Anterior cingulate and prefrontal cortex activity in an FMRI study of

trial-to-trial adjustments on the Simon task

John G. Kerns

Department of Psychological Sciences, 214 McAlester Hall, University of Missouri, Columbia, MO 65211, USA
Received 21 April 2006; revised 30 May 2006; accepted 4 June 2006
Available online 28 July 2006
People alter their task performance on a trial-to-trial basis, for
example after an incongruent trial on tasks involving response conflict.
Previous research has found that these adjustments are most robust in
the Simon task. One explanation for behavioral adjustments is the
conflict-monitoring hypothesis, which posits that the dorsal anterior
cingulate cortex (ACC) responds to conflict and that this serves as a
signal to recruit other brain regions such as the prefrontal cortex
(PFC) to minimize conflict and improve performance. However,
another independently supported explanation for behavioral adjust-
ments on the Simon task is the feature integration view, which can
account for behavioral adjustments as the result of stimulus repetitions
and alternations. Hence, by itself, evidence for behavioral adjustments
on the Simon task does not clearly provide evidence for the conflict-
monitoring hypothesis. However, the conflict-monitoring hypothesis
does predict that behavioral adjustments on the Simon task should
involve ACC conflict activity and PFC post-conflict activity. In the
current study, consistent with the conflict-monitoring hypothesis,
behavioral adjustments in performance on the Simon task were
predicted by ACC conflict-related activity. In addition, subsequent
behavioral adjustments were associated with PFC activity, with
previous trial ACC conflict-related activity predicting greater PFC
activity on subsequent trials. These results provide additional evidence
that behavioral adjustments on the Simon task are due in part to ACC
conflict monitoring and the subsequent recruitment of PFC to minimize
conflict.
2006 Elsevier Inc. All rights reserved.
Keywords: Anterior cingulate; Prefrontal cortex; Response conflict; Post-
conflict adjustments; Cognitive control
Introduction
People adjust their ongoing performance on behavioral tasks in
a number of ways, such as slowing down and being more accurate
after errors (Rabbit, 1966; Laming, 1979). In addition, on tasks
involving response conflict (i.e., the simultaneous activation of at
least two different responses), after incongruent high conflict trials
participants also adjust their performance (Gratton et al., 1992). For
example, a response conflict task such as the Stroop color naming
task can involve high conflict incongruent trials (i.e., the word
RED in green ink, correct response is green) and low conflict
congruent trials (i.e., the word GREEN in green ink). After a high
conflict incongruent trial, responses are slower for a subsequent
congruent trial and faster for a subsequent incongruent trial.
Specifically, on congruent trials responses are slower if preceded
by an incongruent trial (i.e., iC trials) than if preceded by a
congruent trial (i.e., cC trials). Moreover, participants are faster if
an incongruent trial was preceded by an incongruent trial (i.e., iI
trials) than if preceded by a congruent trial (i.e., cI trials). Hence,
after conflict participants adjust their performance, responding
slower for a subsequent congruent trial and faster for a subsequent
incongruent trial.
One explanation for these post-conflict adjustments is the
conflict-monitoring hypothesis (Botvinick et al., 2001, 2004). This
hypothesis posits that at least two brain regions play distinct and
complementary roles in behavioral adjustments. This hypothesis
states that the dorsal anterior cingulate cortex (ACC) is activated
by the occurrence of response conflict (Carter et al., 1998;
Botvinick et al., 1999). The monitoring of response conflict by the
ACC then serves as a signal that results in the recruitment of other
brain regions to minimize the amount of conflict on subsequent
performance (Botvinick et al., 2001; Cohen et al., 2000; Kerns et
al., 2004). The prefrontal cortex (PFC) is one brain region thought
to be engaged after the occurrence of conflict to subsequently
minimize conflict (by providing a contextually or task-set
appropriate biasing signal to poster regions; Botvinick et al.,
2001; Miller and Cohen, 2001). Hence, from the conflict-
monitoring view, ACC conflict monitoring serves as a signal that
results in the recruitment of PFC to minimize subsequent conflict
and improve performance.
However, in addition to the conflict-monitoring hypothesis,
there are other explanations for post-conflict behavioral adjust-
ments, such as the feature integration view. From this perspective,
behavioral adjustments are the result of exact stimulus repetitions
and alternations (Hommel et al., 2004). For example, consider the
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doi:10.1016/j.neuroimage.2006.06.012
Simon task (Simon and Small, 1969; Lu and Proctor, 1995) on
which two stimulus dimensions can vary from trial to trial:
stimulus location (e.g., right or left) and stimulus identity (e.g., red
or green object, necessitating either a right or left response,
respectively). On the Simon task, only cC and iI trials involve
exact stimulus repetitions (e.g., a green object on the left on two
consecutive trials). Given that the post-conflict adjustment effect
involves cC trials being faster than iC trials and iI trials being faster
than cI trials, then faster responses on exact stimulus repetitions for
only cC and iI trials can account for some evidence of post-conflict
behavioral adjustments (Mayr et al., 2003). But in addition, on all
other cC and iI trials not involving an exact stimulus repetition,
they involve an exact stimulus alternation (e.g., green on left
followed by red on right). It has been argued that people process
perceptual stimuli as integrated event files (integration of both
stimulus and response features, i.e., as object on left necessitating a
right-hand response; Hommel et al., 2001; Stoet and Hommel,
1999). If both stimulus dimensions change, this complete
alternation can be used to indicate a faster response. However,
when only one stimulus dimension changes (i.e., red but still on
left), RT has been found to increase because of the partial match to
the previous trial. Hence, from this feature integration view,
behavioral adjustments on the Simon task can be accounted for by
cC and iI trials always involving either exact stimulus repetitions or
alternations (thereby decreasing RT) whereas iC and cI trials
always involve a partial match with the previous trials stimuli
(thereby increasing RT). Importantly, evidence for the feature
integration view has been found in situations without response
conflict (Hommel et al., 2004). This suggests that evidence of
behavioral adjustments on some response conflict tasks may not
necessarily indicate conflict monitoring.
Some research that does support a role for conflict monitoring
in behavioral adjustments comes from brain imaging. The conflict-
monitoring hypothesis predicts that ACC conflict-related activity
should predict adjustments in performance. Moreover, it predicts
that during post-conflict adjustment trials that PFC should be
active. Consistent with this, in a study using the Stroop task, it was
found that ACC conflict activity predicted greater post-conflict
adjustments (Kerns et al., 2004). At the same time, greater
subsequent post-conflict adjustments were associated with greater
PFC activity. Hence, on the Stroop task, it appears that conflict
monitoring may play a role in behavioral adjustments.
However, it is unclear whether these results for the Stroop task
would also be found on other response conflict tasks. In fact, there
is some evidence from previous research that the size of trial-to-
trial adjustments might vary across response conflict tasks. For
example, some studies have not reported significant trial-to-trial
adjustments in the Eriksen flanker task after the removal of
stimulus repetitions (Mayr et al., 2003; Nieuwenhuis et al., in
press). On the Stroop task, conflict adjustments have been reported
without stimulus repetitions in at least three studies (Egner and
Hirsch, 2005a; Kerns et al., 2004, 2005), although in two of these
studies the overall effect was of only moderate to large size and
was not always consistent across both current trial types (e.g., only
iI differed from cI or only iC differed from cC; Kerns et al., 2004,
2005). In contrast, the response conflict task, which seems to
consistently produce the largest conflict adjustment effect sizes, is
the Simon task (Sturmer et al., 2002; and even without exact
stimulus repetitions, Sohn and Carter, 2006). On the surface, this
seems counterintuitive because the Simon task involves a smaller
behavioral RT interference effect (i.e., incongruent RT minus
congruent RT) than either the Eriksen or the Stroop tasks (e.g., in a
recent behavioral study with n>300, interference effects for the
Simon, Eriksen, and Stroop tasks were 26.9, 69.2, and 183.7 ms,
respectively; Kerns, 2006).
In part, one explanation for the variation in the size of post-
conflict adjustment effects across tasks is the extent, consistent
with the feature integration view (Hommel et al., 2004), to which
iC and cI trials involve a partial match to the previous trials
stimuli. On the Simon task, with two possible relevant and
irrelevant stimuli, all iC and cI trials involve a partial match, which
can account for a large adjustment effect. In contrast, the Stroop
task usually involves more than two possible relevant and
irrelevant stimuli, with some iC and cI trials involving exact
stimulus alternations (Kerns et al., 2004), which can account for
the reduced size of the adjustment effect. In support of this, the one
imaging Stroop study that found the largest post-conflict adjust-
ment effect involved only two possible relevant and irrelevant
stimuli and therefore involved all partial matches on iC and cI trials
(Egner and Hirsch, 2005a).
Given evidence for the effect of stimulus repetitions and
alternations on behavioral adjustment, this leaves open the
possibility that behavioral adjustments on the Simon task may
not be due to conflict monitoring. If true, this would suggest that
the conflict-monitoring hypothesis may not apply to all response
conflict tasks, which might suggest important limits on the theory.
The current study examined whether conflict adjustments occurred
on the Simon task and whether they were associated with ACC and
PFC activity. According to the conflict-monitoring hypothesis,
although repetitions and alternations should influence performance
(Cho et al., 2002; Hommel et al., 2004), a previous trials ACC
conflict activity should still predict the subsequent occurrence of
behavioral adjustments (Jones et al., 2002). Moreover, trials
involving large behavioral adjustments (i.e., both speeding up on iI
trials and slowing down on iC trials) should be associated with
PFC activity, which should be associated with amount of ACC
activity on the previous trial.
Materials and methods
Participants
Twenty-six right-handed individuals (14 females) with an
average age of 24.2 years (SD=4.5, range 1836) participated in
this study and were paid $30 for their participation.
Behavioral task
In the scanner, participants performed the Simon task (Lu and
Proctor, 1995; Simon and Small, 1969). On each trial, participants
saw a circle in either red or green ink on the left or right side of
their visual field. Participants needed to respond with their right
index finger for green circles and with their left index finger for red
circles. Each trial began with a fixation cross presented for 250 ms.
Then a green or red circle appeared for 1500 ms, followed by a
blank screen for 750 ms (given an average reaction time of less
than 700 ms, following Wuhr and Ansorge, 2005, the average
response to stimulus interval was a little more than 1800 ms).
Participants completed 8 blocks of 40 trials each. Each trial was
either a congruent or an incongruent trial. On congruent trials, the
location of the stimulus matched the side of response (e.g., a green
circle, necessitating a right-hand response, presented on the right
400 J.G. Kerns / NeuroImage 33 (2006) 399405
side). On incongruent trials, the location of the stimulus did not
match the side of response (e.g., a green circle on the left side). The
proportion of congruent and incongruent trials was 50% each. Both
speed and accuracy was stressed equally. Errors were infrequent
(average of 3.4 error trials per participant) with only 2 participants
(8%) having more than 9 errors. For the basic Simon effect, the
dependent variable was the difference in reaction times between
incongruent and congruent trials.
In analyzing post-conflict adjustments, there are four different
trial types, depending upon the congruence or incongruence of the
preceding and the current trial: (1) iC trials (i.e., a congruent trial
preceded by an incongruent trial), (2) cC trials, (3) iI trials, and (4)
cI trials. The proportion of each of these four types of trials was
25%. The post-conflict adjustment effect is an interaction between
previous and current trial congruency, with iC trials being slower
than cC trials and iI trials being faster than cI trials.
Functional imaging
Functional scans were acquired using a 1.5-T Siemens
Symphony whole body scanner with a standard head coil. Pillows
were used to minimize head movement. Three functional T1-
weighted scouts (in the axial, coronal, and sagittal planes) were
used to localize the AC/PC line. Structural images were obtained
prior to and in the same plane as functional images using a standard
T1-weighted pulse sequence. Twenty-six contiguous 3.8-mm-thick
axial slices with 3.753.75 mm in-plane resolution were obtained
beginning 11.4 mm below the AC-PC line. Functional images were
acquired using an echo-planar sequence (TR 2500 ms, TE 35 ms,
flip angle 70, FOV 24 cm). During the scanning session, there
were 4 BOLD runs, with each BOLD run consisting of two blocks
of 40 trials of the Simon task, with 30 s of baseline fixation (i.e.,
the presentation of a fixation cross) occurring in-between the two
blocks.
Data analysis
Incremental (scan to scan) and total movement were corrected
using FSL (Jenkinson et al., 2002). Participants were excluded
from the study with mean estimated absolute movement from the
reference scan greater than either 4 mm or degrees. Structural
images were cross-registered to a reference brain by minimizing
signal intensity differences with 12-parameter AIR (Woods et al.,
1998). Imaging data were linearly detrended (removing linear trend
within each BOLD run) and then aligned to structural scans using
AIR (Woods et al., 1998), after which images were set to a standard
mean intensity and spatially smoothed (8 mm FWHM). Finally,
functional images were high-pass filtered (50 sec cut off) and pre-
whitened using FSL (Woolrich et al., 2001).
Imaging data were analyzed with a random effects single-
subject general linear model using FSL, AFNI (Cox, 1996), and
NIS software (Fissell et al., 2003). To examine response conflict
activity on the Simon task, one analysis was performed with three
covariates: (1) incongruent correct trials; (2) error trials; and (3) an
all trials covariate (either congruent or incongruent). Hence, in this
analysis, significant conflict activity would be reflected in a
significant incongruent covariate (i.e., in the regression analysis
activity on incongruent trials even after statistically accounting for
general level of activity on all trials). Note that tolerance (a
diagnostic measure of multicollinearity, with values ranging from 0
to 1, with values less than 0.1 indicating problematic multi-
collinearity; Hays, 1988) was 0.70 for this response conflict
analysis (for the overlap between incongruent and all trials
covariates). Z-maps for individual subjects were generated that
reflected the extent to which each voxels activity conformed to an
a priori canonical double gamma hemodynamic response function.
Statistical threshold was p<0.005 and 8 contiguous voxels
(Forman et al., 1995).
To examine whether activity in certain brain regions on conflict
trials predicted making post-conflict adjustments, following Kerns
et al. (2004), iC and iI trials were divided into fast and slow trials
using a median split. Good adjustment for iC trials means going
slower, whereas good adjustment for iI trials means going faster.
To examine whether activity on the preceding trial predicted
activity on iI trials, an analysis was performed with the following
covariates: (1) correct trials immediately preceding correct fast iI
trials; (2) correct trials immediately preceding slow iI trials; (3)
error trials; (4) all other incongruent trials; and (5) an all trials
covariate. Then, activity for trials preceding fast iI trials was
compared to trials preceding slow iI trials. A separate analogous
analysis was completed for to examine whether activity on the
preceding trial predicted activity on iC trials (note that this analysis
was separated from the analysis for iI trials to prevent multi-
collinearity from affecting the results). Finally, to examine which
brain regions predicted post-conflict adjustments both for iI and IC
trials, a conjunction analysis was performed that examined regions
that exhibited significant activity preceding both good iI and good
iC trials, using an a priori p<0.05 threshold within each analysis
(Friston et al., 2005; Kerns et al., 2004).
To examine which brain regions were active on good
adjustment trials, following Kerns et al. (2004), two additional
analyses were performed. In one analysis examining activity on iI
trials, the following covariates were used: (1) correct fast iI trials
that were preceded by a correct trial; (2) correct slow iI trials
preceded by a correct trial; (3) error trials; (4) all other incongruent
trials; and (5) an all trials covariate. Then, activity for fast iI trials
was compared to activity for slow iI trials. A similar separate
analysis was conducted to examine activity on iC trials. Then a
conjunction analysis was performed to examine which brain
regions were active during post-conflict adjustments for both iI and
IC trials.
One final analysis examined whether a brain region that
predicted making good post-conflict adjustments (such as the
ACC) was directly associated with activity in a brain region that
was active during good post-conflict adjustment trials (such as the
PFC). To do this, within each subject, conflict and post-conflict
scans were correlated. Given a TR of 2.5 s, the BOLD response
should be at its peak elevation in between 2 and 4 scans for both
conflict and post-conflict scans, respectively. In order to not
correlate activity for temporally overlapping scans, the second
conflict scan was correlated with the second post-conflict scan
(which was the third conflict scan), the third conflict scan was
correlated with the third post-conflict scan, and the fourth conflict
scan was correlated with the fourth post-conflict scan. These
correlation values were averaged within each subject and then
treated as the dependent variable to test whether the correlation
between conflict and post-conflict scans were significant. In
addition, to rule out general activity across brain regions, partial
correlations were computed two different ways. One set of partial
correlations controlled for activity in a brain region also activated
by response conflict. The other set of partial correlations controlled
for baseline activity (i.e., at scan 1 of the conflict trial). These
401 J.G. Kerns / NeuroImage 33 (2006) 399405
correlation analyses provide information about whether ACC
activity was correlated with PFC activity. However, given the
temporal sluggishness of the BOLD response (Huettel et al., 2004),
one cannot definitively conclude that this correlation reflects only
conflict (ACC) and post-conflict (PFC) trial activity.
Results
Behavioral data
Behavioral data were first analyzed in a 2 (previous trial
type: congruent vs. incongruent) 2 (current trial type: con-
gruent vs. incongruent) ANOVA. As expected, as can be seen
in Table 1 and Fig. 1, the effect of current trial type (i.e., the
basic Simon effect) was significant, F(1,25) =18.80, p<0.001, as
reaction times were slower for current incongruent trials (683.3,
SD=97.9) than for current congruent trials (mean=700.1,
SD=102.8). The effect of previous trial type was not
significant, F(1,25) =2.03, p=0.17. However, there was a
significant previous by current trial type interaction (i.e., the
post-conflict adjustment effect), F(1,25) =102.53, p<0.001. As
can be seen in Fig. 1, iI trials were significantly faster than cI
trials, t(25) =8.96, p<0.001, whereas iC trials were significantly
slower than cC trials, t(25) =7.76, p<0.001. This was true even
after removing exact stimulus repetitions, iI vs. cI, t(25) =5.21,
p<0.001; iC vs. cC, t(25) =4.19, p<0.001.
Imaging data
First, response conflict activity on incongruent trials was
examined. As can be seen in Table 2, as expected and consistent
with previous research (Nee et al., 2004), there was significant
conflict-related activity in the caudal ACC extending superiorly
into supplementary motor areas. Among other regions activated by
conflict were the left PFC and bilateral parietal areas.
Next, a conjunction analysis was conducted to examine which
brain regions on conflict trials predicted making good behavioral
adjustments on both subsequent iI and iC trials. As can be seen in
Fig. 2, only one brain region, the caudal ACC (Brodmanns area,
BA, 32; Talairach coordinates 3, 8, 41), significantly predicted
making better adjustments on the following trial. Interestingly, the
ACC region in the current study is very similar to the ACC
region found in a previous study to predict post-conflict
adjustments on the Stroop task (Kerns et al., 2004; Talairach
coordinates 1, 10, 40).
Next, a conjunction analysis was conducted to examine which
brain regions were active during good adjustment trials for both iI
and iC trials. As can be seen in Fig. 3, two adjacent brain regions in
the left PFC (BA 9/46, coordinates 44, 28, 28; and BA 6/9,
coordinates 52, 5, 36) were significantly active on good
adjustment trials.
Finally, it was examined whether previous trial ACC conflict
activity was directly associated with post-conflict adjustment
dorsolateral (BA 9/46) PFC activity. Previous trial ACC conflict
activity was significantly associated with current trial PFC activity,
p<0.01. This was true even after partialling out activity for a
control brain region, the left inferior parietal region which was also
activated by conflict, or after partialling out baseline PFC activity
(i.e., amount of activity during the first scan of the conflict trial).
Discussion
The results of this study are consistent with the conflict-
monitoring hypothesis. In this study, ACC conflict-related activity
predicted subsequent adjustments in performance. In addition,
Table 1
Post-conflict behavioral adjustment effect on the Simon test
Trial type Mean SD
cC 650.6 102.5
iC 699.3 97.5
cI 712.3 100.7
iI 672.8 100.7
Fig. 1. Simon task behavioral performance, with significant effects of both
current conflict and post-conflict adjustments (i.e., previous trial by current
trial interaction).
Fig. 2. Region of anterior cingulate cortex significantly predicting
subsequent post-conflict behavioral adjustments (BA, 32; coordinates 3,
8, 41).
Table 2
Regions significantly activated during incongruent Simon trials
Region (Brodmann's area) Number
of
voxels
Peak Activity
Talairach coordinates
x y z
Anterior cingulate/supplementary
motor (32/6)
82 7 3 49
Inferior parietal (BA 40) 346 44 52 48
179 41 38 53
Middle frontal gyrus (BA 9) 15 40 30 33
18 33 37 35
402 J.G. Kerns / NeuroImage 33 (2006) 399405
high adjustment trials were associated with increased PFC
activity, with ACC activity on the previous trial being associated
with amount of PFC activity on the subsequent trial. Overall,
these results are consistent with the hypothesis that ACC conflict
monitoring serves as a signal that results in the recruitment of
greater PFC activity and that this contributed to behavioral
adjustments.
The current results using a Simon task are consistent with those
of a previous study using the Stroop task (Kerns et al., 2004).
Moreover, recently a third study has also replicated and extended
this pattern of results using a task-switching paradigm (involving
color and movement direction; Liston et al., 2006). Hence, it
appears that conflict monitoring might contribute to behavioral
adjustments in performance across a number of response conflict
tasks.
One issue in future research testing predictions of the conflict-
monitoring hypothesis is to account for other possible sources of
variance that can contribute trial-to-trial adjustments. For example,
as discussed, post-conflict adjustments after removing exact
repetitions on the Eriksen flanker task have not always been
reported (Mayr et al., 2003). However, it has been argued that
negative priming might reduce adjustment effects on this task
(Ullsperger et al., 2005). Consistent with this, a flanker task that
removed negative priming effects did result in significant
behavioral adjustments (Ullsperger et al., 2005). In the current
study, exact stimulus repetitions and alternations also probably
contributed to post-conflict adjustments (Hommel et al., 2004). In
fact, the crossover interaction, with iI trials being significantly
faster than iC trials in the current study might be accounted for by
the joint effects of both conflict monitoring and repetitions/
alternations. Importantly, recent behavioral research by Wuhr
(2005) has found evidence for significant post-conflict behavioral
adjustments using a modified Simon task even after removing the
effects of repetitions and alternations. Future imaging research
using the Simon task to investigate behavioral adjustments that
removed the effect of repetitions and alternations might produce
even clearer neural evidence of ACC conflict activity predicting
behavioral adjustments and PFC post-conflict activity being
associated with behavioral adjustments.
Another issue for future research on conflict monitoring and
behavioral adjustments is how general the effect of the recruitment
of cognitive control is across different types of tasks. For example,
in the current study significant post-conflict PFC activity occurred
on the left side of the brain, whereas in a previous study PFC
activity was on the right (Kerns et al., 2004; in contrast, the area of
the ACC that predicted post-conflict adjustments in the current
study is very similar to the area reported in a previous study;
current study Talairach coordinates: 3, 8, 41; in Kerns et al.,
2004: 1, 10, 40). As argued previously, different regions of the
PFC might be recruited depending upon the particular nature of the
task (Kerns et al., 2004). In a previous study, the Stroop task,
which involves attending to color, activated right PFC. In the
Simon task, it is possible that participants are more likely to use a
verbal strategy (e.g., right for green, left for red, which were the
exact instructions given to participants), which might be likely to
activate left rather than right PFC. Importantly, several recent
behavioral studies have found evidence that post-conflict adjust-
ments do not transfer across different tasks (Cho et al., 2006; Egner
et al., 2006; Wendt et al., in press). This suggests that the post-
conflict recruitment of PFC activity may be somewhat specific and
not general. This also seems consistent with the view that the role
of the PFC is the maintenance of specific, task-relevant information
(Miller and Cohen, 2001), information that would vary substan-
tially across tasks.
Another issue for future research on trial-to-trial adjustments is
to examine whether results could be influenced by BOLD
nonlinearity. Wager and colleagues have outlined a scenario in
which observed brain activity could differ on a trial-to-trial basis
due to nonlinearity (Wager et al., 2005). If one trial type has greater
activity on a previous trial, there might be relatively reduced
activity on the subsequent trial of that same trial type due to
nonlinearity (i.e., BOLD saturation). This suggests that controlling
for the effects of nonlinearity might be critical in some contexts to
accurately assess neural activity across two trials (Wager et al.,
2005). However, for the current study (and for Kerns et al., 2004),
it is not immediately clear how nonlinearity could account for the
results. The results of the current study differ in two ways from the
scenario outlined by Wager and colleagues. If nonlinearity
produced the trial-to-trial changes in observed BOLD response,
then increased activity on one trial should be followed by
decreased activity on the next trial. However, in the current study,
increased activity on one trial (conflict trial) was followed by
increased (not decreased) activity on the next trial (post-conflict
trial), which is the exact opposite of what would be expected by an
effect of nonlinearity. In addition, another difference from the
scenario outlined by Wager and colleagues is that in the current
study, significant activity was not found across trials in the same
brain region. For example, activity in PFC differed significantly
between trials only on post-conflict trials, but not on the previous
trial. Nonlinearity cannot account for these results because if there
is no difference in previous trial activity then there is no reason to
expect a difference in current trial activity. Nevertheless, given that
nonlinearity could produce BOLD changes across trials, future
imaging research on trial-to-trial effects should take into account
the effects that nonlinearity could have on their results at short ITIs
(Wager and Nichols, 2003; Wager et al., 2005).
One other issue for future research on the nature of post-conflict
adjustments is to more precisely delineate the time course of the
engagement of the PFC and of control processes (Durston et al.,
2003). To answer this question, ideally it would be possible to
conduct research using jittered ITIs and/or at least occasional long
ITIs. However, one can imagine that this type of imaging research
Fig. 3. Left prefrontal regions significantly activated during post-conflict
adjustment trials (BA 9/46, coordinates 52, 5, 36).
403 J.G. Kerns / NeuroImage 33 (2006) 399405
may not be feasible in studying post-conflict adjustments. One
reason is that there might be only a short temporal window in
which greater cognitive control is engaged. In fact, on the Simon
task it has been found that behavioral adjustments decrease with
increasing ITI, although significant adjustments have still been
reported with a 6-s ITI (Wuhr and Ansorge, 2005). A second
reason is that it has been argued that the immediate engagement of
cognitive control may be less likely to occur if there can be a long
delay period before control is needed (Weissman et al., 2005).
However, two studies using a jittered ITI have now reported
significant post-conflict behavioral adjustments and significant
post-conflict PFC activation, one using a jittered ITI of between 3
and 5 s (Egner and Hirsch, 2005b), the other using a jittered ITI of
between 0.5 and 12.5 s (Liston et al., 2006). Hence, it does appear
that the recruitment of the PFC lasts for a number of seconds and
even with variability in the onset of the next trial. One additional
way future research could examine the time course of the
engagement of greater cognitive control is to use ERPs (West,
2004).
Acknowledgments
Thanks to Christy Watson, Joni Schupp, and the rest of the staff
at the Radiology Department at the University of Missouri Medical
School and Hospital. Thanks also to Raymond Y. Cho for helpful
discussions of these research ideas.
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