Personality and Individual Differences 77 (2015) 186192

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Personality and Individual Differences

journal homepage: www.elsevier.com/locate/paid

Review

The extraversion continuum in evolutionary perspective: A review of

recent theory and evidence
Aaron W. Lukaszewski a,, Christopher R. von Rueden b
a
Department of Psychology, Oklahoma State University, Stillwater, OK 74074, United States
b
Jepson School of Leadership Studies, University of Richmond, Richmond, VA 23173, United States

a r t i c l e i n f o a b s t r a c t

Article history: We review research on the ultimate and proximate origins of variation along the extraversion continuum.
Received 30 September 2014 After describing the cost-benet tradeoffs that may have maintained variation in extraversion over
Received in revised form 31 December 2014 human evolution, we consider the evidence bearing on multiple distinct evolutionary hypotheses regard-
Accepted 3 January 2015
ing the causal underpinnings of such variation. On the basis of the reviewed evidence, we argue that uc-
Available online 19 January 2015
tuating selection on specic polymorphic genotypes is unlikely to explain the origins of individual
differences in extraversion. Rather, adaptively patterned variation in extraversion is likely orchestrated
Keywords:
primarily by facultative adaptations designed to calibrate behavioral strategies to cues available in ontog-
Behavior genetics
Evolutionary genetics
eny. For example, emerging research supports the hypothesis that extraversion may be reactively her-
Evolutionary psychology itable by virtue of its calibration to heritable condition-dependent phenotypic features which in turn
Extraversion helps explain extraversions genetic variance, as well as its consistent positive association with reproduc-
Facultative calibration tive success. Finally, evidence suggests that some of the inter-individual variance in extraversion is fun-
Personality damentally noisy, arising as a side effect of mutationselection balance or pleiotropic polymorphisms
Reactive heritability maintained via pathogenhost coevolution. If correct, these conclusions indicate that future research
should focus on elucidating the facultative adaptations designed to regulate the production of behaviors
falling on the extraversion continuum.
2015 Elsevier Ltd. All rights reserved.

1. Introduction Natural selection can theoretically maintain personality varia-

The extraversion continuum is among the aspects of human
personality that have recently been analyzed in evolutionary per-
spective (see Buss & Hawley, 2011; Nettle, 2006; Penke,
Denissen, & Miller, 2007). At the level of phenotypic description,
extraversion is one of several basic higher-order axes of personality
variation, encompassing correlated behavioral facets of sociability,
expressiveness, assertiveness, and status motivation (Ashton & Lee,
2007; Ashton, Lee, & Paunonen, 2002; Denissen & Penke, 2008;
John, Naumann, & Soto, 2008; McCrae & Costa, 2008; Wilt &
Revelle, 2008). In evolutionary perspective, the mystery of extra-
versions origins boils down to the question of why natural selec-
tion has maintained variability along this continuum, rather than
eliminating variation in favor of xed genotypes or behavioral phe-
notypes with the highest net tness (Lukaszewski, 2013;
Lukaszewski & Roney, 2011; Nettle, 2005, 2006, 2011; Verweij
et al., 2012).
Corresponding author at: 116 N. Murray, Oklahoma State University, Stillwater,
OK 74074, United States. Tel.: +1 (405) 744 4277.
E-mail address: aaron.lukaszewski@okstate.edu (A.W. Lukaszewski).
tion via uctuating (including frequency-dependent) selection
pressures that modulate the tness costs and benets of different
personality trait levels (Buss, 2009; Gangestad & Simpson, 2000;
Lewis, 2014; Lukaszewski & Roney, 2011; Nettle, 2006; Penke
et al., 2007; Tooby & Cosmides, 1990). This theoretical perspective
leads to the expectation that personality variation will often be
adaptively patterned in relation to circumstances that predicted
optimal trait levels under ancestral conditions. For example, differ-
ent levels of extraversion may have been optimal for human ances-
tors who varied in their phenotypic features (e.g., physical traits) or
who encountered different socioecological conditions (e.g., levels
of disease prevalence).
There are two broad classes of proximate mechanisms that can
generate personality variation that is adaptively patterned in rela-
tion to individual circumstance. First, uctuating (i.e. balancing)
selection regimes can theoretically maintain specic genetic poly-
morphisms with reliable organizational inuences on the endophe-
notypes (i.e. neural or endocrine substrates) of a behavioral
phenotype (Buss, 2009; Gangestad, 2011; Lukaszewski & Roney,
2011; Penke et al., 2007; Tooby & Cosmides, 1990; Verweij et al.,
2012). Second, uctuating selection pressures can craft universal

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0191-8869/ 2015 Elsevier Ltd. All rights reserved.
 A.W. Lukaszewski, C.R. von Rueden / Personality and Individual Differences 77 (2015) 186192
adaptations that facultatively calibrate personality phenotypes in
response to cues that predicted optimal trait levels under ancestral
conditions (Buss, 2009; Keller & Miller, 2006; Lewis, 2014;
Lukaszewski, 2013; Lukaszewski & Roney, 2011; Nettle, 2011;
Penke, 2011; Sell, Tooby, & Cosmides, 2009; Tooby & Cosmides,
1990).
Whereas specic genetic polymorphisms and facultative adap-
tations, respectively, each produce personality variation that is
adaptively patterned, the evolutionary process can also yield indi-
vidual differences that are fundamentally noisy i.e. randomly
patterned about an optimal level (Tooby & Cosmides, 1990). Thus,
theories of adaptively patterned individual differences must
always be discriminated from the alternative hypothesis that
observed phenotypic variation is patterned randomly.
This article provides a brief review of evidence that bears on
these evolutionary hypotheses regarding the origins of extraver-
sion in humans. We begin by explaining the reproductive cost-ben-
et tradeoffs that likely maintained variation along the
extraversion continuum over human evolution. Next, we consider
data bearing on the hypothesis that individual differences in extra-
version arise from direct organizational effects of polymorphic
genotypes. Thereafter, we review evidence in support of the
hypothesis that variation in extraversion reects the existence of
facultative adaptations designed to calibrate extraversion levels
in response to cues available in ontogeny. Related to this latter
type of model, we then evaluate the reactive heritability hypoth-
esis, which posits that the genetic variance in extraversion is par-
tially explained by its facultative calibration to variations in
condition-dependent phenotypic features. On the basis of the evi-
dence we review, we argue that adaptively patterned variation in
extraversion is likely generated primarily by facultative adapta-
tions, and that some of extraversions inter-individual variance is
probably fundamentally noisy.
2. Cost-benet tradeoffs along the extraversion continuum
The extraversion continuum appears to represent variability in
a coordinated cluster of phenotypic strategies related to social
exchange and hierarchy negotiation (Denissen & Penke, 2008;
Lukaszewski, 2013; Lukaszewski & Roney, 2011; Nettle, 2011;
von Rueden, Lukaszewski, & Gurven, submitted). Recent theories
suggest that variation in extraversion may have been maintained
over human evolution by reproductive cost-benet tradeoffs that
occur at different points along its continuum (Ashton & Lee,
2007; Lukaszewski, 2013; Lukaszewski & Roney, 2011; Nettle,
2005, 2006, 2011).
By denition, highly sociable people tend to participate in social
exchange with a relatively large number of cooperative partners
and groups, whereas people low in sociability are more inclined
to invest in a smaller number of deep engagement relationships
with close relationship partners, as well as solitary activities
(Ashton & Lee, 2007; John et al., 2008). On the benet side, main-
taining a large cooperative network proportionally increases
potential gains in trade via dyadic exchange and collective action
(Lukaszewski, 2013). On the cost side, however, socializing widely
with non-kin opens an individual up to potential exploitation by
cheaters and free riders (Price, Cosmides, & Tooby, 2002), increases
exposure to pathogen vectors (Schaller & Murray, 2008), and takes
up time, energy, and other resources that could be invested in close
relationships or other tness-relevant tasks (e.g., self-maintenance,
skill mastery, parenting) (Ashton & Lee, 2007). Given these trade-
offs, the optimal level of sociability would have varied for human
ancestors along with circumstances that either buffered against
the potential costs, or increased the potential benets, of partici-
pating in social exchange with a large network of cooperative
187
partners. For example, the optimal level of sociability would be
higher for an individual who is relatively unlikely to be exploited
by others (Sell et al., 2009), able to easily attract high quality asso-
ciates (Lukaszewski & Roney, 2011), or unlikely to contract com-
municable illnesses (Schaller & Murray, 2008).
Another facet of the extraversion continuum is the motivation
to attract social attention and assertively compete for positions
of high social status and leadership (Anderson, John, & Keltner,
2012; Anderson, John, Keltner, & Kring, 2001; Ashton & Lee,
2007; Ashton et al., 2002). Recent evolutionary theories to explain
the functional dynamics of human status hierarchies are based on
the recognition that multi-individual cooperative exchanges carry
massive potential benets for all participants, but are undermined
by a number of problems related to the complexity of coordination
and the existence of free riders and rule violators (Anderson &
Kilduff, 2009; Hooper, Kaplan, & Boone, 2010; Price & Van Vugt,
2014; Tooby, Cosmides, & Price, 2006; von Rueden, Gurven,
Kaplan, & Stieglitz, 2014). Within collective actions and coalitions,
leaders are individuals who help solve these logistical problems by,
for example, dening collective goals, delegating tasks to others,
administering collectively sanctioned punishment of free riders,
limiting conict within the group, or enforcing a fair allocation of
resources (Price, 2014; von Rueden et al., 2014). By paying the per-
sonal costs of solving these problems for collective actions and
coalitions, leaders increase the gains in trade experienced by all
participants (von Rueden et al., 2014). In return, followers compen-
sate leaders with respect, deference, or a greater share of the spoils
(Price, 2014; von Rueden et al., 2014).
From this perspective, aspects of extraversion that pertain to
individual differences in status motivation (e.g., assertiveness;
social boldness; desire for attention) can be conceptualized along
a leadership-followership gradient. Leadership-oriented individu-
als must pay the attendant costs of acquiring and maintaining their
inuential positions; but they may also, when successful, enjoy the
benets of high status and respect (Price, 2014; Van Vugt, Hogan, &
Kaiser, 2008; von Rueden, Gurven, & Kaplan, 2011). Symmetrically,
followers of (effective) leaders benet from participation in suc-
cessful collective actions and coalitions, but do not pay the costs
inherent in competing for and implementing leadership. It follows
from these cost-benet tradeoffs that the optimal level of status
motivation would have varied for human ancestors along with cir-
cumstances that altered the potential benets of leadership and
followership, for example, the possession of characteristics that
determine leadership ability (Lukaszewski, 2013; Lukaszewski,
Simmons, Anderson, & Roney, submitted; Nettle, 2011; Price,
2014; von Rueden et al., 2014) or the ratio of leadership-oriented
individuals to followers in the local social world (Van Vugt et al.,
2008).
Phenotypic strategies related to cooperation and hierarchy are
also functionally relevant for the formation of mating relation-
ships. First, extraverted behavior involves proactively approaching
and attracting the attention of others, which likely facilitates the
initiation of mating relationships, all else equal (Ashton & Lee,
2007; Lukaszewski & Roney, 2011; Nettle, 2005). In addition, mem-
bers of both sexes (especially women) exhibit a preference for
resourceful, productive, and high status mates (Buss & Schmitt,
1993). Therefore, extraverted strategies, when successful, can indi-
rectly enhance ones mate value via the acquisition of resources
and social status (Alvergne, Jokela, & Lummaa, 2010; Campbell,
Simpson, Stewart, & Manning, 2003; Nettle, 2005; von Rueden,
Gurven, & Kaplan, 2008; von Rueden et al., 2011). However, com-
peting for access to mates exposes an individual to potential con-
ict with rivals (Puts, 2010; Simpson, Gangestad, Christensen, &
Leck, 1999), and mating with multiple partners is associated with
various costs, including exposure to sexually transmitted disease
and the resource burden of multiple dependent offspring
188 A.W. Lukaszewski, C.R. von Rueden / Personality and Individual Differences 77 (2015) 186192
(Gangestad & Simpson, 2000; Nettle, 2005; Winking, Kaplan,
Gurven, & Rucas, 2007).
Evidence supports the existence of these sorts of cost-benet
tradeoffs along extraversions continuum. For example, relative to
introverts, highly extraverted individuals have larger cooperative
networks (Lee, Dean, & Jung, 2008), are more successful on the
mating market (Nettle, 2005), and are more likely to attain posi-
tions of high social status and leadership (Alvergne et al., 2010;
Anderson et al., 2001; Campbell et al., 2003; Lund, Tamnes,
Mouestue, Buss, & Vollrath, 2007). However, extraverts are also
more likely than introverts to experience antagonistic conict in
competing for social position (Anderson & Shirako, 2008; Lund
et al., 2007), spend limited time and energy socializing (Gurven,
von Rueden, Stieglitz, Kaplan, & Eid-Rodriguez, 2014), contract ill-
nesses (Christakis & Fowler, 2010), and sustain injuries (Field &
OKeefe, 2004).
Although the potential benets of a highly extraverted strategy
are weighed against the potential costs, evidence nonetheless indi-
cates that extraversion positively predicts reproductive success
within societies. Among men, greater manifest extraversion is con-
sistently positively correlated with reproductive success within
both small-scale subsistence societies (Alvergne et al., 2010;
Bailey et al., 2013; Gurven et al., 2014) and modern industrialized
societies (Berg, Lummaa, Lahdenpera, Rotkirch, & Jokela, 2014;
Jokela, 2012). Among women, greater extraversion is less strongly
associated with reproductive success within small-scale societies
(Alvergne et al., 2010; Bailey et al., 2013; Gurven et al., 2014),
but has shown positive association with reproduction in modern
societies (Berg et al., 2014; Jokela, 2012).
In light of the above considerations, a complete theory of extra-
versions origins will explain (i) whether and how individuals
manifest personality strategies are adaptively patterned in relation
to variable circumstances, and (ii) why natural selection has main-
tained variation in extraversion despite its consistent positive
association with reproductive success.
3. Polymorphic genotypes as direct causes of individual
differences in extraversion
Extraversion, like all other aspects of personality that have ever
been studied by quantitative geneticists, exhibits substantial heri-
tability (Bouchard & Loehlin, 2001). In general, heritable variance
in extraversion could be systematically maintained by uctuating
selection pressures on genotype ? personality linkages (Del
Giudice, 2012; Gangestad, 2011; Gurven et al., 2014;
Lukaszewski & Roney, 2011; Nettle, 2005; Penke et al., 2007;
Tooby & Cosmides, 1990; Verweij et al., 2012), or it could arise
as a noisy side effect of pleiotropic polymorphisms maintained in
the context of mutationselection balance or pathogenhost
coevolution (Keller & Miller, 2006; Tooby & Cosmides, 1990).
3.1. Specic polymorphic genes as organizers of adaptively patterned
variation in extraversion
In attempting to explain the maintenance of polymorphic geno-
types for adaptively patterned variation in extraversion that
which is systematically coordinated with circumstances that pre-
dict optimal trait levels uctuating selection is the only viable
type of explanation (Penke et al., 2007). A necessary precondition
for the maintenance of adaptively patterned genotype ? personal-
ity effects via uctuating selection is the existence of common
polymorphic genotypes that organize the endophenotypes of per-
sonality (e.g., neural mechanisms) in consistent ways; in short,
there would need to be alternative alleles that reliably contribute
to higher or lower levels of extraversion (Keller & Miller, 2006;
Lukaszewski & Roney, 2011). Once such genotype ? personality
linkages existed, then selection could maintain non-zero frequen-
cies of those genotypes to the extent that optimal levels of extra-
version uctuate across individuals as a function of sex, time,
space, or the composition of the local social world (Penke et al.,
2007; Tooby & Cosmides, 1990; Verweij et al., 2012).
It is problematic for this type of evolutionary genetic theory
that there is little evidence supporting the existence of common
alleles (present at >.01 frequency in a population) that reliably
explain even a tiny fraction of the variance in extraversion or any
other aspect of personality (De Moor, Costa, Terracciano, Krueger,
& Boomsma, 2012; Johnson, Penke, & Spinath, 2011; Service,
Verweij, Lahti, Congdon, & Freimer, 2012; Verweij et al., 2012;
Vinkhuyzen, Pedersen, Yang, Lee, & Wray, 2012). In recent decades,
candidate gene association studies have sometimes reported links
between specic genotypes and personality trait levels in relatively
small samples (e.g., Canli, 2008; Lukaszewski & Roney, 2011;
Wacker, Mueller, Hennig, & Stemmler, 2012). However, such seem-
ingly promising effects have often failed to replicate (Johnson et al.,
2011). One example is provided by the nding that a single nucle-
otide polymorphism (SNP) in the catechol-O-methyltransferase
(COMT) gene predicts extraversion trait levels (Wacker et al.,
2012). Because COMT is an enzyme that inuences dopamine
metabolism, this gene-personality linkage makes sense on the the-
ory that mesolimbic dopaminergic activity is one of extraversions
neural endophenotypes (Canli, 2008). However, genome-wide
association studies (GWAS) with immense statistical power to
detect even very small genetic effects have found no evidence for
an association between the COMT gene polymorphism and extra-
version (De Moor et al., 2012; Service et al., 2012; Verweij,
Zeitsch, Medland, Gordon, & Wray, 2010).
Moreover, these same GWAS, which together include more than
20,000 subjects, indicate that that are no specic genotypes that
explain a signicant fraction of the variance in extraversion-related
dimensions (De Moor et al., 2012; Service et al., 2012; Verweij
et al., 2010). Even when hundreds of thousands of common SNPs
across the genome are entered as simultaneous predictors, com-
bined effects of these genetic variants account for only 10% of
extraversions variance (Verweij et al., 2012; Vinkhuyzen et al.,
2012). By contrast, (i) studies with similar statistical power have
identied multiple individual common SNPs that reliably predict
human height and (ii) combined effects of common SNPs across
the genome explain nearly half of heights total phenotypic vari-
ance (Yang, Benyamin, McEvoy, Gordon, & Visscher, 2010). Thus,
although effects of specic genetic variants on complex and highly
polygenic trait dimensions can be identied with extant methods,
such effects appear to be of no more than minimal importance in
accounting for the causal underpinnings of individual differences
in extraversion.
In sum, despite some promising ndings in studies using small
samples, the overall weight of the extant evidence does not sup-
port the hypothesis that extraversion levels are organized via
direct effects of polymorphic genes on the neural endophenotypes
of extraverted behavior. In turn, this conclusion suggests that
adaptively patterned variation in extraversion is not likely to have
been maintained by uctuating selection on specic polymorphic
genotypes (Miller, 2011; Verweij et al., 2012).
3.2. Genetic variation as a cause of noisy individual differences in
extraversion
Heritable individual differences in extraversion could also arise
as a noisy side effect of genetic polymorphisms that exist for rea-
sons other than their effects on personality trait levels. First, muta-
tions are constantly introduced into reproducing populations
before they can be removed by selection, and individuals therefore
 A.W. Lukaszewski, C.R. von Rueden / Personality and Individual Differences 77 (2015) 186192
differ in their overall load of mildly deleterious, low-frequency
mutations (Keller & Miller, 2006; Penke et al., 2007; Tooby &
Cosmides, 1990). Importantly, because mutations are copying
errors, their direct organizational effects on personality variation
will be directionally random such that a given mutated allele is
equally likely to organize the individual toward higher or lower
levels of extraversion (Lewis, 2014; Lukaszewski & Roney, 2011;
Penke et al., 2007).
Evidence from large gene association studies is consistent with
the hypothesis that individual differences in extraversion reect
noisy genetic variation arising via mutation. Specically, Verweij
et al. (2012) argued that, because their data indicate that all com-
mon genetic variants collectively explain only a small percentage
of the heritable variance in extraversion, the remaining genetic
variance is probably explained by very rare alleles (i.e. likely recent
mutations). This nding is thus consistent with the view that noisy
variation in extraversion is maintained under mutationselection
balance (Penke et al., 2007; Verweij et al., 2012).
Noisy individual differences in extraversion could also reect
genetic variation being maintained in the context of coevolution-
ary arms races between rapidly-evolving pathogens and their hosts
(Lukaszewski & Roney, 2011; Tooby & Cosmides, 1990; Verweij
et al., 2012). Because pathogens are constantly being selected to
better exploit host tissues, the genotypes that predict pathogen
resistance change across generational time, which in turn leads
to the perpetual maintenance of polymorphisms at gene loci that
inuence pathogen defense (Piertney & Oliver, 2006; Prugnolle,
Manica, Charpientiere, Guegan, & Balloux, 2005). To the degree
that the genes involved in pathogen resistance also have pleiotro-
pic effects on neurotransmitter systems, etc., some heritable varia-
tion in personality traits such as extraversion may arise as a noisy
side-effect of genetic polymorphisms maintained in the context of
pathogenhost coevolution. Like mutations, polymorphic alleles
under pathogen-related selection pressures should have direction-
ally random organizational effects on the endophenotypes of
personality.
In evaluating this hypothesis, it is instructive to reconsider the
evidence discussed above that, although no individual common
genotypes explain a signicant fraction of the variance in extraver-
sion (De Moor et al., 2012; Service et al., 2012; Verweij et al., 2010),
all common polymorphic genotypes together explain 10% of the
phenotypic variance (Verweij et al., 2012; Vinkhuyzen et al.,
2012). These ndings indicate that any direct organizational effects
of specic polymorphic gene loci on extraversion levels that do
exist are so innitesimally small as to evade detection. This is
exactly what might be expected if those polymorphisms were
being evolutionarily maintained for a different functional reason
than their inuence on personality trait levels such as their
effects on pathogen resistance (Tooby & Cosmides, 1990). For
example, if (i) there are alleles at polymorphic loci which oscillate
in frequency between .01 and .50 within pathogenhost coevolu-
tionary arms races, and (ii) these genotypes also have small pleio-
tropic effects on neural or endocrine mechanisms that regulate
social behavior, then (iii) such pathogen-driven polymorphisms
will likely result in noisy inter-individual variation in manifest per-
sonality. Empirically, this hypothesis predicts that common vari-
ants with (extremely small) effects on extraversion levels will
tend to be at gene loci that encode for phenotypic attributes
involved in pathogen-resistance (e.g., immune function; biochem-
ical surface markers).
In sum, the current state of extant evidence suggests that noisy
effects of rare genotypes on extraversion levels could be main-
tained under mutationselection balance. In addition, although
the possible direct effects of common genotypes on extraversion
levels are too small to detect in high-powered studies, any such
effects that do exist could plausibly arise as a noisy side-effect of
189
genetic variation being maintained in the context of pathogen
host coevolution. As evidence accumulates regarding the genomic
regions that encode for particular phenotypes, these related
genetic noise hypotheses will become increasingly testable.
4. Facultative adaptations as calibrators of adaptively patterned
variation in extraversion
Fluctuating selection remains a viable explanation for adap-
tively patterned variation in extraversion when we consider per-
sonality as the variable output of species-typical adaptations for
behavior regulation (Buss, 2009; Gangestad & Simpson, 2000;
Lewis, 2014; Lukaszewski & Roney, 2011; Penke, 2011; Tooby &
Cosmides, 1990). In essence, adaptationist models view variation
in manifest behavioral outputs as arising from evolved conditional
rules of the form, Given condition x, pursue behavioral strategy y0 .
Theoretically, such facultative adaptations for personality calibra-
tion would have been selected for over human evolutionary history
as they improved the functional match between behavioral strate-
gies and individual circumstance. For example, the optimal level
along a given personality dimension might have differed across
ancestral humans who varied in their phenotypic features (e.g.,
physical traits) or who encountered different socioecological con-
ditions (e.g., the prevalence of trustworthy cooperative partners).
If so, such uctuating selection regimes may have crafted adapta-
tions that calibrate manifest personality levels to cues that pre-
dicted these variable circumstances under ancestral conditions.
4.1. Condition-dependent calibration of extraversion
A recent adaptationist model posits that extraversion levels are
facultatively calibrated to variations in phenotypic features whose
expression depends on the overall phenotypic condition of the
organism (Lukaszewski, 2013; Lukaszewski & Roney, 2011; von
Rueden et al., submitted). Phenotypic condition (also referred to as
quality) refers to an individuals ability to efciently convert energy
into tness-enhancing traits and outcomes (Gangestad, Merriman,
& Thompson, 2010; Lewis, 2014; Lukaszewski, Larson,
Gildersleeve, Roney, & Haselton, 2014; Tomkins, Radwan,
Kotiaho, & Tregenza, 2004). Individual differences in condition
are determined by a variety of converging factors, including gen-
ome-wide mutation load, the adaptedness of ones genotypes to
the current coevolutionary state of local pathogens, components
of environmental quality (e.g., nutrition), and exposure to cues that
calibrate an individual toward investment in future (vs. present)
reproduction (Gangestad et al., 2010; Keller & Miller, 2006;
Penke et al., 2007; Tomkins et al., 2004; von Rueden et al.,
submitted). By denition, individuals in better phenotypic condi-
tion, relative to those in poorer condition, need to invest less of
their effective energy budgets into maintenance effort (e.g., operat-
ing the immune system) to maintain adequate health, and will
therefore have larger residual energy budgets to invest in the cul-
tivation of traits that enhance future reproduction; for example,
sexually attractive ornaments, energetically expensive muscle tis-
sue, or the neural tissue required for the embodiment of knowl-
edge and skill.
One of the primary reasons that condition-dependent features
would have ancestrally predicted optimal levels of extraversion
is that such features inuence an individuals relative bargaining
power, which is dened as a joint function of ones ability to confer
benets upon others and to inict costs on others (Lukaszewski,
2013; Sell et al., 2009). For example, physical attractiveness assess-
ments are based on ancestrally valid cues to an individuals health,
fertility or formidability, and physically attractive individuals are
therefore preferred as friends, mates, allies, and leaders
190 A.W. Lukaszewski, C.R. von Rueden / Personality and Individual Differences 77 (2015) 186192
(Anderson et al., 2001; Frederick & Haselton, 2007; Sprecher &
Regan, 2002). According to similar logic, the possession of knowl-
edge, skill, or ability that increases ones ability to generate bene-
ts for others is seen as an indispensible characteristic of high
quality leaders (Anderson & Kilduff, 2009; von Rueden, 2014). Like-
wise, physical formidability is a direct determinant of ones ability
to inict costs on others, and formidable men are correspondingly
shown deference when interests conict (von Rueden et al., 2008),
preferred as relationship partners (Coy, Green, & Price, 2014;
Lukaszewski, 2013), and allocated greater social status
(Lukaszewski et al., submitted; von Rueden et al., 2008; von
Rueden et al., 2014). In short, condition-dependent phenotypic fea-
tures such as physical attractiveness, intelligence, and formidabil-
ity jointly determine overall bargaining power because of their
inuence on others perceptions of ones ability to confer benets
or inict costs.
Because of these widespread effects on others social percep-
tions and affordances, individuals with a greater bargaining power
will be differentially likely to experience the potential benets of
an extraverted behavioral strategy. For example, individuals who
are simultaneously attractive, intelligent, and formidable will tend
to experience a relatively high rate of return on their attempts to
initiate relationships and ascend hierarchies. Additionally, such
individuals will be less likely to experience the potential costs of
an extraverted strategy, including conict with rivals or opportu-
nity costs of unsuccessful investment in courtship, social exchange,
or status pursuit. For these reasons, an individuals relative bar-
gaining power would have positively predicted optimal levels of
extraversion under ancestral conditions (see Lukaszewski, 2013;
Lukaszewski & Roney, 2011; von Rueden et al., submitted).
Evidence supports the hypothesis that extraversion levels are
facultatively calibrated to condition-dependent phenotypic fea-
tures that inuence an individuals bargaining power. For example,
measures of physical attractiveness and physical formidability
have been found to positively predict levels of manifest extraver-
sion among both among citizens of modern societies (Bourdage,
Lee, Ashton, & Perry, 2007; Fink, Neave, Manning, & Grammer,
2005; Lukaszewski, 2013; Lukaszewski & Roney, 2011; Penton-
Voak, Pound, Little, & Perrett, 2006; Pound, Penton-Voak, &
Brown, 2007) and Amazonian hunter-horticulturalists (von
Rueden et al., submitted). In addition, such associations have
obtained when attractiveness and strength were operationalized
via self-perceptive measures (Bourdage et al., 2007; Lukaszewski,
2013; Lukaszewski & Roney, 2011) or more objective measures
such as third party ratings of attractiveness (Loehlin, 2014;
Lukaszewski, 2013; Lukaszewski & Roney, 2011) and measured
upper-body strength (Lukaszewski, 2013; Lukaszewski & Roney,
2011; von Rueden et al., submitted).
4.1.1. The reactive heritability hypothesis
This condition-dependent model of extraversions ontogenetic
calibration has implications for explaining the heritability of this
personality dimension. Overall phenotypic condition is always
under positive directional selection but is highly polygenic and
therefore has a large mutational target size (Penke et al., 2007;
Tomkins et al., 2004). Thus, the substantial heritability of condition
itself largely reects effects of alleles maintained in the context of
mutationselection balance and pathogenhost coevolution, such
that individuals in better condition will tend to have low mutation
loads and genotypes that are well adapted to the current coevolu-
tionary state of local pathogens (Keller & Miller, 2006; Penke et al.,
2007; Prugnolle et al., 2005; Tomkins et al., 2004).
Crucially, if extraversion levels are calibrated to highly heritable
condition-dependent features, it follows that extraversion will also
substantial exhibit heritability even if, as appears to be the case,
there are no specic polymorphic genotypes that organize the
endophenotypes of personality in reliable ways (Lewis, 2014;
Lukaszewski, 2013; Lukaszewski & Roney, 2011). In general, this
phenomenon, wherein a personality dimension is heritable by vir-
tue of its calibration to other heritable phenotypic features, has
been referred to as reactive heritability (Lukaszewski & Roney,
2011; Tooby & Cosmides, 1990).
Emerging research supports the hypothesis that extraversion is
reactively heritable in relation to condition-dependent phenotypic
features. For example, von Rueden et al. (submitted) recently
employed pedigrees to investigate the heritability of multiple phe-
notypes among Amazonian hunter-horticulturalists (see also
Gurven et al., 2014), and found that the estimated heritability of
extraversion (and related prosocial traits) was signicantly
reduced when controlling for condition-dependent features,
including physical strength. Thus, whereas previous ndings link-
ing extraversion to condition-dependent physical features were
merely suggestive, this genetically informative study provides
the rst direct evidence supporting extraversions potential reac-
tive heritability.
This evidence from the Amazon is bolstered by the recent nd-
ing that genetic markers of individuals overall mutation loads are
negatively predictive of both condition-dependent physical fea-
tures such as physical strength (Verweij, Abdellaoui, Veijola,
Sebert, & Zeitsch, 2014) and extraversion-related personality traits
(Verweij et al., 2012) which is exactly what should be expected
on the theory that extraversion is reactively heritable in relation
to condition-dependent features (Lewis, 2014; Lukaszewski &
Roney, 2011; Verweij et al., 2012). In this context, it is important
to note the contrast of these ndings with the predictions of the
genetic noise hypothesis described above. Specically, if muta-
tions explain genetic variance in personality because they have
direct organizational effects on extraversions endophenotypes,
any given mutation should be equally likely to produce higher or
lower manifest trait levels (see Section 3.2). Thus, the fact that
individual differences in mutation load are directionally correlated
with condition-dependent features and extraversion levels is pre-
dicted by the reactive heritability model, but not by the hypothesis
that mutations produce randomly patterned phenotypic variation.
Hypotheses of condition-dependent calibration and genetic
noise are not mutually exclusive. For example, facultative calibra-
tion of extraversion levels to condition-dependent features could
explain (indirect) effects of expressed mutations with a deleterious
impact on phenotypic condition, but which have no direct organi-
zational effects on extraversions endophenotypes. Meanwhile,
there could be other expressed mutations that, independent of
their effects on phenotypic condition, have noisy direct effects on
the neurocognitive mechanisms involved in the production of
extraverted behavior. Importantly, in the rst example, expressed
mutations ultimately become correlated with adaptively patterned
variation in extraversion (via condition-dependent calibration),
whereas the second example describes a scenario wherein muta-
tions have directionally random inuences on extraversion levels
(via direct effects on endophenotypes). Consistent with both types
of effects occurring in parallel, Verweij et al. (2012) found that
markers of mutation load were negatively predictive of extraver-
sion, but that these directional effects only explained part of the
heritable variance that is likely attributable to the inuence of
mutations.
Of the hypotheses evaluated above, only the condition-depen-
dent calibration model readily explains how natural selection can
systematically maintain heritable variation in extraversion within
human populations, despite its consistent positive association with
reproductive success. Overall phenotypic condition, which is
always under positive directional selection, remains heritable in
perpetuity due to the inexorable countervailing forces of muta-
tionselection balance and pathogenhost coevolution. In effect,
 A.W. Lukaszewski, C.R. von Rueden / Personality and Individual Differences 77 (2015) 186192
then, adaptations for condition-dependent personality calibration
provide a solution to the adaptive problem created by the perpet-
ual maintenance of heritable variance in condition.
A basic but heretofore untested prediction of this model is that
individuals whose extraversion levels are miscalibrated in rela-
tion to condition-dependent components of RBP will experience
lower reproductive success than individuals who are properly cal-
ibrated, all else equal. For example, among people who are simul-
taneously very low on physical strength, attractiveness, and
intelligence relative to others in their local community, manifest
extraversion levels should be negatively correlated with tness-
related outcomes (assuming that a sufcient number of such mis-
calibrated individuals exist).
5. Conclusions
We have provided a brief overview of recent research that bears
on distinct evolutionary hypotheses to explain the origins of indi-
vidual differences in extraversion. The evidence discussed above
supports several tentative conclusions:
Adaptively patterned variation in extraversion does not appear
to reect the existence of specic polymorphic genotypes with
direct organizational effects on extraversions endophenotypes.
Therefore, adaptively patterned variation in extraversion is
likely orchestrated primarily by facultative adaptations
designed to calibrate behavioral strategies in response to cues
available in ontogeny.
In particular, emerging evidence supports the hypothesis that
extraversion may be facultatively calibrated to condition-
dependent phenotypic features which in turn helps explain
extraversions genetic variance, as well as its consistent positive
association with reproductive success within human societies.
Some heritable inter-individual variance in extraversion is
likely fundamentally noisy, reecting directionally random
organizational effects of low frequency mutations or pleiotropic
polymorphisms being maintained in the context of pathogen
host coevolution.
At this point in time, the conclusion that adaptively patterned
variation in extraversion must arise primarily from facultative
adaptations is based largely on the absence of specic geno-
type ? personality effects. However, this conclusion makes good
theoretical sense. This is because facultative mechanisms can be
expected to produce a much better functional match between
behavioral strategies and individual circumstances than specic
genotype ? personality effects, given that the latter may often
commit individuals to the expression of trait levels that are mis-
matched with their eventual circumstances (Keller & Miller,
2006; Lewis, 2014; Lukaszewski & Roney, 2011). Moreover, the
behavioral plasticity enabled by calibrational mechanisms is con-
sonant with recent ndings regarding the limited within-person
stability of personality. For example, longitudinal studies indicate
that individuals manifest extraversion levels exhibit substantial
variability across situations (Fleeson, 2001; Fleeson, 2007; Wood
& Roberts, 2006) and over development (Roberts & DelVecchio,
2000; Wood & Roberts, 2006). These ndings are bolstered by
experimental work demonstrating that even brief exposures to sit-
uational cues in the laboratory can lead to immediate recalibra-
tional effects on manifest extraversion levels (Mortensen, Becker,
Ackerman, Neuberg, & Kenrick, 2010). Thus, although extraversion
is often characterized as an inherited trait-like construct that var-
ies in dispositional strength between individuals (McCrae & Costa,
2008), it may be more valid to view it as a continuum of manifest
behavioral outputs whose expression is regulated exibly across
development and immediate situations (Fleeson, 2007;
191
Lukaszewski & Roney, 2011). If correct, these arguments suggest
that future research will be most successful in identifying specic
causes of adaptively patterned variation in manifest extraversion if
it is focused on mapping the species-typical adaptations designed
to regulate the production of behaviors falling on the extraversion
continuum. We hope this article helps stimulate such research.
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