Orienting Attention to Locations in

Internal Representations

Ivan C. Griffin and Anna C. Nobre

Abstract
& Three experiments investigated whether it is possible to same location in the array. Replication of the behavioral costs
orient selective spatial attention to internal representations and benefits of pre-cues and retro-cues in this experiment
held in working memory in a similar fashion to orienting to ruled out changes in response criteria as the only explanation
perceptual stimuli. In the first experiment, subjects were for the effects. The third experiment used event-related
either cued to orient to a spatial location before a stimulus potentials (ERPs) to compare the neural processes involved in
array was presented (pre-cue), cued to orient to a spatial orienting attention to a spatial location in an external versus
location in working memory after the array was presented an internal spatial representation. In this task, subjects
(retro-cue), or given no cueing information (neutral cue). The responded according to whether a central probe stimulus
stimulus array consisted of four differently colored crosses, occurred at the cued location in the array. There were both
one in each quadrant. At the end of a trial, a colored cross similarities and differences between ERPs to spatial cues
(probe) was presented centrally, and subjects responded toward a perception versus an internal spatial representa-
according to whether it had occurred in the array. There were tion. Lateralized early posterior and later frontal negativities
equivalent patterns of behavioral costs and benefits of cueing were observed for both pre- and retro-cues. Retro-cues also
for both pre-cues and retro-cues. A follow-up experiment used showed additional neural processes to be involved in
a peripheral probe stimulus requiring a decision about orienting to an internal representation, including early effects
whether its color matched that of the item presented at the over frontal electrodes. &

INTRODUCTION
slett, 1997; Guariglia, Padovani, Pantano, & Pizzamiglio,
To interact successfully in the spatial world we need 1993). Other studies have demonstrated a link between
both to be able to attend selectively to items in the extra- actual perceptual events and internal ‘‘images,’’ both in
personal world and to those maintained in mind as terms of the influence of ‘‘holding images in mind’’ on
mental representations. Although spatial orienting to- behavioral performance (Downing, 2000; Pashler & Shiu,
ward perceptual events has been intensively investigated, 1999; Farah, 1985; Ishai & Sagi, 1995), and similar regions
spatial orienting to internal representations has remained of neural activation (Kosslyn, Ganis, & Thompson, 2001;
relatively unexplored. Here we investigated the behavior- Ishai, Ungerleider, & Haxby, 2000; O’Craven & Kanw-
al and neural processes involved in orienting spatial isher, 2000). The Downing (2000) study showed that the
attention selectively to information held in internal spatial contents of WM can guide selective attention. Unsurpris-
representations in working memory (WM), and com- ingly, influential theoretical models posit a close link
pared them directly to orienting spatial attention to between WM and the allocation of selective attention
perceptual information. (e.g., Desimone & Duncan, 1995; Baddeley, 1993).
The relationship between attentional operations in Convergent findings from different methodologies
perception and WM has been highlighted by several lines have shown that spatial attention and spatial WM are
of research. Graziano, Hu, and Gross (1997) demon- subserved by partially overlapping large-scale distributed
strated that internal representations can be used to guide systems of frontal and parietal areas (Pollmann & von
action toward previously seen perceptual stimuli that are Cramon, 2000; Awh et al., 1999; LaBar, Gitelman, Parrish,
no longer present. Internal spatial representations can & Mesulam, 1999; McCarthy, 1995). This has led to the
also be impaired in certain cases of neglect (Bisiach & intuitively appealing suggestion that there may be simi-
Luzzatti, 1978), sometimes selectively (Ortigue et al., larities between the process of directing spatial attention
2001; Beschin, Cocchini, Della Sala, & Logie, 1997; Co- to a location, and the process of holding a spatial location
in WM (Awh & Jonides, 2001). Supporting evidence for
this view comes from findings that stimuli appearing at
memorized locations, and attended targets in spatial
University of Oxford attention tasks exhibit a similar pattern of activation and

D 2003 Massachusetts Institute of Technology Journal of Cognitive Neuroscience 15:8, pp. 1176–1194
modulation in posterior extrastriate visual areas in both The present experiments used cues presented after a
brain imaging (Awh et al., 1999) and event-related poten- stimulus array to orient attention selectively to locations
tial (ERP) (Awh, Anllo Vento, & Hillyard, 2000) studies. in internal representations. The cues were presented
Sperling (1960) developed an experimental paradigm within an interval where WM, and not iconic memory,
that enables attention to be directed toward contents of operates. Unlike previous studies, the target stimuli were
mental representations. He used cues to orient partic- not defined before task performance, and the cues there-
ipants to selective aspects of iconic traces of briefly fore did not guide response decisions. The cues were also
presented visual arrays. The present experiments build centrally presented and informative.
on this by using cues to orient attention to locations of In Experiment 1 (see Fig. 1A), subjects were given
arrays held in WM, after the iconic trace period. Spatially spatially informative cues (80% validity) to orient to a
informative cues were presented 1.5–2.5 sec before (pre- spatial location either before a stimulus array was pre-
cue) or after (retro-cue) a stimulus array, which oriented sented (pre-cue), or after the array was presented (retro-
subjects attention to either a perceptual location (pre- cue), or given no cueing information (neutral cue). Sub-
cue), or a specific location in the array that existed only as jects then decided whether a probe stimulus had been
an internal representation in WM (retro-cue). We were present in the array (50% probability). When the probe
firstly interested in whether it was possible to orient stimulus had been present in the array, the informative
attention to selective locations of internal representations pre-cues and retro-cues predicted its correct location 80%
of previously presented stimuli held in WM; and, if so, of the time. Analysis of behavioral data revealed that it is
whether it can provide similar behavioral advantages to possible and advantageous to orient attention selectively
those seen with spatially informative cues presented to locations in internal representations. Experiment 2
before the appearance of a stimulus array (Posner, (see Fig. 1B) followed up from these findings to ensure
1980). Secondly, we wished to compare the neural pro- that the behavioral effects did not reflect solely differ-
cess of orienting attention to internal mental represen- ences in response criteria between valid and invalid trials.
tations to the better known literature relating to the The task and cueing conditions were equivalent to those
orienting of visual attention to upcoming perceptual in Experiment 1, except that a peripheral probe stimulus
stimuli (Corbetta, Kincade, Ollinger, McAvoy, & Shulman, prompted a decision as to whether its color matched that
2000; Hopf & Mangun, 2000; Hopfinger, Buonocore, of the item at the same location in the array (50%
& Mangun, 2000; Nobre, Sebestyen, & Miniussi, 2000; probability). This manipulation eliminated uncertainty
Kastner, Pinsk, De Weerd, Desimone, & Ungerleider, about which location subjects should use in the decision
1999; Yamaguchi, Tsuchiya, & Kobayashi, 1994; Harter, process. In Experiment 3 (see Fig. 1C), ERPs were re-
Miller, Price, LaLonde, & Keyes, 1989), to see whether the corded while subjects performed a simplified version of
same, different, or partially overlapping control systems the tasks with only pre-cue and retro-cue conditions. The
are involved. probe stimuli were presented centrally, and subjects
Many studies have used cues presented after a stimulus responded according to whether the probe stimulus
array to study cognitive processes. For example, Kinchla, occurred at the cued location (50% probability). Analysis
Chen, and Evert (1995) cued peripheral locations to of ERPs to the identical cueing stimuli occurring before
inform subjects which element in a briefly presented versus after the array allowed us to compare the orienting
array was most likely to have been a predefined target of attention to a spatial location of an expected percep-
stimulus. Other studies (Luck et al., 1994; Hawkins et al., tual stimulus with orienting attention to an internal
1990; Downing, 1988) have also used cues presented after spatial representation of the same percept held in WM.
a stimulus array, but to indicate which location in an array Of specific interest was whether the same or different
required a decision about the presence or the absence of processes were involved in the two cases, and whether
a predefined target stimulus. Schmidt, Vogel, Woodman, they had the same time course of operation.
and Luck (2002) also presented cues after a stimulus
array, but in this case the cues were nonpredictive,
peripheral transients, presented immediately after the EXPERIMENT 1: PREDICTING LOCATIONS IN
array, to study the transfer of information into WM. EXTERNAL VERSUS INTERNAL SPACE
Recently, there have been studies examining ERPs to
cues presented after arrays to be memorized, which This experiment established that there were behavioral
instruct subjects what feature dimension is to be retrieved advantages of orienting to locations in both extraperso-
from stimuli held in WM (Bosch, Mecklinger, & Friederici, nal space and internal spatial representations.
2001; Mecklinger, 1998). However, these investigations
have been primarily concerned with differences between Results
retrieving spatial versus object information from WM, and
the memory rehearsal processes involved in each case Reaction Times
(Mecklinger, 1998; Ruchkin, Johnson, Grafman, Can- The pattern of reaction time is shown in Figure 2A. The
oune, & Ritter, 1997). first comparison was between valid and invalid trials for

Griffin and Nobre 1177

Figure 1. Schematics for
experimental tasks. Stimulus
durations and interstimulus
intervals were the same in the
three tasks. All trials contained
a pre-cue (100 msec), an array
of four differently colored
crosses (100 msec), a
retro-cue (100 msec), and a
probe stimulus (100 msec). In
pre-cue trials, the spatially
informative orienting cue (80%
validity) was presented before
the stimulus array. In retro-cue
trials, the spatially informative
orienting cue (80% validity) was
presented after the stimulus
array. In neutral trials, there was
no spatially informative
orienting cue. (A) Experiment
1: In all trials subjects
responded according to
whether the central probe
stimulus had been present in
the array (50% probability).
(B) Experiment 2: The probe
stimulus was presented
peripherally, at one of the four
array locations, and subjects
responded according to
whether the probe stimulus
matched the item presented at
that location in the array (50%
probability). (C) Experiment 3:
There were only pre-cue and
retro-cue trials. The central
probe stimulus was always
present in the array, and
subjects responded according
to whether it occurred at the
cued location (50% probability).

pre-cues and retro-cues. There was a significant main three conditions. Post hoc contrasts revealed that neu-
effect of validity, F(1,9) = 13.774, p = .005, reflecting tral trials had longer reaction times than both pre-cue
shorter reaction times in valid trials. There was no effect and retro-cue trials [pre-cue vs. neutral, F(1,9) = 9.501,
of cue type, or interaction between cue and validity. This p = .013; retro-cue vs. neutral, F(1,9) = 8.666, p = .016].
indicates that the pattern of shortened reaction times Reaction times in pre-cue trials versus retro-cue trials
for valid versus invalid trials was equivalent for pre-cues were equivalent. There was a main effect of response,
and retro-cues. F(1,9) = 13.693, p = .005, reflecting quicker responses
Reaction-time advantages for valid pre-cues and retro- to probes that had been present in the array (‘‘yes’’
cues relative to neutral cues were tested using a two-way probes). There was also a significant interaction between
repeated-measures analysis of variance (ANOVA) with the response and cue factors, F(2,18) = 4.854, p = .032.
factors of response (yes, no) and cue (pre, retro, neu- Post hoc contrasts revealed that the differences in
tral), using only valid pre-cue and retro-cue trials. There reaction time between ‘‘yes’’ and ‘‘no’’ trials were larger
was a main effect of cue, F(2,18) = 8.431, p = .012, for pre-cue and retro-cue trials compared to neutral
indicating differences in reaction times between the trials (although the effect in retro-cue trials only tended

1178 Journal of Cognitive Neuroscience Volume 15, Number 8

toward significance) [pre-cue/neutral  yes/no, F(1,9) = Accuracy advantages for valid pre-cues and retro-cues
7.031, p = .026; retro-cue/neutral  yes/no, F(1,9) = relative to neutral cues were tested using a two-way
4.104, p = .073]. There was no difference between pre- repeated-measures ANOVA with factors of response
cue and retro-cue trials. (yes, no) and cue (pre, retro, neutral), using only valid
Reaction-time costs for invalid pre-cues and retrocues pre-cue and retro-cue trials. There was a main effect of
relative to neutral cues were tested using a similar cue, F(2,18) = 6.209, p = .023, indicating differences in
analysis with factors of response (yes, no) and cue accuracy between the three cue types. Post hoc contrasts
(pre, retro, neutral), using only invalid pre- and retro- revealed that pre-cue trials had greater accuracy than
cue trials. There were no significant effects of cue, or neutral trials [pre-cue vs. neutral, F(1,9) = 7.392, p =
interaction between cue and response. There was only .024]. Otherwise, accuracy did not differ between the
a significant main effect of response, F(1,9) = 6.680, p = conditions. There was no significant effect of response,
.029, again indicating shorter reaction times when the indicating no differences in accuracy between ‘‘yes’’ and
probe was present in the array. ‘‘no’’ trials. There was a significant interaction between
response and cue, F(2,18) = 4.663, p = .029. Post hoc
contrasts revealed that the interaction was due to the
Accuracy
differences in accuracy between ‘‘yes’’ and ‘‘no’’ trials
The pattern of performance accuracy in the experimental being greatest for pre-cue trials compared to neutral trials
conditions is shown in Figure 2B. The first comparison [pre-cue/neutral  yes/no, F(1,9) = 7.943, p = .020].
was between valid and invalid trials for pre-cues and Accuracy costs for invalid pre-cue and retro-cue trials
retro-cues. There was a significant main effect of validity, relative to neutral cues were tested using a similar analysis
F(1,9) = 12.358, p = .007, reflecting greater accuracy in with factors of response (yes, no) and cue (pre, retro,
valid trials. There was no effect of cue, or interaction neutral), using only invalid pre- and retro-cue trials. There
between cue and validity. This indicates that the pattern was no significant effect of cue, indicating no overall
of increased accuracy for valid versus invalid trials was difference in accuracy between the three cue types. There
equivalent for pre-cues and retro-cues. was a significant main effect of response, F(1,9) = 9.820,
p = .012, reflecting decreased accuracy in ‘‘yes’’ trials
compared with ‘‘no’’ trials. There was a significant inter-
action between the response and cue factors, F(2,18) =
4.191, p = .034. Post hoc contrasts revealed that the
difference in accuracy between ‘‘yes’’ and ‘‘no’’ trials (less
accurate in ‘‘yes’’ trials) was greater in pre-cue and retro-
cue trials than neutral trials [pre-cue/neutral  yes/no,
F(1,9) = 5.722, p = .040; retro-cue/neutral  yes/no,
F(1,9)=5.714, p = .041]. There was no difference in the
pattern of results between pre-cue and retro-cue trials.

Summary
The behavioral analyses showed that subjects have de-
creased reaction times and increased accuracy in valid
trials compared to invalid trials. This pattern of results was
the same for both pre-cues and retro-cues. Compared
with neutral trials, subjects showed reaction-time advan-
tages for valid trials, and accuracy disadvantages for
invalid trials. These effects were equivalent for pre-cues
and retro-cues. Pre-cues also showed accuracy advantages
compared to neutral trials. Overall, reaction times were
shorter for trials where the probe stimulus was present in
the array compared with trials where it was not present in
the array for all cue types (pre-cue, retro-cue, neutral).

Figure 2. (A) Experiment 1: Mean reaction time (RT) and standard Discussion
error for probe stimuli in pre-cue, retro-cue, and neutral trials,
The aim of Experiment 1 was to investigate whether it
separated according to the factors of response and cue validity. (B)
Experiment 1: Mean accuracy and standard error for probe stimuli in is possible to orient selective spatial attention to inter-
pre-cue, retro-cue, and neutral trials, separated according to the factors nal representations in WM. We showed that it is
of response and cue validity. possible to orient attention to internal spatial represen-

Griffin and Nobre 1179

tations, at least when the number of items and features is litating the decision about where a target stimulus might
within the limits of WM capacity (c.f. Wheeler & Treis- have been.
man, 2002; Vogel, Woodman, & Luck, 2001; Luck & In a recent study, Schmidt et al. (2002, Experiment 4)
Vogel, 1997). Surprisingly, orienting to internal represen- presented nonpredictive peripheral cues immediately
tations was characterized by an almost identical pattern after stimulus array offset, and found memory for
of behavioral costs and benefits of cueing as orienting to subsequent probe stimuli presented at the cued loca-
spatial locations. tion to be better than memory for stimuli at uncued
The results of Experiment 1 showed that there were locations. Their results suggest that cues appearing after
similar behavioral benefits and costs of orienting atten- the offset of a visual stimulus can influence the transfer
tion to upcoming perceptual stimuli and to internal of perceptual information into visual WM. However,
representations. In the pre-cue condition, the finding Schmidt and colleagues used nonpredictive, peripheral
of faster reaction times to validly cued trials, and transients as cues, presented in the iconic memory
decreased accuracy to invalid trials, relative to neutral period. The present study presented orienting cues
trials replicated the well-known effects of orienting after the iconic trace period, and demonstrated behav-
spatial attention to visual arrays (e.g., Hawkins et al., ioral advantages of predictive, central cues that oriented
1990; Müller & Findlay, 1987; Müller & Rabbit, 1989; subjects’ attention to locations in WM.
Jonides, 1981; Posner, 1980). The behavioral effects in It is important to note that in our experiment the
the pre-cue condition are thus best conceptualized by number of objects and features fell within the limits of
both benefits of valid cueing, and costs of invalid WM capacity (c.f. Wheeler & Treisman, 2002; Vogel et
cueing. Responses were also quicker in the pre-cue al., 2001; Luck & Vogel, 1997). It is possible that the
condition to probe stimuli that had appeared in the findings may have been different if the number of
array than those that did not. Similar findings of faster objects or features in the perceptual array was beyond
responses to ‘‘yes’’ stimuli are common in the visual the capacity of WM. When arrays exceed WM capacity,
search literature (e.g., Chelazzi, 1999; Wolfe, 1994; internal representations of arrays may be insufficient to
Treisman & Gelade, 1980). afford selection by retro-cues. In these cases, pre-cues
The findings of behavioral effects due to valid and may bias significantly the perceptual processing of the
invalid cueing in the retro-cue condition show that it is array in order to confer behavioral advantages. Further
possible and advantageous to orient attention toward experimentation is needed to clarify this issue.
selective attributes of internal representations. Re- The finding of a similar pattern of behavioral benefits
sponses to valid cues were quicker, and responses and costs for pre-cues and retro-cues suggests that a
to invalid cues were less accurate, than neutral trials. reasonable representation of the array was available after
This was exactly the same pattern of behavioral costs and a neutral pre-cue in the retro-cue trials, otherwise retro-
benefits seen in the pre-cue condition. Also, as in the pre- cues would not show the same pattern of effects as pre-
cue condition, responses were quickest for probe stimuli cues. This suggests that the enhancement of early
that had been presented in the array. A similar finding of perceptual representations is not necessary to drive
faster responses to matching probe stimuli is typically the behavioral benefits and costs of attention in the
found in delayed match-to-sample tasks (e.g., Klaver, current study. The results are compatible with a weight-
Smid, & Heinze, 1999; Mecklinger, 1998). ed integration and data-limited view of attention as
This experiment was different to other studies using proposed by Kinchla et al. (1995), in which more weight
cues presented after a perceptual array in two main is given to cued elements in an internal representation
respects. Firstly, the response-relevant targets were not with regard to response selection. Alternatively, it is
predetermined, but were only defined at the end of possible that retro-cues act to enhance portions of the
each trial by a separate probe stimulus. Secondly, retro- internalized representation of the array. As both accura-
cues provided information about the likely location of cy and reaction-time data showed the same pattern of
the relevant target, but did not drive a response deci- results, they rule out explanations based on speed/
sion. In most previous experiments, targets were pre- accuracy tradeoffs.
defined (Kinchla et al., 1995; Luck et al., 1994; Hawkins One shortcoming of the design in Experiment 1,
et al., 1990; Downing, 1988). Post-cues signaled the which limits interpretations of the results, is that it is
location at which a decision was required (Luck et al., not possible to know which array location(s) the sub-
1994; Hawkins et al., 1990; Downing, 1988), or provided jects used to make a decision about the presence or
information about target location (Kinchla et al., 1995). absence of the probe stimulus from the array. This
In the above experiments, the post-cues could facilitate leaves open the possibility that subjects adopted differ-
decision making about whether a target was present ent response criteria for cued and uncued locations in
or absent. The present task design separates orienting the array, as they knew the probe was more likely to
attention to mental representations from response have appeared at the cued location. Validity effects could
decisions, and therefore the behavioral facilitation ob- therefore have been contaminated by differences in
served by retro-cues cannot be attributed merely to faci- response criteria for probes that occurred at cued versus

1180 Journal of Cognitive Neuroscience Volume 15, Number 8

uncued locations—an effect that would be expected to
be similar for both pre-cue and retro-cue trials.1

EXPERIMENT 2: PREDICTING LOCATIONS IN

EXTERNAL VERSUS INTERNAL SPACE
(PERIPHERAL PROBES)
In a follow-up experiment, we presented the probe
stimulus peripherally, at one of the four original array
locations. The probe required a decision to be made
about the item that had occurred at a specific spatial
location of the array. Subjects decided whether the color
of the probe and the item at the same location in the
array matched (50% probability). Behavioral differences
between valid, neutral, and invalid trials using these
peripheral probes are not affected by differential re-
sponse criteria for different array locations, since deci-
sions are probed at each location separately. As before,
the pre-cue and retro-cue stimuli did not afford any
response decisions. Rather, they predicted (80% validity)
the likely location to be probed.
The results established that the behavioral effects of
orienting to locations in both extrapersonal space and
internal representations observed in Experiment 1 were
not due to differences in response criteria between valid
and invalid trials.

Results
Subjects were able to perform both pre-cue and retro-
cue trials while maintaining visual fixation. Analysis of
eye movements indicated a minimal number of devia- Figure 3. (A) Experiment 2: Mean reaction time (RT) and standard
tions from central gaze (an average of six trials pre error for probe stimuli in pre-cue, retro-cue, and neutral trials,
separated according to the factor of cue validity. ‘‘Yes’’ and ‘‘no’’ trials
subject were rejected).
have bee collapsed. (B) Experiment 2: Mean accuracy and standard
error for probe stimuli in pre-cue, retro-cue, and neutral trials,
separated according to the factor of cue validity. ‘‘Yes’’ and ‘‘no’’ trials
Reaction Times
have bee collapsed.
The pattern of reaction-time results is shown in Figure
3A. The first comparison was between valid and invalid trials versus retro-cue trials were equivalent. There was a
trials for pre-cues and retro-cues. There was a signifi- main effect of response, F(1,9) = 49.572, p < .001,
cant main effect of validity, F(1,9) = 207.155, p < .001, reflecting quicker responses to probe stimuli that
reflecting shorter reaction times in valid trials. There matched the item that had been present at the probed
was no effect of cue, or interaction between cue and location in the array (‘‘yes’’ probes). There was also a
validity. This indicates that the pattern of shortened significant interaction between the response and cue
reaction times for valid versus invalid trials was equiv- factors, F(2,18) = 4.181, p = .047. Post hoc contrasts
alent for pre-cues and retro-cues. revealed that the differences in reaction time between
Reaction-time advantages for valid pre-cues and retro- ‘‘yes’’ and ‘‘no’’ trials were larger for pre-cue and retro-
cues relative to neutral cues were tested using a two-way cue trials compared to neutral trials (although the effect
repeated-measures ANOVA with factors of response in retro-cue trials only tended toward significance) [pre-
(yes, no) and cue (pre, retro, neutral), using only valid cue/neutral  yes/no, F(1,9) = 5.860, p = .039; retro-
pre-cue and retro-cue trials. There was a main effect of cue/neutral  yes/no, F(1,9) = 4.876, p = 0.055]. There
cue, F(2,18) = 56.660, p < .001, indicating differences in was no difference between pre-cue and retro-cue trials.
reaction times between the three conditions. Post hoc Reaction-time costs for invalid pre-cues and retro-cues
contrasts revealed that neutral trials had longer reaction relative to neutral cues were tested using a similar
times than both pre-cue and retro-cue trials [pre-cue vs. analysis with factors of response (yes, no) and cue
neutral, F(1,9) = 39.305, p < .001; retro-cue vs. neutral, (pre, retro, neutral), using only invalid pre and retro-
F(1,9) = 65.861, p < .001]. Reaction times in pre-cue cue trials. There were no significant effects of cue, or

Griffin and Nobre 1181

interaction between cue and response. There was only had decreased reaction times and increased accuracy
a significant main effect of response, F(1,9) = 7.098, in valid trials compared to invalid trials. Compared to
p = .026, again indicating shorter reaction times to neutral trials, subjects showed reaction time and accuracy
probe stimuli that matched the item that had been pre- advantages for valid trials, and accuracy disadvantages for
sent at the probed location in the array (‘‘yes’’ probes). invalid trials. These reaction time and accuracy effects
were equivalent for pre-cues and retro-cues. Overall,
reaction times were shorter for trials where the probe
Accuracy stimulus matched the item that had been present at the
probed location in the array (‘‘yes’’ probes) compared to
The pattern of performance accuracy in the experi-
trials where it did not match the item that had been
mental conditions is shown in Figure 3B. The first
present at the probed location in the array (‘‘no’’ probes)
comparison was between valid and invalid trials for
for all cue types (pre-cue, retro-cue, neutral).
pre-cues and retro-cues. There was a significant main
effect of validity, F(1,9) = 105.498, p < .001, reflecting
greater accuracy in valid trials. There was no effect of
cue, or interaction between cue and validity. This in- Discussion
dicates that the pattern of increased accuracy for valid
The aim of Experiment 2 was to replicate the behavioral
versus invalid trials was equivalent for pre-cues and
benefits and costs of orienting attention to upcoming
retro-cues.
perceptual stimuli and internal representations, using a
Accuracy advantages for valid pre-cues and retro-cues
modified version of the task in Experiment 1 that
relative to neutral cues were tested using a two-way
controlled for possible differences in response criteria
repeated-measures ANOVA with factors of response
between cued and uncued array locations. We showed
(yes, no) and cue (pre, retro, neutral), using only valid
that in this task there were behavioral effects associated
pre-cue and retro-cue trials. There was a main effect of
with pre-cues and retro-cues that were almost identical
cue, F(2,18) = 7.580, p = .011, indicating differences in
to those seen in Experiment 1.
accuracy between the three cue types. Post hoc con-
The finding of similar behavioral costs and benefits
trasts revealed that neutral trials had less accuracy than
of pre-cues and retro-cues in Experiment 2 ruled out
both pre-cue and retro-cue trials [pre-cue vs. neutral,
changes in response criteria as the only explanation
F(1,9) = 9.076, p = .015; retro-cue vs. neutral, F(1,9) =
for the effects. Presentation of the probe stimulus at
7.839, p = .021]. Accuracy in pre-cue trials versus retro-
one of the four array locations meant that a decision
cue trials was equivalent. There was no significant effect
about the presence or absence of the probe had to be
of response, indicating no differences in accuracy be-
made at only one location. Therefore, any differences
tween ‘‘yes’’ and ‘‘no’’ trials. There was a significant
in performance between valid and invalid trials cannot
interaction between response and cue, F(2,18) = 4.414,
be attributed to subjects adopting different response
p = .049. Post hoc contrasts revealed that the interac-
criteria for cued and uncued locations in the array
tion was due to the differences in accuracy between
when making their decision about the presence or
‘‘yes’’ and ‘‘no’’ trials being greatest for pre-cue trials
absence of the probe stimulus. The results from
compared to neutral trials [pre-cue/neutral  yes/no,
Experiment 2 suggest a spatially specific enhancement
F(1,9) = 5.170, p = .049].
of portions of the internal representation of the
Accuracy costs for invalid pre-cue and retro-cue trials
stimulus array.
relative to neutral cues were tested using a similar
The behavioral results from Experiments 1 and 2 thus
analysis with factors of response (yes, no) and cue (pre,
demonstrate that we are able to orient selectively to
retro, neutral), using only invalid pre- and retro-cue
spatial locations within internal spatial representations
trials. There was a main effect of cue, F(2,18) = 23.580,
as well as to external perceptual stimuli, with these ef-
p < .001, indicating differences in accuracy between the
fects not being due to differences in response criteria
three cue types. Post hoc contrasts revealed that neutral
between valid and invalid trials. However, one cannot
trials had greater accuracy than both pre-cue and retro-
determine whether the same or different processes
cue trials [pre-cue vs. neutral, F(1,9) = 35.906, p < .001;
underlie these effects in the two cases by behavioral
retro-cue vs. neutral, F(1,9) = 24.656, p = .001]. Accu-
measures alone. ERPs provide a measure of on-line
racy in pre-cue trials versus retro-cue trials was equiva-
information processing during task performance.
lent. There was no significant effect of response, or
interaction between cue and response.

EXPERIMENT 3: IMPERATIVE CUEING OF

Summary EXTERNAL VERSUS INTERNAL SPACE
The pattern of results in Experiment 2 was almost Data from 12 subjects (7 men) were used to characterize
identical to those observed in Experiment 1. Subjects the behavioral and ERP effects of cueing attention to

1182 Journal of Cognitive Neuroscience Volume 15, Number 8

perceptual locations and internal representations. The between pre- and retro-cues, mirror those found in
same 12 subjects were used for both behavioral and Experiments 1 and 2.
ERP analysis. Data were rejected from 7 additional sub- The pattern of accuracy is shown in Figure 4B. There
jects due to excessive eye blinks or saccades, and from was no significant effect of cue, or interaction between
5 additional subjects who had excessive artefacts during cue and response. There was a significant main effect of
EEG recording. response, F(1,11) = 18.691, p = .001, reflecting more
accurate responses to probes that occurred at the cued
location than to probes that were not in the cued
location. As with the reaction-time data, the increased
Results accuracy for ‘‘yes’’ responses compared with ‘‘no’’ re-
Behavioral Results sponses, and the absence of differences between pre-cues
and retro-cues, are similar to the findings from Experi-
The pattern of reaction times is shown in Figure 4A. ments 1 and 2.
There was no significant effect of cue, or interaction
between cue and response, indicating that the pattern of
results was equivalent for pre-cues and retro-cues. There ERP Results
was a significant main effect of response, F(1,11) = Waveforms to the spatial cues were characterized by
60.577, p = .0001, reflecting faster responses to probes early visual responses over lateral posterior sites and a
that occurred in the cued location than probes that were late positive peak broadly distributed over the scalp.
not in the cued location. These two results, faster ‘‘yes’’ Experimental factors exerted three types of effects upon
responses than ‘‘no’’ responses, and no differences waveforms elicited by spatial cues: effects of spatial
orienting that were common for pre-cues and retro-
cues, effects of spatial orienting that were selective
for either pre-cues or retro-cues, and effects of cue
(pre, retro) that were independent of the direction of
attention (left, right).
Common effects of spatial orienting for pre-cues
and retro-cues. Common effects of spatial orienting
were calculated as effects of side of attention that did
not interact with cue type. Pre-cues and retro-cues
elicited a common set of lateralized posterior and frontal
ERP components sensitive to the direction of attention.
There was an early modulation of the visually evoked N1
component over the posterior scalp contralateral to the
direction of attention (see Figures 5 and 6). Statistically,
this effect was evident at the posterior and lateral scalp
regions between 120 and 200 msec, as both two-way
interactions between side of attention and hemisphere,
and three-way interactions between side of attention,
hemisphere, and electrode site, F’s > 5.274, p < .01.
These reflect the fact that the N1 was enhanced over
the right hemisphere when subjects attended left, and
over the left hemisphere when subjects attended right,
and that these effects were maximal at the PO7/8 and
TP7/8 electrode pairings. They also reflect the fact that
this contralateral N1 enhancement was larger over the
left hemisphere.
A later common effect of spatial orienting involved a
contralateral central and frontal negativity between 360
and 480 msec (see Figures 5 and 6). Statistically, these
effects were evident over frontal, central, and lateral
scalp regions as two-way interactions between direction
of attention and hemisphere, and three-way interac-
Figure 4. (A) Experiment 3: Mean reaction time (RT) and standard tions between direction of attention, hemisphere, and
error for probe stimuli in pre-cue and retro-cue trials, separated
according to the factor of response. (B) Experiment 3: Mean accuracy electrode site, F’s > 2.713, p < .05. These interactions
and standard error for probe stimuli in pre-cue and retro-cue trials, with electrode site indicate that the differences were
separated according to the factor of response. maximal at the F5/6, C3/4, and FC5/6 electrode pairs.

Griffin and Nobre 1183

Figure 5. Grand-averaged
waveforms (N = 12 subjects)
elicited by pre-cue stimuli
when subjects attended the left
(thick line) and right (thin line)
visual fields. To isolate effects
of spatial orienting, both
attend-left and attend-right
waveforms have had the
waveform from the neutral cue
stimulus that occurred at the
same time point subtracted
from them. The electrode
montage is shown at the
top-left, with the locations of
the electrode sites for the
sample waveforms shaded
black. The horizontal eye
channel (HEOG) is also shown.
The polarity of the waveforms
is plotted with positive values
upward, and arrows indicate
the presence of statistically
significant effects in this and
subsequent figures. Scalp
topographies of the difference
in grand-averaged ERPs elicited
by pre-cue stimuli when
subjects attended the right
versus left visual fields are
shown at the bottom of the
figure. The maps are shown
from a bird’s-eye perspective.
The color scale (colored bar on
the right) shows the range of
possible voltage values in the
topographies. The anterior
scalp is shown at the top and
the right scalp is shown at the
right side in this and
subsequent figures.

Effects of cue independent of the direction of atten- Selective effects of spatial orienting for retro-cues.
tion. Differences between orienting to perceptual versus Differences in lateralized spatial orienting between pre-
internal representations were revealed by effects of cue cues and retro-cues were revealed by statistical inter-
that did not interact with side of attention. Differences actions between the factors of direction of attention
during the P300 component were observed between (left, right) and cue (pre, retro). The first such interac-
pre-cues and retro-cues. There was an increased relative tion occurred early, between 120 and 200 msec, over the
positivity for retro-cues over posterior scalp regions, and frontal scalp sites. Modulation of ERPs was only present
an increased relative negativity for retro-cues compared for retro-cues, and was a difference in the waveforms
with pre-cues over the anterior scalp (see Figure 7). over the anterior scalp regions. There was an increased
This effect was nonlateralized, and was evident from relative positivity over these regions when subjects
360 msec onwards as both a main effect of cue, and attended right (see Figure 6). This was reflected by
interaction between cue and electrode site over mid- interactions between direction of attention and cue
line, frontal, central, lateral, and posterior scalp sites, over frontal [120–200 msec; F’s > 8.803, p < .013]
F’s > 3.578, p < .046. and central regions [160–200 msec; F(1,11) = 5.953,

1184 Journal of Cognitive Neuroscience Volume 15, Number 8

Figure 6. Grand-averaged
waveforms (N = 12 subjects) elicited
by retro-cue stimuli when subjects
attended the left (thick line) and
right (thin line) visual fields. To
isolate effects of spatial orienting,
both attend-left and attend-right
waveforms have had the waveform
from the neutral cue stimulus that
occurred at the same time point
subtracted from them. Scalp
topographies of the difference in
grand-averaged ERPs elicited by
retro-cue stimuli when subjects
attended the right versus left visual
fields are shown at the bottom of the
figure.

p = .033]. During this time period, there was also a Discussion

main effect of cue over the frontal scalp sites [160 and
ERP recordings during task performance permitted real-
200 msec; F(1,11) = 15.039, p = .003]. Retro-cues
time measurement of the orienting of attention by
evoked a positive potential, which was not evident for
identical cueing stimuli to a location in an upcoming
pre-cues (see Figure 7).
perceptual array, or to the same location in the internal
A later effect was only present for retro-cues, and
representation of the array. The neural correlates of the
was a difference in the relative negativity/positivity
two processes share a common pattern of lateralized
between the two hemispheres depending on the direc-
activity over the posterior then frontal regions. Addition-
tion of attention between 240 and 320 msec over
al processes including early frontal activity are involved
frontal, central, and lateral scalp sites (see Figure 7).
when orienting toward a representation in WM.
This was evident both as two-way interactions of cue
and hemisphere, and three-way interactions between
cue, side of attention, and hemisphere, F’s > 6.139,
Behavioral Results
p < .031. These indicate that for retro-cues, when
attention was directed to the right, the distribution of Behavioral results from Experiment 3, which used imper-
negative charge was greater over the left hemisphere, ative, as opposed to predictive cues, were compatible with
and the distribution of positive charge was greater over those seen in Experiments 1 and 2. Subjects were faster,
the right hemisphere. and more accurate, to respond when probe stimuli had

Griffin and Nobre 1185

Figure 7. Grand-averaged
waveforms (N = 12 subjects)
elicited by pre-cue stimuli (thin
line) and retro-cue stimuli (thick
line). To isolate effects of spatial
orienting, both pre-cue and
retro-cue waveforms have had the
waveform from the neutral cue
stimulus that occurred at the
same time point subtracted from
them. Scalp topographies of the
difference in grand-averaged ERPs
elicited by retro-cue stimuli
versus pre-cue stimuli are shown
at the bottom of the figure.

occurred in the cued location (‘‘yes’’ responses) than optimization of behavior by pre-cues and retro-cues.
when it had occurred at an uncued location. Once again, The task equated perceptual and motor demands
there was no difference in the pattern of results between between the two conditions. Analysis was carried out
the pre-cue and retro-cue conditions. on physically identical cueing stimuli occurring either
before or after the array. Because pre-cues and retro-
cues occurred after different types of events and at
ERP Results
different stages in a trial, direct comparison of their
The analysis of the ERPs elicited by the cueing stimuli waveforms might have reflected differential overlap
revealed that there were both similarities and differ- from preceding events or differential cognitive states
ences in the attentional mechanisms involved in the that were not linked to spatial orienting. Therefore, to

1186 Journal of Cognitive Neuroscience Volume 15, Number 8

minimize contribution from these nonspecific factors, tween 360 and 480 msec. There was increased relative
we subtracted the waveforms elicited by neutral cue negativity over fronto-central left hemisphere sites when
stimuli from those elicited by informative cues that attention was directed to the right. Late frontal activity
occurred at the same time point. To minimize further has been consistently reported in studies of shifting
possible contribution from differential component spatial attention (Hopf & Mangun, 2000; Nobre et al.,
overlap from preceding items, intervals between suc- 2000; Yamaguchi et al., 1994; Harter et al., 1989),
cessive trial events were randomly jittered. Only the however, its distribution and polarity have varied be-
effects of spatial orienting before or after the array are tween these studies. The precise neural processes re-
therefore captured in the resulting waveforms consid- sponsible for these effects, and the reason for their
ered in the analysis. variability between tasks, remain to be understood.
Common processes in pre-cue and retro-cue trials. Late positive potentials over posterior scalp sites linked
Attentional orienting modulated the contralateral visual to directing attention (LDAP) have frequently been re-
N1 component in both pre-cue and retro-cue trials. ported in spatial attention experiments (Eimer, Velzen, &
Contralateral negativities have been reported in most Driver, 2002; Hopf & Mangun, 2000; Harter et al., 1989).
previous studies of attentional orienting (Hopf & Man- The possible contribution of this potential could not be
gun, 2000; Nobre et al., 2000; Yamaguchi et al., 1994; evaluated in the present experiment because of the
Harter et al., 1989) although none of the effects began as restricted time window analyzed after each cue stimulus.
early as ours (120 msec). Such effects have been inter- The occurrence of probe stimuli at a relatively short time
preted as reflecting the initial steps of attentional orient- period after retro-cues (500–1000 msec) precluded the
ing toward a spatial location. There were physical measurement of uncontaminated cue-related activity
differences in the cueing stimuli for attending left and during the late time intervals associated with the LDAP.
right in this study (and some of the others), which may In future experiments, it will be interesting to extend the
have contributed to the earliest parts of the effect. interval between retro-cues and probes in order to exam-
However, the fact that similar early negativities were ine whether this component is elicited when orienting to
found when stimuli were physically equated across internal representations.
cueing conditions (Nobre et al., 2000), suggests that Selective effects of spatial orienting for retro-cues. In
the N1 modulation may reflect part of the process of the retro-cue condition only, there was modulation of
orienting visual attention. Given that the same effect was activity depending on cue direction (left, right) over
found in pre-cue and retro-cue trials, this suggests that frontal scalp sites between 120 and 200 msec. The
some early stages of orienting to spatial locations and earliest reported frontal effects in attentional orienting
mental representations may share the same underlying tasks have begun at 200 msec (Nobre et al., 2000), and
processes. However, the existence of purely attentional no effects as early as this have been seen in studies
early contralateral modulation has been challenged where subjects are cued to retrieve spatial or object
recently by Velzen and Eimer (submitted), who found information from WM (Bosch et al., 2001; Mecklinger,
that N1 modulation depended on physical asymmetries 1998). Awh et al. (2000) found early modulation of
of the predictive properties of central cue stimuli. activity over frontal scalp sites to probe stimuli by both
Further investigation will be required to clarify this spatial attention and rehearsal in WM in a task where
important issue. subjects either had to attend to, or remember, spatial
Inspection of waveforms revealed that the N1 modula- locations. However, this effect, unlike the present find-
tion was much larger over the left hemisphere. Larger left ing, was not restricted to the memory condition. The
hemisphere differences were also seen for the early present finding of early frontal modulation appears to be
negativities in other orienting studies (Hopf & Mangun, specific to orienting to internal mental representations.
2000; Nobre et al., 2000). Initially, this does not seem The most intuitive explanation of the current results
consonant with the notion of right hemisphere speciali- would be selective engagement of frontal regions when
zation for directing attention (Mesulam, 1981). However, orienting to mental representations in WM. There is an
this may indicate that there was greater variability in the enormous literature about frontal involvement in WM
activity in posterior left hemisphere regions between (e.g., Levy & Goldman Rakic, 2000; Owen, 2000; Petrides,
directing attention to the left and right, whereas the right 2000), and it is possible that the present effect, being
hemisphere was more similarly involved in attention shifts restricted to retro-cues, reflects some aspect of orienting
in both directions (c.f. Nobre et al., 2000). This is consis- to internal representations. Its early latency is consistent
tent with behavioral, lesion, and imaging data that have with a source of stimulus-driven top-down biasing signal
suggested that right hemisphere regions are able to direct (Desimone & Duncan, 1995), that could perhaps guide
attention into both visual hemifields, while posterior left the retrieval, maintenance, or filtering of information
hemisphere regions are primarily involved in directing within WM. Its latency makes it less likely to reflect later
attention into the right hemifield (Mesulam, 1981, 1999). monitoring or decision-related processes. However, the
Also common to pre-cues and retro-cues were effects exact functional significance of this effect remains to be
sensitive to the direction of attentional orienting be- resolved. It has been suggested (Vogel & Luck, 2000)

Griffin and Nobre 1187

that such early frontal effects in some cases may reflect by lateralized modulation over anterior scalp locations
overlap with motor potentials linked to response prep- (Hopf & Mangun, 2000; Nobre et al., 2000; Yamaguchi
aration. This is unlikely here, as the response decision is et al., 1994; Harter et al., 1989). This pattern of ERP
not afforded until the probe stimulus is presented. modulation is consistent with parietal and frontal acti-
As this early frontal effect temporally overlaps with vation during the spatial orienting of attention, and
the posterior N1 modulation, this suggests that con- tentatively suggests that parietal activity leads frontal
current activity in multiple brain areas underlie the activity in the control of spatial attentional orienting
observed effects. Although it is hard to pinpoint the (c.f. Velzen & Eimer, submitted).
sources of these modulations, it is possible that they The present results suggest that orienting spatial
may be related, as studies have shown interactions attention to external and internal spatial locations both
between frontal and parietal areas during WM tasks involve this same network reflected in parietal–frontal
(e.g., Chafee & Goldman Rakic, 2000). What can be ERP modulations. Furthermore, orienting attention to
concluded is that brain activity reflected over frontal locations in internal representations engages additional
scalp regions plays an early role in orienting attention processes, including early processing reflected over the
to internal representations. frontal scalp and widely distributed changes in later
The next phase of differences restricted to retro-cues ERP components. These results are consistent with
were hemispheric differences in the distribution of findings from a recent fMRI investigation of spatial
positive and negative charge depending on cue direc- orienting to external versus internal spatial representa-
tion over frontal, central, and lateral scalp sites between tions (Nobre et al., 2002). The task was similar to
240 and 320 msec. Although frontal and central modu- Experiment 3, except that the intervals between suc-
lations dependent on the direction of attention were cessive displays were much longer, in order to resolve
seen in attentional orienting studies (Hopf & Mangun, the brain activity related to cues and arrays separately.
2000; Nobre et al., 2000) around this time period, the Both pre-cues and retro-cues engaged activity in the
fact that here they were only present for retro-cues core posterior parietal, lateral premotor, and dorsal
suggests an explanation based more on spatial WM than prefrontal areas of the spatial orienting network. In
on attention. Possible contributory processes include addition, retro-cues selectively engaged more anterior
interactions between biasing signals and extrastriate prefrontal areas and enhanced common activations in
regions (Desimone & Duncan, 1995), maintenance or the parietal cortex. The present ERP results suggest
retrieval of spatial information from WM, target selec- that the prefrontal contribution, selectively linked to
tion, and filtering of distractors. The wide distribution of orienting spatial attention to internal representations,
the effect (as opposed to only frontal scalp sites) may occur quite early in the orienting process. Early
suggests the activity of a distributed network(s) of brain prefrontal involvement would be in line with a top-
areas, perhaps engaged in multiple temporally overlap- down role in controlling attentional selection guided
ping functions. by memory.
The final difference between pre-cues and retro-cues Complementary findings in other studies. Our re-
was enhancement of the P300 component for retro-cues. sults are consistent with reports of the shared action of
The P300 component can be modulated by many factors, attention and WM upon visual processing. Awh and
although in this case we believe that the enhanced colleagues (2000) found that both spatial attention
positivity for retro-cue trials reflected the context updat- and rehearsal in spatial WM modulated early visual-
ing or closure of the selection process (Donchin & Coles, evoked potentials and frontal activity to probe stimuli
1988; Desmedt, 1980). This is because informative retro- in a task where subjects either had to attend to, or
cues defined both the stimulus location and the stimulus remember, spatial locations. This gave support to the
color to be selected, as the stimulus array had already suggestion that information is maintained in spatial WM
been presented. In contrast, informative pre-cues pro- by focal shifts of spatial attention (e.g., Awh, Jonides, &
vide information about the location, but not the color to Reuter Lorenz, 1998; Awh & Jonides, 2001). In another
be selected. Color information is deferred until array study, pre-cues directed subjects’ attention to a location
presentation. Therefore, the enhanced P300 to retro-cues in a stimulus array, and post-cues probed whether a
could reflect increased closure of the selection process. target was present at the pre-cued location (Luck et al.,
Neural systems for spatial orienting to external and 1994). P1 and N1 modulation of these post-cue stimuli
internal space. Convergent findings from neuropsychol- occurred as a function of pre-cue validity, similarly
ogy, brain imaging, and primate studies have shown that suggesting attentional enhancement of relevant loca-
visual spatial orienting toward extrapersonal space in- tions in WM. However, these findings are not directly
volves a large-scale network of parietal and frontal comparable to the current study, as they reflect the
cortical areas (see Nobre, 2001; Mesulam, 1999 for modulatory effects of spatial attention to a location
reviews). ERPs elicited by cues that direct attention to probed by a peripheral post-cue stimulus, rather than
spatial locations consistently show early lateralized mod- analysis of cues that initiate spatial orienting toward
ulation over posterior parietal scalp locations followed locations in internal representations. Here we extend

1188 Journal of Cognitive Neuroscience Volume 15, Number 8

the view that attention and WM can have similar effects free of neurological disorders. Visual acuity was normal
on stimulus processing by showing that the orienting or corrected to normal.
processes to perceptual stimuli and stimuli in mental
representations can also share similarities.
Stimuli and Task
Our results also complement findings of a strong rela-
tionship between perceptual functions and mental imag- The task is illustrated in Figure 1A. Participants viewed
ery (cf. Kosslyn et al., 2001; Ishai et al., 2000). In our task, arrays of four differently colored crosses either preceded
retro-cue trials are likely to involve mental imagery of the or followed by spatial cues, and made a delayed decision
remembered array. Of interest is the finding of enhanced about the color of the items in the array. There were three
posterior negativity in the N1 time range (150–200 msec) types of trial: pre-cue, retro-cue, and neutral trials. The
to letter stimuli if they match the letter subjects hold as a key difference between the trial types was whether and
mental image, compared to when they are imagining a when in the trial the attentional orienting cue was pre-
different latter (Farah, Peronnet, Gonon, & Giard, 1988). sented. In pre-cue trials, a spatially informative cue (80%
The authors suggested that mental imagery might involve validity) was presented 1500–2500 msec before the visual
stimulus representations in the visual system. Findings of array of colored crosses. In retro-cue trials, the spatially
similar areas of activation during imagery and perception informative cue (80% validity) was presented 1500–2500
(e.g., Kosslyn et al., 2001; Ishai et al., 2000; O’Craven & msec after the array. In neutral trials, there was no
Kanwisher, 2000), as well as the behavioral effects of spatially informative cue, either before or after the array.
imagery upon perceptual performance (Pashler & Shiu, Each trial contained the same sequence of events. Firstly,
1999; Ishai & Sagi, 1995; Farah, 1985), support this view. an asterisk was presented at the center of the screen (200
Our findings of similar early processes in both pre-cue and msec), which indicated the start of the trial. A square (side
retro-cue trials are consistent with the view that attention length 0.88) then appeared at the center of the screen.
to mental imagery and perception may share similarities. After an interval that ranged randomly from 400 to
The additional activity seen in retro-cue trials may in some 600 msec, a pre-cue was presented for 100 msec. The pre-
way be related to mental imagery, the accessing of per- cue was informative in pre-cue trials, but neutral in retro-
ceptual information from memory. cue and neutral trials. An informative cue consisted of two
Summary. In conclusion, we have demonstrated that adjacent sides of the square brightening (forming an
it is possible to orient selective spatial attention to arrow), which instructed the subject to attend covertly
internal representations when within WM capacity. Ori- to that position (e.g., top left). Neutral cues consisted of
enting attention to internal representations was charac- the whole square brightening, which gave no spatial
terized by an equivalent pattern of behavioral costs and information. After a random interval varying between
benefits as orienting to spatial locations across three 1500 and 2500 msec, an array of four crosses of different
experiments. These validity effects were not due to colors appeared for 100 msec. The crosses were any four
changes in response criteria. They may have reflected of the following colors: red, blue, green, yellow, orange,
spatially specific enhancement of representations and/or cyan, pink, gray. Each cross was 0.88 visual angle in size,
increased weighting given to spatially attended stimuli and centered at 38 horizontal and 38 vertical eccentricity.
for response selection. ERP analysis of the identical After another random delay, ranging between 1500 and
cueing stimuli revealed that the neural correlates of the 2500 msec, a retro-cue was presented for 100 msec. The
two processes shared both similarities and differences. retro-cue was informative in retro-cue trials, but neutral in
Orienting spatial attention to both perceptual locations pre-cue and neutral trials. The informative cue again
and internal representations involved lateralized activity consisted of two adjacent sides of the square brightening
over posterior, and then frontal regions, as has been seen (forming an arrow), which instructed the subject to
in previous studies of attentional orienting. Additional attend covertly to that location in the internal mental
processes, including early activity over frontal scalp sites, representation of the array (e.g., bottom left). Neutral
are involved when orienting attention to an internal cues again consisted of the whole square brightening,
representation held in WM. which gave no spatial information. After a random interval
between 500 and 1000 msec, a colored cross (any one of
the eight possible colors; probe stimulus) was presented
at the center of the screen for 100 msec. The task was to
METHODS decide whether the probe stimulus was present or absent
Experiment 1: Predicting Locations in External from the array. When the probe stimulus had been
Versus Internal Space present in the array, the informative pre-cues and retro-
cues predicted its correct location 80% of the time.
Subjects Subjects responded by pressing the left button of the
Ten healthy right-handed (Oldfield, 1971) subjects (age response box if the probe stimulus did appear in the
range 21–36 years, 6 women) took part in the experi- array, and the right button if the probe stimulus did not
ment as paid volunteers. All participants reported being appear in the array.

Griffin and Nobre 1189

 The probe stimulus was present or absent with equal ment as paid volunteers. All participants reported being
(50%) probability. This was true for pre-cue, retro-cue free of neurological disorders. Visual acuity was normal
and neutral trials. All trial types occurred in a random or corrected to normal.
order throughout the experiment. There were 352 trials
in total (160 pre-cue, 160 retro-cue, 32 neutral). Of the
pre-cue trials, 64 were valid (stimulus was in the array, at Stimuli and Task
the cued location), 16 were invalid (stimulus was in the
The task is illustrated in Figure 1B. As before, there
array, at an uncued location), and 80 contained a probe
were three types of trial: pre-cue, retro-cue, and neutral
stimulus that was not in the array. Retro-cue trials
trials. In this experiment the probe stimulus was pre-
contained the same ratio of experimental conditions. Of
sented peripherally, at one of the four locations occu-
the neutral trials, 16 contained a probe that was in the
pied by the array stimuli. The subject’s task was to
array, and 16 contained a probe that was not in the array.
decide whether the probe stimulus matched the item
There were 12 blocks of trials in the experiment, plus one
presented at that location in the array. The informative
additional practice block at the beginning.
pre-cues and retro-cues predicted the location that
would be probed with 80% validity. On neutral trials
Procedure the probe was equally likely to occur at any location.
Subjects responded by pressing the left button of the
Subjects were comfortably seated in a dimly illuminated,
response box if the probe stimulus did match the item
electrically shielded room, facing a computer monitor
at the probed location in the array, and the right button
placed 100 cm in front of them. They were informed, at
if the probe stimulus did not match the item at the
the beginning of the experiment, about the relationship
probed location in the array.
between the cue, array, and probe stimuli. They were
As before, the probability of correct yes or no
asked to maintain fixation on a small cross that was
responses was equal (i.e., 50% of the time the probe
continuously present at the center of the monitor. They
stimulus was the same as the stimulus that had oc-
were instructed to respond as quickly as possible follow-
curred at the probed location in the array, and 50% of
ing probe stimulus onset, while avoiding mistakes. Sub-
the time it was different). This was true for pre-cue,
jects responded with their right hand only.
retro-cue, and neutral trials. In ‘‘no’’ trials, the probe
stimulus was randomly chosen from one of the other
Behavioral Analysis three colors in the array. One constraint was that in
invalid trials, the probe could not be the color from the
Reaction times to probe stimuli and accuracy of perfor-
cued location. All trial types occurred in a random
mance were analyzed by repeated-measures ANOVAs. The
order throughout the experiment. There were 396
first analysis compared valid and invalid trials in which the
trials in total (180 pre-cue, 180 retro-cue, 36 neutral).
probe stimulus was present in the array. This analysis
Of the pre-cue trials, 144 were valid (the probe stim-
tested the factors of cue (pre, retro) and validity (valid,
ulus appeared at the cued location) and 36 were invalid
invalid). The second analysis tested the factors of re-
(the probe stimulus appeared at an uncued location).
sponse (yes, no) and cue (pre, retro, neutral). To evaluate
Retro-cue trials contained the same ratio of experimen-
benefits conferred by valid cues, only valid pre-cue and
tal conditions. Of the neutral trials, 18 contained a
retro-cue trials were considered in this analysis. The third
probe that did match the stimulus that appeared at the
analysis also tested the factors of response (yes, no) and
probed location in the array, and 18 contained a probe
cue (pre, retro, neutral), but considered only invalid pre-
that did not match the stimulus that appeared at the
cues and retro-cues, to assess any attentional costs of
probed location in the array. There were 12 blocks of
invalid cueing. In all analyses, post hoc contrasts were
trials in the experiment, plus one additional practice
carried out to guide interpretation where appropriate.
block at the beginning.

Experiment 2: Predicting Locations in External Procedure

Versus Internal Space (Peripheral Probes)
Eye movements and gaze position were monitored using
Unless stated otherwise, the methods and stimulus an infrared video-based eye tracker with theoretical
parameters used in Experiment 2 were identical to those resolution >0.18 (iView, SMI). Eye movements or devia-
in Experiment 1. tions from central fixation were detected with the eye
tracker using an algorithm that calculated sites and
duration of fixation points during each trial using infra-
Subjects
red tracking. Trials with fixation points further than 18
Ten healthy right-handed (Oldfield, 1971) subjects (age apart from central fixation were excluded from the
range 20–36 years, 4 women) took part in the experi- behavioral analysis.

1190 Journal of Cognitive Neuroscience Volume 15, Number 8

Behavioral Analysis There were an equal number of pre-cue and retro-cue
trials, and they occurred in random order throughout
Reaction times to probe stimuli and accuracy of perfor-
the experiment. To ensure enough data in all relevant
mance were analyzed by repeated-measures ANOVAs,
conditions, subjects participated in two equivalent ex-
in similar fashion to Experiment 1. The first analysis
perimental sessions, separated by at least one week.
compared valid and invalid trials for both pre-cues and
There were 1152 trials in total over the two sessions.
retro-cues. This analysis tested the factors of validity
There were 20 blocks of trials per session, plus one
(valid, invalid) and cue (pre, retro). To assess attentional
additional practice block at the beginning.
benefits, the second analysis tested the factors of
response (yes, no) and cue (pre, retro, neutral) using
only valid pre-cue and retro-cue trials. To assess atten-
Behavioral Analysis
tional costs, the third analysis also tested the factors of
response (yes, no) and cue (pre, retro, neutral), but Reaction times and accuracy were analyzed by repeated-
using only invalid pre-cue and retro-cue trials. In all measures ANOVAs with factors of response (yes, no) and
analyses, post hoc contrasts were carried out to guide cue (pre-cue, retro-cue).
interpretation where appropriate.

ERP Recording
Experiment 3: Imperative Cueing of External
The EEG was recorded continuously from 60 scalp sites
Versus Internal Space
using nonpolarizable tin electrodes mounted on an
Unless stated otherwise, the methods and stimulus elastic cap (Electro-Cap), positioned according to the
parameters used in Experiment 3 are identical to those 10–20 International System (AEEGS, 1991). The mon-
in Experiment 1. tage included 8 midline sites (FPZ, FZ, FCZ, CZ, CPZ, PZ,
POZ, and OZ) and 26 sites over each hemisphere (FP1/
FP2, AF3/AF4, AF7/AF8, F1/F2, F3/F4, F5/F6, F7/F8, FC1/
Subjects
FC2, FC3/FC4, FC5/FC6, FT7/FT8, C1/C2, C3/C4, C5/C6,
Twenty-four healthy right-handed (Oldfield, 1971) sub- T7/T8, CP1/CP2, CP3/CP4, CP5/CP6, TP7/TP8, P1/P2, P3/
jects (age range 20–31 years, 14 women) took part in the P4, P5/P6, P7/P8, PO3/PO4, PO7/PO8, and O1/O2). Ad-
experiment. One subject participated in both Experi- ditional electrodes were used as ground and reference
ments 1 and 3. The experimental methods were nonin- sites. The EEG was referenced to the right mastoid, then
vasive and had ethical approval from the Department of re-referenced off-line to the algebraic average of the
Experimental Psychology, University of Oxford, UK. right and left mastoids. The signal was amplified 20,000
times and digitized at a sampling rate of 250 Hz. Data
were recorded with a band-pass filter of 0.03–100 Hz.
Stimuli and Task
The epoching of ERPs was performed off-line. Epochs
The task is illustrated in Figure 1C. There were only two started 200 msec before and ended 600 msec after cue–
types of trial: pre-cue and retro-cue trials. In this exper- stimulus onset.
iment, the probed item was always present in the array. Horizontal and vertical eye movements were detected
The subject’s task was to decide whether the probe was by recording the horizontal and vertical electrooculo-
presented at the cued location. Subjects responded by gram (HEOG and VEOG) bipolarly with electrodes
pressing the left button of the response box if the probe placed around the eyes. Eye movements were also
stimulus did appear at the cued location, and the right monitored with >0.18 precision using an infrared vid-
button if the probe stimulus did not appear at the cued eo-based eye tracker (iView, SMI).
location. Once again the stimulus array was made up of Epochs containing excessive noise or drift (±100 AV)
four differently colored crosses, however, only four pos- at any electrode between 200 and +600 msec were
sible colors were used for the array (and therefore probe) excluded. Epochs with eye movement artefacts (blinks
stimuli: red, blue, green, and yellow. or saccades) were rejected. Blinks were identified as
As before, the probability of correct yes or no re- large deflections (±50 AV) in the HEOG and VEOG
sponses was equal (i.e., 50% of the time the probe did electrodes. Saccades or breaks in central fixation were
occur at the cued location, and 50% of the time it did detected with the eye tracker using an algorithm that
not). This was true for both pre-cue and retro-cue trials. calculated sites and duration of fixation points during
The possible distribution of colors in the array was fully each trial using infrared tracking as in Experiment 2.
counterbalanced between response and cueing condi- Trials with incorrect behavioral responses were dis-
tions. There were an equal number of trials with all the carded. Trials with RTs faster than 130 msec or slower
potential array types (four different colors, each in a than 1100 msec were regarded as errors and also
different position). The intertrial interval varied random- excluded. To maintain an acceptable signal-to-noise
ly between 1500 and 1800 msec. ratio, subjects with fewer than 40 artefact-free trials

Griffin and Nobre 1191

per condition were excluded from both behavioral and considered significant if they persisted over at least two
ERP analyses. successive time bins in a given region (i.e., 80 msec).

ERP Analysis Acknowledgments

Data analysis was performed using measures of the This study was supported by an award from the James S.
McDonnell Foundation to ACN. We thank Anling Rao for
mean voltage value (mean amplitude) over successive technical assistance and Gianluca Campana for help with
time bins across different scalp regions. Successive time programming.
bins in steps of 40-msec intervals between 0 and
Reprint requests should be sent to A. C. Nobre, Department of
600 msec were used. Five region-specific sets of anal- Experimental Psychology, University of Oxford, South Parks
yses were performed over midline, frontal, central, Road, Oxford, OX1 3UD, UK, or via e-mail: kia.nobre@psy.
lateral, and posterior scalp regions. The midline analysis ox.ac.uk.
included electrode sites FPZ, FZ, CZ, PZ, and OZ. The
frontal analysis used electrodes FP1/2, AF3/4, AF7/8, F3/4, Note
F5/6, and F7/8. The central analysis included sites FC1/2, 1. We thank Steven Luck for pointing this out and for
FC3/4, C1/2, C3/4, CP1/2, and CP3/4. The lateral analysis suggesting Experiment 2 in his signed review.
included electrode sites FC5/6, FT7/8, C5/6, T7/8, CP5/6,
and TP7/8. The posterior analysis used electrode sites:
P3/4, P5/6, P7/8, PO3/4, PO7/8, and O1/2. REFERENCES
The main aim of the experiment was to compare the AEEGS. (1991). American Electroencephalographic Society
processes of orienting attention to a spatial location of guidelines for standard electrode position nomenclature.
an expected perceptual stimulus with orienting atten- Journal of Clinical Neurophysiology, 8, 200–202.
tion to an internal spatial representation held in WM. Awh, E., Anllo Vento, L., & Hillyard, S. A. (2000). The role of
The analysis of ERPs therefore concentrated on compar- spatial selective attention in working memory for locations:
Evidence from event-related potentials. Journal of Cognitive
ing the spatially informative cueing stimuli that occurred Neuroscience, 12, 840–847.
before (pre-cue) or after (retro-cue) the stimulus arrays. Awh, E., & Jonides, J. (2001). Overlapping mechanisms of
These stimuli were physically identical, which afforded attention and spatial working memory. Trends in Cognitive
direct comparison between the two cases. Separate ERP Sciences, 5, 119–126.
averaged waveforms were constructed for the pre-cue Awh, E., Jonides, J., & Reuter Lorenz, P. A. (1998). Rehearsal in
spatial working memory. Journal of Experimental
and retro-cue stimuli in each direction (top-left, top- Psychology: Human Perception and Performance, 24,
right, bottom-left, bottom right). To isolate aspects of 780–790.
neural activity related to attentional orienting, and to Awh, E., Jonides, J., Smith, E. E., Buxton, R. B., Frank, L. R.,
minimize the contribution of general processing differ- Love, T., Wong, E. C., & Gmeindl, L. (1999). Rehearsal in
ences at distinct stages of the trial, each of these spatial working memory: Evidence from neuroimaging.
Psychological Science, 10, 433–437.
conditions then had the ERP waveform for the ‘‘neutral’’ Baddeley, A. D. (1993). Working memory or working attention?
cue condition at the equivalent phase of the trial (whole In A. D. Baddeley & L. Weiskrantz (Eds.), Attention:
square flashing) subtracted from them. To ensure there Selection, awareness, and control: A tribute to Donald
were sufficient trials in each condition, the attend top- Broadbent. (pp. 152–170). New York: Oxford University
left and attend bottom-left conditions were averaged to Press.
Beschin, N., Cocchini, G., Della Sala, S., & Logie, R. H. (1997).
form an attend-left condition. Similarly, attend top-right What the eyes perceive, the brain ignores: A case of pure
and attend bottom-right were averaged to form an unilateral representational neglect. Cortex, 33, 3–26.
attend-right condition. There were therefore four con- Bisiach, E., & Luzzatti, C. (1978). Unilateral neglect of
ditions: pre-cue attend left, pre-cue attend right, retro- representational space. Cortex, 14, 129–133.
cue attend left, and retro-cue attend right. Bosch, V., Mecklinger, A., & Friederici, A. D. (2001). Slow
cortical potentials during retention of object, spatial, and
Differences in mean amplitudes were assessed by verbal information. Brain Research: Cognitive Brain
repeated-measures ANOVAs, using the Greenhouse– Research, 10, 219–237.
Geisser epsilon correction for nonsphericity where ap- Chafee, M. V., & Goldman Rakic, P. S. (2000). Inactivation of
propriate (Jennings & Wood, 1976). Only the corrected parietal and prefrontal cortex reveals interdependence of
probability values and degrees of freedom are reported. neural activity during memory-guided saccades. Journal
of Neurophysiology, 83, 1550–1566.
The ANOVA factors were side of attention (left, right), Chelazzi, L. (1999). Serial attention mechanisms in visual
cue (pre-cue, retro-cue), hemisphere (left, right), and search: A critical look at the evidence. Psychological
electrode (six sites). For the midline analysis, there were Research, 62, 195–219.
five electrode sites and no hemisphere factor. Effects of Corbetta, M., Kincade, J. M., Ollinger, J. M., McAvoy, M. P., &
interest only included main effects or interactions involv- Shulman, G. L. (2000). Voluntary orienting is dissociated
from target detection in human posterior parietal cortex.
ing side of attention and cue factors. Because of the risk Nature Neuroscience, 3, 292–297.
of false-positive effects in the multiple interrelated com- Coslett, H. B. (1997). Neglect in vision and visual imagery: A
parisons in the regional analyses, results were only double dissociation. Brain, 120, 1163–1171.

1192 Journal of Cognitive Neuroscience Volume 15, Number 8

Desimone, R., & Duncan, J. (1995). Neural mechanisms of in human visual short-term memory: An event-related brain
selective visual attention. Annual Review of Neuroscience, potential study. Neuroscience Letters, 268, 65–68.
18193–18222. Kosslyn, S. M., Ganis, G., & Thompson, W. L. (2001). Neural
Desmedt, J. E. (1980). P300 in serial tasks: An essential foundations of imagery. Nature Reviews: Neuroscience, 2,
post-decision closure mechanism. Progress in Brain 635–642.
Research, 54, 682–686. LaBar, K. S., Gitelman, D. R., Parrish, T. B., & Mesulam, M.
Donchin, E., & Coles, M. G. (1988). Is the P300 component a (1999). Neuroanatomic overlap of working memory and
manifestation of context updating? Behavioral and Brain spatial attention networks: A functional MRI comparison
Sciences, 11, 357–427. within subjects. Neuroimage, 10, 695–704.
Downing, C. J. (1988). Expectancy and visual–spatial attention: Levy, R., & Goldman Rakic, P. S. (2000). Segregation
Effects on perceptual quality. Journal of Experimental of working memory functions within the
Psychology: Human Perception and Performance, 14, dorsolateral prefrontal cortex. Experimental Brain
188–202. Research, 133, 23–32.
Downing, P. E. (2000). Interactions between visual working Luck, S. J., Hillyard, S. A., Mouloua, M., Woldorff, M. G., Clark,
memory and selective attention. Psychological Science, 11, V. P., & Hawkins, H. L. (1994). Effects of spatial cuing on
467–473. luminance detectability: Psychophysical and
Eimer, M., Velzen, J., & Driver, J. (2002). Cross-modal electrophysiological evidence for early selection.
interactions between audition, touch and vision in Journal of Experimental Psychology: Human Perception
endogenous spatial attention: ERP evidence on preparatory and Performance, 20, 887–904.
states and sensory modulations. Journal of Cognitive Luck, S. J., & Vogel, E. K. (1997). The capacity of visual working
Neuroscience, 14, 254–271. memory for features and conjunctions. Nature, 390,
Farah, M. J. (1985). Psychophysical evidence for a shared 279–281.
representational medium for mental images and percepts. McCarthy, G. (1995). Functional neuroimaging of memory. The
Journal of Experimental Psychology: General, 114, 91–103. Neuroscientist, 1, 155–163.
Farah, M. J., Peronnet, F., Gonon, M. A., & Giard, M. H. (1988). Mecklinger, A. (1998). On the modularity of recognition
Electrophysiological evidence for a shared representational memory for object form and spatial location: A
medium for visual images and visual percepts. Journal of topographic ERP analysis. Neuropsychologia, 36,
Experimental Psychology: General, 117, 248–257. 441–460.
Graziano, M. S., Hu, X. T., & Gross, C. G. (1997). Coding the Mesulam, M. (1999). Spatial attention and neglect: Parietal,
locations of objects in the dark. Science, 277, 239–241. frontal and cingulate contributions to the mental
Guariglia, C., Padovani, A., Pantano, P., & Pizzamiglio, L. (1993). representation and attentional targeting of salient
Unilateral neglect restricted to visual imagery. Nature, 364, extrapersonal events. Philosophical Transactions of the
235–237. Royal Society of London: Series B, Biological Sciences, 354,
Harter, M. R., Miller, S. L., Price, N. J., LaLonde, M. E., & Keyes, 1325–1346.
A. L. (1989). Neural processes involved in directing attention. Mesulam, M. M. (1981). A cortical network for directed
Journal of Cognitive Neuroscience, 1, 223–237. attention and unilateral neglect. Neurological Progress, 10,
Hawkins, H. L., Hillyard, S. A., Luck, S. J., Mouloua, M., 304–325.
Downing, C. J., & Woodward, D. P. (1990). Visual attention Müller, H. J., & Findlay, J. M. (1987). Sensitivity and criterion
modulates signal detectability. Journal of Experimental effects in the spatial cuing of visual attention. Perception
Psychology: Human Perception and Performance, 16, and Psychophysics, 42, 383–399.
802–811. Müller, H. J., & Rabbit, P. M. A. (1989). Reflexive and voluntary
Hopf, J. M., & Mangun, G. R. (2000). Shifting visual attention in orienting of visual attention: Time course of activation and
space: An electrophysiological analysis using high spatial resistance to interruption. Journal of Experimental
resolution mapping. Clinical Neurophysiology, 111, Psychology: Human Perception and Performance, 15,
1241–1257. 315–330.
Hopfinger, J. B., Buonocore, M. H., & Mangun, G. R. (2000). Nobre, A. C. (2001). The attentive homunculus: Now you see it,
The neural mechanisms of top-down attentional control. now you don’t. Neuroscience and Biobehavioral Reviews,
Nature Neuroscience, 3, 284–291. 25, 477–496.
Ishai, A., & Sagi, D. (1995). Common mechanisms of visual Nobre, A. C., Coull, J. T., Vandenberghe, R., Maquet, P., Frith,
imagery and perception. Science, 268, 1772–1774. C., & Mesulam, M. (2002). Directing spatial attention in
Ishai, A., Ungerleider, L. G., & Haxby, J. V. (2000). Distributed mental representations. Paper presented at the
neural systems for the generation of visual images. Neuron, NeuroImage Human Brain Mapping 2002 Meeting,
28, 979–990. Sendai, Japan.
Jennings, J. R., & Wood, C. C. (1976). Letter: The Nobre, A. C., Sebestyen, G. N., & Miniussi, C. (2000).
epsilon-adjustment procedure for repeated-measures The dynamics of shifting visuospatial attention revealed
analyses of variance. Psychophysiology, 13, 277–278. by event-related potentials. Neuropsychologia, 38, 964–974.
Jonides, J. (1981). Voluntary versus automatic control over the O’Craven, K. M., & Kanwisher, N. (2000). Mental imagery of
mind’s eye’s movement. In J. B. Long & A. D. Baddeley faces and places activates corresponding stimulus-specific
(Eds.), Attention and performance, IX (pp. 187–203). brain regions. Journal of Cognitive Neuroscience, 12,
Hillsdale, NJ: Erlbaum. 1013–1023.
Kastner, S., Pinsk, M. A., De Weerd, P., Desimone, R., & Oldfield, R. C. (1971). The assessment and analysis of
Ungerleider, L. G. (1999). Increased activity in human visual handedness: The Edinburgh inventory. Neuropsychologia,
cortex during directed attention in the absence of visual 9, 97–113.
stimulation. Neuron, 22, 751–761. Ortigue, S., Viaud Delmon, I., Annoni, J. M., Landis, T., Michel,
Kinchla, R. A., Chen, Z., & Evert, D. (1995). Precue effects in C., Blanke, O., Vuilleumier, P., & Mayer, E. (2001). Pure
visual search: Data or resource limited? Perception and representational neglect after right thalamic lesion. Annals
Psychophysics, 57, 441–450. of Neurology, 50, 401–404.
Klaver, P., Smid, H. G., & Heinze, H. J. (1999). Representations Owen, A. M. (2000). The role of the lateral frontal cortex

Griffin and Nobre 1193

 in mnemonic processing: The contribution of functional Sperling, G. (1960). The information available in brief visual
neuroimaging. Experimental Brain Research, 133, 33–43. presentations. Psychological Monographs, 74, 29.
Pashler, H., & Shiu, L. P. (1999). Do images involuntarily trigger Treisman, A. M., & Gelade, G. (1980). A feature-integration
search? A test of Pillsbury’s hypothesis. Psychonomic theory of attention. Cognitive Psychology, 12, 97–136.
Bulletin and Review, 6, 445–448. Velzen, J., & Eimer, M. (submitted). Do early posterior ERP
Petrides, M. (2000). The role of the mid-dorsolateral prefrontal components elicited during spatial orienting reflect the
cortex in working memory. Experimental Brain Research, control of attentional shifts?
133, 44–54. Vogel, E. K., & Luck, S. J. (2000). The visual N1 component as
Pollmann, S., & von Cramon, D. Y. (2000). Object working an index of a discrimination process. Psychophysiology, 37,
memory and visuospatial processing: Functional 190–203.
neuroanatomy analyzed by event-related fMRI. Vogel, E. K., Woodman, G. F., & Luck, S. J. (2001). Storage of
Experimental Brain Research, 133, 12–22. features, conjunctions and objects in visual working
Posner, M. I. (1980). Orienting of attention. Quarterly Journal memory. Journal of Experimental Psychology: Human
of Experimental Psychology, 32, 3–25. Perception and Performance, 27, 92–114.
Ruchkin, D. S., Johnson, R., Grafman, J., Canoune, H., & Ritter, Wheeler, M. E., & Treisman, A. M. (2002). Binding in
W. (1997). Multiple visuospatial working memory buffers: short-term visual memory. Journal of Experimental
Evidence from spatiotemporal patterns of brain activity. Psychology: General, 131, 48–64.
Neuropsychologia, 35, 195–209. Wolfe, J. M. (1994). Guided search 2.0: A revised model of visual
Schmidt, B. K., Vogel, E. K., Woodman, G. F., & Luck, S. J. search. Psychonomic Bulletin and Review, 1, 202–238.
(2002). Voluntary and automatic attentional control of visual Yamaguchi, S., Tsuchiya, H., & Kobayashi, S. (1994).
working memory. Perception and Psychophysics, 64, Electroencephalographic activity associated with shifts of
754–763. visuospatial attention. Brain, 117, 553–562.

1194 Journal of Cognitive Neuroscience Volume 15, Number 8

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42. Paul Zerr, Surya Gayet, Kees Mulder, Yaïr Pinto, Ilja Sligte, Stefan Van der Stigchel. 2017. Remapping high-capacity, pre-attentive,
fragile sensory memory. Scientific Reports 7:1. . [Crossref]
43. Giovanni Mento. 2017. The role of the P3 and CNV components in voluntary and automatic temporal orienting: A high spatial-
resolution ERP study. Neuropsychologia 107, 31-40. [Crossref]
44. Jason Rajsic, Henry Liu, Jay Pratt. 2017. Eye movements can cause item-specific visual recognition advantages. Visual Cognition
25:9-10, 903-912. [Crossref]
45. Freek van Ede. 2017. Mnemonic and attentional roles for states of attenuated alpha oscillations in perceptual working memory:
a review. European Journal of Neuroscience 5. . [Crossref]
46. Ashley F. Curtis, Gary R. Turner, Norman W. Park, Susan J. E. Murtha. 2017. Improving visual spatial working memory in
younger and older adults: effects of cross-modal cues. Aging, Neuropsychology, and Cognition 1, 1-20. [Crossref]
47. Daniel Schneider, Anna Barth, Edmund Wascher. 2017. On the contribution of motor planning to the retroactive cuing benefit
in working memory: Evidence by mu and beta oscillatory activity in the EEG. NeuroImage 162, 73-85. [Crossref]
48. Claudia Poch, Almudena Capilla, José Antonio Hinojosa, Pablo Campo. 2017. Selection within working memory based on a color
retro-cue modulates alpha oscillations. Neuropsychologia 106, 133-137. [Crossref]
49. Bo-Cheng Kuo, Chun-Hui Li, Szu-Hung Lin, Sheng-Hung Hu, Yei-Yu Yeh. 2017. Top-down modulation of alpha power and
pattern similarity for threatening representations in visual short-term memory. Neuropsychologia 106, 21-30. [Crossref]
50. Mario Villena-González, Cristóbal Moënne-Loccoz, Rodrigo A. Lagos, Luz M. Alliende, Pablo Billeke, Francisco Aboitiz,
Vladimir López, Diego Cosmelli. 2017. Attending to the heart is associated with posterior alpha band increase and a reduction in
sensitivity to concurrent visual stimuli. Psychophysiology 54:10, 1483-1497. [Crossref]
51. Claudia Poch, Luis Carretie, Pablo Campo. 2017. A dual mechanism underlying alpha lateralization in attentional orienting to
mental representation. Biological Psychology 128, 63-70. [Crossref]
52. Matthew K. Robison, Nash Unsworth. 2017. Variation in the use of cues to guide visual working memory. Attention, Perception,
& Psychophysics 79:6, 1652-1665. [Crossref]
53. Jason Rajsic, Natasha E. Ouslis, Daryl E. Wilson, Jay Pratt. 2017. Looking sharp: Becoming a search template boosts precision
and stability in visual working memory. Attention, Perception, & Psychophysics 79:6, 1643-1651. [Crossref]
54. Nicholas D’Aloisio-Montilla. 2017. Imagery and overflow: We see more than we report. Philosophical Psychology 30:5, 545-570.
[Crossref]
55. 현현현. 2017. A Review of methodological limitations of change detection task and their theoretical implications for studying visual
working memory. Korean Journal of Cognitive and Biological Psychology 29:3, 287-313. [Crossref]
56. Diego Mendoza-Halliday, Julio C. Martinez-Trujillo. 2017. Neuronal population coding of perceived and memorized visual
features in the lateral prefrontal cortex. Nature Communications 8, 15471. [Crossref]
57. Nicholas E. Myers, Mark G. Stokes, Anna C. Nobre. 2017. Prioritizing Information during Working Memory: Beyond Sustained
Internal Attention. Trends in Cognitive Sciences 21:6, 449-461. [Crossref]
58. Michael J Wolff, Janina Jochim, Elkan G Akyürek, Mark G Stokes. 2017. Dynamic hidden states underlying working-memory-
guided behavior. Nature Neuroscience 20:6, 864-871. [Crossref]
59. Tobias Katus, Anna Grubert, Martin Eimer1. 2017. Intermodal Attention Shifts in Multimodal Working Memory. Journal of
Cognitive Neuroscience 29:4, 628-636. [Abstract] [Full Text] [PDF] [PDF Plus]
60. Anna Heuer, J. Douglas Crawford, Anna Schub?. 2017. Action relevance induces an attentional weighting of representations in
visual working memory. Memory & Cognition 45:3, 413-427. [Crossref]
61. Marcel Niklaus, Anna C. Nobre, Freek van Ede. 2017. Feature-based attentional weighting and spreading in visual working
memory. Scientific Reports 7, 42384. [Crossref]
62. Paula Pazo-Álvarez, Adriana Roca-Fernández, Francisco-Javier Gutiérrez-Domínguez, Elena Amenedo. 2017. Attentional
Modulation of Change Detection ERP Components by Peripheral Retro-Cueing. Frontiers in Human Neuroscience 11. . [Crossref]
63. Nash Unsworth, Matthew K. Robison. 2017. Pupillary correlates of covert shifts of attention during working memory maintenance.
Attention, Perception, & Psychophysics 12. . [Crossref]
64. Jiaxin Peng, Sam C. C. Chan, Bolton K. H. Chau, Qiuhua Yu, Chetwyn C. H. Chan. 2017. Salience of Somatosensory Stimulus
Modulating External-to-Internal Orienting Attention. Frontiers in Human Neuroscience 11. . [Crossref]
65. Chaoxiong Ye, Zhonghua Hu, Tapani Ristaniemi, Maria Gendron, Qiang Liu. 2016. Retro-dimension-cue benefit in visual
working memory. Scientific Reports 6:1. . [Crossref]
66. Jason Rajsic, Sol Z. Sun, Lauren Huxtable, Jay Pratt, Susanne Ferber. 2016. Pop-out and pop-in: Visual working memory
advantages for unique items. Psychonomic Bulletin & Review 23:6, 1787-1793. [Crossref]
67. Benchi Wang, Chuyao Yan, Zhiguo Wang, Christian N. L. Olivers, Jan Theeuwes. 2016. Adverse orienting effects on visual
working memory encoding and maintenance. Psychonomic Bulletin & Review 5. . [Crossref]
68. Bo-Cheng Kuo. 2016. Selection History Modulates Working Memory Capacity. Frontiers in Psychology 7. . [Crossref]
69. Benjamin J. Tamber-Rosenau, René Marois. 2016. Central attention is serial, but midlevel and peripheral attention are parallel
—A hypothesis. Attention, Perception, & Psychophysics 78:7, 1874-1888. [Crossref]
70. Alessandra S. Souza, Klaus Oberauer. 2016. In search of the focus of attention in working memory: 13 years of the retro-cue
effect. Attention, Perception, & Psychophysics 78:7, 1839-1860. [Crossref]
71. Anna Heuer, Anna Schubö, J. D. Crawford. 2016. Different Cortical Mechanisms for Spatial vs. Feature-Based Attentional
Selection in Visual Working Memory. Frontiers in Human Neuroscience 10. . [Crossref]
72. Muhammet Ikbal Sahan, Tom Verguts, Carsten Nicolas Boehler, Gilles Pourtois, Wim Fias. 2016. Paying attention to working
memory: Similarities in the spatial distribution of attention in mental and physical space. Psychonomic Bulletin & Review 23:4,
1190-1197. [Crossref]
73. Thomas C. Sprague, Edward F. Ester, John T. Serences. 2016. Restoring Latent Visual Working Memory Representations in
Human Cortex. Neuron 91:3, 694-707. [Crossref]
74. Daniel Schneider, Christine Mertes, Edmund Wascher. 2016. The time course of visuo-spatial working memory updating revealed
by a retro-cuing paradigm. Scientific Reports 6:1. . [Crossref]
75. Filiz Gözenman, Marian E. Berryhill. 2016. Working memory capacity differentially influences responses to tDCS and HD-tDCS
in a retro-cue task. Neuroscience Letters 629, 105-109. [Crossref]
76. Michael Pilling, Doug J. K. Barrett. 2016. Dimension-based attention in visual short-term memory. Memory & Cognition 44:5,
740-749. [Crossref]
77. Sarah Reaves, Jonathan Strunk, Shekinah Phillips, Paul Verhaeghen, Audrey Duarte. 2016. The lasting memory enhancements
of retrospective attention. Brain Research 1642, 226-237. [Crossref]
78. Atanaska Nikolova, Bill Macken. 2016. The objects of visuospatial short-term memory: Perceptual organization and change
detection. Quarterly Journal of Experimental Psychology 69:7, 1426-1437. [Crossref]
79. Zampeta Kalogeropoulou, Akshay V. Jagadeesh, Sven Ohl, Martin Rolfs. 2016. Setting and changing feature priorities in visual
short-term memory. Psychonomic Bulletin & Review 74. . [Crossref]
80. Haluk Öğmen, Michael H. Herzog. 2016. A New Conceptualization of Human Visual Sensory-Memory. Frontiers in Psychology
7. . [Crossref]
81. Anna Heuer, Anna Schubö. 2016. Feature-based and spatial attentional selection in visual working memory. Memory & Cognition
44:4, 621-632. [Crossref]
82. Markus Janczyk, Heiko Reuss. 2016. Only pre-cueing but no retro-cueing effects emerge with masked arrow cues. Consciousness
and Cognition 42, 93-100. [Crossref]
83. Anna Heuer, Anna Schubö. 2016. The Focus of Attention in Visual Working Memory: Protection of Focused Representations
and Its Individual Variation. PLOS ONE 11:4, e0154228. [Crossref]
84. Sean James Fallon, Nahid Zokaei, Masud Husain. 2016. Causes and consequences of limitations in visual working memory. Annals
of the New York Academy of Sciences 1369:1, 40-54. [Crossref]
85. Robert M. Mok, Nicholas E. Myers, George Wallis, Anna Christina Nobre. 2016. Behavioral and Neural Markers of Flexible
Attention over Working Memory in Aging. Cerebral Cortex 26:4, 1831-1842. [Crossref]
86. Marcel Gressmann, Markus Janczyk. 2016. The (Un)Clear Effects of Invalid Retro-Cues. Frontiers in Psychology 7. . [Crossref]
87. Amanda L. Gilchrist, Audrey Duarte, Paul Verhaeghen. 2016. Retrospective cues based on object features improve visual working
memory performance in older adults. Aging, Neuropsychology, and Cognition 23:2, 184-195. [Crossref]
88. Louis Thibault, Ronald van den Berg, Patrick Cavanagh, Claire Sergent. 2016. Retrospective Attention Gates Discrete Conscious
Access to Past Sensory Stimuli. PLOS ONE 11:2, e0148504. [Crossref]
89. Nina M. Hanning, Donatas Jonikaitis, Heiner Deubel, Martin Szinte. 2016. Oculomotor selection underlies feature retention in
visual working memory. Journal of Neurophysiology 115:2, 1071-1076. [Crossref]
90. Stefan Berti. 2016. Switching Attention Within Working Memory is Reflected in the P3a Component of the Human Event-
Related Brain Potential. Frontiers in Human Neuroscience 9. . [Crossref]
91. Jie Li. 2016. Dissociable loss of the representations in visual short-term memory. The Journal of General Psychology 143:1, 1-15.
[Crossref]
92. Zaifeng Gao, Qiyang Gao, Ning Tang, Rende Shui, Mowei Shen. 2016. Organization principles in visual working memory:
Evidence from sequential stimulus display. Cognition 146, 277-288. [Crossref]
93. Weronika Wojtak, Stephen Coombes, Estela Bicho, Wolfram Erlhagen. Combining Spatial and Parametric Working Memory in
a Dynamic Neural Field Model 411-418. [Crossref]
94. Jonni Hirvonen, Satu Palva. 2016. Cortical localization of phase and amplitude dynamics predicting access to somatosensory
awareness. Human Brain Mapping 37:1, 311-326. [Crossref]
95. Annelinde R. E. Vandenbroucke, Ilja G. Sligte, Jade G. de Vries, Michael X. Cohen, Victor A. F. Lamme. 2015. Neural Correlates
of Visual Short-term Memory Dissociate between Fragile and Working Memory Representations. Journal of Cognitive Neuroscience
27:12, 2477-2490. [Abstract] [Full Text] [PDF] [PDF Plus]
96. 현현현. 2015. A Review of the Debates between Fixed-Resolution Slot and Flexible-Resource Models. Korean Journal of Cognitive
Science 26:4, 453-481. [Crossref]
97. Andria Shimi, Anna Christina Nobre, Gaia Scerif. 2015. ERP markers of target selection discriminate children with high vs. low
working memory capacity. Frontiers in Systems Neuroscience 9. . [Crossref]
 98. George Wallis, Mark Stokes, Helena Cousijn, Mark Woolrich, Anna Christina Nobre. 2015. Frontoparietal and Cingulo-opercular
Networks Play Dissociable Roles in Control of Working Memory. Journal of Cognitive Neuroscience 27:10, 2019-2034. [Abstract]
[Full Text] [PDF] [PDF Plus]
99. Rachel N. Newsome, Audrey Duarte, Carson Pun, Victoria M. Smith, Susanne Ferber, Morgan D. Barense. 2015. A retroactive
spatial cue improved VSTM capacity in mild cognitive impairment and medial temporal lobe amnesia but not in healthy older
adults. Neuropsychologia 77, 148-157. [Crossref]
100. Eren Gunseli, Dirk van Moorselaar, Martijn Meeter, Christian N. L. Olivers. 2015. The reliability of retro-cues determines the
fate of noncued visual working memory representations. Psychonomic Bulletin & Review 22:5, 1334-1341. [Crossref]
101. Daniel Schneider, Christine Mertes, Edmund Wascher. 2015. On the fate of non-cued mental representations in visuo-spatial
working memory: Evidence by a retro-cuing paradigm. Behavioural Brain Research 293, 114-124. [Crossref]
102. Ashleigh M. Maxcey, Keisuke Fukuda, Won S. Song, Geoffrey F. Woodman. 2015. Using electrophysiology to demonstrate that
cueing affects long-term memory storage over the short term. Psychonomic Bulletin & Review 22:5, 1349-1357. [Crossref]
103. Matthew R. Johnson, Gregory McCarthy, Kathleen A. Muller, Samuel N. Brudner, Marcia K. Johnson. 2015. Electrophysiological
Correlates of Refreshing: Event-related Potentials Associated with Directing Reflective Attention to Face, Scene, or Word
Representations. Journal of Cognitive Neuroscience 27:9, 1823-1839. [Abstract] [Full Text] [PDF] [PDF Plus]
104. Tal Makovski, Yoni Pertzov. 2015. Attention and memory protection: Interactions between retrospective attention cueing and
interference. Quarterly Journal of Experimental Psychology 68:9, 1735-1743. [Crossref]
105. Ian A. Clark, Clare E. Mackay. 2015. Mental Imagery and Post-Traumatic Stress Disorder: A Neuroimaging and Experimental
Psychopathology Approach to Intrusive Memories of Trauma. Frontiers in Psychiatry 6. . [Crossref]
106. Michi Matsukura, Shaun P. Vecera. 2015. Selection of multiple cued items is possible during visual short-term memory
maintenance. Attention, Perception, & Psychophysics 77:5, 1625-1646. [Crossref]
107. Mark G. Stokes. 2015. ‘Activity-silent’ working memory in prefrontal cortex: a dynamic coding framework. Trends in Cognitive
Sciences 19:7, 394-405. [Crossref]
108. Joshua J. LaRocque, Adam S. Eichenbaum, Michael J. Starrett, Nathan S. Rose, Stephen M. Emrich, Bradley R. Postle. 2015. The
short- and long-term fates of memory items retained outside the focus of attention. Memory & Cognition 43:3, 453-468. [Crossref]
109. Bradley R. Postle. Activation and Information in Working Memory Research 21-43. [Crossref]
110. Nicholas E. Myers, Lena Walther, George Wallis, Mark G. Stokes, Anna C. Nobre. 2015. Temporal Dynamics of Attention
during Encoding versus Maintenance of Working Memory: Complementary Views from Event-related Potentials and Alpha-band
Oscillations. Journal of Cognitive Neuroscience 27:3, 492-508. [Abstract] [Full Text] [PDF] [PDF Plus]
111. Alessandra S. Souza, Laura Rerko, Klaus Oberauer. 2015. Refreshing memory traces: thinking of an item improves retrieval from
visual working memory. Annals of the New York Academy of Sciences 1339:1, 20-31. [Crossref]
112. Dirk van Moorselaar, Elisa Battistoni, Jan Theeuwes, Christian N. L. Olivers. 2015. Rapid influences of cued visual memories
on attentional guidance. Annals of the New York Academy of Sciences 1339:1, 1-10. [Crossref]
113. Qi Li, Jun Saiki. 2015. Different effects of color-based and location-based selection on visual working memory. Attention,
Perception, & Psychophysics 77:2, 450-463. [Crossref]
114. Charalampos Papadimitriou, Afreen Ferdoash, Lawrence H. Snyder. 2015. Ghosts in the machine: memory interference from the
previous trial. Journal of Neurophysiology 113:2, 567-577. [Crossref]
115. Dirk van Moorselaar, Eren Gunseli, Jan Theeuwes, Christian N. L. Olivers. 2015. The time course of protecting a visual memory
representation from perceptual interference. Frontiers in Human Neuroscience 8. . [Crossref]
116. Andria Shimi, Mark W. Woolrich, Dante Mantini, Duncan E. Astle. 2015. Memory load modulates graded changes in distracter
filtering. Frontiers in Human Neuroscience 8. . [Crossref]
117. Jessica L. Holt, Jean-François Delvenne. 2015. A bilateral advantage for maintaining objects in visual short term memory. Acta
Psychologica 154, 54-61. [Crossref]
118. Tobias Katus, Søren K. Andersen. The Role of Spatial Attention in Tactile Short-Term Memory 275-292. [Crossref]
119. Nahid Zokaei, Shen Ning, Sanjay Manohar, Eva Feredoes, Masud Husain. 2014. Flexibility of representational states in working
memory. Frontiers in Human Neuroscience 8. . [Crossref]
120. Alessandra S. Souza, Laura Rerko, Hsuan-Yu Lin, Klaus Oberauer. 2014. Focused attention improves working memory:
implications for flexible-resource and discrete-capacity models. Attention, Perception, & Psychophysics 76:7, 2080-2102. [Crossref]
121. Rob H.J. Van der Lubbe, Carsten Bundt, Elger L. Abrahamse. 2014. Internal and external spatial attention examined with
lateralized EEG power spectra. Brain Research 1583, 179-192. [Crossref]
122. Qi Li, Jun Saiki. 2014. The effects of sequential attention shifts within visual working memory. Frontiers in Psychology 5. .
[Crossref]
123. Sohee Park, Diane C. Gooding. 2014. Working memory impairment as an endophenotypic marker of a schizophrenia diathesis.
Schizophrenia Research: Cognition 1:3, 127-136. [Crossref]
124. Anastasia Kiyonaga, Tobias Egner. 2014. Resource-sharing between internal maintenance and external selection modulates
attentional capture by working memory content. Frontiers in Human Neuroscience 8. . [Crossref]
125. Jeroen D. Silvis, Kimron L. Shapiro. 2014. Spatiotemporal configuration of memory arrays as a component of VWM
representations. Visual Cognition 22:7, 948-962. [Crossref]
126. Bo-Cheng Kuo, Mark G. Stokes, Alexandra M. Murray, Anna Christina Nobre. 2014. Attention Biases Visual Activity in Visual
Short-term Memory. Journal of Cognitive Neuroscience 26:7, 1377-1389. [Abstract] [Full Text] [PDF] [PDF Plus]
127. Markus Janczyk, Wilfried Kunde. 2014. The role of effect grouping in free-choice response selection. Acta Psychologica 150, 49-54.
[Crossref]
128. Laura Rerko, Alessandra S. Souza, Klaus Oberauer. 2014. Retro-cue benefits in working memory without sustained focal attention.
Memory & Cognition 42:5, 712-728. [Crossref]
129. Claudia Poch, Pablo Campo, Gareth R. Barnes. 2014. Modulation of alpha and gamma oscillations related to retrospectively
orienting attention within working memory. European Journal of Neuroscience 40:2, 2399-2405. [Crossref]
130. Duncan E. Astle, Hannah Harvey, Mark Stokes, Hamid Mohseni, Anna C. Nobre, Gaia Scerif. 2014. Distinct neural mechanisms
of individual and developmental differences in VSTM capacity. Developmental Psychobiology 56:4, 601-610. [Crossref]
131. Kristina C. Backer, Claude Alain. 2014. Attention to memory: orienting attention to sound object representations. Psychological
Research 78:3, 439-452. [Crossref]
132. Helen C. Lückmann, Heidi I.L. Jacobs, Alexander T. Sack. 2014. The cross-functional role of frontoparietal regions in cognition:
internal attention as the overarching mechanism. Progress in Neurobiology 116, 66-86. [Crossref]
133. Andria Shimi, Bo-Cheng Kuo, Duncan E. Astle, Anna C. Nobre, Gaia Scerif. 2014. Age Group and Individual Differences
in Attentional Orienting Dissociate Neural Mechanisms of Encoding and Maintenance in Visual STM. Journal of Cognitive
Neuroscience 26:4, 864-877. [Abstract] [Full Text] [PDF] [PDF Plus]
134. Markus Janczyk, Marian E. Berryhill. 2014. Orienting attention in visual working memory requires central capacity: Decreased
retro-cue effects under dual-task conditions. Attention, Perception, & Psychophysics 76:3, 715-724. [Crossref]
135. Andria Shimi, Anna C. Nobre, Duncan Astle, Gaia Scerif. 2014. Orienting Attention Within Visual Short-Term Memory:
Development and Mechanisms. Child Development 85:2, 578-592. [Crossref]
136. T. Katus, S. K. Andersen, M. M. Muller. 2014. Common Mechanisms of Spatial Attention in Memory and Perception: A Tactile
Dual-Task Study. Cerebral Cortex 24:3, 707-718. [Crossref]
137. Arezoo Pooresmaeili, Dominik R. Bach, Raymond J. Dolan. 2014. The effect of visual salience on memory-based choices. Journal
of Neurophysiology 111:3, 481-487. [Crossref]
138. Bo-Cheng Kuo, Duncan E. Astle. 2014. Neural Mechanisms by Which Attention Modulates the Comparison of Remembered
and Perceptual Representations. PLoS ONE 9:1, e86666. [Crossref]
139. Marcin Leszczyński, Agnieszka Wykowska, Jairo Perez-Osorio, Hermann J. Müller. 2013. Deployment of Spatial Attention
towards Locations in Memory Representations. An EEG Study. PLoS ONE 8:12, e83856. [Crossref]
140. A. Hollingworth, S. Hwang. 2013. The relationship between visual working memory and attention: retention of precise colour
information in the absence of effects on perceptual selection. Philosophical Transactions of the Royal Society B: Biological Sciences
368:1628, 20130061-20130061. [Crossref]
141. 현현현, 현현현. 2013. The Fidelity of Representations in Visual Working Memory Assessed by a Consecutive-Change Detection Task.
Korean Journal of Cognitive and Biological Psychology 25:3, 293-309. [Crossref]
142. Davide Rigoni, Marcel Brass, Clémence Roger, Franck Vidal, Giuseppe Sartori. 2013. Top-down modulation of brain activity
underlying intentional action and its relationship with awareness of intention: an ERP/Laplacian analysis. Experimental Brain
Research 229:3, 347-357. [Crossref]
143. Angela A. Manginelli, Nadine Langer, Diana Klose, Stefan Pollmann. 2013. Contextual cueing under working memory load:
Selective interference of visuospatial load with expression of learning. Attention, Perception, & Psychophysics 75:6, 1103-1117.
[Crossref]
144. Ryan T. Tanoue, Kevin T. Jones, Dwight J. Peterson, Marian E. Berryhill. 2013. Differential Frontal Involvement in Shifts of
Internal and Perceptual Attention. Brain Stimulation 6:4, 675-682. [Crossref]
145. Melonie Williams, Sang W. Hong, Min-Suk Kang, Nancy B. Carlisle, Geoffrey F. Woodman. 2013. The benefit of forgetting.
Psychonomic Bulletin & Review 20:2, 348-355. [Crossref]
146. Alexandra M. Murray, Anna C. Nobre, Ian A. Clark, André M. Cravo, Mark G. Stokes. 2013. Attention Restores Discrete Items
to Visual Short-Term Memory. Psychological Science 24:4, 550-556. [Crossref]
147. Angela A. Manginelli, Florian Baumgartner, Stefan Pollmann. 2013. Dorsal and ventral working memory-related brain areas
support distinct processes in contextual cueing. NeuroImage 67, 363-374. [Crossref]
148. Selene Cansino, Daniela Guzzon, Clara Casco. 2013. Effects of interference control on visuospatial working memory. Journal of
Cognitive Psychology 25:1, 51-63. [Crossref]
149. Claire Sergent, Valentin Wyart, Mariana Babo-Rebelo, Laurent Cohen, Lionel Naccache, Catherine Tallon-Baudry. 2013. Cueing
Attention after the Stimulus Is Gone Can Retrospectively Trigger Conscious Perception. Current Biology 23:2, 150-155. [Crossref]
150. Aki Kondo, Jun Saiki. 2012. Feature-Specific Encoding Flexibility in Visual Working Memory. PLoS ONE 7:12, e50962.
[Crossref]
151. Cheng-Ta Yang, Philip Tseng, Kuan-Yao Huang, Yei-Yu Yeh. 2012. Prepared or not prepared: Top-down modulation on memory
of features and feature bindings. Acta Psychologica 141:3, 327-335. [Crossref]
152. Milan Gnjatović, Marko Janev, Vlado Delić. 2012. Focus tree: modeling attentional information in task-oriented human-machine
interaction. Applied Intelligence 37:3, 305-320. [Crossref]
153. Myriam C. Sander, Ulman Lindenberger, Markus Werkle-Bergner. 2012. Lifespan age differences in working memory: A two-
component framework. Neuroscience & Biobehavioral Reviews 36:9, 2007-2033. [Crossref]
154. Sabrina Trapp, Jöran Lepsien. 2012. Attentional orienting to mnemonic representations: Reduction of load-sensitive maintenance-
related activity in the intraparietal sulcus. Neuropsychologia 50:12, 2805-2811. [Crossref]
155. Ned Block. 2012. The Grain of Vision and the Grain of Attention. Thought: A Journal of Philosophy 1:3, 170-184. [Crossref]
156. Ryan T. Tanoue, Marian E. Berryhill. 2012. The mental wormhole: Internal attention shifts without regard for distance. Attention,
Perception, & Psychophysics 74:6, 1199-1215. [Crossref]
157. Tobias Katus, Søren K. Andersen, Matthias M. Müller. 2012. Nonspatial Cueing of Tactile STM Causes Shift of Spatial Attention.
Journal of Cognitive Neuroscience 24:7, 1596-1609. [Abstract] [Full Text] [PDF] [PDF Plus]
158. K. N. Thakkar, S. Park. 2012. Impaired Passive Maintenance and Spared Manipulation of Internal Representations in Patients
With Schizophrenia. Schizophrenia Bulletin 38:4, 787-795. [Crossref]
159. Angela A. Manginelli, Franziska Geringswald, Stefan Pollmann. 2012. Visual Search Facilitation in Repeated Displays Depends
on Visuospatial Working Memory. Experimental Psychology 59:1, 47-54. [Crossref]
160. 현현현. 2012. Effect of Task-irrelevant Feature Information on Visual Short-term Recognition of Task-relevant Feature. Korean
Journal of Cognitive Science 23:2, 225-248. [Crossref]
161. Tal Makovski. 2012. Are multiple visual short-term memory storages necessary to explain the retro-cue effect?. Psychonomic
Bulletin & Review 19:3, 470-476. [Crossref]
162. Klaus Oberauer, Laura Hein. 2012. Attention to Information in Working Memory. Current Directions in Psychological Science
21:3, 164-169. [Crossref]
163. Jaap Munneke, Artem V. Belopolsky, Jan Theeuwes. 2012. Shifting Attention within Memory Representations Involves Early
Visual Areas. PLoS ONE 7:4, e35528. [Crossref]
164. Marcin Leszczyński, Nicholas E. Myers, Elkan G. Akyürek, Anna Schubö. 2012. Recoding between Two Types of STM
Representation Revealed by the Dynamics of Memory Search. Journal of Cognitive Neuroscience 24:3, 653-663. [Abstract] [Full
Text] [PDF] [PDF Plus]
165. James M. Broadway, Matthew R. Hilimire, Paul M. Corballis. 2012. Orienting to external versus internal regions of space:
Consequences of attending in advance versus after the fact. Psychophysiology 49:3, 357-368. [Crossref]
166. Marian E. Berryhill, Lauren L. Richmond, Cara S. Shay, Ingrid R. Olson. 2012. Shifting Attention among Working Memory
Representations: Testing Cue Type, Awareness, and Strategic Control. Quarterly Journal of Experimental Psychology 65:3, 426-438.
[Crossref]
167. Jean-Francois Delvenne, Jessica L. Holt. 2012. Splitting attention across the two visual fields in visual short-term memory.
Cognition 122:2, 258-263. [Crossref]
168. Adam Gazzaley, Anna C. Nobre. 2012. Top-down modulation: bridging selective attention and working memory. Trends in
Cognitive Sciences 16:2, 129-135. [Crossref]
169. Duncan E. Astle, Anna C. Nobre, Gaia Scerif. 2012. Attentional control constrains visual short-term memory: Insights from
developmental and individual differences. Quarterly Journal of Experimental Psychology 65:2, 277-294. [Crossref]
170. Jarrod A. Lewis-Peacock, Andrew T. Drysdale, Klaus Oberauer, Bradley R. Postle. 2012. Neural Evidence for a Distinction
between Short-term Memory and the Focus of Attention. Journal of Cognitive Neuroscience 24:1, 61-79. [Abstract] [Full Text]
[PDF] [PDF Plus]
171. Bo-Cheng Kuo, Mark G. Stokes, Anna Christina Nobre. 2012. Attention Modulates Maintenance of Representations in Visual
Short-term Memory. Journal of Cognitive Neuroscience 24:1, 51-60. [Abstract] [Full Text] [PDF] [PDF Plus]
172. Duncan Edward Astle, Jennifer Summerfield, Ivan Griffin, Anna Christina Nobre. 2012. Orienting attention to locations in mental
representations. Attention, Perception, & Psychophysics 74:1, 146-162. [Crossref]
173. Tobias Katus, Søren K. Andersen, Matthias M. Müller. 2012. Maintenance of tactile short-term memory for locations is mediated
by spatial attention. Biological Psychology 89:1, 39-46. [Crossref]
174. James R. Brockmole, Christopher C. Davoli, Deborah A. Cronin. The Visual World in Sight and Mind 103-145. [Crossref]
175. Bo-Cheng Kuo, Pia Rotshtein, Yei-Yu Yeh. 2011. Attentional modulation of perceptual comparison for feature binding. Brain
and Cognition 77:3, 335-344. [Crossref]
176. Michi Matsukura, Andrew Hollingworth. 2011. Does visual short-term memory have a high-capacity stage?. Psychonomic Bulletin
& Review 18:6, 1098-1104. [Crossref]
177. Ned Block. 2011. Perceptual consciousness overflows cognitive access. Trends in Cognitive Sciences 15:12, 567-575. [Crossref]
178. Leon Gmeindl, James K. Nelson, Timothy Wiggin, Patricia A. Reuter-Lorenz. 2011. Configural representations in spatial working
memory: modulation by perceptual segregation and voluntary attention. Attention, Perception, & Psychophysics 73:7, 2130-2142.
[Crossref]
179. Holger Schultheis, Thomas Barkowsky. 2011. Casimir: An Architecture for Mental Spatial Knowledge Processing. Topics in
Cognitive Science 3:4, 778-795. [Crossref]
180. Yuji Yi, David Friedman. 2011. Event-related potential (ERP) measures reveal the timing of memory selection processes and
proactive interference resolution in working memory. Brain Research 1411, 41-56. [Crossref]
181. Artem V. Belopolsky, Jan Theeuwes. 2011. Selection within visual memory representations activates the oculomotor system.
Neuropsychologia 49:6, 1605-1610. [Crossref]
182. Alexandra M. Murray, Anna C. Nobre, Mark G. Stokes. 2011. Markers of preparatory attention predict visual short-term memory
performance. Neuropsychologia 49:6, 1458-1465. [Crossref]
183. Bo-Cheng Kuo, Yei-Yu Yeh, Anthony J.-W. Chen, Mark D’Esposito. 2011. Functional connectivity during top-down modulation
of visual short-term memory representations. Neuropsychologia 49:6, 1589-1596. [Crossref]
184. Nelson Cowan. 2011. The focus of attention as observed in visual working memory tasks: Making sense of competing claims.
Neuropsychologia 49:6, 1401-1406. [Crossref]
185. Taiji Ueno, Judit Mate, Richard J. Allen, Graham J. Hitch, Alan D. Baddeley. 2011. What goes through the gate? Exploring
interference with visual feature binding. Neuropsychologia 49:6, 1597-1604. [Crossref]
186. Jöran Lepsien, Ian Thornton, Anna C. Nobre. 2011. Modulation of working-memory maintenance by directed attention.
Neuropsychologia 49:6, 1569-1577. [Crossref]
187. Duncan E. Astle, Gaia Scerif. 2011. Interactions between attention and visual short-term memory (VSTM): What can be learnt
from individual and developmental differences?. Neuropsychologia 49:6, 1435-1445. [Crossref]
188. Tal Makovski, Khena M. Swallow, Yuhong V. Jiang. 2011. Attending to unrelated targets boosts short-term memory for color
arrays. Neuropsychologia 49:6, 1498-1505. [Crossref]
189. Annelinde R.E. Vandenbroucke, Ilja G. Sligte, Victor A.F. Lamme. 2011. Manipulations of attention dissociate fragile visual short-
term memory from visual working memory. Neuropsychologia 49:6, 1559-1568. [Crossref]
190. Ilja G. Sligte, Martijn E. Wokke, Johannes P. Tesselaar, H. Steven Scholte, Victor A.F. Lamme. 2011. Magnetic stimulation
of the dorsolateral prefrontal cortex dissociates fragile visual short-term memory from visual working memory. Neuropsychologia
49:6, 1578-1588. [Crossref]
191. Melissa R. Beck, Amanda E. van Lamsweerde. 2011. Accessing long-term memory representations during visual change detection.
Memory & Cognition 39:3, 433-446. [Crossref]
192. Wayne S. Smith, J. D. Mollon, Rishi Bhardwaj, Hannah E. Smithson. 2011. Is there brief temporal buffering of successive visual
inputs?. Quarterly Journal of Experimental Psychology 64:4, 767-791. [Crossref]
193. 현현현. 2011. Understanding Visual Working Memory Based on Significant Examples of Behavioral Studies. Korean Journal of
Cognitive and Biological Psychology 23:1, 45-90. [Crossref]
194. Teresa Roig Rovira, Marcos Ríos Lago, Núria Paúl Lapedriza. Atención y concentración 29-e6. [Crossref]
195. Gaia Scerif, Ann Steele. Neurocognitive development of attention across genetic syndromes 285-301. [Crossref]
196. Jean-François Delvenne, Axel Cleeremans, Cédric Laloyaux. 2010. Feature Bindings Are Maintained in Visual Short-Term
Memory Without Sustained Focused Attention. Experimental Psychology 57:2, 108-116. [Crossref]
197. San-Cai LIANG, Xu-Qun You. 2010. Conceptual Control of Visual Images Scanning. Acta Psychologica Sinica 42:9, 889-898.
[Crossref]
198. Na Shao, Jie Li, Rende Shui, Xiaojie Zheng, Jiangang Lu, Mowei Shen. 2010. Saccades elicit obligatory allocation of visual working
memory. Memory & Cognition 38:5, 629-640. [Crossref]
199. Eiichi Hoshino, Ken Mogi. Evidence for False Memory before Deletion in Visual Short-Term Memory 255-261. [Crossref]
200. Sabine Schwager, Herbert Hagendorf. 2009. Goal-directed access to mental objects in working memory: The role of task-specific
feature retrieval. Memory & Cognition 37:8, 1103-1119. [Crossref]
201. Pamela M. Greenwood, Ramya Sundararajan, Ming-Kuan Lin, Reshma Kumar, Karl J. Fryxell, Raja Parasuraman. 2009. Both
a Nicotinic Single Nucleotide Polymorphism (SNP) and a Noradrenergic SNP Modulate Working Memory Performance when
Attention is Manipulated. Journal of Cognitive Neuroscience 21:11, 2139-2153. [Abstract] [Full Text] [PDF] [PDF Plus]
202. Yuji Yi, Naomi Driesen, Hoi-Chung Leung. 2009. Behavioral and neural correlates of memory selection and interference resolution
during a digit working memory task. Cognitive, Affective, & Behavioral Neuroscience 9:3, 249-259. [Crossref]
203. Philip C. Ko, Adriane E. Seiffert. 2009. Updating objects in visual short-term memory is feature selective. Memory & Cognition
37:6, 909-923. [Crossref]
204. Hagit Magen, Tatiana-Aloi Emmanouil, Stephanie A. McMains, Sabine Kastner, Anne Treisman. 2009. Attentional demands
predict short-term memory load response in posterior parietal cortex. Neuropsychologia 47:8-9, 1790-1798. [Crossref]
205. Jeffrey S. Phillips, Katerina Velanova, David A. Wolk, Mark E. Wheeler. 2009. Left posterior parietal cortex participates in both
task preparation and episodic retrieval. NeuroImage 46:4, 1209-1221. [Crossref]
206. Michi Matsukura, Shaun P. Vecera. 2009. Interference between object-based attention and object-based memory. Psychonomic
Bulletin & Review 16:3, 529-536. [Crossref]
207. Po-Han Lin, Steven J. Luck. 2009. The influence of similarity on visual working memory representations. Visual Cognition 17:3,
356-372. [Crossref]
208. Shahin Nasr, Ali Moeeny, Hossein Esteky. 2008. Neural Correlate of Filtering of Irrelevant Information from Visual Working
Memory. PLoS ONE 3:9, e3282. [Crossref]
209. María Ruz, Anna C. Nobre. 2008. Attention Modulates Initial Stages of Visual Word Processing. Journal of Cognitive Neuroscience
20:9, 1727-1736. [Abstract] [PDF] [PDF Plus]
210. Ilja G. Sligte, H. Steven Scholte, Victor A. F. Lamme. 2008. Are There Multiple Visual Short-Term Memory Stores?. PLoS
ONE 3:2, e1699. [Crossref]
211. Tal Makovski, Yuhong V. Jiang. 2007. Distributing versus focusing attention in visual short-term memory. Psychonomic Bulletin
& Review 14:6, 1072-1078. [Crossref]
212. Michi Matsukura, Steven J. Luck, Shaun P. Vecera. 2007. Attention effects during visual short-term memory maintenance:
Protection or prioritization?. Perception & Psychophysics 69:8, 1422-1434. [Crossref]
213. J. Lepsien, A. C. Nobre. 2007. Attentional Modulation of Object Representations in Working Memory. Cerebral Cortex 17:9,
2072-2083. [Crossref]
214. Soojung Min, 현현현, Min-Shik Kim. 2007. The Effects of Working Memory and Selection on Social-Emotional Evaluation. Korean
Journal of Cognitive and Biological Psychology 19:2, 171-185. [Crossref]
215. Hoi-Chung Leung, Hwamee Oh, Jamie Ferri, Yuji Yi. 2007. Load response functions in the human spatial working memory
circuit during location memory updating. NeuroImage 35:1, 368-377. [Crossref]
216. Yei-Yu Yeh, Bo-Cheng Kuo, Ho-Ling Liu. 2007. The neural correlates of attention orienting in visuospatial working memory
for detecting feature and conjunction changes. Brain Research 1130, 146-157. [Crossref]
217. Nienke J.H. Korsten, Nikos Fragopanagos, Matthew Hartley, Neill Taylor, John G. Taylor. 2006. Attention as a controller. Neural
Networks 19:9, 1408-1421. [Crossref]
218. Gregory V. Simpson, Corby L. Dale, Tracy L. Luks, William L. Miller, Walter Ritter, John J. Foxe. 2006. Rapid targeting
followed by sustained deployment of visual spatial attention. NeuroReport 17:15, 1595-1599. [Crossref]
219. Jöran Lepsien, Anna C. Nobre. 2006. Cognitive control of attention in the human brain: Insights from orienting attention to
mental representations. Brain Research 1105:1, 20-31. [Crossref]
220. Amir Raz, Jason Buhle. 2006. Typologies of attentional networks. Nature Reviews Neuroscience 7:5, 367-379. [Crossref]
221. Min Bao, Zhi-Hao Li, Xiang-Chuan Chen, Da-Ren Zhang. 2006. Backward inhibition in a task of switching attention within
verbal working memory. Brain Research Bulletin 69:2, 214-221. [Crossref]
222. P. Sauseng, W. Klimesch, W. Stadler, M. Schabus, M. Doppelmayr, S. Hanslmayr, W. R. Gruber, N. Birbaumer. 2005. A shift of
visual spatial attention is selectively associated with human EEG alpha activity. European Journal of Neuroscience 22:11, 2917-2926.
[Crossref]
223. Jöran Lepsien, Ivan C. Griffin, Joseph T. Devlin, Anna C. Nobre. 2005. Directing spatial attention in mental representations:
Interactions between attentional orienting and working-memory load. NeuroImage 26:3, 733-743. [Crossref]
224. J. Richard Jennings, Maurits W. van der Molen. 2005. Preparation for Speeded Action as a Psychophysiological Concept.
Psychological Bulletin 131:3, 434-459. [Crossref]
225. A. C. Nobre, J. T. Coull, P. Maquet, C. D. Frith, R. Vandenberghe, M. M. Mesulam. 2004. Orienting Attention to Locations in
Perceptual Versus Mental Representations. Journal of Cognitive Neuroscience 16:3, 363-373. [Abstract] [PDF] [PDF Plus]