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Current Directions in Psychological
Science
XX(X) 1­–7
© The Author(s) 2012
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DOI: 10.1177/0963721412452557
http://cdps.sagepub.com
The science of behavioral genetics, largely through twin
studies, has made a persuasive case that much of our
identity is stamped on us from conception; to that
extent, our lives seem to be pre-chosen—all we have to
do is live out the script that is written in our genes . . .
Still, it’s confounding to imagine . . . any gardener
knows that rather small variations of water and fertilizer
and soil acidity affect the development of far less com-
plex organisms—are humans so indifferent to the envi-
ronment they are planted in?
Wright, 1997, pp. 143-148
Personality traits are commonly defined as broad, disposi-
tional, and comparative features of psychological individual-
ity. The most prominent and extensively studied bipolar
personality-trait continua are neuroticism–emotional stability
and extraversion–introversion. Compared with more emotion-
ally stable persons, individuals who are high on neuroticism
are more likely to feel anxious, depressed, and guilty. They are
more often self-conscious and are more likely to experience
situations as threatening and stressful. Therefore, not surpris-
ingly, neuroticism is linked to several psychopathological
symptoms (Weinstock & Whisman, 2006). Individuals high on
extraversion are more active, gregarious, and assertive than
are introverts, who prefer to engage in fewer social activities
and to maintain fewer social contacts. Extraverts also engage
in more social and enterprising leisure-time and job-related
interests (Kandler, Bleidorn, Riemann, Angleitner, & Spinath,
2011).
Neuroticism and extraversion, or roughly parallel trait con-
ceptions, are often listed as the first two personality dimen-
sions in models of personality or temperament (see Bouchard
& Loehlin, 2001). Although facets of these traits, such as anxi-
ety (a facet of neuroticism) and activity (a facet of extraver-
sion), are already observable within the first years of life, a
clear structure of personality traits does not begin to appear
before the preteen period (McAdams & Olson, 2010).
Personality traits are enduring features accounting for con-
sistencies in behavior, thoughts, and feelings across situations
and occasions. Longitudinal studies of personality traits have
provided robust evidence that the stability of interindividual
Corresponding Author:
Christian Kandler, Department of Psychology, Bielefeld University,
Universitätsstraße 25, D-33615 Bielefeld, Germany
E-mail: christian.kandler@uni-bielefeld.de
Nature and Nurture in Personality
Development: The Case of Neuroticism
and Extraversion
Christian Kandler
Bielefeld University
Abstract
The stability of interindividual differences (i.e., rank-order continuity) in personality traits tends to increase with age until
it plateaus in middle adulthood and finally decreases in old age. Rank-order continuity also tends to decrease as the time
intervals between occasions of personality assessment increase, irrespective of age. These patterns show that personality
development is a lifelong process. Yet the sources of these patterns are unknown. Theories suggest that personality continuity
and change may result from environmentally mediated processes of identity development due to age-graded social roles and
individual life experiences, but also from biological maturation. Genetically informative longitudinal studies across different
age cohorts allow a differentiated picture of genetic and environmental sources. In this article, I give a short overview of the
genetic and environmental contributions to rank-order continuity and change in neuroticism and extraversion. Both genetic
and environmental factors contribute to personality continuity and change, but genetic factors affect rank-order change
only in younger decades of life, whereas environmental influences appear to represent a lifelong source of interindividual
differences in personality development.
Keywords
genetic continuity, environmental continuity, heritability, neuroticism, extraversion
2		 Kandler
differences over time (referred to hereafter as rank-order con-
tinuity) increases with age. In other words, a person’s specific
position on a personality-dimension continuum relative to the
positions of other people stabilizes with age. Stability coeffi-
cients (i.e., indices of rank-order continuity between occasions
of personality assessment; these coefficients can vary from 0,
no continuity, to 1, perfect continuity), which are low in child-
hood (approximately .30), rise across adolescence and young
adulthood, reaching a plateau (approximately .70) between the
ages of 40 and 70 (Roberts & DelVecchio, 2000; Terracciano,
Costa, & McCrae, 2006). In old age, stability coefficients
appear to decay (Lucas & Donnellan, 2011; Specht, Egloff, &
Schmukle, 2011). Irrespective of the age group studied, stabil-
ity coefficients decline as the time intervals between measure-
ments of personality traits get longer (Fraley & Roberts, 2005).
These patterns of stability and instability in individual ranks
on personality dimensions point to the fact that personality
development is a lifelong individual process. Although these
patterns of rank-order continuity in personality are well
known, their sources remain a matter of controversy.
McCrae and his colleagues (2000) formulated a theory of
biological maturation (McCrae & Costa, 2008). According to
this perspective, personality traits are stable, genetically (or at
least biologically) anchored basic tendencies of thoughts, feel-
ings, and behavior. Deviations from perfect continuity should
be due to short-term environmentally induced fluctuations, but
also to systematic and random measurement error. In fact, con-
trolling for artifacts and occasion-specific effects results in
higher heritability estimates (i.e., estimates of the degree to
which relative differences among individuals are due to
genetic factors) and larger stability coefficients (Kandler et al.,
2010). Furthermore, similar mean-level trends (i.e., trait
change over time averaged across individuals in a sample or
population) for neuroticism and for extraversion have been
found across different nations and cultures (McCrae et al.,
1999, 2000). Neuroticism tends to decrease with age, whereas
extraversion tends to be stable. Individual deviations from
the mean-level trend (i.e., interindividual differences in intra-
individual change) in neuroticism have been found to be
genetically influenced (Bleidorn, Kandler, Riemann, Angleit-
ner, & Spinath, 2009). These results support the hypothesis
that genetically programmed maturation is a propulsive mech-
anism of personality development. Consequently, the increase
in rank-order continuity of personality from childhood to
adulthood may result from individual differences in biological
maturation, and the decrease in such continuity in older age
may be explained by individual differences in biological
degeneration.
Other researchers have formulated different perspectives,
attributing changes in personality to environmental sources,
such as individual requirements in social roles (e.g., worker,
spouse, parent) and related investments (Roberts & Wood,
2006), normative life transitions (e.g., departure from one’s
parental home, the completion of school or an apprenticeship,
retirement), and individual life events that result in major
changes of individuals’ circumstances (e.g., accidents, the
birth of one’s own child, marriage). In other words, personal-
ity development and change are primarily attributable to age-
graded social roles and related experiences (Roberts, Wood, &
Caspi, 2008). Since the average direction of personality
change (i.e., mean-level change in a population) is often posi-
tive with respect to social functionality (e.g., people become
more emotionally stable with age), personality development is
considered to be a process leading to social maturity (Roberts
& Mroczek, 2008). From this perspective, the lower stability
coefficients in younger ages compared with coefficients in
middle adulthood are due to interindividual differences in the
life transitions and role transfers that typically occur during
the younger period of life, such as the completion of an appren-
ticeship and beginning of a job, departure from one’s parental
home, marriage, and new parenthood (Arnett, 2000). In fact,
this period of life is strongly shaped by individually different
developmental courses that are attributable to interindividual
differences in level and duration of education, family forma-
tion, and economic security (McAdams & Olson, 2010).
Social roles and contexts stabilize in midlife. They change
again in old age until social adjustment breaks down in
advanced old age. Consequently, the increase in rank-order
continuity of personality traits from adolescence to adulthood,
the plateau in middle adulthood, and the decrease of continuity
in older age may result from interindividual differences in sta-
bilizing, maintaining, and destabilizing one’s personal identity
in life.
On the basis of these theoretical considerations, I suggest
an integrative perspective. That is, both genetic and environ-
mental influences may explain personality development, at
least in the case of the rank-order continuity of personality
traits, and at least in the case of neuroticism and extraversion.
Longitudinal behavioral genetics studies (e.g., longitudinal
twin or adoption studies) on different age cohorts have allowed
for deeper insight. In the following sections, I briefly review
und discuss the results from these studies with respect to the
patterns of genetic and environmental effects on rank-order
continuity in neuroticism and extraversion.
The Role of Genetic Factors in Personality
Continuity and Change
Using genetically informative designs, the interindividual dif-
ferences in trait scores that are due to genetic differences (i.e.,
heritability) and to environmental influences can be disentan-
gled. Genetically informative longitudinal designs allow a dis-
entanglement of genetic and environmental components on
each of the measurement occasions, as well as the calculation
of the level of continuity of each of these components (i.e.,
genetic and environmental continuity).
It has been consistently shown that genetic continuity is far
from perfect (i.e., stability coefficients are consistently less
than 1) during childhood and adolescence (De Fruyt et al.,
2006; Gillespie, Evans, Wright, & Martin, 2004; Spengler,
Nature and Nurture of Personality Development	 3
Gottschling, & Spinath, 2012). That is, across childhood and
adolescence, new genetic factors emerge that contribute to
interindividual differences and rank-order changes in person-
ality. Like rank-order continuity, genetic continuity tends to
increase across adolescence and young adulthood (Hopwood
et al., 2011; Viken, Rose, Kaprio, & Koskenvuo, 1994) until it
reaches almost perfect stability in middle and late adulthood
(W. Johnson, McGue, & Krueger, 2005; Pedersen, 1993; Read,
Vogler, Pedersen, & Johansson, 2006).
This pattern of genetic continuity has been found for neu-
roticism and extraversion or related variables (see green lines
in Fig. 1 for neuroticism and in Fig. 2 for extraversion). There-
fore, the steady increase in observable rank-order continuity
from childhood to adulthood can be accounted for by stabiliz-
ing interindividual genetic differences. The relatively high sta-
bility of interindividual personality differences in middle
adulthood is accompanied by stability in genetic differences.
In other words, rank-order continuity in neuroticism and extra-
version can be explained by interindividual differences in
genetically influenced maturation.
The Role of Environmental Factors in
Personality Continuity and Change
Stable genetic differences among individuals after age 40,
however, cannot account for the decay of stability coefficients
in older age. Therefore, nongenetic factors may also contribute
to rank-order continuity. Across a broad spectrum of geneti-
cally informative studies, A. M. Johnson, Vernon, and Feiler
(2008) found that the primary nongenetic sources of interindi-
vidual differences in neuroticism and extraversion are specific
to individuals (i.e., not shared by family members reared
together). These effects are known as nonshared environmen-
tal effects. From a behavioral genetics perspective, “environ-
mental” effects include physiologically and biochemically
mediated effects, such as environmentally activated epigenetic
influences. Nonshared environmental factors show different
degrees of stability (i.e., environmental continuity) with
respect to the age studied.
Studying twins between the ages of 12 and 16, Gillespie
et al. (2004) found rather low coefficients for environmental
continuity for both extraversion (.17 for females and .18
for males) and neuroticism (.12 for males and .36 for females).
Hopwood and his colleagues (2011) studied the transition
from adolescence to adulthood (age 17 to 24). The researchers
found environmental-continuity coefficients of .36 for nega-
tive emotionality (neuroticism), .37 for communal positive
emotionality, and .39 for agentic positive emotionality (the
latter two traits can be seen as facets of extraversion). They
also investigated rank-order continuity between the ages of
24 and 29. For this interval, they found higher environmental
continuity, with coefficients ranging from .56 to .60. For
.2
.3
.4
.5
.6
.7
.8
.9
1.0
5 10 15 20 25 30 35 40 45 50 55 60 65 70 75 80 85 90
Age
StabilityCoefficient
Fig. 1. Genetic continuity (depicted by the green line and data points) and environmental
continuity (depicted by the red line and data points) of interindividual differences in neuroticism
(or negative emotionality) as a function of age. Higher stability coefficients (i.e., indices of
continuity between assessments) indicate greater stability over time. The stability coefficients
shown are based on the results of genetically informative longitudinal studies (weighted by sample
sizes): Bratko and Butkovic (2007), De Fruyt et al. (2006), Gillespie, Evans, Wright, and Martin
(2004), Hopwood et al. (2011), W. Johnson, McGue, and Krueger (2005), Kandler et al. (2010),
McGue, Bacon, and Lykken (1993), Pedersen (1993), Read, Vogler, Pedersen, and Johansson
(2006), Spengler, Gottschling, and Spinath (2012), Viken, Rose, Kaprio, and Koskenvuo (1994),
and Wray, Birley, Sullivan, Visscher, and Martin (2007). Across studies and sample cohorts,
intervals between assessments ranged between 2 and 10 years (M = 5.15 years).
4		 Kandler
middle adulthood, W. Johnson and colleagues (2005) reported
5-year environmental-continuity coefficients over .70. How-
ever, other studies have indicated that environmental continu-
ity decreases in later adulthood (Pedersen & Reynolds, 1998;
Read et al., 2006).
These findings and results from other genetically informa-
tive studies (see red lines in Figs. 1 and 2) point to the fact that
changing levels of rank-order continuity in neuroticism and
extraversion are also attributable to changing levels of envi-
ronmental continuity across the life span. Moreover, the
decrease in rank-order continuity in older age appears to be
exclusively due to a decline in the stability of environmental
factors. Therefore, interindividual differences in experiences
of major life transitions or major life events (whether tempo-
ral, quantitative, or qualitative) may contribute to personality
change and, thus, to personality development. Environmental
contributions to change in interindividual ranks may occur
across the entire life span, but they are more pronounced in
younger ages (because of events such as leaving one’s parental
home, starting a job, and forming a new family) and older ages
(because of events such as retirement, illness, death of spouses
or friends). Because “environmental” factors also include bio-
logically mediated effects, the declining rank-order continuity,
especially in later adulthood, might be due to biological causes
(e.g., multi-infarct dementia).
The Interplay of Genetic and
Environmental Factors
As life narratives and memories cumulate across the life span,
effects of life experiences may cumulate, too. Thus, environ-
mental effects on interindividual differences in neuroticism
and extraversion may cumulate with age, leading to an increase
in environmental variance (McCartney, Harris, & Bernieri,
1990). If interindividual differences due to environmental
effects increase with age, stable interindividual differences
accounted for by genetic effects should proportionally
decrease. Consequently, heritability estimates for personality
traits should decrease with age.
Most genetically informative studies that have assessed
heritability estimates for different age cohorts (e.g., Loehlin &
Martin, 2001) have not found significant differences between
age groups. However, other studies that have combined cross-
sectional data (from different age cohorts) and longitudinal
data have suggested that heritability estimates decrease with
age (Kandler et al., 2010; Viken et al., 1994). A meta-analytic
review of fifteen genetically informative studies on different
age groups showed that the level of heritability of neuroticism
and extraversion appears to depend on the age studied (see
Fig. 3). Heritability of extraversion tends to increase until peo-
ple are in their early 40s, but then it declines continuously. For
.2
.3
.4
.5
.6
.7
.8
.9
1.0
5 10 15 20 25 30 35 40 45 50 55 60 65 70 75 80 85 90
Age
StabilityCoefficient
Fig. 2.  Genetic continuity (depicted by the green line and data points) and environmental continuity
(depicted by the red line and data points) of interindividual differences in extraversion (or positive
emotionality) as a function of age. Higher stability coefficients (i.e., indices of continuity between
assessments) indicate greater stability over time. The stability coefficients shown are based on
the results of genetically informative longitudinal studies (weighted by sample sizes): Bratko and
Butkovic (2007), De Fruyt et al. (2006), Gillespie, Evans, Wright, and Martin (2004), Hopwood
et al. (2011), W. Johnson, McGue, and Krueger (2005), Kandler et al. (2010), McGue, Bacon,
and Lykken (1993), Pedersen (1993), Read, Vogler, Pedersen, and Johansson (2006), Spengler,
Gottschling, and Spinath (2012), and Viken, Rose, Kaprio, and Koskenvuo (1994). Across studies
and sample cohorts, intervals between assessments ranged between 2 and 10 years (M = 4.88
years).
Nature and Nurture of Personality Development	 5
neuroticism, heritability progressively decreases after people
enter adulthood.
The changing importance of genetic contributions to inter-
individual differences in neuroticism and extraversion with
age differs from the pattern observed with regard to cognitive
abilities. For cognitive abilities, heritability continuously
increases with age (W. Johnson, 2010). One reason for this
distinction may be that genes and environmental effects dif-
ferentially contribute to how a person typically acts (i.e., his or
her temperament or narrow-sense personality) and how capa-
ble a person is (i.e., his or her abilities or skills).
An increase in the level of personality traits’ heritability
may reflect the increasing importance of active and reactive
gene-environment correlations. People actively select, create,
and change their environments (e.g., by changing their friends,
residences, or jobs) or evoke social reactions that match their
genetically predisposed traits. Environments and social reac-
tions, in turn, may affect personality traits. For example, indi-
viduals who are genetically predisposed to be more extraverted
may perceive life events as more controllable and positive;
experiencing life events as controllable and positive, in turn,
may augment such people’s levels of extraversion (Kandler,
Bleidorn, Riemann, Angleitner, & Spinath, 2012). That is,
genetic influences may partly drive the environmental effects
accounting for the increasing heritability. The processes
involved in these gene-environment correlations may be pro-
pulsive mechanisms of the development of cognitive abilities
across the life span and the development of extraversion in the
first decades of life.
The increasing heritability of extraversion until age 30 and
the cumulatively decreasing heritability of neuroticism and
extraversion after age 30 may accompany midlife tipping
points. For example, young adults often try to expand the self
and gain new information—they actively try to change their
environments to match their goal pursuits; by contrast, older
adults more often try to stabilize and protect the quality of
their lives and ongoing relationships—they change the self to
fit the environment (see McAdams & Olson, 2010, for a
review). Consequently, genetic and environmental effects do
not act separately; rather, they interact in complex ways that
account for the development of personality traits, and the con-
tribution of life experiences to personality development cumu-
lates with age.
Conclusion
I hope that this short review will open the door for further
research identifying the mechanisms of the gene-environment
.05
.15
.25
.35
.45
.55
.65
.75
5 10 15 20 25 30 35 40 45 50 55 60 65 70 75 80 85 90
Age
HeritabilityEstimate
Fig. 3. Heritability estimates (the proportion of interindividual differences accounted for by
genetic differences) for extraversion (depicted by the blue line and data points) and neuroticism
(depicted by the orange line and data points) as a function of age. The heritability estimates shown
are based on the results of genetically informative longitudinal and age-cohort studies (weighted
by sample size): Bratko and Butkovic (2007), De Fruyt et al. (2006), Gillespie, Evans, Wright, and
Martin (2004), Hopwood et al. (2011), W. Johnson, McGue, and Krueger (2005), Kandler et al.
(2010), Loehlin and Martin (2001), McGue, Bacon, and Lykken (1993), Pedersen and Reynolds
(1998), Read, Vogler, Pedersen, and Johansson (2006), Rettew et al. (2006), Rettew, Rebollo-
Mesa, Hudziak, Willemsen, and Boomsma (2008), Spengler, Gottschling, and Spinath (2012),
Viken, Rose, Kaprio, and Koskenvuo (1994), and Wray et al. (2007).
6		 Kandler
interplay in personality development and also focusing on the
patterns and sources of other personality traits (e.g., openness,
agreeableness, and conscientiousness). More genetically and
environmentally informative research is sorely needed. At this
point, we know that genetic factors stabilize with age, leading
to an increase in rank-order continuity of neuroticism and
extraversion and to personality formation across childhood,
adolescence, and young adulthood. Environmental sources
contribute to this increasing continuity but also to decreasing
continuity in old age. Moreover, environmental influences
appear to cumulate across the life span, leading to a decline of
the genetic contribution to variance in neuroticism and extra-
version with age, a pattern pointing to lifelong changes in per-
sonality due to environmental influences. We develop into the
people we are predisposed to become because of our genetic
makeup, but to a large degree, the development of or changes
in our personality will be formed by the environment.
Recommended Reading
Bouchard, T. J., & Loehlin, J. C. (2001). (See References). A com-
prehensive review of research on personality traits and related
variables from a behavioral genetics perspective.
Fraley, C., & Roberts, B.W. (2005). (See References). An compre-
hensive work on patterns of continuity described as mathematical
models.
McAdams, D. P., & Olson, B. D. (2010). (See References). A clearly
written, user-friendly, and empirically founded theoretical per-
spective on personality development of different levels of per-
sonality elements, such as traits, motives, and narratives.
McCrae, R. R., & Costa, P. T. (2008). (See References). A compre-
hensive overview of research and theory on personality traits as
biologically anchored basic tendencies.
Wright, L. (1997). (See References). An interesting and reader-
friendly book on the history of twin studies and the secrets of
our identity.
Declaration of Conflicting Interests
The authors declared that they had no conflicts of interest with
respect to their authorship or the publication of this article.
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Nature, Nurture, & Neuroticism

  • 1. Current Directions in Psychological Science XX(X) 1­–7 © The Author(s) 2012 Reprints and permission: sagepub.com/journalsPermissions.nav DOI: 10.1177/0963721412452557 http://cdps.sagepub.com The science of behavioral genetics, largely through twin studies, has made a persuasive case that much of our identity is stamped on us from conception; to that extent, our lives seem to be pre-chosen—all we have to do is live out the script that is written in our genes . . . Still, it’s confounding to imagine . . . any gardener knows that rather small variations of water and fertilizer and soil acidity affect the development of far less com- plex organisms—are humans so indifferent to the envi- ronment they are planted in? Wright, 1997, pp. 143-148 Personality traits are commonly defined as broad, disposi- tional, and comparative features of psychological individual- ity. The most prominent and extensively studied bipolar personality-trait continua are neuroticism–emotional stability and extraversion–introversion. Compared with more emotion- ally stable persons, individuals who are high on neuroticism are more likely to feel anxious, depressed, and guilty. They are more often self-conscious and are more likely to experience situations as threatening and stressful. Therefore, not surpris- ingly, neuroticism is linked to several psychopathological symptoms (Weinstock & Whisman, 2006). Individuals high on extraversion are more active, gregarious, and assertive than are introverts, who prefer to engage in fewer social activities and to maintain fewer social contacts. Extraverts also engage in more social and enterprising leisure-time and job-related interests (Kandler, Bleidorn, Riemann, Angleitner, & Spinath, 2011). Neuroticism and extraversion, or roughly parallel trait con- ceptions, are often listed as the first two personality dimen- sions in models of personality or temperament (see Bouchard & Loehlin, 2001). Although facets of these traits, such as anxi- ety (a facet of neuroticism) and activity (a facet of extraver- sion), are already observable within the first years of life, a clear structure of personality traits does not begin to appear before the preteen period (McAdams & Olson, 2010). Personality traits are enduring features accounting for con- sistencies in behavior, thoughts, and feelings across situations and occasions. Longitudinal studies of personality traits have provided robust evidence that the stability of interindividual Corresponding Author: Christian Kandler, Department of Psychology, Bielefeld University, Universitätsstraße 25, D-33615 Bielefeld, Germany E-mail: christian.kandler@uni-bielefeld.de Nature and Nurture in Personality Development: The Case of Neuroticism and Extraversion Christian Kandler Bielefeld University Abstract The stability of interindividual differences (i.e., rank-order continuity) in personality traits tends to increase with age until it plateaus in middle adulthood and finally decreases in old age. Rank-order continuity also tends to decrease as the time intervals between occasions of personality assessment increase, irrespective of age. These patterns show that personality development is a lifelong process. Yet the sources of these patterns are unknown. Theories suggest that personality continuity and change may result from environmentally mediated processes of identity development due to age-graded social roles and individual life experiences, but also from biological maturation. Genetically informative longitudinal studies across different age cohorts allow a differentiated picture of genetic and environmental sources. In this article, I give a short overview of the genetic and environmental contributions to rank-order continuity and change in neuroticism and extraversion. Both genetic and environmental factors contribute to personality continuity and change, but genetic factors affect rank-order change only in younger decades of life, whereas environmental influences appear to represent a lifelong source of interindividual differences in personality development. Keywords genetic continuity, environmental continuity, heritability, neuroticism, extraversion
  • 2. 2 Kandler differences over time (referred to hereafter as rank-order con- tinuity) increases with age. In other words, a person’s specific position on a personality-dimension continuum relative to the positions of other people stabilizes with age. Stability coeffi- cients (i.e., indices of rank-order continuity between occasions of personality assessment; these coefficients can vary from 0, no continuity, to 1, perfect continuity), which are low in child- hood (approximately .30), rise across adolescence and young adulthood, reaching a plateau (approximately .70) between the ages of 40 and 70 (Roberts & DelVecchio, 2000; Terracciano, Costa, & McCrae, 2006). In old age, stability coefficients appear to decay (Lucas & Donnellan, 2011; Specht, Egloff, & Schmukle, 2011). Irrespective of the age group studied, stabil- ity coefficients decline as the time intervals between measure- ments of personality traits get longer (Fraley & Roberts, 2005). These patterns of stability and instability in individual ranks on personality dimensions point to the fact that personality development is a lifelong individual process. Although these patterns of rank-order continuity in personality are well known, their sources remain a matter of controversy. McCrae and his colleagues (2000) formulated a theory of biological maturation (McCrae & Costa, 2008). According to this perspective, personality traits are stable, genetically (or at least biologically) anchored basic tendencies of thoughts, feel- ings, and behavior. Deviations from perfect continuity should be due to short-term environmentally induced fluctuations, but also to systematic and random measurement error. In fact, con- trolling for artifacts and occasion-specific effects results in higher heritability estimates (i.e., estimates of the degree to which relative differences among individuals are due to genetic factors) and larger stability coefficients (Kandler et al., 2010). Furthermore, similar mean-level trends (i.e., trait change over time averaged across individuals in a sample or population) for neuroticism and for extraversion have been found across different nations and cultures (McCrae et al., 1999, 2000). Neuroticism tends to decrease with age, whereas extraversion tends to be stable. Individual deviations from the mean-level trend (i.e., interindividual differences in intra- individual change) in neuroticism have been found to be genetically influenced (Bleidorn, Kandler, Riemann, Angleit- ner, & Spinath, 2009). These results support the hypothesis that genetically programmed maturation is a propulsive mech- anism of personality development. Consequently, the increase in rank-order continuity of personality from childhood to adulthood may result from individual differences in biological maturation, and the decrease in such continuity in older age may be explained by individual differences in biological degeneration. Other researchers have formulated different perspectives, attributing changes in personality to environmental sources, such as individual requirements in social roles (e.g., worker, spouse, parent) and related investments (Roberts & Wood, 2006), normative life transitions (e.g., departure from one’s parental home, the completion of school or an apprenticeship, retirement), and individual life events that result in major changes of individuals’ circumstances (e.g., accidents, the birth of one’s own child, marriage). In other words, personal- ity development and change are primarily attributable to age- graded social roles and related experiences (Roberts, Wood, & Caspi, 2008). Since the average direction of personality change (i.e., mean-level change in a population) is often posi- tive with respect to social functionality (e.g., people become more emotionally stable with age), personality development is considered to be a process leading to social maturity (Roberts & Mroczek, 2008). From this perspective, the lower stability coefficients in younger ages compared with coefficients in middle adulthood are due to interindividual differences in the life transitions and role transfers that typically occur during the younger period of life, such as the completion of an appren- ticeship and beginning of a job, departure from one’s parental home, marriage, and new parenthood (Arnett, 2000). In fact, this period of life is strongly shaped by individually different developmental courses that are attributable to interindividual differences in level and duration of education, family forma- tion, and economic security (McAdams & Olson, 2010). Social roles and contexts stabilize in midlife. They change again in old age until social adjustment breaks down in advanced old age. Consequently, the increase in rank-order continuity of personality traits from adolescence to adulthood, the plateau in middle adulthood, and the decrease of continuity in older age may result from interindividual differences in sta- bilizing, maintaining, and destabilizing one’s personal identity in life. On the basis of these theoretical considerations, I suggest an integrative perspective. That is, both genetic and environ- mental influences may explain personality development, at least in the case of the rank-order continuity of personality traits, and at least in the case of neuroticism and extraversion. Longitudinal behavioral genetics studies (e.g., longitudinal twin or adoption studies) on different age cohorts have allowed for deeper insight. In the following sections, I briefly review und discuss the results from these studies with respect to the patterns of genetic and environmental effects on rank-order continuity in neuroticism and extraversion. The Role of Genetic Factors in Personality Continuity and Change Using genetically informative designs, the interindividual dif- ferences in trait scores that are due to genetic differences (i.e., heritability) and to environmental influences can be disentan- gled. Genetically informative longitudinal designs allow a dis- entanglement of genetic and environmental components on each of the measurement occasions, as well as the calculation of the level of continuity of each of these components (i.e., genetic and environmental continuity). It has been consistently shown that genetic continuity is far from perfect (i.e., stability coefficients are consistently less than 1) during childhood and adolescence (De Fruyt et al., 2006; Gillespie, Evans, Wright, & Martin, 2004; Spengler,
  • 3. Nature and Nurture of Personality Development 3 Gottschling, & Spinath, 2012). That is, across childhood and adolescence, new genetic factors emerge that contribute to interindividual differences and rank-order changes in person- ality. Like rank-order continuity, genetic continuity tends to increase across adolescence and young adulthood (Hopwood et al., 2011; Viken, Rose, Kaprio, & Koskenvuo, 1994) until it reaches almost perfect stability in middle and late adulthood (W. Johnson, McGue, & Krueger, 2005; Pedersen, 1993; Read, Vogler, Pedersen, & Johansson, 2006). This pattern of genetic continuity has been found for neu- roticism and extraversion or related variables (see green lines in Fig. 1 for neuroticism and in Fig. 2 for extraversion). There- fore, the steady increase in observable rank-order continuity from childhood to adulthood can be accounted for by stabiliz- ing interindividual genetic differences. The relatively high sta- bility of interindividual personality differences in middle adulthood is accompanied by stability in genetic differences. In other words, rank-order continuity in neuroticism and extra- version can be explained by interindividual differences in genetically influenced maturation. The Role of Environmental Factors in Personality Continuity and Change Stable genetic differences among individuals after age 40, however, cannot account for the decay of stability coefficients in older age. Therefore, nongenetic factors may also contribute to rank-order continuity. Across a broad spectrum of geneti- cally informative studies, A. M. Johnson, Vernon, and Feiler (2008) found that the primary nongenetic sources of interindi- vidual differences in neuroticism and extraversion are specific to individuals (i.e., not shared by family members reared together). These effects are known as nonshared environmen- tal effects. From a behavioral genetics perspective, “environ- mental” effects include physiologically and biochemically mediated effects, such as environmentally activated epigenetic influences. Nonshared environmental factors show different degrees of stability (i.e., environmental continuity) with respect to the age studied. Studying twins between the ages of 12 and 16, Gillespie et al. (2004) found rather low coefficients for environmental continuity for both extraversion (.17 for females and .18 for males) and neuroticism (.12 for males and .36 for females). Hopwood and his colleagues (2011) studied the transition from adolescence to adulthood (age 17 to 24). The researchers found environmental-continuity coefficients of .36 for nega- tive emotionality (neuroticism), .37 for communal positive emotionality, and .39 for agentic positive emotionality (the latter two traits can be seen as facets of extraversion). They also investigated rank-order continuity between the ages of 24 and 29. For this interval, they found higher environmental continuity, with coefficients ranging from .56 to .60. For .2 .3 .4 .5 .6 .7 .8 .9 1.0 5 10 15 20 25 30 35 40 45 50 55 60 65 70 75 80 85 90 Age StabilityCoefficient Fig. 1. Genetic continuity (depicted by the green line and data points) and environmental continuity (depicted by the red line and data points) of interindividual differences in neuroticism (or negative emotionality) as a function of age. Higher stability coefficients (i.e., indices of continuity between assessments) indicate greater stability over time. The stability coefficients shown are based on the results of genetically informative longitudinal studies (weighted by sample sizes): Bratko and Butkovic (2007), De Fruyt et al. (2006), Gillespie, Evans, Wright, and Martin (2004), Hopwood et al. (2011), W. Johnson, McGue, and Krueger (2005), Kandler et al. (2010), McGue, Bacon, and Lykken (1993), Pedersen (1993), Read, Vogler, Pedersen, and Johansson (2006), Spengler, Gottschling, and Spinath (2012), Viken, Rose, Kaprio, and Koskenvuo (1994), and Wray, Birley, Sullivan, Visscher, and Martin (2007). Across studies and sample cohorts, intervals between assessments ranged between 2 and 10 years (M = 5.15 years).
  • 4. 4 Kandler middle adulthood, W. Johnson and colleagues (2005) reported 5-year environmental-continuity coefficients over .70. How- ever, other studies have indicated that environmental continu- ity decreases in later adulthood (Pedersen & Reynolds, 1998; Read et al., 2006). These findings and results from other genetically informa- tive studies (see red lines in Figs. 1 and 2) point to the fact that changing levels of rank-order continuity in neuroticism and extraversion are also attributable to changing levels of envi- ronmental continuity across the life span. Moreover, the decrease in rank-order continuity in older age appears to be exclusively due to a decline in the stability of environmental factors. Therefore, interindividual differences in experiences of major life transitions or major life events (whether tempo- ral, quantitative, or qualitative) may contribute to personality change and, thus, to personality development. Environmental contributions to change in interindividual ranks may occur across the entire life span, but they are more pronounced in younger ages (because of events such as leaving one’s parental home, starting a job, and forming a new family) and older ages (because of events such as retirement, illness, death of spouses or friends). Because “environmental” factors also include bio- logically mediated effects, the declining rank-order continuity, especially in later adulthood, might be due to biological causes (e.g., multi-infarct dementia). The Interplay of Genetic and Environmental Factors As life narratives and memories cumulate across the life span, effects of life experiences may cumulate, too. Thus, environ- mental effects on interindividual differences in neuroticism and extraversion may cumulate with age, leading to an increase in environmental variance (McCartney, Harris, & Bernieri, 1990). If interindividual differences due to environmental effects increase with age, stable interindividual differences accounted for by genetic effects should proportionally decrease. Consequently, heritability estimates for personality traits should decrease with age. Most genetically informative studies that have assessed heritability estimates for different age cohorts (e.g., Loehlin & Martin, 2001) have not found significant differences between age groups. However, other studies that have combined cross- sectional data (from different age cohorts) and longitudinal data have suggested that heritability estimates decrease with age (Kandler et al., 2010; Viken et al., 1994). A meta-analytic review of fifteen genetically informative studies on different age groups showed that the level of heritability of neuroticism and extraversion appears to depend on the age studied (see Fig. 3). Heritability of extraversion tends to increase until peo- ple are in their early 40s, but then it declines continuously. For .2 .3 .4 .5 .6 .7 .8 .9 1.0 5 10 15 20 25 30 35 40 45 50 55 60 65 70 75 80 85 90 Age StabilityCoefficient Fig. 2.  Genetic continuity (depicted by the green line and data points) and environmental continuity (depicted by the red line and data points) of interindividual differences in extraversion (or positive emotionality) as a function of age. Higher stability coefficients (i.e., indices of continuity between assessments) indicate greater stability over time. The stability coefficients shown are based on the results of genetically informative longitudinal studies (weighted by sample sizes): Bratko and Butkovic (2007), De Fruyt et al. (2006), Gillespie, Evans, Wright, and Martin (2004), Hopwood et al. (2011), W. Johnson, McGue, and Krueger (2005), Kandler et al. (2010), McGue, Bacon, and Lykken (1993), Pedersen (1993), Read, Vogler, Pedersen, and Johansson (2006), Spengler, Gottschling, and Spinath (2012), and Viken, Rose, Kaprio, and Koskenvuo (1994). Across studies and sample cohorts, intervals between assessments ranged between 2 and 10 years (M = 4.88 years).
  • 5. Nature and Nurture of Personality Development 5 neuroticism, heritability progressively decreases after people enter adulthood. The changing importance of genetic contributions to inter- individual differences in neuroticism and extraversion with age differs from the pattern observed with regard to cognitive abilities. For cognitive abilities, heritability continuously increases with age (W. Johnson, 2010). One reason for this distinction may be that genes and environmental effects dif- ferentially contribute to how a person typically acts (i.e., his or her temperament or narrow-sense personality) and how capa- ble a person is (i.e., his or her abilities or skills). An increase in the level of personality traits’ heritability may reflect the increasing importance of active and reactive gene-environment correlations. People actively select, create, and change their environments (e.g., by changing their friends, residences, or jobs) or evoke social reactions that match their genetically predisposed traits. Environments and social reac- tions, in turn, may affect personality traits. For example, indi- viduals who are genetically predisposed to be more extraverted may perceive life events as more controllable and positive; experiencing life events as controllable and positive, in turn, may augment such people’s levels of extraversion (Kandler, Bleidorn, Riemann, Angleitner, & Spinath, 2012). That is, genetic influences may partly drive the environmental effects accounting for the increasing heritability. The processes involved in these gene-environment correlations may be pro- pulsive mechanisms of the development of cognitive abilities across the life span and the development of extraversion in the first decades of life. The increasing heritability of extraversion until age 30 and the cumulatively decreasing heritability of neuroticism and extraversion after age 30 may accompany midlife tipping points. For example, young adults often try to expand the self and gain new information—they actively try to change their environments to match their goal pursuits; by contrast, older adults more often try to stabilize and protect the quality of their lives and ongoing relationships—they change the self to fit the environment (see McAdams & Olson, 2010, for a review). Consequently, genetic and environmental effects do not act separately; rather, they interact in complex ways that account for the development of personality traits, and the con- tribution of life experiences to personality development cumu- lates with age. Conclusion I hope that this short review will open the door for further research identifying the mechanisms of the gene-environment .05 .15 .25 .35 .45 .55 .65 .75 5 10 15 20 25 30 35 40 45 50 55 60 65 70 75 80 85 90 Age HeritabilityEstimate Fig. 3. Heritability estimates (the proportion of interindividual differences accounted for by genetic differences) for extraversion (depicted by the blue line and data points) and neuroticism (depicted by the orange line and data points) as a function of age. The heritability estimates shown are based on the results of genetically informative longitudinal and age-cohort studies (weighted by sample size): Bratko and Butkovic (2007), De Fruyt et al. (2006), Gillespie, Evans, Wright, and Martin (2004), Hopwood et al. (2011), W. Johnson, McGue, and Krueger (2005), Kandler et al. (2010), Loehlin and Martin (2001), McGue, Bacon, and Lykken (1993), Pedersen and Reynolds (1998), Read, Vogler, Pedersen, and Johansson (2006), Rettew et al. (2006), Rettew, Rebollo- Mesa, Hudziak, Willemsen, and Boomsma (2008), Spengler, Gottschling, and Spinath (2012), Viken, Rose, Kaprio, and Koskenvuo (1994), and Wray et al. (2007).
  • 6. 6 Kandler interplay in personality development and also focusing on the patterns and sources of other personality traits (e.g., openness, agreeableness, and conscientiousness). More genetically and environmentally informative research is sorely needed. At this point, we know that genetic factors stabilize with age, leading to an increase in rank-order continuity of neuroticism and extraversion and to personality formation across childhood, adolescence, and young adulthood. Environmental sources contribute to this increasing continuity but also to decreasing continuity in old age. Moreover, environmental influences appear to cumulate across the life span, leading to a decline of the genetic contribution to variance in neuroticism and extra- version with age, a pattern pointing to lifelong changes in per- sonality due to environmental influences. We develop into the people we are predisposed to become because of our genetic makeup, but to a large degree, the development of or changes in our personality will be formed by the environment. Recommended Reading Bouchard, T. J., & Loehlin, J. C. (2001). (See References). A com- prehensive review of research on personality traits and related variables from a behavioral genetics perspective. Fraley, C., & Roberts, B.W. (2005). (See References). An compre- hensive work on patterns of continuity described as mathematical models. McAdams, D. P., & Olson, B. D. (2010). (See References). A clearly written, user-friendly, and empirically founded theoretical per- spective on personality development of different levels of per- sonality elements, such as traits, motives, and narratives. McCrae, R. R., & Costa, P. T. (2008). (See References). A compre- hensive overview of research and theory on personality traits as biologically anchored basic tendencies. Wright, L. (1997). (See References). An interesting and reader- friendly book on the history of twin studies and the secrets of our identity. Declaration of Conflicting Interests The authors declared that they had no conflicts of interest with respect to their authorship or the publication of this article. References Arnett, J. J. (2000). Emerging adulthood: A theory of development from the late teens through the twenties. American Psychologist, 55, 469–480. Bleidorn, W., Kandler, C., Riemann, R., Angleitner, A., & Spinath, F. (2009). Patterns and sources of adult personality development: Growth curve analyses of the NEO-PI-R scales in a longitudinal twin study. Journal of Personality and Social Psychology, 97, 142–155. Bouchard, T. J., & Loehlin, J. C. (2001). Genes, evolution, and per- sonality. Behavior Genetics, 31, 243–273. Bratko, D., & Butkovic, A. (2007). Stability and genetic and environ- mental effects from adolescence to young adulthood: Results of Croatian longitudinal twin study of personality. 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