Cerebral Cortex December 2008;18:2811--2819

doi:10.1093/cercor/bhn037
Advance Access publication March 27, 2008

Neural Correlates of True Memory, False Nobuhito Abe1, Jiro Okuda2, Maki Suzuki1,3, Hiroshi Sasaki2,
Tetsuya Matsuda2, Etsuro Mori1, Minoru Tsukada2 and
Memory, and Deception Toshikatsu Fujii1
1
Department of Behavioral Neurology and Cognitive
Neuroscience, Tohoku University Graduate School of Medicine,
Sendai, Japan, 2Brain Science Research Center, Tamagawa
University Brain Science Institute, Tokyo, Japan and 3Division of
Cyclotron Nuclear Medicine, Cyclotron and Radioisotope
Center, Tohoku University, Sendai, Japan

We used functional magnetic resonance imaging (fMRI) to Langleben et al. 2005; Seth et al. 2006). However, it should be
determine whether neural activity can differentiate between true kept in mind that even if we can judge whether or not people
memory, false memory, and deception. Subjects heard a series of are telling a lie, their ‘‘honest’’ responses do not always reflect
semantically related words and were later asked to make ‘‘truthful’’ facts because of occasional memory errors.
a recognition judgment of old words, semantically related non- It is known that human memory is prone to various kinds of
studied words (lures for false recognition), and unrelated new distortions and illusions (Roediger 1996; Schacter 1999; Loftus
words. They were also asked to make a deceptive response to half 2003). Among these memory errors, many researchers have
of the old and unrelated new words. There were 3 main findings. focused on false recognition, whereby people incorrectly claim
First, consistent with the notion that executive function supports that they have recently seen or heard a stimulus they have not
deception, 2 types of deception (pretending to know and pretending encountered (Underwood 1965). In contrast to deception,
not to know) recruited prefrontal activity. Second, consistent with false recognition is not accompanied by a subjective feeling
the sensory reactivation hypothesis, the difference between true that people are responding untruthfully, and therefore
recognition and false recognition was found in the left temporo- researchers need to be able to detect a difference between
parietal regions probably engaged in the encoding of auditorily true and false recognition that is not apparent to the conscious
presented words. Third, the left prefrontal cortex was activated mind. Despite this challenge, many neuroimaging studies have
during pretending to know relative to correct rejection and false shown a difference in brain activities related to true and false
recognition, whereas the right anterior hippocampus was activated recognition (Schacter et al. 1996, 1997; Heun et al. 2000, 2004;
during false recognition relative to correct rejection and pretending Cabeza et al. 2001; von Zerssen et al. 2001; Okado and Stark
to know. These findings indicate that fMRI can detect the 2003; Kahn et al. 2004; Slotnick and Schacter 2004; Umeda et al.
difference in brain activity between deception and false memory 2005; Garoff-Eaton et al. 2006, 2007; Kim and Cabeza 2007).
despite the fact that subjects respond with ‘‘I know’’ to novel The main purpose of this study was directly to compare
events in both processes. brain activity related to deception and memory distortion, both
of which conceal the truth. Specifically, we focused on the
Keywords: false recognition, fMRI, lying, medial temporal lobe,
processes of pretending to know (a type of deception) and
prefrontal cortex
false recognition, in both of which people respond ‘‘I know’’ to
novel events. When distinguishing between these processes, it
is not possible to rely on the difference between subjects’
Introduction responses or the nature (‘‘novel’’ in this situation) of the stimuli.
The development of neuroimaging techniques has enabled us The ability to judge objectively the truthfulness of someone’s
directly to measure brain activity associated with various word based solely on their brain activations would help us to
cognitive functions. In recent years, much attention has been improve existing lie-detection systems on both theoretical and
paid not only to clarifying the neural correlates of cognitive practical bases. In addition, imaging data may provide useful
processes but also to ascertaining what someone is currently evidence for those involved in criminal investigation and
thinking by analyzing patterns of neural activity (Haynes and prosecution in terms of the credibility of eyewitness testimony
Rees 2006). In the context of this ‘‘brain reading,’’ discrimina- (Wells and Olson 2003). Despite this substantial implication,
tion between honest and deceptive responses is one of the there has been no neuroimaging study directly comparing the
most interesting topics in cognitive neuroscience. patterns of brain activities associated with deception and false
The neural correlates of deception have gradually been memory.
delineated in studies using positron emission tomography (Abe We performed an event-related fMRI study with a modified
et al. 2006, 2007), functional magnetic resonance imaging version of the set of word lists developed by Deese (1959) and
(fMRI; Spence et al. 2001; Langleben et al. 2002; Lee et al. 2002, Roediger and McDermott (1995) to produce false recognition
2005; Ganis et al. 2003; Kozel, Padgett, et al. 2004; Kozel, Revell, with high probability. Before fMRI scanning, participants heard
et al. 2004; Nunez et al. 2005; Phan et al. 2005; Mohamed et al. a number of lists consisting of semantic associates (e.g., moon,
2006; Gamer et al. 2007), event-related potential (Johnson et al. light, shine, bright, hot, gleam, etc). During a subsequent test
2003, 2004, 2005, 2008), and, more recently, transcranial direct phase with fMRI scanning, subjects were visually presented
current stimulation (Priori et al. 2008). In addition, researchers with previously studied ‘‘True targets’’ (e.g., hot), nonstudied
have extended their findings to the application of fMRI or ‘‘False targets’’ (e.g., sun) that were semantically related to the
magnetoencephalography as a lie detector on the level of studied items (i.e., lures for false recognition), and unrelated
individual subjects (Davatzikos et al. 2005; Kozel et al. 2005; ‘‘New targets’’ (e.g., building). The subjects’ task was to make an
Ó 2008 The Authors
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old/new judgment in response to these stimuli. In addition to words as ‘‘False targets’’ in the 40 lists of 15 words each. In addition, the
this procedure, the experimental condition of ‘‘Lie’’ was theme word and the 1st, 2nd, and 3rd strongest associates were used as
designed in which participants had to make deceptive ‘‘New targets’’ in the 20 lists of 10 words each. These stimuli were
visually presented to participants in the MRI scanner. Each test word
responses to ‘‘True targets’’ and ‘‘New targets’’ (i.e., ‘‘do not was presented in the center of a screen, and whichever finger of the
know’’ responses for ‘‘True targets’’ and ‘‘know’’ responses for right hand was to be used for old/new button pressing was presented
‘‘New targets’’). on both sides of the word. All the ‘‘False targets’’ and half of the ‘‘True
Before starting this experiment, we presumed the following targets’’ and ‘‘New targets’’ were assigned to ‘‘Truth’’ blocks, in which
hypotheses related to brain activations associated with true subjects were asked to respond honestly to each stimulus. The
memory, false memory, and deceptive responses. First, we remaining halves of the ‘‘True targets’’ and ‘‘New targets’’ were assigned
to ‘‘Lie’’ blocks, in which subjects were asked to make a deceptive
expected that deception would be associated with increased
response to each stimulus. Due to the limited number of stimuli, ‘‘False
brain activity in the lateral prefrontal cortex, which probably targets’’ were not presented in the ‘‘Lie’’ blocks.
reflects executive function such as cognitive control (Spence
et al. 2004; Hughes et al. 2005). Second, we expected that the
regions responsible for auditory processing would be prefer- Tasks
entially active during true versus false recognition due to The experiment consisted of 2 sessions, each of which included 1 study
phase and 1 test phase (i.e., the 1st study phase without fMRI, the
sensory reactivation (Schacter and Slotnick 2004), because true 1st test phase with fMRI, the 2nd study phase without fMRI, and
memories engage perceptual encoding processes that are the 2nd test phase with fMRI). Figure 1 illustrates 1 of the 2 task
presumably not involved in the creation of false memories. In sessions.
fact, recent neuroimaging studies have consistently shown that Before the experiment, the participants were given a thorough
the regions engaged in encoding were reactivated during explanation of the task procedure, and familiarized with the task by
memory retrieval (e.g., Johnson and Rugg 2007; Ueno et al. completing a short practice session. During each study phase, subjects
listened to a total of 300 words (20 lists of 15 words each) at a rate of
2007). The present experimental paradigm is suitable for this
2 s per word. The theme words from the 20 lists were not presented
investigation because the participants were asked to make during the study phase and were used as ‘‘False targets’’ for producing
a recognition judgment to visually presented words that had false recognition in the subsequent test phase. Words were presented
been studied auditorily. Third, we expected that, compared in order of decreasing strength of association with the theme word,
with false recognition and correct rejection (baseline condition except for 2 words to be used as ‘‘True targets,’’ which were shifted to
presenting novel stimuli to participants), the process of positions other than 1, 2, 14, and 15 to prevent primacy and recency
effects. Subjects were instructed to remember the presented words for
pretending to know would be associated with greater activity
the later recognition memory test. Presentation of each list was
in the lateral prefrontal cortex due to the increased demand of separated by a 15-s interval during which subjects performed simple
executive function. Finally, we expected that, compared with arithmetic (e.g., ‘‘5 plus 2’’). The presentation order of the lists was
pretending to know and correct rejection, false recognition randomized across subjects.
would be associated with greater activity in the medial Approximately 10 min after the completion of the study phase, the
temporal lobe structures responsible for subjective familiarity test phase was initiated. During the test phase with fMRI scanning,
and with no reactivation of regions involved in the sensory subjects performed the recognition test, consisting of 4 ‘‘Truth’’ and 4
‘‘Lie’’ blocks. The cues indicating the subsequent ‘‘Truth’’ or ‘‘Lie’’ block
processing of auditory materials. were of 15 s duration and were followed by the target words. In each
‘‘Truth’’ block, 5 ‘‘True targets,’’ 5 ‘‘False targets,’’ and 5 ‘‘New targets’’
were presented in randomized order, and subjects were asked to make
Materials and Methods old/new decisions as quickly as possible by pressing keys on the
response box. In each ‘‘Lie’’ block, 5 ‘‘True targets’’ and 5 ‘‘New targets’’
Participants were presented in randomized order, and subjects were asked to make
Twenty-eight volunteers were recruited to take part in this study. The deceptive responses (i.e., ‘‘new’’ responses for ‘‘True targets’’ and ‘‘old’’
criteria of recruitment for this study were 1) native Japanese speakers, responses for ‘‘New targets’’). Subjects were asked to respond using the
2) no history of neurological or psychiatric diseases, and 3) right- index or middle finger of their right hand. The assignment of these
handedness on the Edinburgh Handedness Inventory (Oldfield 1971). fingers for each old/new decision was counterbalanced across blocks.
All participants gave their written informed consent in accordance with Each word was presented for 2 s, and the intervals between the words,
the Declaration of Helsinki and the guidelines approved by the Ethical during which cross-fixation was constantly presented, ranged between
Committee of Tamagawa University. 2.5 s and 13.5 s to maximize the efficiency of the event-related design
(Dale 1999).
In 1 test phase with fMRI scanning, participants responded to 20
Stimuli ‘‘True,’’ 20 ‘‘False,’’ and 20 ‘‘New’’ target words in ‘‘Truth’’ blocks, and 20
For false recognition (Deese 1959; Roediger and McDermott 1995), we ‘‘True’’ and 20 ‘‘New’’ trials in ‘‘Lie’’ blocks. Thus, across the 2 test
collated 81 word lists, each consisting of semantically related words, phases, participants responded to 40 ‘‘True,’’ 40 ‘‘False,’’ and 40 ‘‘New’’
from the Japanese literature (Hamajima 2000; Takahashi 2001; Hoshino target words in ‘‘Truth’’ blocks, and 40 ‘‘True’’ and 40 ‘‘New’’ trials in
2002; Miyaji and Yama 2002). By removing overlapping lists or words ‘‘Lie’’ blocks.
and including some lists with 1 or 2 words from the Japanese word
association norms (Umemoto 1969), we eventually prepared 60 lists of
semantic associates. Of these, 40 lists consisted of 1 theme word (e.g., Image Acquisition and Data Analysis
sun) and 15 semantically related words (e.g., moon, light, shine, bright, Whole-brain imaging was performed with a 1.5-Tesla MRI scanner
hot, and so forth), and the remaining 20 lists consisted of 1 theme word (Magnetom Sonata; Siemens, Erlangen, Germany). A T2*-weighted echo
and 10 semantically related words. planar imaging (EPI) sequence was used for functional imaging with the
During 2 study phases, a total of 600 words (40 lists of 15 words following parameters: time repetition = 2200 ms, time echo = 45 ms,
each) prerecorded by a male speaker were presented to the flip angle = 90°, 64 3 64 acquisition matrix, field of view = 192 mm, 26
participants through a personal computer. axial slices with 4 mm slice thickness and 1 mm interslice gap, 400
During 2 test phases, a total of 200 words (80 ‘‘True targets,’’ 40 volume acquisitions per run. Head motion was restricted using firm
‘‘False targets,’’ and 80 ‘‘New targets’’) were presented. The 1st and padding that surrounded the head. The cognitive tasks during fMRI
2nd strongest associates were used as ‘‘True targets’’ and the theme scanning were controlled using Cogent 2000 software (Wellcome

2812 False Memory and Deception d

Abe et al.
Figure 1. Depiction of 1 of the 2 task sessions (see Materials and Methods for details). The study-test phase was conducted twice with different stimulus sets. After
participants heard the word lists of semantic associates in the study phase, they were asked to perform the recognition memory task (test phase) consisting of 4 ‘‘Truth’’ and 4
‘‘Lie’’ blocks with fMRI scanning. In the ‘‘Truth’’ blocks, they were asked to respond honestly to ‘‘True targets’’ (old words from the study phase), ‘‘False targets’’ (nonstudied
words that were semantically related to old words), and ‘‘New targets’’ (new words that were not semantically related to old words). In the ‘‘Lie’’ blocks, they were asked to
dishonestly respond to ‘‘True targets’’ and ‘‘New targets.’’ In the ‘‘Truth blocks,’’ true recognition (TR) was defined as an ‘‘old’’ response to a ‘‘True target,’’ false recognition (FR) as
an ‘‘old’’ response to a ‘‘False target,’’ and correct rejection (CR) as a ‘‘new’’ response to a ‘‘New target.’’ In the ‘‘Lie’’ blocks, lying to ‘‘True targets’’ (LT; i.e., pretending not to
know) and lying to ‘‘New targets’’ (LN; i.e., pretending to know) were defined as deceptive responses to each target.

Department of Imaging Neuroscience, London, UK). Visual stimuli individually significant. For all the whole-brain subtraction analyses,
were projected onto a screen and viewed through a mirror attached to the threshold of significance was set at P < 0.001 (uncorrected for
a standard head coil. The subjects’ responses were collected using multiple comparisons) with an extent threshold of 10 contiguous
a magnet-compatible response box. voxels. In the conjunction analysis, the same threshold was used (P <
Data preprocessing and statistical analyses were performed using 0.001), but no extent threshold was applied. To extract the percent
Statistical Parametric Mapping 2 (Wellcome Department of Imaging signal change of activated regions during each task, we also used
Neuroscience). Preprocessing of the image volumes included re- MarsBaR software (Brett et al. 2002).
alignment of head motions, slice-time correction with reference to
the middle slice acquired in time, normalization to the EPI-template
based on the Montreal Neurological Institute (MNI) reference brain
Results
(resampled voxel size 3 3 3 3 3 mm3), and spatial smoothing with
a Gaussian kernel (8 mm at full-width half-maximum). Participants
The fMRI data were analyzed using an event-related model. For each Before the analysis of imaging data, 2 participants were
subject, activity associated with each experimental condition of excluded from the analysis due to excessive head motion
interest (i.e., TR, true recognition to ‘‘True targets’’; CR, correct
during fMRI scanning (approximately 4 mm). An additional
rejection to ‘‘New targets’’; FR, false recognition to ‘‘False targets’’; LT,
lying to ‘‘True targets’’; LN, lying to ‘‘New targets’’) was modeled using 6 participants were excluded due to poor task performance
a canonical hemodynamic response function. Targets that were (i.e., less than 60% accuracy in ‘‘Truth’’ blocks, which was close
incorrectly classified (i.e., error responses) or for which a response to chance level) or an insufficient number of events in at least 1
was omitted were modeled as events of no interest, as were of the conditions used in the imaging contrasts (i.e., fewer
instructions presented during the onset of ‘‘Truth’’ and ‘‘Lie’’ blocks. A than 15), or both. Thus the results of the present study are
high-pass filter of 1/128 Hz was used to remove low-frequency noise, based on the data from the remaining 20 subjects (11 males and
and an AR (1) model corrected for temporal autocorrelation. The
resulting parameter estimates for each regressor at each voxel were
9 females, age range 19--28 years, mean age 21.9 years). These
then entered into a 2nd-level analysis where each participant served participants did not report any difficulty understanding the task
as a random effect in a repeated measures analysis of variance procedure and performing each of the Truth and Lie tasks as
(ANOVA). Appropriate corrections were made for nonsphericity and instructed. There were no pathological findings on MRI of any
correlated repeated measures (Friston, Glaser, et al. 2002; Friston, of the subjects’ brains.
Penny, et al. 2002). The comparisons between experimental conditions
were then performed by appropriately weighted linear contrasts and Comparison between Truthful and Deceptive Responses
determined on a voxel-by-voxel basis. In addition to simple subtraction
analyses, the above procedure allowed us to perform conjunction
Behavioral Data
analyses at the 2nd level, and we identified activated regions with
a conjunction using the minimum statistic (Friston et al. 2005), as All the behavioral data are shown in Table 1. In this analysis, we
suggested by Nichols et al. (2005). This procedure revealed areas in used 2-way repeated measures ANOVA to analyze the
which all the contrasts entered into conjunction analysis were behavioral data of 2 ‘‘Truth’’ and 2 ‘‘Lie’’ tasks except for the

Cerebral Cortex December 2008, V 18 N 12 2813

condition of FR. A main effect of stimulus type (‘‘True targets’’ 0.399). Results for the right superior frontal gyrus [24, –3, 57]
or ‘‘New targets’’) was found in the accuracy of the response, as showed a significant main effect of ‘‘Lie’’ (F1,19 = 23.866, P <
the average accuracy was significantly higher for ‘‘New targets’’ 0.0005), but showed neither a main effect of familiarity of stimuli
than for ‘‘True targets’’ (F1, 19 = 6.462, P < 0.05). A noticeable (F1,19 = 0.694, P = 0.415) nor an interaction (F1,19 = 0.021, P =
trend of response type (‘‘Truth’’ or ‘‘Lie’’ tasks) qualified by 0.886). Results for the left inferior frontal gyrus [–45, 48, –15]
a relatively higher accuracy in ‘‘Truth’’ conditions was also showed a significant main effect of ‘‘Lie’’ (F1,19 = 20.538, P <
found (F1, 19 = 3.662, P = 0.071). For reaction times, a main 0.0005), and a main effect of familiarity of stimuli (‘‘True
effect of stimulus type was evident, as the average reaction targets’’ > ‘‘New targets’’) (F1,19 = 10.911, P < 0.005) without an
time was significantly longer for ‘‘New targets’’ than for ‘‘True interaction (F1,19 = 0.002, P = 0.968). Results for the left
targets’’ (F1, 19 = 24.034, P < 0.0001). There was also a significant supplementary motor area [–12, 21, 57] showed a significant
main effect of response type, characterized by a longer reaction main effect of the ‘‘Lie’’ (F1,19 = 21.678, P < 0.0005) and a
time in ‘‘Lie’’ tasks (F1, 19 = 15.136, P < 0.005). There was no marginal main effect of familiarity of stimuli (‘‘True targets’’ >
interaction between the 2 factors in terms of both accuracy ‘‘New targets’’) (F1,19 = 4.015, P = 0.060), without an interaction
and reaction time. These results indicate that the cognitive (F1,19 = 0.179, P = 0.677). Results for the left middle
demand in ‘‘Lie’’ tasks was higher than that in ‘‘Truth’’ tasks. frontal gyrus [–27, 3, 60] showed a significant main effect of
‘‘Lie’’ (F1,19 = 24.276, P < 0.0001), and a main effect of familiarity
Brain Activation of stimuli (‘‘True targets’’ > ‘‘New targets’’) (F1,19 = 12.683, P <
To examine whether deception was associated with increased 0.005), without an interaction (F1,19 = 0.139, P = 0.713). These
brain activity in the prefrontal cortex, we compared LT and LN results are illustrated in Figure 2.
with TR and CR (i.e., main effect of deception). As predicted,
increased prefrontal activations were observed in this contrast. Comparison between True and False Recognition
Table 2 summarizes these data for anatomical structures and
Brodmann’s area (BA), MNI coordinates, Z-values and cluster Behavioral Data
size of peak activations. In this analysis, we used 1-way repeated measures ANOVA to
Second, to examine the influence of the familiarity of stimuli examine the behavioral data of TR, FR, and CR. Note that the
on fMRI signals in each activated frontal region and whether or accuracy of the task in which participants responded to
not an interaction occurred, we performed the region of nonstudied ‘‘False targets’’ was assessed by the rate of correct
interest (ROI) analysis. The signal changes in each cluster were rejection in response to these stimuli. An ANOVA yielded
extracted and analyzed using 2-way ANOVA with the response a significant effect of target type in accuracy (F2, 38 = 140.698,
to stimuli (‘‘Truth,’’ ‘‘Lie’’) and the familiarity of stimuli (‘‘True P < 0.0001). A post hoc test (Scheffe) revealed significant
targets,’’ ‘‘New targets’’) as factors. Overall, there was no differences between true recognition of ‘‘True targets’’ and
significant interaction between the 2 factors in the entire correct rejection of ‘‘New targets’’ (P < 0.05), between correct
activated regions, indicating that these regions are commonly rejection of ‘‘False targets’’ and correct rejection of ‘‘New
associated with the 2 types of deception. Results of the ANOVA targets’’ (P < 0.0001), and between true recognition of ‘‘True
for the right medial prefrontal cortex [12, 57, –6] showed targets’’ and correct rejection of ‘‘False targets’’ (P < 0.0001). An
a significant main effect of ‘‘Lie’’ (F1,19 = 18.281, P < 0.0005), but ANOVA for reaction time also showed a significant effect of
showed neither a main effect of familiarity of stimuli (F1,19 = target type (F2, 38 = 13.731, P < 0.0001). A post hoc test
0.629, P = 0.438) nor an interaction between the 2 factors (Scheffe) revealed significant differences between TR and CR
(F1,19 = 0.075, P = 0.787). ANOVA for the right superior frontal
gyrus [24, 15, 60] yielded similar results: a significant main
effect of ‘‘Lie’’ (F1,19 = 24.107, P < 0.0001), without a main
Table 2
effect of familiarity of stimuli (F1,19 = 1.943, P = 0.179) or an Brain regions showing main effect of making a deceptive responses {(LT
TR) þ (LN
CR)}
interaction (F1,19 = 1.805, P = 0.195). Results for the right
middle frontal gyrus [42, 6, 51] showed a significant main effect Region (BA) MNI coordinates Z value Cluster size
of ‘‘Lie’’ (F1,19 = 13.112, P < 0.005) and a marginal main effect of x y z
familiarity of stimuli (‘‘True targets’’ > ‘‘New targets’’) (F1,19 = Rt medial prefrontal cortex (10) 12 57
6 3.86 16
3.425, P = 0.080), without an interaction (F1,19 = 0.746, P = Rt insula 33 21 15 4.13 14
Rt superior frontal gyrus (8) 24 15 60 3.90 11
Rt middle frontal gyrus (6) 42 6 51 3.91 24
Rt superior frontal gyrus (6) 24
3 57 3.81 38
Table 1 Rt thalamus 15
27 0 4.22 34
Percent correct and reaction time for all conditions Rt fusiform gyrus (19) 24
69
12 3.80 16
Rt cuneus (19) 18
75 39 3.68 44
Percent correct Reaction time Lt inferior frontal gyrus (47)
45 48
15 3.58 14
Lt supplementary motor area (6)
12 21 57 4.07 73
Mean SD Mean SD Lt insula
30 21 9 3.52 17
Lt insula
48 9 6 3.67 16
Truth Lt middle frontal gyrus (6/8/9)
27 3 60 4.01 140
True targets 71.8 ±8.7 1652 ±351 Lt postcentral gyrus (3)
54
6 39 4.16 22
New targets 81.1 ±9.4 1892 ±417 Lt superior temporal sulcus/superior
42
27 3 3.77 12
False targets 28.9 ±12.2 1647 ±370 temporal gyrus (22)
Lie Lt supramarginal gyrus (40)
30
42 45 4.00 20
True targets 70.6 ±11.4 1959 ±415 Lt superior parietal lobule (7)
27
57 54 4.12 35
New targets 74.8 ±14.3 2168 ±533 Lt angular gyrus (39)
51
60 36 4.00 38
Lt middle occipital gyrus (19)
30
69 30 3.96 59
Note: The accuracy of subjects’ responses to ‘‘False targets’’ was assessed by the rate of correct Lt precuneus (7)
9
78 45 3.86 49
rejection (‘‘new’’ responses), but the reaction time was based on the trials of false recognition
(‘‘old’’ responses). Note: Only the most significant peaks within each area of activation are reported in this table.

2814 False Memory and Deception d

Abe et al.
Figure 2. Regions showing greater activation during lying (LT and LN) relative to truth telling (TR and CR). The signal changes of the following 7 activated regions in the frontal
lobe are depicted (error bars represent SEM). (a) Right medial prefrontal cortex [12, 57,
6], (b) right superior frontal gyrus [24, 15, 60], (c) right middle frontal gyrus [42, 6, 51],
(d) right superior frontal gyrus [24,
3, 57], (e) left inferior frontal gyrus [
45, 48,
15], (f) left supplementary motor area [
12, 21, 57], (g) left middle frontal gyrus [
27, 3,
60]. TR, true recognition; CR, correct rejection; LT, lying to ‘‘True targets’’ (pretending not to know); LN, lying to ‘‘New targets’’ (pretending to know).

(P < 0.0005) and between CR and FR (P < 0.0005). No Table 3

difference was observed between TR and FR (P = 0.996). These Brain regions showing greater responses during TR compared with FR
results show a high rate of false recognition of nonpresented
Region (BA) MNI coordinates Z value Cluster size
‘‘False targets’’ (approximately 70%). The fact that there was no
difference in reaction time between TR and FR indicates that x y z
the difference in brain activity associated with these processes Rt middle temporal gyrus (21/22) 66
21
3 3.88 24
cannot be ascribed to retrieval effort. *Rt/Lt medial superior frontal gyrus (10) 0 60 24 3.53 18
*Rt/Lt cerebellum 0
45
42 3.97 17
*Lt superior temporal sulcus/middle
48
9
15 3.65 11
Brain Activation temporal gyrus (21)
*Lt middle temporal gyrus (21)
60
18
12 3.88 12
Brain activity during TR was compared with that during FR. The *Lt supramarginal gyrus (40)
51
54 36 3.63 13
results are shown in Table 3. Consistent with our hypothesis,
activations in the temporal and parietal lobes were found in the Note: Only the most significant peaks within each area of activation are reported in this table.
*Indicates the region that includes the active voxels detected in the conjunction analysis of TR
bilateral hemisphere. To obtain more reliable evidence for the versus FR and TR versus CR.
interpretation of sensory reactivation during TR, we performed
conjunction analysis of TR versus FR and TR versus CR. If
the difference between TR and FR were a true reflection of superior temporal sulcus/middle temporal gyrus [–48, –9, –15]
the reactivation of sensory information acquired during the showed a significant difference (F2, 38 = 9.502, P < 0.001). A
encoding phase, the observed activations would also be post hoc test (Scheffe) revealed that the activity of TR was
detected in the contrast of TR versus CR. This conjunction higher than that of CR (P < 0.01) and FR (P < 0.005), whereas
analysis revealed that the activations in the left temporoparietal the difference between CR and FR was not significant (P =
regions during TR relative to FR overlapped with the 0.873). Second, results of the ANOVA for the left middle
activations during TR relative to CR. temporal gyrus [–60, –18, –12] showed a significant difference
To confirm further this activation overlap, we extracted the (F2, 38 = 11.293, P < 0.0005). A post hoc test (Scheffe) revealed
signal change in each cluster of 3 left temporoparietal regions that the activity of TR was higher than that of CR (P < 0.005)
during each condition and analyzed it using 1-way repeated and FR (P < 0.0005), whereas the difference between CR and
measures ANOVA. First, results of the ANOVA for the left FR was not significant (P = 0.770). Third, results of the ANOVA

Cerebral Cortex December 2008, V 18 N 12 2815

for the left supramarginal gyrus [–51, –54, 36] showed Brain Activation
a significant difference (F2, 38 = 13.688, P < 0.0001). A post Comparison between LN and FR was the main purpose of this
hoc test (Scheffe) revealed that the activity of TR was higher study. First, we compared the neural activities during LN with
than that of CR (P < 0.0005) and FR (P < 0.001), whereas the those during CR, and found significant activations in the
difference between CR and FR was not significant (P = 0.853). prefrontal cortex. The results are summarized in Table 4. If the
These results are illustrated in Figure 3. prefrontal activities found in LN versus CR were also detected
in LN versus FR, those regions would be a reliable indicator of
Comparison between Deception and False Memory the process of intentional response manipulation characteriz-
ing deception. Conjunction analysis of these 2 contrasts
Behavioral Data revealed that activation in the left middle frontal gyrus
In the analysis of the difference between deception and false detected in LN versus CR overlapped with the activation
memory, we used 1-way repeated measures ANOVA to detected in LN versus FR.
compare the behavioral data of LN, FR, and CR. As mentioned To further confirm this activation overlap, we extracted the
above, the accuracy of the condition in which participants signal change in the cluster of the left middle frontal gyrus [–42,
responded to nonstudied ‘‘False targets’’ was assessed by the 18, 39] during each condition and analyzed it using 1-way
rate of correct rejection of these stimuli. An ANOVA yielded repeated measures ANOVA. Results of the ANOVA showed
a significant effect of accuracy (F2, 38 = 158.569, P < 0.0001). A significant difference of signal change in this region (F2, 38 =
post hoc test (Scheffe) revealed significant differences between 13.209, P < 0.0001). A post hoc test (Scheffe) revealed that
correct rejection of ‘‘False targets’’ and correct rejection of the activity of LN was higher than that of CR (P < 0.0001)
‘‘New targets’’ (P < 0.0001), and between lying in response to and FR (P < 0.05), and a trend was found between CR and FR
‘‘New targets’’ and correct rejection of ‘‘False targets’’ (P < (P = 0.092).
0.0001). No difference was observed between correct rejection As for FR, we compared the neural activities during FR with
of ‘‘New targets’’ and lying in response to ‘‘New targets’’ (P = those during CR, and found significant activation in the right
0.152). An ANOVA for reaction time also showed a significant anterior hippocampus. The results are summarized in Table 5.
effect (F2, 38 = 33.483, P < 0.0001). A post hoc test (Scheffe) If the activations of the medial temporal lobe found in FR
revealed that the reaction time for CR was longer than that for versus CR were also detected in FR versus LN, the region would
FR (P < 0.005), and that the reaction time for LN was longer be a reliable indicator of illusory familiarity characterizing false
than that for CR (P < 0.001) and that for FR (P < 0.0001). These
data for reaction time indicate that although all the stimuli in
these 3 conditions were novel to the participants, the process Table 4
of classifying the stimuli as ‘‘New’’ (i.e., CR) was more Brain regions showing greater responses during LN compared with CR
complicated than that of classifying the stimuli as ‘‘Old’’
Region (BA) MNI coordinates Z value Cluster
(i.e., FR). Furthermore, the longer reaction time for LN relative size
to that for CR suggests that the additional process of lying x y z
further enhanced cognitive demand during the recognition Rt superior frontal sulcus (8) 24 15 39 4.37 14
memory task. Rt thalamus 18
27 3 3.49 12
Lt supplementary motor area (6)
12 21 54 3.32 11
Lt insula
27 21 12 3.75 16
*Lt middle frontal gyrus (9)
42 18 39 4.25 48
Lt superior frontal sulcus (6)
27 9 42 3.62 11
*Lt supramarginal gyrus (40)
51
57 36 3.68 22
Lt middle occipital gyrus (19)
33
69 33 3.84 12

Note: Only the most significant peaks within each area of activation are reported in this table.
*Indicates the region that includes the active voxels detected in the conjunction analysis of LN
versus CR and LN versus FR.

Table 5
Brain regions showing greater responses during FR compared with CR

Region (BA) MNI coordinates Z value Cluster

size
x y z
*Rt basal forebrain/orbitofrontal cortex (12/25) 9 6
9 4.23 35
*Rt hippocampus 36
9
15 3.98 12
*Lt orbitofrontal cortex (11/12)
9 33
18 3.61 25
Lt orbitofrontal cortex (11)
21 27
24 4.28 23
Lt inferior frontal gyrus (45/47)
42 27 0 3.59 15
Lt superior frontal sulcus (8)
24 18 39 3.83 25
Lt insula
27 18 12 4.16 18
Lt caudate nucleus
12 3 9 3.49 15
Figure 3. Statistical parametric map of regions showing greater activation during TR Lt superior parietal lobule (7)
27
63 54 4.05 43
than during CR and FR, displayed on a surface-rendered standard brain. The signal *Lt cuneus (18)
18
63 21 4.04 127
changes of the following 3 activated regions are depicted (error bars represent SEM).
(a) Left supramarginal gyrus [
51,
54, 36], (b) left superior temporal sulcus/middle Note: Only the most significant peaks within each area of activation are reported in this table.
temporal gyrus [
48,
9,
15], (c) left middle temporal gyrus [
60,
18,
12]. *Indicates the region that includes the active voxels detected in the conjunction analysis of FR
TR, true recognition; CR, correct rejection; FR, false recognition. versus CR and FR versus LN.

2816 False Memory and Deception d

Abe et al.
memory. Conjunction analysis of these 2 contrasts revealed lateral temporal and parietal cortices. The conjunction analysis
that the activation in the right anterior hippocampus detected of TR versus FR and TR versus CR, which was performed to
in FR versus CR overlapped with the activation detected in FR obtain more reliable evidence related to sensory reactivation of
versus LN. the regions engaged in encoding of the word lists, further
To confirm further this activation overlap, we extracted the showed left-lateralized activations of the supramarginal gyrus
signal change in the cluster of the right hippocampus [36, –9, –15] and the middle temporal gyrus. Previous neuroimaging studies
during each condition and analyzed it using 1-way repeated employing similar word lists of semantic associates for pro-
measures ANOVA. Results of the ANOVA showed significant ducing false recognition have also reported comparable
difference of signal change in this region (F2, 38 = 10.826, P < findings. For example, Schacter et al. (1996) reported increased
0.0005). A post hoc test (Scheffe) revealed that the activity of blood flow in the left temporoparietal cortex (BA 42/22/40)
FR was higher than that of CR (P < 0.005) and LN (P < 0.001), during true recognition relative to false recognition, and
whereas the difference between CR and LN was not significant Cabeza et al. (2001) reported a difference in activity in the
(P = 0.907). In addition, as we expected, the comparison left parietal cortex (BA 40/39) between true and false
between FR versus CR revealed no reactivation of the regions recognition. Considering the role of the left temporoparietal
responsible for processing the sensory information of auditorily areas in the language processing of words presented auditorily,
presented word lists, such as language-processing areas in the the left temporoparietal activity can be regarded as a reliable
left hemisphere. The signal change of the activation in the left neural signature of true recognition.
middle frontal gyrus and the right hippocampus during each The main purpose of this study was to compare the brain
task is illustrated in Figure 4. activity related to the process of pretending to know and false
recognition in order to identify a reliable indicator of the brain
activations associated with each process. We expected that the
Discussion
activations of the prefrontal cortex detected in the analysis of
In the present study, we aimed to clarify the neural correlates LN versus CR would also be detected in the analysis of LN versus
of true memory, false memory, and deception. As for the FR, and that the activations of medial temporal lobe responsible
cerebral mechanisms underlying deception, the present data for mnemonic processing detected in the analysis of FR versus
show that the process of intentional response manipulation in CR would also be detected in the analysis of FR versus LN. To
deception was characterized by prefrontal activity. These confirm these predictions, we performed 2 conjunction analyses
findings are highly consistent with those of previous neuro- (i.e., LN vs. CR conjunct with LN vs. FR, and FR vs. CR conjunct
imaging studies, indicating a robust contribution of executive with FR vs. LN) and ROI analyses for each activation.
function to deception (Spence et al. 2004; Hughes et al. 2005). As we expected, activation of the left middle frontal gyrus,
Significant activations were also detected in the supplementary possibly reflecting the subjective, intentional cognitive process
motor area, which plays a role in higher motor control (Tanji of response manipulation, was found in the conjunction
1994). This activity may reflect motor regulation during button analysis of LN versus CR and LN versus FR. The middle frontal
pressing when making deceptive responses. gyrus, often referred to as the dorsolateral prefrontal cortex,
In relation to the difference in neural activity between can be regarded as a reliable indicator of pretending to know.
veridical and illusory memories, the comparison of true Expanding on the previous neuroimaging studies on deception,
recognition with false recognition revealed activations of the our data suggest that prefrontal activity reflects not only the
difference between deception and truth telling, but also the
difference between deception and false memory.
Consistent with our hypothesis, conjunction analysis of FR
versus CR and FR versus LN revealed activations of the right
anterior hippocampus without reactivation of the regions
responsible for language processing detected as true recogni-
tion-specific activity. This activation pattern indicates that the
right anterior hippocampus is associated with ‘‘illusory’’
familiarity to novel stimuli. The results also appear to be
consistent with the recent fMRI finding that medial temporal
lobe activity is modulated not only by objective memory
function but also by the subjective confidence level of
recognition memory (Chua et al. 2006). One surprising result
was that hippocampal activity was found not in the left but in
the right hemisphere, despite the fact that verbal materials
have to be retrieved. Although it is difficult to explain this
finding from the available data, there is a possibility that the
right anterior hippocampus plays a role in relatively rough
judgment of episodic familiarity without access to memory
fragments of perceptual traces stored in other cortical areas.
Figure 4. The signal change of increased brain activity in (a) the left middle frontal In conclusion, the present fMRI study provided evidence of
gyrus [
42, 18, 39] during LN in comparison to during CR and FR, and (b) the right neural activities differentiating between true memory, false
hippocampus [36,
9,
15] during FR in comparison to during CR and LN. Error bars
represent standard error. The activation is superimposed onto MRIs of MNI
memory, and deception. The most important finding is that the
templates. CR, correct rejection; FR, false recognition; LN, lying to ‘‘New targets’’ left prefrontal cortex was activated during pretending to know
(pretending to know). relative to both correct rejection and false recognition,

Cerebral Cortex December 2008, V 18 N 12 2817

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Century Center of Excellence (COE) Program by the Japan Psychiatry Res. 99:137--150.
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Notes
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