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Evolution: Mitosis Meiosis Gametes

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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
Main article: Evolution of sexual reproduction
Different forms of anisogamy:
A) anisogamy of motile cells, B) oogamy (egg cell and sperm cell), C) anisogamy of non-motile cells (egg cell
and spermatia).

Different forms of isogamy:


A) isogamy of motile cells, B) isogamy of non-motile cells, C) conjugation.
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, [citation needed] 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
Main article: Sexual reproduction
Further information: Isogamy and Anisogamy

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
Main article: Sexual reproduction in 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
Main article: Plant reproduction
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
Main article: Mating in 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.[citation needed]

Sex determination
Main article: Sex-determination system
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.[citation needed] 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.[vague] 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


Main article: Environmental sex determination
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]

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