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Human Emotions: An Evolutionary Psychological Perspective

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DOI: 10.1177/1754073914565518
Abstract
Evolutionary approaches to the emotions have traditionally focused on a subset of emotions that are shared with other species, characterized by distinct signals, and designed to solve a few key adaptive problems. By contrast, an evolutionary psychological approach (a) broadens the range of adaptive problems emotions have evolved to solve, (b) includes emotions that lack distinctive signals and are unique to humans, and (c) synthesizes an evolutionary approach with an information-processing perspective. On this view, emotions are superordinate mechanisms that evolved to coordinate the activity of other programs in the solution of adaptive problems. We illustrate the heuristic value of this approach by furnishing novel hypotheses for disgust and sexual arousal and highlighting unexplored areas of research.
Emotion Review
Vol. 8, No. 2 (April 2016) 173 –186
© The Author(s) 2015
ISSN 1754-0739
DOI: 10.1177/1754073914565518
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Beginning with Darwin (1872/2009), evolutionary approaches
to the emotions have focused on a delimited subset of psycho-
logical phenomena. They have largely centered on emotions
that solve a subset of adaptive problems, carry distinctive uni-
versal signals, are universally recognized by conspecifics, and
are present in other primates (e.g., Ekman, 1973, 1992; Ekman
& Cordaro, 2011). This approach is empirically fruitful and
has been of great scientific import - Ekman and colleagues’
theoretical and empirical work has been responsible for
immense progress in emotions research over the last several
decades. Nonetheless, for three reasons, the principles of mod-
ern evolutionary psychology suggest that it is too narrow in
scope for a comprehensive theory of the emotions.
First, it places unnecessary emphasis on the subset of emo-
tions that evolved to serve a signaling or communicative func-
tion. In fact, many evolved emotions have no distinct facial
expression (e.g., sexual jealousy; Buss, 2013), or do not appear
to serve a signaling function at all (e.g., sexual regret; Galperin
et al., 2013). We discuss why we expect many evolved emotions
not to have discernible outward signals.
Second, we suggest that emotions have evolved to solve a
much wider range of adaptive problems than typically considered,
even by evolutionary theorists. Differential reproductive success,
not differential survival success, is the “engine” of evolution by
selection. Consequently, emotions may have evolved to solve
adaptive problems in a broad range of domains tributary to repro-
ductive success. This diverse range includes sexual consumma-
tion, intrasexual mate competition, mate retention, mate poaching,
hierarchy negotiation, losses of status, gains of status, punishment
of coalitional free-riders, retribution for social cost infliction, stra-
tegic interference, kin protection, kin investment, food acquisition,
and others.
Third, historically valuable evolutionary approaches have
yet to incorporate the most important modern novel insight into
the evolutionary functions of emotions: that they are hypothe-
sized to have evolved as superordinate mechanisms responsible
for coordinating suites of other information-processing pro-
grams, including those of attention, perception, memory, cate-
gorization, learning, and energy allocation, as well as the more
typically considered elements of physiology and manifest
behavior (Cosmides & Tooby, 2000; Tooby & Cosmides, 2008).
This theoretical approach to the emotions has several impor-
tant consequences. First, it integrates the study of emotion with
domains of psychology such as perception, attention, and learn-
Human Emotions: An Evolutionary
Psychological Perspective
Laith Al-Shawaf
Daniel Conroy-Beam
Kelly Asao
David M. Buss
Department of Psychology, The University of Texas at Austin, USA
Abstract
Evolutionary approaches to the emotions have traditionally focused on a subset of emotions that are shared with other species,
characterized by distinct signals, and designed to solve a few key adaptive problems. By contrast, an evolutionary psychological
approach (a) broadens the range of adaptive problems emotions have evolved to solve, (b) includes emotions that lack distinctive
signals and are unique to humans, and (c) synthesizes an evolutionary approach with an information-processing perspective. On
this view, emotions are superordinate mechanisms that evolved to coordinate the activity of other programs in the solution of
adaptive problems. We illustrate the heuristic value of this approach by furnishing novel hypotheses for disgust and sexual arousal
and highlighting unexplored areas of research.
Keywords
basic emotion, coordinating mechanisms, disgust, emotion, evolutionary psychology, sexual arousal, superordinate mechanisms
Corresponding author: Laith Al-Shawaf, Psychology Department, The University of Texas at Austin, 1 University Station A8000, Austin, TX 78712, USA.
Email: laith.alshawaf@mail.utexas.edu
565518EMR0010.1177/1754073914565518Emotion ReviewAl-Shawaf et al. Evolutionary Psychology of the Emotions
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174 Emotion Review Vol. 8 No. 2
ing, highlighting the principle that emotions pervade nearly
every aspect of psychological functioning. Second, it suggests a
dramatic expansion of the number of adaptive problems emo-
tions evolved to solve, and consequently calls for a theory of
emotions considerably more complex and nuanced than prior
evolutionary theories. Third, it provides a powerful hypothesis-
generation heuristic based on task analyses (Marr, 1982) of the
psychological procedures that must occur in order for an organ-
ism to successfully solve an adaptive problem. And fourth, it
opens new lines of empirical research, ranging from sexual
arousal toward mates exhibiting cues of reproductive value to
the vengeance that motivates ostracism of a coalitional violator.
An evolutionary psychological approach places emotions at the
center of psychological functioning.
Part I. A Modern Evolutionary Psychological
Perspective on the Emotions
Cosmides and Tooby (2000; Tooby & Cosmides, 2008) present
an evolutionary psychological view of the emotions that we
develop and expand upon here. We concur with many aspects of
pioneering evolutionary perspectives on the emotions (Ekman,
1992, 1999; Izard, 1993; Lang, 1995, 2010; Lazarus, 1991;
Nesse, 1990; Oatley & Johnson-Laird, 1987; Plutchik, 1980,
1991; Tomkins, 1984), but our approach differs in several
important respects.
Adaptations Are Solutions to Adaptive
Problems
Organisms face challenges in all domains of life. An adaptive
problem is any challenge—such as avoiding predators, secur-
ing nutritious food, gaining access to mates, resisting infection,
or rearing offspring—that impacts an organism’s chances of
survival or reproduction (Buss, 2012; Cosmides & Tooby,
1995). Natural selection produces solutions to these challenges
called adaptations. An adaptation is a structure or mechanism
that evolved because it helped its bearers solve a problem that
recurrently impacted fitness (roughly, relative reproductive
success) during the evolution of a species. These adaptations
can be found in psychology, physiology, and morphology, and
each is tailored to the solution of a specific adaptive problem
(Confer et al., 2010; Tooby & Cosmides, 1992; Williams,
1966).
Adaptive problems can be structurally complex, often consist-
ing of several subtasks requiring different computational solu-
tions (Kurzban, 2012; Marr, 1982; Tooby & Cosmides, 2005).
Even the seemingly simple problem of predator avoidance
requires the coordination of many distinct processes. Organisms
typically must (a) focus attention on the predator, (b) down-regu-
late attention to distracting stimuli, (c) suppress motivations rel-
evant to other adaptive problems (e.g., approaching mates), (d)
determine whether one has been spotted, (e) identify whether
there are other predators in the environment, (f) accurately recall
the spatial layout of the environment if it is already familiar, (g)
select a propitious escape route, (h) move in the determined route,
and (i) shunt energy away from nonessential physiological pro-
cesses (e.g., immune function) and toward those necessary for
escape (e.g., catabolism in the muscles; Bracha, 2004; Lima &
Dill, 1990; Marks, 1987; Marks & Nesse, 1994). Organisms able
to complete the many subtasks of an adaptive problem would
have outcompeted conspecifics unable to do so. Organisms capa-
ble of producing more efficient solutions would have outcom-
peted those less adept at solving such problems.
The Coordination Problem
Successfully solving an adaptive problem’s subtasks requires
the coordinated operation of many distinct adaptations. For
example, programs responsible for fixing attention on a preda-
tor may need to be coordinated with programs responsible for
suppressing digestion and immune function, down-regulating
mating motivations, and motivating escape behavior. The coor-
dination of these distinct mechanisms presents a new adaptive
problem, one unlike any other. This is the coordination problem.
Haphazard and unsystematic program coordination would be
maladaptive and extraordinarily unlikely to lead to the solution
of an adaptive problem. Uncoordinated program activation can
lead to fatal errors such as continuing to forage for food instead
of escaping a deadly predator. We propose that adaptations
require coordination for at least two reasons: first, adaptations
sometimes have conflicting outputs, and second, the efficiency
of one adaptation’s operation often depends on the operation of
others. We consider these in turn.
The problem of conflicting outputs. One reason that adap-
tations require coordination is that they often have opposing
outputs. Consider approach and avoidance motivations. Some
mechanisms motivate avoidance of infectious stimuli. For
example, disgust promotes avoidance of pathogen-laden sub-
stances such as open sores (Tybur, Lieberman, Kurzban, &
DeScioli, 2012). Other adaptations motivate approach, such as
those promoting pursuit of desirable mates (Sugiyama, 2005).
The activation of two conflicting adaptations may lead to mala-
daptive behavior: indecision, or perhaps intermittent approach
and avoidance. For example, a first glance at an attractive
potential mate might motivate approach, but up close, infected
sores might become unmistakably apparent. Conflicting moti-
vations must therefore be coordinated to result in a successful
solution to the adaptive problem at hand. The problem of con-
flicting outputs can be solved by mechanisms that control the
activation and deactivation of programs so as to minimize inter-
ference with effective problem solving.
The concatenation problem. The second reason that adapta-
tions need coordination is the concatenation problem: many
adaptive problems require that subtasks be completed in a spe-
cific order. In responding to an apparent act of altruism, for
example, the emotion of gratitude should up-regulate the value
placed on a benefactor’s welfare only after determining that the
helpful act was intentional. Mechanism coordination is also
important when the effectiveness of one mechanism depends on
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Al-Shawaf et al. Evolutionary Psychology of the Emotions 175
the operation of another. For example, mechanisms that promote
fleeing from a predator can produce their output more effectively
when other adaptations are concurrently mobilizing energy
toward the muscles. Uncoordinated responses to an adaptive
problem are inefficient at best, disastrously costly at worst, and
frequently unable to solve the adaptive problem at hand.
Emotions as Evolved Solutions to Problems of
Multimechanism Coordination
This approach views the emotions as superordinate mecha-
nisms—programs designed to regulate the activity of other pro-
grams in the solution of an adaptive problem. Such mechanisms
would be designed to process, as input, environmental or
somatic cues that the organism is facing a particular adaptive
problem. Once activated, the emotion’s task is to orchestrate a
number of different programs in the service of solving the adap-
tive problem. To produce an effective solution, an emotion must
(a) activate the right programs, (b) deactivate conflicting pro-
grams, (c) adjust program thresholds to meet task demands, and
(d) manage the sequence and duration of program activation as
well as the point of program termination.
The details of these tasks will differ as a function of context.
For example, a food acquisition problem may call for different
solutions depending on the food’s distance from camp and
whether one is foraging alone or with others. Emotions must
take this contextual variability into account and tailor their acti-
vation patterns to meet the demands of the problem at hand.
Consequently, a given emotion is not expected to invariably
activate the same programs across all contexts (see Tracy, 2014,
for a similar argument about variability in emotion output).
This task is computationally complex and critical to fitness.
Arguably the best solution to the coordination problem—the
most efficient and least likely to lead to errors in function-
ing—comes from coordinating mechanisms specifically
designed for this purpose. Coordinating mechanisms can be
thought of as analogous to regulatory genes, whose function is
to regulate the expression and operation of other genes
(Davidson & Erwin, 2006). Indeed, a variety of fields ranging
from genetics to business to politics exhibit evidence of coor-
dinating bodies expressly designed to manage the activity of
other entities (e.g., Chenhall, 2003). We suggest that this
points to the utility of the superordinate mechanism region of
“design space” (Dennett, 1996).
How Might Emotions Solve the Coordination
Problem?
In principle, an emotion can coordinate any of the mechanisms
in an organism’s body. These include cognitive, perceptual, and
physiological programs, as well as manifest behavior. Emotions
can be thought of as organismic “modes of operation” (Cosmides
& Tooby, 2000; LeDoux, 2003; Oatley & Johnson-Laird, 1987).
On this view, each emotion represents a distinct mode of opera-
tion, characterized by a different profile of cognitive, physiolog-
ical, and behavioral activity.
Cosmides and Tooby (2000) have suggested that the emo-
tions regulate at least 14 types of programs, worth listing again
here: (a) perceptual mechanisms, (b) attention, (c) memory, (d)
goals, (e) motivational priorities, (f) information-gathering
adaptations, (g) categorization or imposed conceptual frame-
works, (h) specialized cognitive inference, (i) communication
and expression, (j) learning, (k) reflexes, (l) mood, energy
level, and effort allocation, (m) physiology, and (n) behavior.
Although no emotion has been the subject of research explor-
ing all of these programs, work on the emotion of sexual jeal-
ousy has made inroads into many (Buss, 2013). For example,
hypothesized sex-differentiated design features of jealousy have
been discovered in studies of attention, speed of processing, and
spontaneous recall (Schützwohl, 2006; Schützwohl & Koch,
2004), information-gathering (Kuhle, 2011), probabilistic infer-
ences (Andrews et al., 2008), physiology (Buss, Larsen, Westen,
& Semmelroth, 1992), and manifest behavior (Buss, 1988;
Kuhle, Smedley, & Schmitt, 2009).
Not every emotion will coordinate all of these programs,
nor will each instance of an emotion activate the same pro-
grams. Fear may activate escape behavior in some contexts
(e.g., a crocodile that can be outrun on dry land) and not in
others (e.g., an adversary who can be overpowered in a fight).
Different environmental circumstances present different
demands. Consequently, the idea of emotions as program
coordinators does not imply that component programs must be
strongly correlated. Evidence showing, for example, that the
components of surprise are not strongly correlated (Reisenzein,
2000) or that smiling exhibits weak–moderate correlations
with certain positive emotions (Reisenzein, Studtmann, &
Horstmann, 2013) is perfectly compatible with our perspective
on the emotions.
Our framework can be used to systematically generate novel
hypotheses using the heuristic tool of task analysis (Marr,
1982). A task analysis for an emotion consists of four key ques-
tions: (1) what adaptive problem, if any, did this emotion evolve
to solve? (2) which subtasks must be solved in the solution of
this adaptive problem? (3) which information-processing pro-
grams are capable of carrying out these subtasks? and (4) how
should these programs be coordinated to facilitate the solution
of these subtasks?
Part II. What Does an Evolutionary
Psychological Perspective Contribute to Our
Understanding of the Emotions?
Nonarbitrary Criteria for Classifying Emotions
The perspective we present here has implications for how emo-
tions are classified. Current perspectives classify emotions as
“basic” or fundamental on the basis of criteria that are mis-
guided from a modern evolutionary perspective (Buss, 2014;
Confer et al., 2010; Tooby & Cosmides, 2008). An evolutionary
psychological perspective suggests conceptual shifts in three
areas: diversity of adaptive problems, distinctive universal sig-
nals, and emotions in nonhuman species.
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176 Emotion Review Vol. 8 No. 2
Emotions solve a wide variety of adaptive problems. Evolu-
tionary approaches have traditionally focused on a limited sub-
set of emotions, many of which solve problems of survival.
Fear, anger, and disgust, for example, evolved to solve survival
problems and are widely regarded as “basic” (Ekman, 1992,
1999; Ekman & Cordaro, 2011; Lazarus, 1991; Plutchik, 1980).
By contrast, evolutionary approaches in the field have largely
neglected emotions such as romantic love (e.g., Buss, 2003;
Hazan & Shaver, 1987), sexual arousal (Symons, 1979, 1995),
sexual jealousy (Buss, 2000; Wilson & Daly, 1992), and paren-
tal love (Daly & Wilson, 1998). There now exists evidence for
the universality of these emotions, pointing to evolved functions
related to challenges of mating, reproduction, and childrearing.
There is no conceptually principled reason why they should be
considered less fundamental than survival-oriented emotions
such as disgust, anger, or fear. In our view, a) the defining crite-
rion for an evolved emotion is whether it coordinates programs
in the solution of an adaptive problem and (b) emotions can
evolve to solve adaptive problems in any domain, including
mate selection, mate retention, parenting, food acquisition, nav-
igating status hierarchies, and many others.
The emphasis on survival in the evolutionary emotions litera-
ture may reflect an earlier view that emphasized natural selection
to the exclusion of sexual selection, and survival to the exclusion
of reproduction. By contrast, a modern evolutionary perspective
emphasizes that the bottom line of evolution by selection is dif-
ferential reproductive success. Evolutionary biologists recognize
that survival is critical, but only to the extent that it is tributary to
reproductive success. When the two conflict, reproduction inevi-
tably trumps survival (Dawkins, 1976; Hamilton, 1964;
Williams, 1966). Adapta tions that promote a n organi sm’s repr o-
duction at a cost to its survival are ubiquitous in the animal king-
dom (Alcock, 2009; Buss, 2012; Dawkins, 1976). Examples
include the brilliant plumage of peacocks (Darwin, 1871), sui-
cidal mating among male honeybees (Woyciechowski, Kabat, &
Kroll, 1994), and adaptations for elevated testosterone that pro-
mote success in mate competition at an on-average cost to male
survival (Trivers, 1985; Williams & Nesse, 1991). Adaptations
that promote an organism’s survival to the detriment of its over-
all reproductive success, on the other hand, are entirely absent
(Alcock, 2009; Dawkins, 1976, 1999).
This suggests a class of emotions that are critical but have
not received much evolutionary attention: those that evolved to
solve a broad range of adaptive problems tributary to reproduc-
tive success. This class of emotions includes states such as envy
(e.g., DelPriore, Hill, & Buss, 2012), pride (e.g., Tracy &
Robins, 2007), embarrassment (e.g., Keltner & Buswell, 1996),
shame (e.g., Tracy & Matsumoto, 2008), guilt (e.g., Tangney &
Dearing, 2003), sexual jealousy (e.g., Buss et al., 1992), sexual
arousal (Symons, 1979, 1995), sexual regret (Galperin et al.,
2013), sexual disgust (Tybur et al., 2012), romantic or passion-
ate love (e.g., Hazan & Shaver, 1987), and parental love (e.g.,
Daly & Wilson, 1998). For example, sexual arousal motivates
the pursuit of intercourse, sexual jealousy protects a mateship
from interlopers, and parental love motivates behaviors that
contribute to the health and success of offspring (Daly & Wilson,
1998; Hatfield & Sprecher, 1986; Symons, 1979). On our view,
these emotions are all part of our fundamental, evolved emo-
tional architecture.
Distinctive universal signals. Historically seminal evolu-
tionary perspectives typically maintain that basic emotions
have distinctive universal signals (e.g., Ekman, 1992, 1999;
Ekman & Cordaro, 2011). This theoretical position, which
has been highly empirically fruitful (Ekman, 1992, 1999;
Ekman & Friesen, 1971, 1986), may be conceptually rooted
in Darwin’s initial work on the emotions. In his classic book
on emotions, Darwin’s central goal was to document continu-
ity in emotion expression between humans and nonhuman
animals—an important focus at the time, but one that came at
the expense of the critical question of adaptive function (Dar-
win, 1872/2009).
By contrast, a contemporary focus on evolved function high-
lights why the “distinctive signal” criterion is too restrictive.
Whether an emotion carries with it a distinctive communication
display depends on whether its evolved function includes sign-
aling to others. For example, one hypothesized function of
anger is to cause other individuals to back down in social con-
flict, which requires a manifest signal. In contrast, one hypoth-
esized function of sexual jealousy is to motivate vigilance of a
partner’s behavior with potential interlopers, which typically
does not require a distinctive signal.
Ultimately, the evolution of a signal is contingent on the
ancestral costs and benefits of conveying that emotional state to
others. During the evolution of a particular emotion, if the costs
of advertising were high and benefits were low, we would not
expect a signal to evolve.
Emotions researchers may benefit from evolutionary biolo-
gists’ distinction between signals and cues. A signal is observa-
ble output whose function is conveying information to, or
influencing, another organism (Krebs & Dawkins, 1984). By
contrast, a cue is observable output that did not evolve to com-
municate information to others. For instance, a fear expression
may have evolved to signal danger to allies. By contrast, fear-
induced sweaty palms or brow may be visible by-products of
physiological arousal that did not evolve to transmit informa-
tion (e.g., Bradbury & Veherncamp, 1998; Maynard Smith &
Harper, 2003, Shariff & Tracy, 2011). An important task for
evolutionary emotions research is therefore to accurately distin-
guish signals and cues. Both may be universal and both impor-
tant, but the distinction is critical in understanding the evolved
function of each emotion.
A modern evolutionary perspective therefore suggests that
the “distinctive universal signal” criterion is needlessly restric-
tive. Instead, some evolved emotions include distinctive sig-
nals, whereas others, such as regret, lack signals altogether.
Evidence that certain emotions may not have signals or may not
be universally recognizable (e.g., Nelson & Russell, 2013) is
perfectly compatible with their status as evolved emotions.
Moreover, some emotions may exhibit context-dependent
signaling or context-dependent suppression of expression (see
Fridlund, 1997, for a similar discussion). For example, in some
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Al-Shawaf et al. Evolutionary Psychology of the Emotions 177
contexts a fear expression may signal a dangerous predator, but in
different circumstances, fear displays may be suppressed to con-
vey bravery. This important context-dependence reinforces the
key point that signaling is linked only to a limited subset of emo-
tions, and even with those, to limited contexts in which signaling
is central to evolved function. A modern evolutionary perspective
acknowledges the importance of signals, but does not support the
notion that a distinctive universal signal—or any signal for that
matter—is a necessary component of evolved emotions.
Cross-species evidence. It is often stipulated that basic emo-
tions must be present in nonhuman species (Ekman, 1992;
Ekman & Cordaro, 2011). Modern evolutionary theory suggests
a different conclusion. Each species has an emotional repertoire
appropriate for dealing with the adaptive problems it recurrently
faced during its evolution. When adaptive problems differ, spe-
cies evolve different solutions. Uniquely human adaptations are
no less basic than those shared with other species. Language and
the ability to accurately throw high-speed projectiles, for exam-
ple, are uniquely human adaptations that are properly regarded
as fundamental aspects of human nature (Pinker, 2007; Pinker
& Bloom, 1992; Roach, Venkadesan, Rainbow, & Lieberman,
2013). Pride, guilt, envy, embarrassment, and love are all exam-
ples of emotions that may have evolved uniquely in humans, or
at a minimum have uniquely human design features. These
evolved human emotions are just as fundamental as emotions
shared with other species.
Even emotions that are shared with other species are likely to
have unique design features in humans. For example, fear and
disgust are widespread across taxa, but differ from species to
species in the cues they take as input, the decision rules they
implement, and the outputs they produce. For instance, the mere
thought of having sex with one’s sibling is repulsive to most
humans (Haidt, 2001; Lieberman, Tooby, & Cosmides, 2007),
but presumably not to species with inbreeding mating systems
(e.g., Hamilton, 1967; Herre, 1985). Humans are revolted at the
prospect of feeding on a rotting deer carcass; scavengers such as
vultures are not. The key point is that an evolved human emotion
can be fundamental regardless of whether it is: (a) shared with
other species, but with many unique features, (b) shared with
other species, with few unique features, or (c) entirely unique to
our species.
In sum, we argue that three features of existing perspectives
require reformulating. Emotions can evolve to solve a broad
array of adaptive problems; do not need to include distinctive
signals; and can be unique to humans (see Table 1).
Beyond these classificatory emendations, an evolutionary
psychological approach makes two additional contributions:
highlighting the centrality of information processing and offer-
ing heuristic value in hypothesis generation.
The Centrality of Information Processing
An evolutionary psychological approach recognizes the impor-
tance of emotion outputs such as behavior—they are indispensa-
ble in the solution of real-life adaptive problems. But it places
outputs in context by situating them in the input-algorithm-
output chain. This approach leads to novel empirical predictions
about the nature of these outputs. In traditional evolutionary
approaches, for example, distinctive signals must be universal
and universally recognizable. The current approach suggests a
different conclusion: some outputs will indeed be universal, but
this isn’t necessary. In most cases, design at the information-
processing level will be universal, but manifest output need not
be. Some emotions may be designed for context-dependent out-
put: that is, designed to produce expressions that differ predict-
ably by context, or to produce signals only in a subset of contexts.
Stated differently, an evolutionary psychological perspective
suggests that universality and evidence of adaptive design are to
be found at the level of computational structure, not manifest
output (Buss, 1995; Symons, 1979; Tooby & Cosmides, 1992).
In the example of facial expressions, the critical question is not
whether the output (emotion expression) is universal or variable,
but rather whether the variability in output is underlain by uni-
formity in the information-processing procedures that produce
the output (e.g., see Tooby & Cosmides, 2005).
This characterization—universal at the information-processing
level, but variable at the manifest level—suggests an analogy
with language. All humans have species-typical language mecha-
nisms, but the particular language learned during ontogeny differs
from culture to culture (Pinker & Bloom, 1992), resulting in the
rich variability of the world’s approximately 6,800 living lan-
guages (Gordon, 2005).
Heuristic Value and Predictive Power
This perspective can be combined with task analysis of adaptive
problems (Marr, 1982) to yield a powerful framework for
hypothesis generation. This entails determining the adaptive
problem that an emotion evolved to solve and identifying the
psychological programs that are capable, in principle, of accom-
plishing the problem’s subtasks. Task analysis can also be used
to identify psychological procedures that would result in a bet-
ter or more efficient solution to the adaptive problem in ques-
tion. This process leads to a series of hypotheses about the
coordinating effects of an emotion on a variety of programs.
Some emotions coordinate a larger subset of programs
than others (Cosmides & Tooby, 2000; Tooby & Cosmides,
2008), and different contexts may reliably elicit different sub-
sets of programs. In principle, these context effects can be
predicted a priori, a theoretical strength that is largely absent
in existing evolutionary approaches. As an illustration, we use
this framework to generate a series of hypotheses for two dis-
tinct emotions.
Part III. Novel Hypotheses and New
Directions
Disgust: An Established Emotion
Disgust is a well-established emotion that primarily coordinates
psychological and physiological mechanisms in the service of
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178 Emotion Review Vol. 8 No. 2
avoiding parasitic infection (Curtis, Aunger, & Rabie, 2004;
Curtis, de Barra, & Aunger, 2011; Rozin, Haidt, & McCauley,
1993; Schaller & Duncan, 2007; Tybur et al., 2012). The particu-
lar pattern of mechanism activation differs by context and by
type of disgust (pathogen disgust, sexual disgust, or moral dis-
gust; see Tybur, Lieberman, & Griskevicius, 2009; Tybur et al.,
2012). We focus here on pathogen disgust, generating hypothe-
ses for how this emotion coordinates programs in the service of
avoiding infection.
Disgust memory. The effects of disgust on memory are
largely unknown, but this perspective furnishes testable hypoth-
eses. We propose that disgust mobilizes memory resources to
encode potentially infectious items, a device that may be useful
in avoiding future exposure. Individuals who appear infectious
may be encoded as such in memory, along with a tag marking
them as unsuitable social partners.
Research on “adaptive memory” shows that survival-related
adaptive problems trigger especially strong encoding, even out-
performing encoding techniques such as visual image genera-
tion (Nairne & Pandeirada, 2008; Nairne, Pandeirada, &
Thompson, 2008). We hypothesize that strong disgust activation
may have similarly powerful effects on memory, allocating
memory resources to information relevant to avoiding future
contamination.
Disgust information-gathering mechanisms. Acquiring infor-
mation can be vital in avoiding contamination. For example,
smelling a foul stench of unknown origin may prompt a search
for the source, enabling a person to identify whether the disgust-
ing object is alive or dead, mobile or immobile, close or distant
- different combinations of which should result in different
behavioral responses.
Disgust may regulate information gathering in other con-
texts as well. For example, seeing a gaping wound on your
child’s leg may activate a search to find out how far the
infection spreads, whether pus is visible, and whether the
wound is worsening over time. This information would have
been useful in reducing the risk of infection for self, off-
spring, and kin. More broadly, an evolutionary psychological
perspective calls attention to the ways in which emotions
adaptively regulate information-gathering mechanisms, a
topic that has thus far received little attention.
Communication and emotional expression. This approach
yields hypotheses about the expression of disgust, as well as
the ways that disgust might regulate communication in other
domains. Facial displays of disgust are hypothesized to
serve at least two distinct functions—self-protective and
communicative (Chapman, Kim, Susskind, & Anderson,
2009; Shariff & Tracy, 2011). We propose that the commu-
nicative value of disgust expressions vary as a function of
context: first, these displays should be less common when
individuals are alone. Second, parents and kin may produce
more pronounced displays if children are present and obser-
vational learning is important for appropriate disgust devel-
opment. Third, overt displays may be adaptively suppressed
in some contexts. For example, men may suppress disgust in
mating contexts because the intensity of one’s disgust reac-
tion reveals information about the quality of one’s immune
system (Fessler, Pillsworth, & Flamson, 2004). These are
precise, testable predictions about the context-dependence
of emotion display.
Disgust communication may also infuse gossip and competi-
tor derogation. In principle, gossip about disgusting individuals
conveys information that can be used to track people’s infection
risk and regulate social interactions accordingly. In the domain
of competitor derogation, Buss and Dedden (1990) found that
men and women derogate their intrasexual rivals in the areas of
hygiene (e.g., he mentioned that his rival never showered) and
disease (e.g., she mentioned that her rival had sexually transmit-
ted diseases).
Table 1. Differing evolutionary perspectives on the emotions.
Traditional evolutionary perspectives: Basic emotions Evolutionary psychological perspective: Superordinate mechanisms
(e.g., Ekman, 1973, 1992; Ekman & Cordaro, 2011) (Al-Shawaf et al, 2015 (this article); Cosmides & Tooby, 2000; Tooby & Cosmides, 2008)
There are 7 basic emotion families (disgust, fear, anger,
surprise, sadness, happiness, contempt).
Evolved emotions are larger in number, including emotions such as love, guilt, jealousy,
and gratitude. Each is defined according to its evolved function.
Focus on subset of adaptive problems. Heavier emphasis on
survival, weaker emphasis on reproductive success.
Focus on broad range of adaptive problems, including any that historically contributed
to reproductive success in any domain, directly or indirectly.
Present in other species, especially nonhuman primate
species.
May be (a) unique to humans, (b) shared with other species, or (c) shared with other
species but with uniquely human features.
Characterized by distinctive physiological activation (e.g.,
distinctive patterns of autonomic nervous system activity).
Evolved emotions are distinct if they are (a) designed to solve different adaptive
problems and (b) characterized by a different overall profile of psychological,
physiological, and behavioral activation.
Include a universal, universally recognizable signal. May (a) include a universal signal, (b) not include a signal at all, or (c) be designed for
context-dependent signaling.
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Al-Shawaf et al. Evolutionary Psychology of the Emotions 179
Specialized disgust inference mechanisms. Disgust acti-
vates specialized inference mechanisms that appear biased
toward false positives (Haselton & Buss, 2000; Haselton & Net-
tle, 2006). For example, humans are disgusted by noninfectious
conditions such as obesity and burn wounds (Park, Faulkner, &
Schaller, 2003), reluctant to eat fake excrement that they know
is really chocolate fudge (Rozin, Millman, & Nemeroff, 1986),
and require less evidence to infer that someone is sick than
healthy (Kurzban & Leary, 2001). We suggest that disgust may
activate other “paranoid” inferential biases as well (Haselton &
Nettle, 2006). To take just one example, knowing that a per-
son’s spouse is ill may lead us to treat that person as infectious
- even in the absence of sufficient evidence.
Disgust behavior. Disgust’s behavioral output must be
coordinated with changes in cognition and perception. Quickly
approaching pathogens, for example, require different motor
responses than immobile contaminants. Behavioral output also
depends on social factors. If the object of disgust is another
human, we expect people to downplay their avoidance because
of the social costs of shunning in-group members. Because
these costs are usually absent for contaminated foods or dead
animals, this suppression should be stronger in response to
humans than to other contaminant vectors. For similar reasons,
we expect greater effort to downplay disgust in response to indi-
viduals of high than low social status. An evolutionary psycho-
logical approach suggests that disgust regulates behavior in
context-specific ways—a richer conceptualization than the
broadband notion of “avoidance,” and one that yields a number
of testable context-specific hypotheses.
Disgust usually produces avoidance, but in some instances it
can motivate approaching and repelling or killing the source.
We suggest that such behavior will be more frequent when there
are vulnerable young kin in proximity, or when one is seeking to
impress mates or other social partners. Lastly, disgust appears to
motivate washing, cleaning, and other hygienic behavior
designed to reduce the likelihood of infection (Curtis, 2007;
Curtis, Cairncross, & Yonli, 2000; Fleischman et al., 2011).
Overriding disgust. In extreme cases, disgust may be sup-
pressed in order to solve a more pressing adaptive problem, such
as consummating a valuable mating opportunity or fending off
starvation. As starvation draws near, we expect a muted disgust
response to pathogenic foods that would otherwise trigger revul-
sion (Hoefling et al., 2009). This disgust suppression should be
specific to food. For example, mounting hunger should not affect
repulsion toward bodily effluvia or open sores.
Hunger-induced disgust suppression may partly explain
human willingness to eat repugnant food during dire circum-
stances. Consider the historical examples of “survival cannibal-
ism” during the 1972 Andes flight crash (Read, 1975), the Donner
Party disaster (Stewart, 1960), or the Nazi siege of Leningrad
(now St. Petersburg; Salisbury, 2009), in which people resorted to
eating human meat to fend off starvation. Eventually, when hun-
ger dissipates, this disgust suppression should lift, reactivating
previous disgust thresholds.
Sexual Arousal: An Unexplored Emotion
An evolutionary psychological approach promotes the inclusion
of states that are traditionally excluded from emotions research.
States such as hunger, pain, curiosity, and sexual arousal may
qualify as emotions as long as they evolved to coordinate other
programs in the solution of an adaptive problem (Cosmides &
Tooby, 2000; Tooby & Cosmides, 2008).
Despite its powerful effects on physiology, psychology,
and behavior, sexual arousal does not appear in previous tax-
onomies of basic emotions and is typically excluded from the
category of emotions altogether. In contrast, an evolutionary
psychological approach regards sexual arousal as a fundamen-
tal emotion that evolved to solve one of the most important
adaptive problems faced by sexually reproducing species—the
coordination of a number of disparate mechanisms in the ser-
vice of conception.
Before generating hypotheses, we address a theoretically
important sex difference. Among humans, the minimum obliga-
tory parental investment is greater for women than for men. In
species characterized by a sex difference in minimum parental
investment, the higher investing sex suffers more severe costs as
a consequence of injudicious mating decisions. Consequently,
easy-to-cross thresholds of sexual arousal are more costly for
women than for men (Symons, 1979). The theoretical optimum
for sexual arousal thresholds is higher for women than it is for
men. Selection therefore favors higher thresholds in women,
making men more sexually excitable in response to more mini-
mal stimulation. Once activated, however, the broad function of
sexual arousal is the same for both sexes: the orchestration of a
diverse array of programs in pursuit of sexual intercourse.
Motivational priorities. Sexual arousal can effect a profound
shift in motivational priorities, elevating sexual activity and
demoting goals such as obtaining food and avoiding pathogens.
For example, sexual arousal can inhibit disgust (Borg & de Jong,
2012; Fleischman, 2014; Stevenson, Case, & Oaten, 2011) and
promote interest in sex with partners one normally finds socially
inappropriate or even undesirable (Ariely & Loewenstein, 2006).
The strength of motivational reprioritization should depend
partly on the intensity of the emotion, with more intense sexual
arousal leading to more dramatic reprioritization.
Sexual arousal should calibrate the extent of motivational
shifts in response to cues that ancestrally were reliably predic-
tive of the costs and benefits of pursuing a particular sexual
goal. Sexual arousal should have a strong impact on motiva-
tional priorities when a potential partner is of high mate value,
the likelihood of a successful mating is high, the temporal prox-
imity to sexual consummation is close, and the costs of pursuing
such an opportunity are low. Because the costs of a missed
sexual opportunity are typically greater for men than for women
(Symons, 1979; Trivers, 1972), motivational shifting should be
stronger, on average, for men than for women, with notable
exceptions such as a rare opportunity for sex with a partner of
unusually high status (Greiling & Buss, 2000) or unusually
good genes (Gangestad & Thornhill, 2008).
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180 Emotion Review Vol. 8 No. 2
Sexual information-gathering adaptations. Sexual arousal
should activate a search for information relevant to the costs and
benefits of pursuing a potential mate. Such programs may seek
information on a potential mate’s relationship status and sexual
receptivity, as well as the presence or absence of mating rivals.
We expect information-gathering programs to seek data about
the potential partner’s preferred mating strategy, personality,
and values and interests.
The information sought and the way in which it is used will
differ across individuals and contexts, and this may be predict-
able in advance. For example, men who are interested in mate
poaching (Schmitt & Buss, 2001) may seek information about a
mated woman’s sexual fidelity. More monogamous individuals
may refrain from searching for this information in the first
place. More broadly, this framework calls attention to informa-
tion-gathering mechanisms as key components of evolved mat-
ing strategies.
Imposed conceptual frameworks and sexual categoriza-
tion. Emotions may regulate conceptual frameworks, compel-
ling division of the world into categories relevant for solving the
adaptive problem at hand. We propose that sexual arousal alters
conceptual frameworks, temporarily categorizing members of
the opposite sex as mateable or unmateable, and members of the
same sex as rivals, nonrivals, protectors, or enablers (“wing-
men” and “wingwomen”).
Sexual memory. Sexual arousal may regulate memory,
encoding the physical features of desirable mates and the cir-
cumstances that led to mating success. When accompanied by
long-term mating intentions, sexual arousal should strengthen
encoding of many types of information about the desired other,
including personal details revealed in the process of courtship.
This information can be useful in courtship and valuable in con-
veying commitment intent.
Memory effects should show sex-differentiated design fea-
tures. Circumstantial evidence for this hypothesis comes from
recent studies of sexual regret. Men recalled and regretted more
acts of sexual omission; women recalled and regretted acts of
sexual comission with inappropriate partners (e.g., men who
were unattractive or uninterested in commitment; Galperin et al.,
2013). Negatively valenced memories presumably provide
guidance about what to do in the present and future, and these
sex differences in regret should influence men and women’s
sexual decision-making.
Sexual learning. We propose that sexual arousal guides
learning to facilitate intercourse. In most species, frequent copu-
lation and sensitivity to sexual opportunities pays greater fitness
dividends to males than to females. Consequently, males should
learn to associate environmental cues linked with copulatory
opportunity more easily and more rapidly than females. Evidence
confirms this prediction—Japanese quail exhibit conditioned
sexual arousal to cues previously associated with copulatory
opportunity, an effect that is stronger among males than females
(Crawford, Holloway, & Domjan, 1993). Male quail even
respond sexually to arbitrary stimuli—such as artificial bright
orange feathers—that were previously associated with copulatory
opportunity in experiments (Domjan, O’Vary, & Greene, 1988).
Humans and rats show similar sexual conditioning effects, and
these are typically weaker or absent in females (Crawford et al.,
1993; Pfaus, Kippin, & Centeno, 2001).
The human male combination of low sexual arousal thresh-
olds and adaptively biased sexual learning mechanisms may
partly explain the greater prevalence of sexual fetishism among
males than females (Laws & Marshall, 1990; O’Donohue &
Plaud, 1994). Evidence shows that paired associations of erotic
stimuli with colored squares or women’s boots can lead men to
become sexually aroused in response to the squares or boots
alone (McConaghy, 1974; Rachman & Hodgson, 1968). We are
not, of course, claiming that these fetishes are adaptations; they
are likely functionless byproducts of the male combination of
easy-to-cross thresholds of sexual arousal and adaptively
biased sexual learning. In sum, an evolutionary psychological
perspective on sexual arousal points to the importance of sex-
differentiated learning mechanisms and reveals a pattern of
greater male sensitivity to copulatory opportunity and cues
associated with sex.
Specialized sexual inference mechanisms. We hypothesize
that sexual arousal activates the sexual overperception bias, the
adaptive male tendency to overestimate the sexual interest of
women displaying ambiguous cues such as a smile (Abbey,
1982; Abbey & Melby, 1986; Haselton & Buss, 2000; Haselton
& Nettle, 2006). By activating the overperception bias, sexual
arousal presumably motivates approach, minimizes missed
opportunities, and may even transform an initially uninterested
woman into an interested one (Haselton & Buss, 2009). Circum-
stantial support for this hypothesis comes from the experimental
finding that physically attractive women, known to preferen-
tially activate men’s sexual arousal, are especially likely to
evoke men’s sexual overperception bias (Perilloux, Easton, &
Buss, 2012).
The commitment skepticism bias refers to women’s tendency
to underestimate the commitment intent of potential mates
(Haselton & Buss, 2000; Haselton & Nettle, 2006). Commitment
skepticism evolved because of a large cost asymmetry in the
types of error a woman could make when inferring a man’s com-
mitment intent: underestimating commitment may result in
wasted time, but overestimating commitment can lead to decep-
tion and exploitation. The latter error can impose the severe fit-
ness costs associated with being impregnated and left to rear
offspring without the provisioning of a committed mate.
Counterintuitively, because the men who are most sexually attrac-
tive to women are also the men who are most unfaithful and least
likely to commit (Buss, 2003; Gangestad & Thornhill, 2008),
sexual arousal may initially amplify the commitment skepticism
bias—a testable prediction that awaits future research.
The orgasmic dissolution hypothesis. Orgasm is an impor-
tant part of the sexual and emotional lives of both sexes. Among
men, orgasm is necessary for conception and indicates the end
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of the primary adaptive problem that sexual arousal is designed
to solve. The same is not true of orgasm in women.
Consequently, this event should have sex-differentiated
effects on sexual arousal. Among men, but not women, orgasm
should lead to a dramatic and precipitous decline in arousal.
And because sexual arousal coordinates the mechanisms dis-
cussed above, orgasm should suddenly terminate these coordi-
nating effects—but again only among men.
We call this the orgasmic dissolution hypothesis because in
males, orgasm suddenly “dissolves” the coordination effects of
sexual arousal. This hypothesis predicts that orgasm should
have the following sudden effects for men but not women:
Prediction 1: Reverse the narrowing of attentional focus,
broadening it back out again.
Prediction 2: Cause memories of other duties and responsi-
bilities to become accessible again.
Prediction 3: Reorder priorities so that previously suppressed
motivations to eat, further one’s career, or avoid pathogens
come online again.
Prediction 4: Deactivate specialized inference mechanisms
activated by sexual arousal (e.g., the sexual overperception
bias).
Prediction 5: Deactivate sexual categorization (e.g., enablers
vs. obstacles).
Prediction 6: Down-regulate information-gathering mecha-
nisms geared toward intercourse.
Prediction 7: Previously suppressed physiological processes
can come online again (e.g., digestion, cell repair).
Prediction 8: Previously suppressed emotions can be reacti-
vated (e.g., disgust, worry).
These effects must come after orgasm, not before—an exam-
ple of the concatenation problem. We expect these effects to be
strikingly sex-differentiated: Among men, they should be pro-
found and near immediate. Among women, they should be
weaker and more gradual, and in some cases entirely absent.
These predictions can be tested experimentally. Participants
can complete psychological assessments while stimulating
themselves sexually (e.g., Strassberg, Mahoney, Schaugaard, &
Hale, 1990). This can be done at progressive stages of arousal
leading up to orgasm and immediately thereafter. The orgasmic
dissolution hypothesis proposes that men, much more than
women, will show a sudden and profound shift from preorgasm
to postorgasm on psychological mechanisms ranging from
motivational priorities to conceptual categorization to attention.
Context-dependent effects of orgasm. Orgasmic dissolution
should be context-dependent. Men pursuing long-term mating
are interested in forging or maintaining a committed relationship
that persists over time. For them, sexual intercourse solves more
than just the problem of conception; it also contributes to the
solution of adaptive problems related to satisfying one’s mate
and forming a stable bond together. The “dissolving” effects of
orgasm should therefore be less dramatic for men seeking com-
mitment. These men may experience a more gradual return to
baseline after orgasm. Committed men may also see sexual
arousal give way to a positive emotion such as affection, whereas
men oriented toward short-term mating appear more likely to
experience a negative affective shift after orgasm (see Haselton
& Buss, 2001).1
The Science of Testing Evolution-Based
Emotion Hypotheses
The collection of hypotheses generated by this framework raises
the question of how to best test hypotheses and interpret results.
The answer depends on the level at which the question is posed.
We h av e p re se nt ed i de as at t hr ee di ff er en t le ve ls i n t he h ie ra rc hi -
cal structure of evolutionary psychology (Buss, 1995): (a) the
“middle-level” theory of emotions as superordinate mechanisms,
(b) broad hypotheses about the function of each individual emo-
tion, and (c) numerous specific hypotheses about the effects of
each emotion on a variety of programs in the mind and body.
The last of these three levels is the most straightforward.
Specific hypotheses about the effects of a given emotion on
memory or attention are used to generate predictions, which are
tested in empirical studies. The results are used to interpret the
truth or falsity of the hypothesis in the standard scientific man-
ner. If a single hypothesis generates multiple predictions, some
may be verified and others falsified. The hypothesis, like all
hypotheses in psychology, must be evaluated according to the
cumulative weight of the evidence.
Broad hypotheses and middle-level theories are evaluated in
much the same way, but two additional points are worth men-
tioning. First, a hypothesis at a particular level of the hierarchy
is evaluated on the basis of the empirical harvest it yields at
more specific levels. For example, the broad hypothesis that dis-
gust coordinates programs in the service of avoiding infection
must be evaluated on the basis of the specific hypotheses it
yields about physiology, memory, attention, and other pro-
cesses. Evidence against the memory hypotheses may falsify
those memory hypotheses, but does not immediately falsify the
broad disgust hypothesis. Instead, this broad hypothesis is falsi-
fied when the cumulative evidence weighs against the multiple
specific hypotheses it yields. The same principle applies to the
middle-level theory of the emotions as superordinate mecha-
nisms. The theory of emotions as superordinate mechanisms is
best evaluated in light of all the broad hypotheses that fall under
its umbrella: disgust, anger, love, sexual arousal, happiness,
gratitude, and so on.
Second, there is no single, definitive evolutionary hypothe-
sis for the effects of disgust on memory, or for the effects of
happiness on information gathering, or for anything else. There
are usually multiple, competing evolutionary hypotheses for
the same phenomenon (e.g., Buss, 1995; Symons, 1979;
Tinbergen, 1963). A broad hypothesis, such as the idea that dis-
gust coordinates programs in the service of avoiding infection,
can be used to generate multiple competing specific hypotheses
about how disgust regulates memory. The specific hypotheses
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182 Emotion Review Vol. 8 No. 2
we propose in this article are by no means the only viable evo-
lutionary hypotheses.
The existence of competing evolutionary hypotheses cau-
tions strongly against assuming a particular hypothesis to be
“the” evolutionary hypothesis, and then equating the verifica-
tion or falsification of that particular hypothesis with the verifi-
cation or falsification of the entire theory on which it is based.
The Role of the Emotions in Conflict and
Manipulation
Conflict and manipulation are ripe but relatively untapped areas
in emotions research. The fact that different individuals have
different fitness interests suggests that social conflict will be
ubiquitous. We expect the emotions to play a crucial role in
these battlegrounds.
Animal signals are best conceptualized as adaptations
designed to influence and manipulate other organisms, rather
than as purely veridical messages (Dawkins & Krebs, 1978;
Krebs & Dawkins, 1984). Emotions can be used to influence or
manipulate others in at least two ways: (a) exploitatively induc-
ing an emotion and (b) communicating false emotion. A wide
array of emotions may be amenable to such tactics, but here we
consider guilt, anger, and empathy as examples.
The more valuable another individual is to oneself, the
more weight one places on that person’s welfare. This relative
weighting of another individual’s welfare to one’s own is
called a welfare trade-off ratio. This internal regulatory vari-
able is hypothesized to play a central role in navigating social
relationships. It is also hypothesized to be critical to “recali-
brational” emotions such as guilt, anger, gratitude, and shame
(Cosmides & Tooby, 2000; Sell, Tooby, & Cosmides, 2009).
For example, evidence suggests that one evolved function of
anger is to induce another individual to recalibrate his welfare
trade-off ratio: feeling slighted motivates an angry response
designed to impel the slighter to recalibrate his welfare trade-
off ratio and treat the victim more favorably in the future (Sell
et al., 2009).
Guilt, too, appears to be a recalibrational emotion. Guilt
functions to recalibrate one’s own welfare trade-off ratio toward
another individual, leading to greater valuation of the other indi-
vidual and better treatment in the future (Tooby & Cosmides,
1990, 2008).
Exploitative induction of emotions. The existence of guilt
suggests the possibility of deliberately inducing guilt in order to
manipulate someone into recalibrating his or her welfare trade-
off ratio. The empirical discovery of guilt induction as a mate
retention tactic (e.g., “I pretended to be mad so that my partner
would feel guilty”; Buss, 1988) may be an instance of this form
of manipulation. More generally, the existence of emotions that
function to recalibrate welfare trade-off ratios renders organ-
isms vulnerable to exploitation.
The evolution of an exploitative strategy, however, selects
for coevolved defenses against exploitation, and this often
results in an antagonistic coevolutionary arms race (Dawkins &
Krebs, 1979; Ridley, 1993). Humans likely have mechanisms
designed to detect and defend against exploitative induction of
emotions.
Exploitative induction of one emotion is sometimes achieva-
ble through the exaggerated display of another. For example,
pity and empathy can be exploitatively induced through exagger-
ated displays of sadness. The very existence of empathy mecha-
nisms renders these mechanisms vulnerable to exploitation,
leading selection to favor the disingenuous amplification of sad-
ness displays (within credible limits). In this way, even if an
emotion such as sadness initially evolved for non-exploitative
purposes, selective pressures can drive it to acquire new exploit-
ative functions.
Disingenuous displays of emotion. Deceptive displays of
emotion can be effective manipulation devices. For instance, an
accused party may mollify his accusers by feigning remorse. A
false display of guilt conveys that one acknowledges one’s cul-
pability, feels remorse, and is in the process of recalibrating
one’s welfare trade-off ratio. When the display is believable, this
can be a useful, minimal-effort manipulative tactic for escaping
punishment.
Exaggerated displays of anger can also be used for manipu-
lation. In antagonistic encounters across the animal kingdom,
exaggerated displays of aggression often work as successful
threats and are used to negotiate social hierarchies (Alcock,
2009; Dawkins, 1976; Zahavi & Zahavi, 1997). In close rela-
tionships, exaggerated displays of anger—for example, feigning
indignation in response to a transgression—may be a useful tac-
tic to induce a relationship partner to place greater weight on
one’s welfare.
In sum, an evolutionary psychological approach points to the
important role of the emotions in conflict. Emotions can be used
for manipulation in at least two ways: exploitative induction and
disingenuous displays. These two may become coupled; for exam-
ple, when exaggerated sadness is used to elicit empathy or exag-
gerated guilt to induce forgiveness. Emotions likely play a critical
role in the battlegrounds of conflict and social manipulation.
Conclusions
Modern evolutionary psychology provides important conceptual
insights into human emotions. We described the coordination
problem, arguing that adaptive problems include a number of
different subtasks, each of which requires the activation of a dif-
ferent program for its solution. These programs must be smoothly
and efficiently coordinated, as the effectiveness of one program
often depends on the prior or concurrent activation of another
(the concatenation problem), and programs sometimes have
conflicting outputs (the problem of conflicting outputs). The
coordination problem can be computationally complex.
Consequently, we propose that selection has fashioned adapta-
tions whose function is to coordinate other mechanisms in the
service of solving an adaptive problem (Cosmides & Tooby,
2000; Tooby & Cosmides, 2008). These are the emotions.
We highlight several conceptual contributions of our
approach: (a) more cogent criteria for classifying the emotions,
(b) an emphasis on information processing in addition to out-
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Al-Shawaf et al. Evolutionary Psychology of the Emotions 183
puts, and (c) systematic hypothesis generation based on task
analyses of adaptive problems. Emotions can evolve to solve
adaptive problems in any domain tributary to fitness, many
evolved emotions lack recognizable signals, and evolved human
emotions need not be present in other species to qualify as basic.
These conceptual revisions enhance classificatory accuracy,
lead to the identification of a broader array of evolved emotions,
and point to a greater role for emotions in psychological func-
tioning than historically envisioned. Our framework also high-
lights many sex-differentiated design features of human
emotions, each corresponding to sex-differentiated adaptive
problems men and women recurrently faced over human evolu-
tionary history—features that have been largely absent from
prior evolutionary theories of the emotions.
Our illustrations of these principles yielded a series of novel
hypotheses about the effects of disgust and sexual arousal.
Many of these hypotheses are context-specific, such as those
pertaining to audience effects on the expression of disgust or
mating strategy effects on orgasmic dissolution. Researchers
can use this framework to generate novel hypotheses about any
evolved emotion. Emotions such as guilt, gratitude, envy, jeal-
ousy, pride, schadenfreude, embarrassment, and curiosity may
have evolved to solve a broad array of adaptive problems tribu-
tary to reproductive success. Such wide-ranging problems
include hierarchy negotiation, sexual consummation, reputation
management, moralistic punishment, childrearing, kin-directed
altruism, and many others.
This approach appears scientifically fruitful: it broadens the
conceptualization of the emotions, advances important theoreti-
cal principles, generates novel hypotheses, and poses new ques-
tions. From the vantage point of evolutionary psychology,
emotions are critical to many domains of life and central to the
science of psychology.
Declaration of Conflicting Interests
None declared.
Note
1. The main author (L.A.S.) presents the following faux limerick as a
humorous summary of the sexual arousal hypothesis and orgasmic
dissolution hypothesis:
Kablam
Arousal leadeth cognitive programs,
Spurring passionate and fertile wham-bams
It coordinates as it whirrs,
With different ends for his and hers –
So only one’s done at the big kablam!
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  • ... An important area for future research is emotion regula- tion. Emotional expression and regulation serve important evolutionary functions [45]. Securely attached individuals' positive expectations of support bolster their well-regu- lated emotional expression and constructive emotion reg- ulation strategies [34]. ...
  • ... A contemporary evolutionary psychological approach to the emotions suggests that they are coordinating mechanismsinformation-processing programs that evolved to coordinate the activity of a variety of psychological and physiological mechanisms in the service of solving an adaptive problem ( Cosmides & Tooby, 2000;Tooby & Cosmides, 1990, 2008). For example, fear regulates programs in order to avoid danger, sexual arousal orchestrates programs in preparation for a valuable sexual opportunity, and pathogen disgust coordinates mechanisms in the service of avoiding infection ( Al-Shawaf, Conroy-Beam, et al., 2015;Al-Shawaf & Lewis, 2017). This perspective suggests that emotions coordinate many different psychological and physiological programs, including perception, attention, memory, specialized inference mechanisms, communication and expression, and behavior ( Cosmides & Tooby, 2000;Tooby & Cosmides, 1990). ...
  • ... The heightened attention toward emotion, particularly negative emotion, serves as a mechanism for individuals to readily identify and extract information that will most benefit the individual within his or her adaptive roles. From an evolutionary perspective, emotions have evolved to assist in solving a range of problems to include: signaling danger and disgust; indicating levels of mood, energy, and effort allocation; and prioritizing motivations ( Al-Shawaf et al. 2016). Memories from experiences that elicited a negative emotion are indicative of situations that should be avoided in the present and future. ...
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