Obstruction increases activation in the right inferior frontal gyrus

Tao Liu2, Hirofumi Saito1, *, and Misato Oi1

Department of Cognitive Informatics, Graduate School of Information Science,

Nagoya University, Furo-cho, Chikusa-ku, Nagoya 464-8601, Japan
2
Department of Psychology, Sun Yat-Sen University, No. 135 Xingang Xi Road,
Guangzhou 510275, China
*

Corresponding author.
E-mail address: saito@is.nagoya-u.ac.jp
Tel: + 81 052 789 5550; fax: + 81 052 789 5452.

The right inferior frontal gyrus (IFG) is involved in intention understanding during
interpersonal interactions. To examine how prior experience of cooperation and
competition affects ones right IFG activation in the subsequent interaction, using
near-infrared spectroscopy (NIRS) we simultaneously measured paired participants
bilateral IFG activations during a turn-taking game. Participant pairs were assigned to
either one of two roles: a Builder taking the initial move to copy a target disk-pattern
on monitor and the Partner taking the second move to aid in (cooperation) or to
obstruct (competition) the Builder. The experiment consisted of two sessions. One
participant (B-P) played as a Builder (B-) in session 1 and changed the role to the
Partner (-P) in session 2, and vice versa for the paired participant (P-B). NIRS data in
competition demonstrated that the Builder (B-) being obstructed in session 1 showed
higher right IFG activation when (s)he took a role of obstructor (-P) in session 2 (the
obstructed effect), whereas the cooperated effect was not revealed in cooperation.
These results suggest that prior experience of being obstructed may facilitate
understanding of the Builder and/or the obstructors tactical move, increasing his/her
right IFG activation when one is meant to obstruct in subsequent competitions.

Keywords: Cooperation and Competition Inferior, frontal gyrus (IFG), Near-infrared

spectroscopy (NIRS), Role-assignment, Turn-based interaction

INTRODUCTION
Interpersonal interaction is defined as an individuals simultaneous or sequential
actions that affect the immediate and future outcomes of the other individuals
involved in the situation (Johnson & Johnson, 2005). According to previous research,
studies of interpersonal interaction are classified into two types (Konvalinka &
Roepstorff, 2012; Mnoret et al., 2014): one is the concurrent interaction that involves
bodily synchrony between two people (e.g., synchronized swimming), and the other is
the turn-based interaction that involves mental synchrony between two people
executing complementary or contrary behaviors (e.g., playing chess).
However, due to technical limitations of brain-imaging such as fMRI, most social
neuroscience research on interpersonal interaction has mainly focused on a single
individuals brain response to computerized programs in cooperative or competitive
contexts (e.g., Polosan et al., 2011). If we consider interpersonal interaction is a
bidirectional process and is concerned with the question of how two minds mutually
shape each other through reciprocal interactions (Singer, 2012), we must thus treat
the interacting members as a coupled unit (Konvalinka & Roepstorff, 2012).
Therefore, growing researchers have proposed to study the brains of two interacting
people in real mutual interactions (Hasson, Ghazanfar, Galantucci, Garrod, & Keysers,
2012; Schilbach et al., 2013).
Hyperscanning is a technique that provides the possibility for simultaneous
acquisition of the cerebral data from two or more participants (Montague et al., 2002).
Several hyperscanning studies have examined the neural mechanisms of bodily
synchrony in concurrent interactions, and reveal that the right frontal cortices are
critical for predicting others action underlying concurrent cooperation (Cui, Bryant,
& Reiss, 2012; Naeem, Prasad, Watson, & Kelso, 2012). In contrast, studies
examining the neural mechanisms underlying turn-based interaction are still scarce,
because of the complex dynamics of interaction situations (Hari, Himberg,
Nummenmaa, Hmlinen, & Parkkonen, 2013; Mnoret et al., 2014). In the present
study, we simultaneously measured paired participants brain activations and
examined the neural correlates of turn-based interaction in a computerized two-person
game.
In an fMRI study, Decety, Jackson, Sommerville, Chaminade, and Meltzoff (2004)
have investigated the neural bases of turn-based cooperation and competition in single
individuals using a computerized game. During the experiment, single participants
and an experimenter were assigned to either one of two roles: a Builder taking the
initial move to copy a target disk-pattern and the Partner taking the second move to
aid in the Builders goal achievement (cooperation condition) or to obstruct it
(competition condition). Their data revealed higher activation in the right orbitofrontal
cortex during cooperation, and higher activation in the medial prefrontal cortex during
competition. Decety et al. (2004) proposed that cooperation may provide a reward or
benefit to interacting people, while competition may require more mentalizing
resources to predict an opponents actions.
3

As the participants in their study took the roles of builder and partner within an
experiment, but an experimenter took the opposite role according to the participants
role, the authors did not compare the difference between builder and partner. Recently,
using near-infrared spectroscopy (NIRS), Liu, Saito, and Oi (in press) have measured
simultaneously pairs of participants brain activations to examine the effect of a
partners behavior (cooperate and obstruct) on the paired players frontoparietal
activations during the computerized two-person game used in Decety et al. (2004).
Their NIRS data demonstrated an interaction of role (builder vs. partner) by task-type
(cooperation vs. competition) in the right inferior frontal gyrus (IFG). The builder in
the cooperation condition showed higher activation than the cooperator, but the same
builder in the competition condition showed lower activation than in the cooperation
condition. Interestingly, the activations in the obstructor-builder pairs showed positive
correlations between their bilateral IFG, but the activations in the cooperator-builder
pairs did not. These results suggest that the builders activation in the right IFG
increases by the cooperators supportive action and decreases by the obstructors
interruptive action. Also, the obstructor may actively trace the builders tactical move,
leading to deeper mental synchrony in the competition condition, while the cooperator
may passively follow the builders move, leading to shallower mental synchrony in
the cooperation condition.
Johnson (1967) has pointed out that role reversal results in more understanding of
the opponents position. That is, prior experience of competing may enable an
individual to learn the viewpoint of the opponent and facilitate the subsequent
interaction (Yaniv, 2012). This may imply that the builders brain activation is not
only affected by the partners behavior (cooperate or obstruct) as demonstrated by Liu
et al. (in press), but also by individuals prior experience. In the present study, we
examined how an individuals IFG activation changes due to his/her prior experience
of cooperation or competition in turn-based interactions.
The right IFG is associated with understanding of the others intention (Hamilton &
Grafton, 2008; Iacoboni et al., 2005) and empathic response (Baird, Scheffer, &
Wilson, 2011; Decety, 2011; Shamay-Tsoory, Aharon-Peretz, & Perry, 2009). Overall,
if an individual learns the partners view from prior experience of cooperation or
competition, (s)he may show higher right IFG activation in the subsequent interaction
than in the prior one, since the learned view facilitates understanding of the partners
tactical behavior.
We used the same computerized turn-taking game as in Liu et al. (in press) and
manipulated the factor of role (Builder and Partner). The experiment consisted of two
successive sessions, in which pairs of participants were asked to reverse their roles.
That is, participant pairs were randomly assigned to either a Builder (B-) or a Partner
(P-) in the session 1. Then the Builder (B-) in the session 1 changed the original role
into the Partner (-P) in the session 2 (termed as B-P), and the Partner (P-) in the
session 1 took the role of a Builder (-B) in the session 2 as P-B vice versa. The Partner
took alternately both roles of Cooperator (C) and Obstructor (O) within a session.
Hence, the participant B-P was further termed as B-C in the cooperation condition and
B-O in the competition condition. Similarly, the participant P-B was termed as C-B
4

and O-B in different conditions, respectively. Based on the finding that the activations
in the obstructor-builder pairs showed positive correlations between their bilateral IFG,
but the cooperator-builder pairs did not (Liu et al., in press), we hypothesized that
prior experience of being obstructed in the session 1 may result in deeper
understanding of the builders tactical move, resulting in higher activation in his/her
right IFG when one is meant to obstruct in the subsequent (second) competition than
in the first competition condition.

METHODS
Participants
Twenty (ten pairs) undergraduate students (16 males, 4 females, age: 19.2 1.2 years)
in Nagoya University participated for course credit. Participants were paired in
same-gender dyads to avoid cross-gender effects (Balliet, Li, Macfarlan, & Van Vugt.
2011). All participants were right-handed as assessed by the Edinburgh Handedness
Inventory (Oldfield, 1971), and had normal or corrected-to-normal vision. They were
informed about the purpose and safety of the experiment, and the written informed
consent was obtained prior to participation. The present study was approved by the
local ethics committee.
Materials
The present study used the same computerized two-person game as in Decety et al.
(2004). Figure 1A shows an example scene of the game. Two participants took
different roles and played the game in a turn-taking style. One was instructed to
consistently control yellow disks to copy a target pattern as a builder, while the other
participant was asked to place blue disks to aid in the builder's task (cooperation
condition) or to obstruct it (competition condition) as a partner. In addition, both
participants also played the game independently under the observation by the partner
as control condition (independent condition).
During the experiment, two participants sat side-by-side in front of a monitor
(LDT321V; Mitsubishi Elec., Japan). The game was displayed on the monitor without
sound, and the participants played the game using two keypads by their right hands
(index finger for moving left and middle finger for moving right). Distance from
participants to monitor was 120 cm.
Experimental procedures
Prior to the experiment, participant pairs received several trainings on the tasks until
they understood each type of game, i.e., cooperative, competitive, and independent
games (totally about 5 min).
The whole experiment consisted of two sessions. Specifically, the participant pairs
5

were randomly assigned to either a builder or a partner in the session 1, and were
instructed to play the game without verbal communication. And then in the session 2,
they switched their builder-partner roles. That is, the builder (B-) in the session 1
changed the role into the partner (-P) in the session 2 (the participant was termed as
B-P through the whole experiment), and the paired participant, who took the role of a
partner in the session 1 and a builder in the session 2, was termed as P-B vice versa.
Figure 1B illustrates the experimental procedure in one session. One experimental
session contained two rest periods and 12 game-sets, including four cooperative, four
competitive, and four independent game-sets (two for Builder, and two for Partner).
The two rest periods, with fixation of a plus mark on the monitor, were assigned to the
middle and the end of the 12 game-sets, respectively.
Each game consisted of eight turns, in which two participants alternately played in
turn-taking style, i.e., four turns per participant. The builder always took the initial
turn, and the partner followed. The yellow or blue disks appear automatically every 2
s on the left-top side of a game board, indicating whose turn it is to play. That is,
participants were prompted to act on the disk movement every 2 s. Totally, each game
lasted about 16 s. Participants could only control the horizontal movement of the disk
until it reaches the aimed column, where the disk will drop.
Both two experimental sessions started with a 10-s black screen. Every game-set
started with a 3-s black screen and followed by a 3-s instruction screen indicating the
game type of the following two contiguous games, and all games were preceded by a
1-s black screen. The presentation order of the game-sets in every session was pseudo
randomized. The performance of the participants and the whole experimental situation
were video-recorded by four cameras. The game program also automatically recorded
the disk-move manipulated by participants.
Apparatus
A multichannel NIRS device (FOIRE-3000/16; Shimadzu Co., Japan) was used to
study the builder-partner pairs functional neural activations by measuring the
concentration changes of oxygenated hemoglobin (Coxy-Hb), deoxygenated
hemoglobin and total hemoglobin in specific cerebral regions. Each participant had 14
optodes which were placed on the bilateral frontoparietal regions in a 2 7 lattice
pattern forming 19 channels (Liu, Saito, Oi, & Pelowski, 2012). The middle channels
in the lowest line were located at the T3 and T4 positions of the international 10-20
system. Each channel consisted of one emitter optode and one detector optode located
3 cm apart. Based on the 3-dimensional probabilistic anatomical cranio-cerebral
correlation, when NIRS probes are placed according to the 10-20 international system
for EEG, the cerebrocranial correlation is considered to have fidelity of within 10 mm,
allowing acceptable resolution at the level of the gyrus (Okamoto et al., 2004). In
addition, NIRS detects hemodynamic changes in the brain with a depth of about 3 cm.
Therefore, the NIRS device covers bilateral frontoparietal regions in close proximity
to scalp tissues. In the present study, we focused on the bilateral IFG covered by the
channels 3 and 18 as the regions of interest. The sample rate was 8.7 HZ.
6

Data analysis
Invalid NIRS data (four P-B participants) and the data obtained from the participants
who did not follow the instruction (one B-P participant) were excluded. Finally, data
obtained from 9 B-P and 6 P-B participants were remained for analysis.
We analyzed Coxy-Hb, since oxygenated hemoglobin is the most sensitive
parameter of regional cerebral blood flow and provided the robust positive correlation
with the BOLD signal (Hoshi, Kobayashi, & Tamura, 2001). The NIRS data of the
builder and the partner were analyzed independently for the left and the right IFG in
each condition. Specifically, the baseline was set 2-s before each game-set. To remove
any longitudinal signal drift, a linear baseline correction was conducted on NIRS raw
data using the mean value of Coxy-Hb during the baseline period. The raw data from
NIRS are all originally relative values and hence cannot be averaged directly across
the participants or channels. To address this issue, raw data were then converted to
z-scores by using the mean value and the standard deviation during the baseline period
(Liu, Saito, & Oi, 2012; Liu et al., 2012; Matsuda & Hiraki, 2006). To avoid the
influences from the error performance, NIRS data obtained from the games that
contained erroneous turns were excluded (less than 5%). The NIRS data without
errors were averaged across 8 turns in each game, and then were further averaged
across 8 games of each type of condition (cooperation, competition, and independent)
separately. Finally, the group-averaged activation for each condition in the bilateral
IFG was obtained, which was analyzed by ANOVA. The statistical analysis was
carried out by means of the Statistical Package for the Social Sciences (SPSS) and the
significance level was set at p < .05.

RESULTS
Behavioral data
To verify that the participants understood the game, we calculated the error rate (%) in
each type of condition as a performance index. An error was recorded if, during a
game, the Builder made one move against the target pattern, or the Partner performed
without obeying instructions, i.e., disrupted the Builder during cooperation (or helped
during competition). Statistical analysis was conducted by means of the Statistical
Package for the Social Sciences (SPSS) and the significance level was set at p < 0.05.
The error rates were less than 5% in both the session 1 (cooperation: 3.75%;
competition: 1.25%; independent: 0%) and the session 2 (cooperation: 2.50%;
competition: 1.25%; independent: 0%).
NIRS data
Independent condition
7

Before comparing the activations (i.e., Coxy-Hb) of the paired participants (builder
and partner) in the cooperation and competition conditions, we firstly examined the
activation differences between the pairs in the independent condition. The activations
of the paired participants were expected to show no differences in the independent
condition.
In the left IFG, the activations of B-P and P-B were 0.79 (SD 0.86) and 0.13 (0.23)
in the session 1, 0.22 (1.19) and -0.12 (0.59) in the session 2, respectively. In the right
IFG, the activations of the same two participants were 0.27 (0.69) and -0.30 (0.36) in
the session 1, 0.20 (0.52) and -0.17 (0.86) in the session 2, respectively. We conducted
a two-way ANOVA (Role-order (B-P vs. P-B) Session (1 vs. 2)) in the bilateral IFG,
independently. In both the left and the right IFG, the results revealed no effects of
Role-order (F(1, 13) = 1.45, ns; F(1, 13) = 2.95, ns) and Session (F(1, 13) = 4.57, p =
0.05; F(1, 13) < 1), and no interaction (F(1, 13) < 1; F(1, 13) < 1).
Cooperation and competition conditions
To examine the activation differences between the builder and the partner through the
sessions 1 and 2 under the cooperation and competition conditions, we conducted a
three-way ANOVA (Role-order (B-P vs. P-B) Game-type (cooperation vs.
competition) Session (1 vs. 2)) on Coxy-Hb in the bilateral IFG, independently.
Figure 2 shows the mean Coxy-Hb in the bilateral IFG under the cooperation and
competition conditions.
In the left IFG, there were no effects of Role-order (F(1,13) < 1), Game-type
(F(1,13) = 3.29, p = 0.93), and Session (F(1,13) < 1), and no first-order interactions of
Role-order by Game-type (F(1,13) = 3.13, p = 0.10), Role-order by Session (F(1,13)
= 4.42, p = 0.56) and Game-type by Session (F(1,13) = 1.28, ns), and no second-order
interaction of all three factors (F(1,13) < 1).
In the right IFG, the analysis reveal no effect of Role-order (F(1,13) = 3.38, p =
0.09), Game-type (F(1,13) < 1), and Session (F(1,13) < 1), and no first-order
interactions of Role-order by Game-type (F(1,13) < 1), Role-order by Session (F(1,13)
= 1.65, ns) and Game-type by Session (F(1,13) < 1), but marginal second-order
interaction of all three factors (F(1,13) = 4.29, p = 0.06).
Although not significant (p = 0.06), the right IFG tended to show different
activation patterns in the cooperation and competition conditions. Hence, we further
conducted a two-way ANOVA (Role-order (2) Session (2)) on Coxy-Hb of the right
IFG in each condition, independently. During cooperation, the right IFG showed no
effects of Role-order (F(1,13) = 4.00, p = 0.07) and Session (F(1,13) < 1), and no
interactions (F(1,13) < 1).
During competition, in contrast, there were no effects of Role-order (F(1,13) < 1)
and Session (F(1,13) < 1), but significant interactions (F(1,13) = 6.11, p < 0.05, p2 =
0.32). The post-hoc analyses (Bonferroni procedure) revealed that in the right IFG,
B-P showed significantly higher activation when (s)he played as the role of obstructor
in the session 2 than when (s)he played as the role of builder in the session 1 (F(1,8) =
5.53, p < 0.05, p2 = 0.41), but P-B did not (F(1,5) = 1.75, ns). There was no
difference between the builder and the obstructor in both the sessions 1 (F(1,13) < 1)
8

and 2 (F(1,13) = 3.50, p = 0.84).

To estimate the robustness of the results in the right IFG, we further conducted
bootstrap analyses in the cooperation and competition conditions separately using R.
Table 1 shows the bootstrap (1000 times) results of confidence interval on the
generalized Eta-Squared measure of effect size.

DISCUSSION
The present study aimed to examine whether or not an individuals prior experience of
role behavior affects his/her right IFG activation in turn-based interaction. To achieve
this goal, using NIRS hyperscanning technique we simultaneously measured pairs of
participants activations in their bilateral IFG during a computerized two-person game.
As Singer (2012) has argued that most social neuroscience studies to date have
focused on understanding the effects of socially relevant stimuli on the mind of a
single person, and an important step forward in social neuroscience is the
development of paradigms allowing for the study of such true, real-time social
interactions. However, due to the complex process of interactive contexts, little is
known about the neural relationship between two interacting persons, who take
different roles and actions to achieve a common or a diverse goal in turn-based
interactions (Hari et al., 2013; Mnoret et al., 2014). The present study thus will
contribute to this limited, but increasing literature in social neuroscience.
The error rates of paired participants (i.e., B-P and P-B groups) were less than 5%
in both the cooperation and competition conditions, indicating that the participants
comprehended the instruction and played the game effectively. On the other hand, B-P
and P-B groups showed no differences in their bilateral IFG activations under the
independent condition, which suggests the equality between B-P and P-B groups.
Importantly, after taking a role of builder who was obstructed by a partner in the
session 1, when (s)he took a role of partner (obstructor) who obstructed a builder in
the session 2, the participant (B-P) showed higher right IFG activation in the session 2
than in the session 1 (hereafter, we call the higher right IFG activation in the second
competition condition than that in first competition condition as the obstructed effect).
However, the obstructed effect was not observed when the participants took a role
of builder who was helped by a partner in the session 1, and took a role of partner who
helped a builder in the session 2. That is, we did not obtain the cooperated effect in
the cooperation condition.
The results suggest the following two things concerning what is learned through
obstruction in B-P and the implication of the right IFG activation in this study. First,
consistent with previous studies, obstruction may need closer understanding of a
series of strategic move in builder (de Vignemont & Haggard, 2008). That is, the
builder in B-O may be intrinsically required to make inferences about the obstructors
actions (Decety et al., 2004), and thus may learn the shared representations of the
dynamic sequences of tactical moves between the roles of builder and obstructor for
final goals (de Vignemont & Haggard, 2008), whereas cooperation can be done by
9

understanding of an immediate turn by builder since they possessed the same final
goal (Decety et al., 2004; Liu et al., in press). The recent experience of being
obstructed in builder may elaborate a skill of understanding the strategic turns in
builder and/or obstructor, which facilitates a dynamic integration of chain-turns
between builder and obstructor.
Second, the IFG is anatomically adjacent to the motor areas, and is a main region of
mirror neuron system, thus is closely associated with imitation, motor sequence
learning, and understanding of actions and intentions (Hamilton & Grafton, 2008;
Iacoboni & Mazziotta, 2007; Molnar-Szakacs, Iacoboni, Koski, & Mazziotta, 2005).
Furthermore, ample lesion studies and patient studies have demonstrated that
dysfunction of the IFG may lead to deficits in social behaviors and empathic ability
(Just, Cherkassky, Keller, Kana, & Minshew, 2006; Kana, Libero, Hu, Deshpande, &
Colburn, 2012; Shamay-Tsoory, Aharon-Peretz, & Perry, 2009). These findings
suggest that the bilateral IFG plays an important role in the modeling of the
behaviour of other people through a mechanism of embodied simulation, i.e., a
direct form of experiential understanding of others (Iacoboni & Dapretto, 2006).
Consistent with these previous studies, the increase of the right IFG activation in the
obstructed effect may reflect that the participant (a builder in B-O) learned concrete
knowledge (obstructive strategy) through the obstructed experience in the session 1,
and (s)he (an obstructor in B-O) applied it to obstruction of partner (builder) in the
session 2. That is, the participant may use his/her obstructed experience to direct more
appropriate actions in the subsequent competition, leading to the increase of activation
in the right IFG (Ghahremani, Monterosso, Jentsch, Bilder, & Poldrack, 2010). The
present study demonstrates that the right IFG is involved in the obstructed effect.
Behrens and colleagues (2008, 2009) have proposed two brain networks that may
underlie social behavior. One is reward-related brain areas, while the other involves
cognitive processing to make estimates of another persons intentions, including the
cingulated cortex gyrus, the dorsomedial prefrontal cortex, the temporoparietal
junction, and the superior temporal cortex. Similarly, Sanfey (2007) has also
suggested that the Theory-of-Mind network, primarily the medial prefrontal cortex
and the anterior paracingulate cortex, is involved in intention detection, i.e.,
assessing the meaning of behavior from others for interpersonal interactions. The
present NIRS data obtained from real dyadic interactions suggests that the right IFG
may be involved in the cognitive network underlying our flexible social behavior.
The remaining question is whether the obstructed effect in the present turn-taking
game genuinely represents distinctive aspects between obstruction (competitive
behavior) and cooperation. That is, why did we obtain an obstructed effect, but not a
cooperated effect? In other words, why did we obtain an asymmetric effect? The
plausible explanation is that a cooperator may achieve their goal by only helping a
builder at each turn, even (s)he does not read a series of tactics of builder from a
birds-eye view. Accordingly, a builder learns less from the cooperated experience
than from the obstructed experience in the present turn-taking game.
It is noteworthy that, however, the player B-C consistently showed higher right IFG
activation than the player C-B in both the sessions 1 and 2 in the cooperation
10

condition, although it is not significant (p = 0.07) due to a lack of statistical power

with small sample sizes. One possible interpretation is that the initial role-assignment
of builder or cooperator in the session 1 may characterize a participant to an initiative
actor (B-) and a secondary actor (C-), and the initial role setting may be sustained
through the cooperation condition. This characterization of cooperator as a secondary
actor (player) may also bedim the cooperated effect (Liu et al., in press).
Taken together, the cooperative role of a partner seems to be passive and secondary
character due to auxiliary role of cooperation in the present turn-taking game, while
the obstructive role of a partner is active and primary character as well as a builder.
The initiative for partnership in turn-based interpersonal relation is an indispensable
factor to be examined in future studies as well as the first and second moves of two
players (Deci, Koestner, & Ryan, 1999; Van Lange, 1999).
CONCLUSION
The present study has demonstrated that in the competition condition the Builder (B-)
being obstructed in the session 1 showed a significantly higher right IFG activation
when (s)he took a role of obstructor (-O) in the session 2 (the obstructed effect). In
contrast, the NIRS data of cooperation condition did not reveal the cooperated
effect. These results imply that the builder in B-O may learn to acquire the shared
representations of actions between the builder and the obstructor through the prior
experience of being obstructed (i.e., the wide scope of achieving a goal), which
facilitates understanding of the opponent, when (s)he obstructs the goal achievement
of the opponent in the subsequent competition.

References
Baird, A. D., Scheffer, I. E., & Wilson, S. J. (2011). Mirror neuron system
involvement in empathy: A critical look at the evidence. Social Neuroscience, 6(4),
327-335. doi:10.1080/17470919.2010.547085
Balliet, D., Li, N. P., Macfarlan, S. J., & Van Vugt, M. (2011). Sex differences in
cooperation: A meta-analytic review of social dilemmas. Psychological Bulletin,
137(6), 881-909. doi:10.1037/a0025354
Behrens, T. E., Hunt, L. T., & Rushworth, M. F. (2009). The computation of social
behavior. Science, 324, 1160-1164. doi:10.1126/science.1169694
Behrens, T. E., Hunt, L. T., Woolrich, M. W., & Rushworth, M. F. (2008). Associative
learning of social value. Nature, 456, 245-249. doi:10.1038/nature07538
Cui, X., Bryant, D. M., & Reiss, A. L. (2012). NIRS-based hyperscanning reveals
increased interpersonal coherence in superior frontal cortex during cooperation.
NeuroImage, 59(3), 2430-2437. doi:10.1016/j.neuroimage.2011.09.003
Decety, J. (2011). Dissecting the neural mechanisms mediating empathy. Emotion
Review, 3(1), 92-108. doi:10.1177/1754073910374662
Decety, J., Jackson, P. L., Sommerville, J. A., Chaminade, T., & Meltzoff, A. N.
11

(2004). The neural bases of cooperation and competition: An fMRI investigation.

NeuroImage, 23(2), 744-751. doi:10.1016/j.neuroimage.2004.05.025
Deci, E. L., Koestner, R., & Ryan, R. M. (1999). A meta-analytic review of
experiments examining the effects of extrinsic rewards on intrinsic motivation.
Psychological Bulletin, 125(6), 627-668. doi:10.1037/0033-2909.125.6.627
Deutsch, M. (1949). A theory of cooperation and competition. Human Relations, 2,
129-152. doi:10.1177/001872674900200204
de Vignemont, F., & Haggard, P. (2008). Action observation and execution: What is
shared?. Social Neuroscience, 3(3), 421-433. doi:10.1080/17470910802045109
Ghahremani, D. G., Monterosso, J., Jentsch, J. D., Bilder, R. M., & Poldrack, R. A.
(2010). Neural components underlying behavioral flexibility in human reversal
learning. Cerebral Cortex, 20(8), 1843-1852. doi:10.1093/cercor/bhp247
Hamilton, A. F., & Grafton, S. T. (2008). Action outcomes are represented in human
inferior frontoparietal cortex. Cerebral Cortex, 18(5), 1160-1168.
doi:10.1093/cercor/bhm150
Hari, R., Himberg, T., Nummenmaa, L., Hmlinen, M., & Parkkonen, L. (2013).
Synchrony of brains and bodies during implicit interpersonal interaction. Trends in
Cognitive Sciences, 17(3), 105-106. doi:10.1016/j.tics.2013.01.003
Hasson, U., Ghazanfar, A. A., Galantucci, B., Garrod, S., & Keysers, C. (2012).
Brain-to-brain coupling: A mechanism for creating and sharing a social world.
Trends in Cognitive Sciences, 16(2), 114-121. doi:10.1016/j.tics.2011.12.007
Hoshi, Y., Kobayashi, N., & Tamura, M. (2001). Interpretation of near-infrared
spectroscopy signals: A study with a newly developed perfused rat brain model.
Journal of Applied Physiology, 90(5), 1657-1662.
Iacoboni, M., & Dapretto, M. (2006). The mirror neuron system and the consequences
of its dysfunction. Nature Reviews Neuroscience, 7(12), 942-951.
doi:10.1038/nrn2024
Iacoboni, M., & Mazziotta, J. C. (2007). Mirror neuron system: Basic findings and
clinical applications. Annals of Neurology, 62(3), 213-218. doi:10.1002/ana.21198
Iacoboni, M., Molnar-Szakacs, I., Gallese, V., Buccino, G., Mazziotta, J. C., Rizzolatti,
G. (2005). Grasping the intentions of others with ones own mirror neuron system.
PLoS Biology, 3(3):e79. doi:10.1371/journal.pbio.0030079
Johnson, D. W. (1967). Use of role reversal in intergroup competition. Journal of
Personality and Social Psychology, 7(2), 135-141. doi:10.1037/h0025001
Johnson, D. W., & Johnson, R. T. (2005). New developments in social
interdependence theory. Genetic, Social and General Psychology Monographs,
131(4), 285-358. doi:10.3200/MONO.131.4.285-358
Just, M. A., Cherkassky, V. L., Keller, T. A., Kana, R. K. & Minshew, N. J. (2007),
Functional and anatomical cortical underconnectivity in autism: Evidence from an
FMRI study of an executive function task and corpus callosum morphometry.
Cerebral Cortex, 17(4), 951-961. doi:10.1093/cercor/bh1006
Kana, R. K., Libero, L. E., Hu, C. P., Deshpande, H. D., & Colburn, J. S. (2012).
Functional brain networks and white matter underlying theory-of-mind in autism.
Social
Cognitive
and
Affective
Neuroscience,
9(1),
98-105.
12

doi:10.1093/scan/nss106
Konvalinka, I., & Roepstorff, A. (2012). The two-brain approach: How can mutually
interacting brains teach us something about social interaction?. Frontiers in Human
Neuroscience, 6:215. doi:10.3389/fnhum.2012.00215
Liu, T., Saito, H., & Oi, M. (2012). Distinctive activation patterns under intrinsically
versus extrinsically driven cognitive loads in prefrontal cortex: A near-infrared
spectroscopy study using a driving video game. Neuroscience Letters, 506(2),
220-224. doi:10.1016/j.neulet.2011.11.009
Liu, T., Saito, H., & Oi, M. (in press). Role of the right inferior frontal gyrus in
turn-based cooperation and competition: A near-infrared spectroscopy study. Brain
and Cognition. doi:10.1016/j.bandc.2015.07.001
Liu, T., Saito, H., Oi, M., & Pelowski, M. (2012). Appraisal of a copresent observer as
supportive activates the left inferior parietal lobule: A near-infrared spectroscopy
study using a driving video game. NeuroReport, 23(14), 835-839.
doi:10.1097/WNR.0b013e328357bb3b
Matsuda, G., & Hiraki, K. (2006). Sustained decrease in oxygenated hemoglobin
during video games in the dorsal prefrontal cortex: A NIRS study of children.
NeuroImage, 29(3), 706-711. doi:10.1016/j.neuroimage.2005.08.019
Mnoret, M., Varnet, L., Fargier, R., Cheylus, A., Curie, A., des Portes, V., &
Paulignan, Y. (2014). Neural correlates of non-verbal social interactions: A
dual-EEG
study.
Neuropsychologia,
55,
85-97.
doi:10.1016/j.neuropsychologia.2013.10.001
Molnar-Szakacs, I., Iacoboni, M., Koski, L., & Mazziotta, J. C. (2005). Functional
segregation within pars opercularis of the inferior frontal gyrus: Evidence from
fMRI studies of imitation and action observation. Cerebral Cortex, 15(7), 986-994.
doi:10.1093/cercor/bhh199
Montague, P. R., Berns, G. S., Cohen, J. D., McClure, S. M., Pagnoni, G., Dhamala,
M., & Fisher, R. E. (2002). Hyperscanning: Simultaneous fMRI during linked
social interactions. NeuroImage, 16(4), 1159-1164. doi:10.1006/nimg.2002.1150
Naeem, M., Prasad, G., Watson, D. R., & Kelso, J. A. (2012). Electrophysiological
signatures of intentional social coordination in the 10-12 Hz range. NeuroImage,
59(2), 1795-1803. doi:10.1016/j.neuroimage.2011.08.010
Okamoto, M., Dan, H., Sakamoto, K., Takeo, K., Shimizu, K., Kohno, S., ... & Dan, I.
(2004). Three-dimensional probabilistic anatomical cranio-cerebral correlation via
the international 1020 system oriented for transcranial functional brain mapping.
Neuroimage, 21(1), 99-111. doi:10.1016/j.neuroimage.2003.08.026
Oldfield, R. C. (1971). The assessment and analysis of handedness: The Edinburgh
inventory. Neuropsychologia, 9(1), 97-113. doi:10.1016/0028-3932(71)90067-4
Polosan, M., Baciu, M., Cousin, E., Perrone, M., Pichat, C., & Bougerol, T. (2011).
An fMRI study of the social competition in healthy subjects. Brain and Cognition,
77(3), 401-411. doi:10.1016/j.bandc.2011.08.018
Sanfey, A. G. (2007). Social decision-making: Insights from game theory and
neuroscience. Science, 318, 598-602. doi:10.1126/science.1142996
Schilbach, L., Timmermans, B., Reddy, V., Costall, A., Bente, G., Schlicht, T., &
13

Vogeley, K. (2013). Toward a second-person neuroscience. Behavioral and Brain

Sciences, 36(4), 393-414. doi:10.1017/S0140525X12000660
Shamay-Tsoory, S. G., Aharon-Peretz, J., & Perry, D. (2009). Two systems for
empathy: A double dissociation between emotional and cognitive empathy in
inferior frontal gyrus versus ventromedial prefrontal lesions. Brain, 132, 617-627.
doi:10.1093/brain/awn279
Singer, T. (2012). The past, present and future of social neuroscience: A European
perspective. NeuroImage, 61(2), 437-449. doi:10.1016/j.neuroimage.2012.01.109
Van Lange, P. A. (1999). The pursuit of joint outcomes and equality in outcomes: An
integrative model of social value orientation. Journal of Personality and Social
Psychology, 77(2), 337-349. doi:10.1037/0022-3514.77.2.337
Yaniv, D. (2012). Dynamics of creativity and empathy in role reversal: Contributions
from neuroscience. Review of General Psychology, 16(1), 70-77.
doi:10.1037/a0026580

14

Table 1. The bootstrap results of confidence interval on the generalized Eta-Squared

measure of effect size for the right IFG activations of cooperation and competition
conditions.

Source

Cooperation

Competition

Low

Up

Low

Up

role-order

0.02

0.47

0.00

0.36

session

0.00

0.12

0.00

0.18

role-order session

0.00

0.18

0.03

0.38

15

Figure 1. (A) A clip from a game in the cooperation condition (the indicative rows and
letters were not displayed in the experimental games). (B) Experimental procedure in
one session (revised from Decety et al., 2004).

16

Figure 2. Average concentration changes in oxygenated hemoglobin (Coxy-Hb) in the

bilateral IFG under the cooperation and competition conditions. IFG: inferior frontal
gyrus. B-P indicates the participant who played the game as a Builder in the session 1
and as a Partner in the session 2, and vice versa for P-B. Error bars represent standard
deviation. The asterisk (*) indicates p < 0.05.

17