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Disgust: Evolved Function and Structure

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DOI: 10.1037/a0030778 · Source: PubMed

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Psychological Review © 2012 American Psychological Association
2013, Vol. 120, No. 1, 65– 84 0033-295X/13/$12.00 DOI: 10.1037/a0030778

Disgust: Evolved Function and Structure

Joshua M. Tybur Debra Lieberman

VU University Amsterdam University of Miami

Robert Kurzban Peter DeScioli

University of Pennsylvania Brandeis University

Interest in and research on disgust has surged over the past few decades. The field, however, still lacks
a coherent theoretical framework for understanding the evolved function or functions of disgust. Here we
present such a framework, emphasizing 2 levels of analysis: that of evolved function and that of
information processing. Although there is widespread agreement that disgust evolved to motivate the
avoidance of contact with disease-causing organisms, there is no consensus about the functions disgust
serves when evoked by acts unrelated to pathogen avoidance. Here we suggest that in addition to
motivating pathogen avoidance, disgust evolved to regulate decisions in the domains of mate choice and
morality. For each proposed evolved function, we posit distinct information processing systems that
integrate function-relevant information and account for the trade-offs required of each disgust system. By
refocusing the discussion of disgust on computational mechanisms, we recast prior theorizing on disgust
into a framework that can generate new lines of empirical and theoretical inquiry.

Keywords: disgust, adaptation, evolutionary psychology, emotion, cognition

Research concerning disgust has expanded in recent years (Ola- selection pressure driving the evolution of the disgust system, but
tunji & Sawchuk, 2005; Rozin, Haidt, & McCauley, 2009), and there has been less precision in identifying the selection pressures
contemporary disgust researchers generally agree that an evolu- driving the evolution of disgust systems unrelated to pathogen
tionary perspective is necessary for a comprehensive understand- avoidance (e.g., behavior in the sexual and moral domains). Al-
ing of the development and function of this emotion (e.g., Chap- though multiple researchers have suggested that pathogen disgust
man, Kim, Susskind, & Anderson, 2009; Curtis, de Barra, & was “co-opted” or “modified” to execute new functions (e.g.,
Aunger, 2011; Kelly, 2011; Oaten, Stevenson, & Case, 2009; Rozin et al., 2008), exactly how these new domains functionally
Rozin, Haidt, & McCauley, 2008; Tybur, Lieberman, & Griskevi- differ from pathogen-related disgust has not always been specified.
cius, 2009). Contributing to this consensus are the cross-cultural Hence, one of our goals here is to articulate the selection pressures
consistency in expressions of disgust (Ekman, 1972), the straight- that led to the evolution of separate disgust adaptations that per-
forward connection between common disgust elicitors and infec- form distinct functions in the domains of pathogen avoidance,
tious disease threats (e.g., rotting meat; Curtis & Biran, 2001), and mate choice, and moral judgment.
the recognition that pathogens have imposed strong selection pres- Further, in contrast to the widespread appeal to function, there
sures on the evolution of most organisms (Fumagalli et al., 2011; has been far less work specifying the underlying psychological
Tooby, 1982). mechanisms that generate the specified disgust responses. In ad-
Despite recent advances, important issues remain. Disgust re- dition to specifying function, then, we examine the computational
searchers have been explicit in identifying pathogens as the key structure of disgust. Whereas an examination of evolved function
addresses the ultimate level of explanation of behavior, that is,
why disgust exists and operates the way it does, a description of
information processing structure provides a proximate level of
This article was published Online First December 3, 2012. explanation, that is, how each disgust system carries out its func-
Joshua M. Tybur, Department of Social and Organizational Psychology, tion (Tinbergen, 1963; Tooby & Cosmides, 1992). For each func-
VU University Amsterdam, Amsterdam, the Netherlands; Debra Lieber- tion disgust performs, psychological mechanisms must take spe-
man, Department of Psychology, University of Miami; Robert Kurzban, cific inputs, integrate them with fitness-relevant moderating
Department of Psychology, University of Pennsylvania; Peter DeScioli, factors, and then set in motion the constellation of behavioral,
Department of Psychology, Brandeis University. cognitive, and physiological processes naturally selected to per-
We thank Vladas Griskevicius for helpful comments on an earlier form
form the function at hand. For instance, pathogen disgust requires
of this article. We also thank Jon Haidt and Randy Thornhill for helpful
comments.
(a) detection systems that take as input cues associated with
Correspondence concerning this article should be addressed to Joshua pathogen presence and (b) integration systems that weigh cue-
M. Tybur, VU University Amsterdam, Department of Social and Organi- based pathogen threat level with other fitness relevant factors (e.g.,
zational Psychology, Van der Boechorstraat 1, 1081 BT Amsterdam, the hunger) and then adaptively regulate withdrawal and avoidance
Netherlands. E-mail: [email protected] behaviors accordingly.

65
66 TYBUR, LIEBERMAN, KURZBAN, AND DESCIOLI

Here we outline the possible information processing structures Fallon, 1987; Rozin, Haidt, & McCauley, 2008, 2009). The RHM
required by pathogen, sexual, and moral disgust systems, with a model is explicitly based on considerations of evolved function,
focus on the trade-offs each system was designed to make (e.g., the and it proposes several “stages” through which disgust has ex-
trade-off between the costs of infection vs. eating made by patho- panded to multiple domains. RHM propose that “core disgust”—
gen disgust; the trade-off between inbreeding vs. having sex made disgust toward food, animals, and body products— evolved from a
by sexual disgust; the trade-off between endorsing or rejecting toxin-based food-rejection system (distaste) and functions to mo-
moralized rules). We argue that the development of such compu- tivate pathogen avoidance. RHM further suggest that core disgust
tational models is a useful tool to help make explicit the informa- was co-opted to address a putative separate function: neutralizing
tion required to perform a particular function, and in so doing it reminders that humans are animals. In particular, the RHM model
can help generate hypotheses regarding functional design, loci of posits that animal-reminder disgust functions to protects us from
individual differences, potential sources of impairment, develop- seeing “ourselves as lowered, debased, and mortal” (Rozin et al.,
ment, and cross-cultural variability. 2008, p. 762) by rejecting any reminders of our animal nature and
To begin, we briefly review the current dominant perspective on mortality, and it is elicited primarily by sex, death (e.g., corpses),
disgust and suggest that limitations associated with this perspec- bad hygiene, and body envelope violations (e.g., gore, wounds,
tive necessitate an alternate approach. We summarize the details of disfigurement). Hence, animal reminder disgust is thought to func-
an evolutionary-computational approach for understanding emo- tion to “protect the body and the soul” (Rozin et al., 2008, p. 764).
tions and apply this to disgust. In so doing, we specify the selection RHM suggest the presence of two additional domains of disgust:
pressures that led to the evolution of pathogen, sexual, and moral interpersonal disgust, which functions to protect the body, soul,
disgust. For each adaptation, we describe a computational structure and social order and is elicited by contact with strangers or
that considers the trade-offs the system was selected to make. undesirables, and moral disgust, which functions to protect the
Finally, we explain how a computational model can generate novel social order and is elicited by moral violations (see Figure 1 for a
testable hypotheses and inform programs of research across mul- comparison of the model proposed by RHM, 2008, and the model
tiple disciplines, including developmental, clinical, personality, proposed herein).
cognitive, and social psychology. Despite its ubiquitous use across disparate fields, the RHM
model has key shortcomings. For example, animal reminder dis-
gust is consistently posited to “protect the soul,” a function pur-
Contemporary Disgust Theory
ported to be unique to humans (Rozin, Haidt, & McCauley, 2008,
There is a growing consensus within the social sciences that 2009; Rozin, Lowery, Imada, & Haidt, 1999). However, avoidance
disgust plays a key role in motivating behavior that probabilisti- of corpses—the prototypical elicitor of animal reminder disgust
cally reduces exposure to pathogens— organisms that exploit their under the RHM model—is ubiquitous across other species (Wag-
hosts for resources critical to the pathogen’s own survival and ner, Stroud, & Meckley, 2011; Yao et al., 2009). The literature on
replication (Calder, Keane, Manes, Antoun, & Young, 2000; Cur- this topic suggests that corpse avoidance in nonhuman animals
tis, Aunger, & Rabie, 2004; Curtis & Biran, 2001; Curtis et al., functions to protect against threats posed by infectious microbes
2011; Fessler, Eng, & Navarrete, 2005; Fessler & Navarrete, that may have killed the other organism or that are rapidly colo-
2003a; Fleischman & Fessler, 2011; Kelly, 2011; Marzillier & nizing the corpse. Other aspects of animal reminder disgust—
Davey, 2004; Oaten et al., 2009; Olatunji & Sawchuk, 2005; Rozin disgust toward sex, poor hygiene, and deformity—likely have
et al., 2008; Stevenson & Repacholi, 2005; Susskind et al., 2008; more specific functions that are well grounded within an evolu-
Tybur, Bryan, Magnan, & Caldwell Hooper, 2011; Tybur, Merri- tionary framework but are not articulated under the RHM model
man, Caldwell, McDonald, & Navarrete, 2010). Many of the (e.g., Fessler & Navarrete, 2003a; Oaten, Stevenson, & Case,
substances that people across cultures find intuitively disgusting, 2011; Ryan, Oaten, Stevenson, & Case, in press; Tybur et al.,
such as feces, dead bodies, and sexual fluids, reliably contain 2009). Moreover, the animal reminder perspective is tied to a
harmful bacteria (see Curtis & Biran, 2001, Table 6, for a sum- questionable assumption about human nature: that “anything that
mary). However, as multiple disgust researchers have noted, many reminds us that we are animals elicits disgust” (Rozin et al., 2008,
objects, acts, and concepts that do not pose objective infectious- p. 761). Nonhuman animals can be readily observed running and
disease threats also elicit disgust. For example, feces-shaped jumping like humans, breathing like humans, sleeping like hu-
fudge, apple juice in bedpans, and sterilized plastic cockroaches mans, and caring for their offspring like humans, yet none of these
elicit reports of disgust, as do many sexual concepts and nonin- behaviors elicit disgust. In fact, humans often view comparisons to
fectious social transgressions (Borg, de Jong, & Schultz, 2010; animals positively rather than with disgust (see Royzman & Sa-
Cannon, Schnall, & White, 2011; Chapman et al., 2009; Hutcher- bini, 2001; Tybur et al., 2009). In general, the rapidly developing
son & Gross, 2011; Koukounas & McCabe, 1997; Rozin, Millman, evolutionary literature—notably the literature on behavioral adap-
& Nemeroff, 1986). A critical issue for disgust researchers has tations for pathogen avoidance (see Curtis, 2007; Curtis & Biran,
been to generate a theoretical model to account for the heteroge- 2001; Curtis et al., 2004, 2011; Hart, 2011; Schaller & Park,
neity of disgust elicitors outside of the domain of pathogen avoid- 2011)—suggests that an examination of functions not articulated
ance. in the RHM model can offer great utility for understanding disgust.
The theoretical model of disgust put forth by Rozin, Haidt, Another shortcoming of the RHM model is that, much like
McCauley, and colleagues, which we refer to as the Rozin–Haidt– Darwin’s (1872/1965) initial account of disgust, it relied on cir-
McCauley (RHM) model, has become the gold standard within the cular functional definitions. Darwin initially defined disgust as a
disgust literature over the past two decades (Haidt, McCauley, & reaction to “something revolting, primarily in relation to the sense
Rozin, 1994; Haidt, Rozin, McCauley, & Imada, 1997; Rozin & of taste, as actually perceived or vividly imagined” (p. 250).
 DISGUST FUNCTION 67

Accounts such as this beg the further question of what renders an

act sleazy. We argue that considerations of evolved function and
structure can help overcome some of these challenges.

An Evolutionary View of Emotion

Our examination of disgust draws heavily on an evolutionary
perspective of emotion (Keltner & Gross, 1999; Keltner, Haidt, &
Shiota, 2006; Nesse & Ellsworth, 2009; Tooby & Cosmides, 1990,
2008; Pinker, 1997; Tooby & Cosmides, 2008). This perspective
has been fruitfully applied to other psychological states, including
anger (Sell, Tooby, & Cosmides, 2009), fear (Öhman & Mineka,
2001; Tooby & Cosmides, 1990), jealousy (Buss & Haselton,
2005), and depression (Andrews & Thompson, 2009; Hagen,
2002; Keller & Nesse, 2006). Here, we apply this logic to three
proposed functionally distinct domains of disgust: pathogen, sex-
ual, and moral disgust.
Accordingly, we provide descriptions of each functional do-
main at two separate levels of analysis. The first, that of
deciphering evolved function, requires a description of the
historically recurring conditions that led to the emotion’s evo-
lution and subsequent maintenance. This level of analysis spec-
ifies the small subset of responses, out of the unbounded set of
Figure 1. Comparison of disgust models. The traditional model proposed all possible responses, that would have overcome the conditions
by Rozin, Haidt, and McCauley (2008; RHM) and the functional model posed by the posited selection pressures and led, on average, to
proposed herein group the elicitors of disgust differently. According to the an increased rate of survival and reproduction. Taken together,
RHM model, multiple domains of disgust emerged from distaste, which a description of selection pressures and subsequent solutions to
functions to protect the body from poisons. RHM’s model proposes four the conditions they posed constitute an analysis of evolved
types of disgust with distinct elicitors: core disgust, interpersonal disgust,
function.
animal nature disgust, and moral disgust. By contrast, a functional per-
Whereas an explanation of evolved function (an ultimate expla-
spective indicates that all of these elicitors, with the exception of sex and
moral offenses, are better interpreted as different sources of infectious nation) requires consideration of the selective conditions that per-
agents. Sex under the traditional model is thought to elicit disgust because sisted over many thousands of generations, our second level of
it reminds humans of their animal nature. From a functional perspective, analysis—that of information processing structure (a proximate
decisions about sexual behavior constitute a separate adaptive problem explanation akin to the algorithmic level of analysis proposed by
related to avoiding sexual contact with fitness-jeopardizing reproductive Marr, 1982)—specifies how the selected design feature operates
partners. Both models propose a separate domain of moral disgust. within an individual organism (Tinbergen, 1963). Given the dif-
Whereas RHM interprets its function as benefiting the group by protecting ferent adaptive problems disgust solves (e.g., decisions regarding
the larger social order, the framework proposed here suggests that moral mate choice vs. pathogen avoidance), many of the procedures that
disgust serves individual fitness interests by communicating condemnation
give rise to disgust are predicted to differ. We suggest that these
of rule violators with other people. In addition, the functional model
computational processes occur nonconsciously but that they lead
suggests that behaviors eliciting sexual and pathogen disgust feed into
systems strategically sifting for behaviors to condemn, addressing the to consciously accessible states (the experienced sensation of
functionally distinct question of why some disgusting acts are perceived as disgust).
morally wrong. The evolutionary analysis of disgust we offer thus builds on
components of the RHM model—most notably the idea that
disgust has multiple functions, one of which is to deter contact
According to this perspective (notably provided before the germ with pathogenic substances— but differs markedly in that we
theory of disease was widely disseminated), disgust is both the specify the selection pressures that led to the expansion of
reason for the response (the object is revolting) and the output of disgust into domains beyond pathogen avoidance and an engi-
the response. Similarly, the animal reminder perspective suggests neering account of how each function is executed.
that people feel disgust toward behaviors that threaten to remind
people that they are animals, but the disgust response appears to Pathogen Disgust: The Avoidance of
also serve as the criteria by which a behavior is identified as Disease-Causing Organisms
reminding people that they are animals (so, some animal behav-
iors, such as urinating and copulating, are argued to remind people Selection Pressures
that they are animals because they elicit disgust, whereas other
behaviors that are no less animal, such as breathing and sleeping, Parasitic microorganisms, though small in scale, pose large
are not argued to remind people that they are animals because they selection pressures on all long-lived, multicellular organisms.
do not elicit disgust). Further, RHM (2008) argued that (in the Pathogens’ rapid reproduction at the expense of their hosts creates
United States, at least) moral disgust is elicited by sleazy acts. a coevolutionary arms race between host avoidance and pathogen
68 TYBUR, LIEBERMAN, KURZBAN, AND DESCIOLI

transmission (Ewald, 1994; Nesse & Williams, 1994). The dy- hosts through ingested food and water. Infectious disease trans-
namic of pathogen– host interactions has been implicated as a mission can also occur during anal penetration (Varghese, Maher,
major selective force in the evolution of phenomena ranging from Peterman, Branson, & Steketee, 2001).
sexual reproduction, cellular differentiation, and an immune sys- 4. The genitals. The sex organs are designed to transport
tem (Fumagalli et al., 2011; Ridley, 1993; Tooby, 1982) to the gametes between organisms, which creates cross-body pathways
decorative plumage on birds (Hamilton & Zuk, 1982), to person- that can be exploited by micropredators. For instance, Chlamydia
ality and cultural variation (Mortensen, Becker, Ackerman, Neu- trachomatis gains access to new hosts via sexual fluids, as do
berg, & Kenrick, 2010; Schaller & Murray, 2008; Thornhill, many other sexually transmitted diseases (Holmes et al., 2007).
Fincher, Murray, & Schaller, 2010). Like a wide range of other To avoid the costs of infection, humans need counteradaptations
taxa, including roundworms, lizards, and chimpanzees, humans to prevent pathogen transmission. These mechanisms must defend
have elaborate defensive physiological, cognitive, and behavioral the key entry points, such as the mouth and skin, and they need to
adaptations that evolved because of the selection pressures posed avoid other individuals’ exit points and the substances leaving
by pathogens (Curtis, 2007; Curtis et al., 2011; Kluger, Ringler, & these points, such as sneezes and feces. Pathogen disgust offers a
Anver, 1975; Schaller & Duncan, 2007; Schaller & Park, 2011; first line of defense by physically steering people away from
Zhang, Lu, & Bargmann, 2005). We argue that pathogen disgust is conspecifics, nonhuman animals, objects, and locations that could
one such adaptation and serves as an initial line of defense against make them vulnerable to contact with pathogens. Naturally, max-
pathogen infection. Pathogen disgust largely captures what others imum pathogen avoidance, such as never eating, touching others,
have called core disgust (Rozin et al., 2008), primary disgust or having sex, would have carried substantial fitness costs. To
(Marzillier & Davey, 2004), theoretical disgust (Nabi, 2002), pure make adaptive trade-offs, psychological mechanisms need to esti-
disgust (Zhong & Liljenquist, 2006), and basic disgust (Chapman mate magnitudes of pathogen-transmission risk and weigh them
et al., 2009; Inbar, Pizarro, & Bloom, 2009). We suggest that the against magnitudes of expected benefits associated with engaging
label pathogen disgust best describes this domain because it refers in these other behaviors. We detail these computational mecha-
to the system’s evolved function and describes the types of cues nisms below.
taken as input.
Infectious microorganisms cannot rapidly travel large distances Information Processing Architecture of Pathogen
the same way that other, larger predators (e.g., tigers) can. To
Avoidance
move to a new host, pathogens need to exploit points of contact
between habitable regions of their current and future hosts. The At least three sets of information processing systems are needed
following areas of animal bodies are particularly useful as exit and to avoid pathogens: (a) perceptual systems that monitor the envi-
entry points. ronment for cues that reliably correlate with pathogen presence, (b)
1. The mouth. The mouth is a primary source of transit intermediate computational processes that integrate the probability
utilized by many micropredators. For example, Mycobacterium of pathogen presence with other factors that should influence the
tuberculosis exits a host via saliva droplets that are expelled by intensity of the pathogen disgust response and then output this
speaking, coughing, and sneezing and enters a new host through integrated information, and (c) a system that sets in motion the
the mouth when airborne droplets are inhaled. The mouth’s diges- specific psychological and physiological processes involved in the
tive function also provides useful avenues of food-borne infection. pathogen avoidance response (see Table 1).
Microbes such as Escherichia coli can gain entry into the body Inputs: Cues to pathogen presence. Although most infec-
through the mouth when infected animal products (e.g., meat, tious microorganisms are too small for humans to observe directly,
wastes) are ingested as food. In addition to exploiting behaviors we are able to detect various properties reliably associated with
such as breathing and eating, pathogens can induce host behaviors their presence. For instance, pathogens flourish in some environ-
that facilitate transmission, such as sneezing and vomiting. For ments more than others— generally those that provide a source of
instance, Vibrio cholerae uses the mouth as a digestive exit nutrients and minimal antipathogen defenses (e.g., corpses).
through vomit. Over evolutionary history, natural selection likely favored per-
2. The skin. As the surface of the body, the skin is an obvious ceptual systems that reliably detect those properties associated
launchpad for infectious attacks. Transmission can occur through with pathogen presence. Color, for example, reliably indicates
skin-to-skin contact as well as through skin lesions that exude pus whether fruit is unripe, ripe, or rotten due to the effects that
or blood. For example, the varicella zoster virus (chicken pox) infectious microorganisms have on fruit pigmentation (Giovan-
travels via contact with infected skin lesions and moreover causes noni, 2001). Other visual cues, including texture and viscosity, can
itching that promote skin-based transmission to another host. Fur- identify potentially pathogenic substances (e.g., blood, pus, semen,
ther, pathogens that do not directly infect via the skin can be feces; see Curtis et al., 2004). Similarly, the visual cues associated
housed on the skin and make their way into the body via the other with “creepy crawlers” (e.g., maggots and worms) can act as a time
entry points discussed. For example, fecal contamination of un- stamp of the length of time an organism has been deceased and
broken skin may not directly cause infection. However, touching hence left undefended against infectious microorganisms. Unlike a
the mouth or eyes with this skin can lead to infection. freshly killed animal, a body that has been dead longer will have
3. The anus. The excretory function of the anus makes it well a greater probability of having been successfully colonized by
suited for pathogen exit. After exiting the body in feces, infectious micro- and macro-organisms. Thus, the presence of maggots, flies,
agents can gain entry into a new host through the mouth, such as and worms indicates a greater likelihood of bacterial contamina-
from contaminated water. For instance, Salmonella enterica, tion. Olfactory, gustatory, and tactile cues also indicate the pres-
which causes typhoid fever, exits the body in feces and enters new ence of pathogens. For instance, research on olfaction and disgust
 DISGUST FUNCTION 69

Table 1
Information Processing Structure of the Functional Domains of Disgust

Cues that indicate

probable presence of Additional factors Computed internal
Adaptive problem adaptive problem modulating response Expected value estimator regulatory variable Output

Avoiding contact Bodily fluids and Nutritional state, Integrates contact Expected value of Pathogen disgust: Sensation
with infectious products, animals, relationship to cue avoidance benefits contact of disgust, disgust facial
disease-causing poor hygiene, source (e.g., (reduced infection risk) expression, motivation to
organisms decomposing or friendship, kinship), with costs (e.g., not avoid contact, and
rotting organic sexual arousal eating, not having sex) deployment of other
matter functionally coordinated
systems according to the
magnitude of the expected
value of contact

Avoiding sexual Sexual interest from Availability of Integrates sexual Expected sexual Sexual disgust: Sensation
contact with individuals with alternative mates, avoidance benefits value of disgust, disgust facial
individuals poor genetic one’s own mate (reproductive expression, motivation to
jeopardizing compatibility (kin) value (opportunity opportunities saved for avoid sexual contact,
fitness and with low mate costs) better partner) with costs and deployment of
value (e.g., missed functionally coordinated
reproductive systems according to
opportunities, impaired the magnitude of the
social relationship with expected sexual value
target)

Communicating Behaviors likely to Relationship to Integrates condemnation Expected Moral disgust: Motivation
and coordinating be condemned by target (e.g., benefits (mitigating condemnation to signal condemnation of
condemnation others (lying, friendship, kinship) conflict escalation) with value the behavior/person and
with other people cheating, stealing, costs (e.g., impaired facial and vocal expressions
rule violations) social relationship with of disgust
target)

has identified the specific chemical compounds in feces that elicit are predicted to be biased toward having a relatively low threshold
disgust (e.g., Wicker et al., 2003). Sour taste buds can detect lactic for acting on ambiguous cues to infection risks (Ackerman et al.,
acid, an indication of elevated bacterial concentrations (DeSimone, 2009; Kurzban & Leary, 2001; Neuberg, Kenrick, & Schaller,
Lyall, Heck, & Feldman, 2001). Touch also provides information 2011; Oaten et al., 2011; Park, Faulkner, & Schaller, 2003; Park,
regarding moisture content and animacy, two factors predictive of Schaller, & Crandall, 2007; Schaller, Miller, Gervais, Yager, &
both pathogenic growth (Lovanh, Cook, Rothrock, Miles, & Sis- Chen, 2010; Schaller & Park, 2011).
tani, 2007) and disgust reactions (Oum, Lieberman, & Aylward, The possibility of pathogen presence should not lead to auto-
2011). matic behavioral avoidance across all contexts. Instead, the ex-
Intermediate computational processes: The probability of pected, probabilistic costs of pathogen infection should be set
pathogen presence and the expected value of contact. Once against other exigencies, such as acquiring nutrients and calories
pathogen detection systems have responded to incoming cues, by ingesting the soon-to-be rotting flesh of a dead animal. For this
additional systems assess the likelihood of the presence of an reason, additional procedures are needed to integrate the computed
actual threat. Much like a blip on a submariner’s sonar, perceptual pathogen threat with other indices that influence the costs versus
activation can be caused by environmental noise or a true signal. benefits of avoiding pathogens in a given context. Consider sexual
For this reason, we posit a set of systems that integrate the detected behavior as an example. For sexual behaviors to take place—
pathogen cues, perform a signal detection analysis, and generate an behaviors that bring individuals into contact with foreign patho-
internal estimate of the probability of pathogen presence. The gens via physical proximity with a conspecific and exposure to
signal detection process that occurs should be tuned to reflect the various bodily fluids—the pathogen avoidance response must be
costs of committing two possible errors: Type I (assuming a threat down-regulated. This could be engineered by a system that inte-
when none is actually present—a false alarm) and Type II (failing grates estimates of pathogen presence with other information re-
to detect a threat when one is present—a miss; Haselton & Nettle, garding sexual arousal and then outputs an expected value of
2006; Nesse, 2005; Tooby & Cosmides, 1990). In the context of contact that reflects the cost– benefit ratio of contact versus avoid-
pathogen detection, a miss involves high costs, including the costs ance in a given context.
of the immune response to fight the infection (e.g., metabolic Under the model we are proposing, the magnitude of pathogen
costs; damage caused by inflammatory responses), any tissue disgust depends on the computed expected value of contact. In-
damage done by the pathogen, and, in some cases, death. In deed, we claim that the sensation of disgust might just be the felt
contrast, the costs of a false alarm, although non-zero, might output of this computation. Critically, this value should reflect the
include what are often relatively small costs, such as proximal trade-offs inherent in avoiding pathogens versus obtaining other
avoidance or declining to ingest a food. Generally, then, humans benefits. For instance, pathogen avoidance might be traded off
70 TYBUR, LIEBERMAN, KURZBAN, AND DESCIOLI

against the benefits of helping kin or obtaining nutrients when be very specific and tailored toward retrieving information
scarce. Stevenson and Repacholi (2005), for example, found that about items ingested because of the causal relationship between
the level of disgust associated with a pathogen-related odorant ingesting food substances and pathogenic infection (Garcia,
depends on the source of the odorant, with kin eliciting signifi- Kimeldorf, & Koelling, 1955). Thus, upon feeling nausea, for
cantly less disgust than strangers. Importantly, such flexibility instance, one’s memory systems should be organized to recall
requires that cues to kinship modulate the pathogen disgust re- substances ingested rather than pathogen-irrelevant conditions,
sponse, an outcome that is possible if the probability of relatedness such as the weather or the color of one’s clothing. In addition,
(see, e.g., Lieberman, Tooby, & Cosmides, 2007) serves as an pathogen disgust should regulate attention (directed toward
additional input to systems computing the expected value of con- avenues of pathogen transmission), conceptual frameworks (to
tact. Similarly, foods with cues to pathogen presence might be activate concepts such as cleanliness, health, and infection),
avoided when alternative foods are available but consumed when reasoning (to infer that if one has been touched, one could be
they are not, as in times of prolonged famine (e.g., Hoefling et al., contaminated), learning mechanisms (to encode cues that pre-
2009). Hence, internal states correlated with hunger (e.g., low dict the transmission of pathogens such as the “ah-ah-ah” that
blood-glucose levels) might serve as yet other inputs to a system precedes the “chooo”), and so on in a manner that would have
that computes the expected value of contact. Internal states indi- aided in the neutralization of pathogen threats (Ackerman et al.,
cating greater vulnerability to pathogens, including progesterone, 2009; Faulkner, Schaller, Park, & Duncan, 2004; Mortensen et
can act as an additional input to modulate the disgust response al., 2010; Navarrete & Fessler, 2006; Park et al., 2003; Tybur,
(Fessler et al., 2005; Fleischman & Fessler, 2011). In general, the Bryan, Magnan, & Caldwell Hooper, 2011).
modular architecture we are proposing allows for flexibility in the Pathogen disgust is also associated with various physiologi-
disgust response by virtue of the variety of inputs required to cal responses posited to reduce the probability that pathogenic
compute an expected value of contact (see Barrett and Kurzban, substances will enter the body and cause infection. The proto-
2006, for a discussion on modularity). typical disgust facial expression appears to restrict airflow
Outputs: The evolved response to pathogen presence. The through the nose, to reduce the exposed surface area of the eyes,
expected value of contact is predicted to initiate the cascade of and to prevent potentially infectious substances from entering
events required to neutralize the threat of pathogen infection. A the mouth (Fessler & Haley, 2006; Rozin, Lowery, & Ebert,
system well designed to perform this function should activate the 1994; Rozin et al., 1995; Susskind et al., 2008). Disgust is often
behavioral, cognitive, and physiological processes that help defend accompanied by nausea, a desire to vomit, and a loss of appetite
against the various modes of pathogen transmission. A key (per- (Rozin & Fallon, 1987; Rozin et al., 2008), all of which may
haps the key) response of pathogen disgust is the motivation to contribute to ridding the body of pathogens that have already
avoid contact with mouths, skin surfaces, anuses, genitals, and been ingested or to decreasing the probability of ingesting
other surfaces, portals, or substances capable of transmitting materials housing pathogens.
disease-causing organisms. Avoiding the spray of sneezes and Perception of pathogen cues also appears to prime our immu-
coughs and contact with skin lesions or body fluids would have nological defense systems. For example, Stevenson, Hodgson,
reduced the probability of pathogen infection. Indeed, comparisons Oaten, Barouei, and Case (2011) recently observed increases in
of action tendencies associated with several affective states indi- salivary tumor necrotizing factor alpha, a cytokine that plays a role
cate that disgust is uniquely related to physical avoidance and in the inflammatory process that clears infection, in participants
pushing away (Hertenstein, Keltner, App, Bulleit, & Jaskolka, exposed to cues to pathogen presence. Similarly, Schaller et al.
2006; Roseman, Wiest, & Swartz, 1994). Myriad studies have (2010) demonstrated that, relative to participants in a control
shown that, when individuals possess cues for pathogen presence, condition, participants who viewed images of faces sneezing or
they are proximally avoided and/or stigmatized (Kleck, 1969; marked with pox produced higher levels of interleukin-6 (IL-6), a
Kurzban & Leary, 2001; Oaten et al., 2011; Schaller & Duncan, cytokine that plays a key role in the inflammatory process that
2007; Worthington, 1974) and that contextual or chronic motiva- clears infections. Hence, visual cues to pathogen presence prepare
tions to avoid disease exacerbate such distancing (Mortensen et al., the body to fight pathogenic invaders. Notably, differences in IL-6
2010; Park et al., 2003, 2007). observed by Schaller et al. were not mediated by differences in
The pathogen disgust system should also adaptively regulate self-reports of disgust toward the stimuli. Instead, self-reports of
other psychological processes that could aid in avoiding contact disgust were inversely related to IL-6 activation among partici-
with pathogens. For instance, (carefully) gathering more informa- pants who viewed pathogenic faces, suggesting that individuals
tion about possible sources of contaminants would have been may follow distinct strategies for countering pathogen threats, with
beneficial, especially under conditions of uncertainty. Additional some preparing a strengthened immune response and others in-
scrutiny with the senses or social communication might help vesting in avoidance.
corroborate suspicions of pathogen presence; this might account In sum, there are numerous psychological abilities and physio-
for the otherwise odd phenomenon in which people, on eating logical systems that could be activated to solve the problem of
something that does not taste right, seem inclined to get a second pathogen transmission. Identifying those that might have assisted
opinion (e.g., “Does this milk taste spoiled to you? Here . . .”). in the avoidance of pathogen threats can help generate hypotheses
In the case that a pathogenic substance has been ingested and about the structure of pathogen disgust. Importantly, activation or
has caused a physiological response such as nausea or expul- deactivation of each system should contribute to the specific
sion, memory systems should be engaged to store the likely function of defending against pathogens. That is, rather than sug-
source of the contaminant (see, e.g., Seligman & Hager, 1972, gesting that disgust functions to generally “protect the self” (e.g.,
on the “sauce béarnaise” effect). These memory traces should Miller, 2004), we posit that the disgust response is tailored to the
 DISGUST FUNCTION 71

particular problem of avoiding pathogens and that evidence of this form of reputational damage and direct aggression from intra-
functionality should be apparent in the activation of the various sexual competitors (see Hoffman et al., 2008; Penn & Smith, 2007;
psychological and physiological programs. Beyond the systems Perilloux et al., 2010; Tybur & Gangestad, 2011). Sexual interac-
mentioned above, we point out, additional features of a pathogen tions with the vast majority of organisms in the ecology impose
disgust system include the calibration processes that guide the such probabilistic costs without yielding any potential reproduc-
learning of which cues are indicative of pathogens during devel- tive benefits. For example, sex with members of other species,
opment and the feedback processes that disengage the pathogen individuals of the same species who are post-reproductive (i.e., the
disgust response. elderly) or pre-reproductive (i.e., children), and members of the
As others have suggested, disgust seems to have expanded to same sex cannot yield reproductive benefits yet still extract non-
operate in domains beyond pathogen avoidance. Here we argue zero (and potentially substantial) costs.
that the pathogen disgust system was co-opted and modified for Perhaps more important, sex entails significant opportunity
the additional function of avoiding biologically costly sexual part- costs. Time and energy spent courting, copulating, and raising
ners, leading to a distinct and distinguishable sexual disgust adap- offspring with one individual generally cannot be invested in
tation. acquiring other mates or other fitness-enhancing behaviors (see
Kenrick, Griskevicius, Neuberg, & Schaller, 2010). Hence, selec-
Sexual Disgust: The Avoidance of Fitness-Jeopardizing tion should have fine-tuned psychological adaptations for estimat-
ing the projected fitness value of a particular individual as a sexual
Sexual Partners
partner given the alternatives present. Critically, males and fe-
Despite the long-standing recognition that disgust plays an males have different opportunity costs on average—that is, they
important role in human sexuality (Angyal, 1941; Mosher & are typically precluding different amounts of prospective alterna-
O’Grady, 1979; Tomkins, 1963) and more recent work demon- tive reproduction by engaging in sex that leads to fertilization
strating empirical links between certain sexual behaviors and dis- (Trivers, 1972). Whereas females, in producing one offspring, are
gust (Ackerman, Kenrick, & Schaller, 2007; Borg et al., 2010; precluded from the production of another offspring due to their
Fessler & Navarrete, 2003a, 2004; Haidt et al., 1994; Koukounas minimum investment of 9 –10 months gestation and subsequent
& McCabe, 1997; Lieberman et al., 2007; Park, 2008; Schaich nursing, a male need only minimally invest the trivial time and
Borg, Lieberman, & Kiehl, 2008; Stevenson, Case, & Oaten, 2011; metabolic resources needed to inseminate the female. These dis-
Tybur et al., 2009; Vonderheide & Mosher, 1988), no consensus parate selective forces should have generated psychological struc-
has been reached on the precise function or functions of sexual tures with different sensitivities to the costs of committing versus
disgust. For example, Goldenberg et al. (2000) implied that sexual forgoing various sexual opportunities (Buss & Schmitt, 1993).
avoidance functions to neutralize existential threats: “Because the Here we discuss two main dimensions that contribute to the
human species has intercourse and reproduces just as other animals opportunity costs involved in selecting one sexual partner over
do, the physical aspects of sex make apparent our animalistic another: genetic compatibility and mate quality (Jennions & Petrie,
creaturely nature” (p. 206). Similarly, Rozin et al. (2008) consid- 2000; Neff & Pitcher, 2005; Zeh & Zeh, 1996). Genetic compat-
ered sex a prototypical example of animal reminder disgust, and ibility has been defined as the nonadditive genetic variance in
they argued that sexual disgust functions to motivate rejection of fitness (Puurtinen, Ketola, & Kotiaho, 2009). Put differently, it is
reminders that human bodies are similar to animal bodies. As a relative measure of how compatible (fitness-promoting) another
discussed above, this perspective runs into difficulties when one individual’s genotype is with one’s own. The most notable in-
considers that humans breathe and sleep just as other animals do; stances of incompatibility involve close kin, who are much more
yet these other “animal” activities are not posited to make apparent likely than nonrelatives to share identical deleterious recessive
our animal nature, nor do they elicit disgust (Royzman & Sabini, alleles that could jeopardize the health of resulting offspring
2001; Tybur et al., 2009). (Bittles & Neel, 1994; Charlesworth & Charlesworth, 1999; Haig,
But the question remains: Why do some sexual acts elicit 1999). By virtue of producing offspring who are more similar
disgust whereas others elicit sexual arousal? For example, a 20- biochemically to their parents, inbreeding also increases the prob-
year-old man might find the thought of sexual intercourse with an ability that pathogens will disrupt offspring development (Tooby,
80-year-old woman disgusting, but the exact same sexual act with 1982). Other factors can influence genetic compatibility, including
an 18-year-old female lingerie model would elicit a markedly immunological compatibility, which can be assessed via chemical
different response. The animal reminder construct promoted by signatures associated with genes that govern the immune response
Rozin et al. (2008) cannot account for these differences, as both (e.g., major histocompatibility complex; see Ober et al., 1997;
acts are “animalistic” (Tybur et al., 2009). We propose an alter- Thornhill et al., 2003; Wedekind & Furi, 1997).
native perspective: that sexual disgust functions to motivate the Mate quality, on the other hand, is a composite of the nonrela-
avoidance of sexual behaviors with partners imposing potentially tive dimensions along which potential sexual partners may vary.
high fitness costs. One such dimension, intrinsic genetic quality—that is, alleles that
increase fitness in an additive fashion— generates what has come
Selection Pressures Imposed by Sexual Decision to be known as “good genes” (Neff & Pitcher, 2005). Evolutionary
biologists and psychologists have proposed that various visible
Making
aspects of the phenotype— generally, what is perceived as “attrac-
Although sexual intercourse is necessary for reproduction, it tive” (Grammer, Fink, Møller, & Thornhill, 2003)— can carry
carries several direct costs, including tissue damage during inter- information regarding intrinsic quality (Møller & Swaddle, 1997;
course, the risk of pathogen transmission, and social risks in the Singh, 1993; Thornhill & Gangestad, 1993, 2006). Other dimen-
72 TYBUR, LIEBERMAN, KURZBAN, AND DESCIOLI

sions also contribute to the magnitude of opportunity costs posed Information Processing Architecture of Sexual
by potential sexual partners. For instance, the ability and willing- Avoidance
ness to invest in offspring, current health, intrasexual competitive
ability, and residual reproductive value (e.g., age) would all factor Sexual avoidance requires a different suite of mechanisms com-
into mate quality (though these dimensions may not be entirely pared to pathogen avoidance. Here we posit: (a) perceptual sys-
independent of good genes). tems that take cues associated with genetic compatibility and mate
In summary, there are multiple selection pressures that would value as input; (b) intermediate computational processes that in-
have shaped sexual decision making. The presence of individuals tegrate information regarding genetic compatibility, mate value,
who varied in terms of their genetic compatibility, genetic quality, and other context-dependent indices to compute for each individ-
and other attributes known to govern sexual attraction (Buss, 1992; ual an expected value as a sexual partner, which reflects the
Ellis, 1992; Thornhill & Gangestad, 1999) created the adaptive expected fitness outcome of avoiding versus selecting that indi-
problem of avoiding those who posed the greatest reproductive vidual as a sexual partner; and (c) a system that uses this expected
threats. We propose that disgust, which initially evolved as a value to set in motion the specific psychological and physiological
motivator of pathogen avoidance, was co-opted to motivate the processes involved in sexual avoidance (see Table 1).
avoidance and rejection of sexual contact with those whose sexual Inputs: Cues to compatibility and quality. Just as pathogen
value was relatively low among the potential pool of mates. avoidance systems monitor for cues associated with pathogen
Critically, given the nature of these adaptive problems, the inputs presence, mate selection systems monitor for cues associated with
that lead to sexual disgust should differ from those involved in the genetic compatibility and mate value. Kinship is one important
pathogen disgust system. Before we turn to an information pro- dimension contributing to genetic compatibility. Recent research
cessing model of sexual disgust, though, we briefly address the has demonstrated that multiple cues are used to assess the relat-
question, Why disgust? edness of others. These cues include childhood coresidence dura-
Sexual attraction and arousal fulfill important functions in mat- tion (Bevc & Silverman, 1993, 2000; Fessler & Navarrete, 2004;
ing decisions: They motivate courtship, copulation, and pair bond- Lieberman, Tooby, & Cosmides, 2003; Lieberman et al., 2007),
ing with individuals of high sexual value. The absence of attraction exposure to one’s mother caring for a newborn (Lieberman et al.,
and arousal could thus potentially perform the function of steering 2007), and perhaps facial and attitudinal similarity (DeBruine,
individuals away from mates of low sexual value. However, the 2005; Park & Schaller, 2005). As suggested by a recent model of
absence of sexual arousal would not prevent that individual from kin detection (Lieberman et al., 2007), kinship cues feed into a
being sexually pursued by other people who possess their own kinship estimator that computes an index of genetic relatedness.
reproductive agendas. To reject and avoid unwanted sexual ad- This index, we suggest, is one input to systems assessing sexual
vances and behaviors another response is required. Emotions such value.
as fear and anger are not well suited to avoiding potentially costly Other aspects of genetic compatibility require dedicated detec-
mates. Fear can lead to immobilization or rapid flight (Öhman &
tion systems. The extent to which one differs immunologically
Mineka, 2001) the former of which likely does not impede sexual
might be computed by a separate system. For instance, humans,
pursuit, and the latter of which is metabolically costly and often
primarily women, prefer the scent of individuals with dissimilar
unnecessary (e.g., if social allies and kin can prevent another’s
ajor histocompatibility complex (MHC) types (Thornhill et al.,
sexual interest from progressing to sexual aggression). Similarly,
2003; Wedekind & Furi, 1997). This preference, at least in mice,
anger often acts as an “approach” emotion (Carver & Harmon-
appears to be calibrated during development (Penn & Potts, 1999;
Jones, 2009), and associated aggression can lead to costly coun-
Yamazaki et al., 1988) and might represent in humans a separate
teraggression (Sell et al., 2009).
dimension along which potential sexual partners are evaluated.
We suggest that the phylogenetically ancient (Curtis, 2007;
Zhang et al., 2005) pathogen disgust was a felicitous system to Other systems should be functionally specialized for detect-
co-opt to perform the function of avoiding biologically costly ing and assessing traits contributing to mate quality. With
mates. First, pathogen disgust motivates withdrawal and physical respect to intrinsic quality, there might be systems that assess
avoidance, cost-effective behaviors that solve the problem of stay- symmetry, skin tone, skin texture, and so forth (Fink, Grammer,
ing out of arm’s reach (we note that flight motivated by fear could & Thornhill, 2001; Grammer et al., 2003; Scott, Pound, Ste-
also contribute to sexual avoidance when a sexual threat involves phen, Clark, & Penton-Voak, 2010; Symons, 1979). Many such
active pursuit). Second, pathogen avoidance is already linked to dimensions will be used to assess the quality of potential mates
sexual behavior. If other individuals and their bodily fluids repre- for both males and females (B. C. Jones et al., 2001; Thornhill
sent planets of potentially infectious microorganisms, then patho- & Gangestad, 2006; Zebrowitz & Rhodes, 2004). Because the
gen disgust must be down-regulated for sex to occur (Angyal, properties that influence mate value vary across the sexes, some
1941). Indeed, sexual arousal appears to dramatically down- of the features that determine mate quality differ between men
regulate pathogen disgust, as evidenced by particular sexual fe- and women (Buss, 2003). Whereas some dimensions might be
tishes (e.g., eproctophilia and coprophilia; Symons, 2007; see also given greater weight by males in assessing the qualities of
Borg & de Jong, 2012). If instead of being down-regulated, disgust females (e.g., cues that relate to youth and fertility, Buss, 1989;
was up-regulated in response to particular traits (e.g., kinship Kenrick & Keefe, 1992; Singh, 1993), others might be given
estimates; see below), evolution would have been well on its way more weight by females in assessing the qualities of males (e.g.,
to fashioning a sexual avoidance mechanism from a pathogen cues of direct intrasexual competitiveness; Gangestad, Simp-
avoidance mechanism. son, Cousins, Garver-Apgar, & Christensen, 2004; Little, Jones,
 DISGUST FUNCTION 73

& DeBruine, 2008; Puts, 2010; Snyder, Kirkpatrick, & Barrett, individual with low sexual value should not. Take, for example, a
2008). woman’s father. Despite having cues for poor genetic compatibil-
Intermediate systems for integrating intrinsic quality and ity, a father should not automatically elicit sexual disgust. Rather,
genetic compatibility. Once cues to genetic compatibility and sexual disgust should be aroused only if sexual behavior is con-
mate quality have been detected and assessed, they must be com- sidered by the offspring or offered by the father (Westermarck,
bined in a manner that allows for the adaptive regulation of mating 1891/1921). The lack of a constant state of sexual disgust toward
decisions. One possibility is that they combine to produce an poor mates reflects the costs associated with avoiding individuals
expected value as a sexual partner (see also Tooby and Cosmides, who are otherwise valuable social partners. Although constant
2008). The expected sexual value system outputs different mag- motivations to avoid poor mates would certainly decrease the
nitudes depending on inputs. When below the minimum threshold probability of reproducing with them, it would also cripple some
for acceptability as a mate, it outputs a magnitude that is felt as beneficial social relationships. Recent research indicates that
sexual disgust, which serves as input to systems motivating sexual avoidance behaviors are indeed not constant but instead might
avoidance. For an individual with a high degree of relatedness and fluctuate in adaptive ways (e.g., Lieberman, Pillsworth, & Hasel-
a high mate quality (e.g., a physically attractive sister), information ton, 2011).
about relatedness should generally trump information regarding In addition to distinct behaviors for sexual avoidance, there are
mate value due to the costs associated with inbreeding, leading to likely domain-specific processes that are activated in response to
a low expected sexual value. This decision should also take as possible or considered sexual engagement with individuals of low
input other factors such as the availability of alternative mates and sexual value. These include specialized systems for reasoning (if
the pathogen load of the environment (Schaller & Murray, 2008). he smiles at me then he is sexually interested in me), attention
The expected value of an individual as a sexual partner should (directed toward the proximity of interested low sexual value
reflect the integration of all these factors and thus represents an targets), memory (of escape routes, individuals who could come to
index of the context-dependent fitness consequences of pursuing one’s aid), and learning (what locations to avoid when alone).
versus avoiding an individual as a sexual partner. But this only Some of these processes may be shared with pathogen disgust. For
addresses systems regulating one’s own sexual interests. Other example, individuals report feeling nauseated when thinking about
individuals compute their own distinct estimates of expected sex- sibling incest (Royzman, Leeman, & Sabini, 2008), and rape
ual value and can act accordingly. For instance, a 50-year-old man victims often report feelings of disgust and being soiled (Isac &
may find an 18-year-old woman sexually attractive despite her Schneider, 1992; Petrak, Doyle, Williams, Buchan, & Forster,
aversion toward him. Perceived sexual intent from a conspecific 1997). Given that sexual disgust likely evolved from an existing
should therefore interact with expected sexual value to output pathogen disgust system and that many cues that elicit sexual
sexual avoidance. disgust pertain to pathogens (e.g., intrinsic mate value depends
Outputs: The evolved response to partners of low sexual cues signaling the presence of infection), the physiological systems
value. In line with our definition of pathogen disgust, we argue that entrained by pathogen avoidance systems may overlap with those
sexual disgust is just the felt output of computational procedures designed to avoid certain individuals as sexual partners as a
estimating expected sexual value. When sexual value estimates reach by-product.
a lower threshold and sexual interest has been detected, behavioral In summary, we suggest that sexual disgust, rather than stem-
strategies, cognitive mechanisms, and physiological systems that ming from a need to avoid reminders of our animal nature (Rozin
facilitate sexual avoidance should be activated. We propose that et al., 2008), functions to motivate the avoidance of mates who
the ancient psychological processes involved in avoiding patho- could potentially jeopardize one’s reproductive success. Critical
gens were co-opted for the function of sexual avoidance because of features of our model of sexual disgust include systems that trade
the delicate nature of avoidance that is optimal for sexual interac- off genetic compatibility against other features such as attractive-
tions. Again, the behavioral avoidance of macropredators moti- ness and resource status in an adaptive manner. The outputs of
vated by fear (e.g., rapid flight) is inefficient for neutralizing many these and other systems combine to generate an expected value of
(perhaps most) poor sexual partners, just as it would be inefficient an individual as a sexual partner; depending on magnitude, this
for neutralizing pathogens. Although rapid flight may neutralize variable can initiate attraction or avoidance behaviors. The cascade
such threats, it is metabolically expensive and may preclude ben- of psychological and physiological systems engaged in sexual
efits associated with interacting with individuals (e.g., kin) in a avoidance likely forms a unique set but might share components
nonsexual manner. Certainly, other cues (e.g., ambient darkness; with the pathogen avoidance system (e.g., facial expression and
pursuit from a sexually aggressive conspecific) may interact with withdrawal behaviors).
cues related to quality and compatibility to output rapid flight, but
threats posed by many poor-quality mates can be neutralized with Moral Disgust: Navigating the Landscape of
less costly avoidance.
Condemnation
Although presumably co-opted from initial pathogen avoidance
mechanisms, the collection of systems entrained to avoid an indi- Cross-culturally, actions that otherwise elicit pathogen or sexual
vidual sexually should differ in important ways. Whereas proximal disgust are often moralized. That is, they are viewed as morally
avoidance (i.e., not touching) mitigates the dangers of pathogens, wrong and subject to punishment. Examples include taboos about
specifically sexual avoidance mitigates the reproductive threats food (especially meat; Fessler & Navarrete, 2003b); sexual acts
posed by individuals with low expected sexual value. Similarly, such as incest, masturbation, bestiality, pedophilia, prostitution,
physical contact with an object possessing pathogen cues should and homosexuality; and behaviors having to do with liquid and
elicit avoidance and disgust, whereas physical contact with an solid bodily wastes (Douglas, 1966; Miller, 1997). Further, people
74 TYBUR, LIEBERMAN, KURZBAN, AND DESCIOLI

report and display disgust toward moralized acts unrelated to basic moral foundation, purity/sanctity, “is based on the emotion of
pathogen threats or sexuality. Danovitch and Bloom (2009), for disgust in response to biological contaminants (e.g., feces or rotten
instance, found that children as young as 6 years old label moral food), and to various social contaminants like spiritual corruption, or
violations related to harm and fairness as “disgusting” both ver- the inability to control one’s base impulses” (Koleva, Graham, Iyer,
bally and via identification of facial expressions. In adults, recent Ditto, & Haidt, 2012, p. 185). Examples of purity violations include
EMG studies have demonstrated the activation of facial muscles a man having sexual intercourse with a chicken carcass, a brother and
associated with pathogen disgust (e.g., the levator labii) after sister having sex, and an avant-garde performance art piece in which
unfair treatment in an ultimatum game (Chapman et al., 2009) and performers act like nonhuman animals and urinate on stage (see Haidt,
while imagining dishonest behavior (Cannon et al., 2011). Finally, 2001, 2006).
fMRI studies have shown common neural activation for responses Haidt (2007, 2012) provided an evolutionary explanation for the
to pathogen disgust and unfair offers in an ultimatum game (San- connection between purity/sanctity and morality, arguing that disgust-
fey, Rilling, Aronson, Nystrom, & Cohen, 2003), theft and vio- based morality functions to improve group cohesion—to “bind” peo-
lence (Schaich Borg et al., 2008), and child abuse (Moll et al., ple into effective groups. Here we propose alternative functions based
2005). These empirical patterns illustrate a close link between on a framework recently used to investigate some of the “mysteries of
disgust and morality, giving rise to a pair of related but separable morality” (DeScioli & Kurzban, 2009, 2012; see also DeScioli, Bru-
questions. The first is why are acts that evoke pathogen or sexual ening, & Kurzban, 2011; DeScioli, Christner, & Kurzban, 2011;
disgust moralized, as in “Having sex with a goat is morally Kurzban, DeScioli, & Fein, 2012; Weeden, 2003).
wrong.” The second is why do violations of moral rules evoke the Selection pressures. Humans live in groups in which viola-
language and facial expression of disgust, as in, “The theft of $23 tions of moral rules are condemned and punished. In addition to
from the Food For Orphans Fund was disgusting.” Here we ad- shaping the evolution of cultures (Boyd & Richerson, 1992; Haidt,
dress both questions. 2012; Richerson & Boyd, 2005), the existence of moralistic pun-
We emphasize that this section is not intended as an extended ishment poses adaptive problems for individuals—avoiding sanc-
treatment of morality (for relevant recent work on this topic, see tions from group members. One solution to this problem is for
DeScioli and Kurzban, 2009, 2012; Haidt, 2007, 2012; Haidt & individuals to strategically influence the content of moral rules by
Kesebir, 2010; Hauser, 2006; Krebs, 2011) but instead as an exami-
supporting or resisting candidate rules based on how the rule, if it
nation of how disgust impacts moral cognition and vice versa. We
were in place, would affect the individual. DeScioli and Kurzban
also note that, in contrast to pathogen and sexual disgust, there is little
(2009, 2012) suggest that decisions to endorse or oppose a partic-
consensus about the measurement, function, and even existence of
ular rule are influenced by the expected impact of the rule on the
moral disgust (see Bloom, 2004; Nabi, 2002; Royzman & Kurzban,
endorser’s fitness (see also Weeden, 2003). Attitudes toward rules
2011; Royzman & Sabini, 2001; Rozin et al., 2008; Rozin, Haidt, &
relating to short-term mating can serve as an example. Rules
McCauley, 2009; Simpson, Carter, Anthony, & Overton, 2006; Tybur
leading to condemnation and punishment of adultery (or, more
et al., 2009). Hence, the ideas discussed in this section are necessarily
generally, casual sex) are especially advantageous for individuals
more exploratory than those in previous sections on pathogen and
invested in a monogamous, pair-bonded mating strategy because
sexual disgust. Nevertheless, the material presented here includes
they recruit the moralized punishment of a group as a barrier to a
three novel insights into the connection between morality and disgust.
First, unlike other treatments on this topic (e.g., Chapman et al., 2009; current or future mate’s infidelity. However, these rules are dis-
Hutcherson & Gross, 2011; Rozin et al., 2008; Rozin, Haidt, & advantageous for individuals whose sexual strategies involve ac-
Fincher, 2009; Tybur et al., 2009), ours separates the relationship quiring multiple mates (Gangestad & Simpson, 2000). Hence,
between disgust and morality into the two questions described above. individuals should endorse rules that favor their fitness interests
Second, we offer an explicit computational perspective, proposing and resist rules that run counter to their fitness interests. Consistent
aspects of the information processing systems required for each pro- with this perspective, individual differences in mating strategy
posed link between disgust and morality (cf. Mikhail, 2007, 2008). predict endorsement of rules constraining others’ sexuality, and
Third, in contrast to recent accounts of moral disgust that have been experimental primes designed to increase the perceived likelihood
couched in group selection processes (e.g., Haidt, 2007, 2012), we of infidelity similarly increase rule endorsement (Kurzban, Dukes,
offer alternative approaches based on individual-level selection pres- & Weeden, 2010; Li, Cohen, Weeden, & Kenrick, 2010; Weeden,
sures (see discussion in Pinker, 2012). Our approach follows the same 2003; Weeden, Cohen, & Kenrick, 2008).
trajectory as the previous sections on pathogen and sexual disgust, Rules can vary tremendously across cultures and times (Sh-
with a consideration of selection pressures and computational pro- weder, Mahapatra, & Miller, 1987), and new rules appear fre-
cesses. quently within groups. For example, in the past century, moral
rules have developed around new technologies such as hormonal
contraceptives, stem cell therapy, cloning, genetically engineered
Disgust as Input to Moral Decisions: Why Are
crops, digital property rights, carbon emissions, cigarette smoking
Disgusting Things “Wrong”? around children, pornography, and taxation on income made from
Haidt (2001, 2006, 2007, 2012) has persuasively argued that moral investments. The introduction of new moral rules heightens the
judgments are often rooted in intuition rather than logical consider- problems associated with endorsing or resisting rules in a manner
ation of the consequences of moral violations (see also Greene, 2007; that favors one’s fitness. Given the rapid introduction of potential
Nichols, 2002). Disgust has been implicated as an important source of moral rules and the general difficulties of forecasting the conse-
these intuitions. Indeed, according to moral foundations theory (Gra- quences of endorsing specific rules, how has selection shaped
ham, Haidt, & Nosek, 2009; Haidt, 2012; Haidt & Joseph, 2004), one moral cognition to endorse rules in a fitness-promoting manner?
 DISGUST FUNCTION 75

As discussed above, pathogen and sexual disgust motivate the ability to enforce the rule, the consistency of the rule with other
avoidance of acts or objects due to fitness costs associated with existing rules, and so on (DeScioli & Kurzban, 2012). Likewise,
infectious disease or mating, respectively. When one contemplates other emotions could provide distinct “intuitions” about the fitness
a given act, then, the extent to which one experiences disgust can costs associated with rule endorsement and explain additional
function as an index of the fitness costs of engaging in the act in features of moral rule content (e.g., Haidt & Joseph, 2004; Rozin
question. The greater the disgust, the more motivated an individual et al., 1999).
should be to avoid that act. Taken together with a consideration of
costs and benefits involved in endorsing rules, this implies that Disgust as Output of Moral Cognition: Why Are
the more the contents of a (potential) moral rule evoke disgust, the
Wrongs “Disgusting”?
greater the probability that the rule would prohibit acts that the
individual would prefer not to do even if the rule were not in place. The previous section proposed an explanation for why people
As a strategic matter, there is little disadvantage to the individual morally condemn acts that elicit (pathogen or sexual) disgust. This
to adopt and spread a rule that prevents such acts. section addresses a separate connection between disgust and moral
Information processing procedures. Our proposal, then, is judgment: the fact that people report being disgusted by acts that they
that disgust intuitions can serve as input to systems that judge the view as morally wrong, even when these acts have no pathogen or
strategic value of endorsing a rule. Computationally, this could sexual content. For instance, people asked to consider behaviors such
occur if felt disgust, an output of systems assessing the expected as stealing a purse from a blind person evaluate such acts as morally
value of contact and expected sexual value described in the patho- disgusting (Hutcherson & Gross, 2011; Nabi, 2002), and individuals
gen and sexual disgust sections above, serves as an input to asked to recall a time that they were “disgusted” often refer to
mechanisms designed to make strategic decisions about which violations of a moral rule related to harm or fairness (Curtis & Biran,
moral rules to endorse and which to resist (see also Pizarro, Inbar, 2001; Haidt et al., 1994, 1997; Tybur et al., 2009). Further, recent
& Helion, 2011.) This computational account is consistent with a evidence suggests that a signature feature of pathogen disgust—the
number of empirical studies investigating the link between disgust canonical facial expression—is observed in domains that have moral
and moral judgment. In general, these investigations have found but no pathogen or sexual content (Cannon et al., 2011; Chapman et
that activating the pathogen or sexual disgust system increases al., 2009). Here, we propose an explanation for this connection
judgments of moral wrongness, presumably because increased between disgust and morality.
disgust marks the decreased fitness costs of condemnation. For Selection pressures: Condemnation coordination. When
example, Schnall, Haidt, Clore, and Jordan (2008) showed that individuals observe other people violate rules that are subject to
participants who smell a disgusting odor judge acts to be more punishment, third parties (observers) are faced with a dilemma.
morally wrong than control participants. Along similar lines, Es- If third parties condemn opposing sides in a dispute, they risk
kine, Kacinik, and Prinz (2011) showed that subjects who experi- creating new enemies and increasing the intensity of the con-
enced a disgusting taste judge acts to be more wrong than subjects flict. That is, as compared to a highly skewed distribution of
who experience a sweet one. Wheatley and Haidt (2005) showed condemners (e.g., when almost everyone condemns the same
that disgust induced under hypnosis similarly increased the extent individual or act), an equal number of supporters on both sides
to which acts are viewed as wrong, and Moretti and di Pellegrino of a dispute can escalate and prolong the costs associated with
(2010) showed that participants rejected low offers in an ultima- the conflict. Coordination of condemnation attenuates costs, but
tum game more frequently after a viewing images that evoked to achieve coordination, individuals must signal (communicate)
pathogen disgust. Activation of sexual disgust similarly shapes whom they condemn to other people, and they must detect other
moral judgments. For instance, Lieberman et al. (2003, 2007) people’s signals of condemnation (see DeScioli and Kurzban,
found that the same kinship cues that predict the development of 2012, for an extensive description of this adaptive problem).
personal sexual aversions toward opposite sex siblings also predict We suggest that expressions of disgust, which are easily,
the strength of one’s moral opposition to third-party sibling incest. rapidly, and cross-culturally recognized (Ekman, 1972; Sauter
This perspective is consistent with moral foundations theory, & Eimer, 2010; Sauter, Eisner, Ekman, & Scott, 2010), might
which suggests that individuals morally judge certain acts because be particularly effective for coordinating condemnation. As
those acts elicit disgust (Graham et al., 2009; Haidt, 2012). proposed above, disgust intuitions act as inputs to wrongness
Importantly, we do not suggest that disgust serves as an input to judgments. Hence, people can use an expression of disgust as a
every moral judgment, nor do we suggest that every act that elicits reliable cue that other people oppose and moralize a given
disgust will inevitably become moralized. There are a multitude of action. In turn, individuals can signal their opposition to an
other computational processes that underlie the moral judgment action by broadcasting a disgust display, even when no patho-
system (Mikhail, 2007, 2008). Some of these processes likely gen or sexual risks are present. By using the vocal and facial
dictate when and how disgust intuitions shape moral rules. For expressions of disgust, third parties signal to others that they
instance, acts must first be assessed as candidates for moralization. oppose the individual who has committed an action they find
When computational processes do not identify an act as a candi- punishable. We contrast this signaling view with the idea that
date for moralization, disgust intuitions ought not to lead to a moral disgust has a function in causing people to avoid some set
moralized disgust. When acts are identified as candidates for of objects, behaviors, or people, as is the case with pathogen
moralization, disgust-driven assessments about the fitness out- and sexual disgust.
come of endorsing versus opposing a particular rule are likely Information processing procedures and expressions of
integrated with information regarding the number and identity/ disgust. There are distinct information processing systems re-
status of group members who agree with one’s assessment, the quired to perform the function of condemnation coordination.
76 TYBUR, LIEBERMAN, KURZBAN, AND DESCIOLI

Naturally, systems must first detect when a rule has been violated such as “a company executive refuses to sit next to a laborer on a
(see Cosmides, 1989, and Delton, Cosmides, Robertson, Guemo, train” (Hutcherson & Gross, 2011). Further, moral violations gen-
& Tooby, 2012, for discussions of detecting cheaters and free erally evoke the motivation to punish wrongdoers as opposed to
riders). Once a violation has been detected, separate systems avoid them (Kurzban, DeScioli, &O’Brien, 2007), although of
requiring additional information must assess the value of broad- course both motivations might simultaneously occur. Finally, a
casting condemnation. One critical input to these systems ought to system designed to identify potentially harmful people might be
be whether other people are present to receive the signal (e.g., expected to attend to the extent to which others intentionally do
Fridlund, 1991). Other inputs might relate to the value of signaling harm, rather than the extent to which they have violated a moral
one’s side in a conflict. For example, when the side one is likely rule. However, a signature feature of moral judgment is that it
to take is uncertain to third parties (perhaps when one’s friends or frequently does not track intended harm (DeScioli & Kurzban,
kin have violated the moral rule), disgust can serve to signal and 2009; Mikhail, 2007).
clarify one’s side. Further, in cases in which one risks condemning A possibility related to this idea and which connects to our
alone, the language of disgust might be deployed as a tool to proposal is that moral disgust functions to motivate social distanc-
recruit punishment from others. That is, because disgust can act as ing, rather than physical distancing, from an individual who has
an input to moral judgment (as detailed above), expressing disgust committed a serious wrong (Curtis & Biran, 2001; Tybur et al.,
toward another individual may help persuade others that the target 2009). If, for instance, those who are perceived as having com-
of disgust expressions should be punished. In general, we suggest mitted moral sins—and their allies—are at risk of being punished
that inputs are integrated to assess an expected value of signaling, by observers (DeScioli & Kurzban, 2012; Neuberg, Smith, Hoff-
as is the case with sexual and pathogen disgust. Once this expected man, & Russell, 1994; Tooby & Cosmides, 2010), there could be
value surpasses a minimum threshold, individuals should activate value in signaling to observers that one condemns the wrongdoer’s
facial and vocal expressions of disgust. actions. Such distancing could defend against being lumped in
Given the arguments detailed above, similarities between the with the condemned individual during subsequent aggression by
pathogen and sexual disgust outputs described previously and moral mobs. However, again, the expressive aspect of disgust may
disgust toward rule violations ought to primarily involve the ex- mitigate this problem as effectively as (if not more effectively
pressive components. That is, the disgust facial expression that than) actual physical distance.
appears designed to shield the body from pathogens or expel
potentially pathogenic material from the mouth (Rozin et al., 2008;
Summary
Susskind et al., 2008) is common across disgust domains, but the
“do not touch” motivation that accompanies pathogen disgust does Here we addressed two separate yet related issues regarding the
not appear to accompany moral disgust. For example, reports of link between disgust and morality. With regard to why disgusting
moral disgust strongly covary with reports of anger, which moti- acts are often moralized, we argued that the pathogen and sexual
vates approach (Carver & Harmon-Jones, 2009), or with desires to disgust systems provide an index of the fitness value of engaging
violently “lash out” at rather than avoid the offending party (Mar- in particular behaviors oneself and that these indices serve as input
zillier & Davey, 2004; Olatunji et al., 2012; Royzman et al., 2008; for systems assessing the costs of condemning third-party behav-
Simpson et al., 2006). Further, although people use some compo- ior. In general, rules prohibiting behaviors that people find dis-
nents of disgust language to describe rule violations, they do not gusting do not substantially encroach on one’s freedom or ability
typically use other terms that correspond with other aspects of to pursue one’s fitness interests. Individuals are thus more likely to
disgust that presumably reflect internal states that function to expel endorse rules that involve punishing behaviors that elicit disgust,
pathogens (e.g., grossed out; see Nabi, 2002). Connections that do and such rules are likely to be agreed upon and prevalent across
seem to support a link between moral disgust and avoidance may groups (see, e.g., Heath, Bell, & Sternberg, 2001). With regard to
be better described in terms of punishment and communication. why violations of a broad array of moral rules elicit expressions of
Chapman et al. (2009), for example, reported some evidence that disgust, we argued that expressions of disgust function to advertise
self-reports and facial signatures of disgust expressed toward un- and coordinate intentions to condemn. Further, if an important part
fair treatment relate to rejection of offers in an ultimatum game. of moral discourse is bringing other third parties around to one’s
Rather than rejecting something in terms of minimizing its contact agenda of imposing costs on others (DeScioli & Kurzban, 2012;
and thus mitigating the possibility of infection, “rejection” in this Tooby & Cosmides, 2010), one would expect moralizers to use
context is a form of punishment. The expressive component of language well suited to recruiting third parties. Because disgust is
disgust might function to communicate intention to condemn and an input to the moral judgment system, agents trying to persuade
potentially recruit others in collective punishment. others about the immorality of an action might use disgust to gain
Alternative views. We emphasize that there are other propos- support against rule violators. On this view, the answer to the
als to explain why people use the language of disgust to refer to question as to why wrong things are “disgusting” is located in the
moral violations. For instance, Hutcherson and Gross (2011) ar- possibility that expressions of disgust serve a persuasive function
gued that “the primary function of both moral disgust and con- on the part of receivers, explaining their use by signalers.
tempt [is] to mark individuals whose behavior suggests that they In closing this section, we note that there is great potential for
represent a threat and avoid them, thereby reducing the risk of future investigations into the link between disgust and morality.
exposure to harm” (p. 720; see also A. Jones & Fitness, 2008; Up to this point, research has primarily focused on establishing
Rozin et al., 2008; Tybur et al., 2009). Some observations seem to whether there is a link between disgust and morality and how that
sit uneasily with this explanation. Moral violations that evoke the link is best measured (e.g., facial expressions, verbal self-reports,
language of disgust do not uniformly relate to harm, as in a case priming effects). Here we differentiate two different disgust and
 DISGUST FUNCTION 77

morality connections, describe these connections using a compu- with other inputs (e.g., cues to kinship, nutritional status, sexual
tational perspective, and propose potential functions for moral arousal), computes expected values of contact, and then outputs
disgust using in a newly developed theory of moral punishment disgust. The model therefore predicts that some of the variation in
and coordination. Future investigations can continue to leverage propensity and intensity of pathogen disgust will stem from indi-
recent advances in evolutionary perspectives on disgust (e.g., vidual differences in (a) the ability to detect cues associated with
Curtis et al., 2011; Tybur et al., 2009) and morality (DeScioli & pathogen threats (e.g., olfactory sensitivity); (b) signal detection
Kurzban, 2009, 2012; Haidt, 2007, 2012) to accelerate our under- analyses for estimating pathogen threat level; (c) how pathogen
standing of moral disgust. threat estimates are traded off against levels of nutrient depletion
(hunger) and other states; (d) how feedback processes return to
Discussion and Future Research Directions baseline after removal of the pathogen threat; and (e) immuno-
competence (e.g., Schaller et al., 2010; Stevenson, Hodgson, et al.,
We began our treatment on disgust with the observation that, 2011). Uncovering potential underlying sources of variation across
although evolutionary theory has implicitly guided much of the the multiple facets of disgust sensitivity might especially benefit
rapidly increasing interest in and research on disgust over the past clinicians dealing with populations suspected of impairments to
20 years, the field has lacked an explicit conceptual framework the disgust system, including individuals with obsessive compul-
that is grounded in evolutionary theory for understanding disgust. sive disorder or individuals with sexual disorders (Borg et al.,
Our goal here has been to address this gap by using a computa- 2010; Olatunji & McKay, 2009; Olatunji & Sawchuk, 2005).
tional approach to understanding emotion (Tooby & Cosmides,
1990, 2008) and by considering research and theory relating to
pathogen avoidance (Curtis et al., 2011; Schaller & Park, 2011), Disgust and Development
sexual decision making (Buss, 1992; Gangestad & Simpson, 2000;
This framework might also help clarify how and when disgust
Thornhill & Gangestad, 2008), and moral condemnation (DeScioli
develops. Pathogen disgust, for example, is notoriously absent (or,
& Kurzban, 2009, 2012; Haidt, 2007, 2012; Mikhail, 2008). In
at least, not fully developed) in young children (see Rozin et al.,
doing so, we have proposed selection pressures and computational
2008). This absence might stem from the fact that, in environments
processes underlying functionally specific disgust domains. Our
similar to those in which humans evolved, infants obtain nutrition
hope is that this attention to the evolved function and associated
directly from their mother in the form of breast milk, often exclu-
computational processes can help synthesize the voluminous ex-
isting literature on disgust and stimulate avenues for novel re- sively until age 2 or 3 (and sometimes until age 4 or 5; Konner &
search. In closing, we briefly sketch some of the implications this Shostak, 1987). In ancestral environments, children were also
framework can have for varied research areas, specifically inves- likely carried throughout the first few years of life, and mothers
tigations into disgust sensitivity, the development of disgust, and might have steered infants away from likely pathogen sources,
cross-cultural variability in disgust. reducing or eliminating the need for the disgust system early in
development. Further, the pathogen disgust system may not func-
tion until an individual reaches other developmental points. For
Individual Differences instance, individuals must acquire culturally evolved information
A substantial portion of disgust research over past decades has regarding those items with high expected benefits of contact (e.g.,
involved empirical observations of covariation between measures edible foods) and the locally adapted hygiene rituals (Schaller &
of disgust sensitivity (i.e., self-reports of how disgusting people Murray, 2008). That is, they may need to acquire information from
find common disgust elicitors) and other constructs such as polit- their social group to deploy disgust in a fitness promoting fashion.
ical ideology (Hodson & Costello, 2007; Inbar et al., 2009; Tybur Considerations such as these can be used to uncover functional
et al., 2010), obsessive and compulsive traits (Olatunji et al., design in the developmental timing of pathogen disgust, which can
2007), and five-factor model personality dimensions (Druschel & in turn contribute to understanding of behavioral antipathogen
Sherman, 1999). Most of these investigations have focused on systems (cf. Schaller & Park, 2011).
domain specificity articulated under the RHM model, an approach Because the other disgust systems described here have different
that has encountered psychometric and conceptual difficulties functions, they should have different developmental trajectories
(Olatunji et al., 2007; Tybur, Bryan, Lieberman, Caldwell Hooper, depending on when in the life span individuals encounter the
& Merriman, 2011; Tybur et al., 2009, 2010). An alternative relevant adaptive problems. For example, sexual disgust might
strategy following the framework we have described involves develop later than pathogen disgust because individuals are typi-
examining variation in three domains independently, separating cally not targeted as sexual partners until they are sexually mature.
pathogen, sexual, and moral disgust. This approach has shown Moral disgust may have yet another distinct trajectory, with de-
early promise, with recent investigations illuminating pathogen velopment coinciding with periods in which rule endorsement and
avoidance, mate preferences, moral inclinations, and aggression condemnation reliably become part of an individual’s social world.
(see, e.g., DeBruine, Jones, Tybur, Lieberman, & Griskevicius, Although previous disgust theories have explicitly referenced do-
2010; Kurzban et al., 2010; Olatujni et al., 2012; Pond et al., 2012; main specificity in function (e.g., Rozin et al., 2008; Tybur et al.,
Tybur et al., 2009, 2010). 2009), investigations have not explicitly tested for domain speci-
Each of the computational processes described here points to a ficity in the timing of development and factors shaping develop-
difference potential source (cause) of individual differences. For ment. The theoretical framework we present here implies very
example, we have suggested that the pathogen disgust system different developmental trajectories for the different varieties of
detects pathogens, integrates the probability of pathogen presence disgust, and it can be used to guide investigations into such domain
78 TYBUR, LIEBERMAN, KURZBAN, AND DESCIOLI

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9280.2005.01614.x Accepted September 11, 2012 䡲

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