Neural Mechanism for Mirrored Self-face Recognition
Cerebral Cortex September
Neural Mechanism for Mirrored Self-face Recognition
Motoaki Sugiura 1 2
Carlos Makoto Miyauchi 0 2
Yuka Kotozaki 2
Yoritaka Akimoto 2
Takayuki Nozawa 2
Yukihito Yomogida 3 4
Sugiko Hanawa 2
Yuki Yamamoto 2
Atsushi Sakuma 2
Seishu Nakagawa 2
Ryuta Kawashima 2
0 Graduate Schools for Law and Politics, The University of Tokyo , Tokyo 113-0033 , Japan
1 International Research Institute of Disaster Science, Tohoku University , Sendai 980-8575 , Japan
2 Institute of Development , Aging and Cancer
3 Tamagawa University Brain Science Institute , Tokyo 194-8610 , Japan
4 Japan Society for the Promotion of Science , Tokyo 102-8472 , Japan
Self-face recognition in the mirror is considered to involve multiple processes that integrate 2 perceptual cues: temporal contingency of the visual feedback on one's action (contingency cue) and matching with self-face representation in long-term memory (figurative cue). The aim of this study was to examine the neural bases of these processes by manipulating 2 perceptual cues using a “virtual mirror” system. This system allowed online dynamic presentations of realtime and delayed self- or other facial actions. Perception-level processes were identified as responses to only a single perceptual cue. The effect of the contingency cue was identified in the cuneus. The regions sensitive to the figurative cue were subdivided by the response to a static self-face, which was identified in the right temporal, parietal, and frontal regions, but not in the bilateral occipitoparietal regions. Semantic- or integration-level processes, including amodal self-representation and belief validation, which allow modality-independent self-recognition and the resolution of potential conflicts between perceptual cues, respectively, were identified in distinct regions in the right frontal and insular cortices. The results are supportive of the multicomponent notion of self-recognition and suggest a critical role for contingency detection in the co-emergence of self-recognition and empathy in infants.
contingency; face; fMRI; recognition; self
Human adults recognize their own faces in a mirror or on any
reflective surface. However, this ability has been demonstrated
in only a limited number of animal species (Gallup 1982; Reiss
and Marino 2001; Plotnik et al. 2006). Because these animals
typically have large brains and show evidence of empathic
behavior, mirrored self-recognition has been understood as an
engram of a unique social-cognitive function of a “self,” which
relies on a unique system in highly evolved brains (Gallup
1982; Marino 2002; Hart et al. 2008; Suddendorf and
CollierBaker 2009). This notion is consistent with the developmental
co-emergence of empathic behavior and mirrored
selfrecognition in human infants (Bischof-Köhler 1988;
ZahnWaxler et al. 1992).
In contrast to this unique self-system notion, several lines of
evidence suggest that multiple (at least 4) “ordinary” cognitive
processes underpin mirrored self-recognition at both
perceptual and semantic levels. At the perceptual level, there are 2
independent processes for detecting different perceptual cues
from a mirrored self-image: contingency and figurative cues.
The contingency cue is the temporal contingency between
one’s intentional facial action and the visually observed
movement feedback of one’s own face. The figurative cue is the
match between the perceived face and the self-face
representation stored in the visual long-term memory. The contingency
cue may trigger 2 additional distinct processes: the sense of
agency underpinning the perceived action (Wegner and
Wheatley 1999; Frith et al. 2000) and the recognition of the mirror,
which involves understanding that the space represented as
being behind the mirror surface is in fact a symmetrically
transformed duplicate of the real space in front (Loveland 1986).
The mechanism for the figurative-cue processing appears to
develop based on agency-related experience derived from the
contingency cue; infants begin self-recognition of
noncontingent images (e.g., photos and recorded videos) after they
acquire abilities related to contingent images (e.g., mirrors and
live videos) (Bigelow 1981; Courage et al. 2004).
At the semantic level, 2 independently processed perceptual
cues as perceptual-level processes are integrated to allow
metalevel self-cognition. In this integration stage, 2 complementary
processes are also at work. One process is the access to amodal
self-representation, which is activated by input from
perceptuallevel processes of any modality or types of self-relevant cues
(i.e., irrespective of the contingency or figurative cue). This
notion seems consistent with the notion of a special-self system.
The second process is the belief-validation process, which
evaluates the consistency between different self-relevant inputs
from a perceptual level (i.e., between contingency and figurative
cues). This hypothesis is based on the observation of
mirroredself misidentification in patients with dementia, in which a false
belief (i.e., that the person in the mirror is not the self)
generated by an impaired perceptual-level process is not corrected by
comparison with other perceptual or contextual information
(Coltheart 2010; Connors and Coltheart 2011).
No study has examined the neural basis of mirrored
selfrecognition by addressing this multicomponent structure,
whereas related studies have suggested the involvement of
different cortical regions, mostly in the right hemisphere (Sugiura
2013). The neuropsychological literature on mirrored-self
misidentification suggests the involvement of the right hemisphere
or frontal regions, but further anatomical specification has been
hampered by the severe dementia and global cortical
impairment of patients (Breen et al. 2001; Villarejo et al. 2011). Many
neuroimaging studies have explored the self-face-specific
response of the brain using noncontingent images (i.e., pictures
or recorded videos) and have thus addressed self-face
recognition based solely on a figurative cue. Indeed, recent studies have
typically reported self-face-specific activation in several regions
in the lateral frontal, parietal, occipital, temporal, and insular
cortices of the right hemisphere (Uddin et al. 2005; Sugiura
et al. 2006, 2008, 2012; Devue et al. 2007; Kaplan et al. 2008;
Oikawa et al. 2012). Neuroimaging studies assessing the sense
of action agency identified the neural response to a contingency
cue by contrasting the real-time and violated visual feedback of
the subject’s hand action; activation has been reported in the
insula and putamen (Farrer and Frith 2002; Farrer et al. 2003;
Leube et al. 2003). However, in these studies, the subjects were
not provided with the figurative cue or the unique visuospatial
experience of being faced with a mirror. Finally, 2 potential
semantic-level self-recognition processes have also been
implicated in the right lateral prefrontal cortex. Amodal
selfrepresentation has been suggested to involve a region in the
right inferior frontal gyrus or sulcus; this region responded not
only to the self-face but also to the self-voice (Nakamura et al.
2001) or to one’s own name being called (Kaplan et al. 2008).
The belief-validation process in general has been speculated
to involve the right middle frontal gyrus, which is responsive to
the violation of predictions based on a belief (Fletcher et al.
2001; Turner et al. 2004).
In this functional magnetic resonance imaging (MRI) study,
we directly examined the neural bases of self-face recognition
in a mirror. We installed a virtual mirror system in a MRI
scanner; the system featured an open-face head coil, video
camera, and a projection system capable of presenting
realtime, delayed, and recorded videos. This system allowed us to
independently manipulate contingency and figurative cues,
enabling us to dissociate the perceptual-level processes for 2 cues
and 2 semantic-level processes. During a simple facial action
task, each subject was presented with a video of the subject’s
own face (Self ) or a similar prerecorded video of an unfamiliar
face (Other). Each type of face stimuli was projected in
realtime (Real-time), with a 500-ms delay (Delayed), or as a still
image (Static). Of these 6 conditions for different types of
visual stimuli, 4 were designed for a 2-by-2 factorial design
composed of the factors Contingency (Real-time, Delayed) and
Face (Self, Other). The perceptual-level process of the
contingency cue was identified as the main effect of Contingency,
and that of the figurative cue as the main effect of Face. The
semantic-level processes were expected to show an interaction
between 2 factors, with the 2 potential processes
demonstrating different activation profiles. Amodal self-representation
should exhibit a lack of additive effect of 2 cues while being
sensitive to either cue alone; that is, an equivalent level of
activation should be observed among Real-time Self, Delayed
Self, and Real-time Other (i.e., Real-time contingency is a
selfrelevant input) compared with the Delayed Other condition.
The belief-validation process should be activated when the
contingency violates the prediction based on the face due to
human expertise in mirrored self-recognition (i.e., real-time
feedback from the self-face).
Materials and Methods
The Ethics Committee of Tohoku University School of Medicine
approved this experimental protocol.
Twenty-seven healthy right-handed male undergraduate or graduate
students (aged 19–25 years) participated, and written informed
consent was obtained from all subjects. No subject had a history of
neurological or psychiatric illness.
Virtual Mirror System
The system is schematically illustrated in Figure 1a. Subjects lay on
the bed of an MRI scanner with their heads fixed in the MRI head
coil using elastic blocks. The head component of a SENSE head spine
coil (Philips, Best, the Netherlands) was used as the open-face head
coil. A high-speed (250 fps) video camera SVS340CUCP (SVS-VISTEK,
Seefeld, Germany) viewed the subject’s face via a half-mirror attached
to the head coil. The image was projected onto a semilucent screen
behind the head coil using an Endeavor Pro4700 (Epson, Suwa, Japan)
and a DLA-HD10K LCD projector (JVC, Wayne, NJ, USA). Custom
software (Physiotech, Tokyo, Japan) allowed presentation of a
quasireal-time (<60-ms delay), delayed-real-time, or prerecorded video. The
subjects viewed the screen via a mirror.
Stimuli and Task
Figure 1b illustrates the task conditions. In all trials, a face image was
presented for 2 s and a small white circle appeared at the position of
the glabella following a 500-ms latency, where it remained for 500 ms;
this was the prompt for the mouth action. Each subject was instructed
to quickly open his mouth as soon as the prompt appeared, and to
then close it immediately. In the scanner, the subjects initially practiced
the task while viewing an example video, and then while viewing
various types of facial images that would be presented during the
experiment. After acquiring the ability to quickly and consistently
perform the task, 3 video clips were recorded: videos of the participant
engaging in regular and delayed performance (cue latency, 1 s), and
without task performance. The first 2 facial action video clips were not
used in the experiment for the subject depicted in the video, but for
other subjects as the clips for Real-time and Delayed Other conditions.
Video clips without task performance were used as the Static Self
condition in experiments for this subject, as well as for the Static Other
condition in experiments for other subjects.
Each subject participated in 3 460-s sessions, each including 10 trials
for each of the 6 conditions, with pseudo-random intertrial intervals of
1–16 s. The image was projected with minimal and 500-ms delays under
the Real-time and Delayed Self conditions, respectively. The recorded
video clips of an unfamiliar subject’s regular and delayed performances
were presented during the Real-time and Delayed Other conditions,
respectively. Videos of the subject himself and another subject not
performing the task were presented during the Static Self and Other
The latency between the onset of the action and the visual prompt
was measured for each subject using the prerecorded video used as a
stimulus. Actions could not be recorded during the fMRI measurement
because the system assigned all the computational resources to visual
presentation to minimize the feedback delay during the Real-time
fMRI Data Acquisition and Preprocessing
Forty-four transaxial gradient-echo images (echo time = 30 ms, flip
angle = 85°, slice thickness = 2.5 mm, slice gap = 0.5 mm, FOV = 192
mm, matrix = 64 × 64, voxel size = 3 × 3 × 3 mm) covering the whole
cerebrum were acquired during all sessions at a repetition time of
2.5 s, using an echo planar sequence and a Philips Achieva (3T) MR
scanner. The following preprocessing procedures were performed
using statistical parametric mapping (SPM8) software (Wellcome
Department of Imaging Neuroscience, London, UK) and MATLAB:
Adjustment of acquisition timing across slices, correction for head
motion, spatial normalization using the EPI-MNI template, and
smoothing using a Gaussian kernel with a full-width at half-maximum
size of 8 mm.
fMRI Data Analysis
Seven subjects were excluded from the analysis; one could not
discriminate self from other faces during the experiment, 4 performed the
action too quickly or too slowly compared with the video of the
respective other (more than a 100-ms difference in action onset), and 2
moved their heads >4 mm within a session. Data from the remaining
20 subjects were analyzed.
A conventional two-level approach for event-related fMRI data was
adopted using SPM8. A voxel-by-voxel multiple regression analysis was
conducted for the first-level within-subject (fixed effects) model.
Expected signal changes were modeled for the 6 conditions. A model of the
expected signal change was constructed using the hemodynamic
response function provided by SPM8. A high-pass filter with a cutoff
period of 128 s was used to eliminate the artifactual low-frequency trend.
A voxel-by-voxel statistical inference on the contrasts of the
parameter estimates was performed on the second-level between-subject
(random effects) model, using one-sample t-tests. First, the main
effects and their interactions were examined in a 2-by-2 factorial
design composed of the Contingency (Real-time, Delayed) and Face
factors (Self, Other) (Fig. 2a). The threshold for significant activation
was initially set at P < 0.001 (uncorrected); it was then corrected to
P < 0.05 for multiple comparisons using cluster size assuming the
whole brain as the search volume. A region-of-interest analysis was
then performed using the activation peaks identified in the contrast
between the main effects of Face and negative interaction; the areas
identified in the 2 contrasts overlapped considerably (see Results). The
analysis was intended for detailed functional segregation using 3
additional contrasts at a liberal threshold (P < 0.05, without correction for
To identify regions involved in the perceptual-level processes, the
main effect of each factor was tested. To identify regions responsive to
the contingency cue, the main effect of Contingency (Fig. 2b) was
tested using the contrast (Real-time Self + Real-time Other) – (Delayed
Self + Delayed Other). To identify regions responsive to the figurative
cue, the main effect of Face (Fig. 2c) was tested using the contrast
(Real-time Self + Delayed Self ) – (Real-time Other + Delayed Other).
For the latter regions, the region-of-interest analysis was performed on
the Self–Other contrast under the Static condition to examine whether
the effect of the figurative cue was dependent on facial motion.
To identify regions involved in the 2 semantic-level processes
together, the negative interaction of 2 factors (Fig. 2d,e) was tested
using the contrasts (Real-time Other + Delayed Self ) – (Real-time Self +
Delayed Other). To identify a moderate degree of negative interaction
among regions that exhibited the main effect of Face, the contrast
Real-time–Delayed under the Other condition was used as an index of
sensitivity to a contingency cue in the absence of a figurative cue in the
region-of-interest analysis. To dissociate 2 semantic-level processes, a
contrast (Real-time Other + Delayed Self ) – 2 × Real-time Self was used
as an index of violation of the predicted contingency; delayed feedback
from the self-face and real-time feedback from the other face should
violate the subject’s prediction. This contrast dissociated the
beliefvalidation process (Fig. 2e) from amodal self-representation (Fig. 2d).
Activation specific to mirrored self-recognition (i.e., Real-time Self), if
any, was explored by testing a positive interaction, namely the contrast
(Real-time Self + Delayed Other) – (Real-time Other + Delayed Self ).
Significantly higher activation in the cuneus was identified
under the Real-time condition than during the Delayed
condition (i.e., the main effect of Contingency) (Fig. 3a, Table 1). The
activation profile showed that the effect was derived from
deactivation under the Delayed condition; this was also observed
under the Static condition.
Significantly higher activation under the Self than under the
Other conditions (i.e., the main effect of Face) is shown in
Figure 3b and Table 1. Activation was observed primarily in the
right lateral cortices. A large activation cluster in the
occipitoparietal region included peaks at the occipito-temporo-parietal
junction (OTPJx), occipitoparietal junction (OPJx), intraparietal
sulcus (IPS), posterior and anterior parts of the supramarginal
gyrus (SMG), and posterior part of the superior temporal gyrus
( pSTG). Activation was also observed in the posterior part of
the right inferior temporal gyrus ( pITG) and in the left IPS. In
the right frontal cortices, a large frontal–insular cluster had
peaks at the most posterior parts of the superior, middle, and
inferior frontal gyri along the precentral sulcus ( pSFG, pMFG,
and pIFG, respectively), the triangular and orbital parts of the
inferior frontal gyrus (tIFG and oIFG, respectively), and the
Peak activation (t-value)
Coordinates x y
“Main effect of contingency” (Real-time > Delayed)
Cuneus 0 −82 22 223
“Main effect of Face” (Self > Other)
OPTJx R 32 −76 24 3169
OPJx R 22 −72 44
IPS R 28 −62 44
L −24 −62 48 882
pSMG R 44 −32 32
aSMG R 62 −18 28
pSTG R 64 −36 18
pITG R 60 −56 −10 188
pSFG R 28 −6 52 2585
pMFG R 44 4 52
pIFG R 58 10 16
tIFG R 42 38 12
oIFG R 44 32 −4
aINS R 30 24 6
mINS R 38 6 4
“Negative interaction” (Real-time Other + Delayed Self > Real-time Self + Delayed Other)
aMFG R 42 52 2 318
aIFS R 42 42 10
pIFS R 30 12 28 733
FOP R 44 10 16
Stereotactic coordinates (x, y, z) of the activation peak, cluster size (number of voxels = 2 × 2 × 2 mm3), P-value (corrected for multiple comparisons), and t-values for the 3 major contrasts (i.e., main
effects and interaction) at the peak are given. Lowercase letters for each cluster indicate activity in the same cluster.
Abbreviations for structures: a, anterior; m, middle; p, posterior; OPT, occipito–parietal–temporal; OP, occipitoparietal; Jx, junction; IPS, intraparietal sulcus; SMG, supramarginal gyrus; STG, superior temporal
gyrus; ITG, inferior temporal gyrus; SFG, superior frontal gyrus; MFG, middle frontal gyrus; IFG, inferior frontal gyrus; t, triangular part; o, orbital part; INS, insula; IFS, inferior frontal sulcus; FOP, frontal
*P < 0.001, **P < 0.05 (uncorrected).
Interaction 0.78 0.16
anterior and middle parts of the insula (aINS and mINS,
respectively). Several of these activation peaks showed a significant
main effect of Contingency or interaction when a liberal (P <
0.05, uncorrected) threshold for statistical significance was used
A significant negative interaction of Contingency and Face
was identified in 2 clusters in the right frontal region (Fig. 3c,
Table 1). One cluster included 2 peaks at the anterior parts of
the middle frontal gyrus (aMFG) and inferior frontal sulcus
(aIFS), and the other involved 2 peaks at the posterior part of
the IFS ( pIFS) and the medial surface of the frontal operculum
(FOP) facing the middle insula.
The results are summarized in Figure 4. The activation peaks
identified in the contrast for the main effect of Face (circles in
Fig. 4a) were divided into 2 groups depending on the
sensitivity to the contingency cue (vertical axis in the upper
panel of Fig. 4a). The peaks that were not sensitive to the
contingency cue (i.e., dedicated to figurative-cue processing;
Fig. 2c) were segregated into 2 groups based on their
sensitivity to the static self-face (horizontal axis in the upper panel of
Fig. 4a): aSMG, pITG, and oIFG were responsive to a static
neutral self-face (green circles), but other occipitoparietal
regions and pSFG were not (beige circles). The peaks that
were sensitive to the contingency cue (i.e., dedicated to
amodal self-representation; Fig. 2d) were all responsive to the
static neutral self-face ( pMFG, pIFG, tIFG, and aINS; light-blue
circles). As expected, these peaks were not sensitive to the
violation of a predicted contingency (vertical axis in the lower
panel of Fig. 4a).
In contrast, the peaks identified in the contrast for negative
interaction (triangles in Fig. 4a) were divided into the aIFS and
3 resting peaks depending on their sensitivity to the static
selfface and violation of a predicted contingency (horizontal and
vertical axes, respectively, in the lower panel of Fig. 4a). The
aIFS was sensitive to the static self-face but not to the violation
of a predicted contingency (light-blue triangle); therefore, it
belonged to a group that exhibited sensitivity to the
contingency cue among the Face-main-effect peaks (i.e., dedicated to
amodal self-representation; light-blue circle; Fig. 2d). In
contrast, the resting 3 peaks aMFG, pIFS, and FOP were sensitive
to the violation of a predicted contingency (i.e., dedicated to
the belief-validation process; Fig. 2e) but not to the static
neutral self-face ( purple triangle).
This fMRI study examined the neural mechanisms underlying
mirrored self-face recognition for the first time. Our data
demonstrated that multiple processes at the perceptual or
semantic level play roles in the processing of 2 perceptual cues
(i.e., contingency and figurative) and their integration. We
demonstrated that different cortical regions were sensitive to
contingency and/or figurative cues. Particularly, several right
frontal–insular regions showed activation reflecting different
types of negative interaction between the 2 cues. These
findings support the multicomponential view of the
Main Effect of Contingency
The observed significant main effect in the cuneus may reflect
the unique visuospatial experience of the mirror confrontation
induced by detecting the real-time contingency in the visual
feedback of one’s own facial action. This region likely
corresponds to V3 or V3A (Tootell et al. 1997) and has been
reported to be involved in 3D depth perception (Paradis et al.
2000) and attention to peripersonal space in front (Weiss et al.
2000; Quinlan and Culham 2007). Upon recognition of the
“mirror,” our subjects may have perceived the space
represented on the image as real and paid attention to its depth in
their peripersonal space. The activation profile in this region
showed deactivation relative to the baseline activity under the
delayed and static conditions; this may be more accurately
described as suppression of 3D depth perception or attention to
peripersonal space during the perception of a virtual image
(i.e., recorded or static video).
Main Effect of Face
Many occipital, parietal, temporal, frontal, and insular regions
were identified, particularly in the right hemisphere. These
regions were largely consistent with those previously reported
as showing self-face-specific activation (Uddin et al. 2005;
Platek et al. 2006; Sugiura et al. 2006, 2008, 2012; Kaplan et al.
2008; Oikawa et al. 2012). Our results segregated these regions
into 3 groups. A group composed of frontal activation peaks
(i.e., pMFG, pIFG, tIFG, and aINS) also exhibited sensitivity to
the contingency cue, suggesting that these regions have a role
in amodal self-representation at the semantic level. Functional
dissociation of these frontal regions from posterior regions
was previously suggested by a different pattern of intersubject
variance in self-specific activation (Sugiura et al. 2006). The
remaining regions, which were specifically sensitive to the
figurative cue, were considered to be involved in
perceptuallevel processes. These were further subdivided into regions
that were sensitive to the static neutral self-face (i.e., aSMG,
pITG, and oIFG), and those that were not (other
occipitoparietal regions and pSFG).
Although the precise processes operating in each group of
perceptual-level regions remain a matter of speculation, the
difference in the roles of the 2 groups seem to be traceable to
different stages of an infants’ acquisition of this ability in front
of a mirror. The regions that do not respond to the static
selfface (e.g., occipitoparietal regions and the pSFG) may be
relevant to experiences during the initial stages of the acquisition
of this ability. Infants show exploratory behavior (e.g., smiling,
moving, and touching a mirror) when they begin to learn
about the contingency between their own actions and visual
feedback from a mirror (Loveland 1986). This visuomotor
learning process involves the parietal and premotor regions
(Ghilardi et al. 2000; Inoue et al. 2000), which largely overlap
with regions in this group. In contrast, the group that showed
sensitivity to the static self-face (i.e., pITG, aSMG, and oIFG)
may be related to the bodily representation of the self-face that
was established after this learning process. Specifically,
regions in this group have been implicated in the sense of
action agency or body ownership (Leube et al. 2003; David
et al. 2007; Schnell et al. 2007; Ionta et al. 2011). Both groups
of regions overlap with the areas that receive vestibular input
(Smith et al. 2012; zu Eulenburg et al., 2012), which is highly
relevant to bodily representation of self-face, as well as its
The expected activation profile was observed in 4 right frontal
peaks ( pMFG, pIFG, tIFG, and aINS) that were identified in the
contrast for the main effect of Face, and in the right aIFS,
which was identified in the contrast for the negative
interaction. Several findings support the notion of the amodal
nature of the self-representation in these regions. The response
to the self-voice (Nakamura et al. 2001) or to one’s own name
being called (Kaplan et al. 2008) was identified previously in
the regions close to the right tIFG and aIFS peaks. A correlation
between social-affective feeling and activation during self-face
viewing has also been reported in a cross-trial correlation
between a feeling of embarrassment and a region close to the
aIFS and the cross-individual correlation of a public
selfconsciousness score and a region close to the aINS and pIFG
(Morita et al. 2008). In contrast, the range or domain of “self”
represented in these regions may be limited considering the
report that they do not respond to a visually presented
selfname, whereas the right tIFG does respond to the self-face
(Sugiura et al. 2008).
A response to the violation of a predicted contingency was
identified in the right aMFG, pIFS, and FOP. The lack of
response to the static neutral self-face may be due to our
expertise in viewing a static self-face. Of these regions, the pIFS and
aMFG are close to a region that has been reported to be
responsive to the violation of a learned association between a drug
and a side effect (Fletcher et al. 2001; Turner et al. 2004). The
aMFG may select processes (i.e., perceptual cues) congruent
with the validated belief; this region has been implicated
in the enhancement and suppression of task-relevant and
task-irrelevant processes, respectively (Sakai and Passingham
2003; Sugiura et al. 2007).
Neural Mechanisms for Mirrored Self-face Recognition
The groups of regions that exhibited different activation
profiles are thus indeed likely to accommodate the different
cognitive processes that have been assumed to underlie mirrored
self-recognition based on the findings of developmental
psychology and clinical observations. This finding may contribute to
the discussion regarding the multiple developmental levels of
the ability for self-recognition, particularly its behavioral index
(Anderson 1984; Brooks-Gunn and Lewis 1984). Historically,
the legitimate index has been the mark test (or rouge test), by
which animals or infants are marked on an unseen part of the
body and examined in front of a mirror to determine whether
they show a behavior directed to the mark rather than to the
mirror (Gallup 1970; Anderson 1984; Brooks-Gunn and Lewis
1984). Some animal species that previously failed to pass this
test were able to pass a modified version of the test; for
example, pigeons passed the test after training (Epstein et al.
1981). Infants typically initially pass this test in the second year
of life, but they pass tests for other self-recognition-relevant
indices earlier; they discriminate between contingent and
noncontingent images of self at <2 months of age (Reddy et al.
2007). It would be interesting to compare these different
indices with the components of mirrored self-recognition and
the related cortical regions.
It is also interesting to focus on the role of contingency
detection in the development of recognition not only of the self
but also of others in an interactive relationship. We attributed
the neural response to the figurative cue in the right pITG,
aSMG, and oIFG to the bodily representation of the self-face
acquired through the experience of contingency testing during
infancy. However, it has been reported that these regions,
particularly the right pITG and aSMG, are also responsive to the
faces of personally familiar people in a social context (Sugiura
et al. 2012). This finding may suggest that the recognition of
personally familiar people in a social context also involves the
representation of contingent relationships. Consistent with
this, it has been proposed that an innate contingency-detection
module analyzes not only the input related to one’s own body
but also the input related to communicative others: the former
carries a perfect contingency between one’s own action and
the perceived motion of the body, whereas the latter carries a
loose contingency (i.e., delay in time, different form and
strength, or different modality) between one’s own social action
and the perceived response of others (Gergely and Watson
1999). In fact, evidence of contingency detection by infants
(e.g., a negative reaction to a recorded video compared with a
real-time video) in response to both the self (Reddy et al. 2007)
and the mother (Nadel et al. 1999) appears at around 2 months
of age. This commonality between the contingency detection
mechanisms for self- and communicative-other recognition
may explain the developmental co-emergence of mirrored
selfrecognition and empathic behavior in infants (Bischof-Köhler
1988; Zahn-Waxler et al. 1992).
To fully understand the roles of the frontal regions during
mirrored self-face recognition, further conceptual elaborations
are necessary. The right lateral prefrontal cortex has been
implicated in the coordination of self-relevant information related
not only to the physical self but also to the social self (Sugiura
2013). The regions assigned to the amodal self-representation
and the belief-validation processes within the right frontal and
insular cortices exhibited mosaic-like distribution. The overlap
of these regions with those reported in previous relevant
studies was often partial or variable. This suggests that further
functional segregation may occur. For example, it is known
that multiple regions in the lateral prefrontal cortex are
dedicated to hierarchically organized cognitive control (Koechlin
and Summerfield 2007; Badre 2008).
Although the absence of a recording of subjects’ facial action
during the fMRI measurement was a methodological limitation
of this study, we believe that it eventually had no significant
effect on our results for the following reasons. Although it was
possible that subjects performed delayed facial actions during
the experiment, this did not affect the Self conditions, which
used real-time feedback under the Real-time and Delayed
conditions. However, it could have affected the Other conditions,
during which recorded videos were presented. The delay
could have reduced and increased the real-time contingency
under the Real-time Other and Delayed Other conditions,
respectively, and thus resulted in a reduction in the contrast
between the 2 conditions. To minimize this possibility, we had
each subject practice the task until he acquired the ability to
consistently perform it. Additionally, during the fMRI
measurement, we monitored the subjects’ performance under the
Realtime Self and Delayed Self conditions in which the real-time
image of the subjects’ action was available. Despite such
efforts, it was still possible that the subjects’ performance was
delayed and that the contingency cue had a reduced effect
under the Other conditions. This may have produced an
artifactual positive interaction between 2 perceptual cues; that is,
the regions that should show the main effect of the
contingency cue may have exhibited artifactual predominance in the
contingency-cue effect under the Self condition. Fortunately,
we did not detect a positive interaction and, therefore, do not
need to discuss the possibility of artifacts.
We successfully identified the neural bases of the processes
underlying mirrored-self-face recognition. Perceptual-level
processes, which were responsive to a single perceptual cue,
were localized primarily in the posterior cortices. Responses to
real-time contingency cues were identified in the cuneus,
which may reflect the unique visuospatial experience of the
mirror confrontation. Responses to the figurative cues of the
self-face, which were assumed to index the self-face
representation, were identified in the bilateral occipitoparietal and right
temporal, frontal, and insular cortices. These regions were
segregated into 2 groups depending on the presence or absence
of the response to the static self-face, and their roles could be
traced to different developmental stages of self-face
recognition. Two regions reflecting semantic-level processes were
identified: Regions for amodal self-representation, which were
sensitive to both perceptual cues and static self-face, and those
for belief-validation, which responded to violations of a
predicted contingency. These 2 region types exhibited mosaic-like
distribution over the right frontal and insular regions,
suggesting the need for further conceptual elaboration in self-relevant
processes at the meta-cognitive level. These results illustrate
the notion of multiple developmental levels of self-recognition
ability, as well as the potential role of contingency detection in
the co-emergence of mirrored self-recognition and empathic
behavior in infants.
This study was supported by MEXT KAKENHI 23119702 and
25560347. Funding to pay the Open Access publication
charges for this article was provided by MEXT KAKENHI
Conflict of Interest: None declared.
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2814 Neural Basis of Mirrored Self-face Recognition