Evolutionary Psychology and the Emotions

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E
Evolutionary Psychology and the
Emotions
Laith Al-Shawaf
1,2
and David M. G. Lewis
3
1
Department of Psychology and Interdisciplinary
Neuroscience Program, Bilkent University,
Ankara, Turkey
2
College of Life Sciences, Institute for Advanced
Study, Berlin, Germany
3
School of Psychology and Exercise Science,
Murdoch University, Murdoch, WA, Australia
Introduction: Evolutionary Approaches
to the Emotions
Charles Darwin launched the evolutionary study
of emotion with his 1872 book, The Expression of
the Emotions in Man and Animals. However,
partly for historical reasons, the book had a nar-
row emphasis, focusing on the continuity of emo-
tional expression between humans and nonhuman
animals. The prevailing view in Darwin’s Victo-
rian England was that God had endowed humans
with specific facial muscles that He had specially
crafted to allow humans to communicate their
emotions to one another (e.g., Keltner et al.
2014). In such a social climate, Darwin’s goal
was to demonstrate to the scientific community
that human facial expressions bore the stamp of
their animal ancestry, and that close examination
of emotion expressions demonstrated the phylo-
genetic continuity between humans and other spe-
cies (Darwin 1872). In the century and a half since
the publication of Darwin’sOn the Origin of
Species (Darwin 1859), things have changed dra-
matically: the fact of evolution has been
established beyond any reasonable doubt, and
the theory of evolution is one of the most parsi-
monious, explanatorily successful, and predic-
tively powerful theories in all of science (Alcock
2009; Coyne 2009; Dawkins 2009; Dennett 1996;
Dobzhansky 1973). Now that there is no longer
any real (scientific) need to demonstrate that
humans share ancestry with all other species on
earth, scientists have found themselves free to
tackle a key question that Darwin largely avoided:
the evolved functions of emotions.
In the last few decades, evolutionists have
made considerable conceptual and empirical pro-
gress in understanding what emotions are and
why they evolved. There are several evolutionary
approaches in this tradition. The current entry
presents a theoretically powerful and empirically
promising contemporary approach that is closely
associated with evolutionary psychology and was
pioneered by John Tooby and Leda Cosmides
(Cosmides and Tooby 2000; Tooby and Cosmides
1990,2008). We can refer to this perspective as
the coordinating mechanism view of the
emotions.
#Springer International Publishing AG 2017
V. Zeigler-Hill, T.K. Shackelford (eds.), Encyclopedia of Personality and Individual Differences,
DOI 10.1007/978-3-319-28099-8_516-1

Coordinating Mechanisms: An
Evolutionary Psychological Approach
This evolutionary psychological perspective sug-
gests that emotions are coordinating mechanisms
whose evolved function is to coordinate a variety
of programs in the mind and body in the service of
solving a specific adaptive problem. For example,
fear coordinates programs in the service of pre-
venting or escaping danger (Tooby and Cosmides
1990; Marks and Nesse 1994), disgust regulates
mechanisms to prevent infection (Curtis et al.
2004; Oaten et al. 2009; Tybur et al. 2009), and
sexual arousal orchestrates physiological and psy-
chological programs for an advantageous mating
opportunity (Al-Shawaf et al. 2015b).
In their presentation of the coordinating mech-
anism approach, Tooby and Cosmides (1990;
Cosmides and Tooby 2000) suggest a list of pro-
grams regulated by the emotions, including:
(1) perceptual mechanisms, (2) attention,
(3) memory, (4) categorization, (5) motivational
priorities, (6) current goals, (7) information-
gathering adaptations, (8) specialized inference
mechanisms, (9) communication and expression,
(10) learning processes, (11) reflexes, (12) energy
level, mood, and effort allocation, (13) physiol-
ogy, and (14) behavior. The idea of the coordinat-
ing mechanisms approach is not that every
emotion is expected to regulate every program in
every instance of that emotion’s activation –
rather, the central idea is that emotions can be
best understood as regulatory mechanisms
whose evolved function is to coordinate a variety
of these programs to ensure their harmonious co-
activation in the service of solving an adaptive
problem.
Fear and Disgust as Illustrative Examples
Fear offers a useful illustration (Tooby and
Cosmides 1990; Cosmides and Tooby 2000).
When an organism is afraid of a potential threat
in the environment, a cascade of changes takes
place: perception becomes heightened (especially
to potential danger); attentional focus is narrowed
so that stimuli of less immediate relevance go
unnoticed; conceptual frameworks that cause
organisms to categorize stimuli as dangerous or
safe are activated while other conceptual frame-
works recede into the background; motivational
priorities change such that less urgent concerns
such as pathogen avoidance and status enhance-
ment fall by the wayside; and physiology shifts so
as to facilitate escape, for example by shunting
energy toward the muscles for flight (Tooby and
Cosmides 1990; Cosmides and Tooby 2000). The
central idea is that fear is a mode of operation for
the whole body and brain; a coordinating mecha-
nism that regulates a variety of physiological and
psychological programs to facilitate the solution
of the adaptive problem at hand.
Disgust, an emotion whose complexity we are
just beginning to understand, provides a second
example. When a pathogen threat activates dis-
gust, many changes take place: attention is
narrowed (van Hooff et al. 2013), sexual arousal
becomes more difficult to achieve (Fleischman
et al. 2015), heart rate and skin conductance are
affected (Schienle et al. 2001; Vrana 1993), par-
ticipants are less willing to engage in potentially
pathogenic behaviors (Tybur et al. 2011), immune
responses are activated (Schaller et al. 2010),
avoidant behaviors are engaged (Mortensen et al.
2010), and people report feeling less extraverted
and less open to new experiences (Mortensen
et al. 2010). The key idea again is the same:
pathogen disgust is a coordinating mechanism
that regulates and synchronizes the activity of
programs in the body and mind to solve the adap-
tive problem of avoiding infection.
How Does This Approach Differ from
Previous Evolutionary Approaches?
This “coordinating mechanisms”view of the
emotions shares features with several other evo-
lutionary approaches to emotion. Key elements of
overlap include the view that emotions are biolog-
ical adaptations that evolved to serve a function,
that adaptive action would be difficult or impos-
sible without emotions, that many emotions are
cross-culturally universal, that our emotions
evolved from our hominin ancestors and are
2 Evolutionary Psychology and the Emotions

phylogenetically related to the emotions of other
species, and that aversive emotions are just as
functional and beneficial as pleasant emotions
despite their inherent subjective aversiveness.
That said, this approach also differs from other
evolutionary approaches in several key respects.
The following six are particularly important.
First, the coordinating mechanisms perspective
suggests that there is no principled, non-arbitrary
reason to draw a distinction between “basic”and
“non-basic”emotions (Al-Shawaf et al. 2015b;
Tooby and Cosmides 1990; Cosmides and
Tooby 2000). There is no evolutionarily compel-
ling reason to privilege some emotions over
others, and no principled reason why the
privileged few should be limited to disgust,
anger, fear, sadness, joy, surprise, and sometimes
contempt (cf. Ekman 1992; Ekman and Cordaro
2011).
Second, most evolutionary accounts of emo-
tion have placed undue emphasis on survival to
the exclusion of reproduction. But evolutionary
scientists recognize that survival is important only
insofar as it facilitates reproduction: differential
reproductive success is the actual engine of the
evolutionary process (Alcock 2009; Hamilton
1964; Williams 1966). Both logic and empirical
evidence demonstrate that whenever survival and
reproduction conflict, the latter trumps the former
(Alcock 2009; Al-Shawaf et al. 2015b; Dawkins
1976; Williams 1966). Consistent with this fun-
damental evolutionary principle, the coordinating
mechanism approach expands the range of adap-
tive problems emotions evolved to solve to
include other adaptive problems tributary to
reproductive success, including problems such
as acquiring mates, retaining mates, competing
with intrasexual rivals, parenting and
childrearing, investing in kin, and more. Conse-
quently, the coordinating mechanism perspective
expands the range of evolved emotions to include
romantic love, parental love, sexual jealousy,
guilt, pride, shame, and many others (Al-Shawaf
et al. 2015b). An emotion does not need to facil-
itate survival per se; if it evolved to solve any
adaptive problem tributary to reproductive suc-
cess, then it is an evolved emotion.
Third, there is no need to stipulate that an
evolved emotion must have a recognizable signal
or facial expression (Tooby and Cosmides 1990).
Whether or not a given emotion evolved an
accompanying signal depends on the ancestral
costs and benefits of displaying that emotion to
others (Al-Shawaf et al. 2015b). Evolved emo-
tions can have accompanying facial expressions
that are fixed and universal, or changeable and
context-dependent, or they may not come with
an accompanying facial expression at all.
Fourth, whereas other evolutionary approaches
often insist that an emotion needs to also exist in
other species for it to qualify as an evolved emo-
tion, an evolutionary psychological approach
does not. Many adaptations are unique to one
species, but this does not make them any less
evolved or “basic.”On this view, an evolved
emotion can be shared with other species, unique
to humans, or shared with other species but with
uniquely human features (Al-Shawaf
et al. 2015b).
Fifth, an evolutionary psychological perspec-
tive is integrative and inclusive –rather than
insisting on the primacy of one particular aspect
of emotions, such as physiology, behavioral out-
put, facial expressions, or subjective feelings
(phenomenology), the coordinating mechanism
approach regards all of these elements as impor-
tant components of emotion. Further, it links them
together with other cognitive and perceptual ele-
ments (e.g., attention, memory, conceptual cate-
gorization) to paint a picture of emotion that is
more comprehensive and less narrowly defined
than approaches that elevate one particular ele-
ment, such as physiology or phenomenology,
above all else (Tooby and Cosmides 1990;
Cosmides and Tooby 2000).
And sixth –obviously but importantly –this
approach differs in its definition of emotions as
evolved neurocognitive programs whose function
is to regulate psychological and physiological
mechanisms in the service of solving an adaptive
problem.
Evolutionary Psychology and the Emotions 3

What Are the Benefits of This
Evolutionary Psychological Approach?
There are at least four major benefits of this
approach. First, it provides a non-arbitrary way
of classifying emotions. In particular, it avoids the
unwarranted basic versus non-basic division and
eschews the arbitrary insistence on particular
emotion elements such as facial expressions, dis-
tinctive physiology, or presence in other species in
order for an emotion to count as “basic”or “fun-
damental.”Second, in broadening the range of
adaptive problems that emotions have evolved to
solve, this perspective has heuristic value,
pointing researchers toward emotions that have
so far received scant attention from evolutionary
emotions researchers, such as romantic love,
parental love, regret, guilt, embarrassment, pride,
shame, and gratitude. Third, and perhaps most
important, this perspective provides a principled,
systematic method for generating a priori predic-
tions about emotions: the method of evolutionary
task analysis (Al-Shawaf et al. 2015b; Marr 1982;
see also Lewis et al. in press). An evolutionary
task analysis for an emotion consists of several
key questions: (1) what adaptive problem, if any,
did this emotion evolve to solve?, (2) which sub-
tasks must be solved in the solution of this adap-
tive problem?, (3) which information-processing
programs are capable of solving these subtasks?,
and (4) how should these programs be coordi-
nated to deliver a well-designed solution to this
adaptive problem? (Al-Shawaf et al. 2015b).
Finally, this perspective can be used to gener-
ate subtle and nuanced predictions about individ-
ual differences in emotion as well as context-
specific effects on emotion (Al-Shawaf et al.
2015b). For example, a task analysis might sug-
gest that individuals who are immunocompro-
mised should exhibit higher levels of disgust
(Fleischman and Fessler 2011; Al-Shawaf and
Lewis 2013); that whether disgust leads to avoid-
ance or approach may depend on who else is
present in one’s immediate environment (e.g.,
approach and kill the pathogen vector if it might
endanger one’s offspring; Al-Shawaf et al.
2015b); or that men’s psychological and physio-
logical programs return to baseline after orgasm
more rapidly than do women’s, but that this dis-
crepancy is attenuated for men who are oriented
toward committed long-term mating (Al-Shawaf
et al. 2015b). The key point is that an evolutionary
psychological approach to the emotions offers a
principled method for generating nuanced
hypotheses about individual differences and con-
text effects. This approach is theoretically power-
ful, empirically promising, and to our knowledge,
unique among theories of emotions in its capacity
to generate truly a priori hypotheses –let alone
such diverse and nuanced ones.
Is This Approach Making Empirical
Progress?
This perspective is relatively new, and only a tiny
fraction of its potential has been tapped. Never-
theless, there has been some important initial pro-
gress. For example, this perspective was recently
used to generate an array of novel hypotheses
about the effects of disgust on specialized infer-
ence mechanisms, information gathering, and
memory (Al-Shawaf et al. 2015b). Many of
these hypotheses include nuanced predictions
about individual differences and the effects of
context. All of these hypotheses were generated
a priori using the coordinating mechanism
approach in conjunction with evolutionary task
analysis.
Importantly, this predictive power extends to
other psychophysiological programs that are not
usually regarded as emotions. For example,
despite its powerful effects on physiology, psy-
chology, and behavior, sexual arousal does not
appear in previous taxonomies of basic emotions
and is typically excluded from the category of
emotions altogether (e.g., Barrett et al. 2016;
Ekman 1992; Keltner et al. 2014). In contrast, an
evolutionary psychological approach regards sex-
ual arousal as a critical emotion that evolved to
solve one of the most important adaptive prob-
lems faced by sexually reproducing organisms –
the coordination of a number of disparate mecha-
nisms in the service of conception. Accordingly,
the coordinating mechanism approach has
recently been used to generate a host of novel
4 Evolutionary Psychology and the Emotions

hypotheses about the effects of sexual arousal and
orgasm on memory, conceptual frameworks,
information gathering, specialized inference, and
learning (Al-Shawaf et al. 2015b).
A similar logic may apply to hunger: hunger is
typically regarded as a “drive”rather than an
emotion, but it is an important psychophysiolog-
ical state that may coordinate a variety of pro-
grams in the body and mind in the service of
acquiring food. Recent work suggests that the
coordinating mechanism approach can be used
to generate a bounty of new hypotheses about
the effects of hunger and eating on perception,
attention, memory, problem-solving, and concep-
tual categorization (Al-Shawaf 2016).
Recent work on anger and shame is also highly
promising (Sell et al. 2009a,b; Sell 2011; Sznycer
et al. 2012,2015,2016). The recalibrational the-
ory of anger suggests that anger is triggered when
an organism detects evidence that another individ-
ual is placing insufficient value on its welfare. The
primary evolved function of anger, on this view, is
to “convince”the target of the anger to upregulate
the value he places on the angry individual’s wel-
fare (Sell et al. 2009). This work is guided by the
coordinating mechanism perspective: the central
idea motivating this research is that anger coordi-
nates physiology, attention, communication,
facial expressions, body posture, behavior, and
other psychological programs in order to cause
another individual to recalibrate the weight he
places on the angry individual’s welfare. This
research is relatively new but has already demon-
strated that: (a) the anger face is not an arbitrary
set of muscular contractions, but rather a specific
constellation of changes that increases the per-
ceived fighting ability of the angry individual
(Sell et al. 2014), (b) the conditions that trigger
anger are exactly what you would expect
according to the welfare-recalibration theory
(e.g., stronger anger is triggered when the cost to
the victim was large, the benefit to the perpetrator
was small, and the perpetrator knew exactly who
he was harming; Sell 2005; Sell et al. under
review), (c) the content of apologies and attempts
to decrease anger are also exactly what you would
expect according this theory (e.g., the benefitto
me was large, I thought the cost to you was small,
and/or I didn’t know you were the person incur-
ring the cost from my behavior; Sell 2005;Sell
et al. under review), and (d) individual differences
in anger thresholds are predictable a priori on the
basis of one’s bargaining power, such that those
who are better able to confer benefits and inflict
costs tend to anger more easily and more readily
(e.g., attractive women and attractive and muscu-
lar men; Sell et al. 2009).
Recent work on shame also points to the utility
of the coordinating mechanism approach. This
approach suggests that shame is a species-wide,
cross-culturally universal mechanism that
evolved as a defense against being devalued by
others (Sznycer et al. 2012,2016). Guided by the
coordinating mechanism perspective, this
research proposes that shame coordinates changes
in physiology, perception, cognition, motivation,
and behavior in order to reduce the likelihood and
costs of social devaluation. This involves
(a) refraining from behaviors that may lead to
devaluation, (b) concealing information that may
lead to devaluation, and (c) minimizing the nega-
tive impact of devaluation when it does occur
(Sznycer et al. 2016). To this end, and consistent
with the coordinating mechanism approach,
researchers have discovered that shame motivates
the avoidance of behaviors and the concealment
of information that lead to devaluation
(Rockenbach and Milinski 2011), and that when
such information is discovered, shame leads an
individual to withdraw (Tangney et al. 1996),
accept subordination (Gilbert 2000), become
more cooperative (Masclet et al. 2003), and
appease social others (Keltner et al. 1997). Addi-
tionally, shame regulates endocrinology and
immunology, raising cortisol levels (Dickerson
and Kemeny 2004) and upregulating pro-
inflammatory cytokines to provide a defense
against infection (Dickerson et al. 2009).
The coordinating mechanism perspective on
the emotions has also led to important advances
in our understanding of pride, but space con-
straints prohibit us from discussing this research
in depth (see Sznycer et al. 2017; Tracy and
Robins 2007; Tracy et al. 2010, for more infor-
mation). Despite the relative youth of this theoret-
ical perspective and the novelty of the research
Evolutionary Psychology and the Emotions 5

programs discussed above, researchers are using
this approach to make rapid gains in our under-
standing of emotions as varied as disgust
(Al-Shawaf et al. 2015b), sexual arousal
(Al-Shawaf et al. 2015b), hunger (Al-Shawaf
2016), anger (Sell et al. 2009a,b,2014), shame
(Sznycer et al. 2012,2015,2016), and pride
(Sznycer et al. 2017; Tracy et al. 2010). The next
few years are sure to bring novel insights and new
discoveries –and to witness the extension of this
coordinating mechanism perspective to other
emotions that are ripe for investigation within its
framework, including romantic love, parental
love, guilt, regret, gratitude, and many more.
Culture, Context, and Individual
Differences
Three theoretical sidenotes bear mentioning here.
First, an evolutionary psychological approach to
the emotions does not imply that emotions are
innate, rigid, and inflexible. As with the rest of
evolutionary psychology, our evolved neurocom-
putational emotion programs are built by the joint
interplay of genes and environment, and are
highly responsive to environmental input (Buss
1995; Confer et al. 2010; Tooby and Cosmides
1992). By definition, these emotions are inextri-
cably tied to what is going on in the environment
around us, and this evolutionary perspective on
the emotions would obviously not make sense if
one were to try to remove the environment from
the equation. An evolutionary psychological
approach to the emotions regards the environment
as critical in driving the evolution of the mecha-
nism in the first place, crucial in the ontogenetic
development of the mechanism, and indispens-
able as a trigger of the mechanism in the immedi-
ate present (see Buss 1995; Confer et al. 2010;
Lewis et al. in press). This evolutionary psycho-
logical perspective is thus environment-centered
at all three stages of the causal process: evolution
across generations, development within a genera-
tion, and immediate causation. Misinformed alle-
gations that evolutionary approaches regard the
emotions as rigid, inflexible, genetically deter-
mined, or environmentally insensitive are simply
incorrect. For example, see Al-Shawaf et al.
2015b for a number of evolutionary hypotheses
about context effects and individual differences in
emotions.
Second, it is worth noting that an evolutionary
psychological approach does not predict complete
invariance in emotions across cultures. Recall that
the computational architecture of a psychological
mechanism consists of three stages of
information-processing: environmental cues are
taken as inputs, which are then stored and oper-
ated upon by computational procedures, which
are subsequently turned into outputs (e.g., behav-
ior). As a general principle, evolutionary psycho-
logical approaches do not predict universality at
the level of output, but rather at the level of com-
putational design (Al-Shawaf et al. 2015b; Lewis
et al. in press; Tooby and Cosmides 1992). Impor-
tantly, culturally variable input combined with
universal neurocognitive mechanisms often yields
culturally variable output (Gangestad and Buss
1993; Lewis et al. in press). This culturally vari-
able output is not necessarily a sign that the mech-
anism in question is not a product of evolution.
Rather, it is often a sign that the inputs that trigger
the mechanism differ across cultures. Stated dif-
ferently, if you start with different cultural or
ecological inputs you will likely get different out-
puts. Consequently, this view of the emotions
does not predict that every aspect of emotions
will be culturally invariant. Instead, it expects
variability in emotion output to be underlain by
cross-cultural uniformity in neurocomputational
design. This is precisely the pattern that seems to
emerge from cross-cultural studies on emotion
(e.g., Neumann et al. 2009; Sznycer et al. 2012,
2017). Perhaps the most interesting aspect of this
discussion is that rather than being arbitrary and
unpredictable, these cultural differences are often
systematically predictable a priori. An evolution-
ary psychological perspective makes it possible to
generate theoretically principled predictions
about what cultural differences one should expect
to observe in advance rather than observing cul-
tural differences and then concocting post hoc
explanations for them (see e.g., Sznycer et al.
2012 for an emotion example or Gangestad and
Buss 1993 for a non-emotion example).
6 Evolutionary Psychology and the Emotions

Third, an evolutionary psychological perspec-
tive does not imply the absence of individual
differences in emotion. On the contrary, individ-
ual differences in emotion are a natural part of the
coordinating mechanisms perspective. Providing
a nice parallel to the case of cultural differences,
an evolutionary psychological approach provides
one with a means for making theoretically
grounded a priori predictions about individual
differences. For example, it has been suggested
that individuals with less robust immune systems
may have lower disgust thresholds (e.g., Fessler
et al. 2004; Al-Shawaf and Lewis 2013;
Al-Shawaf et al. 2015b) and that the emotions
that arise after orgasm are likely different for
men oriented toward short-term mating compared
to those who are oriented toward long-term mat-
ing (Al-Shawaf et al. 2015b). Researchers using
this perspective have also predicted and found that
attractive women anger more easily than their
less-attractive counterparts (Sell et al. 2009b),
that attractive and physically formidable men
anger more readily than their less-attractive and
physically weaker counterparts (Sell et al. 2009),
that individuals with less bargaining power are
more prone to shame (Sznycer et al. 2012), that
individuals with lower relational mobility are
more shame-prone around their friends (but not
around strangers, Sznycer et al. 2012), and that
individuals with a stronger proclivity for short-
term mating have stably lower sexual disgust
(Al-Shawaf et al. 2015a). The key point here is
that, as with cultural differences, individual dif-
ferences are not only consistent with an evolution-
ary psychological approach to the emotions; they
can be predicted a priori according to theoretical
principles.
Future Directions
What are the next key steps for this evolutionary
approach to the emotions? First, researchers
should continue to generate and test novel hypoth-
eses based on this approach. The coordinating
mechanism view is heuristically valuable and pre-
dictively powerful, but given its youth, there is
still a great deal of work to be done at the
hypothesis-generation stage and even more to be
done at the design-and-testing stage. Researchers
have reaped only a small portion of the empirical
fruits of this approach so far, and this suggests an
exciting and promising future. Second,
researchers should continue to generate subtle
and nuanced hypotheses about context effects,
cultural differences, and individual differences in
emotion. As we move toward a more complete
explanation of species-typical mechanisms, sex-
typical mechanisms, cultural differences, and
individual differences, we move toward a more
comprehensive understanding of the mind –both
in the science of emotion and in psychology more
broadly. Finally, it would be valuable for evolu-
tionary psychologists to devote greater attention
to positive emotions. As it currently stands, we
understand the structure and evolved function of
the “negative emotions”a great deal better than
we do the “positive emotions.”This is slowly
starting to change with emotions such as gratitude
(Forster et al. 2016; McCullough et al. 2008) and
pride (e.g., Tracy and Robins 2007; Tracy et al.
2010), but our understanding of these emotions –
along with others such as joy, contentment, and
love –still does not compare to our understanding
of emotions such as anger, fear, and disgust. Great
benefit is likely to come from thinking about the
evolved function of positive emotions in a rigor-
ous, systematic, and theoretically principled way.
We look forward to this development and suggest
that the coordinating mechanism perspective pre-
sented here offers an especially fruitful path
forward.
Conclusion
Summary
An evolutionary psychological perspective sug-
gests that the emotions are coordinating mecha-
nisms whose evolved function is to orchestrate
psychological and physiological programs to
facilitate the solution of a particular adaptive
problem. The current entry presented this
approach, described how it differs from other
evolutionary approaches to the emotions, and
discussed the benefits of this approach. Key
Evolutionary Psychology and the Emotions 7

benefits include a more theoretically principled
method of classifying emotions, a broader consid-
eration of the adaptive problems that emotions
evolved to solve and a broader consideration of
the range of emotions that evolved to solve them,
and –crucially –the use of evolutionary task
analysis to generate a priori hypotheses about
each emotion, including hypotheses about context
effects, individual differences, and cultural vari-
ability. We then detailed the empirical progress
this approach has made in understanding certain
emotions, including anger, pride, shame, disgust,
and sexual arousal. Finally, this entry offers some
useful future directions, including further testing,
theoretical and empirical progress in understand-
ing cultural and individual differences, and
greater attention to positive emotions. We hope
that this entry helps introduce the coordinating
mechanism perspective to non-evolutionists as
well as to those familiar with other evolutionary
approaches to the emotions, and motivates
researchers to use this theoretically grounded
approach to generate and test novel a priori
hypotheses about a wide range of emotions.
Cross-References
▶Anger
▶Basic Emotions
▶Culture
▶Darwin, Charles
▶Desire
▶Disgust
▶Ekman, Paul
▶Embarrassment
▶Facial Expressions and Emotion
▶Fear
▶Guilt and Shame
▶Happiness
▶Human Nature and Human Universals
▶Jealousy
▶Love
▶Pride
▶Reproduction
▶Sadness
▶Sexual Selection
References
Alcock, J. (2009). Animal behavior: An evolutionary
approach. Sunderland: Sinauer Associates.
Al-Shawaf, L. (2016). The evolutionary psychology of
hunger. Appetite, 105, 591–595.
Al-Shawaf, L., & Lewis, D. M. G. (2013). Exposed intes-
tines and contaminated cooks: Sex, stress, & satiation
predict disgust sensitivity. Personality and Individual
Differences, 54, 698–702.
Al-Shawaf, L., Conroy-Beam, D., Asao, K., & Buss, D. M.
(2015a). Human emotions: An evolutionary psycho-
logical perspective. Emotion Review,1–14.
Al-Shawaf, L., Lewis, D. M. G., & Buss, D. M. (2015b).
Disgust and mating strategy. Evolution and Human
Behavior, 36, 199–205.
Barrett, L. F., Lewis, M., & Haviland-Jones, J. M. (Eds.).
(2016). Handbook of emotions. New York: Guilford
Publications.
Buss, D. M. (1995). Evolutionary psychology: A new par-
adigm for psychological science. Psychological
Inquiry, 6(1), 1–30.
Confer, J. C., Easton, J. A., Fleischman, D. S., Goetz,
C. D., Lewis, D. M., Perilloux, C., & Buss, D. M.
(2010). Evolutionary psychology: Controversies, ques-
tions, prospects, and limitations. American Psycholo-
gist, 65(2), 110.
Cosmides, L., & Tooby, J. (2000). Evolutionary psychol-
ogy and the emotions. In M. Lewis & J. M. Haviland-
Jones (Eds.), Handbook of emotions (2nd ed.,
pp. 91–115). New York: Guilford.
Coyne, J. A. (2009). Why evolution is true. New York:
Penguin.
Curtis, V., Aunger, R., & Rabie, T. (2004). Evidence that
disgust evolved to protect from risk of disease. Pro-
ceedings of the Royal Society B, 271, 131–133.
Darwin, C. (1859). On the origin of species by means of
natural selection. London: John Murray.
Darwin, C. (1872). The expression of the emotions in
animals and man. Chicago: Chicago University Press.
(Reprinted 1965).
Dawkins, R. (1976). The selfish gene. Oxford: Oxford
University Press.
Dawkins, R. (2009). The greatest show on earth: The
evidence for evolution. New York: Simon and Schuster.
Dennett, D. C. (1996). Darwin’s dangerous idea: Evolu-
tion and the meanings of life. New York: Simon and
Schuster.
Dickerson, S. S., & Kemeny, M. E. (2004). Acute stressors
and cortisol responses: A theoretical integration and
synthesis of laboratory research. Psychological Bulle-
tin, 130, 355–391.
Dickerson, S. S., Gable, S. L., Irwin, M. R., Aziz, N., &
Kemeny, M. E. (2009). Social evaluative threat and
proinflammatory cytokine regulation. Psychological
Science, 20, 1237–1244.
Dobzhansky, T. (1973). Nothing in biology makes sense
except in the light of evolution. The American Biology
Teacher, 75(2), 87–91.
8 Evolutionary Psychology and the Emotions

Ekman, P. (1992). An argument for basic emotions. Cog-
nition and Emotion, 6(3–4), 169–200.
Ekman, P., & Cordaro, D. (2011). What is meant by calling
emotions basic. Emotion Review, 3, 364–370.
Fessler, D. M., Pillsworth, E. G., & Flamson, T. J. (2004).
Angry men and disgusted women: An evolutionary
approach to the influence of emotions on risk taking.
Organizational Behavior and Human Decision Pro-
cesses, 95(1), 107–123.
Fleischman, D. S., & Fessler, D. M. (2011). Progesterone’s
effects on the psychology of disease avoidance: Sup-
port for the compensatory behavioral prophylaxis
hypothesis. Hormones and Behavior, 59(2), 271–275.
Fleischman, D. S., Hamilton, L. D., Fessler, D. M., &
Meston, C. M. (2015). Disgust versus lust: Exploring
the interactions of disgust and fear with sexual arousal
in women. PloS One, 10(6), e0118151.
Forster, D. E., Pedersen, E. J., Smith, A., McCullough,
M. E., & Lieberman, D. (2016). Benefit valuation pre-
dicts gratitude. Evolution and Human Behavior.
doi:10.1016/j.evolhumbehav.2016.06.003.
Gangestad, S. W., & Buss, D. M. (1993). Pathogen preva-
lence and human mate preferences. Ethology and
Sociobiology, 14(2), 89–96.
Gilbert, P. (2000). The relationship of shame, social anxi-
ety and depression: The role of the evaluation of social
rank. Clinical Psychology & Psychotherapy, 7,
174–189.
Hamilton, W. D. (1964). The genetical evolution of social
behaviour. II. Journal of Theoretical Biology, 7(1),
17–52.
van Hooff, J. C., Devue, C., Vieweg, P. E., & Theeuwes,
J. (2013). Disgust-and not fear-evoking images hold
our attention. Acta Psychologica, 143(1), 1–6.
Keltner, D., Young, R. C., & Buswell, B. N. (1997).
Appeasement in human emotion, social practice, and
personality. Aggressive Behavior, 23, 359–374.
Keltner, D., Oatley, K., & Jenkins, J. M. (2014). Under-
standing emotions (3rd ed.). Hoboken: Blackwell.
Lewis, D. M. G., Al-Shawaf, L., Conroy-Beam, D., Asao,
K., & Buss, D. M. (in press). Evolutionary psychology:
A how-to guide. American Psychologist.
Marks, I. M., & Nesse, R. M. (1994). Fear and fitness: An
evolutionary analysis of anxiety disorders. Ethology
and Sociobiology, 15(5), 247–261.
Marr, D. (1982). Vision: A computational investigation
into the human representation and processing of visual
information. San Francisco: Freeman.
Masclet, D., Noussair, C., Tucker, S., & Villeval, M. C.
(2003). Monetary and nonmonetary punishment in the
voluntary contributions mechanism. American Eco-
nomic Review, 93, 366–380.
McCullough, M. E., Kimeldorf, M. B., & Cohen, A. D.
(2008). An adaptation for altruism: The social causes,
social effects, and social evolution of gratitude. Current
Directions in Psychological Science, 17(4), 281–285.
Mortensen, C. R., Becker, D. V., Ackerman, J. M.,
Neuberg, S. L., & Kenrick, D. T. (2010). Infection
breeds reticence: The effects of disease salience on
self-perceptions of personality and behavioral avoid-
ance tendencies. Psychological Science, 21(3),
440–447.
Neumann, R., Steinhäuser, N., & Roeder, U. R. (2009).
How self-construal shapes emotion: Cultural differ-
ences in the feeling of pride. Social Cognition, 27(2),
327.
Oaten, M., Stevenson, R. J., & Case, T. I. (2009). Disgust
as a disease-avoidance mechanism. Psychological Bul-
letin, 135(2), 303.
Rockenbach, B., & Milinski, M. (2011). To qualify as a
social partner, humans hide severe punishment,
although their observed cooperativeness is decisive.
Proceedings of the National Academy of Sciences,
108, 18307–18312.
Schaller, M., Miller, G. E., Gervais, W. M., Yager, S., &
Chen, E. (2010). Mere visual perception of other peo-
ple’s disease symptoms facilitates a more aggressive
immune response. Psychological Science, 21,
649–652.
Schienle, A., Stark, R., & Vaitl, D. (2001). Evaluative
conditioning: A possible explanation for the acquisition
of disgust responses? Learning and Motivation, 32(1),
65–83.
Sell, A. (2005). Regulating welfare tradeoff ratios: Three
tests of an evolutionary–computational model of
human anger. Dissertation Abstracts International.
Section B: The Sciences and Engineering,66(8-B),
1–244.
Sell, A. (2011). The recalibrational theory and violent
anger. Aggression and Violent Behavior, 16(5),
381–389.
Sell, A., Cosmides, L., Tooby, J., Sznycer, D., von Rueden,
C., & Gurven, M. (2009a). Human adaptations for the
visual assessment of strength and fighting ability from
the body and face. Proceedings of the Royal Society of
London B: Biological Sciences, 276(1656), 575–584.
Sell, A., Tooby, J., & Cosmides, L. (2009b). Formidability
and the logic of human anger. Proceedings of the
National Academy of Sciences, 106(35), 15073–15078.
Sell, A., Cosmides, L., & Tooby, J. (2014). The human
anger face evolved to enhance cues of strength. Evolu-
tion and Human Behavior, 35(5), 425–429.
Sell, A., Cosmides, L., Tooby, J., Lim, J., Krauss, A.,
Feldman, A., Rascanu, R., Al-Shawaf, L., Sznycer,
D., & Sugiyama, L. (under review). The grammar of
anger: Mapping the computational architecture of a
recalibrational emotion.
Sznycer, D., Takemura, K., Delton, A. W., Sato, K., Rob-
ertson, T., Cosmides, L., & Tooby, J. (2012). Cross-
cultural differences and similarities in proneness to
shame: An adaptationist and ecological approach. Evo-
lutionary Psychology, 10(2), 352–370.
Sznycer, D., Schniter, E., Tooby, J., & Cosmides,
L. (2015). Regulatory adaptations for delivering infor-
mation: The case of confession. Evolution and Human
Behavior, 36(1), 44–51.
Sznycer, D., Tooby, J., Cosmides, L., Porat, R., Shalvi, S.,
& Halperin, E. (2016). Shame closely tracks the threat
Evolutionary Psychology and the Emotions 9

of devaluation by others, even across cultures. Pro-
ceedings of the National Academy of Sciences,
2625–2630.
Sznycer, D., Al-Shawaf, L., Bereby-Meyer, Y., Curry,
O. S., De Smet, D., Ermer, E., Kim, S., Kim, S., Li,
N. P., Lopez Seal, M. F., McClung, J., O, J., Ohtsubo,
Y., Quillien, T., Schaub, M., Sell, A., van Leeuwen, F.,
Cosmides, L., & Tooby, J. (2017). Cross-cultural regu-
larities in the cognitive architecture of pride. Proceed-
ings of the National Academy of Sciences, 114(8),
1874–1879.
Tangney, J. P., Miller, R. S., Flicker, L., & Barlow, D. H.
(1996). Are shame, guilt, and embarrassment distinct
emotions? Journal of Personality and Social Psychol-
ogy, 70, 1256–1269.
Tooby, J., & Cosmides, L. (1990). The past explains the
present: Emotional adaptations and the structure of
ancestral environments. Ethology and Sociobiology,
11, 375–424.
Tooby, J., & Cosmides, L. (1992). The psychological foun-
dations of culture. In The adapted mind: Evolutionary
psychology and the generation of culture (pp. 19–136).
Oxford: Oxford University Press.
Tooby, J., & Cosmides, L. (2008). The evolutionary psy-
chology of the emotions and their relationship to inter-
nal regulatory variables. In M. Lewis, J. M. Haviland-
Jones, & L. F. Barrett (Eds.), Handbook of emotions
(3rd ed., pp. 114–137). New York: Guilford.
Tracy, J. L., & Robins, R. W. (2007). Emerging insights
into the nature and function of pride. Current Direc-
tions in Psychological Science, 16(3), 147–150.
Tracy, J. L., Shariff, A. F., & Cheng, J. T. (2010).
A naturalist’s view of pride. Emotion Review, 2,
163–177.
Tybur, J. M., Lieberman, D., & Griskevicius, V. (2009).
Microbes, mating, and morality: Individual differences
in three functional domains of disgust. Journal of Per-
sonality and Social Psychology, 97(1), 103–122.
Tybur, J. M., Bryan, A. D., Magnan, R. E., & Hooper,
A. E. C. (2011). Smells like safe sex olfactory pathogen
primes increase intentions to use condoms. Psycholog-
ical Science, 22(4), 478–480.
Vrana, S. R. (1993). The psychophysiology of disgust:
Differentiating negative emotional contexts with facial
EMG. Psychophysiology, 30(3), 279–286.
Williams, G. C. (1966). Adaptation and natural selection.
Princeton: Princeton University Press.
10 Evolutionary Psychology and the Emotions