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The Dopamine D4 Receptor Gene Shows a Gender-Sensitive Association With Cognitive Empathy: Evidence From Two Independent Samples


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Increasing evidence points to a role of dopaminergic pathways in modulating social behavior. Specifically, a polymorphic region in the third exon of the Dopamine D4 receptor (DRD4) has been associated with a host of social behaviors, often in an environment-sensitive manner. Empathy is thought to be an important motivator of prosocial behaviors and can be seen as multifaceted, combining cognitive empathy (CE) and emotional empathy (EE). In the current study, we analyzed the association between DRD4 and the 2 aspects of empathy, as well as the effect of gender on this association. In Study 1, a large sample of adult participants (N = 477) was inventoried for general empathy, CE, and EE and genotyped for the DRD4 exon 3 polymorphism. Women scored higher than men on all empathy measures and no main effect of genotype was observed. It is important that a significant interaction between genotype and gender emerged specifically for CE, with women carriers of the 7R-allele scoring higher than noncarriers, whereas in men 7R-carriers scored lower than -7R. Notably, these findings were replicated in an independently recruited sample (N = 121) in Study 2. The current report shows that the DRD4 exon3 polymorphism is associated with CE and the direction of the association is gender-sensitive. (PsycINFO Database Record (c) 2014 APA, all rights reserved).
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The Dopamine D4 Receptor Gene Shows a
Gender-Sensitive Association With Cognitive Empathy:
Evidence From Two Independent Samples
Florina Uzefovsky, Idan Shalev, Salomon Israel, Shany Edelman, Yael Raz, Nufar
Perach-Barzilay, David Mankuta, Simone G. Shamay-Tsoory, Ariel Knafo, and Richard P.
Online First Publication, May 26, 2014.
Uzefovsky, F., Shalev, I., Israel, S., Edelman, S., Raz, Y., Perach-Barzilay, N., Mankuta, D.,
Shamay-Tsoory, S. G., Knafo, A., & Ebstein, R. P. (2014, May 26). The Dopamine D4 Receptor
Gene Shows a Gender-Sensitive Association With Cognitive Empathy: Evidence From Two
Independent Samples. Emotion. Advance online publication.
The Dopamine D4 Receptor Gene Shows a Gender-Sensitive Association
With Cognitive Empathy: Evidence From Two Independent Samples
Florina Uzefovsky
Hebrew University Idan Shalev
Pennsylvania State University
Salomon Israel
Duke University Shany Edelman and Yael Raz
Hebrew University
Nufar Perach-Barzilay
University of Haifa David Mankuta
Hadassah Medical Organization, Jerusalem, Israel
Simone G. Shamay-Tsoory
University of Haifa Ariel Knafo
Hebrew University
Richard P. Ebstein
National University of Singapore
Increasing evidence points to a role of dopaminergic pathways in modulating social behavior. Specifi-
cally, a polymorphic region in the third exon of the Dopamine D4 receptor (DRD4) has been associated
with a host of social behaviors, often in an environment-sensitive manner. Empathy is thought to be an
important motivator of prosocial behaviors and can be seen as multifaceted, combining cognitive
empathy (CE) and emotional empathy (EE). In the current study, we analyzed the association between
DRD4 and the 2 aspects of empathy, as well as the effect of gender on this association. In Study 1, a large
sample of adult participants (N477) was inventoried for general empathy, CE, and EE and genotyped
for the DRD4 exon 3 polymorphism. Women scored higher than men on all empathy measures and no
main effect of genotype was observed. It is important that a significant interaction between genotype and
gender emerged specifically for CE, with women carriers of the 7R-allele scoring higher than noncarriers,
whereas in men 7R-carriers scored lower than 7R. Notably, these findings were replicated in an
independently recruited sample (N121) in Study 2. The current report shows that the DRD4 exon3
polymorphism is associated with CE and the direction of the association is gender-sensitive.
Keywords: cognitive empathy, emotional empathy, dopamine, DRD4
Supplemental materials:
Human beings have an incredible ability to understand and
relate to the emotional experiences of others. This capacity is often
referred to as empathy; it is central to smooth social interaction and
is believed to underlie much of moral development and prosocial
behavior (Batson et al., 1988; Hoffman, 2008). The literature
differentiates between the cognitive and emotional components of
empathy, which often occur in tandem, yet are conceptually dis-
tinct. Cognitive empathy (CE) is the ability to understand what the
Florina Uzefovsky, Psychology Department, Hebrew University, Je-
rusalem, Israel; Idan Shalev, Department of Biobehavioral Health,
Pennsylvania State University; Salomon Israel, Department of Psychol-
ogy and Neuroscience and Institute for Genome Sciences and Policy,
Duke University; Shany Edelman and Yael Raz, Neurobiology, Hebrew
University; Nufar Perach-Barzilay, Department of Psychology, Univer-
sity of Haifa, Haifa, Israel; David Mankuta, Hadassah Medical Orga-
nization, Department of Labor and Delivery, Jerusalem, Israel; Simone
G. Shamay-Tsoory, Department of Psychology, University of Haifa;
Ariel Knafo, Psychology Department, Hebrew University; Richard P.
Ebstein, Psychology Department, National University of Singapore,
We are grateful to all the participants and to the research assistants
who recruited them. Financial support (RPE) from the National Uni-
versity of Singapore, Ministry of Education at Singapore, the AXA
research foundation and the Templeton Foundation are gratefully ac-
knowledged. F. Uzefovsky was funded by the Arianne de Rothschild
Correspondence concerning this article should be addressed to Florina
Uzefovsky, Psychology Department, The Hebrew University of Jerusalem,
Jerusalem 91501, Israel. E-mail:
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Emotion © 2014 American Psychological Association
2014, Vol. 14, No. 3, 000 1528-3542/14/$12.00
other is feeling (a construct which is sometimes termed affective
Theory of Mind). Emotional empathy (EE), on the other hand, is
the ability to share the feelings of another, or to experience a
similar emotion, while still maintaining a self-other distinction
(Davis, 1983; Decety & Lamm, 2006; however, see Preston &
Hofelich, 2012). Although it is clear that during a typical empathic
response both the emotional and cognitive aspect of empathy come
into play, the distinction between CE and EE has nevertheless been
strongly supported by neuroimaging and neuropsychological
and clinical findings; for example, anorexia nervosa and autism
spectrum disorder (ASD) are associated with deficits in CE but
not in EE (Calderoni et al., 2013; Jones, Happé, Gilbert, Bur-
nett, & Viding, 2010), whereas anti-social personality disorder
(ASPD), is associated with deficits in EE but not in CE (Jones
et al., 2010). Although much remains to be learned regarding
the neurobiological underpinnings of CE and EE, evidence from
both neuroimaging and lesion studies also support a dual system
model of distinct, albeit somewhat overlapping, brain pathways
(reviewed in Blair, 2005; Shamay-Tsoory, 2011)). It has been
suggested that the broadly defined human Mirror Neuron Sys-
tem (hMNS) is associated with the feeling of emotional empa-
thy. The hMNS is activated both when one experiences a certain
feeling and when one perceives another person experiencing the
same feeling (e.g., pain matrix activated when feeling pain and
when perceiving another feeling pain). For example, activation
in the anterior cingulate cortex and in the anterior insula (both
part of the hMNS) is positively correlated with reported levels
of emotional empathy (Singer et al., 2004). On the other hand,
the mentalizing network (including the ventromedial prefrontal
cortex and the temporoparietal junction) underlies CE; that is,
the ability to understand what another is feeling (Saxe &
Kanwisher, 2003; Shamay-Tsoory, Aharon-Peretz, & Perry,
2009). This network is engaged when participants are asked to
make judgments on the mental states of others (Singer, 2006).
In the current study, we aimed to provide further understanding
into the neurobiological underpinnings of empathy and its two
facets by examining the association between common genetic
variation in an important gene from the dopaminergic system and
individual differences in overall empathy, as well as separately for
cognitive and emotional empathy. It is important that any discus-
sion of individual differences is not complete without examining
the role of gender in empathy. Therefore, we will first discuss the
association between gender and empathy and then turn to the
neurogenetic architecture of empathy.
Gender and Empathy
Gender has been repeatedly reported as a salient predictor of
empathy (Baron-Cohen & Wheelwright, 2004; Mehrabian & Ep-
stein, 1972), with females scoring higher than males on most
empathy measures. Disorders associated with deficits in empathy
such as ASD and ASPD are more prevalent in men than in women,
while disorders that are associated with higher empathy, such as
anxiety and depression, are more prevalent in women than in men
(Baron-Cohen, 2004; Moran, 1999; Zahn-Waxler, Shirtcliff, &
Marceau, 2008). Such gender differences may be explained simply
by hormonal factors, by culturally established gender roles, or by
both culture and hormones; and evidence for all views exists
(Baron-Cohen, 2004; Baron-Cohen, Knickmeyer, & Belmonte,
2005; Ickes, Gesn, & Graham, 2000; Karniol, Gabay, Ochion, &
Harari, 1998; Sagi & Hoffman, 1976). Therefore, irrespective of
whether the underlying mechanisms are social or biological, gen-
der differences must be examined in every investigation into
Genetics of Empathy
Given the importance of empathy to social functioning and the
marked individual differences in this trait, it is of considerable
interest to understand the genetic architecture of empathy. Indeed,
similar to other complex psychological traits, a meta-analysis of
twin studies indicates that the ability to empathize is moderately
heritable. It is important that when emotional and CE were con-
sidered separately, genetic effects of similar magnitude were ob-
served for both CE and EE, but a different pattern of environmen-
tal effects emerged, with individual variability in CE partially
explained by shared-environment effects (17%), and no such effect
for EE (Knafo & Uzefovsky, 2013).
The moderate heritability of empathizing ability suggests that a
search for candidate genes partially contributing to this phenotype
is worthwhile, and a good starting point is the neural pathways that
underlie social cognition in the brain.
Dopamine and Social Behavior
Dopamine is an important neurotransmitter in the “social brain”
neural network, with major influences on animal and human social
behavior (Skuse & Gallagher, 2009). It has been argued that
dopamine is crucial for empathy-motivated prosocial behavior,
which has evolutionary roots in offspring care (Preston, 2013). The
social hormones oxytocin (OT) and arginine vasopressin (AVP)
play a role in modulating dopaminergic receptors in the striatum,
orbitofrontal cortex, and other brain areas involved in social cog-
nition. Together with dopamine they comprise an integrated sys-
tem of social and communication skills (Skuse & Gallagher,
2009). Evidence suggests that social interactions are accompanied
by OT and AVP release, which in turn modulates dopaminergic
activity in the brain reward system. Thus, dopamine activity con-
tributes to experiencing social interactions as positive (Skuse &
Gallagher, 2009).
Collectively, the findings on the role of dopamine in social
interactions suggest that dopaminergic genes might play a crucial
role in empathy. Among the few genes encoding for synaptic
elements of brain dopaminergic activity, the Dopamine D4 Recep-
tor gene (DRD4) is a particularly attractive candidate. First, the
DRD4 gene is expressed in regions that have been shown to play
a role in empathy (Decety & Lamm, 2006), such as the amygdala
and the prefrontal cortex (Oak, Oldenhof, & Van Tol, 2000).
Second, converging evidence from molecular genetics studies sug-
gest that this candidate gene is associated with social cognition
(Kang, Namkoong, & Kim, 2008; Reiner & Spangler, 2011; Skuse
& Gallagher, 2009; Zhong et al., 2010). Most research on the
genetics of DRD4 focuses on a 16 amino acid repeat region (48 bp)
in exon 3 that has functional consequences (Van Craenenbroeck et
al., 2011). The most common repeat in Caucasian populations is 4
(4R), while the second most common variant is the 7 (7R) (Chang,
Kidd, Livak, Pakstis, & Kidd, 1996) that is characterized by a less
efficient receptor. The DRD4 exon3 repeat is robustly associated in
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most but not all studies as a risk allele for attention-deficit/
hyperactivity disorder (ADHD; Faraone, Doyle, Mick, & Bieder-
man, 2001), a disorder that was linked with empathy deficits
(Uekermann et al., 2010). Furthermore, in adults the 4R polymor-
phism has been associated with questionnaire measures of altruism
(Bachner-Melman et al., 2005) and with laboratory-based incen-
tivized “sense of fairness” using the ultimatum game (Zhong et al.,
It is interesting that the association between DRD4 and social
behavior is often contingent on an environmental context. That is,
carriers of the 7R allele seem to be more susceptible to the effects
of the environment on social behavior, and that is also the case
with prosocial behavior. Several studies have reported that a
child’s relationship with his or her parent is associated with proso-
cial behavior only for DRD4-7R carriers. Specifically, one study
showed that securely attached children donated more to UNICEF,
but only if they were carriers of the 7R-allele (Bakermans-
Kranenburg & van Ijzendoorn, 2011). In another study, positive
parenting was associated with mother-rated prosocial behavior,
and unexplained punishment was associated with self-initiated
prosocial behavior, only among 7R carriers (Knafo, Israel, &
Ebstein, 2010). Another study showed that religion priming in-
creased participants willingness to volunteer, but only when par-
ticipants were carriers of the susceptibility alleles (defined as the 7
or 2 repeat alleles; Sasaki et al., 2013). These findings are in line
with a relatively recent line of research that points to a unique role
of the 7R allele as an allele that confers higher sensitivity to
environmental effects:for better or for worse” (Belsky & Pluess,
2009). Notably, the prosocial behaviors associated with the DRD4
gene are likely underpinned by empathy (Batson et al., 1988;
Preston, 2013). Indeed, one study directly examined the associa-
tion between DRD4 and empathy in children, finding that maternal
negativity interacted with the DRD4 genotype to influence chil-
dren’s empathic response to an experimenter feigning physical
pain (Knafo & Uzefovsky, 2013). Taken together, these findings
suggest not only that the DRD4 gene likely contributes to empathy,
but that its role is complex and is modulated by both the internal
(hormonal) and external environment.
It is interesting that it seems that the effects of the DRD4
genotype on behavior are also moderated by gender. Several
studies report on gender-specific effects of the gene (that is,
the gene is associated with a behavior in one of the gender but not
the other) or on gender-contingent effects (that is, the effect of the
gene is different depending on gender; Froehlich et al., 2007;
Laucht, Becker, El-Faddagh, Hohm, & Schmidt, 2005; Laucht,
Becker, & Schmidt, 2006; Tammimäki & Männistö, 2011; Wang,
He, Chen, Huang, & Yeh, 2012; Zhong et al., 2010). These reports,
together with the crucial role that gender plays in individual
differences in empathy, suggests that the two factors (DRD4 and
gender) must be investigated jointly to better understand their
association with empathy.
The likely role of dopamine pathways in empathy as well as the
key role that the DRD4 plays in DA synaptic transmission provide
a promising avenue for research into the genetic architecture of
empathy. However, the complex role of DRD4 in social cognition
combined with the complex nature of empathy itself makes pre-
diction of direction of DRD4 on CE and/or EE uncertain. Never-
theless, the concept of cognitive flexibility (the ability to mentally
switch between different concepts) may be informative. It has been
shown that cognitive, but not emotional empathy is associated with
cognitive flexibility both in clinical (Shamay-Tsoory et al., 2007)
and nonclinical populations (Shamay-Tsoory, Tomer, Goldsher,
Berger, & Aharon-Peretz, 2004). Cognitive flexibility is consid-
ered to be a functional component of empathy and is primarily
associated with the ability to take the mental perspective of the
other; that is, CE (Decety & Jackson, 2004). Given that there is
evidence that the dopaminergic system underlies cognitive flexibly
(Müller et al., 2007), it is possible that the dopaminergic system
also predominantly underlies the cognitive, compared with the
emotional aspects of empathy. Supporting this notion is the recent
study implicating the dopaminergic system, and the DRD4 exon3
polymorphism specifically, in the development of Theory of Mind
(Lackner, Sabbagh, Hallinan, Liu, & Holden, 2012). However, a
direct investigation of the association between DRD4 exon3 and
cognitive and emotional aspects of empathy has not yet been
conducted, and that is a main aim of the current study.
The current investigation was designed to test the following
hypotheses. First, we hypothesized that the DRD4 exon3 polymor-
phism contributes to individual differences in empathy. To test our
hypothesis, we measured empathy and genotyped for the DRD4
exon3 polymorphism in two independently recruited samples of
university students. Second, following recent findings (reviewed
earlier), we hypothesized that the DRD4 exon3 polymorphism
would be more likely to contribute to individual differences in CE
compared with EE. Third, we sought to look at the effect of gender
as a consistent predictor of empathy, predicting that women would
score higher than men on all empathy measures. We also hypoth-
esized that the association between DRD4 and empathy would be
contingent on gender based on numerous findings that show that
the association between this gene and behavioral traits is often
gender-specific (Froehlich et al., 2007; Kang et al., 2008; Laucht
et al., 2005; Laucht et al., 2006; Reiner & Spangler, 2011; Zhong
et al., 2010). To our knowledge, this is the largest candidate gene
study of empathy so far reported.
Study 1
Participants. A total of 477 Hebrew University students (54%
women, mean age 24.42 2.67) were recruited by advertisements
on campus bulletin boards for a study on personality and genetics.
All subjects were aged 18–35; had no self-report history of psy-
chiatric disorders, chronic illness, or drug taking; and were non-
smokers. All participants were of Jewish descent. The project was
approved by the S. Herzog Hospital Institutional Review Board
committee and the Israeli Ministry of Health.
Measures. Participants came to the lab where informed con-
sent was obtained and DNA mouthwash samples were collected.
Participants were then given a demographic questionnaire and a
code name and password for an online website where they filled
out several questionnaire measures, among those three widely used
and highly validated self-report measures of empathy: (1) Inter-
personal Reactivity Index (IRI; Davis, 1980), which consists of 28
items on a 5-point scale, with a higher score conferring a better
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This article is intended solely for the personal use of the individual user and is not to be disseminated broadly.
empathic ability, as is for all the other measures. (2) Empathy
Quotient (EQ; Baron-Cohen & Wheelwright, 2004), which con-
sists of 60 items (40 empathy items and 20 filler items) on a
4-point scale. On each empathy item a person can score 2, 1, or 0.
(3) Questionnaire Measure of Emotional Empathy (QMEE; Meh-
rabian & Epstein, 1972), which consists of 33 items on a 9-point
scale. All these measures tap into emotional and cognitive aspects
of empathy; however, the EQ and the QMEE do not differentiate
between cognitive and emotional aspects of empathy, whereas the
IRI does. The IRI has four validated subscales that tap into CE and
EE. Two of the subscales measure CE (fantasy [F] and perspective
taking [PT]) with items such as “After seeing a play or movie, I
have felt as though I were one of the characters” (F) and “I try to
look at everybody’s side of a disagreement before I make a
decision” (PT). The other two subscale of the IRI tap into emo-
tional empathy (empathic concern [EC] and personal distress
[PD]), with items such as “I often have tender, concerned feelings
for people less fortunate than me” (EC) and “When I see someone
who badly needs help in an emergency, I go to pieces” (PD). An
online version of the questionnaires was used because of its
convenience and the increased sense of anonymity such a medium
provides, without compromising reliability or validity of measures
(Hertel, Naumann, Konradt, & Batinic, 2002).
DNA extraction and genotyping. DNA was extracted from
20 ml of mouthwash samples using the Master Pure kit (Epicenter,
Madison, WI). The DRD4 polymorphism (variable number of
tandem repeats of 48bp) in exon3 was characterized by a poly-
merase chain reaction (PCR) amplification procedure with the
CCTCATGGCCTTGCGCTC–3. PCR reactions were performed
using 5 l of Master Mix (Thermo scientific), 2 l primers (0.5
mol/L), 0.6 l mg/Cl2 (2.5 mM), 0.4 l DMSO 5%, and 1 lof
water to total of 9 l total volume and an additional 1 lof
genomic DNA was added to the mixture. All PCR reactions were
employed on a Biometra T1 Thermocycler (Biometra, Güttingem,
Germany). PCR reaction conditions were as follows: preheating
step at 94.0 °C for 5 min, 34 cycles of denaturation at 94.0 °C for
30s, reannealing at 55 °C for 30s, and extension at 72 °C for 90s.
The reaction proceeded to a hold at 72 °C for 5 min. The reaction
mixture was then electrophoresed on a 3% agarose gel (AMRESCO)
with ethidium bromide to screen for genotypes.
The distribution of genotype frequency was in Hardy-Weinberg
equilibrium (chi-square pvalue .791) and independent of ethnic
origin ((3)
25.05, p.168). See Table 1 for details regarding sex
and age by genotype.
Statistical analysis. Genotype of the DRD4 exon3 repeat was
coded as the presence or absence of the 7-repeat allele, resulting in
a two-level predictor variable. Together gender, DRD4 genotype
and their interaction served as predictors of questionnaire scores.
All statistical tests were carried out using SPSS v19 (Windows).
Study 2
Participants. A total of 121 Haifa University students (62%
female, mean age 24.60 1.97) were recruited through advertise-
ments on campus bulletin boards for a study on personality and
genetics. Selection criteria were identical to those in Study 1. All
participants were of Jewish descent.
Measures and analyses. Measures and genetic analyses were
identical to those in Study 1. The distribution of genotype fre-
quency was in Hardy-Weinberg equilibrium (chi-square pvalue
.488) See Table 2 for details regarding sex and age by genotype.
Study 1
The results section is planned to test our main hypotheses.
Section 1 of the results provides us with a basis for a composite
empathy score, which is used as the phenotype in the subsequent
analyses. In Section 2, we test the hypothesis that there is a
sex-contingent association between DRD4 and empathy. In Sec-
tion 3, we further analyze the role of the DRD4 exon3 polymor-
phism separately in males and females. In Section 4, we analyze
the association between DRD4 and both CE and EE.
(1) Correlation between empathy measures and computa-
tion of a composite score. Scores from all three measures
(IRI, EQ, QMEE) were standardized using Z-scores. As ex-
pected, because all three measures were designed to assess
empathy, the three variables were significantly correlated (all
correlations range from .45 to .70, p.001). Therefore, an
average score was computed for each participant, which was
also standardized using Z-scores. This standardized composite
measure of empathy was used as the phenotype to test associ-
ation with DRD4.
(2) The effect of gender and genotype on composite empathy
scores. We examined the role of genotype and gender on the
composite empathy score using a 2 (7R present vs. absent) 2
(male/female) analysis of variance (ANOVA). The analysis
yielded a significant effect for gender, with women scoring higher
than men, as hypothesized, F(1,402) 51.77,p.001. This
gender difference was qualified by a significant interaction be-
tween gender and DRD4-7R, F(1,402) 5.19, p.023. The main
effect of genotype was not significant, F(1,402) .07, p.79;
see details in Table 3. The interaction reflects the fact that female
carriers of the 7R had higher empathy scores than noncarriers,
whereas the opposite was observed for males. Male carriers of the
7R had lower empathy scores than noncarriers (Figure 1). It is
important that the results hold when comparing 7R carriers with
4R carriers only, Gender, F(1,303) 39.41, p.001; Genotype,
F(1,303) .094, p.76; Genotype Gender, F(1,303) 10.85,
Examination of the simple effects reveals that for both genotype
groups (7R present and 7R absent) women scored significantly
higher than men (p.001). Notably, the effect size (Cohen’s d),
of the difference between women and men on empathy scores was
substantially greater (d1.07) in carriers of the 7R allele than
Table 1
Demographics of the Study 1 Sample by Genotype
Genotype Sex Age
7RMen 157 (49.8%) 24.41 2.75
Women 158 (50.2%)
7RMen 70 (45.8%) 24.48 2.66
Women 83 (54.2%)
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noncarriers (d.51). Examination of the effect of genotype
separately for each sex revealed that the effect of genotype was
only marginally significant for women (p.08) and not signifi-
cant for men (p.15). Thus, the DRD4 genotype appears not to
have a direct across-the-board effect on empathy overall, but it
does interact with gender.
(3) The effect of genotype on cognitive and emotional
empathy. Following our second hypothesis, that the effect of the
DRD4-7R allele might be different for cognitive and emotional
empathy, we next calculated a composite score for CE and EE
separately. The separate scores were computed similarly to that of
the complete empathy score (by averaging Z-scores of the relevant
measures). The CE score was created from the two cognitive
subscales of the IRI: fantasy and perspective taking. The EE score
was created from the two emotional subscales of the IRI; empathic
concern and personal distress.
After extracting composite scores for CE and EE we tested for
an association between the DRD4 7R-allele and each facet of
empathy separately. Gender again had a main effect on both mean
scores; CE, F(1,402) 16.75, p.001; EE, F(1,402) 44.16,
p.001; and there was no main effect of DRD4 genotype; CE,
F(1,402) .63, p.80; EE, F(1,402) .57, p.45. Notably,
the interaction effect, DRD4 Gender, was highly significant for
CE, F(1,402) 13.79, p.001, but not for EE, F(1,402) .44,
p.51. Repeating the analyses with only participants homozy-
gous for the 4R compared with 7R carriers (77 or 47) yielded the
same results for EE; gender, F(1,303) 35.75, p.001; geno-
type, F(1,303) .83, p.37; Genotype Gender, F(1,303)
1.53, p.22; and for CE, Gender, F(1,303) 12.30, p.001;
Genotype, F(1,303) .20, p.65; Genotype Gender,
F(1,303) 19.64, p.001. Leading to the conclusion that
genotype had no main effect on empathy scores, and that the
significant interaction between gender and 7R-allele, which has
been previously observed for the overall empathy score, was
specific for CE.
Intriguingly, for women the presence of the 7R-allele is associ-
ated with higher CE scores whereas for men the opposite is
observed and the 7R-allele is associated with lower CE scores (see
details in Table 3 and Figure 2, as well as online Supplemental
Materials Figure 1). We again turned to examine the simple effects
to better understand the interaction. In the absence of the 7R-allele
there is no significant gender effect on CE, F(1,277) .117, p
.73. In contrast, the gender difference in CE in 7R participants
is highly significant, F(1,125) 21.54, p.001, with a large
effect size (Cohen’s d.83). Examination of the effect of geno-
type separately for men and women revealed that genotype was
significantly associated with CE scores for both sexes, but in
opposite directions. That is, in men, the 7R allele was significantly
associated with lower CE scores, F(1,192) 8.34, p.01,
Cohen’s d.461; whereas for women the 7R allele was associ-
ated with higher CE scores, F(1,210) 5.79, p.05, Cohen’s
Study 2
To check the robustness of the findings reported in Study 1, we
recruited a second, independent sample of participants and ana-
lyzed the association between the DRD4-7R, gender and empathy.
As previously noted, the gender effect was significant (p.001),
but the interaction effect (DRD4 Gender) on the total empathy
score was not significant (p.05), probably because of the
smaller sample size. However, the directionality of the effect was
similar to the effect found in Study 1; males carrying the 7R score
lower than noncarriers (mean scores were .43 .74 and .15
.99, respectively), and females carriers of the 7R score higher than
noncarriers (mean scores were .55 .86 and .33 1.11, respec-
We then turned to examine the effects of DRD4 and gender on
cognitive and emotional empathy separately. Again, as observed in
Study 1, a significant effect of gender was found for both empathy
facets; CE, F(1,121) 3.71, p.056; EE, F(1,121) 14.87, p
.001; and again, there was no main effect of DRD4 genotype; CE,
Table 2
Demographics of the Study 2 Sample by Genotype
Genotype Sex Age
7RMen 33 (38.8%) 24.37 1.80
Women 52 (61.2%)
7RMen 20 (47.6%) 25.05 2.26
Women 22 (52.4%)
Table 3
Means (SD) of Empathy Scores by Genotype and Gender
Genotype Gender Empathy Emotional
empathy Cognitive
7RMen .29 .74 .42 .93 .02 .93
Women .11 .82 .19 .98 .06 .97
7RMen .46 .70 .42 .93 .39 .83
Women .32 .76 .33 .96 .41 1.08
Total Men .34 .73 .42 .93 .11 .92
Women .18 .80 .23 .97 .17 1.02
Figure 1. The effect of gender and DRD4-7R allele on empathy based on
results of Study 1. Error bars indicate SEM.
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F(1,121) .21, p.65; EE, F(1,121) .002, p.96. As
observed previously in Study 1, the DRD4 Gender interaction
was significant for CE, F(1,121) 5.00, p.03, but not for EE,
F(1,121) .288, p.59. See details in Table 4. It is important
that the effects were in the same direction as Study 1, with women
carriers of the 7R-allele scoring higher than women noncarriers,
and men carriers of the 7R-allele scoring lower than men noncar-
riers (see Figure 3 and online Supporting Material Figure 2). As in
Study 1, the same results emerged when genotype was analyzed
for participants homozygous for the 4R compared with 7R carriers
(77 and 47) for EE; Gender, F(1,96) 11.63, p.001; Genotype,
F(1,96) .039, p.85; Genotype Gender, F(1,96) .71, p
.40; and similar results for CE with the exception of a nonsignif-
icant gender effect; Gender, F(1,96) 2.24, p.14; Genotype,
F(1,96) .42, p.52; Genotype Gender, F(1,96) 5.14, p
Again, in the absence of the 7R-allele there is no significant
difference in CE scores by gender, F(1,82) .06,p.80, whereas
the difference in CE for those men and women who carry the
DRD4 7R-allele is significant, F(1,39) 8.53, p.01, with a
large effect size (Cohen’s d.91). A complementary analysis of
the effect of genotype within each gender revealed a significant
effect of genotype on men’s CE scores, with men carriers of the 7R
allele scoring lower than noncarriers, F(1,51) 4.67, p.05,
Cohen’s d.63, and a nonsignificant effect in which women
7R-carriers scored higher than noncarriers on CE, F(1,70) 1.43,
p.24, Cohen’s d.32.
The current findings demonstrate a highly specific association
between the DRD4 exon3 polymorphism and CE in a sex-specific
manner, replicated in two independently recruited groups of stu-
dents from different universities. First, we find that the repeatedly
reported gender difference in empathy scores has been replicated
in both studies, again highlighting the importance of gender to
empathy research. Second, our findings point to an even greater
role of gender in empathy. In both studies, we find that gender
markedly modulates the impact of the 7R allele on CE. In women
the presence of the 7R allele enhances CE, whereas in men the 7R
allele is significantly associated with lower CE.
The sexual dimorphism shown here for the DRD4-7R is con-
sistent with a considerable literature showing that dopaminergic
pathways are often gender-sensitive (Tammimäki & Männistö,
2011; Wang et al., 2012). Indeed, a previous study examined a
related concept-sense of fairness (modeled in the Ultimatum
Game) with participants of East-Asian origin (Zhong et al., 2010).
In this study, the interaction between DRD4 genotype and gender
was significant and the effect of DRD4 was more pronounced for
males than for females. Similar Gene Gender interaction effects
have been reported for other phenotypes as well, including cogni-
tive function (Froehlich et al., 2007), novelty seeking (Laucht et
al., 2006), and smoking behavior (Laucht et al., 2005). In these
studies, effects of genotype were observed in only one of the sexes.
Additional evidence stem from the association between DRD4
genotype and ADHD (Li, Sham, Owen, & He, 2006), a disorder
with a strong sex bias that is characterized by deficits in empathy
(Marton, Wiener, Rogers, Moore, & Tannock, 2009). Taken to-
gether, these findings show that the observed effects of the DRD4
genotype, and specifically the 7R-allele, are often sex-specific or
sex contingent, with the direction of the effect dependent on the
phenotype examined.
The current gender-specific findings beg further consideration
into the underlying mechanisms of the sex-contingent effects of
DRD4 exon3 on empathy. First, does the DRD4 genotype moder-
ate the effects of gender, or is it the other way around and gender
moderates the effect of DRD4? Second, what does our measure of
In the Asian population the 7R allele is extremely rare and the 2R allele
is the second most common allele. Studies examining social phenotypes
such as novelty seeking, other-regarding emotions and ADHD (Kang,
Namkoong, & Kim, 2008; Leung et al., 2005; Reist et al., 2007) show that
in this population, the 2R allele serves a similar role to that of the 7R allele
in Caucasian populations.
Table 4
Means (SD) of Empathy Scores by Genotype and Gender
Genotype Gender Emotional empathy Cognitive empathy
7RMen .28 .83 .12 .92
Women .34 1.09 .18 1.18
7RMen .39 .85 .65 .78
Women .43 .93 .17 1.01
Total Men .32 .83 .32 .90
Women .36 1.04 .08 1.13
Figure 2. The effect of DRD4-7R genotype and gender on (A) cognitive empathy and (B) emotional empathy
based on results of Study 1. Error bars indicate SEM.
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sex/gender reflects (i.e., biological sex or social gender)? Alto-
gether, the currently observed Gene Gender interaction can be
interpreted in one of three ways, with each interpretation contain-
ing an underlying presupposition regarding these two questions.
First, we can consider gender as reflecting the genetic sex (i.e.,
XX or XY) of the participant. According to this view, the Gene
Gender interaction is in fact a Gene Gene interaction, making
the direction of the moderation (gene moderates sex or sex mod-
erates gene) less important.
The second possibility is to interpret gender as determining the
internal hormonal environment in which genetic products operate.
For example, women have higher levels of estrogen than men, and
these differences were shown to affect social behavior (van Anders
& Gray, 2007). Moreover, evidence suggests that sex hormones
such as estrogen affect dopamine neural pathways. Of particular
note is a study by Dreher et al. (Dreher et al., 2007), which
demonstrated augmented reactivity of the dopaminergic reward
system in women during the midfollicular phase, when estrogen is
unopposed by progesterone. In addition, behaviors associated with
addiction are mediated by dopaminergic brain reward network, and
are often different for men and women, presumably because of the
interaction between sex hormones and the dopaminergic reward
system (Becker & Hu, 2008). Thus, the Gene Gender interaction
reported in our study can be viewed as an interaction between a
gene and the internal-hormonal environment in which genes op-
erate. Future studies would benefit from examining empathy vis-
`-vis levels of hormones such as estrogen and testosterone, espe-
cially in light of the studies showing that empathy, facial mimicry,
and prosocial behavior are directly affected by sex hormones
(reviewed in Hastings, Zahn-Waxler, & McShane, 2006). Such
investigations can also shed more light on the direction of the
moderation observed in the current findings. For example, can a
manipulation of male hormone levels (e.g., testosterone) change
the direction of the interaction?
Finally, we can interpret gender as those behaviors that are
affected by social norms and perceptions regarding gender-
normative behavior, suggesting the interpretation of the interaction
as a Gene Social environment interaction. This view is sup-
ported by several findings that identify empathy as part of the
female societal gender role. For example, men and women tend to
define women in terms of relationships, and men in terms of
agency and independency (Guimond, Chatard, Martinot, Crisp, &
Redersdorff, 2006). The way gender is perceived is greatly influ-
enced by society, and therefore gender can be viewed as an
environmental factor that can in turn influence empathy. In a
meta-analysis of studies that assessed CE, it was found that only
after invoking female demand characteristics did women outper-
form men on this task (Ickes et al., 2000). A direct test of the
meta-analysis findings (Thomas & Maio, 2008) revealed that
women whose female stereotypical supremacy in CE was threat-
ened, had increased motivation to perform better at the task which
resulted in better performance when compared with women who
did not undergo the manipulation. It is interesting that when the
female gender role was evoked in men, they too perceived emo-
tions more accurately. Hence, increasing female demand charac-
teristics causes in turn an increase in motivation to perform better
in these types of tasks. From this point of view, it is possible to
consider the sex-contingent effect of DRD4 as reflecting a Gene
Environment interaction.
It is important to note here that although questionnaire measures
have the advantage of revealing a complex and stable character-
istic, they are also prone to reporter bias and demand characteris-
tics, which might have had an influence on the findings. For
example, perhaps carriers of the 7-repeat are more prone to rate
themselves (as opposed to really feel or act) according to gender
related cultural norms. Hence, future studies might employ other
measures of empathy (i.e., laboratory-based tasks) that are less
affected by these biases. Nevertheless, although cultural and ste-
reotypical differences in the role men and women take on in
society undoubtedly contribute to gender differences in empathiz-
ing ability, this influence might be moderated, as our results
suggest, by specific molecular genetic endowments. Thus, accord-
ing to this interpretation and our results, genotype enhances exist-
ing sex differences in those people who are more affected by the
environment, so that the DRD4 genotype moderates gender effects
on empathy. Indeed, future research might profit from exploring
how cultural attitudes interact with genotype in determining em-
pathizing ability.
The current finding that DRD4 interacts with gender to affect
cognitive but not emotional empathy supports the theoretical dis-
tinction between EE and CE and, furthermore suggests a biological
mechanism for this distinction. Although research in social and
developmental psychology had consistently differentiated between
the two aspects of empathy, the biological underpinnings of the
Figure 3. The effect of DRD4-7R genotype and gender on (A) cognitive empathy and (B) emotional empathy
based on results of Study 2. Error bars indicate SEM.
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two systems remained unclear. Recent advances in neuroimaging
and lesion research provided insights into the brain structures that
underlie EE and CE, but much remains to be learned. The current
findings contribute to a better understanding of the biological
mechanism of empathy by using a neurogenetic strategy. The
neurotransmitter dopamine is associated with a host of social
behaviors and emotional responses, many of them basic mecha-
nisms that do not involve higher order emotional or cognitive
processing (Preston, 2013). Therefore, it was important to under-
stand if the DRD4 gene is associated with emotional and cognitive
aspects of empathy. Indeed, the current study lends support to a
recent model that suggested dopamine as a key neurotransmitter in
cognitive but not emotional empathy (see Shamay-Tsoory, 2011
and Lackner et al., 2012). Moreover, this finding echoes our
previous finding of different etiologies for cognitive and emotional
empathy derived from a meta-analysis of twin studies, whereby
cognitive, but not emotional empathy, was found to be influenced
by shared environment (Knafo & Uzefovsky, 2013). In addition,
brain lesion studies (Shamay-Tsoory et al., 2009) and imaging
studies (Cox et al., 2011) strongly suggest that different neuronal
networks are involved in emotional and CE. This anatomical and
functional separation between emotional and CE is also supported
by some behavioral studies (Jones et al., 2010; Schwenck et al.,
2011). The current genetic evidence strengthens the notion that the
two empathy systems are distinct at the molecular and neural
levels. It is interesting that CE is both more complex and thought
to be phylogenetically more recent than emotional empathy (Pres-
ton & de Waal, 2003). It also depends on cognitive flexibility,
which is mediated by phylogenetically recent prefrontal cortices.
CE involves the ability to create a theory about the other’s mental
state and cognitively take the perspective of others, a feat almost
exclusively carried out by Homo sapiens with some possible
primate exceptions (Call & Tomasello, 2008). It is important that
the DRD4-7R allele is also considered to be a relatively recent allele
(Wang et al., 2004), perhaps explaining the specific association be-
tween this allele and the more phylogenetically advanced ability of
CE. Further research is needed to understand the intricate association
between empathy and the dopaminergic system. It is plausible that
other pathways within the dopaminergic system are associated with
the emotional aspects of empathy.
We currently identify one gene (of presumably many) under-
pinning the molecular genetic architecture of CE, providing a
partial understanding of the neural pathways likely accounting for
individual differences in this phenotype. Taken together, the cur-
rent findings not only open a window into the molecular architec-
ture of CE, but could also add to the Nature versus Nurture debate
regarding gender differences in empathizing abilities. Our findings
strengthen the argument that the sexual dimorphism observed in
empathy is at least partially genetically influenced. Moreover, they
add another possible interpretation of the gene by gender interac-
tion, fueled by recent research into Gene Environment effects,
that the term gender reflects both the biological and the social
environment. This interpretation, albeit somewhat speculative,
aims to propose another way of looking at the decades old debate
among researchers regarding the mechanisms, either biological or
social, underlying observed differences between men and women
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Received March 21, 2013
Revision received February 10, 2014
Accepted February 27, 2014
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... For example, genetically based dopaminergic and oxytocinergic activations affect stress and reward processing, thus affecting reactivity and approach-avoidance behavior (DeYoung, 2015a; Rodrigues et al., 2009;Yarkoni, 2015). Accordingly, they should stand at the basis of traits like neuroticism, extraversion, and openness to experience (Caspi & Shiner, 2007;Chong et al., 2019;DeYoung, 2015a), as well as empathy (Gong et al., 2017;Pearce et al., 2017;Uzefovsky et al., 2014Uzefovsky et al., , 2015. Indeed, a Genome-Wide Association Study (GWAS) found a genetic overlap between cognitive empathy and openness to experiences, as well as a marginal genetic overlap with brain volume in the dorsal striatum, a region enriched with dopaminergic input (Warrier et al., 2018). ...
... The behavioral content of the empathic personality profiles directs attention to systems affecting stress reactivity (for emotional empathy), affiliation, and approach tendencies, such as the oxytocinergic and dopaminergic systems. Indeed, genetic variations which encode for differences in the expression of oxytocin and dopamine were associated with empathy (Ben-Israel et al., 2015;Chong et al., 2019;Gong et al., 2017;Pearce et al., 2017;Rodrigues et al., 2009;Uzefovsky et al., 2014Uzefovsky et al., , 2015 as well as with affiliation, extraversion, and stressreactivity tendencies (Anacker et al., 2013;Fischer et al., 2018;Pearce et al., 2017;Rodrigues et al., 2009;Zheng et al., 2020). Although the specific effects of candidate genes do not consistently replicate, possibly due to low statistical power, small effect sizes, and numerous gene-environment interaction processes (Halldorsdottir & Binder, 2017;Montag et al., 2020), this body of literature and the present study's findings highlight the potential of investigating oxytocinergic and dopaminergic genetic contributions to both empathy and personality. ...
Objective How do genetic and environmental processes affect empathy during early adolescence? This study illuminated this question by examining the aetiology of empathy with the aetiology of other personality characteristics. Method Israeli twin adolescents rated their empathy and personality at ages 11 ( N = 1176) and 13 ( N = 821) (733 families, 51.4% females). Parents rated adolescents' emotional empathy. Adolescents performed an emotion recognition task, indicating cognitive empathy. Results Using a cross‐validated statistical learning algorithm, this study found emotional and cognitive “empathic personality profiles,” which describe and predict self‐reported empathy from nuanced Big‐Five personality characteristics, or “nuances” (i.e., individual items). These profiles predicted empathy moderately ( R ² = 0.17–0.24) and were stable and robust, within each age and between ages. They also predicted empathy in a new sample of older nontwin adolescents ( N = 96) and were validated against non‐self‐report empathy measures. Both emotional and cognitive empathy were predicted by nuances representing positive attitudes toward others, trust, forgiveness, and openness to experiences. Emotional empathy was also predicted by nuances representing anxiousness and negative reactivity. Twin analyses revealed overlapping genetic and environmental influences on empathy and the empathic personality profiles and overlapping environmental influences on empathy–personality change. Conclusions This study demonstrates how addressing the complexity of individuals' personalities can inform adolescents' empathy development.
... Indeed, a few candidate gene studies have shown relations between dopaminergic genes and self-reported empathic concern (Pearce et al., 2017;Ru et al., 2017). In addition, two studies found relations between the dopamine receptor gene D4 and cognitive empathy (Uzefovsky et al., 2014;Ben-Israel, Uzefovsky, Ebstein, & Knafo-Noam, 2015). However, in these two studies the effects were moderated in opposite directions by the participants' sex, which highlights the need of further research. ...
A range of empirical and theoretical perspectives on the relationship between biology and social cognition from infancy through childhood. Recent research on the developmental origins of the social mind supports the view that social cognition is present early in infancy and childhood in surprisingly sophisticated forms. Developmental psychologists have found ingenious ways to test the social abilities of infants and young children, and neuroscientists have begun to study the neurobiological mechanisms that implement and guide early social cognition. Their work suggests that, far from being unfinished adults, babies are exquisitely designed by evolution to capture relevant social information, learn, and explore their social environments. This volume offers a range of empirical and theoretical perspectives on the relationship between biology and social cognition from infancy through childhood. The contributors consider scientific advances in early social perception and cognition, including findings on the development of face processing and social perceptual biases; explore recent research on early infant competencies for language and theory of mind, including a developmental account of how young children become moral agents and the role of electrophysiology in identifying psychological processes that underpin social cognition; discuss the origins and development of prosocial behavior, reviewing evidence for a set of innate predispositions to be social, cooperative, and altruistic; examine how young children make social categories; and analyze atypical social cognition, including autism spectrum disorder and psychopathy. Contributors Lior Abramson, Renée Baillargeon, Pascal Belin, Frances Buttelmann, Melody Buyukozer Dawkins, Sofia Cardenas, Michael J. Crowley, Fabrice Damon, Jean Decety, Michelle de Haan, Ghislaine Dehaene-Lambertz, Xiao Pan Ding, Kristen A. Dunfield, Rachel D. Fine, Ana Fló, Jennifer R. Frey, Susan A. Gelman, Diane Goldenberg, Marie-Hélène Grosbras, Tobias Grossmann, Caitlin M. Hudac, Dora Kampis, Tara A. Karasewich, Ariel Knafo-Noam, Tehila Kogut, Ágnes Melinda Kovács, Valerie A. Kuhlmeier, Kang Lee, Narcis Marshall, Eamon McCrory, David Méary, Christos Panagiotopoulos, Olivier Pascalis, Markus Paulus, Kevin A. Pelphrey, Marcela Peña, Valerie F. Reyna, Marjorie Rhodes, Ruth Roberts, Hagit Sabato, Darby Saxbe, Virginia Slaughter, Jessica A. Sommerville, Maayan Stavans, Nikolaus Steinbeis, Fransisca Ting, Florina Uzefovsky, Essi Viding
... Thus, we have not equalized the gender and ages of our subsamples. Future studies could collect data for more genes, such as the oxytocin receptor (OXTR), which is associated with empathy traits (Luo et al., 2015), or the dopamine D4 receptor (DRD4) related to social behavior (Uzefovsky et al., 2014) and others.. We also suggest that larger samples could be examined in order to expand upon the knowledge of the association between genetics and personality. ...
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Background. Emotional intelligence is the ability to quickly and correctly recognize the emotional expressions of other people and to express and manage one’s own emotions. It contributes to the success of a person in activities related to communication and interaction with people. Emotional intelligence has been studied largely in the context of organizational and education psychology, but less is known about the influence of genetics on it. Objective. We aim to study emotional intelligence in carriers of different СОМТ, BDNF, DRD2, and HTR2A genotypes. Design. We used three methods to measure emotional intelligence. Mayer-Salovey-Caruso Emotional Intelligence Test is a set of tasks with forced choice and frequency-based correct responses. We also applied two self-report questionnaires by Lyusin and Hall. We recruited 280 participants who took part in all three measures. We also identified their genotypes of the СОМТ, BDNF, DRD2, and HTR2A genes. Results. Carriers of the Val/Met genotype of the COMT gene, A/A genotype of the HTR2A gene and C/C genotype of the DRD2 gene showed the highest level of emotional intelligence, while no differences were found between carriers of the BDNF genotypes. These data were obtained by using the Mayer-Salovey-Caruso Emotional Intelligence Test. Self-report scores of emotional intelligence did not differ between carriers of different genotypes across all four of the genes in question. Conclusion. Mayer-Salovey-Caruso Emotional Intelligence Test scores were differed for carriers of some genotypes, whereas self-reported emotional intelligence scores did not differ between according to genotype.
... A recent meta-analysis of twin studies (k = 23; Abramson et al., 2020) revealed that the dissociation between EE and CE is also supported at the genetic level with EE more influenced by heritability than CE (estimated 48.3% for EE and 26.9% for CE), and CE, unlike EE, depending on shared-environment (Abramson et al. suggested that cultural norms and beliefs about emotions were possible shred-environmental factors). There is some indication that the difference in heritability is subserved by different genes related to each trait (Pearce et al., 2017;Uzefovsky et al., 2014;Uzefovsky et al., 2015). Further support is provided by neuroimaging studies showing that CE and EE differ in brain activation and structure (de Waal & Preston, 2017;Shamay-Tsoory et al., 2008;Uribe et al., 2019). ...
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A large body of research showed that autistic people have intact emotional (affective) empathy alongside reduced cognitive empathy. However, there are mixed findings and these call for a more subtle understanding of empathy in autism. Empathic disequilibrium refers to the imbalance between emotional and cognitive empathy and is associated with a higher number of autistic traits in the typical population. Here we examined whether empathic disequilibrium predicts both the number of autistic traits and autism diagnosis. In a large sample of autistic ( N = 1905) and typical individuals ( N = 3009), we examined empathic disequilibrium and empathy as predictors of autistic traits and autism diagnosis, using a polynomial regression with response surface analysis. Empathy and autistic traits were measured using validated self‐report questionnaires. Both empathic disequilibrium and empathy predicted linearly and non‐linearly autism diagnosis and autistic traits. Specifically, a tendency towards higher emotional than cognitive empathy (empathic disequilibrium towards emotional empathy) predicted both autism diagnosis and the social domain of autistic traits, while higher cognitive than emotional empathy was associated with the non‐social domain of autism. Empathic disequilibrium was also more prominent in autistic females. This study provides evidence that beyond empathy as was measured thus far, empathic disequilibrium offers a novel analytical approach for examining the role of empathy. Empathic disequilibrium allows for a more nuanced understanding of the links between empathy and autism. Lay summary Many autistic individuals report feelings of excessive empathy, yet their experience is not reflected by most of the current literature, typically suggesting that autism is characterized by intact emotional and reduced cognitive empathy. To fill this gap, we looked at both ends of the imbalance between these components, termed empathic disequilibrium. We show that, like empathy, empathic disequilibrium is related to autism diagnosis and traits, and thus may provide a more nuanced understanding of empathy and its link with autism.
... In 1996 two studies indicated that people with the DRD4 7R allele engage in more 'novelty seeking' (impulsivity, thrill seeking) behavior presumably to stimulate more dopamine release in reward pathways because of the low binding affinity for dopamine in their D4 receptors (Benjamin et al., 1996;Ebstein et al., 1996). Since these publications, an explosion of studies indicating having 7 repeats or more on at least one DRD4 allele may be linked to: attention deficit hyperactivity disorder in children (Rowe et al., 1998), alcohol and drug abuse (Kotler et al., 1997), spirituality (Comings et al., 2000), empathy in women (Uzefovsky et al., 2014), delinquency, anger and thrill seeking in adolescent boys (Dmitrieva et al., 2011), infidelity and sexual promiscuity (Garcia et al., 2010) and increased creativity (Mayseless et al., 2013) among others. ...
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DNA segments with variable number tandem repeats (VNTR) serve as a model for students to learn DNA extraction and polymerase chain reaction (PCR) techniques in biology laboratory courses from the high school to the graduate level. Because of a growing interest in the neurosciences among undergraduates, we have developed a PCR exercise with a focus on the nervous system and behavior, with the aim of inspiring students from all aspects of the neurosciences to appreciate the central dogma and neurogenetics. DRD4 was a good candidate to provide a lab exercise that would be more engaging than VNTR analysis of a non-coding segment. DRD4 encodes for the dopamine D4 receptor and has been controversially associated with 'novelty seeking' or 'wanderlust' behavior. DRD4 has three common variants of the 48 bp sequence on exon 3, easily differentiated through gel electrophoresis. The 2 repeat (2R), 4 repeat (4R) and 7 repeat (7R) of the 48 bp sequence are the most common alleles. The 7R sequences result in the expressed dopamine D4 receptor having less affinity for dopamine binding, which was proposed to be the reason individuals engage in 'novelty seeking' behavior, to increase dopamine release to facilitate more binding to the receptor. Here we demonstrate an enjoyable and simple two lab sequence to analyze DRD4 genotypes that is appropriate for neuroscience and genetics courses.
... Adult women who carry the R7 allele have higher cognitive empathy than ones without it. It has also been shown that this correlation is reversed in adult males, meaning that this polymorphism is related to gender [152]. In children aged 3.5 and 5 years, respectively, a study focused on level of affective knowledge in two age and gender subgroups. ...
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Everyday life's hygiene and professional realities, especially in economically developed countries, indicate the need to modify the standards of pro-health programs as well as modern hygiene and work ergonomics programs. These observations are based on the problem of premature death caused by civilization diseases. The biological mechanisms associated with financial risk susceptibility are well described, but there is little data explaining the biological basis of neuroaccounting. Therefore, the aim of the study was to present relationships between personality traits, cognitive competences and biological factors shaping behavioral conditions in a multidisciplinary aspect. This critical review paper is an attempt to compile biological and psychological factors influencing the development of professional competences, especially decent in the area of accounting and finance. We analyzed existing literature from wide range of scientific disciplines (including economics, psychology, behavioral genetics) to create background to pursuit multidisciplinary research models in the field of neuroaccounting. This would help in pointing the best genetically based behavioral profile of future successful financial and accounting specialists.
... The selected SNPs have been previously reported to have an impact on social behavior [30] and other-oriented emotional and cognitive empathy [31][32][33][34][35]. Nevertheless, research in this field should be considered at the exploratory stage as the meta-analysis regarding selected SNPs of OXTR does not support the above-mentioned effects [36]. Thus, a novel perspective is the inclusion of more SNPs and analyzing empathy in the particular context-here, in couples [37][38][39]-and parental sensitive responsiveness [40][41][42][43]. ...
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Sensitive responsiveness refers to parents’ ability to recognize and respond to infants’ cues and has been linked to parental empathy. Additionally, oxytocin (OT) and vasopressin (AVP) are hormones important for sensitivity and empathy. The aim of this study is to test the links between dispositional empathy along with changing OT and AVP levels and responsiveness to a life-like doll in couples and to verify whether these factors are predictors of responsiveness to a child’s cues. Exploratory analyses include predictors of sensitive responsiveness: polymorphisms of OXTR, AVPR1a and CD38 genes, personal characteristics and relational factors. The project employs standardized experimental settings that can be used with non-parents and the assessment of parental sensitive responsiveness towards their child. The participants are couples expecting their first child (111) and childless couples (110). The procedure involves caretaking of a life-like doll. Salivary samples and questionnaire data are collected in a planned manner. In the second part, the expectant couples are invited for the assessment of their sensitivity to their own child (Free Play episodes). Parental sensitivity is assessed using the Ainsworth Sensitivity Scale. This paper presents an interdisciplinary research project that reaches beyond the questionnaire measurement, considering many factors influencing the dynamics of adult–infant interaction.
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We have studied important differences between the urbanization of "wild" species of birds and mammals and the domestication of domestic ones, together with the difference in stressors to which they adapt. In the first case, these are the most general characteristics of the urban environment - an extreme level of heterogeneity, instability and variability compared to any non-urban landscapes. It requires "urban" birds to quickly change nesting sites, feeding habitats, feeding methods and other features of biology in the wake of habitat changes, local and citywide, with the development of the ability to predict them, live, constantly "jumping from ice floe to ice floe" as opposed to sustainability existence in "rural" or "forest" populations. In the second, it is simply a change in the reaction to people, equipment, and animal care from an anxious-defensive to a friendly-interesting one. In the process of urbanization of "wild" species of birds, the brain increases, as in other variants of extreme habitats. Cognitive progress is achieved by each "urban" individual independently, due to the developing impact of the urban environment on its psyche, thanks to the growth of the possibilities for evaluating and predicting its dynamics using precursor signals. Therefore, it is preceded by an increase in the courage of individuals, a better differentiation of stimuli by them, a separation of significant ones from all the others (to which indifference is growing). On the contrary, during domestication, the brain decreases, cognitive progress in the new environment is achieved due to the “cooperative thinking”, social “prompts” of people and relatives. Behavioral changes during the urbanization of "wild" species are also sharply different from domestic ones. An analysis of aggression, defensive, exploratory behavior, attitudes towards the novelty of "urban" birds and mammals shows that they do not become "kind" or "trustful" (as happened with domestic animals). On the contrary: their aggression increases, along with courage and a better response to potential danger, a more accurate differentiation of it from “just anxiety”, to which they become more indifferent. As the species urbanizes, the behavior of each of the individuals becomes more diverse. Aggression, courage, flight from potential danger or vice versa, taking risks, exploring new places and objects no longer characterize individuals, but situations. All behavior is made as flexible and contextual as possible, with better recognition of the specifics of the situation, better choice of the mode of action, more accurate “dosing” of reactions according to the goal. This is the developing role of the urban environment. Urbanization destroys the behavioral syndrome: the correlations existing in the original populations between exploration, courage, aggression, risk taking, and in some cases other parameters. The launches of the forms of behavior characterized by each of these features in "urban" birds are mutually independent, in contrast to "rural" individuals. This maximizes the accuracy of the choice of behavior in a problem situation and its switching to another according to the situation, which is opposite to the changes associated with domestication and inconsistent with their explanation based on the D.K. Belyaev model. Urbanization changes the life strategy in a completely different way than domestication. The strategy of the newly formed urban population in the r-K-continuum shifts towards a “more pronounced K” compared to the initial one, due to a set of changes that mutually determine and reinforce each other: 1) Population growth occurs to a greater extent due to the lengthening of the average life expectancy, while reducing the reproduction of individuals (partly similar to the demography of Homo sapiens); 2) The primacy of future reproduction in the best conditions, with a directed movement to search for them “following the dynamics of the urban environment”, compared with the maximum reproductive effort “here and now”; 3) Greater “fractionality” of the reproductive potential of “urban” individuals, subdivided into a greater number of breeding attempts compared to the original population, with greater mobility and contextuality of each of them. Domesticated species behave in the opposite way and shift the strategy towards the "r-pole" of the continuum. A partly similar result during urbanization and domestication is achieved by opposite changes in the life strategy, behavior, cognitive characteristics, attitudes towards humans, and other aspects of the "natural history" of the species. The “individuality genes” DRD4 and SERT, which are under positive selection during urbanization, do not participate in domestication-related changes in the genome. On this basis, an evolutionary scenario for the urbanization of "wild" species is proposed, explaining on the same basis both differences and similarities with domestication. Modern cities, their expansion and association into groups (urbanization) are very interesting as arenas of the fastest microevolutionary processes that "separate" the urban population from the original and adapt it mainly to the most general features of the urban environment, but only gradually, then, and not completely - to specific influences that it "dumps" on individuals of this species. This happens in each region separately: the newly formed populations in different cities change parallel to each other (the same is true for urbanization changes in different, completely unrelated species), but do not approach each other in any way, remaining genetically closer to the local initial ones than to urban populations in other regions. Human-modified ("man-made") landscapes create extreme habitats, cities are its quintessence. We consider the adaptation of "wild" species of birds, partly mammals, to such changes, occurring through directed invasion into it and rapid changes in the population structure, ecology and behavior of individuals, restoring viability in new conditions and facilitating their even greater development, with penetration into areas, all more and more modified by man. The settlement of the urban environment is the culmination of all processes of this kind. Microevolution goes faster here than in natural landscapes (the faster, the stronger the transformation, with a maximum in the urban environment), but "gets stuck" at the stage of adaptation. Form formation - the appearance of "urban" subspecies, species, etc. - does not happen, despite the growing separation of "urban" populations from the original ones.
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Many people enjoy sad music, and the appeal for tragedy is widespread among the consumers of film and literature. The underlying mechanisms of such aesthetic experiences are not well understood. We tested whether pleasure induced by sad, unfamiliar instrumental music is explained with a homeostatic or a reward theory, each of which is associated with opposite patterns of changes in the key hormones. Sixty‐two women listened to sad music (or nothing) while serum was collected for subsequent measurement of prolactin (PRL) and oxytocin (OT) and stress marker (cortisol and adrenocorticotropic hormone) concentrations. Two groups of participants were recruited on the basis of low and high trait empathy. In the high empathy group, PRL and OT levels were significantly lower with music compared with no music. And compared to the low empathy group, the high empathy individuals reported an increase of positive mood and higher ratings of being moved with music. None of the stress markers showed any changes across the conditions or the groups. These hormonal changes, inconsistent with the homeostatic theory proposed by Huron, exhibit a pattern expected of general reward. Our findings illuminate how unfamiliar and low arousal music may give rise to pleasurable experiences. In this study, we tested whether pleasure induced by sad, unfamiliar instrumental music is explained with a homeostatic or a reward theory. Sixty‐two women, divided into high and low empathy groups, listened to sad music while serum prolactin (PRL) and oxytocin (OT) and stress marker concentrations were collected. In the high empathy group, PRL and OT levels were significantly lower with music, compared with silence. This group also reported an increase of positive mood with music and higher ratings of being moved, compared with the low empathy group.
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A surfeit of research confirms that people activate personal, affective, and conceptual representations when perceiving the states of others. However, researchers continue to debate the role of self-other overlap in empathy due to a failure to dissociate neural overlap, subjective resonance, and personal distress. A perception-action view posits that neural-level overlap is necessary during early processing for all social understanding, but need not be conscious or aversive. This neural overlap can subsequently produce a variety of states depending on the context and degree of common experience and emotionality. We outline a framework for understanding the interrelationship between neural and subjective overlap, and among empathic states, through a dynamic-systems view of how information is processed in the brain and body.
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Perception-action approaches are sometimes criticized because empathy takes cognitive forms and people do not overtly imitate or feel all observed states. These complaints reflect a misunderstanding of the framework, which we tried to clarify through a review that bridged social and neuroscientific views. Far from "simple fixes," these misunderstandings appear to reflect deeply rooted differences in the way that each discipline conceptualizes science and the mind. We address the important points made by the commentators and reiterate the need to incorporate rich, phenomenological descriptions into academic works so that we may prevent such conceptual cross-talk in the future. The open exchange of ideas across fields is often difficult, but essential to an integrated, scientific view of empathy.
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Background: A growing, but conflicting body of literature suggests altered empathic abilities in subjects with anorexia nervosa-restricting type (AN-R). This study aims to characterize the cognitive and affective empathic profiles of adolescents with purely AN-R. Methods: As part of a standardized clinical and research protocol, the Interpersonal Reactivity Index (IRI), a valid and reliable self-reported instrument to measure empathy, was administered to 32 female adolescents with AN-R and in 41 healthy controls (HC) comparisons, matched for age and gender. Correlational analyses were performed to evaluate the links between empathy scores and psychopathological measures. Results: Patients scored significantly lower than HC on cognitive empathy (CE), while they did not differ from controls on affective empathy (AE). The deficit in CE was not related to either disease severity nor was it related to associated psychopathology. Conclusion: These results, albeit preliminary, suggest that a dysfunctional pattern of CE capacity may be a stable trait of AN-R that should be taken into account not only for the clinical management, but also in preventive and therapeutic intervention.
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The current review aims to unify existing views of altruism through an examination of the biological bases of a fundamental form of giving: altruistic responding. Altruistic responding is most salient during heroic acts of helping but is also observed any time one perceives another's distress or need, which in turn motivates one to help at a current cost to the self. Such aid is simple, observable across species, and rooted in the instincts and circuits that evolved to maximize inclusive fitness through the care of helpless offspring. By design, the system already biases aid to both kin and nonkin under conditions that are largely adaptive. These inherent benefits are also buttressed in primates and humans by known, later-arriving benefits to helping in group-living animals. Evidence for the proposed homology between altruistic responding and offspring retrieval is presented through 10 key shared factors. Conceptually, both require (a) participation by nonmothers, (b) motor competence and expertise, (c) an adaptive opponency between avoidance and approach, and a facilitating role of (d) neonatal vulnerability, (e) salient distress, and (f) rewarding close contact. Physiologically, they also share neurohormonal support from (g) oxytocin, (h) the domain-general mesolimbocortical system, (i) the cingulate cortex, and (j) the orbitofrontal cortex. The framework intermixes ultimate and proximate levels of analysis and unifies existing views by assuming that even complex human behaviors reflect ancient mammalian neural systems that evolved to solve key problems in adaptive ways, with far-reaching consequences for even our most venerated human traits. (PsycINFO Database Record (c) 2013 APA, all rights reserved).
Research on the associations between hormones and human partnering represents an exciting advance in understanding human behavior, relationships, and sexuality. We review empirical literature on circulating hormones and human partnering (i.e., pair bonding, sexual/romantic relationships, pairing). We begin by reviewing steroids (e.g., androgens, estrogens, cortisol) and peptides (e.g., oxytocin, vasopressin). We then move on to androgen-partnering associations, attending to early stages of relationships, and partnering behaviors. A major focus includes relevant theoretical frameworks, possible directional or causal associations, and related controversies. We next cover associations between pair bonding, partnering behaviors, and both peptide (e.g., oxytocin) and stress hormones (e.g., cortisol). Following is a discussion of some contextual factors that might be important to understanding hormone- partnering associations, such as pregnancy or menopausal status. We conclude by summarizing and highlighting the main findings of partnering-hormone links and their implications; and we close by describing some of the challenges facing the field and some future directions given the field's current trajectory.
The dopamine D4 receptor (DRD4) is a candidate gene for schizophrenia because the dopaminergic system has been implicated in this neuropsychiatric disorder. Several research groups have reported an association between allelic variants at DRD4 and schizophrenia, while others have been unable to replicate that finding. Knowledge of the appropriate gene frequencies in the underlying populations may resolve these inconsistencies. We have determined the frequencies of 8 different alleles of the 48 bp imperfect tandem repeat of exon 3 at the DRD4 locus in samples from 33 populations around the world. The frequencies vary considerably in the different populations with the most common allele ranging from 16% to 95%. Frequencies and Fst values will be presented for the 3 most common alleles (4-, 7-, and 2- repeat) by continental groupings, but the individual populations vary significantly around the averages. The populations averaged 4.3 alleles (range 2 to 7).
To assess the relative contribution of genderand gender-role orientation to empathy and itsdevelopment, 8th and 11th grade Israeli adolescentscompleted Davis' empathy scale and Bem's gender-roleorientation inventory, which yields a femininity score anda masculinity score for each participant. Withmasculinity and femininity treated as a within subjectsvariable, masculinity/femininity interacted with grade, such that whereas masculinity in both boys andgirls increased with grade, femininity decreased ingirls and increased in boys. Empathy was unrelated tograde or to masculinity but was related to gender and to femininity. When the contribution ofmasculinity/femininity was covaried, empathy was foundto be unrelated to gender. This pattern was found forthree of the empathy subscales: Perspective Taking, Empathic Concern, and Personal Distress, whichwere positively correlated. On the basis of a mediansplit, participants were then classified as Masculine,Feminine, Undifferentiated, or Androgynous. Androgynous individuals did not differ from Feminineindividuals on any of the empathy subscales. Thefindings were discussed in terms of the socialization ofemotions and gender-role orientation.