Sex

Organisms of many species are

specialized into male and female
varieties, each known as a sex.[1][2]
Sexual reproduction involves the
combining and mixing of genetic traits:
specialized cells known as gametes
combine to form offspring that inherit
traits from each parent. The gametes
produced by an organism define its sex:
males produce small gametes (e.g.
spermatozoa, or sperm, in animals) while
females produce large gametes (ova, or
egg cells). Individual organisms which
produce both male and female gametes
are termed hermaphroditic.[2] Gametes
can be identical in form and function
(known as isogamy), but, in many cases,
an asymmetry has evolved such that two
different types of gametes
(heterogametes) exist (known as
anisogamy).

Physical differences are often associated

with the different sexes of an organism;
these sexual dimorphisms can reflect the
different reproductive pressures the
sexes experience. For instance, mate
choice and sexual selection can
accelerate the evolution of physical
differences between the sexes.

Among humans and other mammals,

males typically carry an X and a Y
chromosome (XY), whereas females
typically carry two X chromosomes (XX),
which are a part of the XY sex-
determination system. Humans may also
be intersex. Other animals have various
sex-determination systems, such as the
ZW system in birds, the X0 system in
insects, and various environmental
systems, for example in reptiles and
crustaceans. Fungi may also have more
complex allelic mating systems, with
sexes not accurately described as male,
female, or hermaphroditic.[3]

Overview

The male gamete (sperm) fertilizing the female

gamete (ovum)

One of the basic properties of life is

reproduction, the capacity to generate
new individuals, and sex is an aspect of
this process. Life has evolved from
simple stages to more complex ones,
and so have the reproduction
mechanisms. Initially the reproduction
was a replicating process that consists in
producing new individuals that contain
the same genetic information as the
original or parent individual. This mode
of reproduction is called asexual, and it is
still used by many species, particularly
unicellular, but it is also very common in
multicellular organisms, including many
of those with sexual reproduction.[4] In
sexual reproduction, the genetic material
of the offspring comes from two
different individuals. As sexual
reproduction developed by way of a long
process of evolution, intermediates exist.
Bacteria, for instance, reproduce
asexually, but undergo a process by
which a part of the genetic material of an
individual donor is transferred to another
recipient.[5]

Disregarding intermediates, the basic

distinction between asexual and sexual
reproduction is the way in which the
genetic material is processed. Typically,
prior to an asexual division, a cell
duplicates its genetic information
content, and then divides. This process
of cell division is called mitosis. In sexual
reproduction, there are special kinds of
cells that divide without prior duplication
of its genetic material, in a process
named meiosis. The resulting cells are
called gametes, and contain only half the
genetic material of the parent cells.
These gametes are the cells that are
prepared for the sexual reproduction of
the organism.[6] Sex comprises the
arrangements that enable sexual
reproduction, and has evolved alongside
the reproduction system, starting with
similar gametes (isogamy) and
progressing to systems that have
different gamete types, such as those
involving a large female gamete (ovum)
and a small male gamete (sperm).[7]

In complex organisms, the sex organs

are the parts that are involved in the
production and exchange of gametes in
sexual reproduction. Many species, both
plants and animals, have sexual
specialization, and their populations are
divided into male and female individuals.
Conversely, there are also species in
which there is no sexual specialization,
and the same individuals both contain
masculine and feminine reproductive
organs, and they are called
hermaphrodites. This is very frequent in
plants.[8]

Evolution

Differen Differen
t forms t forms
of of
anisoga isogam
my: y:
A) A)
anisoga isogam
my of y of
motile motile
cells, B) cells, B)
oogam isogam
y (egg y of
cell and non-
sperm motile
cell), C) cells, C)
anisoga conjuga
my of tion.
non-
motile
cells
(egg
cell and
sperma
tia).
Sexual reproduction first probably
evolved about a billion years ago within
ancestral single-celled eukaryotes.[9] The
reason for the evolution of sex, and the
reason(s) it has survived to the present,
are still matters of debate. Some of the
many plausible theories include: that sex
creates variation among offspring, sex
helps in the spread of advantageous
traits, that sex helps in the removal of
disadvantageous traits, and that sex
facilitates repair of germ-line DNA.

Sexual reproduction is a process specific

to eukaryotes, organisms whose cells
contain a nucleus and mitochondria. In
addition to animals, plants, and fungi,
other eukaryotes (e.g. the malaria
parasite) also engage in sexual
reproduction. Some bacteria use
conjugation to transfer genetic material
between cells; while not the same as
sexual reproduction, this also results in
the mixture of genetic traits.

The defining characteristic of sexual

reproduction in eukaryotes is the
difference between the gametes and the
binary nature of fertilization. Multiplicity
of gamete types within a species would
still be considered a form of sexual
reproduction. However, no third gamete
type is known in multicellular plants or
animals.[10][11][12]

While the evolution of sex dates to the

prokaryote or early eukaryote stage, the
origin of chromosomal sex determination
may have been fairly early in eukaryotes
(see evolution of anisogamy). The ZW
sex-determination system is shared by
birds, some fish and some crustaceans.
XY sex determination is used by most
mammals,[13] but also some insects,[14]
and plants (Silene latifolia).[15] The X0
sex-determination is found in most
arachnids, insects such as silverfish
(Apterygota), dragonflies (Paleoptera)
and grasshoppers (Exopterygota), and
some nematodes, crustaceans, and
gastropods.[16][17]

No genes are shared between the avian

ZW and mammal XY chromosomes,[18]
and from a comparison between chicken
and human, the Z chromosome appeared
similar to the autosomal chromosome 9
in human, rather than X or Y, suggesting
that the ZW and XY sex-determination
systems do not share an origin, but that
the sex chromosomes are derived from
autosomal chromosomes of the
common ancestor of birds and
mammals. A paper from 2004 compared
the chicken Z chromosome with platypus
X chromosomes and suggested that the
two systems are related.[19]

Sexual reproduction

The life cycle of sexually reproducing organisms

cycles through haploid and diploid stages
Sexual reproduction in eukaryotes is a
process whereby organisms produce
offspring that combine genetic traits
from both parents. Chromosomes are
passed on from one generation to the
next in this process. Each cell in the
offspring has half the chromosomes of
the mother and half of the father.[20]
Genetic traits are contained within the
deoxyribonucleic acid (DNA) of
chromosomes—by combining one of
each type of chromosomes from each
parent, an organism is formed containing
a doubled set of chromosomes. This
double-chromosome stage is called
"diploid", while the single-chromosome
stage is "haploid". Diploid organisms can,
in turn, form haploid cells (gametes) that
randomly contain one of each of the
chromosome pairs, via meiosis.[21]
Meiosis also involves a stage of
chromosomal crossover, in which
regions of DNA are exchanged between
matched types of chromosomes, to form
a new pair of mixed chromosomes.
Crossing over and fertilization (the
recombining of single sets of
chromosomes to make a new diploid)
result in the new organism containing a
different set of genetic traits from either
parent.

In many organisms, the haploid stage

has been reduced to just gametes
specialized to recombine and form a new
diploid organism. In plants the diploid
organism produces haploid spores that
undergo cell division to produce
multicellular haploid organisms known
as gametophytes that produce haploid
gametes at maturity. In either case,
gametes may be externally similar,
particularly in size (isogamy), or may
have evolved an asymmetry such that the
gametes are different in size and other
aspects (anisogamy).[22] By convention,
the larger gamete (called an ovum, or
egg cell) is considered female, while the
smaller gamete (called a spermatozoon,
or sperm cell) is considered male. An
individual that produces exclusively large
gametes is female, and one that
produces exclusively small gametes is
male.[23] An individual that produces both
types of gametes is a hermaphrodite; in
some cases hermaphrodites are able to
self-fertilize and produce offspring on
their own, without a second organism.[24]

Animals
Hoverflies mating

Most sexually reproducing animals

spend their lives as diploid, with the
haploid stage reduced to single-cell
gametes.[25] The gametes of animals
have male and female forms—
spermatozoa and egg cells. These
gametes combine to form embryos
which develop into a new organism.
The male gamete, a spermatozoon
(produced in vertebrates within the
testes), is a small cell containing a single
long flagellum which propels it.[26]
Spermatozoa are extremely reduced
cells, lacking many cellular components
that would be necessary for embryonic
development. They are specialized for
motility, seeking out an egg cell and
fusing with it in a process called
fertilization.

Female gametes are egg cells (produced

in vertebrates within the ovaries), large
immobile cells that contain the nutrients
and cellular components necessary for a
developing embryo.[27] Egg cells are
often associated with other cells which
support the development of the embryo,
forming an egg. In mammals, the
fertilized embryo instead develops within
the female, receiving nutrition directly
from its mother.

Animals are usually mobile and seek out

a partner of the opposite sex for mating.
Animals which live in the water can mate
using external fertilization, where the
eggs and sperm are released into and
combine within the surrounding water.[28]
Most animals that live outside of water,
however, use internal fertilization,
transferring sperm directly into the
female to prevent the gametes from
drying up.

In most birds, both excretion and

reproduction is done through a single
posterior opening, called the cloaca—
male and female birds touch cloaca to
transfer sperm, a process called "cloacal
kissing".[29] In many other terrestrial
animals, males use specialized sex
organs to assist the transport of sperm—
these male sex organs are called
intromittent organs. In humans and other
mammals this male organ is the penis,
which enters the female reproductive
tract (called the vagina) to achieve
insemination—a process called sexual
intercourse. The penis contains a tube
through which semen (a fluid containing
sperm) travels. In female mammals the
vagina connects with the uterus, an
organ which directly supports the
development of a fertilized embryo within
(a process called gestation).

Because of their motility, animal sexual

behavior can involve coercive sex.
Traumatic insemination, for example, is
used by some insect species to
inseminate females through a wound in
the abdominal cavity—a process
detrimental to the female's health.

Plants

Flowers are the sexual organs of flowering plants,

usually containing both male and female parts.

Like animals, plants have specialized

male and female gametes.[30] Within
seed plants, male gametes are produced
by extremely reduced multicellular
gametophytes known as pollen. The
female gametes of seed plants are
contained within ovules; once fertilized
by male gametes produced by pollen
these form seeds which, like eggs,
contain the nutrients necessary for the
development of the embryonic plant.

Female (left) and male (right) cones are the sex

organs of pines and other conifers.
Many plants have flowers and these are
the sexual organs of those plants.
Flowers are usually hermaphroditic,
producing both male and female
gametes. The female parts, in the center
of a flower, are the pistils, each unit
consisting of a carpel, a style and a
stigma. One or more of these
reproductive units may be merged to
form a single compound pistil. Within the
carpels are ovules which develop into
seeds after fertilization. The male parts
of the flower are the stamens: these
consist of long filaments arranged
between the pistil and the petals that
produce pollen in anthers at their tips.
When a pollen grain lands upon the
stigma on top of a carpel's style, it
germinates to produce a pollen tube that
grows down through the tissues of the
style into the carpel, where it delivers
male gamete nuclei to fertilize an ovule
that eventually develops into a seed.

In pines and other conifers the sex

organs are conifer cones and have male
and female forms. The more familiar
female cones are typically more durable,
containing ovules within them. Male
cones are smaller and produce pollen
which is transported by wind to land in
female cones. As with flowers, seeds
form within the female cone after
pollination.

Because plants are immobile, they

depend upon passive methods for
transporting pollen grains to other plants.
Many plants, including conifers and
grasses, produce lightweight pollen
which is carried by wind to neighboring
plants. Other plants have heavier, sticky
pollen that is specialized for
transportation by animals. The plants
attract these insects or larger animals
such as humming birds and bats with
nectar-containing flowers. These animals
transport the pollen as they move to
other flowers, which also contain female
reproductive organs, resulting in
pollination.

Fungi

Mushrooms are produced as part of fungal sexual

reproduction

Most fungi reproduce sexually, having

both a haploid and diploid stage in their
life cycles. These fungi are typically
isogamous, lacking male and female
specialization: haploid fungi grow into
contact with each other and then fuse
their cells. In some of these cases, the
fusion is asymmetric, and the cell which
donates only a nucleus (and not
accompanying cellular material) could
arguably be considered "male".[31] Fungi
may also have more complex allelic
mating systems, with other sexes not
accurately described as male, female, or
hermaphroditic.[3]

Some fungi, including baker's yeast, have

mating types that create a duality similar
to male and female roles. Yeast with the
same mating type will not fuse with each
other to form diploid cells, only with
yeast carrying the other mating type.[32]

Many species of higher fungi produce

mushrooms as part of their sexual
reproduction. Within the mushroom
diploid cells are formed, later dividing
into haploid spores. The height of the
mushroom aids the dispersal of these
sexually produced offspring.

Sex determination
Sex helps the spread of advantageous traits
through recombination. The diagrams compare
evolution of allele frequency in a sexual population
(top) and an asexual population (bottom). The
vertical axis shows frequency and the horizontal
axis shows time. The alleles a/A and b/B occur at
random. The advantageous alleles A and B, arising
independently, can be rapidly combined by sexual
reproduction into the most advantageous
combination AB. Asexual reproduction takes longer
to achieve this combination, because it can only
produce AB if A arises in an individual which already
has B, or vice versa.
The most basic sexual system is one in
which all organisms are hermaphrodites,
producing both male and female
gametes. This is true of some animals
(e.g. snails) and the majority of flowering
plants.[33] In many cases, however,
specialization of sex has evolved such
that some organisms produce only male
or only female gametes. The biological
cause for an organism developing into
one sex or the other is called sex
determination. The cause may be genetic
or non-genetic. Within animals and other
organisms that have genetic sex
determination systems, the determining
factor may be the presence of a sex
chromosome or other genetic
differences. In plants also, such as the
liverwort Marchantia polymorpha and the
flowering plant genus Silene that have
sexual dimorphism (dioicy or dioicy,
respectively), sex may be determined by
sex chromosomes.[34] Non-genetic
systems may use environmental cues,
such as the temperature during early
development in crocodiles, to determine
the sex of the offspring.[35]

In the majority of species with sex

specialization, organisms are either male
(producing only male gametes) or female
(producing only female gametes).
Exceptions are common—for example,
the roundworm C. elegans has an
hermaphrodite and a male sex (a system
called androdioecy).

Sometimes an organism's development

is intermediate between male and
female, a condition called intersex.
Sometimes intersex individuals are
called "hermaphrodite"; but, unlike
biological hermaphrodites, intersex
individuals are unusual cases and are not
typically fertile in both male and female
aspects.
Genetic

Like humans and other mammals, the common fruit

fly has an XY sex-determination system.

In genetic sex-determination systems, an

organism's sex is determined by the
genome it inherits. Genetic sex-
determination usually depends on
asymmetrically inherited sex
chromosomes which carry genetic
features that influence development; sex
may be determined either by the
presence of a sex chromosome or by
how many the organism has. Genetic
sex-determination, because it is
determined by chromosome assortment,
usually results in a 1:1 ratio of male and
female offspring.

Humans and other mammals have an XY

sex-determination system: the Y
chromosome carries factors responsible
for triggering male development. The
"default sex," in the absence of a Y
chromosome, is female-like. Thus, XX
mammals are female and XY are male. In
humans, biological sex is determined by
five factors present at birth: the presence
or absence of a Y chromosome (which
alone determines the individual's genetic
sex), the type of gonads, the sex
hormones, the internal reproductive
anatomy (such as the uterus in females),
and the external genitalia.[36]

XY sex determination is found in other

organisms, including the common fruit
fly and some plants.[33] In some cases,
including in the fruit fly, it is the number
of X chromosomes that determines sex
rather than the presence of a Y
chromosome (see below).
In birds, which have a ZW sex-
determination system, the opposite is
true: the W chromosome carries factors
responsible for female development, and
default development is male.[37] In this
case ZZ individuals are male and ZW are
female. The majority of butterflies and
moths also have a ZW sex-determination
system. In both XY and ZW sex
determination systems, the sex
chromosome carrying the critical factors
is often significantly smaller, carrying
little more than the genes necessary for
triggering the development of a given
sex.[38]
Many insects use a sex determination
system based on the number of sex
chromosomes. This is called X0 sex-
determination—the 0 indicates the
absence of the sex chromosome. All
other chromosomes in these organisms
are diploid, but organisms may inherit
one or two X chromosomes. In field
crickets, for example, insects with a
single X chromosome develop as male,
while those with two develop as
female.[39] In the nematode C. elegans
most worms are self-fertilizing XX
hermaphrodites, but occasionally
abnormalities in chromosome
inheritance regularly give rise to
individuals with only one X chromosome
—these X0 individuals are fertile males
(and half their offspring are male).[40]

Other insects, including honey bees and

ants, use a haplodiploid sex-
determination system.[41] In this case,
diploid individuals are generally female,
and haploid individuals (which develop
from unfertilized eggs) are male. This
sex-determination system results in
highly biased sex ratios, as the sex of
offspring is determined by fertilization
rather than the assortment of
chromosomes during meiosis.
Nongenetic

Clownfishes are initially male; the largest fish in a

group becomes female

For many species, sex is not determined

by inherited traits, but instead by
environmental factors experienced
during development or later in life. Many
reptiles have temperature-dependent sex
determination: the temperature embryos
experience during their development
determines the sex of the organism. In
some turtles, for example, males are
produced at lower incubation
temperatures than females; this
difference in critical temperatures can be
as little as 1–2 °C.

Many fish change sex over the course of

their lifespan, a phenomenon called
sequential hermaphroditism. In
clownfish, smaller fish are male, and the
dominant and largest fish in a group
becomes female. In many wrasses the
opposite is true—most fish are initially
female and become male when they
reach a certain size. Sequential
hermaphrodites may produce both types
of gametes over the course of their
lifetime, but at any given point they are
either female or male.

In some ferns the default sex is

hermaphrodite, but ferns which grow in
soil that has previously supported
hermaphrodites are influenced by
residual hormones to instead develop as
male.[42]

Sexual dimorphism
Common pheasants are sexually dimorphic in both
size and appearance.

Many animals and some plants have

differences between the male and female
sexes in size and appearance, a
phenomenon called sexual dimorphism.
Sex differences in humans include,
generally, a larger size and more body
hair in men; women have breasts, wider
hips, and a higher body fat percentage. In
other species, the differences may be
more extreme, such as differences in
coloration or bodyweight.

Sexual dimorphisms in animals are often

associated with sexual selection—the
competition between individuals of one
sex to mate with the opposite sex.[43]
Antlers in male deer, for example, are
used in combat between males to win
reproductive access to female deer. In
many cases the male of a species is
larger than the female. Mammal species
with extreme sexual size dimorphism
tend to have highly polygynous mating
systems—presumably due to selection
for success in competition with other
males—such as the elephant seals. Other
examples demonstrate that it is the
preference of females that drive sexual
dimorphism, such as in the case of the
stalk-eyed fly.[44]

Other animals, including most insects

and many fish, have larger females. This
may be associated with the cost of
producing egg cells, which requires more
nutrition than producing sperm—larger
females are able to produce more
eggs.[45] For example, female southern
black widow spiders are typically twice
as long as the males.[46] Occasionally
this dimorphism is extreme, with males
reduced to living as parasites dependent
on the female, such as in the anglerfish.
Some plant species also exhibit
dimorphism in which the females are
significantly larger than the males, such
as in the moss Dicranum[47] and the
liverwort Sphaerocarpos.[48] There is
some evidence that, in these genera, the
dimorphism may be tied to a sex
chromosome,[48][49] or to chemical
signalling from females.[50]

In birds, males often have a more

colourful appearance and may have
features (like the long tail of male
peacocks) that would seem to put the
organism at a disadvantage (e.g. bright
colors would seem to make a bird more
visible to predators). One proposed
explanation for this is the handicap
principle.[51] This hypothesis says that, by
demonstrating he can survive with such
handicaps, the male is advertising his
genetic fitness to females—traits that will
benefit daughters as well, who will not be
encumbered with such handicaps.

See also
Sex and gender distinction
Sex assignment
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Further reading
Ainsworth, Claire (2015). (19 February
2015). "Sex redefined: The idea of two
sexes is simplistic. Biologists now think
there is a wider spectrum than that".
Nature. 518 (7539): 288–291.
doi:10.1038/518288a . PMID 25693544 .
Arnqvist, G.; Rowe, L. (2005). Sexual
conflict. Princeton University Press.
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Alberts, B.; Johnson, A.; Lewis, J.; Raff, M.;
Roberts, K.; Walter, P. (2002). Molecular
Biology of the Cell (4th ed.). New York:
Garland Science. ISBN 978-0-8153-3218-3.
Ellis, Havelock (1933). Psychology of Sex.
London: W. Heinemann Medical Books.
 N.B.: One of many books by this pioneering
authority on aspects of human sexuality.
Gilbert, S.F. (2000). Developmental Biology
(6th ed.). Sinauer Associates, Inc.
ISBN 978-0-87893-243-6.
Maynard-Smith, J. (1978). The Evolution of
Sex . Cambridge University Press.
ISBN 978-0-521-29302-0.

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