553001

research-article2014
EMR0010.1177/1754073914553001Emotion ReviewLench et al. Evolution of Emotion

ARTICLE

Emotion Review
Vol. 7, No. 1 (January 2015) 90­–98
© The Author(s) 2014
ISSN 1754-0739

A Functionalist Manifesto: Goal-Related DOI: 10.1177/1754073914553001

er.sagepub.com

Emotions From an Evolutionary Perspective

Heather C. Lench*
Shane W. Bench*
Kathleen E. Darbor
Melody Moore
Department of Psychology, Texas A&M University, USA

Abstract
Functional theories posit that emotions are elicited by particular goal-related situations that represented adaptive problems and
that emotions are evolved features of coordinated responses to those situations. Yet little theory or research has addressed the
evolutionary aspects of these theories. We apply five criteria that can be used to judge whether features are adaptations. There
is evidence that sadness, anger, and anxiety relate to unique changes in physiology, cognition, and behavior, those changes are
correlated, situations that give rise to emotions are consistent, and emotions are complex. To date, there is little experimental
evidence regarding whether discrete emotions resolve adaptive problems and do so relatively efficiently. Evidence supporting all
criteria is required to claim that discrete emotions are evolved features.

Keywords
constructionist, discrete emotions, evolution, functional theories

This article examines functional theories of goal-related emo- and to evaluate evidence for these assumptions in light of what
tions from the perspective of evolutionary theory. Functional is reasonable to expect within constraints of evolutionary pro-
theories have guided predictions in studies of emotion for sev- cesses. This article is an attempt to identify what types of evi-
eral decades and the core tenet of these theories—that different dence would support emotions as adaptations and to examine
emotions represent adaptations to particular problems—went related evidence for sadness, anger, and anxiety.
largely unquestioned. Recently, however, this tenet has been
challenged, in part because of the lack of clear theory regarding
how and why emotions are adaptations (Barrett, 2006).
Functional Accounts
Unfortunately for these accounts, the adapted nature of emo- There has been growing consensus that emotions are relatively
tions was assumed and not dealt with explicitly. As a result, it is brief and intense reactions to specific events (Eich, Kihlstrom,
difficult to evaluate the validity of this core tenet in light of Bower, Forgas, & Niedenthal, 2000). In contrast, moods are
available evidence. For functional theories to remain viable, it is relatively long-lasting states less tied to specific events (Russell,
imperative that the evolutionary assumptions of these theories 2003). In this review, we focus on sadness, anger, and anxiety,1
be stated and evaluated. This review represents a manifesto in as an example of applying evolutionary criteria in the study of
the sense that we intend to make a call to action for functional emotion, because they represent adapted responses to particular
accounts to be more explicit about addressing whether and in goal states according to functional theories. These emotions
what sense emotions are adaptations. We offer one view of how were also selected because they are all negative in valence and
this might be done that incorporates multiple functional theories have been the target of multiple studies and reviews. Arguments
of goal-related emotion and provide an example using sadness, have been made that many other emotions are discrete adapted
anger, and anxiety. Our goal is to identify shared assumptions reactions. It was necessary to limit the scope of this review to a

Author note: *Authors contributed equally. Thanks to EM lab and to Linda Levine and Brandon Schmeichel for comments.
Corresponding author: Heather Lench, Department of Psychology, Texas A&M University, 4235 TAMU, College Station, TX, 77843, USA. Email: hlench@tamu.edu

Downloaded from emr.sagepub.com at UNIV FEDERAL DE SERGIPE on December 12, 2016


small number of emotions, but the following criteria can be used
to evaluate evidence that any emotion is an adaptation.
Functional theories typically posit that emotions met specific
adaptive problems during human evolutionary history. Although
it is not necessary that emotions result from evolution to be
functional, most theories appeal to natural selection directly or
indirectly. The majority of functional theories share the follow-
ing assumptions:
1. Emotions are elicited by particular situations.
Some accounts (not included in this review) focus on
stimuli thought to evoke reflexive responses, such as
fear in response to snakes. Most functional accounts,
however, focus on the perceived discrepancy between
the desired and current status of goals (Carver, 2004;
Frijda, 1987; Lench, Flores, & Bench, 2011; Levine,
1996; Roseman, Antoniou, & Jose, 1996). Despite
variation in the particulars, across theories sadness
occurs when goals are irrevocably lost; anger when
attainment of goals is obstructed; anxiety when goals
are threatened. Emotions follow appraisals—rapid,
often unconscious, evaluations of goal status and the
environment (Arnold, 1960; Ellsworth & Scherer,
2003; Roseman & Smith, 2001).
2. These situations represented adaptive problems that
affected reproductive success and emotions are
evolved features of coordinated responses to those
problems (Mauss, Levenson, McCarter, Wilhelm, &
Gross, 2005; Pinker, 1997). Accordingly, the responses
associated with each discrete emotion are theorized to
address the adaptive problem that gave rise to that emo-
tion. Responses to sadness should promote disengage-
ment from lost goals and avoidance of future goal
failure; anger should promote motivation to overcome
obstacles to goals; anxiety should promote motivation
to avoid potential threats.
How tenable is the proposition that goal-related emotions are
adaptations that resulted from natural selection? Emotional
responses that occur across species and in response to biologi-
cally significant stimuli (joy while eating sweets; fear of snakes)
seem plausibly related to adaptation in early environments. But
many emotions experienced by modern humans, which are the
focus of this review, are less tied to specific stimuli and instead
result from the perceived status of current goals. It is unlikely, for
example, that rush-hour traffic impeding progress represented an
adaptive problem faced by early man that triggers reflexive
anger. Instead, anger in this situation depends on relatively
advanced evaluations of goals and situations that likely became
possible only after broad expansion in human cognitive ability,
thought to have occurred in the Pleistocene environment (Buss,
Haselton, Shackelford, Bleske, & Wakefield, 1999; Panksepp &
Panksepp, 2000). Evidence suggests that emotions were already
present at that time and across species (Panksepp & Panksepp,
2000). As a result of the chronology—that expanded cognitive
function evolved after emotional reactions were present—it is
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Lench et al. Evolution of Emotion 91
logically impossible that emotions resulting from evaluations of
goal status preceded this cognitive expansion.
It is plausible to assert, however, that goal-related emotions
are exaptations. Exaptations are features that took advantage of
earlier adaptation by expanding functionality (Buss, Haselton,
Shackelford, Bleske, & Wakefield, 1998; Gould & Vrba, 1982).
Exaptations result from changes wrought by expressive behav-
iors of an organism (the phenotype), which can later be passed
through genes if those behaviors improve reproductive success
(Gould, 1982). Gould and Vrba (1982) give the example of
bones, likely developed for nutrient storage, but were co-opted
for support as animals moved to land, after which natural selec-
tion acted on bones for that purpose. The argument, then, would
be that humans who could link cognitions about goal status to
emotional systems, and take advantage of resulting motivation,
would have been more successful than humans who lacked this
ability and, as a result, emotions in response to particular goal
situations became common. For example, an early human who
linked appraisals of goal obstruction to the physiological,
behavioral, and cognitive responses associated with anger,
might be more adept at obtaining resources than humans who
lacked this link. One implication of distinguishing between phy-
logenetically earlier and later emotions is that findings related to
one cannot necessarily be taken as evidence for the other (Izard,
2007). For simplicity, we continue to use the term “adaptation”
next, but we are referring to potential exaptation.
Evaluating Functional Accounts
Testing evolutionary theories is notoriously difficult given the
inability to recreate early environments and humans, but theo-
rists have described several lines of evidence that support claims
that features are adaptations. We identified these criteria in
frameworks from evolutionary psychologists (Cosmides, Tooby,
& Barkow, 1992) and developed them through consultation of
work by evolutionary biologists (Darwin, 1872; Dawkins, 1986;
Sommerhoff, 1950; Symons, 1992; Williams, 1966).2 Figure 1
presents an overview of these criteria applied to the study of
goal-related emotions. Williams (1966) argued that features
should only be described as adaptations if there is compelling
evidence for all of these criteria. When there is evidence related
to these criteria for sadness, anger, and anxiety, we briefly
review that evidence; when there is scant evidence, we suggest
experimental paradigms that would address the criterion.
Precision
Precision reflects mapping of suspected adaptations onto unique
causes and consequences that support inference of a biological
basis for a feature. The precision of discrete emotions can be
reflected in unique causal physiological mechanisms and in the
degree to which discrete emotions relate to changes in physiol-
ogy, behavior, and cognition. These aspects of precision are
similar to, but distinct from, those used to evaluate whether dis-
crete emotions are categorical/“specific” in a philosophical
sense.3 Much empirical work has focused on these issues and
92 Emotion Review Vol. 7 No. 1
Constancy of situa on
predic ng emo on
Constancy of correlated
outcomes
Situa onal
challenge
Physio.
mechanism Emo on
Precision of physiological
mechanism associated
with emo on
Precision of emo on
resul ng in changes
Figure 1. Evolutionary criteria applied to the study of functional goal-related emotions
Note. Although emotion is represented as a separate construct for the purposes of this figure, we do not intend to claim that emotions are separate from organized changes in
outcomes. In most cases, however, emotion would be separate empirically in that it would be experimentally manipulated.
therefore we describe the logic related to precision and refer to
existing reviews of related evidence.
Precision of physiological mechanism. If emotions are
adaptations, then they must have specific physiological con-
comitants that can be passed through genes. Therefore precision
can be recognized in identification of physiological structures
or mechanisms, presumably in the brain, that occur with an
emotion. Importantly, it is not necessary that each emotion be
completely unique in all physiological concomitants, such that,
for example, each is associated with one and only one brain area
(Levenson, 2011). Instead, the requirement is that there is struc-
tural or functional physiology associated with an expressed fea-
ture (the emotion) that could be selected for.
There is evidence that precise structural physiological
mechanisms underlie sadness, anger, and anxiety. Medications
are often effective in reducing these chronic emotions experi-
enced in mood disorders and drugs that target specific areas
can elicit these emotions (Buck, 2010; Carver & Harmon-
Jones, 2009). For example, amytal injections in the left hemi-
sphere produce sadness. Further, there is evidence that
impairments in specific brain regions result in discrete emo-
tional experiences (Berlin, Rolls, & Kischka, 2004; Carver &
Harmon-Jones, 2009; Cohen, Paul, Zawacki, Moser, & Sweet,
2001; Van der Velde et al., 2013). Patients with neurodegen-
erative or psychiatric diseases that impair brain function expe-
rience changes in the frequency of negative emotions, although
specific negative emotions are seldom targeted and multiple
brain regions are often affected (Levenson, Sturm, & Haase,
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Efficiency of emo on
compared to other
constructed “emo ons”
Behavior
Greater success
in func onally
relevant
situa ons
Cogni on
Errors when
mismatched
Physiology
Economy of emo on in
resolving adap ve challenges
2014; Marsh, 2013). The strongest such evidence comes from
studies where specific brain areas are compromised. In case
studies, anger was experienced more often in patients with
orbitofrontal cortex lesions than those with different or no
lesions (Berlin et al., 2004).
Evidence of functional physiological mechanisms that co-
occur with discrete emotions comes from neuroimaging stud-
ies. Meta-analyses have concluded that there is evidence of
precision, particularly for anxiety and the amygdala, sadness
and the subcallosal cingulate, and anger and the lateral orbito-
frontal cortex (OFC), but there was variability in activation
(Brosch & Sander, 2013; Fuser-Poli et al., 2009; Murphy,
Nimmo-Smith, & Lawrence, 2003; Phan, Wager, Taylor, &
Liberzon, 2002; Vytal & Hamann, 2010). Three other meta-
analyses concluded that there is no support for precision
(Kober et al., 2008; Lindquist, Wager, Kober, Bliss-Moreau, &
Barrett, 2012; Sergerie, Chochol, & Armony, 2008). The dis-
crepancy in conclusions is likely due to differing assumptions
(Hamann, 2012a). Most neuroimaging work focuses on entire
areas; however, from an evolutionary perspective, it is only
necessary that emotions relate to one specific mechanism, not
an entire neural region. Indeed, research suggests that emo-
tions, like other processes, relate to neural networks that cross
multiple regions (Hamann, 2012b). Further, areas identified in
neuroimaging studies are typically broad and serve multiple
functions, making it difficult to predict areas theoretically
related to emotions (Armony, 2013). These limitations pre-
clude conclusions about the degree to which functional brain
processes relate to emotions.

Precision of emotion resulting in changes. Precision can
also be examined through relation of a feature to consequences,
which presumably occur after the underlying physiological
causal mechanism is activated. This can be represented as the
degree to which emotions are uniquely associated with changes
in physiology, behavior, and cognition.4 Importantly, expressed
behaviors of any adapted feature result from both biologically
based features (the genotype; nature) and experiences (nurture)
that alter expression of features (the phenotype; Cacioppo,
Berntson, Sheridan, & McClintock, 2000; Gould & Lewontin,
1979). From an evolutionary perspective, then, discrete emo-
tions should probabilistically associate with unique conse-
quences, but it would not be reasonable to expect perfect
associations because of the influence of environmental factors
and individual learning (Buck, 2010; Levenson, 2011; Weis-
feld & Goetz, 2013; see, however, Lindquist, Siegel, Quigley,
& Barrett, 20133). It is also not reasonable to predict com-
pletely unique consequences for each emotion because there is
some overlap in the theorized function (Lench, Bench, & Flo-
res, 2013). Anger and anxiety, for example, are both theorized
to facilitate responses to problems (obstructions or threats), and
should therefore be associated with physiology that facilitates
action.
There is evidence of precision in physiological reactions in
the autonomic nervous system, although many changes that
relate to proposed functions of discrete emotions have not
been investigated (e.g., sweating or blood flow changes;
Levenson, 2014). The behavioral response that has received
the most empirical attention is facial expression. Studies have
revealed precision in the degree to which a combination of
facial movements is associated with discrete emotions,
including sadness, anger, and anxiety, and this pattern is reli-
ably present across individuals and cultures (Ekman, 1992,
although people may not reliably identify emotions in others;
Widen, Christy, Hewett, & Russell, 2011). A meta-analysis of
the effects of discrete emotions revealed that, across nearly
700 experimental studies, there were small to moderate dif-
ferences in physiology, cognition, and behavior, among sad-
ness, anger, and anxiety, in directions predicted by functional
theories (Lench et al., 2011). The reliability of consequences
of emotion is also supported by evidence that computer algo-
rithms incorporating changes in behavior and physiology pre-
dict reported emotions (Bailenson et al., 2008; Kragel &
LaBar, 2013). In other words, programs can determine what
emotion people are experiencing by looking at how they
behave and their physiology.
Summary. Sadness, anger, and anxiety do have precision in
their relation to physiological mechanisms and expression in
physiology, behavior, and cognition. This is not to say that these
emotions are completely unique in associated causes and conse-
quences, and such an expectation would be unreasonable from
evolutionary or emotion perspectives3 (Tooby & Cosmides,
1992). Rather, these emotions are associated with neurological
mechanisms and phenotypical expressions, consistent with pre-
cision in adapted features.
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Lench et al. Evolution of Emotion 93
Complexity
Natural selection can give rise to very complex features, such as
the eye, that involve numerous changes in structure and func-
tion over long periods. Thus complexity can be used to evaluate
the likelihood that features are adaptations (Dawkins, 1986;
Williams, 1966). Emotions are certainly complex and, if
adapted, likely evolved over a long period, resulting in exapted
goal-related emotions. As reviewed before, sadness, anger, and
anxiety have been related to physiological changes throughout
the human body, behaviors from muscle movements to social
interactions, and cognitions from attentional processes to mem-
ory, in ways consistent with functional theories (Lench et al.,
2011). From an evolutionary perspective, the question is
whether this degree of complexity is likely to have arisen by
chance alone (Dawkins, 1986), and this seems unlikely in the
case of emotions.
Constancy
Constancy can be reflected in the consistency of responses. If
discrete emotions evolved to meet adaptive problems, then
there should be consistency in situations that give rise to par-
ticular emotions. There should also be correlated changes in
phenotypical expressions of discrete emotions. The precision
of changes in physiology, behavior, and cognition was
addressed in the previous lines; constancy reflects not the
uniqueness of these changes, but rather the degree to which
these changes co-occur.
Constancy of situations. If discrete emotions evolved to
coordinate responses to particular adaptive problems, as posited
by functional theories, then particular situations should evoke
those emotions. Identifying relevant situations is particularly
difficult for goal-related emotions because it is evaluations of
situations (appraisal), rather than objective situations, that elicit
emotions. For example, not everyone will react to failing tests
with sadness, but everyone who appraises that their goal is
irrevocably lost, regardless of the specific situation, should
experience sadness. A similar challenge is faced by biologists
when they cannot directly manipulate situations related to adap-
tations, and in such cases it is the correlational pattern that is
examined (Sommerhoff, 1950). If emotions are adaptations,
there should be patterns in the appraisals that result in emotions
across individuals and cultures. There is evidence that the emo-
tions people report can be predicted from their appraisals
(Scherer & Meuleman, 2013). Multiple studies have revealed
that appraisals correlate with organized changes in physiology,
behavior, and cognition associated with sadness, anger, and
anxiety (Moors, Ellsworth, Scherer, & Frijda, 2013). There is
also evidence that apparent variation in emotional response
across individuals and groups, which appear to contradict evo-
lutionary accounts, is accounted for by appraisals (which result
from learning; Moors et al., 2013; Stein & Hernandez, 2007).
Thus there appears to be constancy in appraisals of situations
that relate to sadness, anger, and anxiety.
94 Emotion Review Vol. 7 No. 1
Constancy of correlated outcomes. If emotions evolved to
coordinate responses to particular adaptive problems, then there
should be constancy in organized changes in physiology, behav-
ior, and cognition (Barrett, 2006; Tooby & Cosmides, 1990). It
is a common misunderstanding that adaptation implies genetic
determinism, such that adapted features will always result in
particular expression of that feature. Modern evolutionary the-
ory, however, recognizes the fact that individual experiences
modify expression of any biological adaptation (Cacioppo et al.,
2000; Gould & Lewontin, 1979). Therefore changes associated
with emotions are likely to be only moderately correlated due to
this variability (Izard, 2007; Panksepp, 2007).
Across numerous studies, emotional expressions have small
to moderate correlations with physiological changes and experi-
ence (Lewis, 2011; Reisenzein, Studtmann, & Horstmann,
2013). Few studies have focused on correlations across other
responses, but two investigations found moderate to high cor-
respondence during sadness, anger, and anxiety among cogni-
tion, behavior, and physiology (Bonanno & Keltner, 2004;
Mauss et al., 2005). A meta-analysis attempted to address this
issue by correlating effects in studies that measured more than
one outcome (Lench et al., 2011). They found small to moderate
correlations among effects of sadness, anger, and anxiety on
physiology, experience, behavior, and cognition (cognition was
not significantly correlated with behavior and experience).
There is also evidence that responses, including physiology, are
more highly correlated with one another during emotional expe-
riences compared to nonemotional experiences, consistent with
the theorized organizing function of emotion (Hsieh et al., 2011;
Levenson, 2014; Mauss et al., 2005).
Summary. Sadness, anger, and anxiety show reasonable
constancy. They are reliably elicited by appraisals of situations.
Changes in physiology, behavior, and cognition co-occur as a
result of these discrete emotions in a coherent manner.
Economy
The economy of a feature is reflected in how well it resolves
the adaptive problem thought to have given rise to the feature,
in what is sometimes called a “performance evaluation”
(Tooby & Cosmides, 1992). Features should spread through
populations only when they improve outcomes (Cosmides et al.,
1992; Levenson, 2011; Sommerhoff, 1950; Symons, 1992;
Williams, 1966). The economy of emotions would be sup-
ported by evidence that they resolve problems. Critically, the
improvement in outcomes must be in situations that match the
adaptive challenge thought to have given rise to the emotion
(e.g., evidence that sadness increases contemplation is not suf-
ficient unless contemplation is shown to resolve failure). To
date, there is shockingly little experimental evidence that
emotions improve outcomes in situations that match the adap-
tive problems proposed in functional theories. This is surpris-
ing given that such evidence is considered the most compelling
demonstration that features are adaptations (Sommerhoff,
1950; Williams, 1966).
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There is evidence that emotional expression can improve
outcomes during social interactions. Expressing anger
improves compliance in negotiations (Tamir, Mitchell, &
Gross, 2008; van Kleef, De Dreu, & Manstead, 2004) and can
be useful in competitions (Tamir & Ford, 2012). Expressing
sadness during calls for help increases support from others
(Lelieveld, van Dijk, van Beest, & van Kleef, 2013). Yet most
functional theories focus on the goal-related function of emo-
tions, and evidence that emotions improve outcomes in these
situations is sparse. As described before, there is evidence
that discrete emotions precisely influence physiology, behav-
ior, and cognition in ways that seem as though they would be
functional, such as anger increasing attention to rewards
(Ford, Tamir, Brunye, Shirer, & Mahoney, 2010). Intuitively,
then, it would seem that these consequences are functional,
but this has not been shown. There is some correlational evi-
dence that sadness predicts goal adjustment (Wrosch &
Miller, 2009), that anger predicts persistence during chal-
lenge and higher scores (Lench & Levine, 2008; Mikulincer,
1988), and that anxiety predicts plans to avoid threats (Safer,
Levine, & Drapalski, 2002).
But, to demonstrate economy, studies would need to experi-
mentally manipulate one emotion and show improvement in
resolution of the theorized adaptive problem compared to other
emotions (particularly other similarly valenced states and neu-
tral conditions). For example, one could manipulate anger
(compared to controls) and examine the degree to which people
persist during challenge and, as a consequence of persistence,
overcome obstacles. Without such studies, the link between the
consequences of emotion and actual improvement in outcomes
is largely undemonstrated. Further, particularly strong support
for the economy of emotions would be provided by evidence
that emotions result in systematic errors when situations do not
match the theorized adaptive problem that gave rise to the emo-
tion (Dawkins, 1986). Similar to people overeating sugar in
environments where it is plentiful because of evolved prefer-
ences for sweet tastes when sugar was rare, emotions should
lead to problematic responses when situations do not resemble
the adaptive problems that gave rise to the emotion. For exam-
ple, persistence related to anger should result in errors when
goals are not attainable. In contrast, if emotions are not adapta-
tions, then people should learn from experience and no longer
be prone to systematic errors in situations where the emotion
would not be helpful.
Efficiency
Similar to principles of parsimony, evolutionary accounts are
supported by evidence that features are more efficient in resolv-
ing an adaptive problem than other possible solutions. Efficiency
is typically difficult to address because theorists have trouble
imagining solutions other than those that seem to have occurred.
For example, how might one respond to failure in an adaptive
manner other than feeling sad and associated changes of reduced
arousal, disengagement from goal pursuit, and rumination over
causes of failure?

One alternative that has been put forth, in what are some-
times called constructionist accounts, is that all of the changes
associated with emotion could occur, but without the organizing
construct of the emotion (Barrett, 2006, 2014 Clore & Ortony,
2013). In other words, changes in physiology, behavior, and
cognition are separate learned reactions to environmental stim-
uli and should be highly variable. For example, people might
react with reduced arousal, disengagement, and rumination after
failure, but these reactions entirely depend on learning history
and are not organized in such a way that they should reliably
co-occur. Some constructionist accounts consider the ability to
learn these responses and valenced reactions (feeling positive/
negative) to be adaptations, but not discrete emotions (Barrett,
2013, 2014). To examine efficiency, then, the question would
be: Which feature most efficiently resolves the adaptive prob-
lem—emotions that organize responses to challenges or learn-
ing the different responses in physiology, behavior, and
cognition necessitated by particular situations?
Prima facie, an organizing entity appears more efficient in
responding to adaptive problems than learning a multitude of
responses in a multitude of situations, because it would allow
for fast and efficient responses across similar situations and
could be organized from an early age. Discrete emotion accounts
are sometimes called reductionistic (Barrett, 2006, 2013), but
biology is also reductionistic in the sense that efficient adapta-
tions are favored (Williams, 1966). Although learning responses
would offer maximum flexibility across situations, the evidence
suggests constancy in response rather than complete variability.
From an efficiency perspective, it is not clear why evolution
would favor extreme flexibility in responses if that flexibility is
rarely utilized (Stewart-Williams & Thomas, 2013).
But we suggest that the relative efficiency of functional ver-
sus constructionist explanations can be addressed empirically.
One possibility would be to teach individuals novel responses
to challenges and compare the efficacy of these responses to
theorized discrete emotions. If emotions are constructed, these
learned responses should be efficient; if emotions are adapta-
tions, they should be more efficient than learned responses.
This would involve not just teaching people to modify existing
emotional responses, but rather teaching them to experience
completely novel responses. It is difficult to imagine how to
create novel “emotional” responses, but this might be done by
collating a list of changes in cognition, behavior, and physiol-
ogy associated with existing emotions, and then randomly
picking from those lists to create a new profile. One would then
teach people to have these reactions in a particular situation.
Such an investigation would also demonstrate whether or not
people, including infants, are capable of learning to alter and
coordinate the responses that have been associated with dis-
crete emotions, including hormonal and neurotransmitter
responses (Sullivan & Lewis, 2003; Tooby & Cosmides, 1992).
If results supported the efficiency of emotions as adaptations
rather than learned constructions, however, the role of learning
is still important. Expression of emotions, even if adaptations,
will be modified by learning and experiences, as is the case
with all adapted features.
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Lench et al. Evolution of Emotion 95
Are They Adaptations?
We conclude from the available evidence that the jury is still out
about whether sadness, anger, and anxiety are adaptations. To
make the case that features are adaptations, compelling evi-
dence must be provided for all of the aforementioned criteria
(Williams, 1966). There is compelling evidence for some crite-
ria, including precision and constancy, but support for other cri-
teria is lacking. Particularly problematic is the dearth of
evidence that these emotions are functional and resolve the
adaptive challenges thought to have given rise to them.
Evidence Not Required
For a functional account to be supported, there are several
streams of evidence that are not required from an evolutionary
perspective. First, even if one judges, based on the available
evidence, that sadness, anger, and anxiety are adaptations, this
judgment cannot automatically extend to all emotions. An
explanation that involves adaptation should only be offered in
the face of compelling evidence and at the lowest level of
organization supported by evidence (Williams, 1966).
Theorists have argued, for example, that emotions that develop
later in life and arise in response to culturally learned situa-
tions, such as shame, guilt, and embarrassment, might be con-
structions rather than adaptations (Izard, 2007). It is likely that
emotions vary in the degree to which they reflect biological
adaptations versus learned responses and the relative contribu-
tions of nature and nurture would need to be parsed for each
emotion (Sommerhoff, 1950). Researchers who believe emo-
tions are exaptations would need to garner evidence regarding
the criteria described before.
Second, for functional accounts to be supported, it is not nec-
essary that emotions occur completely separately from cultures
or concepts transmitted through culture. Culture is the product
of individual minds that are influenced by evolutionary pro-
cesses (Tooby & Cosmides, 1992), and the fact that cultures
convey information about emotions does not negate emotions as
adaptations. For example, one culture might prescribe disgust at
the idea of eating dogs, while another culture enjoys the dish.
This does not indicate that disgust is not an adaptive response to
appraisals of noxious stimuli, but rather that disgust is not an
adaptive response to dog meat. Cultures might similarly dis-
courage certain emotions based on social standards, such as
restrictions about experiencing or expressing anger. This does
not indicate that anger is not an adapted response to appraisals
of obstructed goals. In fact, the continuity of these cultural pro-
scriptions provides indirect evidence that emotions are not
learned; otherwise, the emotions would be eliminated within a
few generations and there would be no need for continued pro-
scriptions (Cole, Shrestha, & Tamang, 2006).
Third, functional accounts are not disproven by evidence
that discrete emotions can be mapped onto dimensions such as
valence and arousal. Natural selection influences organisms
over long periods, but it does not plan features (Dawkins, 1986).
As a result, adapted features build upon what is already present.
96 Emotion Review Vol. 7 No. 1
As discussed previously, goal-related emotions are plausibly
exaptations that built upon emotional reactions that were more
specific to stimuli (fear of snakes). Yet these reactions are prob-
ably also exaptations and were likely derived, through natural
selection, from valenced reactions to approach beneficial stim-
uli and avoid noxious stimuli. Valenced reactions are important
organizing components of human thought and behavior
(Damasio, 2003). However, it does not follow that discrete emo-
tions are not adapted. It does mean that discrete emotions should
be related to phylogenetically earlier valenced reactions.
Therefore there should be overlap in the causes and conse-
quences of positive emotions and of negative emotions.
Although not required, there are two additional sources of
evidence that could support functional accounts. It is impossible
to recreate evolutionary pressures by placing people back in the
Pleistocene, but one way to examine the importance of evolu-
tionary pressures is to compare groups of people that vary in
their exposure to those pressures. Evidence that people with
particular evolutionary histories (because of gender or particu-
lar group experiences) have different propensities toward emo-
tional experience consistent with those histories could support
functional accounts. For example, Taylor (2006) argued that dif-
ferential investment in offspring contributed to gender differ-
ences in reactions to threat (“tend and befriend” vs. “fight or
flight”), and physiological evidence supports this proposition.
There may be additional instances where group differences in
evolutionary pressures might have altered emotional experience
or expression.
Conclusions
Scientific theories also evolve, with those garnering the most
support emerging as the consensus and those with less support
fading away. If functional accounts of emotion are to remain
viable explanations, it is critical that they empirically address
whether emotions are adaptations.
Notes
1 This review focused on anxiety perception of potential threats to goals
(an upcoming exam), rather than fear perception of present, often
physical, threats (Lench et al., 2011) because: (a) although anxiety and
fear are both goal-related, anxiety is typically described in functional
accounts (the focus of this review) and (b) most empirical studies
include manipulations consistent with anxiety (anticipating a difficult
speech); thus it was accurate to describe the experimental evidence in
relation to anxiety.
2 Some theorists combine several criteria into a single criterion. For
example, Darwin’s “principle of serviceable associated habits”
includes features of precision and economy. For clarity, we separated
criteria in these cases.
3 As an example of the discrepancy between specificity in a philosophi-
cal sense versus precision in an evolutionary sense, Barrett (2006)
often describes specificity as “carving nature at its joints” whereas an
evolutionary approach would not attempt to carve nature because evo-
lution is viewed as continuous. This distinction has implications for
how evidence is evaluated and whether one would expect, for exam-
ple, completely unique emotion profiles (specificity) or a probabilistic
association with particular outcomes (precision).
Downloaded from emr.sagepub.com at UNIV FEDERAL DE SERGIPE on December 12, 2016
4 Physiological changes could include neurological changes that occur
after the physiological mechanism that causes an emotion is activated.
We do not explicitly address this possibility because, at present, tech-
nology and analyses do not permit identification of the time course of
activation.
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