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Citation: Töllner, T., Conci, M., Rusch, T., & Müller, H. J. (2013). Selective manipulation of target identification demands in visual
search: The role of stimulus contrast in CDA activations. Journal of Vision, 13(3):23, 1–13, http://www.journalofvision.org/
content/13/3/23, doi:10.1167/13.3.23.
doi: 10 .116 7 /1 3. 3. 2 3 Received December 31, 2012; published August 2, 2013 ISSN 1534-7362 Ó 2013 ARVO
Journal of Vision (2013) 13(3):23, 1–13 Töllner, Conci, Rusch, & Müller 2
Support for this notion (Töllner, Rangelov, & field potentials from macaque, Reinhart et al. (2012)
Müller, 2012) stems primarily from the differential recently revealed a distributed neural network to
activation patterns of the posterior contralateral generate the CDA. In particular, they found especially
negativity (PCN)2 and the lateralized readiness poten- prefrontal areas (FEF, SEF) to exhibit close relation-
tial (LRP), two asymmetric waveforms of the event- ships, in terms of timing and activation levels, to
related potential (ERP) generally accepted to reflect memory-based behavioral performance, indicative of a
space-specific allocation of focal attention (e.g., Luck & contribution of these areas to the sustained surface
Hillyard, 1994; Eimer, 1996; Woodman & Luck, 1999) CDA measureable at posterior electrode sites. Vogel
and effector-specific production of motor responses and Machizawa (2004; see also Klaver, Talsma, Wijers,
(e.g., Coles, 1989; Eimer, 1998), respectively. In Heinze, & Mulder, 1999; Ikkai, McCollough, & Vogel,
particular, in a comparison of several task set 2010), in one of the most influential studies of CDA
conditions (i.e., localization vs. detection vs. discrimi- involvement in working memory tasks, revealed CDA
nation vs. compound) that required different stimulus- activation to increase systematically with the number of
response mappings to be performed on physically objects maintained in vSTM, with the amplitude
identical pop-out search stimuli, Töllner, Rangelov, reaching an asymptotic level when an individual
and Müller (2012) found no influence of task type reached her/his vSTM storage capacity limit. This
whatsoever on the elicitation of the PCN. Such task pattern demonstrates a direct relationship between
set–insensitive PCN responses can be taken as evidence neural CDA activity and the capacity of visual working
that any task-relevant target is selected automatically memory.
with the same speed whether or not the task set engaged Of importance for the present study, the CDA has
requires deeper stimulus analysis to reveal the target’s also been reported in a growing body of studies that
featural identity. By contrast, the stimulus-locked LRP required participants to initially search for the task-
was substantially delayed when the task set demanded relevant target among distracter objects (e.g., Mazza,
precise identification, as compared to simple localiza- Turatto, Umiltà, & Eimer, 2007; Emrich, Al-Aidroos,
tion, of the target. Hence, the RT cost attributable to Pratt, & Ferber, 2009; Carlisle, Arita, Pardo, &
the identification task demands must originate at some Woodman, 2011; Luria & Vogel, 2011; Woodman &
stage subsequent to focal-attentional selection but prior Arita, 2011; Wiegand, Finke, Müller, & Töllner, 2013;
to motor-response production. From the combined Anderson, Vogel, & Awh, 2013). For instance, Mazza
analysis of PCN and LRP responses, however, it et al. consistently observed CDA activations immedi-
remained unclear to what degree postselective percep- ately following PCN responses in visual search, but
tual versus response selection processes contributed to only when the task set required finer, in-depth analysis
this RT cost. of the selected item. In particular, by comparing the
sensory-driven event-related EEG lateralizations elicit-
ed in localization and discrimination tasks, Mazza et al.
Contralateral delay activity as an index of in- found the CDA to be triggered selectively for the latter
depth analysis of visual input task only; whereas, the PCN was virtually indistin-
guishable between the two types of tasks. To further
However, there is another electroencephalographic verify that the CDA elicited in such visual attention
brain response that promises to be informative about tasks is comparable to that observed in visual memory
the involvement of postselective perceptual processing tasks but functionally different from the PCN, Joli-
in the behavioral stimulus identification effect in a more coeur et al. (2008) examined PCN and CDA responses
fine-grained fashion, namely, the contralateral delay as a function of memory load in a visual choice
activity (CDA). This neural marker is a negative response task. Participants were required to encode
potential most prominent over parietal, occipital, and (i.e., identify) and respond to either one or two out of
temporal electrodes contralateral to the side of an four possible colored digits presented bilaterally in the
attended visual location or hemifield (e.g., Vogel & visual displays. Mirroring the pattern of Vogel and
Machizawa, 2004; Gazzaley & Nobre, 2011; Reinhart Machizawa (2004), the authors found that the number
et al., 2012; Wiegand et al., in press). In studies that of items to be processed had a significant impact on the
require the active encoding and maintenance of activation and duration of the CDA but not the PCN
information in visual short-term memory (vSTM), this component. This electro-cortical dissociation was taken
potential shows a sustained activation during the as evidence for two functionally distinct cognitive
retention interval (which is why one pool of researchers functions underlying the PCN and CDA. Together with
prefer the term Sustained Posterior Contralateral recent PCN reports (e.g., Brisson & Jolicoeur, 2007;
Negativity; see Jolicoeur, Brisson, & Robitaille, 2008, Töllner, Zehetleitner, Gramann, & Müller, 2010, 2011;
for a detailed review). Combining ERP recordings from Conci, Töllner, Leszczynski, & Müller, 2011; Mc-
macaque and humans, together with intracranial local Donald, Green, Jannati, & Di Lollo, in press), these
Journal of Vision (2013) 13(3):23, 1–13 Töllner, Conci, Rusch, & Müller 3
results provide additional support for the view that the previous findings (e.g., Sagi & Julesz, 1985b; Mazza et
PCN reflects the engagement of visuo-spatial attention al., 2007; Töllner, Rangelov, & Müller, 2012). Second,
on laterally presented target objects. By contrast, the because we used the identical physical stimulus
CDA is specifically related to the location-specific material for both the localization and identification
encoding and/or in-depth analysis of visual input in tasks, we expected the PCN timing, indexing focal-
vSTM, thus reflecting similar cognitive operations to attentional selection of the target, to be unaffected by
those involved in classical working memory tasks. the respectively engaged task set and stimulus-
background orientation contrast—as the latter should
affect only the salience of the response-defining
Logic of the present study feature. By contrast, assuming the recruitment of an
additional recurrent process in which participants are
The present study was designed to investigate a required to extract target identity information, we
direct implication of the functional distinction between expected a CDA wave to be manifest when the task
the PCN and CDA responses as drawn by Jolicoeur demanded target identification; whereas, the CDA was
and colleagues (2008, see also Mazza et al., 2007), expected to be reduced, or entirely absent, when the
namely, that the CDA indexes postselective in-depth task required simple target localization. Third, for the
analysis of information at already attended stimulus identification task, we expected a gradual increase in
locations. On this assumption, CDA activations should the CDA response with decreasing stimulus-back-
increase gradually as more, relative to less, recurrent ground orientation contrast, indicative of increasing
processing needs to be engaged in order to extract the post-selective processing demands for extracting the
object identity information required for subsequent target’s exact orientation feature value (vertical vs.
response selection. To systematically examine this horizontal).
prediction, we combined mental chronometry data with
event-related EEG lateralizations in two visual search
tasks that differed in the depth of signal processing Methods
required for determining the correct response. The first
compound-search task required participants to initially
search for a pop-out target (defined variably across Participants
trials by the color red or green) before they could
decide upon the appropriate motor response (defined Fifteen volunteers (five female) took part in this study
by the target’s vertical or horizontal orientation). Note for either monetary payment or course credit. Two
that this compound task logic (e.g., Kingstone, 1992; participants had to be excluded from the analyses due to
Pollmann, Weidner, Müller, Maertens, & von Cramon, excessive horizontal eye movement artifacts. The ages of
2006; Töllner, Gramann, Müller, Kiss, & Eimer, 2008) the remaining 13 participants ranged from 20 to 30
elegantly dissociates target-defining features, which (median 25) years. All had normal or corrected-to-
determine PCN responses, from response-critical fea- normal vision, and none reported a history of neuro-
tures, which presumably influence CDA activations. logical disorders. Participants gave their written in-
Accordingly, this design permitted us to selectively formed consent prior to performing the experiment. The
manipulate the ease/difficulty with which participants experimental procedure was approved by the ethics
could extract the response-critical target orientation, committee of the Department of Psychology, University
namely, by introducing three different stimulus-back- of Munich, in accordance with the Code of Ethics of the
ground contrast levels created by Gaussian ‘‘blurring’’ World Medical Association (Declaration of Helsinki).
of orientation information (see Figure 1); henceforth,
this variable will be referred to as stimulus-background
orientation contrast. Exactly the same physical stimu- Stimuli and study design
lus material was additionally employed in a second
control task requiring participants to make simple Except for the colors and contrast modulations used
localization (left vs. right position), rather than precise for the present stimuli (see below), the search displays
identification (see above), responses. were identical to those of Töllner et al. (2008), Töllner
In both experiments, we analyzed RT performance et al. (2010), and Töllner, Rangelov, and Müller (2012).
together with both PCN and CDA responses to That is, search displays consisted of eight colored shape
dissociate preattentive from postselective perceptual stimuli presented on a gray background and arranged
processes as a function of task set and stimulus- in a circular formation (radius: 4.68 of visual angle)
background orientation contrast. First, we expected to around a central white fixation point (see Figure 1). In
find slower behavioral response latencies for identifi- each trial, one of the six lateral locations contained a
cation relative to localization responses, replicating feature singleton target, equally likely defined by the
Journal of Vision (2013) 13(3):23, 1–13 Töllner, Conci, Rusch, & Müller 4
Figure 1. Examples of search displays used in the present study. The color (red vs. green), orientation (vertical vs. horizontal),
stimulus-background contrast (high, intermediate, low) as well as position (at one of the six lateral locations) of the target varied
randomly on a trial-by-trial basis. Participants were required to give a speeded forced-choice response indicating either the featural
identity (vertical vs. horizontal orientation; Experiment 1) or location (left vs. right position; Experiment 2) of the color-defined
feature singleton.
color red (i.e., a single red circle/disk; CIE .622, .344, 6) presented on a 17-in. CRT screen (85-Hz refresh rate),
or green (i.e., a green circle/disk; CIE .293, .589, 6), mounted at a viewing distance of approximately 75 cm.
together with seven homogenous distracter items (i.e., Each experiment consisted of nine blocks of 72 trials
blue circles/disks; CIE .144, .066, 6). Each stimulus each, resulting in a total of 1,296 trials. In both
outline contained a grating composed of three gray experiments, target-defining features and stimulus-
bars (0.48 · 2.48) separated by two gaps (0.38 · 2.48), background orientation contrast varied randomly on a
which were randomly oriented either vertically or trial-by-trial basis throughout all blocks. A trial started
horizontally. Three different levels of stimulus-back- with the presentation of a white central fixation cross
ground orientation contrast (high, intermediate, low) for 500 ms, which was immediately followed by the
were realized by applying Gaussian (blurring) filters search array shown for 100 ms. Trials were terminated
with radii of 100, 110, and 128 pixels, respectively, to by the participant’s response or after a maximum time
the stimuli. These transformations particularly affected limit of 1000 ms. Response times were measured
the conspicuousness of the bar’s contours without, relative to the onset of the search displays. In case of a
however, modulating the salience of the respective response error or if no response was given within the
colors. Within a given trial, both target and distracter maximum time allowed, a red minus sign (i.e., ‘‘–’’) was
items were presented with identical contrast levels. centrally presented for 1000 ms, signaling erroneous
Participants were instructed to maintain central eye behavior. The subsequent intertrial interval displayed a
fixation throughout the experimental trials and to make white central fixation cross for a randomly chosen
a two-alternative forced-choice response as quickly and duration of either 950, 1000, or 1050 ms. Prior to the
accurately as possible. Participants were required to start of each individual experiment, at least one block
identify the target’s orientation (vertical vs. horizontal) of practice was administered in order for participants
in one experiment and to localize the target’s posi- to become familiarized with the stimulus material and
tioning relative to the midline of the search display (left the required stimulus-response mapping. After each
vs. right) in the other (control) experiment by pressing block, participants received summary performance
the respectively assigned mouse buttons with their left statistics (mean error rate and RT) as feedback
or right thumb. information.
The two experiments (identification/localization
task) were controlled by an IBM-PC compatible
computer using Matlab routines and Psychophysics EEG recording and data analysis
Toolbox extensions (Brainard, 1997; Pelli, 1997) and
were performed consecutively in random order in a The electroencephalogram was continuously sam-
dimly lit, sound-attenuated, and electrically shielded pled at 1 KHz using Ag/AgCl active electrodes (acti-
experimental booth (IAC). Search displays were CAP system, Brain Products, Munich) from 64 scalp
Journal of Vision (2013) 13(3):23, 1–13 Töllner, Conci, Rusch, & Müller 5
Figure 3. Electrophysiological results. (A) Grand-average target-synchronized contralateral versus ipsilateral ERP waves in the 500-ms
interval following stimulus onset, relative to a 200-ms prestimulus baseline, at electrodes PO7/PO8. (B) Topographical scalp
distribution maps computed by mirroring the contralateral-minus-ipsilateral difference waves (to obtain symmetrical voltage values
for both hemispheres based on spherical spline interpolation) for the successive time intervals of 180–240 ms, 240–300 ms, 300–360
ms, and 360–420 ms poststimulus. (C) The corresponding PCN difference waves obtained by subtracting ipsilateral from contralateral
activity as a function of stimulus-background contrast (high, middle, low), separately for the localization (left panel; Experiment 2) and
identification tasks (right panel, Experiment 1).
feature contrast increases. Exactly this pattern was expectancy (e.g., Töllner, Strobach, Schubert, &
established by Töllner, Zehetleitner, Gramann, and Müller, 2012): The PCN signal has been found to be
Müller (2011). amplified for predictable (vs. nonpredictable) targets as
Although attention-guiding saliency representations well as for cross-trial repetitions (vs. changes) of the
are derived largely in bottom-up fashion, they can also target-defining dimension. According to salience sum-
be biased by internal system settings relating to mation models (e.g., Müller et al., 1995, 2010), these
intertrial history (e.g., Töllner et al., 2008) or target boosted activations originate from a ‘‘top-down’’
Journal of Vision (2013) 13(3):23, 1–13 Töllner, Conci, Rusch, & Müller 8
condition), or partially identical/different (i.e., ‘‘inter- of all items within the pre-cued hemifield had to be
mediate’’ condition) within the task-relevant visual precisely identified and memorized in this working
hemifield (for further methodological details, see Luria memory task, as participants were asked to respond
& Vogel, 2011). Using this procedure, Luria and Vogel whether the memory and test arrays were identical or
found CDA responses to be enhanced as the search not. Using this design, however, Ikkai et al. observed
difficulty increased, which they interpreted in terms of a exactly the opposite pattern: Reduction of the stimulus
stronger reliance on working memory representations contrast had no influence on the CDA (despite
with difficult relative to easy searches. However, there significant behavioral effects) while yielding signifi-
is one fundamental difference between Luria and cantly attenuated PCN amplitudes. Based on this
Vogel’s and the present study: The difficulty manipu- pattern, the authors reasoned that the CDA might be
lation employed by Luria and Vogel did substantially immune to any sensory factors, being driven exclusively
affect the salience of the target relative to the items in by the number of objects represented in vSTM.
its surround, thereby changing the nature of the task In the light of these and other results (Vogel et al.,
from an efficient, spatially parallel pop-out search 2005), an alternative explanation for the enhanced
(‘‘easy’’ condition) to an inefficient, serial search CDA amplitudes with lower stimulus-background
(‘‘difficult’’ condition). It should be noted that this contrast observed in the present study may be that this
manipulation does, in the first instance, influence the pattern simply results from poor filtering of nearby
elicitation of the preceding PCN wave (which is distracters when the target contrast is low.5 Despite
indicative of visuo-spatial selection) as is evident in these, at first glance, close similarities between Ikkai et
Luria and Vogel’s experiments 2 and 3. Thus, as al. (2010) and our study, it seems rather improbable
suggested by these authors, the enhanced CDA that this alternative explanation can account for the
triggered with serial relative to pop-out searches may current data set. Note that in the study of Ikkai et al.,
be attributable to the increased demands of matching there were always multiple task-relevant items defined
candidate target items (i.e., distracters from the search by the same color features that subsequently also
array) against the target template. Because there are determined the required response, namely, to indicate
likely several candidate targets in serial search as whether or not the test array was the same as the
compared to just one in pop-out search, there would be memory array. In the current study, by contrast, there
iterative matching operations in this condition, for was always only one task-relevant pop-out item defined
which candidate items would need to be buffered in by a feature (i.e., red vs. green color) entirely unrelated
vSTM (owing to the brief display duration). to that determining the subsequent response (i.e.,
At marked variance with Luria and Vogel’s (2011) vertical vs. horizontal orientation). Accordingly, be-
design, the difficulty manipulation used in the present cause our contrast manipulation affected selectively the
study did not change the target’s salience relative to its response-critical target attribute but not the feature
surround at all, thus ensuring the pop-out nature of the that singled out the target from its distracter surround
search task for all stimulus-background contrast (with displays disappearing after 100 ms), it would
conditions. In agreement with this, the present PCN appear rather unlikely that the current CDA results
responses were wholly unaffected by the difficulty may be driven by differential, contrast-dependent
manipulation—in marked contrast to the CDA wave, filtering of nearby distracters. The view that the current
which increased gradually as a function of stimulus- contrast manipulation did not affect the filtering of
background contrast. Given that there was always only distracters is further supported by the fact that the
a single, highly salient ‘‘pop-out’’ item in our search PCN (which indexes spatial filtering) was insensitive to
displays, it is unlikely that this CDA amplification was stimulus-background contrast. Ikkai et al., by contrast,
due to increased demands for temporary storage of found decreased PCN activations for reduced stimulus
candidate targets (i.e., distracters from the search contrast (see above), in line with the notion that the
array) and more frequent template matching operations PCN reflects the target’s saliency signal (see, e.g.,
for low-contrast, relative to high-contrast, pop-out Töllner, Zehetleitner, Gramann, & Müller, 2011).
searches. In sum, the current study revealed stimulus-
Finally, our study design appears also linked to background contrast as a critical determinant of the
another recent working memory study (Ikkai et al., CDA in visual search scenarios that require partic-
2010), which, likewise, manipulated the contrast of the ipants to search for and identify a single, highly
items to be stored in vSTM. In more detail, the arrays salient ‘‘pop-out’’ target defined by a feature that
employed by Ikkai et al. were either of high or low differs from the response-critical target attribute (i.e.,
contrast and consisted of either two or four differently compound search). Based on the current data set,
colored items, with the task-relevant visual hemifield however, it remains an open issue whether or not this
being pre-cued by an arrow (pointing to the left or factor of stimulus contrast is additive to or interacts
right) on a trial-by-trial basis. Importantly, the colors with an individual’s memory storage capacity in
Journal of Vision (2013) 13(3):23, 1–13 Töllner, Conci, Rusch, & Müller 10
Figure 4. Schematic of the suggested approach for temporally disentangling four functionally distinct sensory- and motor-related
substages of the human information-processing stream: (1) the PCN latency indexes the time required for focal-attentional target
selection in visual space; (2) the CDA (minus PCN) timing indexes the time required for extracting detailed stimulus identity
information from vSTM; (3) the stimulus-locked LRP (minus CDA) timing indexes the time required for selecting the appropriate motor
response; (4) finally, the response-locked LRP timing reflects the time required by motor-response production processes.
determining the CDA response. Future studies may demands of this postselective extraction process are
systematically address this question in order to derivable from the CDA timing, which indicates the
establish whether the revealed pattern generalizes point in time at which the target’s identity is revealed
across paradigms from visual search to working from vSTM representations. Next, response selection
memory tasks and vice versa. processes (indexed by the stimulus-locked LRP) deter-
mine the appropriate motor response required by the
respectively extracted stimulus attribute, as defined by a
Quartering RTs via event-related lateralizations previously established task set (i.e., stimulus-response
of the EEG mapping rule). Finally, response production processes
(indexed by the response-locked LRP) generate and
The characteristics of the CDA as revealed by the execute the motor response thus selected. It should be
current study have fundamental implications with noted, however, that this approach of quartering
regard to the precision of dissociating RT effects on the reaction times is only feasible when the shape of the
basis of event-related EEG lateralizations in visual CDA (contralateral minus ipsilateral difference) wave
search. When analyzed together with the PCN, the is elicited monophasic rather than being sustained in
stimulus-locked LRP, and the response-locked LRP, nature so as to allow the extraction and usage of its
the temporal processing demands of the following four temporal information. Accordingly, an adequate sig-
functionally distinct substages of the human informa- nal-to-noise ratio (see, for instance, Luck, 2005, for a
tion-processing stream can be electro-cortically disso- detailed overview of the ‘‘standard’’ rules) in combi-
ciated: (a) preattentive perception, (b) postselective nation with precise target identification demands (see
perception, (c) motor-response selection, and (d) above) seem to be an inevitable prerequisite for this
motor-response production (see Figure 4). approach being employable.
In particular, preattentive perceptual processes
encode the whole stimulus array initially in parallel and
determine when and where the attentional spotlight will
be engaged. Accordingly, the temporal information of Conclusions
the PCN, which reflects the point in time when the
target becomes focally selected in visual space (e.g., In conclusion, the present findings advance our
Luck & Hillyard, 1994; Eimer, 1996; Töllner, Zehet- understanding of the processes underlying the CDA
leitner, Gramann, & Müller, 2011; Töllner, Müller, & wave in studies of visual search. Here, we could directly
Zehetleitner, 2012), can be used to estimate the times demonstrate that this neural marker does not simply,
demanded by feedforward visual processing6 to deter- or only, reflect the amount of information to be stored
mine the presence and location of the target. Post- and maintained in vSTM (e.g., Vogel & Machizawa,
selective perceptual processes extract then, via 2004, Jolicoeur et al., 2008; Wiegand et al., in press),
recurrent feedback connections, the response-critical but also the degree of postselective perceptual pro-
stimulus information at the respectively attended cessing recruited for extracting detailed object identity
location. As elaborated above, the temporal processing information from vSTM representations. However,
Journal of Vision (2013) 13(3):23, 1–13 Töllner, Conci, Rusch, & Müller 11
whether this factor of stimulus contrast is additive to or example: the left thumb) and, thus, not cancelled out by
interacts with memory storage capacity in the elicita- the averaging process. Notably, the manifestation of
tion of the CDA remains an open question to be the LRP immediately following the PCN provides
addressed in future studies. Nonetheless, our data further support for the notion that master map activity
clearly demonstrate that systematic analysis of the (as indexed by the PCN) is sufficient to indicate the
CDA together with multiple event-related EEG later- target’s location, with the location information being
alizations—such as the PCN, stimulus-locked LRP, transferred directly to motor response–related stages
and response-locked LRP—can help to improve the (see Töllner, Rangelov, & Müller, 2012).
precision of electro-cortically dissociating RT effects in 4
Prior to the PCN response, the data suggest an
visual search tasks. additional posterior contralateral positivity (Ppc; as
Keywords: attention, stimulus contrast, recurrent recently observed by Jannati, Gaspar, & McDonald, in
processing, electroencephalography press) in the localization task, which appears to be most
pronounced for high-contrast targets and gradually
decreasing for intermediate- and low-contrast targets.
However, this observation could not be substantiated
Acknowledgments statistically as indicated by the absence of a significant
effect, F(2, 24) ¼ 1.47, p . 0.25, of stimulus-
We would like to thank Edward Vogel, John background contrast on Ppc amplitudes. (We thank
McDonald, and one anonymous reviewer for their John McDonald for suggesting this analysis.)
5
helpful comments on an earlier version of this We thank Edward Vogel for suggesting this
manuscript. This research was supported by German alternative explanation.
6
Research Foundation (DFG) grant (Nr. CO 1002/1-1). This is not to say that ‘‘feedforward processing’’ in
the present sense equals the very first sweep of visual
Commercial relationships: none. processing through the brain. Concerning this issue,
Corresponding author: Thomas Töllner. there is also a debate as to the exact mechanisms that
Email: ttoellner@gmail.com. drive the PCN: While the currently dominant view
Address: Department of Experimental Psychology, holds that the PCN reflects re-entrant processes (e.g.,
Ludwig-Maximilians-Universität München, Munich, McDonald et al., in press), others have recently argued
Germany. in favor of a feedforward-driven process (e.g., Töllner,
Rangelov, & Müller, 2012).
Footnotes
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