In approximately 300 words explain the topic in your own words:
Sexual selection and mate choice
Malte Andersson1 and Leigh W. Simmons2
1Department of Zoology, University of Gothenburg, SE 405 30 Gothenburg, Sweden
2Centre for Evolutionary Biology, School of Animal Biology (M092), The University of Western Australia, Crawley 6009, WA,
Australia
The past two decades have seen extensive growth of
sexual selection research. Theoretical and empirical
work has clarified many components of pre- and
postcopulatory sexual selection, such as aggressive
competition, mate choice, sperm utilization and sexual
conflict. Genetic mechanisms of mate choice evolution
have been less amenable to empirical testing, but
molecular genetic analyses can now be used for incisive
experimentation. Here, we highlight some of the
currently debated areas in pre- and postcopulatory
sexual selection. We identify where new techniques
can help estimate the relative roles of the various
selection mechanisms that might work together in the
evolution of mating preferences and attractive traits,
and in sperm–egg interactions.
Introduction
Twenty years ago, when sexual selection was reviewed in
the first issue of TREE [1], it was a rapidly growing field.
Darwin’s idea of female preferences for male ornaments
was still controversial, although his theory had received
new support from two directions. First, empirical studies
showed that male ornaments are favoured by female
choice in some fishes and birds [1]. Second, a major
problem left open by Darwin, the reasons why females
prefer ornamented males, was clarified when genetic
models [2,3] verified the logical coherence of Fisherian
self-reinforcing coevolution of male ornaments and female
preferences (Box 1).
These results posed as many interesting questions as
they answered, and inspired much new research. In the
two decades since 1986, sexual selection theory has been
corroborated [4] and enriched with exciting new ideas and
discoveries, some of which we highlight here. We also
point to new possibilities for testing genetic mechanisms
of sexual selection in the era of functional genomics.
Genetic analyses of sexual selection by mate choice have
worked so far on a mainly top-down basis, inferring
genetic causes from phenotypic patterns, based on few-
locus genetics or quantitative genetics theory [5]. These
remain excellent tools for the analysis of preference–
display coevolution and for other purposes in sexual
selection research. However, quantitative trait locus
(QTL) identification and sequencing combined with
functional genomics now provide the opportunity for
bottom-up approaches, based on the precise
characterization of genes and their effects, from DNA
sequences via protein to phenotypic expression at the level
of the individual, with possible consequences at the
population level and above.
Evolution of mate choice
Although mate choice occurs in males and females [4], for
convenience we refer here to female choice of male traits.
As experimental evidence accumulated, mate choice
became widely recognized, but the genetic mechanisms
underlying its evolution remain the subject of debate
(Box 1). Showing how mating preferences evolve geneti-
cally is harder than showing that they exist, and the
problem is aggravated by the possibility that several
mechanisms co-occur (Box 1). Moreover, conflicts between
the sexes can add further selection pressures on pre-
ference and the preferred trait [6,7]. Opinions differ over
the relationships between Fisherian and genetic indicator
mechanisms of mate choice (e.g. [5,8–13]). Given that
there are qualitative differences between them, we think
there are good reasons to keep the distinction clear
[9,11,12].
Costs of mate choice, such as the loss of energy and
time, might prevent Fisherian self-reinforcing coevolution
of the trait and the preference for it, but need not do so; the
outcomes of models depend on the details of several
assumptions, for instance about sex linkage [10] (reviewed
in [5]). In spite of being theoretically plausible, the
Fisherian genetic mechanism has been difficult to
demonstrate empirically. There is corroborating pheno-
typic evidence (e.g. [14,15]), and this is also the case for
indicator mechanisms (e.g. reviewed in [5,11,16]). These
and other mechanisms (Box 1) are compatible and might
co-occur (e.g. [4,12]), and thus a challenging task is to
gauge their relative roles. Estimating the effects of the
Fisherian sexy son mechanism might help us to decide
whether it explains why male ornaments and displays are
often extreme, and differ more than do other traits
between closely related species, apparently evolving
rapidly and perhaps being involved in speciation
[2,17,18]. These questions have been with us since Charles
Darwin’s time and have yet to be answered satisfactorily.
Testing the evolution of trait and preference
Molecular genetic and genomic tools enable the detailed
characterization of genes and their effects (e.g. reviewed in
[19–21]). Combined with selection experiments that
quantify genetic evolution over generations, they offer
tools for detailed genetic analyses and tests of whether
Corresponding author: Andersson, M. (malte.andersson@zool.gu.se).
Available online 3 April 2006
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mating preferences and preferred traits evolve as pre-
dicted by sensory bias, Fisherian, indicator, or
other mechanisms.
There is evidence for sensory bias in an increasing
number of species,wherephylogenetic analyses have shown
that femalepreference for amaledisplay trait is alsopresent
in other related species lacking themale trait (e.g. [22–24]).
In such cases, it seems likely that the female preference
arosefirst for other reasons, suchasattractionto food (which
needs to be tested [12]), hereafter the male trait evolved,
favoured by the pre-existing female sensory bias [22].
The Fisherian mechanism is, in essence, based only on
genes that influence mating preferences and preferred
traits, and it can lead to their rapid coevolution if the most
preferred males are polygynous (e.g. [2,5,10,17]). Genetic
indicator mechanisms, however, depend on overall quality
and hence on much or most of the functional genome; they
can also work in monogamous systems where all males
obtain a mate [11,12]. These differences suggest ways in
which the two processes can be distinguished and their
relative importance estimated.
A prediction from the Fisherian mechanism is that
the expression of trait and corresponding preference
are positively correlated, and this has been found in
several species. However, such a correlation could
also arise from other mechanisms of male–female coevolu-
tion [4,8] (Box 1).
Bottom-up genetic testing
One possible approach to testing is to identify, sequence,
and characterize in functional detail a gene locus with
several alleles that influence the expression of an
attractive male trait, and to do likewise for the
corresponding female preference. Although there are
many difficulties, a combination of molecular genetic
tools can make such analyses feasible in suitable model
organisms (Box 2). Chromosomal regions hosting genetic
variation with substantial influence on the expression of
trait or preference can be identified by QTL analysis
[19,25], followed by nucleotide characterization of the
alleles at the loci. Techniques such as DNA microarrays,
which can detect expression-level polymorphism, are
useful in gene hunting for variable loci that influence
the phenotypic traits of interest (e.g. [19,21,26]), as are
candidate gene approaches [20,26].
If several alleles can be sequenced that encode for
different degrees of expression of the trait, and likewise for
the preference, genetic tests of the Fisherian process
might be possible, for example by finding out whether
alleles for trait and preference coevolve as predicted.
Selective breeding over several generations from those
males with greatest (or smallest) trait expression could
show whether and how preference alleles coevolve with
trait alleles, and vice versa. In combination with
microsatellite analysis of parentage, it might also be
possible to measure mating and fertilization success, and
to quantify over several generations the Fisherian sexy
sons advantage in relation to male trait size.
How can the genetic effects of Fisherian and indicator
processes be distinguished in such experiments? In
species where only males express the attractive trait and
do not provide parental care or other phenotypic
resources, daughters can be tested for genetic viability
effects that are correlated with the indicator trait of their
father, to find out whether it reflects a viability advantage,
which is not predicted by the Fisherian mechanism.
Maternal half-sib designs [27] can be particularly useful.
Experimental designs could also be used where Fisherian
mating advantages are prevented, enabling a male to
mate only with a single female. There are other
possibilities; for example, if an attractive male trait is
also expressed to some extent in females, although at a
cost, the added effect of intralocus sexual conflict [7] might
Box 1. Mechanisms of mate choice evolution
Several mechanisms have been put forward to explain mate choice:
(i) Direct phenotypic effects. Female preference for a male ornament
can evolve as a result of direct phenotypic benefits if the ornament
reflects the ability of the male to provide material advantages, such
as a high-quality territory, nutrition, parental care or protection.
There is considerable empirical support for this mechanism [76].
Female choice might also evolve as a result of resistance to direct
costs imposed by males [7,38].
(ii) Sensory bias. Female preference favouring a male ornament can
initially evolve under natural selection for other reasons, for instance
in the context of foraging or predator avoidance (e.g. [22,77]). Males
evolving traits that exploit this bias then become favoured by mate
choice. There is increasing phenotypic evidence that some male
ornaments initially evolved through female sensory biases
[22,77,78], but the evolution of female sensory bias itself requires
more testing [12].
(iii) Fisherian sexy sons. If there are genetic components to variance
in female preference and male trait, a female choosing a male with a
large trait bears daughters and sons that can both carry alleles for a
large trait, and for the preference for it. This genetic coupling might
lead to self-reinforcing coevolution between trait and preference
[2,3,5,10,12,17]. Direct critical testing of this mechanism is difficult,
but molecular genetics offers new possibilities (see main text).
(iv) Indicator mechanisms (‘good genes’ or ‘handicap mechanisms’)
suggest that attractive male traits reflect broad genetic quality.
Inherent in such mechanisms is the maintenance of genetic
variation, the ‘paradox of the lek’, and parasite- and pathogen-
mediated mechanisms have been suggested as potential solutions
(Box 4 [79,80]). In addition, other advantageous genes and relative
freedom from deleterious mutations might lead to high male
condition and expression of sex traits (e.g. [11,81–83]). Female
preference for such traits can provide genetic benefits to those of her
offspring that inherit favourable alleles from their father (e.g.
reviewed in [5,11,13,84], but see [85]). The resolution of the lek
paradox remains a crucial area for sexual selection research.
(v) Genetic compatibility mechanisms. As well as additive genetic
benefits reflected by indicator traits, there might be non-additive
benefits from choosing a mate with alleles that complement the
genome of the chooser [37,65–68,84]. Examples have been found for
instance in major histocompatibility complex genes, and compat-
ibility advantages might be one adaptive reason for multiple mating
by females (see main text).
The evolution of mate choice is based either on direct selection of
a preference that gives a fitness advantage (mechanisms i–ii) or on
indirect selection of a preference as it becomes genetically correlated
with directly selected traits (mechanisms iii,iv) [4,5,10,12]. In
addition, rather than favouring any particular display trait, mate
choice might evolve because it conveys non-additive genetic
benefits (mechanism v) [65,66]. These mechanisms are mutually
compatible and can occur together, rendering the evolution of
mating preferences a multiple-causation problem, and calling for
estimation of the relative roles of individual mechanisms [4]. Several
diagnostic differences among the mechanisms suggest ways in
which they can be tested by quantitative genetic analyses [5,12].
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be estimated by measuring the fitness effect of the trait in
females as well as in males.
Prospects for success in such tests seem best in model
species (Box 2), wheremuch genetic information is already
available (also see [28]). For example, in Drosophila, QTL
analyses have identified loci that cause variation in
courtship traits, such as male song and female mate
recognition [29,30]. Ecologically interesting phylogenetic
relatives of genomic model species could also be used by
taking advantage of a candidate gene approach, based on
knowledge of the model species [19,20], because gene
function is sometimes phylogenetically conserved. For
instance, information from the genome databases of
pufferfish Tetraodon nigroviridis and zebrafish Danio
rerio could be useful in searching for trait and/or
preference loci in two frequent targets of sexual and
natural selection analyses, the guppy (e.g. [23,31–35],
Box 2) and the threespined stickleback (e.g. [24,27]).
Mate choice and sexual conflict
The importance of conflict between the sexes is becoming
increasingly clear through a flood of exciting recent
research [7]. Early insights of Williams [36], Trivers [37]
and, especially Parker [6], have recently received much
support, and it now appears that sexual conflict between
males and females is the rule rather than the exception,
sometimes leading to sexually antagonistic coevolution
(e.g. [38,39]).
In terms of mate-choice evolution (Box 1), sexual
conflict can impose direct selection on the female
preference that leads to increased female resistance to
the male trait, because its consequences are negative for
females [40]. Many aspects of sexually antagonistic
coevolution, such as its co-occurrence with indirect
selection on mate choice (Box 1), are debated (e.g.
[7,9,13,38,39,41]). Combinations of genetic and phenoty-
pic approaches might help resolve these issues (reviewed
in [7]).
Mate choice or paternity choice?
One of the main advances in sexual selection theory since
Darwin was the realization by Parker [42] that selection
can continue after copulation. If a female mates with more
than one male, there will be competition among sperm of
different males to fertilize available ova. In the 35 years
since Parker’s paper, sperm competition has become a
major branch of sexual selection research [43].
Initially, the evolution of traits such as male genital
morphology and ejaculate size were attributed to sexual
selection for male engagement in sperm competition.
Traits such as mating plugs, seminal fluid peptides that
influence female reproductive physiology, and postcopula-
tory mate guarding, were interpreted as male adaptations
for the avoidance of sperm competition. Multiple mating
by females is a prerequisite for sperm competition, yet it is
only in the past decade that postcopulatory sexual
selection has been considered from the female perspective
[44,45]. Bateman [46] suggested that female fitness is
maximised by mating with one or a few males. However,
this view is now shifting rapidly to one in which females
are thought to often obtain direct phenotypic and indirect
genetic benefits from multiple mating [47]. Whereas
sperm competition is viewed as the postcopulatory
equivalent of male contest competition, the corresponding
equivalent of female choice is referred to as cryptic female
choice, because we are unable to observe directly female
Box 2. Model systems in sexual selection
The properties desirable in a model system for analysis of sexual
selection and mate choice include:
† A model species should be amenable to comprehensive study in
the natural environment, as a major goal is to understand how
sexual (and other) selection works in the wild, where species evolve
and acquire their characteristics. Genetic and other experimental
approaches, many of which are feasible only in the laboratory, can
then be interpreted against a solid background of ecology, selection
pressures and adaptations of the species in its natural
evolutionary context.
† Amodel system should involve conspicuous sexual dimorphism in
structure or behaviour, as these are common correlates of strong
sexual selection.
† Model organisms should have small body size and short
generation time, as these aremost suitable for experimental analysis
of selection and its evolutionary consequences over a sufficient
number of generations.
† Ideally the species, or a close phylogenetic relative, should already
be genetically well known, to facilitate crucial microevolutionary
tests of sexual selection.
These properties are available in several insects and fishes, such
as Drosophila fruit flies (Figure Ia, reproduced with permission from
Tracey Chapman), Cyrtodiopsis stalk-eyed flies (Figure Ib, repro-
duced with permission from Sam Cotton), threespined stickleback
Gasterosteus aculeatus (Figure Ic, reproduced with permission from
Massimo Lorenzoni) and guppy Poecilia reticulata (Figure Id,
reproduced with permission from Ann Houde). These are all
subjects of incisive research programs, as they make excellent
models for combined genetic and phenotypic analyses of evolution
by sexual and other forms of natural selection. For example,
experimental studies of life-history evolution and tradeoffs between
sexual selection and predation have been done in wild populations
of guppies (reviewed in [34,35]). Reproducing readily and with short
generation time in captivity, the guppy has also proved ideal for
microevolutionary analyses in the laboratory. Recently, it was
shown that the orange colour spots in males preferred by females
might be colour mimics of important fruit food for the guppy, a
possible example of sensory bias (Box 1) [23]. Orange coloration
also correlates with postcopulatory processes that bias paternity
towards preferred males [32]. When a genome database becomes
available for the guppy (which should be high priority), important
steps are likely to be taken towards a deeper understanding of the
sexual and other selection mechanisms that are driving genetic
microevolution in nature.
(a) (b)
(c) (d)
Figure I.
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preferences for sperm from particular males. Rather, they
manifest as biases in paternity (Box 3).
Thus, mate choice by females can be viewed as the first
step in the process of paternity choice. The incorporation
of female perspectives into postcopulatory sexual selection
has generated theoretical models for the evolution of
multiple mating by females that in some ways parallel the
mechanisms of mate choice evolution in Box 1. For
example, the ‘sexy-sperm’ hypothesis parallels the Fish-
erian sexy sons mechanism in proposing that multiply
mating females will have sons with high fertilization
success, because they inherit traits that caused high
fertilization success for their fathers. These traits can
include genital morphology, number and/or quality of
sperm, or seminal fluid composition [48].
Similar to Fisherian sexy sons, the evolutionary
potential of the sexy-sperm mechanism depends on the
genetic architecture of traits that contribute to fertiliza-
tion success [49]. Nevertheless, it has been argued that
such a Fisherian mechanismmight underlie the rapid and
divergent evolution of male genital morphology, if females
are more likely to store and use sperm from males able to
provide the appropriate genital stimulation during copu-
lation [50]. Under this scenario, male genitalia, tradition-
ally viewed as primary sexual traits, can become
postcopulatory equivalents of the peacock’s train, ques-
tioning the utility of a distinction between primary and
secondary sexual traits. There is increasing experimental
support for such a notion [51,52].
The ‘good-sperm’ hypothesis suggests that the ability of
a male to gain high fertilization success is correlated with
his underlying genetic quality, so that males successful in
sperm competition sire offspring with generally high
viability [53]. The good-sperm hypothesis in this respect
parallels indicator mechanisms for the evolution of male
sexual ornaments. Male sexual ornaments subject to
precopulatory sexual selection are often condition-depen-
dent indicators, developing in proportion to male condition
[11,16]. Perhaps directional postcopulatory sexual selec-
tion might likewise favour condition-dependent traits that
influence paternity and thereby provide indirect genetic
benefits for offspring (Box 4). There are few tests of this
‘good-sperm’ hypothesis, but the males of bulb mites [54]
and dung flies [55] that are most successful in sperm
competition sire offspring of higher fitness, and quanti-
tative genetic studies of traits thought to be important in
sperm competition suggest condition dependence in a
beetle species [56].
Interacting pre- and postcopulatory choice
Although theoretically possible, there is currently little
evidence that sperm competition and/or cryptic female
choice can amplify precopulatory mate choice. Suggestive
evidence comes from studies of guppies in which males
with more orange body colouration, a trait subject to
precopulatory female choice, have a paternity advantage
following artificial insemination of females with equal
numbers of sperm from two competing males [32]. Given
that females had not met the sperm donors in these
experiments, the most parsimonious explanation seems to
be that males of intrinsically high quality have attractive
sexual traits and also produce highly competitive sperm
Box 4. Male germ line control: an opportunity for
postcopulatory female choice?
The immunocompetence handicap hypothesis (ICHH) proposes that
elaboration of secondary sexual traits under sexual selection, and
success in sperm competition, have a common denominator in a
male tradeoff with investment in immune function [79,92]. Immune
responses to pathogens and parasites are suggested to be costly, so
that individuals subject to high levels of infection are likely to have
reduced resources available for secondary sexual display. The ICHH
is thus an indicator mechanism of sexual selection, and proposes
that females can obtain heritable resistance to disease for their
offspring from those males who can afford to invest in large sexual
displays. A postcopulatory extension of the ICHH suggests that
immune defenses also compromise the ability of a male to produce
high-quality ejaculates.
This hypothesis was originally formulated for vertebrates and
assumed that the immunosuppressive properties of testosterone are
required to counter attack by the diploid immune system of the male
on his haploid sperm cells [92]. Similar to the precopulatory version
of the ICHH, the hypothesis can be generalised if we assume a
resource allocation tradeoff between sperm production and immune
function [93]. Males that are forced to fight infection by upregulating
immune function pay a cost of reduced sperm quality [94]. Mating
with multiple males and enabling sperm competition to filter the
most successful, females might therefore produce offspring with the
resistance to disease that enabled their fathers to produce sperm of
superior quality. There is increasing evidence that males face a
phenotypic tradeoff between investment in secondary sexual traits
and immune function [95–97]. A recent quantitative genetic analysis
in crickets provides evidence for a genetic tradeoff between sperm
quality and immune function, as required by the postcopulatory
extension of the ICHH [98].
Box 3. Problems in demonstrating cryptic female choice
Females could bias paternity toward preferred mates by a range of
potential mechanisms [45], the easiest of which to demonstrate
occur during copulation. Females of some species expel sperm
following copulations with some males yet accept sperm from
others, biasing paternity (e.g. [43,86]). More often, only the numbers
of sperm found in storage after single matings, or patterns of
paternity following multiple matings, can be determined. The sperm
stored might reflect biases of cryptic female choice imposed during
the travel of sperm within her reproductive tract [43,45]. The
numbers of sperm might also reflect adaptive male responses to
the reproductive value of the female (i.e. cryptic male choice [87]). As
regards relative paternity among several males, it is equally difficult
to distinguish between male and female processes [43,88] and to
show unequivocally whether paternity reflects sexual selection at
the gametic level.
An increasing number of studies have found patterns of paternity
in offspring that can be interpreted as selective fertilization by
genetically superior (e.g. [32,89]) or compatible sperm haplotypes,
either among sperm from different [90] or single males [91]. Such
studies typically assign paternity using DNA or morphological
markers scored from newly born or adult offspring. The patterns of
paternity could reflect the outcome of interactions between sperm
and egg, male success in sperm competition, or preferential use of
particular sperm haplotypes by females. However, this interpretation
can be confounded by viability consequences of genetic incompat-
ibilities, or by intrinsic sire effects on offspring viability [47]. To
demonstrate unequivocally that patterns of paternity reflect sperm
competition or cryptic female choice, paternity should be deter-
mined at a very early stage, before the onset of embryo mortality.
Nevertheless, recent work on free-spawning invertebrates provides
good evidence that sperm–egg interactions can generate selective
fertilization [47].
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(but see Box 3). Later behavioural experiments found that
females accept more sperm from attractive males [33].
Both sperm competition and cryptic female choice might
amplify precopulatory sexual selection in guppies.
Measurement of the relative sexual selection intensities
during pre- and postcopulatry phases of paternity choice
seems a worthwhile endeavour, yet has rarely been
attempted [57,58].
Molecular genetic studies of parentage have revolutio-
nised our views of avian mating systems, and multiple
mating by avian females is known to result in widespread
and highly variable rates of extra-pair paternity (EPP)
[59]. In several species, EPP is most common in broods of
females whose social mate is relatively unattractive and/
or of lower social status in precopulatory sexual selection
[59,60]. If females engage in extra-pair copulations to
trade up from their social mate and maximize offspring
fitness, we might expect pre- and postcopulatory mate
choice to work synergistically in favouring the evolution of
male sexual ornaments. Comparative evidence suggests
that sexual dichromatism correlates with rates of EPP
across bird species, particularly with respect to structural
colours used in sexual display [61]. But this relationship
can also be explained by females timing their copulation
behaviour to ensure that offspring are sired by extra-pair
males. Thus, patterns of EPP might reflect precopulatory
choice of extra-pair males rather than postcopulatory
mate choice. An exciting new avenue for research on EPP
in birds would be to explore correlations between male
display traits and the fertilization capacity of their sperm
in controlled experiments, to determine the relative
importance of extra-pair copulations compared with
postcopulatory processes in determining EPP. Artificial
insemination techniques offer a promising approach.
Postcopulatory sexual selection can attenuate precopu-
latory sexual selection. It is now recognized that, counter
to Bateman’s principle [46], the reproductive success of a
male is sometimes limited by the number of sperm that he
can produce. Males who are successful in precopulatory
sexual selection can have higher costs of sperm production
and become sperm depleted, so that female fertility is
sperm limited, and successful males can suffer reduced
competitive fertilization success [62–64]. Moreover, one of
the putative genetic benefits of postcopulatory female
choice is the avoidance of genetic incompatibility between
maternal and paternal haplotypes, so that the preferred
sire of a female depends on her own genotype [65,66].
Thus, although attractive males might gain more copula-
tions, their paternity can be reduced among females with
whom their genes are incompatible. As a result, the
strength of sexual selection on male ornamental traits can
be reduced [67]. Integrating the effects of precopulatory
female choice of males carrying good genes with post-
copulatory choice of compatible genes is a new challenge in
mate choice evolution [68].
Postcopulatory sexual conflict
Despite an early focus on male perspectives in sperm
competition, Parker [42] recognized that females are not
inert environments in which male adaptations to sperm
competition arise. Traits such as male genital morphology,
or accessory gland products that enhance male fitness in
sperm competition, can impose costs on female fitness [69]
and generate antagonistic coevolution between the sexes
(e.g. [7,38,39]). Molecular genetic and genomic tools are
now being used to clarify the proximate mechanisms of
potentially strong postcopulatory sexual selection and
conflict, for example by identifying gene loci and proteins
of Drosophila seminal fluids. These proteins induce
changes in the female that are beneficial to the male,
such as increased egg laying and decreased receptivity to
other males. Some of the substances are, however,
harmful to and reduce life span in females, and so are
generating selection via sexual conflict (reviewed in
[7,70]). Nonetheless, such harmful male adaptations
might also evolve via a Fisherian sexy sons process, if
the net fitness benefit to the female from producing
harmful sons outweigh the costs of harm to her (e.g. [41]).
Many male and female reproductive proteins involved
in sperm–egg recognition and fertilization evolve rapidly,
and differ strongly between closely related taxa (reviewed
in [7,18,71]). For example, among 987 genes expressed at
different times during mouse spermatogenesis, protein
evolution was fastest for testis-specific genes expressed
during late stages of sperm maturation and involved in
sperm–egg interactions [72]. Although the mechanisms
are not yet known, many of these genes show evidence of
positive selection [72], as expected if there is sexual
selection leading to rapid coevolution of male and female
reproductive proteins [7,72]. Genetically engineered lines
of Drosophila enable the estimation of the relative
strengths of selection via antagonistic coevolution and
indirect genetic benefits to females through their sons
[73–75]. These studies illustrate how combinations of
behavioural, biochemical and genetical experiments can
produce further insights into sexual selection and conflict
and their potential roles in speciation [18,40].
Conclusions
Twenty years on we no longer think of sexual selection
simply acting on male sexual ornaments. Traits as diverse
as behavioural and structural displays, genital mor-
phology and reproductive proteins can be subject to a
multitude of selection pressures imposed by mate choice.
The genetic mechanisms of pre- and postcopulatory sexual
selection are still far from being fully understood, but
molecular genetic and genomic tools enable their detailed
experimental testing. This goal is likely to be achieved
first in model organisms with sequenced genomes, or in
some of their phylogenetic relatives. Progress will not be
quick and easy, but we would not be surprised if the 40th
anniversary issue of TREE is able to review detailed
genetic clarification of Fisherian, indicator, mate compat-
ibility, sperm competition, cryptic female choice, sexual
conflict and other mechanisms of mate choice evolution.
We thus look forward to the future of sexual selection with
much curiosity and expectation.
Acknowledgements
We thank R. Marshall and the referees for constructive comments on the
article, and the Swedish Research Council, the Swedish Foundation for
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International Cooperation in Research and Higher Education (M.A.) and
the Australian Research Council (L.W.S.) for financial support.
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Endeavour
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Female scientists in films by B.A. Jones
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