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Research
Cite this article: Ebstein RP, Monakhov MV,
Lu Y, Jiang Y, Lai PS, Chew SH. 2015
Association between the dopamine D4 receptor
gene exon III variable number of tandem
repeats and political attitudes in female Han
Chinese. Proc. R. Soc. B 282: 20151360.
http://dx.doi.org/10.1098/rspb.2015.1360
Received: 9 June 2015
Accepted: 3 July 2015
Subject Areas:
behaviour, genetics
Keywords:
DRD4, dopamine, candidate gene,
social attitudes, political ideology
Author for correspondence:
Richard P. Ebstein
e-mail: rpebstein@gmail.com
Electronic supplementary material is available
at http://dx.doi.org/10.1098/rspb.2015.1360 or
via http://rspb.royalsocietypublishing.org.
Association between the dopamine D4
receptor gene exon III variable number of
tandem repeats and political attitudes in
female Han Chinese
Richard P. Ebstein1, Mikhail V. Monakhov1, Yunfeng Lu2, Yushi Jiang2,
Poh San Lai3 and Soo Hong Chew2
1
Department of Psychology, National University of Singapore, AS4, 1 Arts Link, 117570, Singapore
2
Department of Economics, National University of Singapore, AS2, 1 Arts Link, 117570, Singapore
3
Department of Paediatrics, National University of Singapore, 1E Kent Ridge Road, 119228, Singapore
Twin and family studies suggest that political attitudes are partially deter-
mined by an individual’s genotype. The dopamine D4 receptor gene
(DRD4) exon III repeat region that has been extensively studied in connection
with human behaviour, is a plausible candidate to contribute to individual
differences in political attitudes. A first United States study provisionally
identified this gene with political attitude along a liberal–conservative axis
albeit contingent upon number of friends. In a large sample of 1771 Han Chi-
nese university students in Singapore, we observed a significant main effect of
association between the DRD4 exon III variable number of tandem repeats and
political attitude. Subjects with two copies of the 4-repeat allele (4R/4R) were
significantly more conservative. Our results provided evidence for a role of the
DRD4 gene variants in contributing to individual differences in political atti-
tude particularly in females and more generally suggested that associations
between individual genes, and neurochemical pathways, contributing to
traits relevant to the social sciences can be provisionally identified.
‘There’s not a liberal America and a conservative America—there’s the United
States of America.’ —Barack Obama
1. Introduction
The liberal–conservative ideological divide has become pervasive in American
politics viz., red and blue states, and understanding the determinants of this
enduring gulf is of keen interest not only to political scientists and psychologists
but also to the media and the public at large. Nor is the American liberal–conser-
vative divide peculiar to the United States but characterizes political attitudes
across the globe. Although the liberal–conservative distinction is by no means
watertight, it remains one of the best ways to grade political attitudes since the
French Revolution [1]. Two core dimensions have been suggested to underlie
the liberal–conservative continuum: (i) attitudes towards inequality and (ii) atti-
tudes towards social change versus tradition [2,3]. Indeed, it appears that
ideology is doggedly salient in the life of nations and can be defined as an ‘inter-
related set of moral and political attitudes that possesses cognitive, affective, and
motivational components’ [1, p. 653].
Individual differences in political attitude have been attributed to both
situational and dispositional factors [4]. Situational factors such as 9/11 pro-
duce a collective shift in political views [5] and are clearly relevant for
understanding differences in attitudes. However, biology cannot be ignored.
Starting with a study of Madsen [6] showing a role of serotonin in dominance
behaviour, biology has increasingly become a focus of explanation in under-
standing ideas related to political and other social attitudes. Among
& 2015 The Author(s) Published by the Royal Society. All rights reserved.
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dispositional factors related to broad biological processes
that have captured recent attention are neural structures
[7– 10] and cognitive processes [11] including physiological
response to threats [12], facial expression processing [13]
and sensitivity to disgust.
Accumulating evidence shows that social attitudes towards
many contemporary issues are moderately heritable [14–17].
The total contribution of genetic factors to social attitudes
such as ‘conservatism’ is significant. In the large twin study
of Eaves et al. [15], ‘conservatism’ shows a heritability of
64.5% for men and 44.7% for women. Indeed, heritability
of social attitudes is comparable to heritability of personality
traits. A few studies have also reported association between
specific genes and political attitudes and behaviour [18–23].
Dopaminergic neurons date back at least 600 million
years to the stem metazoan, an archaic crawling annelid
[24]. This ancient neurotransmitter has acquired over the
course of evolution an important role in brain function
including risk-based decision-making, reward prediction
and punishment learning [25–27], as well as risky behaviours
[28,29]. Animal models of the role of dopamine in cognition,
learning and memory, have provided invaluable clues as
to the role of dopamine in these same human behaviours
[30 –37].
A particular gene, encoding the dopamine D4 receptor
(DRD4), that we first provisionally identified with the per-
sonality trait of Novelty Seeking [38– 40], (but see [41 –43]),
was recently identified by Fowler and his group [18] as con-
tributing to individual differences in political attitude but
only contingent on number of friends. The DRD4 gene con-
tains a variable number of tandem repeats (VNTR) locus in
exon III, where a 48 bp sequence may be repeated two to
11 times [44]. Settle et al. [18] tested association between
DRD4 exon III VNTR and political attitude using data from
the ‘Add Health’ survey study [45]. Self-identified political
attitude was ascertained from responses to a widely used
survey questionnaire [46]. Settle et al. [18] used a family-
based design to minimize population stratification in a
group of young adults and found a significant interaction
between self-reported number of friends and the DRD4 7
repeat (7R) allele. In short, the number of friendships moder-
ates an association between 7R and political attitude, and
having more 7R alleles and more friends is associated with
being more liberal.
By leveraging our ongoing investigation of decision-
making [47– 53], we were positioned to undertake an inde-
pendent study of association between DRD4 and political
attitude in 1771 Han Chinese university students in Singa-
pore. We employed not only a similar self-report question
as did Settle et al. [18] to measure political attitude, but an
additional more detailed questionnaire [54] to further vali-
date our first findings. Our aim was to test association
between DRD4 exon III polymorphism and political attitude
in a non-Caucasian population living in a different cultural
and political milieu, characterized by distinct DRD4 allele fre-
quencies [55] and using in addition a more detailed measure
of political attitudes [54].
We further hypothesized that there would probably be a
gender difference in political attitudes. Notably, in an East
Asian study, a greater number of young men have outright
support for democracy compared with young women. This
corresponds to the view that more young women in East
Asia support authoritarian rule and are more cynical towards
whether the type of political system matters [56]. (See also 2
[57] who showed that Japanese females were more conserva-
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tive than Japanese males.) In another study, which is
consistent with many others, traditional morality and atti-
tudes prevail especially among women in China compared
with the UK [58].
Finally, a replication in Singaporean Han Chinese would
considerably strengthen the provisional role of the DRD4 in
partially explaining individual differences in political attitude
along the liberalism –conservatism continuum.
Proc. R. Soc. B 282: 20151360
2. Material and methods
(a) Participants and questionnaires
Study subjects were recruited via email advertisement among
students of the National University of Singapore (NUS). They
participated in an incentivized laboratory economic experiment
session, donated samples of blood or saliva for DNA extraction
and then were invited to complete online surveys with psycho-
logical questionnaires. To minimize population stratification,
we recruited only students with Han Chinese ancestry.
NEO-PI-3 [59] and TCI-R [60] questionnaires were used for
assessment of personality traits.
Political attitude was indexed by self-placement scores, as
described in [18,54] (see the electronic supplementary material,
table S1).
Number of friends was indexed by question: ‘how many
close friends/confidants do you have?’ with options being:
‘none’, ‘1 or 2 friends’, ‘3 to 5 friends’, ‘6 to 9 friends’, ‘10 or
more friends’.
(b) Genotyping
DNA was extracted either from blood samples using QIAamp
DNA Blood Midi Kit (Quiagen), or from saliva samples, collected
with Oragene DNA OG-500 tubes (DNA Genotek Inc., Ontario,
Canada). The DRD4 exon III VNTR was analysed by PCR with
0.3 units of HotStar Plus DNA polymerase (Qiagen) per reaction,
1 Q-solution, 1 CoralLoad buffer, 200 mM of each dNTP,
200 nM of each primer and 10–20 ng of genomic DNA per
reaction, in a volume of 10 ml. Primer sequences were: forward
50 -GCGACTACGTGGTCTACTCG-30 , reverse 50 -AGGACCCTCAT
GGCCTTG-30 [61]. Thermal protocol comprised activation step—
958C for 5 min; 40 cycles of 948C for 30 s, 558C for 30 s, 728C for
40 s; and final hold at 728C for 5 min.
(c) Genotype classification scheme
The 7R variant of DRD4 exon III VNTR is present in very low fre-
quencies in East Asian populations [55]. In East Asian groups,
the second most common variant is the 2R which has been
suggested to functionally ‘substitute’ for the 7R [62]. Recently,
it was suggested that any allele other than most common
( presumably ancestral) 4R probably confers decreased functiona-
lity of the DRD4 gene [63]. Molecular studies of DRD4 receptor
synthesis [64,65] strengthen this notion and show that the
folding efficiency is rate-limiting in the biogenesis of DRD4
and importantly, the receptor with only two repeats (2R) is less
upregulated compared with the receptor with four repeats
(4R). Importantly, the folding of DRD4 in the endoplasmic reticu-
lum forms the bottleneck of receptor biosynthesis. Altogether, in
East Asian groups, it therefore makes sense, to avoid losing any
genetic information, to test association by comparing 4R/4R sub-
jects against all others (mostly 2R carriers) as we do in this study.
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(d) Statistics
Statistical analysis was performed with STATA/IC 11 (StataCorp LP,
College Station, TX, USA). Deviations from Hardy–
Weinberg equilibrium were tested with command -genhw- [66].
Associations with political attitude were tested using ordered
logistic regression (OLR), associations with personality traits
were tested using linear regression. In regression models, the
DRD4 exon III VNTR 4R/4R genotype was coded as ‘1’; all
other genotypes were coded as ‘0’ (recessive genetic model). Sex
was coded as ‘1’ for males and ‘2’ for females. All regressions
used a robust variance estimator. Cliff’s deltas were calculated
using the method from [67]. Permutation analysis on the basis of
Monte Carlo simulations was conducted using command—
permute- in STATA/IC 11. The values of dependent variable
(political orientation) were permuted 1000 times and regression
tests with several genetic models (4R additive, dominant, reces-
sive, 2R additive, long alleles (more than 4R) dominant) were
performed at each permutation.
3. Results
(a) Summary statistics
In this study, 1771 (883 males) NUS undergraduates indi-
cated their political views on a 5-item Likert scale from
‘highly conservative’ to ‘highly liberal’ (higher scores corre-
spond to more liberal attitude). The political attitude
distribution is shown in the electronic supplementary
material, table S2.
The mean age of these subjects was 21 years, s.d. ¼ 1.4,
range 18 –28 (males: mean 21.7, s.d. 1.3, range 18– 28;
females: mean 20.2, s.d. 1.2, range 18 –28). OLR indicated
that the effect of age on political attitude in this student
sample was not significant (odds ratio (OR) ¼ 1.04, p ¼
0.22), but the distinction between sexes was evident, with
females being less liberal than males (OR ¼ 0.698, p ¼ 6.3 
1025) (see the electronic supplementary material, table S3).
(b) Association between personality traits and political
attitude
Subjects were also inventoried with the NEO-PI-3 (Big Five
model) and TCI-R (Temperament and Character Inventory).
NEO-PI-3 scores were available for 1257 subjects (71% of the
sample), and TCI-R scores—for 959 subjects (54%). Using OLR,
we found that liberal political attitude was positively correlated
with Openness (coeff. ¼ 0.036, p ¼ 3.5  10210), inversely with
Agreeableness (coeff. ¼ 20.33, p ¼ 9  10210) and to a lesser
degree with Neuroticism (coeff. ¼ 20.018, p ¼ 0.02) domains
of the NEO (electronic supplementary material, table S4).
Moreover, liberalism was positively associated with TCI
novelty seeking (coeff. ¼ 0.032, p ¼ 1.7  1026), negatively
related with self-transcendence (coeff. ¼ 20.018, p ¼ 0.0035)
and to a lesser extent with reward dependence (coeff. ¼
20.013, p ¼ 0.03) (electronic supplementary material, table S5).
(c) Association between political attitude and DRD4
exon III variable number of tandem repeats
Genotype and allele frequencies of DRD4 exon III VNTR are
shown in the electronic supplementary material, table S6. As
expected, the most common allele was the 4R followed by the
2R in this Han Chinese population in which the prevalence of
the 7R allele is known to be low [55]. No deviation from
Hardy–Weinberg equilibrium was detected ( p . 0.05). Nota- 3
bly, the frequency of all alleles of more than six repeats were
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extremely rare as is typical of Chinese populations (less than
0.5%).
We initially examined the role of both DRD4 and sex for
association with political ideology. As shown in table 1
(model a), a significant association was observed between
the DRD4 exon III VNTR and political attitude (OR ¼ 0.820,
p ¼ 0.027), the 4R/4R carriers were less liberal than other
subjects. Comparison of 4R/4R with all other DRD4 geno-
types is not an uncommon practice [68–73]. Additionally,
presence of a single 7R trumps the effect of the 4R in many
Proc. R. Soc. B 282: 20151360
studies hence it is not surprising that the 4R/4R grouping
has special significance. Moreover, the 4R/4R is the ancestral
genotype [74], again suggesting its distinct status. Finally, the
4R/4R is more efficient in forming heterodimers than the 4R/
7R [64], suggesting a possible biochemical mechanism under-
lying the grouping of the 4R/4R homozygote versus all
others as we have herein done. The effect of sex on political atti-
tude remained significant in the model including DRD4 (OR ¼
0.700, p ¼ 7.7  10205) with females being less liberal. We next
included the interaction term (DRD4  sex) in the model and
observed a significant interaction (OR ¼ 0.675; p ¼ 0.03)
between DRD4 and sex (table 1, model 2); this implied that
the effect of DRD4 was modulated by sex.
To have a closer look at the interaction effect, we next
examined each sex separately. As shown in table 1, the associ-
ation between political attitude and DRD4 was highly
significant for females (OR ¼ 0.658, p ¼ 0.001), but not for
males (OR ¼ 1, p ¼ 1). Cliff’s delta (effect size measure for
ordinal data [67]) was 0.058 in the combined sample and
0.115 in the female group, indicating a small effect size is
almost always observed for complex traits from diabetes
[75] to educational achievement [76]. Among females in the
highly conservative category 37.5% of subjects carried non-
4R/4R genotypes, whereas this proportion was as high as
62.1% in the highly liberal group. We further examined the
robustness of the association results by carrying out permu-
tation analysis (1000 iterations) with several genetic models
(4R additive, 4R recessive, 4R dominant, 2R additive and
long alleles (more than 4R) dominant) towards minimizing
the problem of multiple testing [77]. In the combined
sample (males and females), the empirical p-value showed
only borderline significance ( p ¼ 0.076), whereas the associ-
ation remained highly significant in the female sample ( p ¼
0.003).
(d) Association of DRD4 with a broader range of
political attitudes
Whether people are liberal or conservative is an aggregate
measure of political attitude, and individuals may have differ-
ent views on specific economic, social and political issues.
Hence, we also used an adapted version of the questionnaire
in Benjamin et al. [54], which inventoried a broader range of
such attitudes. The original version of the questionnaire had
34 questions, which focused on subjects living in Sweden.
We modified the questionnaire and chose 22 questions,
which were more relevant to attitudes on economic, social
and political policies in Singapore.
We applied the OLR model to test the association between
subject’s attitude on each question and DRD4, with sex as a
covariate (electronic supplementary material, table S7).
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Table 1. Effect of DRD4 exon III VNTR on political attitudes. (Ordered logistic regression: odds ratios indicate odds of being in higher political attitude category 4
with one unit increase of independent variable. (a) Model without interaction term. (b) Model with DRD4  sex interaction term. DRD4 ¼ 1 if DRD4 exon III

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VNTR genotype is 4R/4R and DRD4 ¼ 0 for all other genotypes. Sex ¼ 1 for males and 2 for females. Higher values of political ideology scale correspond to
liberal attitude. Minus sign at the coefficients, and odds ratios less than 1 imply that 4R/4R carriers were less liberal. p-values in parentheses. *p , 0.10;
**p , 0.05; ***p , 0.01.)

both sexes

(a) (b) males females

coeff. odds ratio coeff. odds ratio coeff. odds ratio coeff. odds ratio

Proc. R. Soc. B 282: 20151360

DRD4 20.199** 0.820** 0.394 1.482 20.0001 1.000 20.419*** 0.658***
(2.7  1022) (0.18) (1) (1.1  1023)
sex 20.356*** 0.700*** 20.134 0.874
25
(7.7  10 ) (0.32)
DRD4  sex 20.393** 0.675**
(3.0  1022)
n 1771 1771 883 888

Among the 22 items, there were three which we judged were with DRD4 genotype ( p ¼ 0.00069). This suggests that politi-
very relevant to our Singapore student population, and that cal attitude in Singapore may be indexed more accurately by
appeared to strongly index Singapore political orientation attitude on environmental protection which has a high profile
(Question 13, ‘strengthen animal rights’; Question 15, ‘decrease focus on campus that is unusually pervasive.
carbon dioxide emissions’; Question 14, ‘invest more to pre-
vent environmental damages’; see the electronic
supplementary material, figure S1). The associations of these
(e) Interaction between DRD4 and number of friends
As Settle et al. [18] reported a significant association between
items with DRD4 polymorphism were significant ( p , 0.05;
DRD4 and political attitude only as an interaction between
electronic supplementary material, table S7). The coefficients’
gene and number of friends, we also examined the role of
signs were consistent with our main results: those subjects
friendship in this study. Our subjects were asked to report
who have genotype 4R/4R tended to have less liberal views.
their number of close friends. The distribution is shown in
To minimize issues of multiple testing, we carried out a
the electronic supplementary material, table S8.
PCA analysis on all 22 questions of the broad political
As shown in the electronic supplementary material, table
attitudes questionnaire. In model 1 of table 2, the main com-
S9, the effect of number of friends, coded either as a ordinal
ponent from PCA was correlated with DRD4 (coeff. ¼ 0.298,
or dichotomized variable, i.e. ‘number of friends’ into ‘0–5
p ¼ 0.003). When we further checked the loadings of the
friends’ and ‘more than 5’ friends categories, was not signifi-
main component, we found that Question 13, Question 14
cant (electronic supplementary material, table S9, models 1
and Question 15 had the highest loadings (0.40, 0.47 and
and 2). Nor was there any significant interaction between
0.47, respectively). These were all questions that were directly
friendship and DRD4 genotype (electronic supplementary
related with environmental problems. In Singapore, political
material, table S9, models 3 and 4). The only significant inter-
discussion and activity is distinct from Western democracies
action term, after including DRD4, sex and friendship in the
[78], but students do care about environmental issues [79].
model, was the interaction between sex and DRD4 (electronic
Broad concerns about the habitat in Singapore and beyond
supplementary material, table S9, model 5). In short, we
are as much of a social issue as an economic issue. People
found no evidence for a role of friendship in the role of
here suffer from forests burning in Indonesia fairly fre-
DRD4 in contributing to political attitudes in our Singaporean
quently, and hence the environment is naturally a hot
Han Chinese subjects.
button issue (e.g. [79]). Moreover, specifically at the NUS
campus, the environment has a very high profile. Indeed,
the NUS has an Office of Environmental Sustainabi-
lity (http://www.nus.edu.sg/oes/prog/prog.html). The 4. Discussion
campus is also distinguished by a ‘Green Carnival’ (http:// The main finding of this study was an association between
www.nus.edu.sg/oes/prog/envedu/green_carnival.html). DRD4 and political attitude along the liberal–conservative
The Cronbach’s a-value of these three questions is 0.8, which axis. Singapore is characterized by a distinct political culture
suggests that they are consistently measured. We group the compared with the United States and we further examined a
three questions together, and add up the three responses of homogeneous ethnic group of Han Chinese with distinct
the questions into an aggregate score of attitude towards DRD4 exon III VNTR allelic frequencies. Despite these notable
environmental protection. In table 2, the model 2 (environ- differences in the two samples, our results also indicated that
mental score) shows that the score is significantly associated variation in DRD4 affects variation in political attitudes.
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Table 2. Effects of DRD4 on major principal component and environmental
problem attitude score from political attitudes battery. (DRD4 ¼ 1 if DRD4
exon III VNTR genotype is 4R/4R and DRD4 ¼ 0 for all other genotypes.
Sex ¼ 1 for males and 2 for females. p-values in parentheses. *p , 0.10;
**p , 0.05; ***p , 0.01.)
(1) (2)
main component
variables of PCA environmental score
DRD4 0.298*** 0.353***
(6.4  1024) (6.9  1024)
sex 20.509*** 20.832***
(1.5  10 )27
(1.5  10212)
n 1178 1178
However, we observed a main effect of DRD4 on political atti-
tude, whereas Settle et al. [18] observed the effect of the gene
in the context of number of friends. Importantly, we further vali-
dated our findings with a more detailed political attitudes
questionnaire adapted for Singapore’s unique political climate.
In this study, we observed a highly significant correlation
between TCI novelty seeking and political attitude in the
expected direction. Individuals high on novelty seeking
tended to be more liberal politically. We however, observed
highly significant correlations between NEO agreeableness
(negative) and openness ( positive) with political liberalism.
We observed a weaker negative correlation between liberal-
ism and neuroticism. Gerber et al. [80] provides a good
discussion of personality and political attitudes. Some studies
have examined political attitudes and agreeableness [81,82]
with unclear results. Hirsh et al. [83] examined personality
and political attitudes and observed that one key subpart of
agreeableness ( politeness) positively correlates with conser-
vatism, while the other main subpart (empathy) positively
correlates with liberalness. These results suggest that if agree-
ableness is reported as an aggregate score, then the two
subscales would tend to cancel each other out perhaps
explaining in part the discrepancies reported in the literature
between this trait and political ideology.
Although we found that political attitude was strongly
correlated with TCI novelty seeking, DRD4 was solely associ-
ated with political attitude and not with personality traits
in the current sample (electronic supplementary material,
tables S10 and S11). That being said, other work from our
laboratory with this same group of subjects showed an
association between financial risk attitude modelled in
behavioural economic tasks and the DRD4 exon III VNTR
(Y. Jiang, M. Monakhov, R. P. Ebstein, S. Zhong, S. H.
Chew 2015, unpublished data) and notably, risk attitude is
an important correlate of novelty and sensation seeking.
We suggested the notion that risk attitude may not be
easily captured by pencil and paper questionnaires and that
behavioural economic tasks are perhaps a more efficient
strategy in looking for specific gene associations (see our
review [38]).
In the current report, we observed a significant sex 
DRD4 interaction on political attitude. Additionally, Singa-
porean females were also more conservative than males. It
is interesting that dopaminergic function is also enhanced
in females [84–86]. These studies suggest that females may
have higher dopaminergic activity in striatum and higher 5
extra-striatal dopamine receptor density and availability com-
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pared with males [86]. We have proposed a neurochemical
model linking low dopamine tone to risk proneness and
high tone to risk aversion [87]. The high dopaminergic tone
that characterizes females, in particular in women with the
4R/4R genotype, could perhaps predispose them to low
risk attitudes as well as political conservatism.
Several studies show that associations between DRD4 and
behaviour, including behavioural economic tasks [52], per-
sonality [88] and smoking [89] are contingent on sex.
Altogether, these considerations suggest that the current find-
Proc. R. Soc. B 282: 20151360
ings indicating a main effect of DRD4 on political attitude in
females, are consistent with a number of reports not only
showing gender-specific interactions with this gene, but
also reflecting the importance of gender on political attitudes
[90].
An important difference between Singapore and the
United States that might offer some explanation of the differ-
ence between our results and the results reported by Settle
et al. [18] regarding ‘friendship’ is that Singapore compared
with the United States is a considerably more collectivist
society. In Hofstede’s model [91], Singapore scores 20 on indi-
vidualism, whereas the United States scores 91. In societies
characterized as collectivist, connections with other people
are long-term due to social bonds that are made more concrete
by reciprocal commitments between members of the family,
tribe or religion. Indeed, such connections are so relevant
that the self is defined by them and self for members of a col-
lectivist culture is fundamentally relational [92]. These
cultural distinctions between Singapore and the United
States suggested to us that the association between DRD4
and political attitude, not surprisingly, would be character-
ized by nuanced differences. After all, gene effects on
behaviour are sometimes contingent on culture [93–95].
This study is especially important in the context of the
Risch et al. [96] criticism, that warned against ‘attempts to
rescue an unsuccessful candidate gene disease association,
no matter how strong the candidate, by invoking an inter-
action with a common environmental exposure’, as our
study showed a main effect (no need for interaction) of
association of the DRD4 exon III VNTR on political attitude.
Nevertheless, contrasting our study with Settle et al. [18]
suggested that cultural context was important. Finally, the
current findings suggested that the Settle et al. [18] study of
an interaction between the DRD4 and social context, despite
the absence of a main effect, was probably a true finding and
importantly, consistent with the current results pointing at
DRD4 as a possible correlate of political views.
To our knowledge, this is the first attempt to replicate the
Settle et al. findings. We now have two reports linking DRD4
exon III VNTR and political attitude which include a first
study of 1941 individuals [18] and the current report that
examined 1771 subjects, are among the largest single studies
of this gene and altogether give us some measure of confi-
dence that the association will continue to be successfully
extended in future studies.
Ethics. Subjects gave informed written consent prior to participating.
The Institution Review Board of the National University of Singapore
approved the study. Subjects were reimbursed for participation in the
project (S$25 per hour on average).
Data accessibility. Data can be downloaded from the Dryad Digital
Repository: http://doi.org/10.5061/dryad.c5m95.
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Authors’ contributions. R.P.E., M.M., Y.L., Y.J., P.S.L. and S.H.C. per- Neuroimaging and Behavioral Study of Human Decision Making’) 6
formed research. R.P.E., M.M., Y.L., P.S.L. and S.H.C. analysed the and National University of Singapore (‘Decision Making Under
data. R.P.E., M.M., Y.L., Y.J. and S.H.C. wrote the paper. Urbanization: A Neurobiological and Experimental Economics

Competing interests. We declare we have no competing interests.
Funding. This study was supported by grants from AXA Research
Fund (‘The Biology of Decision Making Under Risk’), John Temple-
ton Foundation (‘Genes, God and Generosity: The Yin Yang of
DNA and Culture’), Singapore Ministry of Education (‘The Genetic,
rspb.royalsocietypublishing.org
Approach’ and Start-Up grants to R.P.E. and S.H.C.).
Acknowledgements. We thank Aileen Pang Yu Wen, Lye Hui Jen, Ping
Yuan, Zhu Qingdi and Xiong Gaogao for assistance with DNA extrac-
tion and genotyping; Anne Chong, Tang Rong and Zhang Xing for
collection of samples and carrying out experimental sessions.

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