In biology, sexual reproduction is a process of combining and mixing genetic traits, often resulting in the specialization of organisms into a male or female variety, each known as a sex.[1] Sexual reproduction involves combining specialized cells (gametes) to form offspring that inherit traits from both parents. Gametes can be identical in form and function (known as isogametes), but in many cases an asymmetry has evolved such that two sex-specific types of gametes (heterogametes) exist: male gametes are small, motile, and optimized to transport their genetic information over a distance, while female gametes are large, non-motile and contain the nutrients necessary for the early development of the young organism.
An organism's sex is defined by the gametes it produces: males produce male gametes (spermatozoa, or sperm) while females produce female gametes (ova, or egg cells); individual organisms which produce both male and female gametes are termed hermaphroditic. Frequently, physical differences are associated with the different sexes of an organism; these sexual dimorphisms can reflect the different reproductive pressures the sexes experience.
It is considered that sexual reproduction in eukaryotes first appeared about a billion years ago and evolved within ancestral single-celled eukaryotes.[2] The reason for the initial 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 evolved as an adaptation for repairing damages in DNA (see Evolution of sexual reproduction).
While there are a number of theories, there are two main alternative views on the evolutionary origin and adaptive significance of sex. The first view assumes that sexual reproduction is a process specific to eukaryotes, organisms whose cells contain a nucleus and mitochondria. In addition to sex in animals, plants, and fungi, there are other eukaryotes (e.g. the malaria parasite) that also engage in sexual reproduction. On this first view, the adaptive advantage that maintains sexual reproduction (in competition with asexual modes of reproduction) is the benefit of generating genetic variation among progeny. Furthermore, on this view, sex originated in a eukaryotic lineage. The earliest eukaryotes and the bacterial ancestors from which they arose are assumed to have lacked sex. For instance, some bacteria use conjugation to transfer genetic material between cells; and while not the same as sexual reproduction, this also results in the mixture of genetic traits. The reason that bacterial conjugation is not the same as sexual reproduction is that the numerous genes necessary for conjugation are not located on the bacterial chromosome, but on small circular DNA self-replicating parasitic elements called conjugative plasmids. Thus, conjugation arises from an adaptation of parasitic DNA for its own transmission.[3]
The second alternative view on the evolutionary origin and adaptive significance of sex is that sex existed in early bacteria as the process of natural transformation, a well studied DNA exchange process still in existence in many present day bacterial species (see Transformation (genetics)). Transformation involves the transfer of DNA from a donor to a recipient bacterium. Recipient bacteria must first enter a special physiological state, termed competence, to receive donor DNA (see Natural competence). The numerous genes necessary for establishment of competence are located on the bacterial chromosome itself. Thus the process of transformation is likely beneficial to bacteria, and can be regarded as a simple form of sex. In general, competence is induced under stressful conditions, such as nutrient limitation and exposure to DNA damaging agents, as reviewed by a number of authors.[4][5][6] Sex, on this view, was present in the earliest single-celled eukaryotes because they were descended from ancestral bacteria capable of transformation. Sex was maintained as an adaptation for repairing DNA damage (see Evolution of sexual reproduction). In particular, meiosis the key stage of the sexual cycle of eukaryotes, in which genetic information derived from different individuals (parents) recombines, was likely derived from the analogous, but simpler, genetic information exchange and DNA repair process that occurs during transformation in bacteria[7][8][9][10] (and also see Meiosis, section: Origin and function of meiosis). Thus, by this view, sex appears to have evolved in bacteria as a way of repairing DNA damages induced by environmental stresses, was maintained through the prokaryote/eukaryote boundary, and continued to evolve in higher multicellular eukaryotes, in part, as a DNA repair process.
What is considered defining 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 is known in multicellular animals.[11][12][13]
While the evolution of sex itself dates to the prokaryote or early eukaryote stage, the origin of chromosomal sex determination may have been fairly early in eukaryotes. The ZW sex-determination system is shared by birds, some fish and some crustaceans. Most mammals, but also some insects (Drosophila) and plants (Ginkgo) use XY sex-determination. X0 sex-determination is found in certain insects.
No genes are shared between the avian ZW and mammal XY chromosomes,[14] 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.
Sexual reproduction in eukaryotes is a process whereby organisms form 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.[16] 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.[17] 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 others, the gametes are capable of undergoing cell division to produce multicellular haploid organisms. 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).[18] 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. 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.[19]
Animals
Hoverflies engaging in sexual intercourse
Most sexually reproducing animals spend their lives as diploid organisms, with the haploid stage reduced to single cell gametes.[20] 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 within a testicle), is a small cell containing a single long flagellum which propels it.[21] 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 within ovaries), large immobile cells that contain the nutrients and cellular components necessary for a developing embryo.[22] 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.[23] Most animals that live outside of water, however, must transfer sperm from male to female to achieve internal fertilization.
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".[24] 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.
Main article: Plant reproduction
Like animals, plants have developed specialized male and female gametes.[25] Within most familiar plants, male gametes are contained within hard coats, forming pollen. The female gametes of plants are contained within ovules; once fertilized by pollen these form seeds which, like eggs, contain the nutrients necessary for the development of the embryonic plant.
Pinus nigra cone.jpg Pine cones, immature male.jpg
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 carpels—one or more of these may be merged to form a single pistil. Within carpels are ovules which develop into seeds after fertilization. The male parts of the flower are the stamens: these long filamentous organs are arranged between the pistil and the petals and produce pollen at their tips. When a pollen grain lands upon the top of a carpel, the tissues of the plant react to transport the grain down into the carpel to merge with an ovule, eventually forming seeds.
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 insects. The plants attract these insects with nectar-containing flowers. Insects transport the pollen as they move to other flowers, which also contain female reproductive organs, resulting in pollination.
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".[26]
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.[27]
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.
An organism's sex is defined by the gametes it produces: males produce male gametes (spermatozoa, or sperm) while females produce female gametes (ova, or egg cells); individual organisms which produce both male and female gametes are termed hermaphroditic. Frequently, physical differences are associated with the different sexes of an organism; these sexual dimorphisms can reflect the different reproductive pressures the sexes experience.
It is considered that sexual reproduction in eukaryotes first appeared about a billion years ago and evolved within ancestral single-celled eukaryotes.[2] The reason for the initial 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 evolved as an adaptation for repairing damages in DNA (see Evolution of sexual reproduction).
While there are a number of theories, there are two main alternative views on the evolutionary origin and adaptive significance of sex. The first view assumes that sexual reproduction is a process specific to eukaryotes, organisms whose cells contain a nucleus and mitochondria. In addition to sex in animals, plants, and fungi, there are other eukaryotes (e.g. the malaria parasite) that also engage in sexual reproduction. On this first view, the adaptive advantage that maintains sexual reproduction (in competition with asexual modes of reproduction) is the benefit of generating genetic variation among progeny. Furthermore, on this view, sex originated in a eukaryotic lineage. The earliest eukaryotes and the bacterial ancestors from which they arose are assumed to have lacked sex. For instance, some bacteria use conjugation to transfer genetic material between cells; and while not the same as sexual reproduction, this also results in the mixture of genetic traits. The reason that bacterial conjugation is not the same as sexual reproduction is that the numerous genes necessary for conjugation are not located on the bacterial chromosome, but on small circular DNA self-replicating parasitic elements called conjugative plasmids. Thus, conjugation arises from an adaptation of parasitic DNA for its own transmission.[3]
The second alternative view on the evolutionary origin and adaptive significance of sex is that sex existed in early bacteria as the process of natural transformation, a well studied DNA exchange process still in existence in many present day bacterial species (see Transformation (genetics)). Transformation involves the transfer of DNA from a donor to a recipient bacterium. Recipient bacteria must first enter a special physiological state, termed competence, to receive donor DNA (see Natural competence). The numerous genes necessary for establishment of competence are located on the bacterial chromosome itself. Thus the process of transformation is likely beneficial to bacteria, and can be regarded as a simple form of sex. In general, competence is induced under stressful conditions, such as nutrient limitation and exposure to DNA damaging agents, as reviewed by a number of authors.[4][5][6] Sex, on this view, was present in the earliest single-celled eukaryotes because they were descended from ancestral bacteria capable of transformation. Sex was maintained as an adaptation for repairing DNA damage (see Evolution of sexual reproduction). In particular, meiosis the key stage of the sexual cycle of eukaryotes, in which genetic information derived from different individuals (parents) recombines, was likely derived from the analogous, but simpler, genetic information exchange and DNA repair process that occurs during transformation in bacteria[7][8][9][10] (and also see Meiosis, section: Origin and function of meiosis). Thus, by this view, sex appears to have evolved in bacteria as a way of repairing DNA damages induced by environmental stresses, was maintained through the prokaryote/eukaryote boundary, and continued to evolve in higher multicellular eukaryotes, in part, as a DNA repair process.
What is considered defining 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 is known in multicellular animals.[11][12][13]
While the evolution of sex itself dates to the prokaryote or early eukaryote stage, the origin of chromosomal sex determination may have been fairly early in eukaryotes. The ZW sex-determination system is shared by birds, some fish and some crustaceans. Most mammals, but also some insects (Drosophila) and plants (Ginkgo) use XY sex-determination. X0 sex-determination is found in certain insects.
No genes are shared between the avian ZW and mammal XY chromosomes,[14] 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.
Sexual reproduction in eukaryotes is a process whereby organisms form 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.[16] 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.[17] 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 others, the gametes are capable of undergoing cell division to produce multicellular haploid organisms. 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).[18] 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. 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.[19]
Animals
Hoverflies engaging in sexual intercourse
Most sexually reproducing animals spend their lives as diploid organisms, with the haploid stage reduced to single cell gametes.[20] 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 within a testicle), is a small cell containing a single long flagellum which propels it.[21] 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 within ovaries), large immobile cells that contain the nutrients and cellular components necessary for a developing embryo.[22] 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.[23] Most animals that live outside of water, however, must transfer sperm from male to female to achieve internal fertilization.
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".[24] 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.
Main article: Plant reproduction
Like animals, plants have developed specialized male and female gametes.[25] Within most familiar plants, male gametes are contained within hard coats, forming pollen. The female gametes of plants are contained within ovules; once fertilized by pollen these form seeds which, like eggs, contain the nutrients necessary for the development of the embryonic plant.
Pinus nigra cone.jpg Pine cones, immature male.jpg
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 carpels—one or more of these may be merged to form a single pistil. Within carpels are ovules which develop into seeds after fertilization. The male parts of the flower are the stamens: these long filamentous organs are arranged between the pistil and the petals and produce pollen at their tips. When a pollen grain lands upon the top of a carpel, the tissues of the plant react to transport the grain down into the carpel to merge with an ovule, eventually forming seeds.
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 insects. The plants attract these insects with nectar-containing flowers. Insects transport the pollen as they move to other flowers, which also contain female reproductive organs, resulting in pollination.
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".[26]
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.[27]
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.
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