Invariances in the architecture of pride across
, Dimitris Xygalatas
, Sarah Alami
, Xiao-Fen An
, Kristina I. Ananyeva
, Shintaro Fukushima
, Alexander N. Kharitonov
, Jeremy M. Koster
, Charity N. Onyishi
, Ike E. Onyishi
Pedro P. Romero
, Kosuke Takemura
, Jin-Ying Zhuang
, Leda Cosmides
, and John Tooby
Department of Psychology, University of Montreal, Montreal, QC, Canada H3C 3J7;
Center for Evolutionary Psychology, Department of Psychological and
Brain Sciences, University of California, Santa Barbara, CA 93106-9660;
Department of Anthropology, University of Connecticut, Storrs, CT 06269;
Department of Anthropology, University of California, Santa Barbara, CA 93106-3210;
School of Psychology and Cognitive Science, East China Normal
University, Shanghai 200062, China;
Institute of Psychology, Russian Academy of Sciences, 129366 Moscow, Russia;
School of Cultural and Creative Studies,
Aoyama Gakuin University, Tokyo 150-8366, Japan;
Faculty of Humanities, Fukuoka University, Fukuoka 814-0180, Japan;
Department of Anthropology,
University of Cincinnati, Cincinnati, OH 45221-0380;
Department of Human Behavior, Ecology and Culture, Max Planck Institute of Evolutionary
Anthropology, 04103 Leipzig, Germany;
Department of Sociology and Anthropology, University of Nigeria, 41000 Nsukka, Nigeria;
Psychology, University of Nigeria, 41000 Nsukka, Nigeria;
Department of Economics, Universidad San Francisco de Quito, Pichincha 17-0901, Ecuador;
Faculty of Economics, Shiga University, Shiga 522-8522, Japan; and
Center for Evolutionary Psychology, Department of Anthropology, University of
California, Santa Barbara, CA 93106-3210
Edited by Steven Pinker, Harvard University, Cambridge, MA, and approved July 6, 2018 (received for review May 16, 2018)
Becoming valuable to fellow group members so that one would
attract assistance in times of need is a major adaptive problem. To
solve it, the individual needs a predictive map of the degree to
which others value different acts so that, in choosing how to act, the
payoff arising from others’valuation of a potential action (e.g., show-
ing bandmates that one is a skilled forager by pursuing a hard-to-
acquire prey item) can be added to the direct payoff of the action
(e.g., gaining the nutrients of the prey captured). The pride system
seems to incorporate all of the elements necessary to solve this adap-
tive problem. Importantly, data from western(-ized), educated, indus-
trialized, rich, and democratic (WEIRD) societies indicate close
quantitative correspondences between pride and the valuations of
audiences. Do those results generalize beyond industrial mass socie-
ties? To find out, we conducted an experiment among 567 participants
in 10 small-scale societies scattered across Central and South America,
Africa,andAsia:(i) Bosawás Reserve, Nicaragua; (ii ) Cotopaxi, Ecua-
dor; (iii) Drâa-Tafilalet, Morocco; (iv) Enugu, Nigeria; (v)LeMorne,
Mauritius; (vi) La Gaulette, Mauritius; (vii ) Tuva, Russia; (viii) Shaanxi
and Henan, China; (ix) farming communities in Japan; and (x)fishing
communities in Japan. Despite widely varying languages, cultures, and
subsistence modes, pride in each community closely tracked the valu-
ation of audiences locally (mean r=+0.66) and even across commu-
nities (mean r=+0.29). This suggests that the pride system not only
develops the same functional architecture everywhere but also oper-
ates with a substantial degree of universality in its content.
Evidence from behavioral ecology, archaeology, and contem-
porary forager societies suggests that our hominin ancestors
evolved in an ecology characterized by high rates of mortality,
scarcity and high variance in food acquisition (1), high incidence
of disease and injury (2), and attacks by predators and conspe-
cifics (3, 4). Reliance on fellow group members, including non-
kin, for the assistance necessary to survive and reproduce is a
distinctively human characteristic (5). Indeed, mutual aid has
been such a universal and basic feature of forager subsistence
that it is believed to be central to the evolutionary biology of our
species. In this social ecology, it would have been essential to
incentivize mates, cooperative partners, and fellow group mem-
bers to value one’s welfare so that they would be inclined to
render assistance in times of hunger, incapacitation, and in-
terpersonal conflict (2). The extent to which fellow group
members valued, helped, and refrained from exploiting an in-
dividual and the extent to which they deferred to the individual
in conflicts of interests would have sensitively impacted whether
that individual reproduced successfully, struggled, or died
In general, there are two classes of bargaining tactics organisms
have available for influencing others’choices. First, they can
conditionally inflict costs—aggression; second, they can bestow (or
withhold) benefits—altruism. The first causes individuals to be
respected (or feared). The second causes individuals to be valued.
Thus, it might be advantageous to put weight on another’s welfare,
(i) because the individual is formidable and could inflict costs if
not propitiated or (ii) because the individual’s actions or existence
make positive fitness contributions to the valuer, which would be
diminished or lost if assistance was not given. Here, we call these
two components respect (for formidability) and valuation (for
positive fitness contributions)—also referred to as dominance and
prestige (7). Being respected and being favorably valued by others
were resources, and selection on our ancestors would have shaped
the human motivational system to cost-effectively promote access
to both of those different types of resources.
Because nonhumans are far more limited in the kinds of as-
sistance that they can render each other, almost all nonhuman
It has been proposed that one key function of pride is to guide
behavior in ways that would increase others’valuation of the
individual. To incline choice, the pride system must compute for
a potential action an anticipated pride intensity that tracks the
magnitude of the approval or deference that the action would
generate among local audiences. Data from industrial mass
societies support this expectation. However, it is presently not
known whether those data reflect cultural evolutionary pro-
cesses or a panhuman adaptation. Experiments conducted in 10
traditional small-scale societies with widely varying cultures
and subsistence modes replicate the pattern observed in mass
societies. This suggests that pride is a universal system that is
part of our species’cooperative biology.
Author contributions: D.S. designed research; D.X., S.A., X.-F.A., K.I.A., S.F., H.H., A.N.K.,
J.M.K., C.N.O., I.E.O., P.P.R., K.T., and J.-Y.Z. performed research; D.S. analyzed data; and
D.S., D.X., L.C., and J.T. wrote the paper.
The authors declare no conflict of interest.
This article is a PNAS Direct Submission.
Published under the PNAS license.
To whom correspondence should be addressed. Email: email@example.com.
This article contains supporting information online at www.pnas.org/lookup/suppl/doi:10.
www.pnas.org/cgi/doi/10.1073/pnas.1808418115 PNAS Latest Articles
bargaining is based on aggression. Differences in the ability to
inflict costs (formidability or resource holding power) led to
adaptations for the advertisement of formidability and adapta-
tions for assessing own and others’formidability (8, 9). In group-
living species, dominance hierarchies emerge from patterns of
deference to those with more formidability—individuals cede
resources or rank to avoid being harmed (10).
Although humans fully retain and exploit phylogenetically
ancient adaptations for aggression and dominance [including
systems for threat, fighting, display, and assessment (8, 10–12)]—
as seen in groups of children, adolescents, and adults (13–16)—
human evolution was distinctive in the greatly expanded role that
mutual assistance played in daily group living, and hence in the
reproductive fortunes of individuals (2, 17). The hominin entry
into the cognitive niche (involving the emergence and integration
of intelligence, language, tool use, coordination, and culture)
greatly amplified the opportunities for mutually advantageous
prosocial interactions (18, 19). As our ancestors entered the
cognitive niche and became hunter-gatherers, there would have
been novel and intense selection for adaptations designed to
make the self valuable to others, and hence recruit assistance
from others. We hypothesize that the emotion of pride functions
as an evolved guidance system that modulates behavior to cost-
effectively manage and capitalize on the propensities of others to
both respect and value the actor.
Mechanisms favoring the valuation of others evolved through
several distinct selection pressures, including kin selection (20),
reciprocation (21), reputation (22), risk pooling (1), externality
management (23), and (substituting respect for valuation) the
asymmetric war of attrition (24). These selection pressures, in
turn, crafted an array of specialized choice architectures to
promote altruistic (or selfish) decisions given the information
available to the actor about a potential recipient [e.g., how to
respond to cues of the recipient’s relatedness, skills, trustwor-
thiness, or ability to defend her interests (25)]. This implies that
humans will have evolved a neurocognitive architecture for
computing the social value of others, which governs altruistic
behavior (26). We note that formidability—the ability to inflict
costs through aggression—commonly incentivizes others (in
bargaining contexts) to place more weight on the welfare of the
more formidable, even when such aggressive capacity is not
deployed in ways that help others. Hence, both the ability to
confer benefits (e.g., skills, the emission of positive externalities)
and the ability to inflict costs should act as inputs to the systems
that compute the social value of others (7, 11, 15).
In short, others’assessments of the acts and characteristics of a
focal individual lead them to value (or disvalue) her. When
others (an audience) detect new information about an individual
that is at odds with their current level of valuation, their valua-
tion is recalibrated either upward or downward, with corre-
spondingly positive or negative effects on the individual’s fitness
(26). This would have selected on the recipient’s end for moti-
vational adaptations to cost-effectively manage the flow of in-
formation about the self to others (27). Indeed, cross-cultural
evidence has recently provided support for the hypothesis that the
emotion of shame is a neurocognitive adaptation that evolved to
prevent audiences from receiving negative information about the
individual and to limit the degree and costs of devaluation [e.g., by
signaling submission to avoid aggression from audiences (10, 12)]
if negative information does spread (28–32).
Reciprocally, the neurocomputational system that organizes
the emotion of pride seems to be an adaptation that evolved to
pursue and advertise acts or traits leading to enhanced respect
and valuation of the individual in the minds of others. A system
designed for this function should orchestrate a suite of cognitive
mechanisms that (i) motivate the pursuit of acts or the cultivation of
traits that would increase others’respect and valuations of the indi-
vidual; (ii) motivate the advertisement of acts or characteristics that,
when discovered by others, would lead them to increase their respect
and valuations of the individual; and (iii) mobilize the individual to
profit from the resulting enhanced social landscape (e.g., by pursuing
gainful activities previously beyond reach or pressing for better
treatment from others). This “advertisement–recalibration theory of
pride”(33; see also refs. 10, 12, 34) deductively emerges from the
integration of the dynamics of audience recalibration with evolu-
tionary models of human dominance and valuation, which specify the
direction and magnitude of those recalibrations.
Existing findings on pride are strongly consistent with this
theory of its functional architecture. Pride-like behavior is tax-
onomically widespread [including primates (35, 36), cervids (37,
38), canids (39), and invertebrates (40)], and therefore phylo-
genetically ancient. Pride occurs in every known culture (41) and
it appears reliably and early in development—as early as in
toddlers (42, 43). Pride is triggered by achievements (42), ag-
gressive formidability (44), and other socially valued character-
istics. Pride is a highly pleasant emotion (45); this internal reward
can incentivize people to undertake and persevere at costly but
socially valued courses of action (46, 47). Pride has a full-body
display featuring an erect and expanded posture and gaze di-
rected at the audience (12, 42, 48), and thus it appears to generate
common knowledge about the individual’s enhanced value (49).
This display conveys achievement or dominance (10, 12, 50, 51), is
produced by congenitally blind individuals (45), and is recognized by
young children (52) and by adults within and across cultures (53).
Thus, pride and related indicators of being respected and valued
affect second and third parties in lawful fashion: They appeal to
potential mates (54, 55) (presumably because they indicate good
genes, health, resource holding potential, and other types of em-
bodied, social, and material capital); guide social learning through
imitation (56, 57); elicit submissiveness (58); and intimidate rivals
(10, 59), which reduces agonistic interactions (24) and stabilizes
dominance hierarchies (60).
We note that human pride and its obverse, shame, are evo-
lutionarily derived from physiological and behavioral features
undergirding dominance and submission (10, 12, 17, 61, 62)—as
articulated by the Dominance Hierarchy Model of pride and
shame (10)—and various aspects of those emotions (e.g., the
displays) are homologous with those of nonhuman primates (10).
For example, receiving a pride display may elicit submission,
while receiving a shame display may terminate aggression. Thus,
these two complementary systems reduce overt conflict and
subsequent attacks (refs. 10 and 63; nonhuman primate examples
are in refs. 37, 64, and 65). Pride provides an internal reward for
competitive success, whereas shame punishes failure; since much
animal competition, including human competition, is ultimately
over reproductive opportunities (40, 66–68), this may account for
the heightened hubristic pride and, to a lesser extent, shame
observed during adolescence and early adulthood (69).
The decision-making architecture of a social organism should
evaluate and integrate two kinds of payoffs to regulate behavior
adaptively: (i) the direct payoff of the potential action (e.g., the
value of foraging for a food item) and (ii) the social valuation payoff
[e.g., showing bandmates that one is a skilled forager by pursuing a
hard-to-acquire prey item (70)]. According to the advertisement–
recalibration theory, the feeling of pride is an internal signal of the
estimated social valuation payoff—a payoff that can motivate, for
example, status-seeking behavior.
One central prediction of the theory then is that the intensity
of the feeling of pride will track the magnitude of audience
evaluations incrementally and closely for each kind of informa-
tion. This calibration is necessary if the intensity of the internal
signal (anticipatory pride) is used prospectively to compute
whether the benefit of enhanced audience valuation or respect
outweighs the cost of engaging in a given act—and to decide
whether the likely net payoff of a candidate act will make that
act worth pursuing. An internal pride signal that is too weak
www.pnas.org/cgi/doi/10.1073/pnas.1808418115 Sznycer et al.
compared with the prevalent magnitude of audience valuation
would lead to maladaptive choices where the relevant act is in-
sufficiently pursued (or if achieved, underadvertised), the in-
crease in valuation in the audience is less than what it would be
under more complete knowledge resulting from fuller adver-
tisement, and the individual foregoes valuation that would have
been cost-effective to acquire. A pride signal that is too strong
yields diminishing or even negative returns, as beneficial courses
of action are pursued in excess of their actual return, and
moreover, audiences are designed to resist and devalue entitled
actions that exceed the individual’s actual social value (71, 72).
To avoid these errors, the pride system should estimate the
magnitude of valuation that a given act would cause among local
audiences and calibrate the intensity of its internal signal in
proportion to those estimates. This internal signal is expected to
be equally well-calibrated for traits (e.g., physical formidability)
and other attributes (e.g., sibling of chief) for the individual to
know the right degree of advertisement and entitlement afforded
by those attributes. Pride is sometimes referred to as a self-
conscious (73) or self-focused (74) emotion; however, the pre-
ceding analysis suggests that a well-designed pride system must
be coupled to the evaluative psychology of others. Importantly,
because the internal pride signal is used by the systems that
decide how to act, the intensity of felt pride should track the
magnitude of audience valuation even when there is no com-
munication between audiences and the individual who is evalu-
ating alternative courses of action based on anticipated pride.
The internal pride signal is useful for promoting audience valu-
ation and respect by choosing certain acts, displays, and modes of
conduct over others. The system generating this signal would be
handicapped if it needed to observe audience valuation to know
its magnitude instead of computing those magnitudes in advance.
These predictions were tested experimentally in 16 countries: the
United States, Canada, the United Kingdom, France, Belgium,
The Netherlands, Switzerland, Italy, Turkey, Israel, India, Singapore,
the Philippines, South Korea, Japan, and Australia (33). Subjects
were given a set of scenarios that tapped situations likely to vary in
how much valuation the actions or traits that they described might
elicit. One group of subjects rated how positively they would eval-
uate the person described in each scenario. A second independent
group of subjects rated how much pride they would feel if they were
the person described in the situation. As predicted, the intensity of
anticipated pride for a given act or trait closely tracked the corre-
sponding magnitude of audience valuation. This result replicated
Fishing communities, Japan
Farming communities, Japan
Shaanxi and Henan, China
Le Morne, Mauritius
La Gaulette, Mauritius
Bosawás Reserve, Nicaragua
Fig. 1. Map of the 10 field sites.
Table 1. Demographic information (samples A–J)
Community Economy Religion NNfemale Age, y (SD)
Bosawás Reserve, Nicaragua Foraging, horticulture Syncretic Catholicism 46 23 40 (12)
Cotopaxi, Ecuador Subsistence agriculture, pastoralism Evangelism 34 25 41 (18)
Drâa-Tafilalet, Morocco Subsistence agriculture Sunni Islam 75 43 32 (13)
Enugu, Nigeria Subsistence agriculture Catholicism 80 39 34 (8)
Le Morne, Mauritius Fishing, farming, wage labor Catholicism 80 33 40 (13)
La Gaulette, Mauritius Fishing, farming, service sector Hinduism 80 53 35 (12)
Tuva, Russia Seminomadic pastoralism Shamanism, Buddhism 29 22 36 (13)
Shaanxi and Henan, China Farming Mostly nonreligious 41 17 41 (12)
Farming communities, Japan* Farming, wage labor Buddhism, Shintoism 50 23 68 (11)
Fishing communities, Japan
Fishing, farming, wage labor Buddhism, Shintoism 52 18 66 (12)
Means (SDs in parentheses).
*Participants sampled from 13 communities (in three prefectures) where at least 25% of the residents are farmers.
Participants sampled from 13 communities (in three prefectures) where at least 25% of the residents are fishers.
Sznycer et al. PNAS Latest Articles
in each of the 16 countries. Importantly, valuation was tracked
specifically by pride. Excitement, amusement, and happiness—
three other positively valenced and arousing emotions and states
that coactivate with pride—failed to track audience valuation.
Although this 16-nation experiment is suggestive, those pop-
ulations are all western(-ized), educated, industrialized, rich, and
democratic mass societies (75) and importantly, are in close media
contact, sharing many norms, values, and attitudes. Hence, the goal
of these studies is twofold. (i) The claim being evaluated is that the
pride system is a fundamental part of human biology, and therefore,
the signature of its operation should be detectable in all human
societies, no matter how widely distributed and mutually unfamiliar
they are. (ii) By hypothesis, the pride system evolved in small-scale
face-to-face social groups wherepeoplekneweachother,and
therefore, it is important to assess the evidence for its operation in
small coresidential social ecologies.
Is the tracking of audience valuation by pride limited to in-
dustrial mass societies? Or does this tracking occur throughout
the range of human societies, potentially reflecting the operation
of a panhuman pride system? To answer this question, we con-
ducted an experiment with 567 adult participants from 10 small-
scale communities living in widely different physical ecologies
and featuring very different languages, cultures, and modes of
subsistence: (i) Bosawás Reserve, Nicaragua; (ii ) Cotopaxi, Ecua-
dor; (iii) Drâa-Tafilalet, Morocco; (iv) Enugu, Nigeria; (v)Le
Morne, Mauritius; (vi) La Gaulette, Mauritius; (vii)Tuva,Russia;
(viii) Shaanxi and Henan, China; (ix) farming communities in Japan;
and (x) fishing communities in Japan (Fig. 1 and Table 1). We
created 10 scenarios in which someone’s acts, traits, or circum-
stances might lead her to be viewed positively. The scenarios were
designed to elicit reactions in a variety of evolutionarily relevant
domains, such as generosity, social exchange, dominance contests,
skills, and health. They were expressed at a level of abstraction that
was not culturally particular (e.g., “You have many skills”rather
than “You know how to bake and how to pilot airplanes”).
The experimental design was adapted from Sznycer et al. (33).
Participants were randomly assigned to either an audience
condition or a pride condition. Participants in the audience con-
dition were asked to provide their reactions to 10 scenarios in-
volving a third party: a same-sex individual other than themselves
(e.g., “He has many skills,”“He is generous with others,”“He can
defend himself, so people never push him around”). These par-
ticipants were asked, for each scenario, to “indicate how you
would view this person if this person was in those situations”; they
indicated their reactions using scales ranging from one (I would
not view them positively at all) to four (I would view them very
positively). These ratings provide situation-specific measures of
the degree to which members of a given population would posi-
tively evaluate the individual described in the scenarios.
In the pride condition, a different set of participants was asked to
“indicate how much pride you would feel if you were in those sit-
uations”(i.e., in each of the 10 scenarios; e.g., “You have many
skills,”“You are generous with others,”“You can defend yourself,
so people never push you around”), with scales ranging from one
(no pride at all) to four (a lot of pride; the exceptions being Bosawás
Reserve, Nicaragua and Drâa-Tafilalet, Morocco, where valuation
and pride were measured on 1–3and1–7 scales, respectively). The
stimuli in the audience condition and the pride condition were
identical on a scenario-by-scenario basis, the only difference being
the perspective from which the events are described.
Within-Community Results. First, we report the valuation and pride
results for each community (SI Appendix,SI Text and Tables S1–
S2j). There was widespread agreement on how valuation-enhancing
these situations are relative to one another: mean intraclass cor-
relation (ICC) across the 10 communities: ICC (2,n)=0.70 (SI
Appendix,TableS3). In other words, participants agreed about
the extent to which they would positively view the individual de-
scribed in these scenarios. Participants also agreed about the relative
degree to which these various situations would elicit pride:
mean ICC (2,n)=0.61 (SI Appendix,TableS3). To test the main
prediction that pride tracks audience valuation, we calculated, for
each scenario, the mean pride ratings provided by participants in
the pride condition and the mean valuation ratings provided by
participants in the audience condition. Pride and valuation means
were highly correlated with one another within each community,
with a mean r=0.66 (SD =0.24; minimum r=0.36; maximum r=
0.92; Nrvalues =10) and Pvalues =0.0002–0.31 (Fig. 2 and Table
2, diagonal values). Recall that the pride ratings and the valuation
ratings originate from different participants. Consequently, these
high correlations cannot be attributed to participants matching
their pride ratings and valuation ratings. This is consistent with the
Between-Community Results. The pride system evolved for making
decisions in—and tracking the values of—one’s local group and
not people from other cultures. Obviously, there would have
been no selection to map the valuations of persons with whom
one has never interacted. However, if there is a human-universal
system of social valuation, then scenarios that tap this system
: .78 r
: .41 r
: .85 r
: .62 r
: .17 r
Fig. 2. Scatterplots: pride as a function of valuation (samples A–J). Each point
represents the mean pride rating and mean valuation rating of one scenario.
Pride ratings and valuation ratings were given by different participants. Non
which the correlations are based is the number of scenarios =10. Effect size:
linear. (A) Bosawás Reserve, Nicaragua; (B) Cotopaxi, Ecuador; (C)Drâa-
Tafilalet, Morocco; (D) Enugu, Nigeria; (E) Le Morne, Mauritius; (F) La Gaulette,
Mauritius; (G) Tuva, Russia; (H) Shaanxi and Henan, China; (I) farming commu-
nities in Japan; and (J) fishing communities in Japan.
www.pnas.org/cgi/doi/10.1073/pnas.1808418115 Sznycer et al.
may elicit agreement across cultures about what is worthy of
valuation and pride, and pride in a given culture may track val-
uation in other cultures, despite a lack of contact between them.
Are there situations that provoke valuation and elicit pride
across cultures? To test for between-community agreement in
valuation, in pride, and in the pride–valuation link, we com-
puted the extent to which the mean valuation ratings and the
mean pride ratings are correlated across communities. There is
between-community agreement on average on the extent to
which a given situation would elicit valuation: mean r=0.37
(SD =0.30; minimum r=−0.26; maximum r=0.91; Nrvalues =
45) and Pvalues =0.0002–0.96 (SI Appendix, Table S4). There is
also between-community agreement on the extent to which a
given situation would elicit pride: mean r=0.20 (SD =0.38;
minimum r=−0.80; maximum r=0.92; Nrvalues =45) and P
values =0.0002–0.99 (SI Appendix, Table S5). Furthermore, the
pride elicited in each of 10 communities is positively correlated
on average with the valuation from the other 9 communities:
mean r=0.29 (SD =0.34; minimum r=−0.62; maximum r=
0.87; Nrvalues =90) and Pvalues =0.001–0.98 (Table 2, off-
diagonal values)—71 of these 90 correlations (79% of them)
have a positive sign. Although there is substantial variation in the
extent to which pride tracked valuation across communities, in-
cluding null and negative correlations, the pride elicited by these
scenarios in one community (e.g., Mayangna forager–horticulturalists
of the Bosawás Reserve, Nicaragua) tended to track how positively
people viewed these scenarios in the other communities (e.g.,
pastoralists from Tuva, Russia; Amazigh farmers from Drâa-
Tafilalet, Morocco; and farmers from Enugu, Nigeria). Of course,
some actions, traits, and situations elicit valuation and pride in
some cultures but not others (33, 70).
A cross-culturally replicable, close quantitative correspondence
between anticipated pride and the valuation of local audiences is
what one expects of a computational system that is well-designed
for furthering the social value of the individual in the minds of
others. Features causing this close calibration assist the individ-
ual in balancing the competing demands of effectiveness and
restraint by steering between an internal pride signal that is too
strong (which would lead to, for example, the overpursuit of
socially valued acts) and one that is too weak (which would, for
example, insufficiently motivate acts that are socially valued).
This match is not limited to industrial mass societies but gener-
alizes across populations with widely different cultures, subsistence
modes, institutions, and languages. Thus, this feature is more likely
to originate in a human-universal adaptation designed by natu-
ral selection than in cultural evolutionary processes (76). The
agreement across cultures, and not just within them, on pride,
valuation, and their interrelationship is noteworthy. According to
some accounts, different cultures are richly and arbitrarily different
from each other (77). If this were true, then what cultures value
and what members of different cultures are proud about should be
radically different. Cultural differences in pride and in the un-
derlying items granted respect and valuation do exist, as shown
here and elsewhere (45, 78, 79). However, these cultural differ-
ences can be adaptively patterned (33), and therefore cultural
variation is not necessarily divorced from the logic of adaptive
functionality. Moreover, regularities across vastly disparate cultures
can emerge when pride is analyzed from the standpoint of its
probable function and target domain. These data contribute to
a growing body of findings indicating that theories of adaptive
function are a powerful tool for identifying regularities in the
structure and content of human emotion.
The study procedures were approved by the institutional review boards at the
University of California, Santa Barbara; East China Normal University; the
University of Nigeria, Nsukka; Universidad San Francisco de Quito, and
the University of Cincinnati, and the research ethics committee of the In-
stitute of Psychology, Russian Academy of Sciences. All of the participants
gave informed consent. The data and study materials are included in Dataset
S1 and SI Appendix, respectively.
Participants. We collected data from 567 participants in Bosawás Reserve,
Nicaragua (sample A); Cotopaxi, Ecuador (sample B); Drâa-Tafilalet, Morocco
(sample C); Enugu, Nigeria (sample D); Le Morne, Mauritius (sample E); La
Gaulette, Mauritius (sample F); Tuva, Russia (sample G); Shaanxi and Henan,
China (sample H); farming communities in Japan (sample I); and fishing
communities in Japan (sample J). Sample sizes and demographic information
are described in Table 1.
Procedure. The 10 scenarios are shown in SI Appendix, Tables S2a–S2j. Partici-
pants were randomly assigned to either the audience condition or the pride
condition. The language in the scenarios was gendered according to participants’
stated gender, except for at the two Japan sites. At both Japan sites, data col-
lection was through self-administered questionnaires sent by mail; here, we used
gender-neutral pronouns and instructed respondents in the audience condition
to imagine the target individual was someone of their same sex and age. Sample
size, order in which the scenarios were administered, method of stimuli ad-
ministration, and language of stimuli are listed in SI Appendix,TableS1.
ACKNOWLEDGMENTS. We thank two anonymous reviewers. This research
was supported by funding from Federal Agency for Scientific Organizations,
Russian Federation Grant 0159-2016-0001 (to K.I.A. and A.N.K.), Japan Soci-
ety for the Promotion of Science KAKENHI Grant 26780343 (to K.T.), and
John Templeton Foundation (JTF) Grant 29468 (to L.C. and J.T.). The opinions
expressed in this publication are those of the authors and do not necessarily
reflect the views of the JTF or the other funding agencies.
Table 2. Correlations between pride and valuation within and between communities (samples A–J)
ABCDE F GHI J
(A) Bosawás Reserve, Nicaragua 0.88* 0.07 0.77* 0.60 0.56 0.57 0.87* 0.46 0.16 0.33
(B) Cotopaxi, Ecuador 0.62 0.64* 0.16 −0.01 0.06 0.05 0.24 0.35 0.77* 0.86*
(C) Drâa-Tafilalet, Morocco 0.77* 0.03 0.87* 0.77* 0.63 0.59 0.80* 0.22 0.32 0.31
(D) Enugu, Nigeria 0.11 −0.35 0.76* 0.92* 0.39 0.33 0.68* −0.16 −0.04 −0.09
(E) Le Morne, Mauritius 0.60 0.11 0.56 0.37 0.79* 0.72* 0.65* 0.47 0.33 0.48
(F) La Gaulette, Mauritius 0.37 0.26 0.43 0.52 0.77* 0.87* 0.64* 0.51 −0.09 0.10
(G) Tuva, Russia 0.51 0.04 0.39 0.39 −0.05 0.45 0.36 0.32 −0.05 0.12
(H) Shaanxi and Henan, China 0.20 0.05 0.39 0.24 0.80* 0.58 0.53 0.42 0.03 0.23
(I) Farming communities, Japan 0.10 0.43 −0.42 −0.62 −0.06 −0.19 −0.29 0.48 0.36 0.36
(J) Fishing communities, Japan −0.02 0.44 −0.41 −0.52 −0.10 −0.20 −0.30 0.46 0.51 0.48
Coefficients are Pearson’srvalues. Non which the correlations are based is the number of scenarios =10. Pride ratings and
valuation ratings were given by different participants. Grey cells, within-community correlations.
*Correlations meet P<0.05 or less.
Sznycer et al. PNAS Latest Articles
1. Kaplan H, Hill K (1985) Food sharing among ache foragers: Tests of explanatory hy-
potheses. Curr Anthropol 26:223–239.
2. Sugiyama LS (2004) Illness, injury, and disability among Shiwiar forager-
horticulturalists: Implications of health-risk buffering for the evolution of human
life history. Am J Phys Anthropol 123:371–389.
3. Keeley LH (1997) War Before Civilization (Oxford Univ Press, New York).
4. Treves A, Naughton-Treves L (1999) Risk and opportunity for humans coexisting with
large carnivores. J Hum Evol 36:275–282.
5. Clutton-Brock T (2009) Cooperation between non-kin in animal societies. Nature 462:
6. von Rueden CR, Jaeggi AV (2016) Men’s status and reproductive success in 33 non-
industrial societies: Effects of subsistence, marriage system, and reproductive strat-
egy. Proc Natl Acad Sci USA 113:10824–10829.
7. Henrich J, Gil-White FJ (2001) The evolution of prestige: Freely conferred deference as
a mechanism for enhancing the benefits of cultural transmission. Evol Hum Behav 22:
8. Sell A, et al. (2009) Human adaptations for the visual assessment of strength and
fighting ability from the body and face. Proc Biol Sci 276:575–584.
9. Sell A, et al. (2010) Adaptations in humans for assessing physical strength from the
voice. Proc Biol Sci 277:3509–3518.
10. Weisfeld GE, Dillon LM (2012) Applying the dominance hierarchy model to pride and
shame, and related behaviors. J Evol Psychol 10:15–41.
11. Sell A, Tooby J, Cosmides L (2009) Formidability and the logic of human anger. Proc
Natl Acad Sci USA 106:15073–15078.
12. Fessler DMT (1999) Toward an Understanding of the Universality of Second Order
Emotions, ed Hinton AL (Cambridge Univ Press, New York), pp 75–116.
13. Hawley PH (2007) Social dominance in childhood and adolescence: Why social com-
petence and aggression may go hand in hand. Aggression and Adaptation: The Bright
Side to Bad Behavior, eds Hawley PH, Little TD, Rodkin P (Erlbaum, Hillsdale, NJ),
14. De Bruyn EH, Cillessen AH (2006) Popularity in early adolescence: Prosocial and an-
tisocial subtypes. J Adolesc Res 21:607–627.
15. Cheng JT, Tracy JL, Foulsham T, Kingstone A, Henrich J (2013) Two ways to the top:
Evidence that dominance and prestige are distinct yet viable avenues to social rank
and influence. J Pers Soc Psychol 104:103–125.
16. Hold BC (1980) Attention-structure and behavior in G/wi San children. Evol Hum
17. Gilbert P (1997) The evolution of social attractiveness and its role in shame, humili-
ation, guilt and therapy. Br J Med Psychol 70:113–147.
18. Tooby J, DeVore I (1987) The Reconstruction of Hominid Behavioral Evolution
Through Strategic Modeling, ed Kinzey WG (State Univ New York Press, Albany,
NY), pp 187–237.
19. Pinker S (2010) Colloquium paper: The cognitive niche: Coevolution of intelligence,
sociality, and language. Proc Natl Acad Sci USA 107:8993–8999.
20. Hamilton WD (1964) The genetical evolution of social behaviour. I. J Theor Biol 7:
21. Trivers R (1971) The evolution of reciprocal altruism. Q Rev Biol 46:35–57.
22. Nowak MA, Sigmund K (1998) Evolution of indirect reciprocity by image scoring.
23. Tooby J, Cosmides L (1996) Friendship and the Banker’s Paradox: Other pathways to
the evolution of adaptations for altruism. Evolution of Social Behaviour Patterns in
Primates and Man, Proceedings of the British Academy, eds Runciman WG, Maynard
Smith J, Dunbar RIM (The British Academy, London), pp 119–143.
24. Hammerstein P, Parker GA (1982) The asymmetric war of attrition. J Theor Biol 96:
25. Lieberman D, Tooby J, Cosmides L (2007) The architecture of human kin detection.
26. Tooby J, Cosmides L, Sell A, Lieberman D, Sznycer D (2008) Internal Regulatory
Variables and the Design of Human Motivation: A Computational and Evolutionary
Approach, ed Elliot AJ (Lawrence Erlbaum Associates, Mahwah, NJ), pp 251–271.
27. Leary MR, Kowalski RM (1990) Impression management: A literature review and two-
component model. Psychol Bull 107:34–47.
28. Sznycer D, et al. (2016) Shame closely tracks the threat of devaluation by others, even
across cultures. Proc Natl Acad Sci USA 113:2625–2630.
29. Sznycer D (2010) Cognitive adaptations for calibrating welfare tradeoff motivations,
with special reference to the emotion of shame. Doctoral dissertation (University of
California, Santa Barbara, CA).
30. Sznycer D, Schniter E, Tooby J, Cosmides L (2015) Regulatory adaptations for de-
livering information: The case of confession. Evol Hum Behav 36:44–51.
31. Sznycer D, et al. (2012) Cross-cultural differences and similarities in proneness to
shame: An adaptationist and ecological approach. Evol Psychol 10:352–370.
32. Robertson TE, Sznycer D, Delton AW, Tooby J, Cosmides L (2018) The true trigger of
shame: Social devaluation is sufficient, wrongdoing is unnecessary. Evol Hum Behav
33. Sznycer D, et al. (2017) Cross-cultural regularities in the cognitive architecture of
pride. Proc Natl Acad Sci USA 114:1874–1879.
34. Tracy JL, Shariff AF, Cheng JT (2010) A naturalist’s view of pride. Emot Rev 2:163–177.
35. Weisfeld GE, Beresford JM (1982) Erectness of posture as an indicator of dominance
or success in humans. Motiv Emot 6:113–131.
36. Smuts BB, Cheney DL, Seyfarth RM, Wrangham RW (2008) Primate Societies (Univ
Chicago Press, Chicago).
37. Barrette C (1977) Fighting behavior of muntjac and the evolution of antlers. Evolution
38. Clutton-Brock TH, Albon SD (1979) The roaring of red deer and the evolution of
honest advertisement. Behaviour 69:145–170.
39. Fox MW (1969) The anatomy of aggression and its ritualization in Canidae: A de-
velopmental and comparative study. Behaviour 35:242–258.
40. Huntingford F, Turner A (1987) Animal Conflict (Chapman and Hall, London).
41. Brown DE (1991) Human Universals (Temple Univ Press, Philadelphia).
42. Lewis M, Alessandri SM, Sullivan MW (1992) Differences in shame and pride as a
function of children’s gender and task difficulty. Child Dev 63:630–638.
43. Stipek D (1995) The development of pride and shame in toddlers. Self-Conscious
Emotions: The Psychology of Shame, Guilt, Embarrassment, and Pride, eds Tangney JP,
Fischer KW (Guilford, New York), pp 237–252.
44. Tracy JL, Matsumoto D (2008) The spontaneous expression of pride and shame: Evi-
dence for biologically innate nonverbal displays. Proc Natl Acad Sci USA 105:
11655–11660, and erratum (2008) 105:20044.
45. Mauro R, Sato K, Tucker J (1992) The role of appraisal in human emotions: A cross-
cultural study. J Pers Soc Psychol 62:301–317.
46. Williams LA, DeSteno D (2008) Pride and perseverance: The motivational role of pride.
J Pers Soc Psychol 94:1007–1017.
47. Riskind JH (1984) They stoop to conquer: Guiding and self-regulatory functions of
physical posture after success and failure. J Pers Soc Psychol 47:479–493.
48. Tracy JL, Robins RW (2007) The prototypical pride expression: Development of a
nonverbal behavior coding system. Emotion 7:789–801.
49. Thomas KA, DeScioli P, Pinker S (2018) Common knowledge, coordination, and the
logic of self-conscious emotions. Evol Hum Behav 39:179–190.
50. Shariff AF, Tracy JL (2009) Knowing who’s boss: Implicit perceptions of status from the
nonverbal expression of pride. Emotion 9:631–639.
51. Tiedens LZ, Ellsworth PC, Mesquita B (2000) Sentimental stereotypes: Emotional ex-
pectations for high-and low-status group members. Pers Soc Psychol Bull 26:560–575.
52. Tracy JL, Robins RW, Lagattuta KH (2005) Can children recognize pride? Emotion 5:
53. Tracy JL, Robins RW (2008) The nonverbal expression of pride: Evidence for cross-
cultural recognition. J Pers Soc Psychol 94:516–530.
54. Sadalla EK, Kenrick DT, Vershure B (1987) Dominance and heterosexual attraction.
J Pers Soc Psychol 52:730–738.
55. de Bruyn EH, Cillessen AH, Weisfeld GE (2012) Dominance-popularity status, behavior,
and the emergence of sexual activity in young adolescents. Evol Psychol 10:296–319.
56. Chudek M, Heller S, Birch S, Henrich J (2012) Prestige-biased cultural learning: By-
stander’s differential attention to potential models influences children’s learning.
Evol Hum Behav 33:46–56.
57. Martens JP, Tracy JL (2013) The emotional origins of a social learning bias: Does the
pride expression cue copying? Soc Psychol Personal Sci 4:492–499.
58. Tiedens LZ, Fragale AR (2003) Power moves: Complementarity in dominant and
submissive nonverbal behavior. J Pers Soc Psychol 84:558–568.
59. Mazur A (1985) A biosocial model of status in face-to-face primate groups. Soc Forces
60. Savin-Williams RC (1977) Dominance in a human adolescent group. Anim Behav 25:
61. Darwin C (1872) The Expression of the Emotions in Man and Animals (John Murray,
62. Barkow JH, et al. (1975) Prestige and culture: A biosocial interpretation. Curr
63. Omark DR, Strayer FF, Freedman DG (1980) Dominance Relations: An Ethological
View of Human Conflict and Social Interaction (Garland, New York).
64. Altmann SA (1962) A field study of the sociobiology of rhesus monkeys, Macaca
mulatta. Ann N Y Acad Sci 102:338–435.
65. Deag JM (1977) Aggression and submission in monkey societies. Anim Behav 25:
66. Trivers RL (1972) Parental Investment and Sexual Selection, ed Campbell B (Aldine-
Atherton, Chicago), pp 136–179.
67. Wilson M, Daly M (1985) Competitiveness, risk taking, and violence: The young male
syndrome. Ethol Sociobiol 6:59–73.
68. Daly M, Wilson M (1988) Homicide (Aldine de Gruyter, New York).
69. Orth U, Robins RW, Soto CJ (2010) Tracking the trajectory of shame, guilt, and pride
across the life span. J Pers Soc Psychol 99:1061–1071.
70. Blurton Jones N, Hawkes K, O’Connell JF (1997) Why do Hadza children forage?
Uniting Psychology and Biology: Integrative Perspectives on Human Development,
eds Segal NL, Weisfeld GE, Weisfield CC (American Psychological Association, New
York), pp 297–331.
71. Anderson C, Ames DR, Gosling SD (2008) Punishing hubris: The perils of over-
estimating one’s status in a group. Pers Soc Psychol Bull 34:90–101.
72. Schlenker BR, Leary MR (1982) Audiences’reactions to self-enhancing, self-
denigrating, and accurate self-presentations. J Exp Soc Psychol 18:89–104.
73. Tracy JL, Robins RW (2004) Putting the self into self-conscious emotions: A theoretical
model. Psychol Inq 15:103–125.
74. Simon-Thomas ER, et al. (2012) An fMRI study of caring vs self-focus during induced
compassion and pride. Soc Cogn Affect Neurosci 7:635–648.
75. Henrich J, Heine SJ, Norenzayan A (2010) The weirdest people in the world? Behav
Brain Sci 33:61–83.
76. Boyd R, Richerson PJ (1988) Culture and the Evolutionary Process (Univ Chicago Press,
77. Geertz C (1973) The Interpretation of Cultures: Selected Essays (Basic Books, New
78. von Fürer-Haimendorf C (1967) Morals and Merit: A Study of Values and Social
Controls in South Asian Societies (Univ of Chicago Press, Chicago).
79. Dong Q, Weisfeld G, Boardway RH, Shen J (1996) Correlates of social status among
Chinese adolescents. J Cross Cult Psychol 27:476–493.
www.pnas.org/cgi/doi/10.1073/pnas.1808418115 Sznycer et al.