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Abstract

Current theoretical and empirical findings suggest that mate preferences are mainly cued on visual, vocal and chemical cues that reveal health including developmental health. Beautiful and irresistible features have evolved numerous times in plants and animals due to sexual selection, and such preferences and beauty standards provide evidence for the claim that human beauty and obsession with bodily beauty are mirrored in analogous traits and tendencies throughout the plant and animal kingdoms. Human beauty standards reflect our evolutionary distant and recent past and emphasize the role of health assessment in mate choice as reflected by analyses of the attractiveness of visual characters of the face and the body, but also of vocal and olfactory signals. Although beauty standards may vary between cultures and between times, we show in this review that the underlying selection pressures, which shaped the standards, are the same. Moreover we show that it is not the content of the standards that show evidence of convergence – it is the rules or how we construct beauty ideals that have universalities across cultures. These findings have implications for medical, social and biological sciences.
Darwinian aesthetics: sexual selection and the
biology of beauty
KARL GRAMMER
1
,BERNHARDFINK
1
,
*, ANDERS P. MØLLER
2
and RANDY THORNHILL
3
1
Ludwig-Boltzmann-Institute for Urban Ethology, Althanstrasse 14, A-1090 Vienna, Austria.
2
Universite
´
Pierre et Marie Curie, Laboratoire de Parasitologie Evolutive, CNRS UMR 7103, Ba
ˆ
timent A,7e
`
me e
´
tage,
7 quai St. Bernard, Case 237, FR-75252 Paris Cedex 05, France.
3
University of New Mexico, Castetter Hall, Department of Biology, Albuquerque, NM 87131-1091, USA.
(Received 30 January 2002; revised 4 September 2002 ; accepted 13 September 2002 )
ABSTRACT
Current theoretical and empirical findings suggest that mate preferences are mainly cued on visual, vocal and
chemical cues that reveal health including developmental health. Beautiful and irresistible features have evolved
numerous times in plants and animals due to sexual selection, and such preferences and beauty standards provide
evidence for the claim that human beauty and obsession with bodily beauty are mirrored in analogous traits and
tendencies throughout the plant and animal kingdoms. Human beauty standards reflect our evolutionary distant
and recent past and em phasize the role of health assessment in mate choice as reflected by analyses of the
attractiveness of visua l characters of the face and the body, but also of vocal and olfactory signals. Although beauty
standards may vary between cultures and between times, we show in this review that the underlying selec tion
pressures, which shaped the standards, are the same. Moreover we show that it is not the content of the standards
that show evidence of convergence it is the rules or how we construct beauty ideals that have universalities across
cultures. These findings have implications for medical, social and biological sciences.
Key words : attractiveness, beauty standards, Darwinian aesthetics, face, humans, mate choice, sexual select ion.
CONTENTS
I. Introduction ....................... ......... ......... ...... .......... ......... ......... ...... ......... .......... ......... ...... ......... ......... .......... . 386
II. Sexual selection and mate choice .............................. ......... ......... ...... .......... ......... ......... ...... ......... .......... . 386
III. Sexual selection and why beauty matte rs ....................... ......... ......... .......... ...... ......... ......... ......... ....... .... 387
IV. Human beauty and sexual selection ....................... ......... ......... ...... .......... ......... ......... ...... ......... .......... .... 387
V. Attractiveness and daily life .............................................. ......... ...... .......... ......... ......... ...... ......... .......... .... 388
VI. Health and beauty perception in humans and other animals ....................... ......... ......... ...... .......... .... 389
VII. Attractiveness and physical features ....................... ......... ......... ...... .......... ......... ......... ...... ......... .......... .... 389
(1 ) Theories of featur e processing : pro ........... ......... ...... ......... ......... .......... ...... ......... ......... ......... ....... .... 390
(2 ) Theories of featur e processing : contra ...... ......... ...... ......... ......... .......... ...... ......... ......... ......... .......... . 391
VIII. The attractive prototype: faces ........................................... ......... ......... ....... ......... ......... ......... ...... .......... . 392
IX. The attr active prototype: bodies ................................ ......... ......... ...... .......... ......... ......... ...... ......... .......... . 393
(1 ) Theories of prototype processing : pro ................ ......... ......... ......... .......... ...... ......... ......... .......... ...... . 393
(2 ) Theories of prototype processing : contra ........................................................ ...... ......... .......... ....... 394
X. Developmental stability and beauty ................................ ......... ...... .......... ......... ......... ...... ......... .......... .... 394
(1 ) Theories of symmetry and attractiveness: pro ................................................ ......... ......... .......... .... 395
(2 ) Theories of symmetry and attractiveness: contra .............................................. ......... ......... .......... . 396
* Address for correspondence: Ludwig-Boltzmann-Institute for Urban Ethology, University of Vienna, Althanstra sse 14,
A-1090 Vienna, Austria. Tel: +43 1 4277 54769. Fax : +43 1 4277 9547. E-mail : bernhard.fink@ieee.org
Biol. Rev. (2003), 78, pp. 385–407. f Cambridge Philosophical Society 385
DOI: 10.1017/S1464793102006085 Printed in the United Kingdom
XI. Cross-sensory modalities: body odour, voices, decoration and movement ....................................... 397
XII. The beauty of boundaries and boundaries of beauty ................................. ......... ...... ......... ......... ......... 399
XIII. The future of the adapted mind ......................................... ......... ......... .......... ...... ......... ......... ......... ....... .. 401
XIV. Directions for future research ....................................... ...... ......... ......... .......... ...... ......... ......... ......... ....... .. 402
XV. Conclusions .......................... ...... ......... ......... .......... ...... ......... ......... ......... ....... ......... ......... ......... ...... .......... .. 402
XVI. Acknowledgments ............ ......... ...... ......... ......... .......... ...... ......... ......... .......... ...... ......... ......... ......... ....... ..... 403
XVII. References .................................. ......... ......... .......... ...... ......... ......... ......... ....... ......... ......... ......... ...... .......... .. 403
I. INTRODUCTION
Human assessments of beauty and human beauty
standards have attracted considerable attention in re-
cent years. Given the interest of this subject to biologists,
psychologists, social workers, medical doctors and lay
people, it seems surprising how little general emphasis
has been put on interpreting these phenomena in a
sexual selection and general evolutionary context. Here
we review the subject. We start out by putting the
study of beauty standards and assessment of beauty into
a sexual selection context. Next, we address the re-
lationship between beauty and health and describe the
consequences of such assessment for individuals. In the
following sections we address research on attractiveness
of beauty of different parts of the human body and the
functional significance of such attractiveness by pre-
senting arguments supporting (pros) and criticizing
(contras) current theories. We end the review by
discussing the ways in which beauty is perceived and
the consequences of such general assessment. Finally,
we present a list of topics for future research.
II. SEXUAL SELECTION AND MATE CHOICE
It is a widespread notion that humans differ funda-
mentally from all other animals and so much that
comparisons are invalid. It is also a widespread belief
that somewhere in the world it is possible to find a
culture where people live in harmonious, non-com-
petitive, altruistic bliss with each other, and were it not
for the existence of Western culture we would be able
to achieve this ideal state. Both claims are erroneous.
Humans carry an incredibly large baggage of evol-
utionary history, and the mere fact that our DNA se-
quences are similar to those of our nearest relatives
among the great apes by as much as 99% makes it a
highly unlikely claim that we could just step out of our
ape dress. Human nature is to a large extent universal.
This includes certain beauty standards and the ways
in which males and females interact, as we will show
below.
Sexual selection theory is concerned with the ad-
vantages that certain individuals have over others of the
same sex and species, in exclusive relation to repro-
duction (Darwin, 1871). What is sexual selection and
why is it important for judgments of human beauty
standards? Sexual selection arises from sexual compe-
tition among individuals for access to mates and has
given rise to the evolution of such bizarre traits as the
antlers of stags, the horns of antelopes, the tail of the
peacock (Pavo cristatus), bird song, frog croaks, and
the extravagant colours of many fish and birds. Darwin
in his 1871 treatise was the first person to realize the
explanation for the evolution and the maintenance of
these bizarre traits that obviously do not enhance the
survival prospects of individuals and therefore cannot
be explained by natural selection. On the contrary,
extravagant secondary sexual characters are costly,
often reduce survival prospects and can only be main-
tained by sexual selection. Two mechanisms are in-
volved in sexual selection: mate competition between
individuals of the chosen sex, usually males, for access to
females has resulted in the evolution of weaponry such
as antlers and horns, but also increases in mere male size
that provides some individuals with an advantage over
others for access to females. The second mechanism is
mate choice by individuals of the choosy sex, usually
females, that has resulted in the evolution of many bi-
zarre traits such as the tail of the peacock, beautiful
coloration in birds and fish and many kinds of bird
vocalizations (Andersson, 1994). Humans are not much
different from other organisms by having evolved sexual
size dimorphism due to male–male competition [more
than 90% of all same-sex homicide involves men in
their early twenties when mate competition is intense
(Daly & Wilson, 1988)], musculature and other features
due to the effects of testosterone at puberty, and female
breasts and facial beauty due to the effects of oestrogens
and male choice.
Extravagant secondary sexual characters in other
species are considered to be beautiful by humans and
perhaps also by animals in general. If both non-human
animals and humans find similar structures attractive,
the likely reason is that animal and human psychologies
have evolved to perceive and become agitated by and
interested in these impressions. Sugar is only perceived
to be sweet by humans because the pleasant and
powerful feeling of sweetness during our evolutionary
386 Karl Grammer and others
past has been shaped by the benefits that we obtained in
terms of energy and nutrition from eating fruits. In the
same way, particular features of faces of women and
particular proportions of waists and hips are only
considered to be beautiful because our ancestors with
such preferences left more healthy offspring than the
individuals in the population without the preferences.
III. SEXUAL SELECTION AND WHY BEAUTY
MATTERS
Sexual selection can work in a number of different ways
because sexual signals may provide different kinds of
information to potential receivers. Human evolutionary
psychological studies across a wide range of cultures
have shown that in consideration of mates men rank
female beauty higher than women rank male looks,
while women rank male resources higher than men rank
female resources (Buss, 1994). Female beauty signals
youth, fertility and health while male resources signal
male competitive ability and health.
The advantages of sexual selection as seen from the
point of view of the choosy partner may derive from the
following (review in Andersson, 1994). Females may
choose males with exaggerated features simply because
such signals indicate the presence of direct fitness
benefits that enhance the reproductive success of choosy
individuals. Males with a high-quality territory or
nuptial gift, males without contagious parasites, and
males with sperm of better fertilizing ability all pro-
vide females with such benefits (review in Møller &
Jennions, 2001). Male displays may also signal benefits
that females do not acquire directly, but only indirectly
in the next generation through the mating success of
the offspring (Fisher, 1930). If the male signal and the
female preference both have a genetic basis, choosy
females will on average pair up with males with
exaggerated secondary sexual characters, and the mate
preference and the signal will become genetically
coupled as a result of this process. The male trait and the
female preference will coevolve to even more extreme
versions that enhance male mating success until the
mating benefit is balanced by an oppositely directed
natural selection pressure, or until the genetic variance
in either female preference or male trait become
depleted. There is little empirical evidence for this
mechanism (Andersson, 1994), but it is likely to work in
most contexts although it will work better in mating
systems with an extreme skew in male mating success.
An alternative model of female mate preferences that
gives rise to indirect fitness benefits is the so-called good
genes hypothesis, which is based on the handicap
principle. Since secondary sexual characters are costly,
only individuals in prime condition may be able to
develop and carry such displays. It is only the differ-
ential ability of certain individuals due to their genetic
constitution that allows them to develop seriously
handicapping and costly traits (Zahavi, 1975). The
honesty and reliability of such displays is maintained by
their costs and their greater cost to low-quality in-
dividuals. A choosy female will, by preferring the most
extravagantly ornamented male, produce offspring of
high viability simply because low-quality individuals
with an inferior genetic constitution will not be able to
cheat and produce an extravagant character. A par-
ticular kind of handicap is the revealing handicap of
Hamilton & Zuk (1982), suggesting that males cannot
help reveal their infection status by virulent parasites
because the presence of such parasites automatically
will be discernible from the expression of their sec-
ondary sexual characters. Thus, females may obtain
reliable information about genetically based parasite
resistance by using male secondary sexual characters as
a basis for their mate choice. There are a number of
studies consistent with this mechanism of sexual selec-
tion (Andersson, 1994), and, on average across species,
approximately 1–2% of the variance in offspring vi-
ability is explained by the expression of male secondary
sexual characters (Møller & Alatalo, 1999).
IV. HUMAN BEAUTY AND SEXUAL SELECTION
Charles Darwin (1871) was the first person to think ex-
tensively and write about human beauty standards from
a biological point of view. The main problem with
Darwin’s approach was that he relied extensively on
correspondence with missionaries in order to obtain
information about the beauty standards in different
human cultures. These data often were collected by
persons with a British beauty standard and thus do not
give evidence for a cross-cultural standard of beauty.
Contrary to most other fields of evolutionary biology,
which were actually advanced by Darwin’s treatments,
Darwin actually stagnated studies of human beauty for
a century by the claims about lack of general principles.
It is only recently that features of human facial and
bodily beauty have been cross-culturally validated
(Singh, 1993; Perrett, May & Yoshikawa, 1994; Thorn-
hill & Gangestad, 1999; Thornhill & Grammer, 1999).
Darwin’s claims about the lack of a general beauty
standard were at odds with the sheer magnitude of the
beauty industry. Although feminist claims may suggest
that this obsession with beauty is an outcome of male-
initiated capitalist activities (see Wolf, 1992), there is
Darwinian aesthetics 387
plenty of evidence for females putting lots of time and
effort into their looks as far back as archaeological and
historical information can date. The human obsession
with beauty in modern Western societies is not much
different from similar efforts in other societies, and the
mere success of the industry is a reflection of the im-
mense strength of the relevant psychological adap-
tations and mate preferences. The strong beliefs among
women in the wonders of cosmetics and their ability to
provide eternal youth obviously are based on the
presence of the same psychological adaptations. Any
book on the use of cosmetics is a manual of how to
accentuate the features that are known to be reliable
health and fertility indicators: oestrogenized faces, and
symmetric facial features. With the development of
plastic surgery these much desired and admired features
of human female beauty can be acquired in a more
permanent state as compared to the temporary state of
cosmetics. Not surprisingly almost all plastic surgery
attempts to correct asymmetries and exaggerate traits
that are considered to be generally beautiful and reliable
indicators of health and fertility.
V. ATTRACTIVENESS AND DAILY LIFE
The human obsession with beauty is not different from
similar obsessions in other organisms. Thus it is quite
likely that human mate selection criteria, which have
evolved through human evolutionary history, are re-
sponsible for the shaping of our perception of attract-
iveness and beauty. In such a view, perception of
attractiveness will be sex-specific because both sexes
have different aspirations for mates. These different as-
pirations are a result of a statistical accumulation of
problems our ancestors have encountered in our evol-
utionary past. If those algorithms which were able to
process information and solve everyday problems better
than others produced more offspring through natural
and sexual selection, we are quite likely to have basic
adaptations in our thinking (Cosmides, Tooby & Bar-
kow, 1992).
Within cultures the generality of attractiveness is
easily accepted. Several rating studies, especially those
by Iliffe (1960) have shown that people of an ethnic
group share common attractiveness standards. In this
standard, beauty and sexual attractiveness seem to be
the same, and ratings of pictures show a high congru-
ence over social class, age and sex. This work has been
replicated several times by Henss (1987, 1988). Thus it
seems to be a valid starting point when we state that
beauty standards are at least shared in a population.
Moreover, recent studies (Cunningham et al., 1995)
suggest that the constituents of beauty are neither ar-
bitrary nor culture bound. The consensus on which a
female is considered to be good looking or not is quite
high in four cultures (Asian, Hispanic, Black and White
women rated by males from all cultures).
Although we are all legally equal’, everyone knows
that people are often treated differently according to
their physical appearance. This differential treatment
by others starts early in life. Three-month-old children
gaze longer at attractive faces than at unattractive faces.
Slater et al. (1998) report two experiments where
pairings of attractive and unattractive female faces were
shown to newborn infants (in the age range 14–151 h
from birth). In both experiments the infants looked
longer at the attractive faces. Following an earlier
suggestion by Langlois, Roggman & Reiser-Danner
(1990) these findings can be interpreted either in terms
of an innate perceptual mechanism that detects and
responds specifically to faces or in terms of rapid
learning of facial features soon after birth. Attractive
children receive less punishment than unattractive
children for the same kinds of misbehaviour. Differ-
ential treatment goes on at school, college and into
university (Baugh & Parry, 1991). In this part of our
lives attractiveness is coupled to academic achieve-
ments. It is common knowledge that attractive students
receive better grades. Moreover female students even
build dominance hierarchies according to attractiveness
(Weisfeld, Bloch & Ivers, 1984). Even when we apply
for jobs, appearance may dominate qualification
(Collins & Zebrowitz, 1995). This differential treatment
reaches its culmination perhaps in the judiciary where
attractiveness can lead to better treatment and easier
convictions. But this is only the case if attractiveness
did not play a role in the crime (Hatfield & Sprecher,
1986). We even believe that attractive people are
better what is beautiful is good is a common stan-
dard in our thinking (Dion, Berscheid & Walster, 1972).
According to evolutionary considerations on a meta-
theoretical level females experience higher cost than
males in opposite-sex interactions because they have the
higher investment in their offspring (Trivers, 1972).
Since females invest more per offspring, their potential
fertility is lower than that of males. Females are thus the
limiting factor in reproduction and males compete for
them. Females in turn choose among males. In humans,
sex differences are most prominent in the role that status
and physical attractiveness play in mate selection (Buss
& Schmitt, 1993). Females value men’s socioeconomic
status, social position, prestige, wealth and so forth and
use these as indicators, more than male attractiveness.
By contrast, men attach greater value to women’s
physical attractiveness, healthiness, and youth; all cues
388 Karl Grammer and others
linked more with reproductive capacity than to female
social status. These sex-specific differences in pre-
ferences have been found in 37 cultures (Buss, 1989).
Men are also more inclined to pursue multiple short-
term mates (that is philandering) and are less dis-
criminating in their mate choices (Buss, 1994).
The final piece of evidence consistent with the hy-
pothesis that evolved human mate selection criteria
shaped our attractiveness standards and created an
obsession with attractiveness would be that attractive
people have more or better offspring in the future. But
there are several caveats for an approach like this:
attractiveness has to be a flexible concept. The reason
for this is that a fixed template for attractiveness could
unnecessarily narrow down the possibilities in mate
selection, as we will show.
VI. HEALTH AND BEAUTY PERCEPTION IN
HUMANS AND OTHER ANIMALS
Parasites and diseases have played an important role in
human evolution, and perhaps even more so than in
many of our close relatives. Parasites exert tremendous
selection pressures on their hosts by reducing their
longevity and reproductive success. It has been known
for a long time that individuals differ in their suscepti-
bility to parasites because of genetically determined host
resistance, and sexual selection for healthy partners
would obviously provide choosy individuals with po-
tentially important fitness benefits (Hamilton & Zuk,
1982). Parasite-mediated sexual selection may benefit
choosy individuals by preventing them from obtaining
mates with contagious parasites that could spread both
to themselves and their offspring, obtaining mates that
are efficient parents, and obtaining mates that are
genetically resistant to parasites (Møller et al., 1999a).
There is considerable evidence for secondary sexual
characters in a wide variety of organisms reliably re-
flecting levels of parasite infections (Møller et al.,
1999a). Studies of a diverse array of plants and animals
show that parasites render their hosts more asymmetric
and hence less attractive than unparasitized individuals
(Møller, 1996b). While secondary sexual characters
may reveal parasite infection status, there is an even
stronger relationship between host immune response
and the expression of secondary sexual characters
(Møller & Alatalo, 1999). While virtually any host
species may be exploited by more than 100 species of
parasites, each with their peculiar ecology, life history
and transmission dynamics, hosts should be expected to
have evolved generalized immune responses to cope
with the most debilitating parasites. This appears to be
the case given that immune responses are much better
predictors of the expression of secondary sexual charac-
ters than are the prevalence or intensity of parasite
infections (Møller et al., 1999a). This is also the case in
humans: people throughout the cultures of the world
value physical attractiveness, but the importance of
beauty is the highest in cultures with serious impact of
parasites such as malaria, schistosomiasis and similarly
virulent parasites (Gangestad & Buss, 1993).
Hosts may reliably avoid the debilitating effects of
parasites by evolving efficient immune defences, and the
immune system in humans is one of the most costly only
equalled by that of the brain. Immune defence may play
a role in host sexual selection because secondary sexual
characters reliably may reflect the immunocompetence
of individuals (Folstad & Karter, 1992). Many sec-
ondary sexual characters develop under the influence of
testosterone and other sex hormones. However, hor-
mones have antagonistic effects on the functioning of
the immune system (e.g. Thornhill & Gangestad, 1993;
Service, 1998), and only individuals in prime condition
may be able to develop the most extravagant secondary
sexual characters without compromising their ability to
raise efficient immune defences. An alternative version
of this model just assumes that both secondary sexual
characters and immune defences develop in response to
condition, and the reliability of the signalling system is
therefore not based on negative interactions between
androgens and immunocompetence (Møller, 1995).
There is some empirical experimental support for the
immune system being involved in reliable sexual sig-
nalling in birds, but tests for humans are still unavailable
(Møller et al., 1999a).
VII. ATTRACTIVENESS AND PHYSICAL
FEATURES
Early approaches to assess physical attractiveness were
done by measuring different distances in faces, having
these faces rated for attractiveness and comparing the
facial distances to these ratings. Features like a high
forehead, large eyes, small nose and a small chin have
been mentioned in many studies as traits of babyness
(Rensch, 1963; Cunningham, 1986; Johnston &
Franklin, 1993). Other studies could not replicate the
appeal of babyness features (Grammer & Atzwanger,
1994; Grammer & Thornhill, 1994). A female trait,
which is linked to attractiveness, replicated by all the
above authors, is a small size of the lower face. Another
feature that could be replicated several times for female
faces is high and prominent cheekbones’. This ma-
turity feature clearly contradicts the presence of an
Darwinian aesthetics 389
attractive babyness feature (Zebrowitz & Apatow,
1984), which would consist of high foreheads, big eyes
and blown up cheeks. There is only one male facial
feature where a positive correlation with attractiveness
has been replicated several times: wide jaws and big
chins and generally bigger lower faces (Grammer &
Thornhill, 1994; Mueller & Mazur, 1997; Thornhill
& Gangestad, 1999).
When we move on to single attractive features of the
body, there are some hints from the literature, e.g. that
female breast size (Hess, Seltzer & Shlien, 1965) and
male shoulder width may correlate with attractiveness
for the other sex (Horvath, 1979, 1981). We will come
back to these two measures later. In addition to this a
positive pelvic tilt in females is one of the bodily fea-
tures judged as being most attractive by males. In regard
to females judging males we mainly find negative as-
pects in judgments: male bellies and male overall fatness
are judged as unattractive (Salusso-Deonier, Markee &
Pedersen, 1991).
(1) Theories of feature processing: pro
Many researchers have taken measurement data and
tried to put them into an evolutionary framework, but
the general approach has changed in recent years. Most
new hypotheses are no longer post-hoc explanations of
existing phenomena, so-called evolutionary story tell-
ing’, instead most of what we know today is derived
from empirical testing of ad-hoc hypotheses generated
from evolutionary theory on a meta-level which uses
biological constraints. If attractiveness has any relation
to mate selection then we would expect two basic dif-
ferences in the evaluation of traits in the opposite sex:
first, traits which guarantee optimal reproduction, i.e.
youth, should be valued, and second, these traits should
be basically those which are sexually dimorphic. This
should be the case because sexual dimorphism is a result
of sex-specific adaptation of a body to the requirements
of the evolutionary past, i.e. survival and reproduction.
In the human face the basic proportions are sexually
dimorphic, male traits develop under the influence of
testosterone (male sex hormones) and female traits
develop under the influence of oestrogens (female sex
hormones). In the case of the broad male chin as a
feature of attractiveness the constraints seem to be
known. If females want dominant males, broad chins
may signal a tendency to dominate others. This is in-
deed the case. Keating, Mazur & Segall (1981) have
shown that males with broad chins are perceived in
eight cultures as those who are likely to dominate others.
Comparable results have been put forward by Mazur,
Mazur & Keating (1984). These authors describe
careers of West Point cadets those with broad chins
at entrance to West Point rose higher in the military
hierarchy than others. In addition, college men with
broad chins copulate more often and have more girl-
friends (Mueller & Mazur, 1997). Winkler & Kirch-
engast (1994) have shown among the !Kung San
bushmen that those males with broad chins and a more
robust body build had a higher reproductive success.
A broad chin could, however, also signal a handi-
cap (Zahavi & Zahavi, 1997) because testosterone pro-
duction might be costly due to the suppression of
immune function and thereby increased disease sus-
ceptibility during puberty (Folstad & Karter, 1992).
Immunocompetence is highly relevant because steroid
reproductive hormones may negatively impact immune
function (Folstad & Karter, 1992). Extreme male fea-
tures, which are triggered by testosterone, thus advertise
honestly that their bearer was sufficiently parasite re-
sistant to produce them. But male facial features cannot
become extreme, as we would expect in a run-away
selection (Fisher, 1930). Perrett et al. (1998) have shown
that adding a feminine touch to a male face can make it
more attractive to some females. The reason seems to be
clear: broad jaws signal high testosterone levels and thus
also possible aggressiveness. If females rely on stable
relationships male aggressiveness may also turn against
them. Thus there is an upper limit for male jaw width
this is when the feature might also become disadvan-
tageous to females. In addition, female chins and lower
faces are small when they are attractive this probably
signals the absence of male sex hormones and the
presence of female sex hormones, which are a necessary
prerequisite for reproduction. Johnston et al. (2001)
examined the facial preferences of female volunteers
at two different phases of their menstrual cycle. In
agreement with prior studies (e.g. Penton-Voak et al.,
1999), their results suggest that women prefer more
masculinized male faces. That is, the attractive male
face possesses more extreme testosterone markers, such
as a longer, broader lower jaw, and more pronounced
brow ridges and cheekbones than the average male
face. This finding suggests that women consider such
testosterone markers to be an index of good health and
that important health considerations may underlie
their aesthetic preference (see for recent review Fink &
Penton-Voak, 2002). However, pronounced testoster-
one facial markers were considered to be associated
with dominance, unfriendliness, and a host of negative
traits (threatening, volatile, controlling, manipulative,
coercive, and selfish). The causal relationship between
testosterone levels and these behavioural attributes
is still controversial (see for review Mazur & Booth,
1998). If such relationships are valid, then the aesthetic
390 Karl Grammer and others
preference of human females may be an adaptive
compromise between the positive attributes associated
with higher than average testosterone (health cues) and
the negative attributes associated with more extreme
masculinization.
Jones (1996) favours the sensory bias theory of sexual
selection as an explanation for human female facial
attractiveness. He shows that the impression of rela-
tively neotenous female faces, i.e. faces that appear to be
younger than the actual age of the face, based on certain
facial proportions small lower face, lower jaw and
nose, and full large lips are rated as more attractive by
male raters from five populations. He also found that
US women models have neotenous faces compared to
female undergraduate students. Thus, markers of high
oestrogens levels may reliably signal an immune system
of such high quality that it can deal with the handicap of
levels of high oestrogens (Thornhill & Møller, 1997).
Also, there is evidence that oestrogens’ by-products are
toxic to the body (Eaton et al., 1994; Service, 1998).
Thus, markers of oestrogens may honestly signal ability
to cope with toxic metabolites (Fink, Grammer &
Thornhill, 2001; Fink & Penton-Voak, 2002).
Sexually dimorphic traits in the human body can be
found in the distribution of body fat (breast and but-
tocks), the general structure of the skeleton, i.e. bigger
and more massive shoulders in males, and bigger pelvis
in females, and finally muscular build. Male muscular
build is a main difference. Again muscles are built under
the influence of male sex hormones, and muscles will be
of use for males in male competition.
The signalling value of body features in the case of
females seems to be linked to reproductive stage. Im-
portant sex differences in our bodies depend on fat
distribution. The amount of fat in the female body is
responsible for a stable level of female sex-steroids (fat
equals 25% of body mass; Frisch, 1975). Thus the
amount of visible fat can predict if a female is receptive
or not. In order to strengthen its signalling value, body
fat must be distributed over prominent places like
breasts and buttocks. Otherwise its signalling value may
be lowered and thus body fat may simply restrict the
biomechanical abilities of the body. Indeed, breast size
correlates with overall body fat (Grammer, 1995).
Furthermore, overall weight is linked to fertility: heavier
mothers have more children (Grammer, 1995; Singh &
Zambarano, 1997). Appreciation of heavier women in
various cultures seems to depend on environmental
stability (Anderson et al., 1992). In unstable environ-
ments plumpness is linked to status and attractiveness.
Besides being sexually dimorphic, the distribution of
body fat can also signal youth and neoteny. Firm breasts
with small aureole and erect nipples and the breast axis
pointing upward in a V-angle are rated as attractive
(Grammer et al., 2001).
Another signal is the absence and presence of body
hair, which is also sexually dimorphic, and thus a fea-
ture of attractiveness. Females appreciate body hair
developed under male sex hormones but males prefer its
absence on females. Removal of female body hair thus is
more prominent. Length and colour of females’ hair of
the head are attractiveness traits. Rich & Cash (1993)
have shown that blonde hair, although only infrequent
in most populations, dominates in pictures of females
presented in magazines for males.
Thelen (1983) showed that preference for hair colour
depends on the distribution of hair colour in a popu-
lation. Males prefer the rare colour. The reason for
this situation might be a quest for rare genes, which
could help in the host–parasite race discussed below. In
addition males seem to prefer long hair (Grammer et al.,
2001) and female hair growth on the head is more stable
than that of males. Indeed hair loss and baldness are a
result of male sex hormones (Muscarella & Cunning-
ham, 1996; Anderson, 2001; Choi, Yoo & Chung,
2001). Long hair thus is again sexually dimorphic. The
general function of hair (on the head, in the armpits and
pubic hair) may be the distribution of human pher-
omones produced in the apocrine glands. Hair will give
a greater surface for its distribution into the air (see also
Stoddart, 1990). We will see below that sexual pher-
omones play a major role in attractiveness. Long female
hair thus would be a pheromone-distribution organ’.
Body forms also have an inherent signal value: they
create regions of high contrast, which in return keep
attention. In a study where an eye-mark recorder re-
vealed male fixations on a female body, males tended to
fixate the body contours and regions of high contrast
like the shadows under the breasts (Santin, 1995).
(2) Theories of feature processing: contra
There are, however, many objections to a simple fea-
ture-based approach to the decoding of attractiveness.
One of them is methodological. Most researchers
measure many features (up to hundreds) and then they
correlate them with attractiveness. There is often no
correction for a large number of statistical tests, and
hence the replication rate of many attractiveness traits
from these studies is very low. Other reasons for in-
consistent results regarding the attractiveness of exag-
gerated facial traits may have arisen from differences in
samples of perceivers used. For example, Little et al.
(2001) explored how female self-rated attractiveness
influences male face preference by females and found
that there is an increased preference for masculinity and
Darwinian aesthetics 391
symmetry by women who regard themselves as at-
tractive. Some other studies considered the position
in the menstrual cycle when females rated male faces,
and this has repeatedly been shown to affect ratings
significantly (e.g. Penton-Voak et al., 1999; Johnston
et al., 2001). A lot of previous studies did not control for
these variables, although this is likely to obscure any
relationships due to noise. However, there is a strong
demand for consistent research designs in future studies
to make results comparable (see Fink & Penton-Voak,
2002).
One of the earliest assumptions in beauty research
was that innate templates are involved in recognition of
attractiveness, like the one for babyness (Zebrowitz,
McArthur & Apatow, 1984). Babyness is normally
perceived as positive and females react to babyness with
an increase in frequency of smiles (A. J. Friedlund &
J. M. Loftis, unpublished observations). This almost
automatic reaction has led to the assumption that ba-
byness could also be involved in adult attractiveness
perception, because it could signal neoteny or youth.
But the main characteristics of babyness like big puffy
cheeks are unappreciated by males. High cheekbones,
as a sign of maturity, must be added for a female face to
be viewed as attractive (Grammer & Atzwanger, 1994).
But it is not this simple. Research finds that babyness
in its original expression is not attractive at all, because
males attribute to it negative behavioural tendencies: a
babyface seems to imply being babyish (Grammer &
Atzwanger, 1994). So, if you remove one part of a
template (the puffy cheeks) it is not a template any
more there seems to be a completely new Gestalt’,
and thus a new template. We call this template sexy-
scheme’, because it is a combination of parts of the
babyness feature (signalling youth) and high and promi-
nent cheekbones (signalling maturity). The prominent
cheekbones themselves seem to be a trait developed
under the influence of female sex hormones (Symons,
1995) and thus possibly signal an immunocompetence
handicap comparable to male jaw size.
Moreover the presence of babyness features in a fe-
male face leads to a transfer of personality traits. Per-
sonality traits coupled to babyness are positive and
negative: babyness can signal submission and elicit-
ing parental care but it also signals incompetence’,
which would be the last trait in a partner one would look
for. Raising offspring requires competence. If a female is
very young and at the optimal age of reproduction, she
is likely to be an incompetent mother. The optimal age
of reproduction is reached at age 24 (Buss, 1989). If
attractiveness judgments vary with age, 24-year old
females should receive the highest scores. This seems to
be the case (Grammer & Thornhill, 1994). All features
described as attractiveness features are sex-specific and
are also responsible for gender recognition. When one
examines how many measures are necessary to dis-
criminate between faces of different sexes we find that a
combination of only 16 different measurements reaches
sufficient reliability (Bruce et al., 1993). So gender
recognition per se seems to be more than simple feature
analysis.
Comparable ideas relate to the perception of bodies.
In most studies we find a curvilinear relation between
features and attractiveness. This means that attractive
features are neither too small nor too big. Legs should
be neither too short, thick, thin or long (Ronzal, 1996).
The same applies to attractive breasts. Moreover,
what is attractive seems to be gender prototypical,
i.e. sexually dimorphic. Rensch (1963) recognized
this relation between stimuli and attractiveness quite
early, after studying facial attractiveness and he came
to the conclusion that those features which are gender-
prototypical are those which are rated as attractive.
VIII. THE ATTRACTIVE PROTOTYPE : FACES
What could the Gestalt we use for attractiveness and
beauty decoding then be? A basic feature of human
cognition is the creation of prototypes (Rosch, 1978).
This means that we constantly evaluate stimuli from our
social and non-social environment and classify them
into categories and concepts, thus reducing the amount
of environmental information into pieces’, which can
be used or stored very economically. For a first ap-
proach let us assume that prototypes are some kind of
average representation of stimuli of one class.
There are some hints that our brain solves the
problem of storing faces with the help of prototypes. We
seem to build facial prototypes and then simply assess
the deviations of a single face from these prototypes.
Children build such facial prototypes very early and
when confronted with average faces in recognition tests
children give false alarms to them (Bruce, 1988). They
behave as if they had seen them before, although they
have not. Moscovitch, Winocur & Behrmann (1997)
put forward the idea that there is a holistic processing
involved in face recognition. The spatial relations
among its components define the Gestalt but this Ge-
stalt is more than the sum of its parts. From this starting
point basically three hypotheses emerge. The first is
norm-based coding (Rhodes, Brennan & Carey,
1987), where averaging a large number of faces in the
brain derives the norm. The second hypothesis is the
density alone hypothesis’, where the Gestalt is a point-
by-point representation in a multidimensional space
392 Karl Grammer and others
(Valentine, 1991). The third hypothesis is the tem-
plate hypothesis, which suggests that the brain analyses
the single parts with templates and then reintegrates
them (e.g. Farah, 1990; Corballis, 1991).
Moscovitch, Winocur & Behrmann (1997) analysed
the three hypotheses using the performance in face
recognition of a patient who suffered from object ag-
nosia after a brain trauma but was able to recognize
faces. Interestingly this patient could recognize atypical
faces, cartoons, family resemblance, and he had a good
memory for unfamiliar faces. However, he was unable
to recognize a face when it was inverted, when single
features were inverted, when spatial features were dis-
torted and when faces were misaligned. These results
suggest that we indeed process faces via norm-based
coding: the patient could process faces only as a
whole’. If this is so, norm-based coding will be one of
the main processes involved in the assessment of beauty.
As soon as prototypes are present they can be used for
learning. We learn very fast and almost irreversibly to
link personality traits with facial prototypes (Henss,
1992). This helps us to decode behavioural tendencies
of people we meet, and thus we are able to structure our
behaviour accordingly. Indeed, several studies have
repeatedly shown that computer-generated proto-
typical faces are more attractive than the single faces
which have been used to generate them (Galton, 1878;
Kalkofen, Mu
¨
ller & Strack, 1990; Langlois & Rogg-
man, 1990; Mu
¨
ller, 1993; Grammer & Thornhill,
1994; Perrett, May & Yoshikawa, 1994). But there
are two caveats again: this is only replicable for female
faces and all researchers find that there are some indi-
vidual faces which are more attractive than the proto-
types.
IX. THE ATTRACTIVE PROTOTYPE : BODIES
Prototyping does not only apply to faces. Comparable
results are reported for the attractiveness of averageness
for female body features. The waist-to-hip ratio (WHR)
has been suggested to be a good predictor of the ability
of women to produce male offspring. Thus, an an-
drogynous body shape may be judged as most attractive
in cultures that value male children. Several studies
have described WHR in women as a single measure
linked consistently across studies to bodily attractiveness
(Singh, 1993, 1995). There is a curvilinear relationship
to attractiveness with a maximum attractiveness at 0.71.
Surprisingly this maximum is related to many health
features in women. Moreover there is a direct link to
fertility: females with an optimal WHR become more
often and more quickly pregnant through artificial
insemination. It has been taken for granted for a long
time that the preference for body shapes at the popu-
lation mean is cross-culturally stable. Research in Great
Britain and Uganda showed similar results (Furnham &
Baguma, 1994). Recent studies, however, found that
male preferences for a low WHR is not culturally uni-
versal (Yu & Shepard, 1998). Furthermore, Tove
`
e et al.
(2001) suggested that differences in attractiveness pre-
ferences between different ethnic groups appear to be
based on weight scaled for height (the body mass
index or BMI) rather than WHR. Although there is
a preferred optimal BMI for each ethnic group, which
will balance environmental and health factors, this
optimal BMI may differ between groups and environ-
ments.
One problem of these studies is that women included
in the samples do not represent the average of the actual
female population at the age of optimal reproduction.
German measurements of 10 000 young adult females
show a much higher average WHR (Grammer, 1995).
Generally, waists have higher measures in the popu-
lation than perceived as optimal and attractive.
For instance, in Playboy centrefolds, breast measure-
ments are around the population mean (population
mean=88.4 cm in Germany; 88.8 cm in Playboy
centrefolds), but waist measures are 7.2 cm smaller in
Playboy centrefolds than the population mean for
German females (see Garner et al., 1980). The con-
clusion up to this point is that beauty is averageness, but
with exceptions.
(1) Theories of prototype processing: pro
If our brain uses prototypes, averageness might well be
coupled with being prototypical’. Thus there might be
a better fit of the stimulus onto the prototypical tem-
plate. As a result, prototypes are recognized faster and
better and thus might create higher nervous excitation.
This could be the reason that averageness is preferred.
Our brain could accept more willingly better fitting
stimuli. Mu
¨
ller (1993) has called this process neuro-
aesthetics’.
The fact that female attractiveness should be the
average was predicted by Symons (1979) on completely
different grounds. He proposed that males should avoid
mating with females who are at the extremes of a
population, because these females may carry disad-
vantageous genes. Prototypes do portray some genetic
information. With respect to features with additive
genetic variance, homozygous individuals tend to be
over-represented at the extreme tails of the distri-
butions. By contrast, heterozygous individuals tend to
be over-represented at the middle of such distributions
Darwinian aesthetics 393
(Soule
´
& Cuzin-Roudy, 1982). However, the traits
studied by Soule
´
& Cuzin-Roudy (1982) were not
secondary sexual characters, so it remains unclear to
which extent their observations can be extended to this
category of traits.
By contrast, male gender prototypes are not average
(Grammer & Thornhill, 1994). We have seen that large
facial traits are attractive for males. The most interesting
feature of prototypes is that attractive prototyping al-
lows two things: first, we are able to adjust our beauty
standards to the mean of the population. This is nicely
demonstrated by the Farrah-effect’. Men who see
films with beautiful women adjust their beauty stan-
dards accordingly as compared to controls (Kenrick &
Gutierres, 1980; Kenrick, Gutierres & Goldberg, 1989).
They then have higher aspiration to attractiveness in
a dating experiment. Media thus can create unreal
beauty standards. Second, prototyping opens our
possibilities of mate-choice. If we had an innate tem-
plate for attractiveness, we could run the danger of
either never meeting somebody fitting the template,
or being frustrated by the non-fitting mates we find.
Through prototyping our beauty standard is adjusted to
the population in which we live. If the distribution of
traits is a normal one, there are simply more average
people than extremes. This creates a bigger population
of possible mates. In view of this we should expect some
learning mechanisms involved in beauty standards, and
an almost automatic fitting of these standards to the
population in which we live which again increases our
chances of finding a mate.
(2) Theories of prototype processing: contra
One problem for prototyping is created by the fact that
we recognize average faces poorly (Bruce, 1988), be-
cause they do not deviate from the templates we use to
store faces. With individual recognition as the basis for
social interaction, attractive people would have a
handicap when it comes to pro-social exchange. If this is
true, we should expect deviations from averageness in
beautiful faces. One has to be recognizable and distinct.
Adding an individual touch to averageness could thus
make an attractive face beautiful.
There are still more problems with this approach.
First, the computer-generated prototypes lack skin
blemishes and faces appear much softer than in normal
pictures. Second, there are single faces that are not
prototypical at all, and they are constantly rated as more
attractive (Grammer & Thornhill, 1994; Perrett et al.,
1994). Third, symmetry could play a role; composite
faces are much more symmetrical than the single faces,
which are used to form the prototype. Computed
averageness is symmetrical; thus we have to control for
symmetry and determine what role it plays for proto-
types and attractiveness.
X. DEVELOPMENTAL STABILITY AND BEAUTY
Developmental stability reflects the ability of individuals
to maintain stable development of their morphology
under given environmental conditions (Møller &
Swaddle, 1997). While developmental noise and vari-
ous developmental upsets tend to destabilize develop-
ment, developmental control adaptations have the
opposite effects on the phenotype. Measures of devel-
opmental instability include fluctuating asymmetry
and the frequency of phenodeviants, but also other
measurements. A character demonstrates fluctuating
asymmetry when symmetry is the norm and deviations
from symmetry are randomly distributed with respect
to side (Ludwig, 1932). Phenodeviants are relative
large deviations from normal phenotypes such as a
position of the heart in the right side of the body
cavity or the presence of an even number of fingers
on a hand.
Fluctuating asymmetry is a particularly useful mea-
sure of developmental control ability for several reasons.
First, we know the optimal solution a priori: it is sym-
metry. Second, fluctuating asymmetry develops in
response to an enormous range of genetic and en-
vironmental factors that tend to upset developmental
processes (review in Møller & Swaddle, 1997). Third,
fluctuating asymmetry can be measured accurately with
practice and we can investigate plants, insects, birds and
humans using the same simple and uncostly tool, a
precise ruler. Fourth, we cannot investigate how plants
and animals feel about or perceive their environment,
but we can answer this question indirectly by measuring
their asymmetry because asymmetry reliably inte-
grates the consequences of many disruptive effects of
the environment. Since the optimal phenotype is the
symmetric one because it promotes performance, any
deviation from perfect symmetry can be considered a
sub-optimal solution to a design problem that will result
in performance problems in the future. It was probably
difficult for a pre-historic human to escape from a lion,
but it was even more difficult to escape with two legs
of unequal length. Indeed, skeletal remains from pre-
historic Indians have shown that individuals that were
old had more symmetric bones than individuals that
died young (Ruff & Jones, 1981). This finding is par-
ticularly interesting because continuous re-modelling
of bones during life generally gives rise to increasing
asymmetry among older humans.
394 Karl Grammer and others
It is perhaps not surprising that asymmetry has been
found to be important for plants and animals including
humans when faced with the realities of life, the struggle
for survival, mates and reproduction (Møller & Swad-
dle, 1997). The continuous selection against asymmetry
starts already among sperm and eggs within females
of species with internal fertilization: developmental
selection against deviant gametes and zygotes appears
to be a very widespread phenomenon. Fruit and seed
abortion is extremely common in plants. Experimental
work has demonstrated that in the flowering plant
fireweed (Epilobium angustifolium) around three-quarters
of all embryos are aborted during the first few cell div-
isions because of irregular developmental patterns
(Møller, 1996a). Interestingly, the abortion frequency is
directly related to the symmetry of the flowers of both
the pollen donor and the pollen recipient. Similar
phenomena have been described among a wide range of
organisms spanning invertebrates and vertebrates in-
cluding humans (Møller, 1997). Infanticide has been
and is still a common practice in many human societies
mainly directed towards children with deviant pheno-
types. This behaviour has obviously been adaptive by
avoiding wastage of costly resources on offspring with
poor survival prospects. Evolutionary psychological
studies of parental reactions to newborns have dem-
onstrated that modern human beings still carry
psychological adaptations towards this end by reacting
with strongly negative feelings that best can be de-
scribed as disgust and aggression when confronted
with children with increasingly deviant appearances
(Daly & Wilson, 1988).
Asymmetry also matters when it comes to the mating
game. Developmental stability and sexual selection are
closely associated in a wide variety of organisms ranging
from plants, flies, grasshoppers and fish to birds and
mammals (Møller & Thornhill, 1998; Møller & Cuervo,
in press). For example, women prefer men with sym-
metric faces and bodies (Grammer & Thornhill, 1994;
Thornhill & Gangestad, 1994; Jones et al., 2001), and
the number of sexual partners during life is directly
related to skeletal asymmetry in men (Thornhill &
Gangestad, 1994; Gangestad, Bennett & Thornhill,
2001). Since symmetry relates to performance in gen-
eral, choosy females that prefer symmetric males will
obtain mates that are better able to provide resources,
but also able to provide genes for developmental health
to the offspring. Given the intense developmental
selection against asymmetric offspring, females will also
benefit in terms of increased fecundity.
Some bodily and facial asymmetries manifest them-
selves very early in human development and remain
stable during lifetime (Thornhill & Gangestad, 1996;
Thornhill & Møller, 1997). These minor physical
anomalies (MPAs) seem to be the result of develop-
mental instabilities during early embryonic develop-
ment. MPAs are formed in the first trimester of ges-
tation, and fluctuating asymmetries develop throughout
life. However, several studies have shown positive
correlations between the frequency of MPAs and fluc-
tuating asymmetry. At this point we have to distinguish
between MPAs and bodily laterality. Usually the sides of
the body differ, but the vertical body symmetry line can
still be a straight line (despite laterality being present).
Asymmetries in the face distort this straight line into a
zigzag line.
Thus MPA or fluctuating asymmetry may be a sig-
nificant negative predictor of attractiveness and used as
a negative scale for prototype beauty. Comparable re-
sults can be found for the rating of bodily attractiveness
in relation to breast asymmetry (Singh, 1995). Sym-
metrical breasts are more attractive than asymmetrical
breasts. Moreover, breast asymmetry is a significant
negative predictor of lactation ability and even repro-
ductive success (Møller, Soler & Thornhill, 1995). Thus
bodily and facial symmetry seems to be important in
ratings of attractiveness.
(1) Theories of symmetry and
attractiveness: pro
Host–parasite co-evolutionary cycling predicts that
parasite resistance should be a trait that is valued in
mate selection (Møller et al., 1999a). One defence
against parasites is the production of substantial poly-
morphism: when a parasite adapts to one allele, alterna-
tive alleles may be advantageous. Pathogens are a major
environmental perturbation underlying developmental
instability, and developmental stability may be related
to additive genetic variance in disease resistance, which
in turn may relate to fitness. Thus symmetry, which
cannot be faked, may be an honest signal of mate
quality. Symmetry as a mate-selection criterion has been
shown in many species, from insects, birds and mam-
mals to humans (reviews in Møller & Thornhill, 1998;
Møller & Cuervo, in press). Moreover attractiveness
plays a prominent role in mate selection in those human
societies where parasites are prominent (Gangestad &
Buss, 1993). The basic finding is that if symmetry is
present in the face or the body an individual is judged as
being relatively attractive, and if the body is asymmetric
the face is rated unattractive, even if the rater never sees
the body (Thornhill & Gangestad, 1993, 1994; Gang-
estad, Thornhill & Yeo, 1994).
Although this statement should be qualified, let us
simply assume that symmetry seems to be influential in
Darwinian aesthetics 395
ratings of attractiveness and thus mate choice. In ad-
dition, we note that symmetry may be more salient than
averageness (Grammer & Thornhill, 1994).
(2) Theories of symmetry and
attractiveness: contra
There are several objections against a theoretical con-
nection between fitness and facial and bodily symmetry.
Most of these objections can be subsumed under the
term sensory exploitation’. Research on the perception
and computation of stimuli has shown that symmetry of
stimuli is one of the main factors in recognition and
reaction to stimuli. Several computer simulations have
shown that neural networks, when confronted with
stimuli, respond better and easier to symmetrical ones
( Johnstone, 1994). Enquist & Arak (1994) have shown
that symmetry preferences may arise as a by-product
of the need to recognize objects irrespective of their
position and orientation in the visual field. Another ob-
jection is that templates can only be constructed sym-
metrically (Eibl-Eibesfeldt, 1997), although this poses
the question why such a mechanism came about, if not
for the utility of processing information about biological
entities. Symmetrical stimuli thus would exploit the
sensory system of the receiver. Symmetrical patterns are
attractive for humans in contexts unrelated to signal-
ling, although it remains unclear whether such pre-
ferences arose before or after the preferences for
symmetry of sexual signals. Such general preferences
for symmetry serve no obvious function (Rensch, 1963).
Preferences for symmetry might have evolved for
completely different reasons, but independent from this,
they have an effect on mate choice, related to repro-
ductive success. Using photographs of men’s faces, for
which facial symmetry had been measured, Scheib,
Gangestad & Thornhill (1999) found a relationship
between women’s attractiveness ratings of these faces
and symmetry. Interestingly, subjects could not rate
facial symmetry accurately. Moreover, the relationship
between facial attractiveness and symmetry was still
observed, even when symmetry cues were removed by
presenting only the left or right half of faces. These
results suggest that attractive features other than sym-
metry can be used to assess phenotypic condition. The
authors identified one such cue, facial masculinity.
Further empirical results are also strongly contrary to
this incidental effect hypothesis for the human facial
symmetry preference and strongly consistent with the
good genes hypothesis for the preference (i.e. the pre-
ference favours healthy individuals). Little et al. (2001)
found that experimental manipulations of facial sym-
metry have a greater influence on opposite-sex facial
attractiveness ratings than on same-sex ratings. In their
study comparing females who did and did not consider
themselves to be physically attractive, those who con-
sidered themselves physically attractive showed a
greater preference for two proposed markers of quality
in male faces: masculinity and symmetry. Little et al.
(2001) consider this a condition-dependent mating
strategy analogous to behaviours found in other species.
In other words, the absence of a preference for markers
of good genes may be adaptive in women of low mate
value to avoid the costs of decreased parental invest-
ment from the owners of such characteristics. This was
repeated by Jones et al. (2001). The latter study also
found the same bias when judging apparent health:
facial symmetry variation affects opposite-sex health
ratings more than same-sex ratings, and symmetry is
rated as healthy. These sex-specific patterns are in-
consistent with the claim that symmetry is found at-
tractive as a by-product of the ease with which the
recognition system can process symmetric stimuli. As
strongly inconsistent with this suggestion was Jones
et al.’s (2001) report that the relationship between
facial attractiveness and facial symmetry is mediated
by a link between judgments of apparent health and
symmetry. When the effect of attractiveness ratings
was controlled statistically, the positive relationship be-
tween facial symmetry ratings and health remained
significant. This negates the view that the relationship
between facial symmetry and rated health may be
caused by an attractiveness halo effect. But when the
effect of judgments of apparent health was controlled,
the relationship between facial symmetry and attract-
iveness disappeared. This latter result establishes
that the facial attractiveness–symmetry relationship is
mediated by the link between facial symmetry and
judgments of health.
Another feature which should be mentioned here is
the role of skin texture because blemishes in the skin can
also be related to the perception of symmetry. Blemishes
can appear and disappear and distort symmetry. The
role of skin texture and facial symmetry has not been
subject to much scientific research. No research has
been done to solve the problem of what causes the
higher attractiveness of some faces compared with a
symmetric composite. Both average faces and more
symmetrical faces could be recognized simply by tex-
ture. In a recent study, Fink et al. (2001) used the co-
occurrence-matrix (Haralick, Shanmugam & Dinstein,
1973) to classify female facial skin texture. With this
image segmentation method, Fink et al. (2001) showed
that human skin provides reliable cues of its health
condition. Skin texture plays a role in normal faces the
more homogenous these faces, the more attractive they
396 Karl Grammer and others
are. But this could be due to a correlation between
symmetry, averageness and texture. Symmetry was
controlled to ensure that a significant correlation be-
tween attractiveness and texture was not caused by the
influence of symmetry. For this purpose, Fink et al.
(2001) used the techniques of morphing and warp-
ing the original faces to remove their individual shape
and thus standardize them. Skin texture still had a
significant influence on the judgment of female facial
attractiveness: a homogeneous skin texture was con-
sidered more attractive. This effect might be caused by
cues of immune function that we derive from skin
condition. There is a relationship between dermatoses
and elevated levels of sex hormones (testosterone,
oestrogens), and this is often correlated with ovarian
dysfunction (e.g. polycystic ovary syndrome; Stein-
berger et al., 1981; Schiavone et al., 1983). Accordingly,
Mackintosh (2001) evaluated the antimicrobial prop-
erties of melanocyctes. He presented evidence that
melanization of skin and other tissues forms an im-
portant component of the innate immune defence
system following previous findings by Duke-Cohan et al.
(1998) and Duke-Cohen, Tang & Schlossman (2000)
on attractin, a protein that seems to have regulating
functions on melanization and immunity. Melanization
and the innate immune response are likely to be func-
tionally, biochemically and genetically linked. Mack-
intosh (2001) suggested that melanized tissue acts to
seal off vulnerable nutrient-rich tissues from microbial
attack. Consequently, the greater the melanosome and
melanin content, the more protective will be the effect.
This may underlie the finding by Fink et al. (2001)
who showed that among Caucasian females, a darker
skin complexion is considered more attractive than a
light one.
XI. CROSS-SENSORY MODALITIES:
BODY ODO UR, VOICES, DECORATION
AND MOV EMENT
A pheromone is a chemical signal emitted by one in-
dividual that alters either the behaviour or physiology of
another individual (Luscher & Karlson, 1959). Several
investigations on human body odour revealed the rel-
evance of olfactory communication in humans and its
implications for sexual behaviour (Schaal & Porter,
1991). Human body odour has been reported to in-
fluence female mate choice and may allow finding a
partner who possesses complementary immune re-
sponses (Wedekind et al., 1995). Females find the body
odour of those males attractive whose major-histo-
compatibility complex (MHC) is different from their
own. In this case sexual attractiveness of body odour
would lead to heterozygotic offspring. Moreover, as
suggested by Grammer (1993), an important possible
function of a pheromone would be the induction of
effects, because emotions may change information
processing in the receiver. Via a pathway to the limbic
areas of the brain, the chemical signals carried by
odours have a direct influence on emotions. Odours
induce negative or positive moods and feelings. Thus,
odours allow direct manipulation of cognition. Indeed,
odours modify the social perception of other persons
(Cowley, Johnson & Brooksbank, 1977). Emotions
and moods again drastically change social perception
and information processing (Forgas & Moylan, 1991).
Isen (1984) found that people who are in a positive
mood often use more heuristic, truncated processing,
use larger and more inclusive categories, and are more
likely to take risks, provided that such risks do not
threaten their positive mood state. These people seem to
use an effort-minimizing, simplifying processing style.
The possible effect of pheromones on mood makes
pheromones a highly likely candidate for the alteration
of attractiveness assessments. In humans, we have seen
that sexually asymmetric parental investment leads to
gender-specific mate-selection criteria. If those criteria
are part of the adapted mind, they become prone to
exploitation. This is the case especially for male mate-
selection criteria, because males rely so importantly on a
single optical stimulus: female attractiveness.
In their vaginal secretions females produce a sample
of fatty acids with behavioural effects, referred to as
copulins (Curtis et al., 1971). They were originally
discovered by Michael & Keverne (1968) in rhesus
monkeys (Macaca mulatta). Although normally motiv-
ated to copulate, when sexually inexperienced rhesus
males were made anosmic, they showed no further
sexual motivation despite a powerful visual cue: the
female’s sexual swellings (Michael & Keverne, 1968).
Furthermore, rhesus males show no interest in ovar-
iectomized rhesus females, presumably because ovar-
iectomized rhesus females lose the odour characteristic
of ovulation. Rhesus males regained interest in
copulation when the vaginal secretions from non-
ovariectomized females were applied to ovariectomized
females. Studies on menstrual cycle fluctuations in the
fatty-acid composition of women’s vaginal fluids in-
dicated that a similar type of signalling system might
also exist in humans (Waltman et al., 1973; Michael,
Bonsall & Warner, 1974; Michael, Bonsall & Kutner,
1975; Preti & Huggins, 1975). Further investigation
on the function of copulins in humans remain to be
conducted, but we may assume that ovulation is not
concealed, and that men could use ovulation-linked
Darwinian aesthetics 397
odours in their mate selection. This has now been
shown in four separate studies (Gangestad & Thornhill,
1998; Rikowski & Grammer, 1999; Thornhill &
Gangestad, 1999; R. Thornhill, S. W. Gangestad,
R. Miller, G. Scheyd, J. Knight & M. Franklin, in
preparation).
The strongest pattern in human sex pheromone
research pertains to the body scent of symmetric
men. Such men’s body scent is attractive to women,
especially women at peak of fertility in their menstrual
cycles. Although men do not show a consistent
preference for the body scent of symmetric women, two
studies have found that men prefer the body scent of
women at ovulatory menstrual cycle phases (Singh &
Bronstad, 2001; R. Thornhill, S. W. Gangestad, R.
Miller, G. Scheyd, J. Knight & M. Franklin, in prep-
aration).
In addition to smell many other signals might con-
tribute to attractiveness. Voice quality is another can-
didate. Unfortunately there is not much research in this
direction. Zuckerman & Driver (1989) found attract-
iveness prototypes in voices. People tend to agree what
an attractive voice is. They tried to disentangle the
physical qualities of voices in relation to attractiveness.
The only variable found is frequency: deep male voices
are rated attractive. The biological background for such
an attractiveness rating might lie in the fact that usually
the size of the voice-producing apparatus (the larynx
and the oral cavity) correlate with body size, which is
sexually dimorphic and thus again prototypical for
males. Vocalizations may also provide reliable infor-
mation about health status as is shown by evidence from
studies of bird song reflecting current health status
(Saino et al., 1997) and the calls of nestling birds re-
flecting their current health status (Sacchi, Saino &
Galeotti, 2002). Hertrich & Ziegelmayer (1988) have
examined to what extent the speaker’s body size and
shape are betrayed in his speech signal and thus can be
recognized by listeners. Contrary to earlier consti-
tutional studies only size and not shape correlates with
acoustical parameters of speech; comparing listening
experiments with acoustical analysis gives some evi-
dence that the average sound spectrum is used by lis-
teners to judge the speaker’s body size. This effect may
be direct, or it may be an indirect signal of symmetry
since symmetric males tend to have more attractive calls
in jungle fowl Gallus gallus (Furlow, Kimball & Marshall,
1998). Furthermore, large men are more symmetric
than small men (Manning, 1995).
Another point often put forward in attractiveness
research is the role of decoration and subsequent body
changes. Decoration allows primarily two things: first
decoration can alter perception of attractiveness and
second, decoration can be used for in-group identifi-
cation. Grammer (1998) showed that advertisement by
females is mostly about body improvement and dec-
oration. Here culturally determined ways of body im-
provement operate on a biological theme. Grammer
(1995) showed that although body decoration may
significantly change ratings of attractiveness and per-
sonality, it cannot change the overall perception of a
person. For instance, if a female is rated as dominant
and not romantic, only the overall perception of the
magnitude of the rating changes, but the female will
not be rated subordinate and romantic. Low’s (1979)
research suggests an interesting link to decoration and
parasite resistance. Decoration may be most prevalent
in those societies where parasite load is high. In this view
decoration would follow a handicap principle and it
would signal that the decorated person is able to spend
time on decoration. Recent studies of perfume use in
western society have suggested that perfume is a means
of enhancing or exaggerating differences in natural
odour related to MHC genotype, again suggesting a
clear link to parasite-mediated sexual selection (Milinski
& Wedekind, 2001).
Last but not least we will look at the beauty of human
movement. It is obvious that movement plays a role in
person perception. In German expression psychology
the assessment of movement quality played an im-
portant role as early as at the beginning of the 20th
century. Fischer (1911) introduced an objective method
for the analysis of movements from films by measuring
the co-ordinates of joints in every film picture. Conse-
quently, Flach (1928) postulated: a symbol alone, a
gesture is ambiguous. In contrast, the dynamics of a
movement are unambiguous and convincing (p. 461,
authors’ translation).
Unfortunately, this objective approach was dropped
in favour of subjective interpretations of the objective
measurement data. Fifty years later, Johansson (1973,
1976) fixed point-light displays to the joints of partici-
pants and filmed their movements in the dark. If such
point-light films were shown to raters, they would re-
cognize sex, age and movement patterns (e.g. walking),
but if presented as a fixed image, the points would
appear randomly distributed (Cutting & Proffitt, 1981;
Runeson & Frykholm, 1983). Observers are able to
detect effort, intention and deception from body
movements. Berry et al. (1991) used quantized videos,
which obscured the individual information and left
only the movement visible for rating. In this approach
the constraints of point-light displays are not present.
The new method allowed the above results to be re-
plicated. In neither approach was an empirical de-
scription of movement quality carried out. A substantial
398 Karl Grammer and others
gap therefore exists between the fact that we know
that participants are able to deduce information from
movement and the description of the kind of infor-
mation the participants use. Nonetheless, the results of
the above work also suggest that movement alone can
carry a meaning.
Thus movement carries at least one type of infor-
mation for certain: gender identification, which is one
source for attractiveness ratings. But there are two more
likely candidates we have to look at: symmetry and
hormone profiles. Møller, Sanotra & Vestergaard
(1999 b) showed that symmetric chickens show more
coordinated and more efficient walking behaviour.
Thus symmetry could not only be an indicator of de-
velopmental stability; it could also be an indicator of
movement efficiency and thus bodily efficiency of an
individual. However, since external developmental in-
stability only reflects one part of developmental insta-
bility, effects of stress on the neural system might just as
well account for the development and the expression of
external asymmetries. Studies of animal behaviour have
indicated that the fractal dimension of repeated be-
haviour such as movement differs between healthy
and sick individuals (Esco
´
s, Alados & Emlen, 1995).
Thus, the ability to repeat behaviour in a consistent way
may provide important information about condition
and represent a behavioural equivalent of morpho-
logical developmental instability (Møller, 1998). The
same also holds for faces: on symmetrical faces for in-
stance it could be easier to detect and see emotions.
Moreover, asymmetric movement in facial expression is
one indicator of deception (Ekman & Friesen, 1969).
Thus facial and bodily symmetry are not only indi-
cators of developmental stability, they might also
guarantee honesty (even if this is not functionally
related to it).
In recent years, although there is still controversy
about the findings, some studies have shown that female
movement might be related to sex-hormone profiles.
Self-reports of Olympic women athletes and other
sportswomen indicate a pre-menstrual decline in ath-
letic performance. Hampson & Kimura (1988) showed
that there is a significantly better mid-luteal than
menstrual performance on several measures of manual
speed and co-ordination. Grammer, Fieder & Filova
(1997) have also shown that a neural network, which
was fed with digitally analysed movement data, was able
to discriminate between women at high and low oes-
trogen levels in relation to menstrual cycle point. Thus
we can speculate that the beauty of movement could be
related to signalling optimal oestrogen levels, but also
that oestrogen could affect asymmetry, which in turn
might affect attractiveness.
XII. THE BEAUTY OF BOUNDARIES AND
BOUNDARIES OF BEAUTY
What we find in research on attractiveness is a differ-
ential reaction to humans according to their appear-
ance, which is biologically based. Yet it remains unclear
how attractiveness itself is decoded. We have discussed
many variables which may influence the decoding of
attractiveness: form of faces and bodies, structure, skin
texture, gender prototypicality, body movements, voi-
ces, age, decoration, cosmetics, body scent, hair colour,
hair style, cultural dynamics, normative comparisons
and finally temporal dynamics. Considerable evidence
has accumulated in recent years supporting the hy-
pothesis that both facial and bodily physical attract-
iveness are health certifications and thus represent
honest signals of phenotypic and genetic quality. The
hypothesis that beauty connotes health was first pro-
posed by Westermarck (1921) and later by Ellis (1926)
and Symons (1979, 1995). There is no doubt, regarding
all these variables, that beauty or attractiveness, are
cognitive constructs in the eye of the beholder. Thus
beauty, its signal values and cognitive processes are
interlinked to a high degree.
An approach to solve the problem of integration of
many features of signals into one meaning was suggested
by Schleidt & Crawley (1980) as an n-dimensional
vector approach to communication. They assumed that
meaning could be encoded in the form of pulse rate
modulation. Here the sender sends a signal of uniform
height and duration repeatedly at distinct intervals. The
receiver then applies some kind of low-pass filter in
order to integrate the signals over time. The effect on
the receiver then is a slowly accumulating tonic one.
Schleidt & Crawley (1980) suggested describing be-
haviour as a change of an organism over time, which
can be done by describing the momentary states of
organisms at any point in time. This approach demands
an evaluation of features of the individual at its surface,
the orientation of the individual and its location. The
result would be data in an n-dimensional feature space,
which can include information on the internal features
of an organism, i.e. physiological state, motivation,
emotions, etc.
Grammer et al. (2001) analysed attractiveness ratings
on 92 American females and developed a list of 36
features. These features range from simple measured
traits (e.g. lip size, breast size or eye size, body height,
body mass index, waist-to-hip ratio) to digitally ana-
lysed descriptors (e.g. body colour, hair structure, skin
colour) to more sophisticated digitally analysed mea-
sures (symmetry, skin texture and averageness). This
multidimensional vector room was then transformed
Darwinian aesthetics 399
with principal component analysis to four factors
explaining 56% of the variance in the population.
The basic factors are: a body mass index factor which
is related negatively to attractiveness, a nubile factor
which is related positively, a positively related sym-
metry and skin colour factor, and a babyness-andro-
gynous factor which is related negatively to attractive-
ness. This shows that it is possible to extract basic
features.
If such a multidimensional feature space is linked to
the perception of beauty, and if beauty is an honest
signal, this feature space needs to have a specific internal
structure. Each feature could also be a prototype. In
order to reach its communicative purpose, all features
have to point in the same direction because otherwise
the receiver would be unable to decode the signal clearly
and unmistakably. Møller & Pomiankowski (1993)
come to a comparable conclusion in their analysis of
why birds have multiple sexual ornaments. Thornhill &
Grammer (1999) showed that independent ratings of
faces in Austria and the USA, body fronts with faces
covered, and backs of the same women are significantly
positively correlated, as predicted by the health certi-
fication hypothesis and the n-dimensional feature ap-
proach. The correlation between the ratings of different
pictures implies that women’s faces and bodies comprise
a single ornament of honest mate value, apparently
developed during puberty under the influence of oes-
trogens.
This is not a single piece of evidence. If we are right,
features from different communicative channels also
have to point in the same direction. In order to study
this, Rikowski & Grammer (1999) investigated whether
smell could signal general mate quality like other cues in
sexual selection. They compared ratings of attractive-
ness and measurements of bodily asymmetry with the
evaluated attractiveness of body odour from male and
female subjects. Each subject wore a T-shirt on three
consecutive nights under controlled conditions. Im-
mediately after use the T-shirts were deep frozen and
heated up to 37 xC just before the evaluation of odour,
then 15 subjects of the opposite sex rated the smell of
each T-shirt on intensity, pleasantness and sexiness on a
seven-point scale.
Another 22 men and women evaluated portraits of
the subjects on attractiveness. For the assessment of
bodily asymmetry seven bilateral traits of each subject’s
body were measured. The results showed significant
positive correlations between facial attractiveness and
sexiness of body odour for female subjects. Moreover,
the more symmetric the body of a woman, the more
sexy her smell. Men rated the smell of women as
more erotic, the more attractive their faces had been
evaluated. Positive relations were found between body
odour and attractiveness for males only when female
odour raters were in their most fertile phase of their
menstrual cycle. In other words, these fertile women
tend to prefer the odour of physically attractive and
symmetric men. As mentioned above, the finding that
symmetric men’s scent is attractive to fertile women has
been reported in four separate studies.
One point we want to emphasize is that every theory
in this field has to take the construction of the cognitive
apparatus into account. We have shown that the de-
coding of attractiveness depends on our abilities to
create prototypes. Humans seem to use single cues,
prototypical cues and overall constructive features of
the body. Our brain combines these n-features; their
common direction makes decoding a little easier. Now
we can also suggest a possible method for the decoding
procedure itself. If, and when, the features have the
same orientation, only their size needs to be compared.
Gigerenzer & Goldstein (1996) suggests that people use
fast and frugal algorithms, which produce the same
results as more complex decision-making algorithms in
many every-day decision-making problems. Such a
simple algorithm would be the worst (or best) feature
approach’. This means that signal receivers simply
compare the size of the best or worst feature in an n-
dimensional feature space (regardless of the feature
content) in order to come to a decision that one person is
more attractive than another. Grammer et al. (2001)
showed that when the lowest value of the regressed
factor scores on the above mentioned four principal
component analysis factors is used as an attractive-
ness descriptor, the correlation between attractiveness
rating and factor size exceeds 0.60. Note that this
method only takes the size of the factor into account,
not the content. This also suggests that beauty per-
ception is not a positive concept it could be that it is
reversed: avoid ugliness. Yet even more simple methods
are possible. When there is no direct comparison
available a simple threshold model could be used; the
worst feature then has to be over a certain threshold
before the whole person is rated as attractive. Other
models might be parallel-distributed models of cogni-
tive processing as suggested by Haken (1996) through
synergetics, where one feature might strongly affect the
perception of other features and then create a coherent
signal value.
The main feature is that although different people
might have different templates for beauty, these tem-
plates underlie common construction principles. This
also explains the high cultural and temporal varia-
bility of beauty standards. What we know is that
mate-selection criteria play a role in attractiveness
400 Karl Grammer and others
ratings: females rate males as attractive when they are
dominant and healthy, males rate females as attractive
when they are healthy, receptive at the optimal age of
reproduction, and when they promise high-quality off-
spring. The question of what exactly is communicated is
still open. For the time being we suggest that it is mainly
gender identification: appearing as a typical male or
a typical female in a certain environment will cause
attractive ratings. This means that sexually dimorphic
traits that signal youth are valued. These traits then
form the respective prototypes for the cognitive evalu-
ation of attractiveness.
In addition, each theory of attractiveness has to take
into account that a great deal of learning is involved.
Different cultures indeed have different standards, if we
look at the content of these standards (although they
might agree on faces of a single population). The effect
of learning is that we adapt our standards to our
population and ecological niche. This changes as soon
as we look at the construction rules: features, aver-
ageness and symmetry. Thus the concept of attract-
iveness can be filled with different contents, as long as
these contents follow the given rules.
One condition posed at the beginning of this review is
the fact that attractiveness and beauty should be
linked to reproductive success. If this is so, why do
people not become more and more attractive and
beautiful? This question leads to Van Valen’s (1973)
Red-Queen-Hypothesis based on an experience Alice
had in the novel Alice Through the Looking Glass by Lewis
Caroll. In this experience, Alice proposes to race against
the red queen on the chessboard in order to become
queen herself. Unfortunately, one of the principles
behind the mirrors is that you have to be twice as
fast as you can simply in order to move and leave
your place. One such race is host–parasite co-evolution.
In this view only rare genotypes have a selective ad-
vantage in terms of survival and reproductive success.
Genotypes that are attractive today may not be so
for a long time because their increase in abundance
will cause evolution by parasites towards efficient
exploitation of this increasingly more common host
genotype.
We see that Darwin was wrong, but he was also right.
We saw that cultures and their media might change
beauty standards, but these standards are biologically
based, not their actual content but the rules which
determine these standards. If we assume that beauty
brings a certain amount of status in a society, we have
started another race behind the mirrors. This time
people will race against the media and surely also
against other people. The future of the adapted mind is
the creation of artificial people.
XIII. THE FUTURE OF THE ADAPTED MIND
A feature of the adapted mind which is rarely addressed
is the fact that adaptation is prone to exploitation, and to
what extent this exploitation occurs in the modern so-
ciety. This does not mean that we are ill- or maladapted
to the modern environment. It may simply be the case
that, for instance, new cultural developments can out-
run biological adaptations. One example is the speed of
information transfer. Many of our adaptations are
possibly tuned to slow and small information changes in
our environment, and the invention of modern media
has created a completely new situation. The literature
shows that beauty brings status and success (and nat-
urally, reproductive success, at least in the evolutionary
past). If so, artificial body enhancement, which am-
plifies beauty perception will be widespread. In fact,
almost all cultures use such measures. Yet beauty is
limited through evolution we cannot become more
beautiful. In the future, if a certain beauty enhancement
generates status, this enhancement will lose its advan-
tage when used by too many people, and new en-
hancements will be made. This is the eternal circle of
new fashion and beauty product inventions. Now this
trickle-down mechanism meets prototyping. If beauty
standards are a result of what people perceive in the
average media, exposure to media will change the
prototypes. As the media themselves will use beauty for
the status quest among different types of media, beauty
standards will automatically trickle down in the media
and then a quest for more beauty will start. As human
beauty is limited, it is plastic surgery and hormonal
treatments which come into play.
Beauty surgery, especially breast augmentation, is
increasingly common. Surveys suggest that more than
800 000 American women have breast implants. The
majority is not done for medical reasons. Surveys sug-
gest that women desiring surgical breast augmentation
are as psychologically stable as other women. They
differ from other women only in limited areas pri-
marily in their negative evaluation of their breasts and
their greater emphasis on dress and physical attract-
iveness (Shipley, O’Donnell & Bader, 1977); increase in
desire for attractiveness is the reason for plastic surgery.
Indeed in very young (pubescent) girls who have breast
surgery, the removal of asymmetries make up more
than 60% of the cases (Grolleau et al., 1999). This
correction would be expected if we take the role of
symmetry in beauty detection into account. In a study of
women who had received augmentation mammaplasty
an interesting pattern occurred (Cook et al., 1997).
Women with breast implants were more likely to drink a
greater average number of alcoholic drinks, be younger
Darwinian aesthetics 401
at first pregnancy, be younger at first birth, have a
history of terminated pregnancies, have used hair dyes,
and have had a greater lifetime number of sexual
partners than women without implants. This difference
between women with and without breast implants
suggest that breast augmentation may lead to higher
attractiveness for males, a larger number of possible
choices of high status males, and finally possibly higher
reproductive success. But as soon as this circle is started
and success comes by surgery, its use will spread and
trickle down to more and more surgery, until plastic
people emerge. But there is also a down side to this
game. When the media raise attractiveness standards by
prototyping beauty, then unreal expectations to mate
quality (beauty) will emerge. If the mean is more
beautiful than reality, no mate selection can occur on
realistic grounds leading to a high proportion of singles.
The second even more problematic development is
coupled to the emergence of relatively new diseases like
anorexia and similar conditions. Feminist approaches
(Crogan, 1999) to understanding women’s dissatis-
faction with their bodies claim that social pressure on
women causes them to strive for the slender, toned body
shape which is associated with youth, leading in its
extreme form to such diseases. Actually, this social
pressure is caused by other women who compete for
resources, and not by the beauty industry. It can be
viewed as a trickle-down phenomenon with a biological
basis, which with the help of fast information transfer
causes a beauty rat-race. However, independent of any
artificial changes we may still infer that such modi-
fications generally will not have the desired effects. The
reason is that the various dimensions of attractiveness
tend to signal the same qualities. Although external
phenotype can be modified, the message arising from
such a signal will prove to be incongruent with other
signals such as movement, vocalizations, and pher-
omones. Hence, we predict that experimentally ma-
nipulated phenotypes will prove unsatisfactory, even for
the individuals themselves because of being incon-
gruently perceived by others. There are some hints
consistent with such effects in the studies of con-
sequences of plastic surgery (e.g. Simis, Verhulst &
Koot, 2001; Muzaffar & Rohrich, 2002).
XIV. DIRECTIONS FOR FUTURE RESEARCH
Although the field of sexual selection and studies of
attractiveness in humans has made tremendous strides
forward during the last decade, there is still much that
needs to be done. We have emphasized the large
number of different categories of traits that play a role in
sexual selection but it is a salient question how these
different kinds of traits are related to each other. Do they
reflect different aspects of quality, are they providing
redundant information, or are some signals simply not
providing any information (Møller & Pomiankowski,
1993)? How are these different traits related to each
other when fully developed? What is their order of
ontogeny? What determines their development? And
how are they integrated? We have described the ap-
parent links between human beauty and parasites and
disease. We still need to know which kinds of diseases
are reflected by signals of beauty? How well do different
signals predict risks of disease and parasitism? These are
important questions of potentially great medical im-
portance. Numerous studies of animals have already
addressed how sexual selection relates to immune
function (review in Møller et al., 1999a). However, we
know virtually nothing about the links between beauty
and immunity in humans. Again, such knowledge could
potentially have important implications for medicine
and public health.
Finally, while almost all studies of animals have tied
research on sexual signals to components of fitness, that
has rarely if ever been done in humans. How does at-
tractiveness relate to reproductive success? We need
many more studies that investigate the benefits of mate
choice in terms of reproductive success measured as the
number of reproductive offspring in the next gener-
ation, and even in the grand-parental generation. Be-
fore such estimates have been obtained, conclusions still
remain extremely tentative.
XV. CONCLUSIONS
(1) Sexual selection is the study of what might be
called beauty perception and its role in mating com-
petition in plants and animals.
(2) Human obsession with beauty is no different
from similar obsessions in other organisms.
(3) Humans are sexually size dimorphic with males
on average being larger than females, females usually
limiting male reproductive success, and male resources
being a primary goal for female mate preferences. The
basic features of human beauty in faces and bodies are
symmetry, averageness, and sex-hormone markers.
These features reflect sex-prototypical design of traits,
developmental stability and immuno-handicaps and
are linked directly to optimal reproduction. The basic
processes are biological universals for humans, animals
and even plants.
(4) Single features from these three categories can be
placed in two groups. The first group gives information
402 Karl Grammer and others
about the general reproductive capabilities of an indi-
vidual, while the second group consists of transient
features which give information about the current
physiological state of an individual.
(5) Features that comprise the attractiveness com-
plex are intricate parts of an n-dimensional feature
space. This feature space is organized such that all
features point in the same direction. Attractiveness thus
follows the redundant signalling hypothesis.
(6) The features themselves are cross culturally
universal but their importance can be modified cul-
turally depending on socio-economic factors.
(7) We do not assume innate beauty detectors; we
rather propose that the brain has an innate tendency
and basic rules on how to create beauty templates,
which then are filled up during ontogeny.
XVI. ACKNOWLEDGMENTS
The authors have contributed equally to this publication.
K.G., A.P.M. and R.T. wish to thank the Mindship
Foundation Copenhagen for its kind contribution to the
symposium in 1996.
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