Temperament and Evolution

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Temperament and Evolution

Kevin MacDonald

Department of Psychology

California State University–Long Beach1250 Bellflower Blvd

Long Beach, CA 90840-0901

Phone: 562 985-8183; Fax: 562 985-8183

Email: [email protected]

In Marcel Zentner (Ed.), Handbook of Temperament . New York: Guilford Press


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Introduction

Evolution is now a mainstream paradigm for understanding human psychology. The promise

and challenge of evolutionary psychology are to chart the set of human psychologicaladaptations—mechanisms designed by natural selection over the course of evolution that solve

particular adaptive problems. At a fundamental level, these mechanisms are conceptualized as

adaptive systems that served a variety of social and nonsocial functions in the environment of

evolutionary adaptedness (EEA) — the environment that humans evolved in and which presented the set of problems whose solutions are the set of human adaptations. This perspective

expects to find homologous (i.e., inherited from a common ancestor) systems in animals that

serve similar adaptive functions, and it expects that these systems will be organized within the brain as discrete neurophysiological systems (see Buss [2008] for a review focused on

personality psychology). It expects that each system will be responsive to particular

environmental contexts and that different temperament and personality systems will be in

competition with each other within individuals, leading at times to psychological ambivalence(MacDonald, 2005).

Here I review theory and data on temperament from an evolutionary perspective. Standard

definitions of temperament acknowledge the centrality of biology. Buss and Plomin (1984)defined temperament as a set of personality traits that originate early in life and have a

substantial biological (genetic) component. Rothbart’s definition focuses on the two broad

functional domains of temperament: constitutionally-based individual differences in reactivityand self-regulation (see Rothbart & Bates, 2006). Rothbart and Bates distinguish temperament

from personality by defining temperament as the affective, activational, and attentional core of

personality — all of which are strongly biological, while personality is a larger category thatincludes also beliefs, social cognition, morals, skills, habits, etc. and is more characteristically

human. For example, proneness to the emotion of fear is a function of temperament, but the

cognitive biases of fearful people are included in the study of personality.The term ‘trait’ implies that individual differences are critical to temperament. This requires

comment because in general, evolutionary psychologists regard adaptations as species-wide

universals. However, genetic variation is ubiquitous, even for adaptations (e.g., West-Eberhard,

2003), leading to the evolution of appraisal mechanisms in which the value of different personality traits may be appraised differently depending on the perceived interests of evaluators

— potential spouses, lovers, employees, employers, friends, leaders, etc. (Lusk, MacDonald &

Newman, 1998). For example, finding mates is an evolutionarily ancient problem for both sexes,so it is not surprising that sex-specific mate-detection mechanisms have evolved or that there is

substantial cross-cultural commonality in what people seek in spouses (Buss, 1989).

From an evolutionary perspective, individual differences within the normal range are seen asvariation in evolved systems. Research in neuroscience has revealed that the mammalian brain

contains highly specific emotional and motivational systems (e.g., Panksepp, 1998). These are

adaptations designed to carry out fundamental biological functions. However, there is important

variation among people in these systems, ranging, for example, from relatively restrained torelatively unrestrained attitudes and behavior regarding sex (Schmitt, 2005).

The most accepted proposal for why genetic and phenotypic variation in adaptive systems

remains in populations is the need to confront environmental heterogeneity (MacDonald, 1995;


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Nettle, 2006; Penke, Denissen, & Miller, 2007). The importance of environmental heterogeneity

in maintaining variation in personality-related traits is well established. For example, Nettle

(2006) used the example of the Galapagos finches: Variation is maintained in the population because different traits are more or less adaptive at different times (e.g., drought versus normal

rainfall; Grant & Grant, 1986). Further, maintaining a heterogeneous feeding environment results

in more genetic variation in Drosophila (MacKay, 1981). Nettle uses the example of research onthe great tit (Dingemanse, Both, Drent, & Tinbergen, 2004) to illustrate the point that a

personality profile that is adaptive in resource rich-environments is not adaptive in resource-poor

environments: Traits that are beneficial in some environments impose costs in others depending

on local environmental conditions. For example, in humans, extraversion may have been beneficial because it was associated with a greater number of sexual partners, but it also exposes

people to increased risk from accidents and law breaking.

An evolutionary theory seeks to establish the set of adaptations that underlie temperamentand personality. This is not the same as showing that a temperament trait has a biological basis or

that it is genetically influenced. It seeks to “carve nature at its joints” — on the basis of

functional units—systems that have been the focus of natural selection. An illustrative example

of a trait that shows genetic variation but is not an adaptation is proneness to divorce. McGueand Lykken (1992) found that proneness to divorce is heritable. However, proneness to divorce

does not reflect variation in an adaptation. Different people are prone to divorce for different

reasons (e.g., emotional instability [high neuroticism], selfishness, or proneness to philandering).Finding genetic variation for proneness to divorce is not without scientific interest; but it likely

represents the influence of variation in a number of different adaptations for different people. It

is not carving nature at its joints.The standard psychometric approach is not ideal for discovering the adaptations underlying

personality and temperament. For example, Freeman and Gosling’s (2010) review of studies of

Primate personality found 14 categories: Sociability, fearfulness, playfulness,confidence/aggression, activity, excitability, curiosity, dominance, agreeableness, irritability,

intelligence, impulsiveness, anxiousness, and independence. Support for putative personality

dimensions depends on inter-rater reliability as well as convergent and discriminant validity, allof which are at least promising.

From an evolutionary perspective, finding reliable and valid traits is an essential first step.

However, an evolutionary analysis requires in addition evidence that these traits are real foci of

natural selection. Ideally, one could find independent evidence that the traits represent variationin adaptive systems designed to solve particular problems. Was variation in curiosity or

playfulness a focus of natural selection or is it simply “noise” as Tooby and Cosmides (1990)

proposed for personality variation in general? Are these traits like the divorce example —complexly influenced by a variety of evolved systems and where different animals may be

playful or curious for different reasons? Are these traits byproducts of adaptive systems — for

example, the color of bone or blood is the result of selection for bone structure, not color per se.Is the variation linked to differences in adaptive outcomes in the different environments that the

animal encounters, or is it merely variation that human observers find interesting?

Tools of an Evolutionary Theory: Evolution, Sex, and Age.

An important tool for carving nature at its joints is the evolutionary theory of sex. The

evolutionary theory of sex is a powerful predictive theory of sex differences in fundamental


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extractions and rotations result in a lack of consensus. The hope here is that an evolutionary

perspective would provide some grounding for the field.

Variation in personality and temperament represents variation in the most basic systemsunderlying a person’s interactions with the social and non-social environment. These systems are

designed to achieve fundamental goals of survival and reproduction.

In general, evolutionary psychologists are “splitters” when it comes to studying adaptations.That is, they study each putative adaptation separately; the emphasis is on universality, so

typically there is no concern for whether the trait exhibits genetic variation or how people

evaluate this variation or how the genetic variation results in different outcomes in different

environments. Nor is there concern for correlations among individual differences amongdifferent adaptations. For example, evolutionary psychologists study anger as an adaptation (Sell,

Cosmides, & Tooby, 2009) without considering how individual differences in anger are

correlated with individual differences in other traits.Because of the centrality of individual differences, personality psychology has naturally

been interested in the correlational patterns among traits, leading to an important role for higher-

order factor analyses, such as the Five Factor Model. This review conceptualizes such models as

complexly related to several general adaptive spaces. These adaptive spaces are meaningfullyrelated to the Five Factor Model but they cannot be mapped in a 1:1 manner with the results of

factor analytic studies.

The concept of an adaptive space is an abstraction in the sense that it does not refer to a particular adaptation but to a suite of adaptations with the following attributes:

• There are phenotypic correlations among individual differences in thetraits detectable by factor analysis;

• There is shared genetic variation reflected in the heritability of thefactors;

• There are broadly similar functions among this suite of adaptations;

•

There are shared motivational, attentional, perceptual and, in additionfor personality, cognitive mechanisms; (There may also be motivational,attentional, perceptual and cognitive mechanisms that are unique to a particular

sub-system.)

• This suite of adaptations results from an evolutionary history ofelaboration and differentiation from systems that existed in common ancestors.

All of these characteristics of an adaptive space are falsifiable empirical propositions. That

is, it is possible that there are phenotypic correlations among individual differences in a set oftraits even though there is no common evolutionary history or genetic overlap. Such overlap

could occur if, for instance, people tended to group quite different types of negative affect

together (e.g., fear and anger) even though fear and anger are quite different functionally and atthe level of neuropsychology (see below). Nevertheless, the adaptive space idea is compatiblewith how evolution typically works: Gradual differentiation and elaboration of existing

structures rather than evolving structures de novo. This results in the expectation that personality systems should be conceptualized hierarchically,

from more specific to more general, within a generally defined adaptive space. For example, the

Behavioral Approach Adaptive Space is at the highest level of evolutionary analysis — the

highest level of meaningful adaptive space, with more specific systems arrayed beneath it. An


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evolutionary interpretation suggests that these different sub-systems accrued over evolutionary

time as primitive foraging and mate attraction systems became elaborated and somewhat

differentiated in response to specialized features of the “approach” adaptive space, effectivelyresulting in sub-systems or “facets” of temperament and personality: mechanisms that share

anatomical and neurological structures as well as genetic and phenotypic variance with each

other and may therefore be nested under one or more of the superfactors of higher order models, prototypically the Five Factor Model. Thus, for example, testosterone influences aggression,

dominance, sexual activity, mating effort, anti-social behavior, rough and tumble play, and

personality traits (extraversion, sociability, disinhibition, sensation seeking, and instrumental

effort) (Archer, 2006). Thus despite important differences among these behaviors anddispositions as well as the mechanisms underlying them, they share common mechanisms as

well.

The following sketches six “adaptive spaces” that are navigated by humans and animals andreflected in the temperament and personality literature. The procedure is to sketch out these

adaptive spaces and then, by using the evolutionary tools mentioned above, discuss how the

results of factor analytic studies map onto these adaptive spaces.

The Behavioral Approach Adaptive Space. Perhaps the most basic adaptive space that

animals and humans must navigate is to obtain resources from the environment. Among even the

most primitive mammals, there must be systems designed to approach the environment to obtainresources, prototypically foraging and mate attraction systems. For example, Panksepp (1998)

has argued that the mammalian brain contains a “foraging / exploration / investigation / curiosity

/ interest/ expectancy / SEEKING” system (p. 145).The behavioral approach personality systems evolved from systems designed to motivate

approach toward sources of reward (e.g., sexual gratification, dominance, control of territory)

that occurred as enduring and recurrent features of the environments in which animals or humansevolved. It overlaps anatomically and neurophysiologically with aggression—not surprising,

since aggression is a prepotent way of dealing with the frustration of positive expectancies

(Panksepp, 1998, p. 191); aggression is an evolutionarily ancient, if not necessarily honorable,way of securing resources. Several of the traits studied by Freeman and Gosling (2010) —

confidence/aggression, dominance, impulsiveness, activity, curiosity and perhaps playfulness —

have a surface plausibility as components of the Behavioral Approach Adaptive Space for

Primates.Important components of the Behavioral Approach Adaptive Space are dopaminergic

reward-seeking mechanisms (Cloninger, 1987; Gray, 1987, 2000; Panksepp, 1982, 1998;

Zuckerman, 1991). In rats, the dopaminergic reward-seeking mechanism involves energeticsearching, investigating, and sniffing objects in the environment as possible sources of reward,

but this seeking behavior is motivationally generalized: It can be directed at any of a variety of

specific rewards depending on the context (Panksepp, 1998). Gray (1987, 2000) proposes closelinkages between behavioral approach mechanisms and positive emotions, and Heller (1990)

notes that the left hemisphere contains high levels of dopamine reward mechanisms and there are

massive projections from the dopamine receptors to the left frontal areas associated with positive

affect.The centrality of reward-seeking mechanisms underlying behavioral approach reflects the

typical manner in which evolution shapes positive motivation to engage in behavior (Wilson,

1975). Evolution has resulted in affective motivational systems that are triggered by specific


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types of stimulation (e.g., the taste of sweet foods, the pleasure of sexual intercourse, the joy of

the infant in close, intimate contact with its mother), and it is difficult conceptualize how it could

have done otherwise. The evolutionary basis of the motivation to approach the environment in pursuit of necessary resources is the evolution of affective systems underlying particular adaptive

behaviors in the EEA.

The evolutionary hypothesis is that behavioral approach reflects common and overlapppingneurophysiological structure among these different components, but it is also compatible with

differences among them, as between aggression and other aspects of behavioral approach or

between exploration and interest.

There are also species differences in behavioral approach related to the animal’s ecology; forexample, predatory aggression is a component of behavioral approach in cats, but not in rats

(Panksepp, 1998, p. 194). In other words, over evolutionary time, the approach adaptive space

becomes elaborated and differentiated according to the unique adaptive demands of each species.For example, the foraging components of behavioral approach in carnivores are very different

from herbivores, the former requiring mechanisms involved in stalking and taking down the

prey, the latter requiring mechanisms for locating edible plants. Dominance mechanisms are an

important component of behavioral approach for many social species, but not for solitaryspecies.

The behavioral approach system is related to Surgency/Extraversion in the FFM and

Dominance in the Circumplex Model of Interpersonal Descriptors (Trapnell & Wiggins, 1996;Wiggins, 1991). At the heart of behavioral approach is Dominance/Sensation Seeking which

consists of individual differences in social dominance as well as several other highly sex-

differentiated behaviors, including sensation seeking, impulsivity, and sensitivity to reward. Newman (1987; see also Derryberry, 1987) found that compared to introverts, extraverts have a

stronger response to reward. Among adults, behavioral approach is also associated with

aggressiveness and higher levels of sexual experiences (Gray, 1987, 2000; Zuckerman, 1991),while impulsivity, “High Intensity Pleasure,” and aggressiveness are components of behavioral

approach in young children (Rothbart et al., 2001). Fox (1991) found that approach tendencies

are associated with positive emotions and anger, while avoidance tendencies are associated withthe negative emotions of distress, fear, and disgust. This indicates that aggression and sensitivity

to reward are associated not only at the level of neurobiology, but also at the level of individual

differences—a point to be returned to below.

Differences in attraction to reward are thus central to behavioral approach. Newman (1987;see also Avila, 2001; Derryberry, 1987) found that compared to introverts, reward has a

relatively greater effect on responding among extraverts and especially among disinhibited

subjects (psychopaths). The responding of some subjects was actually facilitated by punishment.Similarly, Frick and colleagues (Frick & Ellis, 1999; Frick et al., 1993) found that a subset of

children diagnosed with conduct disorder were strongly attracted to reward and sensation seeking

as well as being highly aggressive.The most sexually-differentiated aspects of behavioral approach are maximized during late

childhood and early adulthood while non-sex-differentiated aspects of behavioral approach

appear early in infancy and are strongly associated with positive emotionality. Sensitivity to

reward emerges very early in life as a dimension of temperament and is independent of measuresof behavioral inhibition, the latter system developing in the second half of the first year (Bates,

1989; Rothbart, 1989a; Rothbart & Bates,2006). In early infancy there are individual differences

in the extent to which infants approach rewarding stimulation, as indicated by attraction to sweet


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food, grasping objects, or attending to novel visual patterns. This trait is sometimes labeled

“exuberance,” defined as an “approach-oriented fact of positive emotionality (Pfiefer et al.,

2002; see also Fox et al., 2001). Children who are high on behavioral approach are prone to positive emotional response, including smiling, joy, and laughter available in rewarding

situations and the pleasant social interaction sought by sociable children.

Sensation seeking, including the promiscuous sexual activity loading on the Disinhibitionsubscale (Zuckerman, 1979), and aggression (Wilson & Daly, 1985) peak in late adolescence and

young adulthood, followed by a gradual decline during adulthood. As noted above, this “young

male syndrome” is highly compatible with evolutionary thinking: Sex-differentiated systems are

expected to be strongest at the time of sexual maturation and maximum divergence of male andfemale reproductive strategies. Because mating is theorized to involve competition with other

males, the male tendencies toward sensation seeking, risk taking and aggression are expected to

be at their peak during young adulthood when males are attempting to establish themselves in thewider group and accumulate resources necessary for mating.

However, boys are higher on behavioral approach even during infancy in cross cultural

samples (see Rothbart, 1989a for a review), and sex differences in aggression (Eagly & Steffan,

1986), externalizing psychiatric disorders (conduct disorder, oppositional/defiant disorder), risk-taking (Klein, 1995) and rough and tumble play (which is often associated with aggression) can

be seen beginning in early childhood (DiPietro, 1981; MacDonald & Parke, 1986; Humphreys &

Smith, 1987; O’Brien & Huston, 1985). Beginning in infancy, boys engage in more large-motor, physically intense activity (Eaton & Enns, 1986; Eaton & Yu, 1989). Increases in activity level

are the clearest effect of prenatal exposure of genetic females to androgens (Ehrhardt, 1985;

Ehrhardt & Baker, 1974). In factor analytic work activity level appears to line up in the samearea as dominance and sensation seeking (see Larsen & Diener, 1992).

Moreover, the social interactions of boys are more characterized by dominance interactions

and forceful, demanding interpersonal styles (Charlesworth & Dzur, 1987; Cowan & Avants,1988; LaFreniére & Charlesworth, 1983; Savin-Williams, 1987). On the other hand, females are

more prone to depression which is associated with low levels of behavioral approach (Davidson,

1993; Fox, 1994). Indeed, anhedonia and negative mood are primary symptoms of depressionwithin the DSM IV classification.

Taken together, the data on behavioral approach indicate that over development there is

differentiation of behavioral approach from a relatively simple dimension involving differences

in activity level, approach to novel objects, visual displays and sensory stimulation, to sociabilityand positive emotionality during early infancy, to include aggression, dominance, sensation

seeking, and rough and tumble play during early childhood, and undergoing further

differentiation and intensification as children approach reproductive competence and behavioralapproach begins to include attraction to sexual gratification.

Research in personality strongly supports the proposal that variation in intercorrelated

systems of behavioral approach is sex-differentiated in a manner consistent with the evolutionarytheory of sex. Males are more prone to physical aggression, dangerous sensation seeking and

risk-taking, and social dominance.

The Behavioral Withdrawal Adaptive Space. Another important general adaptive space iscoping with environmental threats. While behavioral approach systems respond to environmental

opportunities, specialized systems are required to respond to environmental threats,

prototypically by withdrawal or defensive aggression. The paradigm for a system designed to


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respond to environmental threats is the well-studied fear system (Gray 1987; LeDoux, 1996). It

is well established that the fear system results in reflexive defensive behavior prior to any

conscious processing (LeDoux, 1996, 2000). This evolutionarily ancient pathway for fear processing routes stimuli directly to the amygdala, bypassing the cortex entirely. For example, in

humans a loud noise or the sudden appearance of a threatening snake results in a reflexive

response directly triggered through the amygdala. Freeman and Golsing’s (2010) fearfulness andanxiousness are plausibly considered as components of the Behavioral Withdrawal Adaptive

Space in a wide range of Primates.

The Behavioral Inhibition System functions to monitor the environment for dangers and

impending punishments (Gray, 1987, 2000; LeDoux, 1996). The BIS responds with the emotionsof fear and anxiety to signals of uncertainty or anticipated punishment. Individual differences in

behavioral inhibition are observable beginning in the second half of the first year of life with the

development of the emotion of fear and expressions of distress and hesitation in the presence ofnovelty (Rothbart, 1989a; Rothbart & Bates,2006). Children who are high on behavioral

inhibition respond negatively to new people and other types of novel stimulation (Kagan et al,

1987; Fox et al. 2001). Physiological research on behaviorally inhibited children indicates that

these children generally have a more responsive sympathetic nervous system. This sympatheticdominance can be seen by the finding that behaviorally inhibited children tend to have a high and

stable heart rate in unfamiliar situations, indicating that these children are highly aroused by

unfamiliarity. Inhibited children also appear to have a highly sensitive amygdala, a limbicstructure implicated in fear reactions (Fox et al., 2001; Kagan & Snidman, 1991). Many

behaviorally inhibited children respond intensely to novel situations, and in particular, they tend

to be highly prone to tension, anxiety, and fear in these situations.Evolutionary theory predicts that females will be more sensitive than males to physical

dangers. Females are more prone to most anxiety disorders, including agoraphobia and panic

disorder (e. g., Weissman, 1985; DSM IV). Girls report being more fearful and timid in uncertainsituations than boys and are more cautious and take fewer risks than boys (Christopherson, 1989;

Ginsburg & Miller, 1982).

The Reactivity/Affect Intensity Adaptive Space. A third important adaptive space is

reactivity (affect intensity). Variation in arousal is needed in order to energize the animal to meet

environmental challenges or opportunities; in the absence of such a system, the animal would

either be permanently aroused — which would needlessly consume resources, or it would be permanently underaroused and less able to meet environmental challenges.

Affect Intensity functions to mobilize behavioral resources by increasing arousal in acutely

demanding situations in the service of either approach or withdrawal. Affect intensity may beviewed as a general behavioral “engine” that is utilized both in the service of behavioral

approach and behavioral avoidance. It is a behavioral scaling system that allows the organism to

scale its responses to current environmental opportunities and threats. This system is well studiedat the neurophysiological level where research implicates catecholamine systems that energize

both positive and negative emotion systems (Panksepp, 1998, pp. 109–110, 117). Freeman and

Gosling’s (2010) excitability is clearly related to the Reactivity/Affect Intensity Adaptive Space.

The Reactivity/Affect Intensity Adaptive Space is fundamentally concerned with reactivitywhich, along with self-regulation, comprise the two fundamental realms of temperament in

Rothbart’s scheme (see, e.g., Rothbart & Bates,2006). Children who are highly reactive respond

intensely to stimulation, reach peak arousal at lower stimulus intensity, and have a relatively low


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threshold for arousal (Rothbart, 1989a,b; Strelau, 1989). These children are often viewed as

having a weak nervous system in the sense that they are easily aroused and easily overstimulated.

In the presence of high levels of stimulation, these high-reactive individuals inhibit theirresponding and tend to withdraw from the source of stimulation. On the other hand, they respond

very intensely to even low levels of stimulation. Low-reactive children, on the other hand, may

be said to have relatively strong nervous systems in the sense that they have a relatively highthreshold of stimulation and do not become aroused by stimulation which would overwhelm a

high-reactive individual. These low-reactive individuals are thus more likely to be found in

highly stimulating environments, although at extremely high levels of stimulation, even these

individuals begin to inhibit their responding and withdraw from stimulation.Emotionally intense individuals respond relatively strongly to emotional stimulation

independent of the emotion involved, including both positive and negative emotions (Larsen &

Diener 1987)—an important aspect of the analysis of the fit between psychometric data and anevolutionary account discussed below. This is also implied in models of mood in which

activation (arousal) forms one dimension, while positive and negative affect is the other (Russell,

2003). Thus the degree of activation (arousal) is independent of the quality of the feeling state

(pleasure versus displeasure) but serves to accentuate or minimize a mood or emotion (see Figure1). As a result sad/gloomy and serene/contented are at the same level of low arousal but opposite

in valence; similarly, excited/ebullient and tense/jittery are at the same level of high arousal but

opposite in valence.

Place Figure 1 (= Fig. 1 of Russell, 2003) about here.

People high on affect intensity are prone to fast and frequent mood changes and lead varied

and variable emotional lives (Larsen & Diener 1987). Clinically, affect intensity is related to

cyclothymia, bipolar affective disorder, neurotic symptoms, and somatic complaints(nervousness, feeling uneasy, shortness of breath). Several developmental studies have found

that proneness to both positive and negative emotions under moderate levels of stimulus intensity

is associated with reactivity as indicated by measures of vagal tone and event-related potentials(Fox, 1989; Gunnar & Nelson, 1994; Porges, 1991). Garey et al. (2003) identified a generalized

arousal component in the behavior of mice across experiments, investigators, and mouse

populations. This factor accounts for about 1/3 of the variance in arousal-related measures.

Affect intensity may be viewed as a generalized motivation-enhancement system which can be directed toward behavioral approach (exhibited as positive emotionality) as well as behavioral

avoidance and checking for possible threats in the environment (exhibited as negative

emotionality, including irritability; note that Freeman and Gosling [2010] include irritability as aPrimate trait). The catecholamine systems underlying arousal are non-specific; they induce

arousal in a wide variety of systems (Panksepp, 1998). Individuals high on affect intensity are

thus highly motivated to intensive interaction with the environment, whether approach orwithdrawal. For example, Fox et al. (2001) found that highly reactive children who showed

continuity of behavioral inhibition (fearfulness) were prone to negative emotional responding

and had a pattern of right frontal asymmetry in their EEG patterns. On the other hand, highly

reactive exuberant, positively emotional children had a pattern of left frontal asymmetry.Reactive children are thus prone to intense emotional response, but they may be biased toward

positive or negative emotions. Again, these data illustrate the independence of reactivity from the

Behavioral Withdrawal and the Behavioral Approach adaptive spaces.


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Affect intensity is most closely associated with Neuroticism in the FFM (Larsen & Diener,

1992). Watson and Clark (1992) show that Neuroticism is associated with all four of their

dimensions of negative affect—guilt, hostility, fear, and sadness. However, these negativeemotions also tend to be associated with the other systems associated with the FFM: hostility

(negatively) with Nurturance/Love, sadness with introversion, and guilt with Nurturance/Love

and Conscientiousness. Neuroticism also appears to be related to a wide range of personalitydisorders which also load on other systems (Costa & McCrae, 1986; Widiger & Trull, 1992; see

also below). High affect intensity thus energizes negative emotional responding in general.

However, affect intensity also provides a powerful engine for positive emotional responses

which are central to other physiologically and psychometrically independent systems (Aron &Aron, 1997; Panksepp, 1998, p. 117; see below).

The Nurturance and Pair Bonding Adaptive Space. Mammalian females give birth to andsuckle their young. This has led to a host of adaptations for mothering, an outgrowth of which

are pair bonding mechanisms present also in males (MacDonald, 1992). For species that develop

pair bonds and other types of close relationships involving nurturance and empathy, one expects

the evolution of a system designed to make such relationships psychologically rewarding. Theadaptive space of Nurturance/Pair Bonding therefore becomes elaborated into a mechanism for

cementing adult relationships of love and empathy, prototypically within the family.

Variation in Nurturance/Love, the second factor emerging from the Circumplex model, isassociated with relationships of intimacy and other long-term relationships, especially family

relationships involving reciprocity and transfer of resources to others (e. g., investment in

children) (Kiesler, 1983; Trapnell & Wiggins, 1990; Wiggins, Trapnell, & Phillips, 1988).Recently, models of temperament have included a dimension of affiliativeness. Rothbart and

others (Derryberry & Rothbart, 1997; Evans & Rothbart, 2007; Gartstein & Rothbart, 2003;

Oldehinkel, Hartman, de Winter, Veenstra, & Ormel, 2004; Rothbart, 1994; Rothbart, Ahadi, &Evans, 2000; Rothbart & Bates, 2006) have proposed affiliativeness as a dimension of

temperament. Affiliativeness involves warmth, love, closeness, empathic concern and a desire to

nurture others. (This is not the same as sociability, which is linked to extraversion and is anaspect of the behavioral approach adaptive space. Sociability refers to a preference for

gregariousness — conversing, interacting, and approaching others. See below.)

Individual differences in warmth and affection observable in early parent-child relationships,

including secure attachments, are conceptually linked with this dimension later in life(MacDonald, 1992; 1997; 1999a). Secure attachments and warm, affectionate parent-child

relationships have been found to be associated with a high-investment style of parenting

characterized by later sexual maturation, stable pair bonding, and warm, reciprocally rewarding,non-exploitative interpersonal relationships (Belsky, Steinberg, & Draper, 1991).

The physiological basis of pair bonding involves specific brain regions (Bartels & Zeki,

2000) and the hormone oxytocin (Insel et al., 1998; Panksepp, 1998; Turner et al., 1999). In prairie voles—a monogamous species, oxytocin receptors are found in brain regions associated

with reward (Insel et al., 1998), supporting the proposal that pair bonding is a reward-based

system that functions to facilitate intimate family relationships and parental investment

(MacDonald, 1992). The stimuli that activate this system act as natural clues (in the sense ofBowlby [1969]) for pleasurable affective response. Intimate relationships and the nurturance of

the objects of affection are pleasurable, and such relationships are sought out by those sensitive

to the reward value of this stimulation.


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Such interconnections do not imply mechanisms of central, top-down control — merely

reflexive interactions among systems. However, an important potential general adaptive space is

the possibility of top-down control which would enable coordination of specialized adaptations,including all of the mechanisms associated with the four general adaptive spaces discussed

above. For many mammals, the prefrontal cortex or its analogues underlie executive control of

behavior that takes into account not only subcortically generated affective cues that are routedthough the orbitofrontal cortex (OFC), but also sensory input and other information (e.g., learned

contingencies) available to working memory (Uylings et al., 2003).

Humans have greatly elaborated this general adaptive space, resulting in top-down effortful

control mechanisms able to control a very wide array of mechanisms encompassed in the fourgeneral adaptive spaces mentioned above, but also capable of incorporating explicit contruals of

context in generating behavior, most notably linguistic and symbolic information (MacDonald,

2008). For example, affective states resulting from evolutionary regularities place people in a prepotently aggressive state—an emotional state that is one of the subsystems of the behavioral

approach adaptive space discussed above. Buss’s (2005) evolutionary theory of aggression

proposes that evolutionary regularities in the context of mating result in affective cues of sexual

jealousy and anger at romantic rivals that are prepotent cues for aggression. However, whether ornot aggression actually occurs may also be influenced, at least for people with sufficient levels of

effortful control, by explicit evaluation of the wider context, including explicit evaluation of the

possible costs and benefits of the aggressive act (e.g., penalties at law, possible retaliation, etc.;see below). These explicitly calculated costs and benefits are not recurrent over evolutionary

time but are products of the analytic system evaluating current environments and producing

mental models of possible consequences of behavior. Figure 2 illustrates the top down control ofapproach and withdrawal, based on MacDonald (2008).

Place Figure 2 about here.

Mary Rothbart has pioneered the idea that effortful control is a fundamental aspect of

temperament related to self-regulation (e.g., Posner & Rothbart, 1985). There is increasingcoherence between 22 and 33 months of age among a variety of tasks assessing the ability to

suppress dominant socioaffective responses—for example, waiting for a signal before eating a

snack, not peeking while a gift is wrapped, not touching a wrapped gift until the experimenter

returns (Kochanska, Murray, & Harlan, 2000). In general, effortful control increases with age,with girls superior to boys (Kochanska & Knaack, 2003; Kochanska et al., 2000; Lamb, Chuang,

Wessels, Broberg, & Hwang, 2002; Olson, Sameroff, Kerr, Lopex, & Wellman, 2005). The

superior performance of girls on effortful control fits well with the evolutionary theory of sexdiscussed above. Males are expected to be higher on behavioral approach systems (sensation

seeking, impulsivity, reward seeking, aggression) and therefore on average be less prone to

control prepotent approach responses (see Figure 3 above).Developmentally, the increasing efficiency of effortful control parallels developmental

changes in the prefrontal cortex. In general there is linear development of PFC from childhood to

adulthood; however, age changes in sensation seeking and reward-oriented behavior are

nonlinear because behavior is also influenced by the degree of maturation of limbic structuresunderlying the behavioral approach (Casey, Jones, & Hare, 2008; see Figure 3). This illustrates

well the complex, dynamically interactive nature of temperament systems as well as normative

changes over age in the relative strength of temperament systems.


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Place Figure 3 (=Figure 3 of Casey, Jones, & Hare, 2008) about here.

Beginning at about 10 to 12 months of age, there are individual differences in focused

attention and in the ability to inhibit inappropriate approach tendencies (Rothbart et al., 2000).

However, effortful control predicts ability to modulate both positive and negative emotionsassociated with behavioral approach (anger as measured by less angry reaction to tight seat belt

restraint; joy as measured by less joyful reaction to a puppet show) (Kochanska et al., 2000).

Effortful control is also associated with control over emotions linked to behavioral withdrawal

(fear, sadness, rumination as a symptom of depression; reviewed in MacDonald [2008]).Effortful Control is also negatively associated with Neuroticism (Evans & Rothbart, 2007),

results compatible with data showing central effortful inhibitory control over withdrawal

emotions.Several authors have proposed that the personality system most closely associated with

effortful control is conscientiousness (Caspi, 1998; Kochanska & Knaack, 2003; Rothbart et al.,

2000). The only temperament factor of Rothbart’s Adult Temperament Questionnaire that is

correlated with conscientiousness is the effortful control factor, which includes measures ofattention shifting from reward and from punishment (MacDonald, Figueredo, Wenner, &

Howrigan, 2007; Rothbart et al., 2000).

There are also strong conceptual links between conscientiousness and the effortful control of prepotent socioaffective responses. Conscientiousness is a dimension in the Five Factor model of

personality (Costa & McCrae, 1992; Digman, 1990, 1996; Goldberg, 1981; John, Caspi, Robins,

& Moffitt, 1994) referring to “socially prescribed impulse control that facilitates task and goal-directed behavior” (John & Srivastava, 1999, p. 121; italics in original). Conscientiousness

involves variation in the ability to defer gratification in the service of attaining long-term goals,

persevere in unpleasant tasks, pay close attention to detail, and behave in a responsible,dependable, cooperative manner (Digman & Takemoto-Chock, 1981; Digman & Inouye, 1986).

Conscientiousness is associated with academic success (Digman & Takemoto-Chock; Dollinger

& Orf, 1991; John et al., 1994), an area where there are sex differences favoring femalesthroughout the school years, including college (King, 2006). Correlations between high school

grades and assessments of conscientiousness performed 6 years previously were in the .50 range.

Similar correlations occurred for occupational status assessed when subjects were in their mid-

20s.At the conceptual level, these associations suggest that conscientious people are able to

control short-term pleasure seeking and other types of impulsive, affectively tinged responding

stemming from subcortical psychological adaptations. Conscientiousness does indeed predictactual performance tendencies related to impulse control. Persons low in conscientiousness tend

to have worse job performance and proficiency (Barrick & Mount, 1991), less restraint from

dishonest activities (Murphy & Lee, 1994), poorer health habits, and increased mortality(Friedman, 2000).

Psychopathology is associated with being extreme on personality systems (MacDonald,

1995; 2005; Widiger & Trull, 1992). The low end of conscientiousness is associated with a

variety of sex-differentiated externalizing behaviors (Eisenberg et al., 2004), conduct disorder(Krueger, Caspi, Moffitt, White, & Stouthamer- Loeber, 1996), aggression (Krueger et al., 1996;

Pulkkinen, 1986), adolescent drug use (Block, Block, & Keyes, 1988), delinquency (Krueger et

al.; Robins, John, & Caspi 1994; White et al., 1994), and antisocial personality disorder—for


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example, irresponsible and delinquent acts, failure to honor obligations or plan ahead (Widiger,

Trull, Clarkin, Sanderson, & Costa, 2002; Widiger & Trull), all of which are more common

among males. The greater involvement of males in such behavior is expected to be particularlyacute in adolescence and young adulthood when evolutionarily prepotent behavioral approach

tendencies related to aggression and sexuality mature and play a prominent role in sexual

competition among males — the young male syndrome noted above. Several theorists have proposed that adolescent antisocial behavior, reward seeking, and risk taking are at least partly

caused by the relative underdevelopment of prefrontal mechanisms compared to behavioral

approach mechanisms at this age (Casey et al., 2008; Raine, 2002; see Figure 3 above).

The Orienting Sensitivity Adaptive Space. Recent models of temperament have added

dimensions in order to make them more congruent with the Five Factor Model. The addition of

affiliativeness noted above provides a temperament basis for close relationships. Similarly,Evans and Rothbart (2007) propose that the temperamental basis for Openness is Orienting

Sensitivity, a trait that taps perceptual sensitivity, and is substantially correlated with standard

personality measures of Openness. Markon et al. show that Openness splits off from Positive

Emotionality when moving from a four-factor to a five-factor solution. Moreover, openness andextraversion appear on the same factor in 2-, 3- and 4-factor models but they split off in the 5-

factor solution (Caruso & Cliff, 1997; DeYoung, 2006; Digman, 1997; Rushton & Irwing, 2009).

This suggests that openness is part of the behavioral approach adaptive space, but that it became a unique adaptive space of its own as a result of differentiation and elaboration. This is

intuitively plausible because people high in Orienting Sensitivity are intensely engaged with the

environment in a positive manner, a posture that would naturally lead to approaching sources ofnovel aesthetic, perceptual, and intellectual experience characteristic of Openness. It also seems

likely that people high on Orienting Sensitivity are high on reactivity/affect intensity, since many

of the items indicate sensitivity to low-level stimulation. As indicated above, highly reactive people are prone to strong reactions to even relatively weak stimulation; and reactivity/affect

intensity does not appear as a separable factor in Five-Factor analyses of temperament and

personality questionnaires because at the phenotypic level it is intertwined with behavioralapproach and behavioral withdrawal (see Figure 5 below). The following are the sub-scales of

Orienting Sensitivity with sample items indicating the importance of sensitivity to environmental

cues:

Neutral Perceptual Sensitivity: Detection of slight, low intensity stimuli from both

within the body and the external environment. Example: “When I am barefoot, I am often

aware of the different textures of the surfaces that my feet touch.Affective Perceptual Sensitivity: Spontaneous emotionally valenced, conscious

cognition associated with low intensity stimuli. Example: “Sometimes when I hear a song

that I have not heard in a long time, I experience memories that are charged withemotion.”

Associative Sensitivity: Spontaneous cognitive content that is not related to

standard associations with the environment. Example: “I sometimes experience

sudden intuitive insights before I have actually thought things out.”


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Fitting the Adaptive Spaces to the Results of Factor Analytic Studies

Animal research. Individual differences in personality among chimpanzees can beunderstood within the FFM framework (Figueredo & King, 1996; King & Figueredo, 1994).

Reviewing the data for 12 quite different species, Gosling and John (1999) found evidence for

Extraversion (E), Neuroticism (N), and Agreeableness (A) in most species: E was found in 10species (but not rats and hyenas); N was found in 9 species (but not in vervet monkeys, donkeys,

and pigs); A was found in 10 species (but not in guppies and octopi). Conscientiousness (C) was

found only in humans and chimpanzees.

As noted above, Extraversion is associated with the Behavioral Approach Adaptive Space.Gosling and John’s (1999) results surely do not mean that rats and hyenas do not have behavioral

approach systems designed to obtain resources or that vervet monkeys do not have fear systems

or systems of reactivity/affect intensity. These findings may indicate that although these animalshave these systems, individual differences are not observable. For example, Figueredo and King

(1996, 2001) have hypothesized that social species are more likely to show individual differences

than nonsocial species.

On the other hand, it would not be surprising that guppies and octopi do not havemechanisms of pair bonding and close relationships, since such relationships are not part of these

animals’ ecology. Nor would it be surprising that humans and other relatively advanced animals

were uniquely involved in long-term projects requiring delay of gratification and close attentionto detail (i.e., Effortful Control/Conscientiousness); less cognitively advanced species — i.e.,

species that respond to environmental challenges mainly via preprogrammed responses — may

fail to exhibit differences in focused effort. The point is that the systems perspective expectsanimal personality psychology to mirror the ecology of the animal.

Human Research. Rothbart and Bates (2006) review studies that yielded from 2–8 factorsdepending on the items in the item pool. They note support for strong conceptual similarities to

three of the dimensions of the Five Factor Model: Negative Emotionality, Positive Emotionality,

and Effortful Control. Since that time, Rothbart has added items intended to tap the otherdimensions of the FFM, as reviewed above: Affiliation, tapping the Nurturance/Pair Bonding

Adaptive Space, and Orienting Sensitivity, intended to tap the temperamental basis for Openness

on the FFM (see Evans and Rothbart, 2007). As indicated above, Orienting Sensitivity may be

considered a sub-system of the Behavioral Approach Adaptive Space.These developments indicate a powerful convergence between research on temperament

and personality centered around the FFM. In seeking to determine how this factor analytic

research fits with an adaptationist perspective, I consider an important paper by Markon,Kreuger, and Watson (2005). This paper may be considered paradigmatic of a factor analytic

approach that could potentially be incorporated into an evolutionary account. This is because it

shows an orderly sequence in factor solutions, from two factors to five factors. The question is:Is it reasonable to view this result as mapping a 1:1 congruence between adaptive spaces and

personality factors? In particular, could the branching pattern noted in Figure 4 reflect a real

evolutionary sequence of elaboration and differentiation of primitive structures?

Place Figure 4 (equal to Markon et al.’s Figure 2) about here.


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Markon et al.’s α- and β-factors clearly refer to withdrawal (negative emotionality) and

approach (positive emotionality) respectively. The three-factor solution adds disinhibition, and

the four-factor solution distinguishes between disagreeable disinhibition and unconscientiousdisinhibition. As mentioned above, the fifth factor arises when openness splits off from

extraversion.

The differentiation of Negative Emotionality into Negative Emotionality, DisagreeableDisinhibition and Unconscientious Disinhibition reflects clinical categorizations rather than a

reasonable interpretation of evolutionary adaptive spaces. Disagreeable Disinhibition is related to

the reverse of the Nurturance/Pair Bonding Adaptive Space, and Unconscientious Disinhibition

is the reverse of Effortful Control/Conscientiousness within the Prefrontal Executive ControlAdaptive Space. It is more likely that evolution has resulted in selection for these positive traits

rather than their reverse. In the case of pair bonding, there is a clear evolutionary rationale for the

development of nurturance and pair bonding mechanisms derived from parental investmenttheory, as noted above. In the case of effortful control, the prefrontal machinery of top-down

control is one of the wonders of evolutionary engineering with clear adaptive benefits deriving

from planning and impulse control. On the other hand, the failure of these mechanisms is

powerfully linked to psychopathology, also discussed above.One gap between an evolutionary perspective and factor analyses such as presented by

Markon et al. is that an evolutionary perspective is much more compatible with a factor rotation

yielding factors of Dominance/Sensation Seeking and Nurturance/Love rather than Extraversionand Agreeableness (MacDonald, 1995, 1999a,b). Markon et al.’s Five-Factor solution is typical

of many others emphasizing extraversion-like traits as a basic factor. For example, Depue &

Collins (1999) advocate Gregarious/Aloof and Arrogant/Unassuming as fundamental causaldimensions of personality covering the same factor space. Extraversion is also a factor in the

NEO and the SNAP questionnaires utilized by Markon et al. in their study.

As Trapnell and Wiggins (1990) note, the difference amounts to a rotational difference between two ways of conceptualizing the same interpersonal space. Nevertheless, an

evolutionary perspective is better conceptualized with Dominance/Sensation Seeking and

Nurturance/Love as the primary axes of interpersonal space, since this conceptualizationmaximizes theoretically important sex differences and is thus likely to have been the focus of

natural selection. As noted above, evolutionary theory predicts that in species with sex-

differentiated patterns of parental investment, the sex with the lower level of parental investment

(typically the males) is expected to pursue a more high-risk strategy compared to females,including being prone to risk taking and reward seeking, and less sensitive to cues of

punishment. Depue and Collins (1999) claim that the traits associated with behavioral approach

(i.e., dominance, aggression, sensation seeking, risk-taking, boldness, sensitivity to reward, andimpulsivity are heterogeneous. But within the evolutionary theory of sex, they form a natural

unit: They all involve risky behavior that would benefit males more than females; they are

indeed heterogeneous at the level of mechanism but they also have mechanisms in common,notably testosterone (Archer, 2006). They are thus much more likely to be the focus of natural

selection than are Extraversion and Agreeableness. And, as noted above, there is a clear

evolutionary logic for supposing that mechanisms that promote parental investment are a

critically important adaptive space, again with clear implications for sex differences favoringfemales.

While there are robust sex differences favoring males in Dominance and Sensation Seeking

(Trapnell & Wiggins, 1990; Zuckerman, 1991), sex differences in Extraversion are relatively


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modest and actually favor females (McCrae et al., 2002; Srivastava, John, Gosling, & Potter,

2003). This is because Extraversion scales include items related to dominance and

venturesomeness, which are higher among males, as well as items related to warmth andaffiliation, which are higher among females (see discussion in Lucas, Deiner, Grob, Suh, &

Shao, 2000). In the Markon et al. (2005) study, warmth loads approximately equally on

extraversion and (negatively) on Disagreeable Disinhibition, and similar results were obtained byEvans and Rothbart (2007). From the evolved systems perspective developed here, it is unlikely

that combining warmth and affiliation with dominance, sensation seeking, and exploratory

behavior cuts nature at its joints.

Moreover, as discussed above, at the level of brain functioning, these systems are quiteseparate: There are unique neurochemical and neuroanatomical substrates for Nurturance/Love

and for Behavioral Approach, respectively (Archer, 2006; Bartels & Zeki, 2000; Depue &

Morrone-Strupinski, 2005; Panksepp, 1998). Focusing on the highly sex-differentiated traits ofdominance, aggression and sensation seeking, on one hand, and nurturance and love, on the

other, is not only much more compatible with a theoretical understanding of how evolution must

have worked but is also compatible with what we know of the systems actually found in the

brain.A related reason for focusing on these highly sex-differentiated traits is that they exhibit

theoretically expected age changes, while there is little evidence for mean age changes in

Extraversion (McCrae & Costa, 1990; McCrae et al., 2002). The “young male syndrome”describes the pattern in which sensation seeking, impulsivity, and aggression—all associated

with the behavioral approach systems—peak in young adulthood exactly at the time when young

males must compete for mates and establish themselves in the dominance hierarchy.Personality psychology is based on ratings of people by themselves and others, so that the

most socially salient features of people are emphasized and these may bear only indirectly on the

underlying systems. For example, the factor of Neuroticism refers to a tendency toward negativeemotionality, but at the system level, the research discussed above reveals separate systems of

reactivity/affect intensity (involving a general tendency toward both positive and negative

emotionality) and the Behavioral Withdrawal Adaptive Space dominated by the emotions of fearand anxiety. Reactivity/Affect Intensity should be understood to be a separable component of

temperament systems apart from motivation (MacDonald, 1988; Rothbart & Bates,2006). At the

motivational core of Behavioral Withdrawal are the emotions of fear and anxiety, while

Reactivity/Affect Intensity makes an independent contribution, heightening these emotions in people who are high on Reactivity/Affect Intensity.

This implies that the psychological salience of Neuroticism in everyday evaluations of self

and others provides an imperfect guide to the underlying systems. Similarly, the emergence ofExtraversion in factor analysis may well reflect social evaluations in everyday life: Extraverts

combine warmth and gregariousness with assertiveness and excitement seeking (See, e.g.,

Markon et al., Five-Factor solution, Table 10, p. 151.) As noted above, warmth loads moderatelyon both Extraversion and Disagreeable Disinhibition. But at the systems level, extraverts

combine the traits of two quite separate evolved systems designed for two quite different

purposes (behavioral approach and nurturance/pair bonding) with two quite different,

evolutionarily expected patterns of sex and age differences.Moreover, in the model presented by Markon et al., the Reactivity/Affect Intensity Adaptive

Space is split into positive emotionality and negative emotionality. Negative emotionality

includes hostility and aggression, both loading on the Disagreebale Disinhibition (reverse of


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Again, it is important to distinguish the arousal component of temperament systems as

separate from motivational components (MacDonald, 1988, 1995; Rothbart & Bates,2006). For

example, motivation for behavioral approach includes mechanisms such as sensitivity to rewarddiscussed above. As a result, people can be high on behavioral approach without being intensely

emotional. This is another example of the lack of a 1:1 mapping between the factors revealed by

psychometric research and an adaptationist perspective.Indeed, there is a very large literature showing that many people who are highly aggressive

and prone to sensation seeking are emotionally hyporeactive. For example, Adrian Raine and

colleagues (e.g., Ortiz & Raine, 2004; Raine, 2002) have provided evidence that reduced

adrenergic function as indicated by low resting heart rate is the best biological correlate ofaggression, anti-social behavior, and sensation seeking. As noted above, the biological substrate

of Reactivity/Affect Intensity is the adrenergic arousal system, indicating that these children are

low on Reactivity/Affect Intensity while nevertheless high on Behavioral Approach. (Deficits in prefrontal structures associated with Effortful Control/Conscientiousness are also implicated.)

Low resting heart rate at age 3 predicts aggressive behavior at age 11 and is heritable; sex

differences are in the expected direction: males are more likely to have low resting heart rate.

Such results are compatible with proposals that people low on autonomic arousal useaggression and sensation seeking to attain an optimal level of arousal (Eysenck, 1997; Raine,

1997; Quay, 1965). These results are also compatible with the idea that people high in

reactivity/affect intensity would avoid sensation seeking and aggression because these activitieswould be emotionally overwhelming (MacDonald, 1995): that is, highly reactive people—people

with weak nervous systems—withdraw in the presence of even moderate levels of stimulation.

These findings fit well with the common distinction between aggression accompanied by anger(‘hostile or reactive aggression’) and aggression without anger (commonly termed ‘instrumental

or proactive aggression’). For example, Frick and Ellis (1999) shows that children with reactive

aggression are prone to anger and emotional dysregulation, but this is not the case for childrendiagnosed with proactive aggression. Children labeled as callous/unemotional were found to

have the most severe type of conduct disorder. Such children are not only low on empathy, guilt,

and concern for others (associated with Nurturance/Love), they are generally low on emotionalexpressiveness (“does not show emotion”), including fearfulness and anxiety. Similarly,

psychopathic adults show a pattern of “lower anxiety, less fearfulness, and other evidence for

deficits in their processing of emotional stimuli” (p. 160). This low emotional sub-type is also

prone to sensation seeking and reward seeking— indicating they are high on behavioralapproach.

Bushman and Anderson (1999; see also Anderson & Bushman, 2002) note that emotionally

charged, angry aggression and aggression unaccompanied by anger may have similar motives(e.g., harming another, reclaiming self-esteem). In terms of the present framework, people who

combine high levels of behavioral approach with high reactivity/affect intensity are prone to

emotionally charged, angry, hostile aggression; they are also prone to reward seekingaccompanied by positive emotionality. On the other hand, combining high behavioral approach

with low reactivity/affect intensity is associated with aggression unaccompanied by anger

(instrumental, proactive aggression) and reward seeking unaccompanied by strong positive

emotionality.

Discussion


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to experience. Similarly, Rushton and colleagues have provided evidence for a general factor of

personality using five-factor model questionnaires (Rushton, Bons, & Hur, 2008; Rushton &

Irwing, 2009).Life history theorists propose that these loadings on a single factor derive from natural

selection for a coordinated suite of traits. That is, people with similar levels of extraversion,

conscientiousness, emotional stability, agreeableness and openness mate assortatively so thatthese traits become somewhat intercorrelated while nevertheless also yielding the lower level

analyses emphasized here. This is a promising direction that is compatible with the present

analysis.

References

Anderson, C. A., & Bushman, B. J. (2002). Human aggression. Annual Review of Psychology,

53, 27–51.

Archer, J. (2006). Testosterone and human aggression: an evaluation of the challenge hypothesis. Neuroscience Biobehavioral Reviews, 30, 319–345.

Aron, E. N., & Aron, A. (1997). Sensory-processing sensitivity and its relation tointroversion and emotionality. Journal of Personality and Social Psychology, 73,

345–368.

Avila, C. (2001). Distinguishing BIS-mediated and BAS-mediated disinhibitionmechanisms: A comparison of disinhibition models of Gray (1981, 1987) and of

Patterson and Newman (1993). Journal of Personality and Social Psychology,80, 311–324.

Barrick, M. R., & Mount, M. K. (1991). The Big Five personality dimensions and job

performance: A meta analysis. Personnel Psychology, 44, 1–26.

Bartels, A., & Zeki, S. (2000). The neural basis of romantic love. NeuroReport, 11(17), 3829– 3834.

Belsky, J., Steinberg, L., & Draper, P. (1991) Childhood experience, interpersonal development,

and reproductive strategy: An evolutionary theory of socialization. Child Development,62, 647–670.

Berndt, T. J. (1986). Children’s comments about their friendships. In M. Perlmutter

(Ed.), Minnesota Symposia in Child Development: Vol. 18. Cognitive perspectives on children’s social and behavioral development (pp. 189–212).Hillsdale, NJ: Erlbaum.

Buhrmester, D., & Furman, W. (1987). The development of companionship and intimacy. Child Development, 58, 1101–1113.

Bushman, B. J., & Anderson, C. A. (2001). Is it time to pull the plug on the hostile versus

instrumental aggression dichotomy? Psychological Review, 108 , 273–79

Betzig, L. (1986). Despotism and dmifferential reproduction. Hawthorne, NY: Aldine.Block, J., Block, J. H., & Keyes, S. (1988). The role of ego-control and ego-resiliency in the

organization of behavior. In W. A. Collins (Ed.), The Minnesota symposia on child

psychology (Vol. 13, pp. 39–101). Hillsdale, NJ: Erlbaum.

Bowlby, J. (1969). Attachment and loss: Vol. I. Attachment. London: Hogarth Press and theInstitute of Psychoanalysis.

Buss, A. H., & Plomin, R. (1984). Temperament: Early developing personality traits. Hillsdale,

NJ: Erlbaum.


23/35

Buss, D. M. (1989). Sex differences in human mate preferences: Evolutionary hypotheses tested

in 37 cultures. Behavioral & Brain Sciences, 12, 1–49.

Buss, D. M. (2005). The murderer next door: Why the mind is designed to kill. New York:Penguin.

Buss, D. M. (2008). Human nature and individual differences: Evolution of Human personality.

In O. John, R. W. Robins, & L. Pervin (Eds.), Handbook of Personality: Theory and Research (pp. 29–61). New York: Guildford Press.

Buss, D. M., & Schmitt, D. P. (1993). Sexual strategies theory: An evolutionary

perspective on human mating. Psychological Review, 100, 204–232.

Caspi, A. (1998). Personality development across the lifespan. In N. Eisenberg (Ed.), Handbook

of child psychology (Vol. 3, pp. 105–176). New York: John Wiley.

Caruso, J. C., & Cliff, N. (1997). An examination of the Five-Factor Model of normal

personality variation with reliable component analysis. Personality and Individual

Differences, 23(2), 317–325.

Casey, B. J., Jones, R. M., & Hare, T. A. (2008). The adolescent brain. Annals of the New York Academy of Sciences, 1124, 111–126.

Charlesworth, W., & Dzur, C. (1987). Gender comparisons of preschoolers’ behavior andresource utilization in group problem solving. Child Development, 58, 191–200.

Christopherson, E. R. (1989). Injury control. American Psychologist, 44, 237–241.

Clark, L. A., & Watson, D. (2008). Temperament: An ongoing paradigm for trait psychology. InO. John, R. W. Robins & L. Pervin (Eds.), Handbook of personality: Theory and

research, pp. 265–286. New York: Guildford Press.

Cloninger, C. R. (1987). A systematic method for clinical description and classification of personality. Archives of General Psychiatry, 44, 573–588.

Costa, P. T., & McCrae, R. R. (1986). Personality stability and its implications for clinical

psychology. Clinical Psychology Review, 6 , 407–423.Costa, P. T., Jr., & McCrae, R. R. (1992). Normal personality assessment in clinical practice:

The NEO Personality Inventory. Psychological Assessment, 4, 5–13.

Cowan, G., & Avants, S. K. (1988). Children’s influence strategies: Structure, sex differences,and bilateral mother-child influences. Child Development, 59, 1303–1313.

Davidson, R. J. (1993). The neuropsychology of emotion and affective style. In M. Lewis & J.

M. Haviland (Eds.), Handbook of emotions (pp. 143–154). New York: Guilford Press.

Dawson, G. (1994). Development of emotional expression and emotion regulation in infancy:Contributions of the frontal lobe. In G. Dawson & K. W. Fischer (Eds.), Human Behavior

and the developing brain, pp. 346–379. New York: Guilford Press.

Deaner, R. O., Barton, R. A., & van Schaik, C. P. (2003). Primate brains and life histories:Renewing the connection. In P. M. Kappeler & M. E. Pereira (Eds.), Primate life

histories and socioecology, pp. 233–265. Chicago: University of Chicago Press.Depue, R. A., & Collins, P. F. (1999). Neurobiology of the structure of personality: Dopamine facilitation

of incentive motivation and extraversion. Brain and Behavioral Sciences, 22, 491–569.Depue, R. A., & Morrone-Strupinsky, J. V. (2005). A neurobehavioral model of behavioral bonding:

Implications for conceptualizing a human trait of affiliation. Behavioral and Brain Sciences,28(3), 313–378.

Derryberry, D. (1987). Incentive and feedback effects on target detection: A chronometricanalysis of Gray's theory of temperament. Personality and Individual Differences, 8 ,

855–865.

http://homepage.psy.utexas.edu/homepage/group/busslab/pdffiles/SexDifferencesinHuman.PDFhttp://homepage.psy.utexas.edu/homepage/group/busslab/pdffiles/SexDifferencesinHuman.PDFhttp://homepage.psy.utexas.edu/homepage/group/busslab/pdffiles/SexDifferencesinHuman.PDFhttp://homepage.psy.utexas.edu/homepage/group/busslab/pdffiles/SexDifferencesinHuman.PDFhttp://homepage.psy.utexas.edu/homepage/group/busslab/pdffiles/SexDifferencesinHuman.PDFhttp://homepage.psy.utexas.edu/homepage/group/busslab/pdffiles/SexDifferencesinHuman.PDF


24/35

Derryberry, D., & Rothbart, M. K. (1997). Reactive and effortful processes in the organization of

temperament. Development and Psychopathology, 9, 633–652.

DeYoung, C. G. (2006). Higher-order factors of the Big Five in a multi-informant sample. Journal of Personality and Social Psychology, 91, 1138–1151.

Digman, J. M. (1990). Personality structure: Emergence of the five-factor model. Annual Review

of Psychology, 41, 417–440.Digman, J. M. (1996). The curious history of the five-factor model. In J. S. Wiggins (Ed.) The

five-factor model of personality: Theoretical perspectives, pp. 1–20. New York: Guilford.

Digman, J. M. (1997). Higher-order factors of the Big Five. Journal of Personality and SocialPsychology, 73, 1246–1256.

Digman, J. M., & Inouye, J. (1986). Further specification of the five robust factors of

personality. Journal of Personality and Social Psychology, 50, 116–123.

Digman, J. M., & Takemoto-Chock, N. K. (1981). Factors in the natural language of personality:Re-analysis, comparison, and interpretation of six major studies. Multivariate Behavioral

Research, 16, 149–170.

Dingemanse, N. J., Both, C., Drent, P. J., & Tinbergen, J. M. (2004). Fitness consequences of

avian personalities in a fluctuating environment. Proceedings of the Royal Society of London, Series B: Biological Sciences, 271, 847–852.

Digman, J. M. (1997). Higher-order factors of the Big Five. Journal of Personality and SocialPsychology, 73, 1246–1256.

DiPietro, J. A. (1981). Rough and tumble play: A function of gender. Developmental

Psychology, 17, 50–58.

Dollinger, S. J., & Orf, L. A. (1991). Personality and performance in “personality”:Conscientiousness and openness. Journal of Research in Personality, 25, 276–

284.

Douvan, E. A., & Adelson, J. (1966). The adolescent experience. New York: JohnWiley.

Draper, P., & Harpending, H. (1988). A sociobiological perspective on the development of

human reproductive strategies. In K. MacDonald (Ed.), Sociobiological perspectives onhuman development, pp. 340–372. New York: Springer-Verlag.

Eagly, A. H., & Steffan, V. J. (1986). Gender and aggressive behavior: A meta-analytic review

of the social psychological literature. Psychological Bulletin, 100, 283–308.

Eaton, W. O., & Enns, L. R. (1986). Sex differences in human motor activity level.Psychological Bulletin, 100, 19–28.

Eaton, W. O., & Yu, A. P. (1989). Are sex differences in child motor activity level a

function of sex differences in maturational status? Child Development, 60, 1005– 1011.

Ehrhardt, A. A., & Baker, S. W. (1974). Fetal androgens, human central nervous

system differentiation, and behavioral sex differences. In R. C. Friedman, R. M.Rickard, & R. L. Van de Wiele (Eds.), Sex differences in behavior. New York:

John Wiley.

Ehrhardt, A. A. (1985). The psychobiology of gender. In A. S. Rossi (Ed.), Genderand the life course. New York: Aldine.

Eisenberg, N., Spinrad, T. L., Fsabes, R. A., Reiser, M., Cumberland, A., Shepard, S.

A., et al. (2004). The relations of effortful control and impulsivity to children’s

resiliency and adjustment. Child Development, 75, 25–46.


25/35

Ellis, B. J. (2004). Timing of Pubertal Maturation in Girls: An Integrated Life History

Approach. Psychological Bulletin, 130, 920–958.

Ellis, B.J., Figueredo, A.J., Brumbach, B.H., & Schlomer, G.L. (2009). Fundamental dimensionsof environmental risk: The impact of harsh versus unpredictable environments on the

evolution and development of life history strategies.

Evans, D. E., & Rothbart, M. K. (2007). Developing a model for adult temperament. Journal of Research in Personality, 41, 868–888.

Human Nature, 20, 204–268.

Eysenck, H. J. (Ed.) (1982). Personality, genetics, and behavior. New York: Praeger. Eysenck,

H. J. (1997). Personality and the biosocial model of antisocial and criminal behavior. In

A. Raine, P. Brennan, D. P. Farrington, & S. A. Mednick (Eds.), Biosocial bases of

violence, pp., 21–38. New York: Plenum.

Eysenck, H.J., (1997), Personality and the biosocial model of antisocial and criminal behavior. In

A. Raine, P. Brennan, D. P. Farrington, & S. A. Mednick (Eds.) Biosocial bases of

violence (pp. 21–38). New York: Plenum.

Figueredo, A. J., & King, J. E. (1996). The evolution of individual differences in behavior.Western Comparative Psychological Association Observer, 2(2), 1–4.

Figueredo, A. J., Vásquez, G., Brumbach, B. H., & Schneider, S. M. R. (2004). The heritabilityof life history strategy: The K-factor, covitality, and personality. Social Biology, 51, 121–

143.

Figueredo, A. J., Vásquez, G., Brumbach, B. H., Schneider, S. M. R., Sefcek, J. A., Tal, I. R., etal. (2006). Consilience and life history theory: From genes to brain to reproductive

strategy. Developmental Review, 26 , 243–275.

Figueredo, A. J., Vásquez, G., Brumbach, B. H., & Schneider, S. M. R. (2007). The K-factor,covitality, and personality: A psychometric test of life history theory. Human Nature, 18 ,

47–73.

Fox, N. A. (1989). Psychophysiological correlates of emotional reactivity during the first year of

life. Developmental Psychology, 25, 364–372.Fox, N. A. (1991). If it’s not left, it’s right: Electroencephalograph asymmetry and the

development of emotion. American Psychologist, 46 , 863–872.

Fox, N. A. (1994). Dynamic cerebral processes underlying emotion regulation. In N. Fox (Ed.),

The development of emotion regulation: Biological and behavioral considerations.

Monographs of the Society for Research in Child Development, 59(2/3, Serial No. 240),

152–166.

Fox, N. A., Henderson, H. A., Rubin, K. H., Calkins, S. D., & Schmidt, L. A. (2001). Continuity

and discontinuity of behavioral inhibition and exuberance: Psychophysiological and

behavioral influences across the first four years of life. Child Development, 72, 1–21.

Freeman, H. D., & Gosling, S. D. (2010). Personality in nonhuman primates: A review and

evaluation of past research. American Journal of Primatology, 72, 653–671.Frick, P. J., & Ellis, M. (1999). Callous-unemotional traits and sub-types of conduct disorder.

Clinical Child and Family Psychology Review, 2,

Frick, P. J., Lahey, B. B., Loeber, R., Tannenbaum, L., Van Horn, Y., Christ, M. A. G., et al.

(1993). Oppositional defiant disorder and conduct disorder: A meta-analytic review of factor

analyses and cross-validation in a clinic sample. Clinical Psychology Review, 13, 319–340.

149–168.

http://cals.arizona.edu/fcs/sites/cals.arizona.edu.fcs/files/EllisFigueredoetal2009.pdfhttp://cals.arizona.edu/fcs/sites/cals.arizona.edu.fcs/files/EllisFigueredoetal2009.pdfhttp://cals.arizona.edu/fcs/sites/cals.arizona.edu.fcs/files/EllisFigueredoetal2009.pdfhttp://cals.arizona.edu/fcs/sites/cals.arizona.edu.fcs/files/EllisFigueredoetal2009.pdfhttp://cals.arizona.edu/fcs/sites/cals.arizona.edu.fcs/files/EllisFigueredoetal2009.pdfhttp://cals.arizona.edu/fcs/sites/cals.arizona.edu.fcs/files/EllisFigueredoetal2009.pdf


26/35

Friedman, H. S. (2000). Long-term relations of health: Dynamisms, mechanisms, and tropisms.

Journal of Personality 68, 1089–1107.

Garey, J., Goodwillie, A., Frohlich, J., Morgan, M., Gustafsson, J.-A.,Smithies, O., Korach, K.

S., Ogawa, S., & Pfaff, D. W. (2003). Genetic contributions to generalized arousal of brain

and behavior. Proceedings of the National Academy of Science, 100, 11019–11022.

Gartstein, M. A., & Rothbart, M. K. (2003). Studying infant temperament via the Revised Infant

Behavior Questionnaire. Infant Behavior and Development, 26 , 64–86.

Goetz, P., & D. Walters. (1997). The dynamics of recurrent behavior networks. Adaptive

Behavior, 6 (2), 245–282.

Goldberg, L. R. (1981). Language and individual differences: The search for universals in

personality lexicons. In L. Wheeler (Ed.), Review of personality and social psychology, Vol.

2, pp. 141–165. Beverly Hills, CA: Sage.

Gosling, S. D., & John, O. P. (1999). Personality dimensions in nonhuman animals: A cross-species

review. Current Directions in Psychological Science, 8 (3), 69–75.

Gray, J. A. (1987). The psychology of fear and stress (2nd

Gray, J. A. (2000). The neuropsychology of anxiety: An enquiry into the functions of the septo-hippocampal System (2

ed.). Cambridge: Cambridge

University Press.

nd

Grant, B. R., & Grant, P. R. (1989). Evolutionary dynamics of a natural population: The large

cactus finch of the Galapagos. Chicago: University of Chicago Press.

ed.). Oxford: Oxford University Press.

Gunnar, M. R., & Nelson, C. A. (1994). Event-related potentials in year-old infants: Relations

with emotionality and cortisol. Child Development, 65, 80–94.Heller, W. (1990). The neuropsychology of emotion: Developmental pattems and implications

for psychopathology. In N. L. Stein, B. Leventhal, & T. Trabasso (Eds.), Psychologicaland biological approaches to emotion (pp. 167–211). Hillsdale, NJ: Lawrence Erlbaum.

Humphreys, A. P., & Smith, P. K. (1987). Rough and tumble, friendship, and dominance inschool children: Evidence for continuity and change with age. Child Development, 58,

201–212.Insel, T. R., Winslow, J. T., Wang, Z., & Young, L. J. (1998). Oxytocin, vasopressin, and the

neuroendocrine basis of pair bond formation. Advances in Experimental Medicine and

Biology, 449, 215–224.John, O., Caspi, A., Robins, R. W., Moffitt, T. E., & Stouthamer-Loeber, M. (1994). The “little

five”: Exploring the nomological network of the five-factor model of personality in

adolescent boys. Child Development, 65, 160–178.

John, O. P., & Srivastava, S. (1999). The Big Five trait taxonomy: History, measurement, andtheoretical perspectives. In L. A. Pervin & O. P. John (Eds.), Handbook of personality:

Theory and research (2nd ed., pp. 102–138). New York: Guilford.Kagan, J., & Snidman, N. (1991). Infant predictors of inhibited and uninhibited profiles.Psychological Science, 2, 40–44.

Kernberg, O. F. (1986). Hysterical and histrionic personality disorders. In R. Michaels (Ed.),Psychiatry, Vol. 1 (19): 1–11). Philadelphia: J. B. Lippencott Co.

Kiesler, D. J. (1983). The 1982 interpersonal circle: A taxonomy for complementarity in human

transactions. Psychological Review, 90, 185–214.


27/35

King, J. (2006). Gender equity in education: 2006. Washington, DC: American Council on

Education.King, J. F., & Figueredo, A. J. (1994, April). Human personality factors in zoo chimpanzees? Paper

presented at the Western Psychological Association Convention, Kona, Hawaii.

Kochanska, G., & Aksan (1995). Mother-child mutually positive affect, the quality of child

compliance to requests and prohibitions, and maternal control as correlates of early

internalization. Child Development, 66 , 236–254.Kochanska, G., & Knaack, A. (2003). Effortful control as a personalit characteristic of young

children: Antecedents, correlates, and consequences Journal of Personality, 71, 1087–

1112.Kochanska, G., Murray, K. T., & Harlan, E. (2000). Effortful control in early childhood:

Continuity and change, antecedents, and implications for social development.

Developmental Psychology, 36, 220–232.

Krueger, R. F., Caspi, A., Moffitt, T. E., White, J. L., & Stouthamer-Loeber, M., (1996). Delayof gratification, psychopathology, and personality: Is low self-control specific to

externalizing problems? Journal of Personality, 64, 107–129.

LaFreniere, P. J., & Charlesworth, W. R. (1983). Dominance, affiliation and attention in a preschool group: A nine-month longitudinal study. Ethology and Sociobiology, 4, 55–67.

Lamb, M. E., Chuang, S. S., Wessels, H., Broberg, A. G., & Hwang, C. P (2002). Emergence

and construct validation of the Big Five factors in early childhood: A longitudinalanalysis of their ontogeny in Sweden. Child Development, 73, 1517–1524.

Lang-Takoc, E., & Osterweil, Z. (1992). Separateness and connectedness: Differences between

the genders. Sex Roles, 27, 277–289.

Larsen, R. J., & Diener, E. (1992). Problems and promises with the circumplex model ofemotion. Review of Personality and Social Psychology, 13, 25–59.

LeDoux, J. (1996). The emotional brain: The mysterious underpinnings of emotional life. New

York: Simon & Schuster.LeDoux, J. (2000). Emotion circuits in the brain. Annual Review of Neuroscience, 23, 155–184.

Lorenz, K. (1981). Foundations of ethology. New York: Springer-Verlag.Lucas, R. E., Deiner, E., Grob, A., Suh, E. M., & Shao, L. (2000). Cross-cultural evidence for the

fundamental features of extraversion. Journal of Personality and Social Psychology, 79, 452– 468.

Lusk, J., MacDonald, K., & Newman, J. R. (1998). Resource Appraisals among Self, Friend and

Leader: Implications for an Evolutionary Perspective on Individual Differences and a

Resource/Reciprocity Perspective on Friendship. Personality and Individual Differences,24, 685–700.

MacDonald, K. B. (1983). Stability of individual differences in behavior in a litter of wolfcubs (Canis lupus). Journal of Comparative Psychology, 2, 99–106.

MacDonald, K. B. (1988). Social and personality development: An evolutionary

synthesis. New York: Plenum.MacDonald, K. B. (1991). A perspective on Darwinian psychology: Domain-general

mechanisms, plasticity, and individual differences. Ethology and Sociobiology,

12, 449–480.

MacDonald, K. B. (1992). Warmth as a developmental construct: An evolutionaryanalysis. Child Development, 63, 753–773.

MacDonald, K. B. (1995). Evolution, the five-factor model, and levels of personality.

Journal of Personality 63, 525–567.


28/35

MacDonald, K. B., & Parke, R. D. (1986). Parent-child physical play: The effects of

sex and age of children and parents. Sex Roles, 15, 367–378.

MacDonald, K. B. (1997). The coherence of individual development: An evolutionary perspective on children’s internalization of cultural values. In J. Grusec & L.

Kuczynski (Eds.), Parenting strategies and chi

Temperament and Evolution

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