NeuroImage 213 (2020) 116681

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NeuroImage
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Genetic susceptibility to parenting style: DRD2 and COMT

influence creativity
Si Si, Yukang Su, Shun Zhang **, Jinghuan Zhang *
Department of Psychology, Shandong Normal University, Jinan, China

A R T I C L E I N F O A B S T R A C T

Keywords: The present study systematically investigated if genetic variations in the DRD2 and COMT interacted with
Creativity parenting style to predict individual differences in creativity. In a sample of young adults, we first examined the
DRD2 interaction of each individual polymorphism and parenting style on creativity. Then, we aggregated the contri-
COMT
bution of these susceptibility polymorphisms into a Cumulative Genetic Score (CGS) which combines the plas-
Cumulative genetic score (CGS)
Gene-environment interaction
ticity conferred by two or more single nucleotide polymorphisms (SNPs) of DRD2 and COMT into a single score.
Differential susceptibility model When polymorphisms were examined in isolation, the results indicated there were only two polymorphisms from
COMT (rs5993882 and rs5993883), which were found to interact with mother authoritativeness to predict
creativity. However, a CGS comprising of several potential susceptibility polymorphisms from DRD2 and COMT
genes did indeed interact with parenting style to explain a significant amount of variance in the creativity.
Furthermore, the analysis of regions of significance (Ros) showed supporting evidences for differential suscep-
tibility model, such that the individuals with high CGS were more likely to be affected by their both negative and
positive parenting style, demonstrating higher and lower creativity. These results further provide evidence for the
involvement of dopaminergic genes in the relationship between parenting style and creativity and suggest in-
dividuals with the greater susceptibility are the ones who are more susceptible to environmental influences.

1. Introduction extremely tolerate their impulses. Authoritative parents tend to behave

between the two extremes, encouraging personal independence and
Creativity has been defined as the way of thinking that leads to feelings expression but also setting reasonable limits. Many theoretical
original, yet potentially useful outcomes (Mumford, 2003). It brings perspectives have argued, creativity is fostered in environments that
humans numerous benefits, such as leading or adapting to changes, provide psychological safety and freedom (Erikson, 1963; Rank, cited in
increasing social standing, and making high quality decisions (Nijstad Harrington et al., 1987; Rogers, 1970). Therefore, parental authorita-
et al., 2010; Runco, 2004; Sternberg, 1999). Therefore, nurturing the tiveness and permissiveness which are characterized as high level of
creativity is essential in preparing individuals for a better life and has autonomy are inclined to lead to higher creativity; parental authoritari-
been the long-standing goal of the whole society. Given its importance, a anism, by contrast, which is characterized as high level of control has
number of studies have been conducted to identify the underlying been demonstrated to result in lower creativity (Mozafari, 2014; Meh-
mechanism that contributes to creative thinking and extensive research rinejad et al., 2015; Nichols, 1964; Miller et al., 2012). However, the
has found that family, especially the parenting style, is one well studied relationships between parenting styles and creativity are still inconclu-
source of influence of creativity (Kemple and Nissenberg, 2000; Sen and sive. For example, there was evidence showing that not all individuals
Sharma, 2013). exposed to authoritative and permissive parenting would be more crea-
Parenting style refers to the emotional and behavioral interaction tive (Fearon et al., 2013; Mehrinejad et al., 2015). Some individuals with
made by parents with their children. According to Baumrind (1966), permissive parenting were even found to be less creative than their
there were three main types of parenting, namely authoritarian, counterparts (Mozafari, 2014).
permissive and authoritative. The authoritarian parents over-control One of the possible reasons for these inconsistent results is that in-
their child’s behavior and attitudes while permissive parents often dividuals are differently affected by their rearing experience. Different

* Corresponding author. Department of Psychology, Shandong Normal University, No. 88 East Wenhua Road, Jinan, 250014, China.
** Corresponding author. Department of Psychology, Shandong Normal University, No. 88 East Wenhua Road, Jinan, 250014, China.
E-mail addresses: sisi52144@163.com (S. Si), yukangsu@126.com (Y. Su), yinxingren1986@hotmail.com (S. Zhang), zhangjinghuan@sdnu.edu.cn (J. Zhang).

https://doi.org/10.1016/j.neuroimage.2020.116681
Received 31 July 2019; Received in revised form 21 February 2020; Accepted 24 February 2020
Available online 28 February 2020
1053-8119/© 2020 The Authors. Published by Elsevier Inc. This is an open access article under the CC BY-NC-ND license (http://creativecommons.org/licenses/by-
nc-nd/4.0/).
S. Si et al. NeuroImage 213 (2020) 116681

genotypes might moderate how individuals experience environment polymorphism
parenting style interactions to be investigated are in-
instead of having a direct effect on creativity. For example, potential role clined to have small effects (Kendler, 2005; Duncan and Keller, 2011),
in creativity is suggested for the serotoninergic genes (Bachner-Melman such that roles of many functional variants with minor effects cannot be
et al., 2005; Volf et al., 2009). Among them, the tryptophan hydroxylase detected.
1 gene (TPH1) is the most extensively studied. However, previous studies Fortunately, recent researchers have found that investigating the role
examining the association of TPH1 and creativity yielded incompatible of multiple genetic polymorphisms using a cumulative genetic score
results (Reuter et al., 2006; Runco et al., 2011; Zhang and Zhang, 2017). (CGS) approach may be a way to address these concerns (e.g., Dobewall
Researchers argued that the effect of genetics may only become apparent et al., 2018; Mehta et al., 2016; Mitchell et al., 2011). A CGS models
if certain environmental factors are present (Rutter, 2006). Recent cumulative genetic influences and combines the susceptibility conferred
research on gene
environment interaction (G
E) highlights that the by two or more polymorphisms into a single score. Use of a CGS provides
negative effect of parental authoritarianism on creativity is only promi- several notable advantages, including increased power, as compared
nent in individuals with TPH1 rs1799913 CC genotype (Zhang et al., with the small effect sizes of individual polymorphisms and the ability to
2018). This suggests that some individuals are more susceptible to model more variability in the dopamine system. For the present study, we
environment than others because of different genetic background. Thus are using a CGS approach to determine whether the CGS of COMT and
understanding the genetic factors that influence individual sensitivity to DRD2 influences the relationship between parenting style and creativity.
environment is critical for a better understanding the effect of parenting These two genes, and the dopaminergic functions that they influence,
style on creativity. appear to interact in prefrontal and striatum brain regions (Zhang et al ,
Besides TPH1, two other genes (DRD2 and COMT) have also been 2014b). This finding suggests that the DRD2.
implicated in creativity. The DRD2 gene encodes a G protein-coupled And COMT may be biologically and behaviorally linked. Hence,
receptor that plays an important role in mediating synaptic DA dopaminergic genes which strongly affect frontal and striatal activities
signaling in striatum. The COMT gene encodes the key metabolic enzyme may tend to jointly influence creativity. This has been further elucidated
responsible for the degradation of catecholamines (including dopamine by a recent study which found creativity can also be predicted from in-
and norepinephrine). Extensive molecular genetic studies of creativity teractions between genetic polymorphisms related to frontal (COMT) and
have shown that variations of DRD2 and COMT as well as the interactions striatal (DAT) DA pathways (Zabelina et al., 2016).
between these variations were closely related to individual differences in To explain why the effect of rearing experience is more pronounced in
creativity (e.g., Murphy et al., 2013; Reuter et al., 2006; Runco et al., some individuals with particular genetic make-up, two competing
2011; Zhang et al , 2014a; Jiang et al., 2015; Zhang et al, 2014b; Han models of G
E interactions: diathesis-stress and differential suscepti-
et al., 2018), suggesting the importance of dopamine and bility have been formulated. The traditional diathesis-stress model was
norepinephrine-related genes (as well as the potential interactions be- first put forward which suggest that individuals who are “vulnerable”
tween dopamine and norepinephrine-related genes) in creativity genetically are worse off in unfavorable environments (Monroe and Si-
(Beversdorf, 2018). mons, 1991). As the dysfunctional “vulnerable alleles” were found to
Single variations in dopaminergic genes do not work independently, provide advantages over other genotypes, at least in certain contexts,
but might interact with environment to influence creativity. For example, researchers recently proposed the differential susceptibility model, sug-
in previous work, a study has shown a gene by environment interaction gesting individuals with certain alleles may be not only adversely
between family stress and a variation in the DRD2 on cognitive functions affected by negative environments, but also benefit the most from posi-
(Berman and Noble, 1997). It was found that the family stress score tive environmental factors (Belsky and Pluess, 2009). This model is
negatively associated with cognitive abilities only in individuals with considered to be much more consistent with recent G
E data results.
A1þ allele (rs1800497). Besides, there was also a study demonstrating a Therefore, it would seem more appropriate to speak of “plasticity genes”
significant DRD2 (rs1800497)
strict maternal disciplinary style inter- rather than “vulnerability genes” (Belsky et al., 2009). More recently, a
action in predicting novelty seeking (Keltikangas-J€arvinen et al., 2009). cumulative genetic plasticity model has been proposed, which predicts
With regard to COMT polymorphisms, G
E interaction effects for many that more “plasticity” alleles individuals carry, the more susceptible they
cognitive abilities have been reported in previous studies. For example, will be to environmental influences (Belsky et al., 2007; Belsky et al.,
many researchers (e.g., Enoch et al., 2009; Goldberg et al., 2013; 2009; Belsky and Pluess, 2009). Several studies have provided support
O’Donnell et al., 2017) have investigated whether COMT variant rs4680 for this model (LeClair et al , 2014; Disner et al., 2014).
moderates the influence of adverse environments on creativity-related Considering that the relationship between parenting style and crea-
cognitive functions (e.g., working memory, decision making, atten- tivity has been closely associated with dopaminergic functioning, the
tion). Furthermore, a few studies did find rs4680 would interact with present study sets out to systematically identify how variants in DRD2
positive versus more adverse parenting on decision making and attention and COMT moderate this critical association. We first examined whether
(Voelker et al., 2009; He et al., 2012). Therefore, individuals might be each individual polymorphism interacts with parenting style to predict
differentially susceptible to parenting style due to their DRD2 and COMT individual differences in creativity. Then, we aggregated the contribution
genotype, potentially accounting for individual differences in creativity. of these polymorphisms in DRD2 and COMT into a CGS which combines
To further identify the genetic factors that moderate context sensi- the susceptibility conferred by two or more single nucleotide poly-
tivity which are likely to influence the relationship between parenting morphism (SNPs) into a single score. Finally, to further clarify the G
E
style and creativity, it would be important to systematically investigate if interaction pattern for creativity as well as to better explain the potential
variations in DRD2 and COMT genes might interact with parenting style interaction of genetic profile (CGS) and parenting styles, the present
to predict creativity. Despite no study available to illustrate COMT study also examined whether the potential gene
parenting interaction
polymorphisms moderating roles in the relationship between parenting would coincide with the diathesis-stress model or the differential sus-
style and creativity, a recent study showed gene by environment (G
E) ceptibility model.
interaction between parenting style and a variation in DRD2 (Si et al.,
2018). As a preliminary result, this study does provide important insight 2. Materials and methods
into the role of dopamine-related single genes that influence individual
sensitivity to parenting style in a better understanding of individual 2.1. Participants and procedures
difference in creativity; however, it should also be noted that, one
polymorphism does not sufficiently explain the variation in gene func- The main characteristics of participants and study procedure have
tion, possibly limiting our range of understanding of biological suscep- been described previously (Si et al., 2018). Briefly, 427 Chinese under-
tibility to environmental influence. Besides, single genetic graduate students ages 18–22 (mean age ¼ 18.92, SD ¼ 0.84,

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S. Si et al. NeuroImage 213 (2020) 116681

male/female ¼ 101/326) took part in the present study. All participants perceived parenting styles of our sample, an adapted version of PAQ was
were of Han Chinese origin. This study was approved by the institutional employed. It consists of 21 items and yields permissive, authoritarian and
review board of Shandong Normal University; and a written consent form authoritative scores for both the fathers and the mothers. Each item was
was obtained from each participant after a full explanation of the study scored on a Likert-type scale from 1 (strongly disagree) to 5 (strongly
procedure. Participants first completed psychometric tests (creativity agree). The construct validity of the adapted version of PAQ was inves-
test, parental authority questionnaire and general intelligence test) tigated by examining the confirmation of the dimensionality of the PAQ
which were administered in a randomized order to control for potential using a confirmatory factor analysis (CFA) with maximum likelihood
test order effects, then peripheral venous blood samples were collected estimation in LISREL version 8.70, for the fathers and mothers. According
for genotyping. to the fit indices, the 3-factor structure model of the adapted PAQ did
display acceptable fit to the sample of the present study (father: RMSEA
2.2. Measures ¼ 0.16, NNFI ¼ 0.76, CFI ¼ 0.78, GFI ¼ 0.58; mothers: RMSEA¼.067,
NNFI¼.94, CFI¼.95, GFI¼.89). All subscales of the adapted PAQ
2.2.1. Creativity demonstrate good reliability: mother authoritative parenting α ¼ 0.81,
Creativity levels were measured with three figural divergent thinking father authoritative parenting α ¼ 0.86, mother authoritarian parenting
(DT) tasks which were selected from the Runco Creativity Assessment α ¼ 0.83, father authoritarian parenting α ¼ 0.86, mother permissive
Battery (rCAB; Creativity Testing Service, creativitytestingservices.com). parenting α ¼ 0.70, father permissive parenting α ¼ 0.72.
They were widely used in previous creativity research (Murphy et al.,
2013; Zhang et al., 2014a) and were commonly identified as culture-free 2.2.3. Intelligence
tasks. DT tasks provide a measure of (mainly) capacity for idea genera- General fluid intelligence was measured by Raven’s Advanced Pro-
tion. Since idea generation is critical to creativity across domains, DT task gressive Matrices (APM; Raven et al., 1998). This test is a
scores can be understood as indicators of creative potential, and they well-established culture-free measure of general intelligence. In this test,
have been found to be predictive of creative achievement (Guilford, participants were asked to choose the correct entry to complete the 3
3
1966; Kim, 2008; Plucker, 1999; Runco et al., 2010). Moreover, as DT matrix for each item. Participants were given 20 min to complete the 36
tasks suffer from a poor alternate-form reliability (Barbot et al., 2016; items, and the score was calculated as the total number of items correctly
Barbot, 2019), the importance of relying on a range of DT tasks (rather answered. It has been demonstrated that the 20 min-timed score was a
than a single one) has been highlighted. Accordingly, we used three tasks reasonable and adequate predictor of the untimed APM score (Hamel and
simultaneously. Schmittmann, 2006). The IQ scores range 9–31 (M ¼ 20.56,SD ¼ 3.968)
Participants were required to complete the tasks according to the and no participant’ IQ was 3 SD below or above the mean. So no
following constructions: This is a visual game. Below, you will find a line participant was excluded.
drawing. We would like you to make a list of all of the things that each
drawing could be. The more things you write down, the better! There are 2.2.4. SNP selection
no incorrect answers, no grades, no points, and spelling does not matter. The selections of DRD2 SNPs and COMT SNPs have been described in
This is a game, NOT a test. You have 12 min to complete these questions detail previously (Zhang et al., 2014a, 2014b). They were selected based
(Four min. for each task). Now look at the first figure below and write on both allele frequency and function.
down as many things as you can for what that figure might be. What does To capture most of the common polymorphisms in DRD2 and COMT,
it look like? What could it be? seven and nine tag SNPs were selected respectively from HapMap
When the tasks were finished, experienced raters who blinded to (http://hapmap.ncbi.nlm.nih.gov) genotype data for the CHB (Han
other information about the participants, received training prior to the Chinese in Beijing, China) population (Data Rel 27 Phase II þ III, Feb09,
final ratings, as well as familiarized with the creativity literature were to on NCBI B36 assembly, dbSNP b126) by applying the Tagger program as
evaluate all responses. Creativity was assessed by three dimensions of the implemented in Haploview (Version 4.2) software (Barrett et al., 2005)
task—fluency, flexibility, originality, according to the guideline of with the following criteria: pairwise tagging only, r2 > 0.80 and minor
Creativity Testing Service. Briefly, fluency scores were calculated by the allele frequency (MAF) > 5%. Specifically, for the genomic region of
number of ideas each participant gave. Irrelevant ideas were ignored and DRD2 (chr11: 112785528.112851091, based on NCBI Genome Build
excluded from scoring. Flexibility referred to the divided categories in 36.3), we selected seven tag SNPs (rs4938019, rs4245148, rs4648319,
one participant’s response. Scoring of flexibility is based on a classifi- rs4436578, rs7122246, rs1076560, rs6279), which captured 59 out of 66
cation standard formulated by raters for each task. Originality was the (89%) common alleles (MAF > 5%) with a mean maximal r2 ¼ .95. With
number of unique ideas given by each participant. Scoring of originality regard to COMT, nine tag SNPs (rs737865, rs174675, rs5993882,
is usually based on the statistical frequencies of each response in the rs5993883, rs4646312, rs6267, rs4680, rs769224, and rs174697) were
study sample (Forthmann et al., 2017; Mouchiroud and Lubart, 2001). In selected which captured 80% of common alleles (MAF > 5%) within the
the present study, unique ideas were those ideas given by less than 5% of genomic region of COMT (chr22:18309309.18336528, based on NCBI
the sample. All three indexes demonstrated reliability with internal Genome Build 36.3) with a mean maximal r2 of 0.966.
consistency αs of 0.88, 0.83 and 0.69, respectively. For each task, two Besides, for DRD2, seven putative functional SNPs (rs1799978,
trained raters (both were psychology graduate students from Shandong rs1799732, rs4648317, rs2283265, rs6277, rs6276, and rs6278) as well
Normal University) were engaged to score all the responses. The as rs1800497 were also genotyped. As for COMT, three putative func-
inter-rater reliabilities for all the three DT indexes were higher than 0.95. tional SNPs (rs6269, rs4633, and rs4818) were selected as well.

2.2.2. Perceived parenting styles 2.2.5. Genotyping

Parental Authority Questionnaire (PAQ) was developed for the pur- Methods for DNA extraction and genotyping have been described
pose of measuring Baumrind (1971) permissive, authoritarian and previously (Zhang et al., 2014a). Briefly, DNA samples were prepared
authoritative parental authority prototypes. PAQ is appropriate for both from peripheral venous blood cells by using the QIAamp DNA Mini Kit
women and men who are older adolescents or young adults (Bud, 1991) (Qiagen, Valencia, CA, USA). Genotyping were performed at the Beijing
and it is useful to measure their current perception of their parents’ Genomics Institute-Shenzhen (BGI-Shenzhen, Shenzhen, China) by using
parenting styles. The PAQ has shown good validity in Asian societies the Sequenom® MassARRAY® iPLEX system (Sequenom, San Diego, CA,
(Ang and Goh, 2006) and has been a reliable indicator of parenting styles USA) according to the manufacturer’s instructions. For quality control,
in Hong Kong adolescents and college students in previous research 5% random DNA samples were genotyped twice for each gene to calcu-
(Chan and Chan, 2009; Lai and McBride-Chang, 2001). To measure late genotyping error. The genotyping accuracy was 100%.

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S. Si et al. NeuroImage 213 (2020) 116681

2.3. Statistical analysis Table 2

Characteristics of the genotyped single-nucleotide polymorphisms (SNPs) of
Hierarchical regression models were used to examine the interactive DRD2.
effects of variations in DRD2 and COMT SNPs and parenting styles on SNPa Positionb Location Allele MAF HWE
creativity. The analysis was first conducted in targeted SNP of DRD2 and (minor/ (%) p
COMT respectively. The procedure was as follows: participants’ gender major)

and intelligence were entered into the first step. The main effects of rs1799978 112851561 50 Promoter region G/A 19.1 .275
parenting style and SNP were entered in the second step. Finally, the rs1799732 112851462 50 Promoter region Del/C 10.4 .438
rs4938019 112846601 Intron 1 C/T 39.1 .685
interaction terms for parenting style and SNP were entered in the third
rs4648317 112836742 Intron 1 T/C 40.5 .616
step. Then, similar data processing was also done in CGS analysis. All rs4245148 112825629 Intron 1 T/C 14.3 .232
analyses were conducted in IBM SPSS Statistics Software Version 22. rs4648319 112819573 Intron 1 T/C 36.5 .145
Then we performed sensitivity analyses when significant CGS x rs4436578 112811975 Intron 1 C/T 43.0 .844
parenting style effect was found. Regions of significance (RoS) tests were rs7122246 112809667 Intron 1 A/G 5.3 .326
rs2283265 112790746 Intron 5 T/G 45.1 .434
performed to probe interactions in differential susceptibility research rs1076560 112788898 Intron 6 A/C 44.5 .625
through an online application (http://www.yourpersonality.net/int rs6277 112788669 Exon 7 T/C 5.5 .366
eraction/, see Roisman et al.). To estimate the differential susceptibil- rs6276 112786607 30 UTR A/G 48.0 .561
ity model, the RoS analysis provides five indexes (Roisman et al., 2012). rs6279 112786283 30 UTR G/C 47.9 .628
rs6278 112785934 30 UTR T/G 42.6 .922
First, the lower and higher bound, where the effect of CGS on creativity is
rs1800497 112776038 30 flanking region T/C 42.6 1
significant, is recommended to be within 2 Standard Deviation (SD). (also ANKK1 Exon
Second, the crossover point, where the regression lines intersection is 8)
expected to be closed to zero. Third, the proportion of interaction (PoI),
Note:MAF ¼ minor allele frequency; HWE ¼ Hardy–Weinberg equilibrium; UTR
which is represented on the right side of the crossover point, indicating ¼ untranslated region.
how much a interaction is “for better”, should be near 0.50. Fourth, the a
SNPs are listed down the column in sequential order from the 50 end to the 30
proportion affected (PA), which represents the proportion of the popu- end of the sense strand of DRD2.
lation that is differentially affected by the moderator, should be greater b
Physical position is based on NCBI Genome Build 36.3.
than 16% and near 50%. Finally, to avoid the nonlinear diathesis-stress
phenomenon, the X2 or ZX2 term was tested, which should be not Disner et al., 2014), we employed an additive model to assign the score
significant. for each variation, that is genotypes with zero, one and two copies of the
minor allele were coded as 0, 1 and 2, respectively. However, as the
3. Results number of participants with Del/Del genotype of rs1799732 (DRD2), AA
genotype of rs7122246 (DRD2), TT genotype of rs6277 (DRD2), GG ge-
3.1. Preliminary analyses notype of rs5993882 (COMT), TT genotype of rs6267 (COMT), AA ge-
notype of rs769224 (COMT) was small, a dominant model was used to
Table 1 shows means, standard deviations and cross-correlations of code each of these corresponding genotype. That is, the genotype with at
psychometric variables for the total sample. least one minor allele was assigned 1, and 0 was assigned to those with
Table 2 and Table 3 summarize the MAFs and the results of Hardy- none.
Weinberg equilibrium tests for DRD2 and COMT SNPs. All DRD2 15
SNPs were polymorphic with MAF>5% and no deviation from Hardy-
3.2. Regression analyses
Weinberg equilibrium was observed. All COMT 12 SNPs were poly-
morphic with MAF>5%, but the frequency distribution of rs4680 and
3.2.1. Single Polymorphsim
rs174697 genotype differed slightly from Hardy-Weinberg equilibrium
To determine whether DRD2 and COMT genetic variants interacted
(p¼.048; p¼.021). Haploview was used to calculate linkage disequilib-
with parenting styles to predict creativity, we first conducted regression
rium (LD) between SNPs. The LD patterns of the genotyped SNPs for
analysis using 9 SNPs of DRD2 and 9 SNPs of COMT as the candidate
DRD2 and COMT were shown in Fig. 1 and Fig. 2 respectively.
markers, respectively. The 9 SNPs of DRD2 and 9 SNPs of COMT were
Besides, the genotype distribution of each SNP was demonstrated in
selected from our genotyped SNPs (15 SNPs of DRD2 and 12 SNPs of
supplementary materials (see Supplementary Table 1 and Table 2). Ac-
COMT) which included both the common alleles and the functional al-
cording to previous research (Masarik et al., 2014; Pearson et al., 2014;
leles. According to analysis results of linkage disequilibrium (LD), several

Table 1
Ms, SDs and cross-correlations of psychometric variables for the total sample.
1 2 3 4 5 6 7 8 9 10 11

1.Gender
2.Intelligence .00
3.F-authoritativeness .01 .02
4.F-authoritarianism .16** -.07 -.33**
5.F-permissiveness -.11* -.06 .41** -.39**
6.M-authoritativeness .03 .02 .49** -.16** .21**
7.M-authoritarianism .05 -.05 -.13** .41** -.10* -.37**
8.M-permissiveness -.08 -.08 .23** -.17** .48** .48** -.45**
9.Fluency -.24** .10* .06 -.11* .10* .08 -.05 .06
10.Originality -.17** .05 .03 -.09 .09 .07 -.03 .02 .93**
11.Flexibility -.20** .15** .06 -.11* .08 .04 -.03 .03 .81** .74**
M — 20.56 21.06 13.05 9.92 20.89 12.14 9.82 10.24 4.97 5.13
SD — 3.97 5.53 6.99 2.92 5.13 6.05 2.88 4.24 3.09 1.26

Note. *p<.05, **p<.01; male ¼ 1, female ¼ 0; F-authoritativeness ¼ father authoritativeness, F-authoritarianism ¼ father authoritarianism, F-permissiveness ¼ father
permissiveness, M-authoritativeness ¼ mother authoritativeness, M-authoritarianism ¼ mother authoritarianism, M-permissiveness ¼ mother permissiveness. M ¼
mean, SD ¼ standard deviation.

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S. Si et al. NeuroImage 213 (2020) 116681

Table 3 only one of each of them was included in single variants
parenting style
Characteristics of the genotyped single-nucleotide polymorphisms (SNPs) of analysis. rs1800497, rs6276, rs6277, rs2283265, rs1076560 and
COMT. rs4938019 were excluded. For COMT, strong LDs were observed for
SNP Position Location Allele (minor/ MAF HWE p rs6269 and rs4646312 (r2¼.92), rs4818 and rs4646312 (r2¼.92), rs4818
major) (%) and rs6269 (r2¼.92), rs4633 and rs4680 (r2¼.92). Hence, rs4646312,
rs737865 18310121 Intron 1 C/T 29.7 .907 rs4818 and rs4680 were excluded. Finally, 9 SNPs of DRD2 (rs6278,
rs174675 18314051 Intron 1 T/C 39.3 .612 rs6279, rs7122246, rs4648317, rs1799978, rs1799732, rs4245148,
rs5993882 18317533 Intron 1 G/T 12.1 1 rs4648319, rs4436578) and 9 SNPs of COMT (rs6269, rs4633, rs737865,
rs5993883 18317638 Intron 1 G/T 41.3 .690
rs174675, rs5993882, rs5993883, rs6267, rs769224, rs174697) were
rs4646312 18328337 Intron 1 C/T 37.6 .217
rs6269 18329952 Exon 3 G/A 37.9 .182 selected to conduct single variants
parenting style analysis.
rs4633 18330235 Exon 3 T/C 26.2 .104 To minimize the likelihood of a Type I error, we adjusted our alpha
rs6267 18330263 Exon 3 T/G 7.5 .721 based on the number of genetic polymorphisms we planned to examine in
rs4818 18331207 Exon 4 G/C 37.2 .254 a gene. Given that we are investigating nine polymorphisms in each gene,
rs4680 18331271 Exon 4 A/G 26.6 .048*
rs769224 18331804 Exon 5 A/G 6.9 .710
we used a conservative Bonferroni adjusted alpha of .006 (a ¼ .05/9) to
rs174697 18333832 Intron 5 A/G 32.9 .021* determine statistical significance.
The significant results for each model testing the individual variant

Note:*p<.05; MAF ¼ minor allele frequency; HWE ¼ Hardy-Weinberg equilib-
parenting style interactions for the prediction of fluency, originality and
rium; UTR ¼ untranslated region.
a
SNPs are listed down the column in sequential order from the 50 end to the 30 end
flexibility are shown in supplementary materials (Supplementary Table 3
of the sense strand of COMT. and Table 4). While neither single polymorphism nor parenting style was
b
Physical position is based on NCBI Genome Build 36.3. independently significant in the model including rs5993882 and
rs5993883, the two-way interaction term of mother authoritativeness
SNPs were excluded in single variants
parenting style analysis because and COMT genotype, controlling for participant gender and intelligence,
of their strong LDs with other SNPs. For DRD2, strong LDs were observed was significant in predicting creativity (fluency: β ¼ 0.14, p ¼ 0.002;
for rs6278 and rs1800497 (r2¼.97), rs6276 and rs6279 (r2¼.99), rs6277 flexibility: β ¼ 0.14, p ¼ 0.004). However, none of the models including
and rs7122246 (r2¼.91), rs2283265 and rs1076560 (r2¼.97), rs1076560 other polymorphisms in DRD2 and COMT passed the Bonferroni adjusted
and rs6278 (r2¼.90), rs4938019 and rs4648317 (r2¼.92). Therefore, threshold (p < 0.006) for statistical significance though there were seven

Fig. 1. Linkage disequilibrium (LD) pattern of the 15 DRD2 SNPs analyzed in this study.

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S. Si et al. NeuroImage 213 (2020) 116681

Fig. 2. | Linkage disequilibrium (LD) pattern of the 12 COMT SNPs analyzed in the present study. Numbers in squares designate the degree of LD (r2) between any two
SNPs. The color from white to black represents the degree of LD from low to high. LD blocks were defined using the solid spine of LD algorithm implemented
in Haploview.

Table 4
Effects of CGSs and parenting styles on creativity.
CGS Combinations of SNPs Parenting style Dimension β △R2 p

CGSa rs4648317(DRD2) rs5993882(COMT) rs5993883(COMT) rs6269(COMT) rs769224(COMT) M-authoritativeness Originality 0.166 0.027 0.001
CGSb rs7122246(DRD2) rs5993882(COMT) rs5993883(COMT) rs6269(COMT) M-authoritativeness Flexibility 0.189 0.036 0.000
CGSc rs7122246(DRD2) rs6279(DRD2) rs6269(COMT) F-authoritativeness Flexibility 0.152 0.023 0.001
CGSd rs7122246(DRD2) rs5993883(COMT) F-permissiveness Flexibility 0.159 0.025 0.001
CGSe rs7122246(DRD2) rs737865(COMT) F-authoritarianism Flexibility 0.118 0.013 0.014

Note: F-authoritativeness ¼ father authoritativeness, F-authoritarianism ¼ father authoritarianism, F-permissiveness ¼ father permissiveness, M-authoritativeness ¼
mother authoritativeness.

and six candidate genetic polymorphisms respectively approached sta- Thus, after the single variants analyses, we further computed five
tistical significance at 0.10 before multiple measure adjustment. As different CGSs of DRD2 and COMT, and examined their interactions with
complex phenotypes are controlled by many genes of small effect, these parenting style on creativity.
polymorphisms also have the potential to contribute to creativity. Through the DRD2 and COMT single variants analyses, susceptibility
variants and plasticity alleles were identified. Among them, DRD2
3.2.2. Cumulative genetic score (CGS) rs4648317(T), COMT rs5993882(G), COMT rs5993883(G), COMT
According to the cumulative genetic plasticity model, individuals will rs6269(G), COMT rs769224(A) might moderate the relationship between
be more susceptible to the environment if they carry more plasticity al- mother authoritativeness and originality; DRD2 rs7122246(A), COMT
leles. It made us hypothesize that the effect of parenting style on crea- rs5993882(G), COMT rs5993883(G), COMT rs6269(G) might moderate
tivity might be determined by individuals’ cumulative genetic plasticity. the relationship between mother authoritativeness and flexibility; DRD2

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S. Si et al. NeuroImage 213 (2020) 116681

rs7122246(A), DRD2 rs6279(G) and COMT rs6269(G) might moderate creativity, using single genetic polymorphism and cumulative genetic
the relationship between father authoritativeness and flexibility; DRD2 score analyses. When polymorphisms were examined in isolation, there
rs7122246(A) and COMT rs5993883(G) might moderate the relationship were only two polymorphisms from COMT, rs5993882 and rs5993883,
between father permissiveness and flexibility; DRD2 rs7122246(A) and which were found to interact with mother authoritativeness to predict
COMT rs737865(C) might moderate the relationship between father creativity. Specifically, maternal authoritative parenting was only asso-
authoritarianism and flexibility. Therefore, those potential variants sus- ciated with creativity in individuals with rs5993883 GG and rs5993882
ceptible to the same parenting style for the same creativity dimension GG genotype. Since both rs5993882 and rs5993883 are in intron, the
were combined and then five genetic profiles- CGS (a-e) were developed. exact biochemical effects of them are largely not clear. A recent G
E
Each of CGS (a-e) for individuals was computed by summing the number study found that, the second generation antipsychotic (SGA) treatment
of plasticity alleles (alleles indicated in parentheses above; 0, 1, or 2) for bipolar disorder was only associated with low verbal and cognitive
across the corresponding susceptibility variants. See Supplementary control in individuals with rs5993883 GG genotype (Flowers et al.,
Table 5 for the composition and distribution of CGS(a-e). 2016). This probably implied that individuals with rs5993883 GG ge-
Regression analyses indicated there was a significant two-way inter- notype were more susceptible than others to environmental influence.
action between CGSa and mother authoritativeness for predicting origi- Hence, the effect of parenting style on creativity-related cognitive func-
nality (β ¼ 0.17, p ¼ 0.001). The interaction in isolation accounted for tion may be more prominent in those people who are more likely to be
2.7% of the variance in originality. Additionally, the interaction between affected. As for rs5993882, no study has been conducted to reveal its
CGSb and mother authoritativeness (β ¼ 0.19, p ¼ 0.000), CGSc and fa- change due to plasticity. Since rs5993882 also located in intron 1, it is
ther authoritativeness (β ¼ 0.15, p ¼ 0.001), CGSd and father permis- assumed to have similar influence to rs5993883 in the relationship be-
siveness (β ¼ 0.16, p ¼ 0.001), CGSe and father authoritarianism (β ¼ tween parenting style and creativity. Future studies focusing on these two
0.12, p ¼ 0.014) on flexibility were also significant, accounting for polymorphisms are warranted. Together, COMT variants may interact
3.6%、2.3%、2.5% and 1.3% of the variance in flexibility respectively. with parenting style to predict creativity.
(see Table 4). However, the results of CGS analyses further indicated that these
interactions were moderated by DRD2 polymorphisms. Because it was
3.3. Sensitivity analyses found that CGS comprising of several potential susceptibility poly-
morphisms from DRD2 and COMT genes did indeed interact with
Sensitivity analyses were then performed to check whether the above parenting style to explain a significant amount of variance in the crea-
described significant CGS
parenting style interactions conformed to tivity. For flexibility, we found four genetic profiles (CGSb-e) that may
diathesis-stress model or differential susceptibility model. See Table 5 influence the relationship between parenting style and creativity. Spe-
and Fig. 3 for complete results. Take CGSa and mother authoritativeness cifically, the interaction of CGSe and authoritarian parenting style on
as an example, RoS on X and accompanying simple slopes tests revealed flexibility was significant. The negative effect of authoritarian parenting
significant effects of CGSa on originality at both high and low levels of style was only present in individuals with high CGSe, suggesting that
mother authoritativeness (within 2 SD). The RoS on Z test revealed individuals with high CGSe might be more susceptible to the adverse
significant effects of mother authoritativeness on originality for partici- environments, such as high father authoritarianism. This extends previ-
pants with high CGSa. Simple slopes of low CGSa and high CGSa differ ous findings by showing that the effect of father authoritarianism on
significantly from each other. Furthermore, the crossover point (origi- flexibility was moderated by both DRD2 and COMT. Besides, we for the
nality: -.37) was near 0 for mother authoritativeness which has been first time found the significant interaction of genetic profile (CGSb, CGSc,
standardized, PoI (originality: 0.45) was near 0.50, and PA (originality: CGSd) and positive parenting style (parental authoritativeness and father
0.64) was greater than 16% (Roisman et al., 2012). Nonlinear terms were permissiveness) on creativity. The post-hoc analyses further indicated
nonsignificant. Similar findings were found for CGSb, CGSc and CGSd on that the positive effects of mother authoritativeness, father authorita-
flexibility, although the G
E pattern regarding CGSe did not conform to tiveness and permissiveness were respectively more prominent in in-
diathesis-stress model or differential susceptibility model (see Supple- dividuals with high CGSb, CGSc and CGSd, although there were statistical
mentary Table 6 and Fig. 1). Therefore, generally speaking, all these possibilities that mother authoritativeness negatively predicted flexi-
statistical indexes provided support for the hypothesis of differential bility for individuals with low CGSb and father authoritativeness nega-
susceptibility over diathesis-stress. tively predicted flexibility for individuals with low CGSc. These probably
highlight flexibility will be affected by both negative and positive envi-
4. Discussion ronments. And the effect of environment on flexibility may be deter-
mined by several genetic profiles derived from COMT (frontal) and DRD2
The current study examined the interactive effects of variants in (striatal). Because the striatum and prefrontal cortex are strongly inter-
DRD2 and COMT genes and parenting styles (mother/father authorita- connected and conditioned by the neurotransmitter dopamine (Alex-
tiveness, authoritarianism, permissiveness) on individual differences in ander et al., 1986), CGS may influence flexibility—a key cognitive

Table 5
Regression estimates, regions of significance (RoS), and proportion of interaction index (PoI) for statistically significant (p<.05) CGS
parenting styles interactions.
RoS of X Simple slopes at 2 RoS of Z Simple slopes for two PoI PA X2 or ZX2 Crossover
SD X groups of Z

Lower bound Higher bound 2 SD þ2 SD Lower bound Higher bound 1 SD þ1 SD

CGSa
MA 1.18 .17 -.27** .40*** 1.32 .10 -.10 .24*** .45 .64 ns -.37
CGSb
MA -.96 .17 -.31** .43*** -.77 .33 -.15* .22** .43 .62 ns -.31
CGSc
FA -.85 .42 -.33** .38** 1.06 .27 -.13y .22** .36 .55 ns -.14
CGSd
FP 1.20 .16 -.27** .39*** 1.35 .08 -.10 .24*** .45 .65 ns -.37

Note. RoS, regions of significance; PoI, the proportion of interaction; PA, the proportion affected; MA, mother authoritativeness; FA, father authoritativeness; FP, father
permissiveness; X, parenting style; Z, cumulative genetic score (CGS); RoS on X indicates that outside of noted bounds there is a significant effect of CGS on creativity;
simple slopes of the effect of CGS on creativity at 2 SD of X are also presented. RoS on Z indicates that outside of noted bounds there is a significant effect of X on
creativity; simple slopes of the effect of X on creativity for two groups of CGS (low and high CGS) are also presented; X2 or ZX2 represents whether X2 or ZX2, or the set of
both nonlinear terms together was statistically significant in the equation Y ¼ b0 þb1X þ b2Z þ b3XZ þ b4X2þb5ZX2; ns, Not significant; Crossover denotes the value of X
(parenting style, standardized to M ¼ 0, SD ¼ 1) at which the regression lines intersected.

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S. Si et al. NeuroImage 213 (2020) 116681

Fig. 3. Interaction of CGS and parenting style on creativity. The gray shaded areas represent regions of significance (RoS) on X and denote where the two lines differ
significantly from each other within  2 SD. The triangular light blue shaded areas represent regions of significance (Ros) on Z and denote where the effect of X on
creativity is significant. The triangular pink shaded areas depict the proportion of the interaction (PoI). Values in the RoS are significant (see Table 5).

processes that support creativity (Nijstad et al., 2010) by modulating also affected flexibility. Since there is evidence that flexible processing
dopaminergic functioning of such fronto-striatal circuitries. benefits originality (Nijstad et al., 2010), our finding possibly makes us
For originality, we only found the interaction of CGSa and mother assume these polymorphisms might enhance originality by promoting
authoritativeness. And the following analysis revealed the positive effect flexibility which provides more possibilities for individuals to generate
of mother authoritativeness was only present in individuals with high new and different ideas (Baas et al., 2008; Nijstad et al., 2010). Besides,
CGSa. This also implies the effect of parenting style, particularly mother two polymorphisms (DRD2 rs4648317, COMT rs769224) were found
authoritativeness, on originality should be determined by aggregated merely influence originality—the hallmark of creativity, suggesting the
polymorphisms from COMT and DRD2. Despite less being known about genetic basis of originality is at least partly distinct from flexibility.
the underlying biological mechanisms for the observed interaction, two Therefore, these inspired us to suggest investigating G
E of creativity
important issues have to be mentioned. Firstly, three polymorphisms separated by each of its dimensions.
(COMT rs5993882, COMT rs5993883, COMT rs6269) included in CGSa Taken together, the effect of parenting style on creativity is not

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S. Si et al. NeuroImage 213 (2020) 116681

determined by a single variant of DRD2 or COMT, but combinations of between them. This to certain extent coincides with the vantage sensi-
them (CGS). They may influence creativity through their effects on DA tivity model that was newly proposed by Manuck (2011) and Sweitzer
neurotransmission. A wide range of evidence has established the role of et al. (2013). It mainly describes individuals’ different response to pos-
DA in motivation (Carlson, 2001). And a theory based on this evidence itive environments, arguing some individuals tend to be more susceptible
stating that the dopaminergic neural system may facilitate creativity to beneficial environments but are relatively unaffected by adverse sit-
through motivation (Flaherty, 2005). A recent empirical study showed uation (Pluess and Belsky, 2013). Nonetheless, a low level of father
motivation acted as a mediator between dopamine related gene and authoritarianism may not be the same as the exclusively positive envi-
divergent thinking (Takeuchi et al., 2015). These probably make us ronment described by Kegel et al., (2011). So it is still uncertain whether
speculate that DA mediated by DRD2 and COMT might influence crea- this finding has the potential to provide support for vantage sensitivity
tivity by affecting motivation or other dopamine-related cognitive pro- model. Another reason why we did not identify the interaction pattern
cess. According to Self-Determination Theory (SDT; Deci and Ryan, 2000, might be the serious skewed distribution of CGSe data in our sample.
2008), social environments (including parenting) influence individuals’ Future studies with larger sample need to retest this initial finding.
motivation which in turn, affects cognitive, affective, and behavioral In contrast to previous studies, we examined combinations of poly-
outcomes; therefore, motivation may be one of the mechanisms under- morphisms from DRD2 and COMT genes by using CGS which can help us
lying the interactive effect of CGS and parenting style on creativity. understand the dopamine cumulative effect of genetic influence. More
Future studies are warranted to test this hypothesis. concretely, we developed five CGSs consisting of two or more SNPs with
However, just as mentioned before, COMT not only affects dopamine, a consistent direction of effects with regard to a specific parenting style
but also norepinephrine (Beversdorf, 2018). Since the combined effect of that models the cumulative genetic plasticity on creativity. It helps us
norepinephrinergic and dopaminergic system were also found to mod- identify a multi-locus genetic profile that putatively influences dopamine
erate the association of environmental factors with creativity-related transmission and individual sensitivity to environment. This can to a
personality (Kazantseva et al., 2009), the present findings could not certain extent reduce multiple-testing problems of analyzing individual
exclude the possibility that both dopaminergic and norepinephrinergic polymorphisms (Docherty et al., 2011). Moreover, the present study
system may be involved in the relationship among CGS, parenting style pointed out CGS models presented here accounted for more variance
and creativity. Consequently, future studies should take into account the than did individual variants alone. This suggests even a relatively simple
effect of norepinephrinergic genes or their interactions with dopami- CGS, such as the one we created, can explain some of the genetic influ-
nergic genes in parenting style and creativity. Furthermore, it is generally ence that may have led to inconsistent discoveries in individual genetic
known that serotonin and dopamine neurotransmitters are closely variation analysis. Therefore, aggregating polymorphisms may become a
related with cognitive function, and neurotransmission mediated by se- better way to estimate genetic influence on phenotypes and could also be
rotonin transporter could be affected to some extent by DA and vice versa utilized in G
E research.
(Gonzalez-Burgos & Feria-Velasco, 2008). Although there was evidence This study is not without limitations. Relatively small sample size is
that one particular serotonergic gene might act as a moderator between arguably the most important. Replication of these preliminary findings
parenting style and creativity (Zhang et al., 2018), no study has consid- with a larger sample size is clearly warranted. In addition, we did not use
ered incorporating the potential role of dopaminergic genes among them. an objective measure of the parenting style, such as observations of
Finally, creativity is a complex trait determined by multiple genes (Power parenting behaviors. Future studies simultaneously employing both the
et al., 2015), as all of the dopamine, serotonin and norepinephrine sys- subjective and objective measures of parenting style would provide more
tem are involved in creative cognitive process, it is reasonable to suppose convincing results. Also, the findings may only be generalized to diver-
the relationship between parenting style and creativity might be gent thinking. According to Guilford (1950), convergent thinking is also
moderated by interaction between all of them. a main ingredient of creativity. It is considered a process of generating
Follow up tests using the Roisman criteria provided evidence for the one possible solution to a particular problem. There was evidence that
differential susceptibility model. Compared with individuals with low the relationship between individual spontaneous eye blink rate (EBR)—a
CGS, high CGS carriers are more likely to be affected by parenting style. clinical marker of dopaminergic functioning and convergent thinking is
This is in accordance with previous studies which found the more negative (more or less linear), implying that convergent thinking is
“plasticity” allele individuals carry, the more susceptible they will be to increasingly impaired by higher dopamine levels (Chermahini and
environmental influences (Belsky et al., 2007; Belsky et al., 2009; Belsky Hommel, 2010). Therefore, in the future studies, we’d like to investigate
and Pluess, 2009). In our present study, individuals with more plasticity whether there will be a negative relationship between convergent
alleles showed higher level of creativity under conditions of high thinking and COMT/DRD2. Last but not the least, since all variants
authoritativeness or permissiveness while demonstrated lower level of included are not likely to interact with the environment equivalently,
creativity when they are experiencing low authoritativeness or permis- future research should develop weighted CGS using separate samples,
siveness. In other words, the higher CGS makes individuals more sus- and the weights for each variant used in the CGS should be determined by
ceptible to an adverse environment with low authoritativeness or the first sample (Purcell et al., 2009).
permissiveness, but also to a more favorable environment with high In conclusion, the current study provides evidence of the combined
authoritativeness or permissiveness, for creativity. The link between effect of polymorphisms from DRD2 and COMT genes and parenting style
higher CGS and increased environmental influence on divergent thinking on creativity. Furthermore, it extends these findings to suggest the pos-
is also similar to existing examples of differential susceptibility for sibility of differential susceptibility, such that the individuals with high
cognition which suggests that individuals with more susceptibility alleles CGS were more likely to be affected by their both negative and positive
are more likely to be affected by environment in a better or worse manner parenting style, demonstrating higher and lower creativity. These results
(Gibbons et al., 2012). Thus combinations of DRD2 and COMT variations further provide evidence for the involvement of dopaminergic genes (i.e.
that are associated with environmental sensitivity may moderate the DRD2, COMT) in the relationship between parenting style and creativity
extent to which an individual is influenced by his or her environment. and suggest individuals with the greater susceptibility are the ones who
Notably, the interaction pattern of CGSe and father authoritarianism are more susceptible to environmental influences.
on creativity did not conform to either diathesis-stress or genetic sus-
ceptibility model. The result of Ros analysis indicated, when experi- Declaration of competing interest
encing a low level of father authoritarianism, individuals with high CGSe
demonstrated higher level of creativity than those individuals with low The authors declare that the research was conducted in the absence of
CGSe; however, when they are experiencing a high level of father any commercial or financial relationships that could be construed as a
authoritarianism, no significant difference in creativity was found potential conflict of interest.

9
S. Si et al. NeuroImage 213 (2020) 116681

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