of
priming (see Voss and Paller, 2008). At longer latencies, a
parietal old/new effect around 400500 ms has been interpreted as a correlate of recollection, since its magnitude is
modulated according to whether the item is associated with
correct or incorrect source judgments and whether the item
is judged as remembered or known. In line with this
literature, the present study found an early, frontally distributed old/new effect (starting at around 200 ms) followed
by a more parietal effect (Fig. 3, left), even for items not
correctly recognized as old. While this constitutes an implicit
memory effect, it is unclear whether it specically reects an
implicit familiarity signal in the absence of conscious recollection (Jacoby, 1991) or a form of implicit priming (Schacter,
1990; Voss and Paller, 2008). Since the present study was not
designed for dissociating the contribution of different memory mechanisms, it is difcult to ascribe specic processes to
these ERP effects; this would have required additional manipulation of encoding conditions and behavioral measures of
priming or recognition (e.g. source memory or remember/
know judgments). Thus, the old/new effects reported here
may reect to some extent implicit perceptual priming,
familiarity-based recognition, as well as recollection. However, given the similarity to effects reported as correlates of
familiarity-based recognition, it is tentative to assume that
the early old/new effect found here reects a correlate of a
familiarity signal (Rugg et al., 1998). Several studies have also
reported a late right-frontal old/new effect with latencies of
500 ms and longer. The nature of this late positive complex
has remained somewhat elusive. Some authors have associated this effect to retrieval of contextual information
(Wilding and Rugg, 1996), to strategic, post-retrieval processes
(Mecklinger and Meinshausen, 1998), or to the participants'
decision bias (Windmann et al., 2002). Nessler et al. (2001)
found that the right LPC was larger for recognized old items
relative to entirely new items, but also for new lure items
when erroneously reported as old. Consistent with these
reports, the present study found a late (8001200 ms) rightfrontal ERP effect that was more positive-going for items
which participants reported as old, independent of the
accuracy of this report. Therefore, this effect seems to

brain research 1520 (2013) 107115

indicate the involvement of post-retrieval and possibly postdecisional monitoring and evaluation processes (Nessler
et al., 2001).
The present study found evidence for implicit object
memory traces under change blindness: a robust old/novel
ERP effect that was independent of change detection and
recognition of the changing object. The extant literature on
change blindness has been inconclusive as to whether implicit object representations under change blindness actually
exist. A number of behavioral studies have addressed this
question by asking whether observers can guess the location
of a change or whether their performance in a subsequent
task is modulated by the identity of the changing object, even
when they are not aware of any change. Some studies have
reported implicit effects on behavior in spite of change
blindness (Fernandez-Duque and Thornton, 2003, 2000;
Thornton and Fernandez-Duque, 2000; Laloyaux et al., 2006),
while others have argued that these effects can be explained
without implicit representations (Mitroff and Simons, 2002;
Mitroff et al., 2002). Likewise, EEG and fMRI studies have
provided only limited empirical support for implicit object
representations under change blindness, with some studies
nding neural signatures of implicit change detection
(Schankin and Wascher, 2008, 2007; Fernandez-Duque et al.,
2003), but with as many failures (Pourtois et al., 2006; Busch
et al., 2010b, 2010a; Koivisto and Revonsuo, 2003; Niedeggen
et al., 2001; Henderson and Orbach, 2006; Eimer and Mazza,
2005) or inconclusive ndings (Beck et al., 2001). It is important to note that most previous ERP studies that searched for
implicit or preserved object representations under change
blindness have looked at neural activity at the time the
change occurred (usually time-locked to the onset of the
changed display). These analyses have focused on the difference between trials without any change and change trials in
which subjects were blind to the change. However, comparing undetected changes to no-changes can only reveal implicit detection of change; this procedure is not suited for
revealing implicit object memory representations under
change blindness. Thus, this is the rst ERP study demonstrating electrophysiological correlates of implicit object
memory traces under change blindness by combining a
change detection paradigm with a subsequent recognition
memory task. This nding is in line with behavioral studies
that found preserved memory for objects even when a
change to that object had not been detected (Varakin and
Levin, 2006; Hollingworth, 2005; Hollingworth and Henderson,
2002; Beck et al., 2007; Beck and van Lamsweerde, 2011). It
should be noted that old items in the present study were, in
fact, always old and changed items. Thus, the old/novel ERP
effect shows that participants stored information about these
objects even under change blindness. However, this comparison does not reveal whether such memories include information about the object's status as changed or unchanged.
Future studies should compare old and changed items to
old, but unchanged items to clarify this question.
In sum, the ndings of the present study suggest that
visual representations may not be as sparse and volatile as
previously thought. Rather, change blindness may point to a
failure to retrieve and use these representations for change
detection. Thus, the change blindness phenomenon shows

113

that we see less than we believe, but we also see more than
we know.

4.

Experimental procedures

4.1.

Participants

Behavioral and EEG data were recorded from 22 participants


(mean age: 28.7; 13 women; 2 left-handed). All had normal or
corrected-to-normal visual acuity and were free of neurological or psychiatric disorders. All participants gave signed
informed consent. Three participants had to be excluded due
to excessive noise or artifacts in the EEG data. The experimental protocol was approved by the ethics committee of the
German Psychological Society (DGPS).

4.2.

Stimuli and procedure

The experiment was written in Matlab (Mathworks Inc.) using


the Psychophysics Toolbox (Pelli, 1997; Brainard, 1997). Participants were seated in a dark, electrically shielded, sound
attenuated chamber. Stimuli were presented on a calibrated
19 CRT monitor with a resolution of 1280  1024 pixels and
refresh rate 100 Hz, placed 53 cm from the participants eyes.
Head position was stabilized using a chin rest.
Images of nameable everyday objects were collected from
a commercially available database (Hemera Photo-Objects,
Vols. I and II). The stimulus set was restricted to images of
familiar objects that were easy to recognize. All images were
presented at a size of 2.31 visual angle on a grey background.
On each trial, a scene was presented consisting of 9
different objects and a central xation cross (see Fig. 1). The
objects were arranged on two concentric circles (invisible to
the participant) with eccentricities of 2.41 and 6.41. Each scene
was presented for 700 ms, briey interrupted by a blank
screen for 250 ms, and then presented a second time for
another 700 ms. On each trial, one of the objects was replaced
during the interruption. I therefore refer to the scene before
and after the interruption as the pre-change and postchange scene, respectively. Immediately after the offset of
the post-change scene, the objects were replaced by grey
placeholders. Participants were required to indicate where
the change had occurred by clicking on the corresponding
placeholder using the mouse. Following the localization
response, an empty screen was presented for 1000 ms followed by two consecutive probes. Each probe was a single
object presented centrally for 500 ms. The two probes were
separated by an empty screen for 1000 ms. One of the two
probes was a novel object that had not been presented
previously. The other object was an old object from the
previous scene, either (on 50% of all trials) the object from
the pre-change scene or the modied object from the postchange scene. I refer to these stimuli as novel probes, prechange probes, and post-change probes. After 1000 ms
following the offset of the second probe, a text appeared on
the screen which asked participants to indicate with a mouse
click which of the two objects had been part of the scene. The
response started the presentation of the next trial. Participants were instructed to respond as accurately as possible,

114

brain research 1520 (2013) 107115

but no instruction for response speed was given. Participants


were also instructed to always maintain central xation.
The location of the changes was randomized across trials.
The order across trials in which pre-change and post-change
probes appeared and the order of novel and old probe within
a trial was counterbalanced.

analysis avoids the problem of circularitythe use of the


same data set for selection and selective analysis
(Kriegeskorte et al., 2009).
To test whether the topographical distributions of the
early and late old/novel effect were signicantly different, I
computed their global dissimilarity index (see Murray et al.,
2008) as

4.3.

v
u
u1 n
earlyi
latei 2

GDIearly;late t 
n i 1 GFPearly GFPlate

EEG recording and analysis

EEG was recorded with a BioSemi Active-Two amplier


system from 64 Ag/AgCl electrodes arranged according to
the international 10-10 system and the two mastoids. To
monitor for eye movements and blinks, the horizontal and
vertical electro-oculogram (EOG) was recorded from three
additional channels. Two additional electrodes were used as
reference and ground. Signals were sampled at 512 Hz with
24-bit conversion resolution and a 0.16 Hz high-pass and a
100 Hz low-pass lter.
After recording, data were downsampled to 256 Hz, lowpass ltered at 30 Hz, converted to an averaged mastoid
reference, and epoched from 200 to 1200 ms time-locked
to onset of probe stimuli. Independent-component analysis
(ICA) was applied to the unaveraged raw data to correct for
eye blinks and eye movements using the MATLAB toolbox
EEGLAB and the extended infomax ICA algorithm (Delorme
and Makeig, 2004).
The aim of the EEG analysis was to identify electrophysiological correlates of memory traces in the proberecognition task. Memory traces were indicated by a difference in the ERP waveforms evoked by old (pre-change or
post-change) probe objects vs. novel probe objects. In particular, the objective of the study was to identify implicit
memory effects: old/novel differences under change blindness when the old object was not recognized. Implicit
memory effects if they exist at all were expected to be
small and difcult to detect. Thus, the rst step of the ERP
analysis consisted of the selection of relevant time ranges
and subsets of channels, at which explicit memory effects
were strongest. To this end, the data were rst inspected on
trials on which the change was correctly detected and the old
probe object (pre-change or post-change) was correctly discriminated from the novel probe object. This selection then
served as a spatio-temporal region of interest. Based on the
time course and the topography of the explicit old/novel
effect, two such regions of interest with a robust old/novel
effect were identied: from 200 to 700 ms at central electrodes (FC1, FCz, FC2, C1, Cz, C2, CP1, CPz, CP2) and from 800 to
1200 at right fronto-central electrodes (F4, FC2, FC4, FC6, Cz,
C2, C4, C6). ERPs were then quantied for statistical analysis
by averaging across time points and channels within these
regions of interest. For each of these two spatio-temporal
regions of interest, a three-factor ANOVA was computed with
the factors probe novelty (old probe vs. novel probe), detection
(correct vs. incorrect) and recognition (correct vs. incorrect).
Note that this selection of time points and channels
introduced a strong bias to yield a statistical effect of probe
novelty for trials with correct detection and recognition.
However, the critical question was whether any such effect
would be observed at the same time points and electrodes as
well for incorrect detection and recognition. Thus, this

where earlyi/latei are the average-referenced voltages at electrode i, and GFP refers to the global eld power, which is
equivalent to the standard deviation of the voltages across
electrodes (Murray et al., 2008). The GDI is an index of
conguration differences between two electric elds, independent of their strength. To test this measure of topographical difference, a permutation test was computed as
follows: (1) single-subject topographies were randomly reassigned to the two conditions, (2) the grand-average topography was recalculated, and (3) the GDI of this new grandaverage was calculated. This procedure was repeated 5 times.
This permutation tests the null hypothesis that all differences between the two topographies are not due to actual
condition effects, but rather due to random variance across
participants. A p-value was calculated by comparing the
empirical GDI to the distribution of GDI s obtained by the
permutation procedure.

Acknowledgments
This work was supported by the German Research Foundation (Grant # BU2400/1-1). I thank Irena oreva for help with
stimulus collection and data acquisition.

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