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Sex is a trait that determines an individual's reproductive function, male or female, in animals and plants that propagate their species through sexual reproduction. [1] [2] The type of gametes produced by an organism defines its sex. Commonly in plants and animals, male organisms produce smaller gametes (spermatozoa, sperm) while female organisms produce larger gametes (ova, often called egg cells). [3] [4] Organisms that produce both types of gametes are called hermaphrodites. [2] [5] During sexual reproduction, male and female gametes fuse to form zygotes that develop into offspring that inherit a selection of the traits of each parent.


Male and female individuals of a species may be similar, or have physical differences (sexual dimorphism). The differences 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.

The terms "male" and "female" typically do not apply in sexually undifferentiated species in which the individuals are isomorphic and the gametes are isogamous (indistinguishable in size and morphology), such as the green alga Ulva lactuca . If there are instead functional differences between gametes, such as with fungi, [6] they may be referred to as mating types. [7]

Sex is genetically determined in most mammals by the XY sex-determination system, where male mammals carry an X and a Y chromosome (XY), whereas female mammals carry two X chromosomes (XX). Other chromosomal sex-determination systems in animals include the ZW system in birds, and the X0 system in insects. Various environmental systems include temperature-dependent sex determination in reptiles and crustaceans. [8]

Evolution of sex

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.

Gametes may be externally similar (isogamy), or may differ in size and other aspects (anisogamy). [7] Oogamy is an extreme example of anisogamy, in which a large, non-motile gamete is fused with a smaller, usually motile one. [9]

Anisogamy most likely evolved from isogamy, but its evolution has left no fossil evidence. [10] Its evolution was due to disruptive selection leading to two gamete sizes. [11] In anisogamous species an intermediate gamete is unable to persist [12] due to disruptive selection. [13] When oogamy has evolved males and females typically differ in many ways.[ clarification needed ] [14] The evolution of anisogamy is viewed as corresponding to the origin of male and female. [15]

Evolution of sex determination

Chromosomal sex determination may have evolved early in the history of eukaryotes. [13] No genes are shared between the avian ZW and mammal XY chromosomes [16] and the chicken Z chromosome is similar to the human autosomal chromosome 9, rather than X or Y. This suggests not that the ZW and XY sex-determination systems share an origin but that the sex chromosomes are derived from autosomal chromosomes of the common ancestor of birds and mammals. In the platypus, a monotreme, the X1 chromosome shares homology with therian mammals, while the X5 chromosome holds an avian sex-determination gene, further suggesting an evolutionary link. [17]

Breeding systems


Approximately 95% of animal species are gonochoric (also known as dioecious), [18] with about 5% of animals being hermaphroditic. However, this is due to hermaphroditism being absent in insects. [19] Hermaphroditism occurs in 70% of animal phyla. [20]

In gonochoric species, individuals are either male or female throughout their lives. [21] Gonochorism is very common in vertebrate species, with 99% being gonochoric; the other 1% is hermaphroditic, with almost all of them being fishes. [22] All birds and mammals are gonochoric. [23]


Roughly 5 to 6% of flowering plants are dioecious, resulting from between 871 and 5000 independent origins. [24] Consequently the majority are bisexual, [20] either hermaphrodite (with both stamens and pistil in the same flower) or monoecious (with separate male and female flowers on the same plant). [25] [26] In dioecious species male and female sexes are on separate plants. [27] Dioecy is common in gymnosperm species with 65% being dioecious, but most conifers are monoecious. [28]

Mixed breeding systems

Androdioecy, gynodioecy, and trioecy are sometimes called mixed breeding systems. [29] The roundworm Caenorhabditis elegans has a hermaphrodite and a male sex - a system called androdioecy. [30] The flowering plant Salvia pratensis is gynodioecious, where a species has females and hermaphrodites. [31] Although rare, a species can have males, females, and hermaphrodites - a system called trioecy. [32] Trioecy occurs in about 3.6% of flowering plants, such as Pachycereus pringlei and Fraxinus excelsior . [33]

Sexual reproduction

The life cycle of sexually reproducing organisms cycles through haploid and diploid stages Sexual cycle.svg
The life cycle of sexually reproducing organisms cycles through haploid and diploid stages

Sexual reproduction in eukaryotes produces offspring that inherit a selection of the genetic traits from both parents. In this process, chromosomes are passed on from one generation to the next. Each of the offspring's cells has half the chromosomes of the mother and half of the father. [34] The codes for genetic traits are contained within the deoxyribonucleic acid (DNA) of chromosomes. By combining one set of chromosomes from each parent, an organism is formed containing a double 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. [35] Meiosis also involves a stage of chromosomal crossover in which regions of DNA are exchanged between matched types of chromosomes, to form new pairs of mixed chromosomes, each of which is a blend of the genes of both parents. This process is followed by a mitotic division, producing haploid gametes that contain one set of chromosomes. Crossing over to make new recombinant chromosomes and fertilization (the fusion of two gametes) [36] result in the new organism containing a different set of genetic traits from either parent.

In the life cycle of most multicellular organisms, there is no multicellular haploid phase and the gametes are the only haploid cells, specialized to recombine to form a diploid zygote that develops into a new multicellular diploid organism.

In the life-cycle of plants and algae, diploid and haploid multicellular phases alternate. The diploid organism is called the sporophyte because it produces haploid spores by meiosis, which, on germination, undergo mitotic cell division to produce multicellular haploid organisms, the gametophytes. [37]

Isogamy is very common in unicellular organisms while anisogamy is common in multicellular organisms. [15] An individual that produces exclusively large gametes is female, and one that produces exclusively small gametes is male. [38] [4] [39]

An individual that produces both types of gametes is a hermaphrodite. [5] Some hermaphrodites, particularly hermaphroditic plants, are able to self-fertilize and produce offspring on their own, without a second organism. [40] However, some hermaphrodite animals such as Helix pomatia and Cepaea cannot self-fertilize. [41]


Hoverflies mating Hoverflies mating midair.jpg
Hoverflies mating

Sexually reproducing animals are diploid, and their single-celled gametes are the only haploid cells in their life cycles. [42] Animals have two gamete types: male spermatozoa (sperm) and female ova (egg cells). [43] During fertilization, the gametes combine to form diploid zygotes that develop into embryos, which in turn develop into new organisms. Animals are usually mobile and seek out a partner of the opposite sex for mating.

A spermatozoon, produced in vertebrates within the testes, is a small cell containing a single long flagellum which propels it. [44] Spermatozoa are extremely reduced cells, lacking many cellular components that would be necessary for embryonic development. They are specialized for motility, seeking out and fertilizing an egg cell.

Egg cells (ova) are produced within the ovaries. They are large, immobile cells that contain the nutrients necessary for a developing embryo. [45] Egg cells are often associated with other cells which support the development of the embryo, forming an organic vessel called an egg.

All animals that live outside of water use internal fertilization to transfer sperm directly into the female, thereby preventing the gametes from drying up. [46] Specialized organs called intromittent organs are commonly used by males to assist the transport of sperm.[ citation needed ]


With the exception of monotremes, mammals are viviparous, where the fertilized egg develops into an embryo within the female, receiving nutrition directly from its mother. [47] In mammals the female reproductive tract, called the vagina, connects with the uterus, an organ which directly supports the development of a fertilized embryo within, a process called gestation. In humans and other mammals the equivalent male organ is the penis, which enters the vagina to achieve insemination in a process called sexual intercourse. The penis contains a tube through which semen (a fluid containing sperm) travels.


In 97% of bird species, males do not have a penis. [48] Instead in most birds, both excretion and reproduction are done through a single posterior opening called the cloaca. Male and female birds touch cloacae to transfer sperm, a process called "cloacal kissing". [49]

Aquatic animals

Most aquatic animals such as fish and corals mate using external fertilization, where the eggs and sperm are released into, and combine within, the surrounding water. [50] However, some species like crustaceans use internal fertilization. [46] In seahorses, females use their ovipositors to deliver eggs into the males’ underside for fertilization and gestation. Pipefish and seahorses are the only species that entail male pregnancy. [51]


Most insects reproduce through oviparity, where a female mates with a male and the females lays the egg outside of her body. [52] A few groups of insects such as the Strepsiptera reproduce through traumatic insemination, where a male pierces a female's exoskeleton with his aedeagus. [53] In some harvester ants, a queen needs to mate with two types of males: one to reproduce queens and another to reproduce worker ants. Some biologists say harvester ants could be deemed to have three or four sexes. [54]


In the green seaweed genus Ulva , there is no sexual specialization among the isomorphic individual plants, their sexual organs, or their isogamous gametes. [55] However, the majority of plants and animals have specialized male and female gametes. [56] [14]

The male gametes are the only cells in plants and green algae that contain flagella. They are motile, able to swim to the egg cells of female gametophyte plants in films of water. Seed plants other than Cycads and Ginkgo have lost flagella entirely. Once their pollen is delivered to the stigma of flowering plants, or the micropyle of gymnosperm ovules, their gametes are delivered to the egg cell by means of pollen tubes produced by one of the cells of the microgametophyte. Many plants, including conifers and grasses, are anemophilous producing lightweight pollen which is carried by wind to neighboring plants. Other plants, such as orchids, [57] have heavier, sticky pollen that is specialized for zoophily, transportation by animals. Plants attract 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.


In seed plants, male gametes are produced by extremely reduced multicellular microgametophytes known as pollen. The female gametes (egg cells) of seed plants are produced by larger megagametophytes contained within ovules. Once the egg cells are fertilized by male gametes produced by pollen, the ovules develop into seeds which contain the nutrients necessary for the initial development of the embryonic plant. [58] :175

Pinus nigra cone.jpg
Pine cones, immature male.jpg
Female (left) and male (right) cones contain the sex organs of pines and other conifers. Most conifers are monoecious, [28] producing separate male and female cones on the same plant.

In pines and other conifers, the sex organs are contained in the cones. The female cones (seed cones) produce seeds and male cones (pollen cones) produce pollen. [59] The female cones are longer lived and typically much larger and more durable. The ovules attached to the cone scales are not enclosed in an ovary, giving rise to the name gymnosperm meaning 'naked seed'. The smaller male cones produce pollen which is transported by wind to land in female cones. Naked seeds form after pollination, protected by the scales of the female cone. [60]

Flowers contain the sexual organs of flowering plants. They usually contain both male and female parts, organs to attract pollinators and organs that provide rewards to pollinators. Mature flower diagram.svg
Flowers contain the sexual organs of flowering plants. They usually contain both male and female parts, organs to attract pollinators and organs that provide rewards to pollinators.

The flowers in flowering plants contain their sexual organs. The majority of them are hermaphroditic and produce both male and female gametes on the same plant, most often from the same flowers. [19]

Bisexual flowers that contain both male and female sexual organs are said to be perfect. [61] [25] Angiosperms may also have imperfect flowers that lack one or other type of sex organs. Sometimes, as in the tree of heaven, Ailanthus altissima the panicles can contain a mixture of functionally unisexual flowers and functionally bisexual flowers. [62]

The female parts in the flower, are the pistils, composed of one or more carpels. Carpels consist of an ovary, a style and a stigma. The male parts of the flower are the stamens, which consist of the filaments supporting the anthers that produce the pollen. [63] [64]

Within the angiosperm ovary are ovules, which contain haploid megagametophytes that produce egg cells. 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 the egg cell in an ovule that eventually develops into a seed. At the same time the ovary develops into a fruit. [65] Because flowering plants are immobile, they evolved flowers to attract animals to help in fertilization. [66]


Mushrooms are produced as part of fungal sexual reproduction Shiitake mushroom.jpg
Mushrooms are produced as part of fungal sexual reproduction

Most fungi are able to reproduce sexually and asexually and have both haploid and diploid stages in their life cycles. [6] :214 Many fungi are typically isogamous, lacking male and female specialization. [67] Even fungi that are anisogamous are all hermaphroditic, which is why even anisogamous fungi are considered to be mating types rather than sexes. [68] There has been a long debate if sex or mating type should be applied to fungi, on the grounds that mating types lack differences.[ clarification needed ] However, following studies on Phycomyces blakesleeanus there may be some reconsideration of this.[ vague ] [69] :182

Fungi may have complex allelic mating systems and many species of fungi have two mating types. [70] However, Coprinellus disseminatus has been estimated to have about 123 mating types, and in some species there are thousands of mating types. [67] For example, Schizophyllum commune has about 28,000 or more mating types. [71]

Some fungi, including that used as baker's yeast, have mating types that create a duality similar to male and female roles.[ citation needed ] Yeast with the same mating type do not fuse to form diploid cells, only with yeast carrying another mating type. [72]

Many species of higher fungi [ clarification needed ] produce mushrooms as part of their sexual reproduction. Within the mushroom diploid cells are formed, later dividing into haploid spores.


Sexual reproduction is common among parasitic protozoa but rare among free-living protozoa, which usually reproduce asexually unless food is scarce or the environment changes drastically. Both anisogamy and isogamy are found in free-living protoza. [73] Ciliates are all isogamous such as Tetrahymena thermophila , which has 7 mating types. [74]

Sex determination systems

Sex helps the spread of advantageous traits through recombination. The diagrams compare the 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. Evolsex-dia2a.svg
Sex helps the spread of advantageous traits through recombination. The diagrams compare the 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 biological cause of an organism developing into one sex or the other is called sex determination. The cause may be genetic, environmental, haplodiploidy, or multiple factors. [19] Within animals and other organisms that have genetic sex determination systems, the determining factor may be the presence of a sex chromosome. In plants that are sexually dimorphic, such as the liverwort Marchantia polymorpha or the dioecious species in the flowering plant genus Silene , sex may also be determined by sex chromosomes. [75] Non-genetic systems may use environmental cues, such as the temperature during early development in crocodiles, to determine the sex of the offspring. [76]

Sex determination is often distinct from sex differentiation, sex determination is the designation for the development stage towards either male or female while sex differentiation is the pathway towards the development of the phenotype. [77]


Like humans and most other mammals, the common fruit fly has an XY sex-determination system. Drosophila XY sex-determination.svg
Like humans and most 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 carrying genetic features that influence development.[ citation needed ][ vague ] Sex may be determined either by the presence of a sex chromosome or by the number of sex chromosomes 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.[ citation needed ]

XY sex determination

Humans and most other mammals have an XY sex-determination system: the Y chromosome carries factors responsible for triggering male development, making XY sex determination mostly based on the presence or absence of the Y chromosome. It is the male gamete that determines the sex of the offspring. [78] In this system XX mammals typically are female and XY typically are male. [19] However, individuals with XXY or XYY are males, while individuals with X and XXX are females. [8]

XY sex determination is found in other organisms, including insects like the common fruit fly, [79] and some plants. [80] In some cases, it is the number of X chromosomes that determines sex rather than the presence of a Y chromosome. [8] In the fruit fly individuals with XY are male and individuals with XX are female; however, individuals with XXY or XXX can also be female, and individuals with X can be males. [81]

ZW sex determination

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. [82] In this case, ZZ individuals are male and ZW are female. It is the female gamete that determines the sex of the offspring. This system is used by birds, some fish, and some crustaceans. [8]

The majority of butterflies and moths also have a ZW sex-determination system. In groups like the Lepidoptera, females can have Z, ZZW, and even ZZWW. [83]

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. [84] [ better source needed ]

XO sex determination

In the X0 sex-determination system, males have one X chromosome (X0) while females have two (XX). All other chromosomes in these diploid organisms are paired, but organisms may inherit one or two X chromosomes. This system is found in most arachnids, insects such as silverfish (Apterygota), dragonflies (Paleoptera) and grasshoppers (Exopterygota), and some nematodes, crustaceans, and gastropods. [85] [86]

In field crickets, for example, insects with a single X chromosome develop as male, while those with two develop as female. [87]

In the nematode Caenorhabditis elegans , most worms are self-fertilizing hermaphrodites with an XX karyotype, but occasional abnormalities in chromosome inheritance can give rise to individuals with only one X chromosome—these X0 individuals are fertile males (and half their offspring are male). [88]

ZO sex determination

In the Z0 sex-determination system, males have two Z chromosomes whereas females have one. This system is found in several species of moths. [89]


For many species, sex is not determined by inherited traits, but instead by environmental factors such as temperature experienced during development or later in life.

The bonelliidae larvae can only develop as males when they encounter a female. [19]

In the fern Ceratopteris and other homosporous fern species, the default sex is hermaphrodite, but individuals which grow in soil that has previously supported hermaphrodites are influenced by the pheromone antheridiogen to develop as male. [90]

Sequential hermaphroditism

Clownfishes are initially male; the largest fish in a group becomes female Ocellaris clownfish.JPG
Clownfishes are initially male; the largest fish in a group becomes female

Some species can change sex over the course of their lifespan, a phenomenon called sequential hermaphroditism. [91] Teleost fishes are the only vertebrate lineage where sequential hermaphroditism occurs. In clownfish, smaller fish are male, and the dominant and largest fish in a group becomes female; when a dominant female is absent, then her partner changes sex. In many wrasses the opposite is true—the fish are initially female and become male when they reach a certain size. [92] Sequential hermaphroditism also occurs in plants such as Arisaema triphyllum .

Temperature-dependent sex determination

Many reptiles, including all crocodiles and most turtles, have temperature-dependent sex determination. In these species, the temperature experienced by the embryos during their development determines their sex. [19] In some turtles, for example, males are produced at lower temperatures than females; but Macroclemys females are produced at temperatures lower than 22 °C or above 28 °C, while males are produced in between those temperatures. [93]


Other insects, including honey bees and ants, use a haplodiploid sex-determination system. [94] Diploid bees and ants 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 (arrhenotoky or pseudo-arrhenotoky resulting in males) rather than the assortment of chromosomes during meiosis. [95]

Sex ratio

Most organisms which reproduce sexually have a 1:1 sex ratio of male and female births. The English statistician and biologist Ronald Fisher outlined why this is so in what has come to be known as Fisher's principle. [96] This essentially says the following:

  1. Suppose male births are less common than female.
  2. A newborn male then has better mating prospects than a newborn female, and therefore can expect to have more offspring.
  3. Therefore parents genetically disposed to produce males tend to have more than average numbers of grandchildren born to them.
  4. Therefore the genes for male-producing tendencies spread, and male births become more common.
  5. As the 1:1 sex ratio is approached, the advantage associated with producing males dies away.
  6. The same reasoning holds if females are substituted for males throughout. Therefore 1:1 is the equilibrium ratio.

Sex differences

Biologists agree that gamete size is the fundamental difference between the male and female sexes, [97] [98] this difference is also considered the first sex difference that predates many sex differences. [48] It has even been said that all the differences between the sexes stem from the differences in the gametes. [99]

Sex differences in humans include a generally larger size and more body hair in men, while women have larger breasts, wider hips, and a higher body fat percentage. In other species, there may be differences in coloration or other features, and may be so pronounced that the different sexes may be mistaken for two entirely different taxa. [100]

Sex differences in behavior

Males in many species typically invest more in finding mates and invest less in parental care. [101] Males in many species are more competitive than females for reproduction. [15] It is said that females are typically more choosy regarding who to mate with due to their large gametes. [102]

Sex characteristics

Primary sex characteristics are structures directly involved in reproduction such as the testes or ovaries, while secondary sex characteristics in humans for example are body hair, breasts, and distribution of fat. [103]

In some species, a few individuals may have a mixture of characteristics from both sexes, a condition called intersex. [104] This can be caused by extra sex chromosomes or by a hormonal abnormality during fetal development. [105] The term intersex typically applies to abnormal members of gonochoric species rather than to hermaphroditic species. [106] Some species[ which? ] can have gynandromorphs. [105]

Sexual dimorphism

Common pheasants are sexually dimorphic in both size and appearance. Male and female pheasant.jpg
Common pheasants are sexually dimorphic in both size and appearance.

In many animals and some plants, individuals of male and female sex differ in size and appearance, a phenomenon called sexual dimorphism. [30] Sexual dimorphism in animals is often associated with sexual selection—the mating competition between individuals of one sex vis-à-vis the opposite sex. [100] 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 drives sexual dimorphism, such as in the case of the stalk-eyed fly. [107]

Females are the larger sex in a majority of animals. [30] For instance, female southern black widow spiders are typically twice as long as the males. [108] This size disparity 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. [109] [30]

Sexual dimorphism can be extreme, with males, such as some anglerfish, living parasitically on the female. Some plant species also exhibit dimorphism in which the females are significantly larger than the males, such as in the moss Dicranum [110] and the liverwort Sphaerocarpos . [111] There is some evidence that, in these genera, the dimorphism may be tied to a sex chromosome, [111] [112] or to chemical signalling from females. [113]

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. [114] 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

Related Research Articles

Asexual reproduction Reproduction without a sexual process

Asexual reproduction is a type of reproduction that does not involve the fusion of gametes or change in the number of chromosomes. The offspring that arise by asexual reproduction from either unicellular or multicellular organisms inherit the full set of genes of their single parent. Asexual reproduction is the primary form of reproduction for single-celled organisms such as archaea and bacteria. Many eukaryotic organisms including plants, animals, and fungi can also reproduce asexually. In vertebrates, the most common form of asexual reproduction is parthenogenesis, which is typically used as an alternative to sexual reproduction in times when reproductive opportunities are limited.

Gamete Cell that fuses during fertilisation, such as a sperm or egg cell

A gamete is a haploid cell that fuses with another haploid cell during fertilization in organisms that reproduce sexually. Gametes are an organism's reproductive cells, also referred to as sex cells. In species that produce two morphologically distinct types of gametes, and in which each individual produces only one type, a female is any individual that produces the larger type of gamete—called an ovum— and a male produces the smaller tadpole-like type—called a sperm. Sperm cells or spermatozoon are small and motile due to the flagellum, a tail-shaped structure that allows the cell to propel and move. In contrast, each egg cell or ovum is relatively large and non-motile. In short a gamete is an egg cell or a sperm. In animals, ova mature in the ovaries of females and sperm develop in the testes of males. During fertilization, a spermatozoon and ovum unite to form a new diploid organism. Gametes carry half the genetic information of an individual, one ploidy of each type, and are created through meiosis, in which a germ cell undergoes two fissions, resulting in the production of four gametes. In biology, the type of gamete one produces determines the classification of their sex.

Gametophyte Haploid stage in the life cycle of plants and algae

A gametophyte is one of the two alternating multicellular phases in the life cycles of plants and algae. It is a haploid multicellular organism that develops from a haploid spore that has one set of chromosomes. The gametophyte is the sexual phase in the life cycle of plants and algae. It develops sex organs that produce gametes, haploid sex cells that participate in fertilization to form a diploid zygote which has a double set of chromosomes. Cell division of the zygote results in a new diploid multicellular organism, the second stage in the life cycle known as the sporophyte. The sporophyte can produce haploid spores by meiosis that on germination produce a new generation of gametophytes.

Meiosis Type of cell division in sexually-reproducing organisms used to produce gametes

Meiosis is a special type of cell division of germ cells in sexually-reproducing organisms used to produce the gametes, such as sperm or egg cells. It involves two rounds of division that ultimately result in four cells with only one copy of each chromosome (haploid). Additionally, prior to the division, genetic material from the paternal and maternal copies of each chromosome is crossed over, creating new combinations of code on each chromosome. Later on, during fertilisation, the haploid cells produced by meiosis from a male and female will fuse to create a cell with two copies of each chromosome again, the zygote.

Ploidy Number of sets of chromosomes in a cell

Ploidy is the number of complete sets of chromosomes in a cell, and hence the number of possible alleles for autosomal and pseudoautosomal genes. Somatic cells, tissues, and individual organisms can be described according to the number of sets of chromosomes present : monoploid, diploid, triploid, tetraploid, pentaploid, hexaploid, heptaploid or septaploid, etc. The generic term polyploid is often used to describe cells with three or more chromosome sets.

Reproduction Biological process by which new organisms are generated from one or more parent organisms

Reproduction is the biological process by which new individual organisms – "offspring" – are produced from their "parent" or parents. Reproduction is a fundamental feature of all known life; each individual organism exists as the result of reproduction. There are two forms of reproduction: asexual and sexual.

Fertilisation Union of gametes of opposite sexes during the process of sexual reproduction to form a zygote

Fertilisation or fertilization, also known as generative fertilisation, syngamy and impregnation, is the fusion of gametes to give rise to a new individual organism or offspring and initiate its development. Processes such as insemination or pollination which happen before the fusion of gametes are also sometimes informally called fertilization. The cycle of fertilisation and development of new individuals is called sexual reproduction. During double fertilisation in angiosperms the haploid male gamete combines with two haploid polar nuclei to form a triploid primary endosperm nucleus by the process of vegetative fertilisation.

Alternation of generations Reproductive cycle of plants and algae

Alternation of generations is the type of life cycle that occurs in those plants and algae in the Archaeplastida and the Heterokontophyta that have distinct haploid sexual and diploid asexual stages. In these groups, a multicellular haploid gametophyte with n chromosomes alternates with a multicellular diploid sporophyte with 2n chromosomes, made up of n pairs. A mature sporophyte produces haploid spores by meiosis, a process which reduces the number of chromosomes to half, from 2n to n.

Biological life cycle Life cycle of living species

In biology, a biological life cycle is a series of changes in form that an organism undergoes, returning to the starting state. "The concept is closely related to those of the life history, development and ontogeny, but differs from them in stressing renewal." Transitions of form may involve growth, asexual reproduction, or sexual reproduction.


In biology, mating is the pairing of either opposite-sex or hermaphroditic organisms for the purposes of sexual reproduction. Fertilization is the fusion of two gametes. Copulation is the union of the sex organs of two sexually reproducing animals for insemination and subsequent internal fertilization. Mating may also lead to external fertilization, as seen in amphibians, fishes and plants. For the majority of species, mating is between two individuals of opposite sexes. However, for some hermaphroditic species, copulation is not required because the parent organism is capable of self-fertilization (autogamy); for example, banana slugs.

Anisogamy is the form of sexual reproduction that involves the union or fusion of two gametes, which differ in size and/or form. The smaller gamete is male, a sperm cell, whereas the larger gamete is female, typically an egg cell. Anisogamy is common and widespread in multicellular organisms. It is thought to have evolved from isogamy. Since the biological definition of male and female is based by gamete size the evolution of anisogamy is viewed as the evolutionary origin of male and female.


Isogamy is a form of sexual reproduction that involves gametes of similar morphology, found in most unicellular organisms. Because both gametes look alike, they generally cannot be classified as male or female. Instead, organisms undergoing isogamy are said to have different mating types, most commonly noted as "+" and "−" strains. Isogamous species often have two mating types. But some isogamous species have more than two mating types, but the number is usually lower than ten, though in some extremely rare cases a species can have thousands of mating types. In all cases, fertilization occurs when gametes of two different mating types fuse to form a zygote.

Dioecy is a characteristic of a species, meaning that it has distinct individual organisms that produce male and female gametes, either directly or indirectly. Dioecious reproduction is biparental reproduction. Dioecy has costs, since only about half the population directly produces offspring. It is one method that excludes self-fertilization and promotes allogamy (outcrossing), and thus tends to reduce the expression of recessive deleterious mutations present in a population. Flowering plants have several other methods of excluding self-fertilization, such as self-incompatibility.

Male Sex of an organism which produces sperm

Male (♂) is the sex of an organism that produces the gamete known as sperm, which fuses with the larger female gamete, or ovum, in the process of fertilization. A male organism cannot reproduce sexually without access to at least one ovum from a female, but some organisms can reproduce both sexually and asexually. Most male mammals, including male humans, have a Y chromosome, which codes for the production of larger amounts of testosterone to develop male reproductive organs. Not all species share a common sex-determination system. In most animals, including humans, sex is determined genetically; however, species such as Cymothoa exigua change sex depending on the number of females present in the vicinity.

Haplodiploidy Biological system where sex is determined by the number of sets of chromosomes

Haplodiploidy is a sex-determination system in which males develop from unfertilized eggs and are haploid, and females develop from fertilized eggs and are diploid. Haplodiploidy is sometimes called arrhenotoky.

Plant reproduction is the production of new offspring in plants, which can be accomplished by sexual or asexual reproduction. Sexual reproduction produces offspring by the fusion of gametes, resulting in offspring genetically different from the parent or parents. Asexual reproduction produces new individuals without the fusion of gametes, genetically identical to the parent plants and each other, except when mutations occur.

Selfing or self-fertilization is the union of male and female gametes and/or nuclei from the same haploid, diploid, or polyploid organism. It is an extreme degree of inbreeding.

Hermaphrodite An organism that has complete or partial male and female reproductive organs

In reproductive biology, a hermaphrodite is an organism that has both kinds of reproductive organs and can produce both gametes associated with male and female sexes. Many taxonomic groups of animals do not have separate sexes. In these groups, hermaphroditism is a normal condition, enabling a form of sexual reproduction in which either partner can act as the female or male. For example, the great majority of tunicates, pulmonate snails, opisthobranch snails, earthworms, and slugs are hermaphrodites. Hermaphroditism is also found in some fish species and to a lesser degree in other vertebrates. Most plants are also hermaphrodites. A species having different sexes, male and female, is called gonochoric, which is the opposite of hermaphrodite.

Female Sex of an organism which produces ova

Female is the sex of an organism that produces non-mobile ova, which is the gamete that fuses with the male gamete during sexual reproduction. Most female mammals, including female humans, have two X chromosomes. Female characteristics vary between different species with some species having pronounced female characteristics, such as the presence of pronounced mammary glands in mammals. There is no single genetic mechanism behind sex differences in different species and the existence of two sexes seems to have evolved multiple times independently in different evolutionary lineages.

Sexual reproduction Reproduction process that creates a new organism by combining the genetic material of two organisms

Sexual reproduction is a type of reproduction that involves a complex life cycle in which a gamete with a single set of chromosomes (haploid) combines with another to produce an organism composed of cells with two sets of chromosomes (diploid). Sexual reproduction is the most common life cycle in multicellular eukaryotes, such as animals, fungi and plants. Sexual reproduction does not occur in prokaryotes, but they have processes with similar effects such as bacterial conjugation, transformation and transduction, which may have been precursors to sexual reproduction in early eukaryotes.


  1. Stevenson A, Waite M (2011). Concise Oxford English Dictionary: Book & CD-ROM Set. OUP Oxford. p. 1302. ISBN   978-0-19-960110-3. Archived from the original on 11 March 2020. Retrieved 23 March 2018. Sex: Either of the two main categories (male and female) into which humans and most other living things are divided on the basis of their reproductive functions. The fact of belonging to one of these categories. The group of all members of either sex.
  2. 1 2 Purves WK, Sadava DE, Orians GH, Heller HC (2000). Life: The Science of Biology. Macmillan. p. 736. ISBN   978-0-7167-3873-2. Archived from the original on 26 June 2019. Retrieved 23 March 2018. A single body can function as both male and female. Sexual reproduction requires both male and female haploid gametes. In most species, these gametes are produced by individuals that are either male or female. Species that have male and female members are called dioecious (from the Greek for 'two houses'). In some species, a single individual may possess both female and male reproductive systems. Such species are called monoecious ("one house") or hermaphroditic.
  3. Adkins-Regan E (18 November 2010). "Sexual behavior: conflict cooperation, and coevolution". In Székely T, Moore AJ, Komdeur J (eds.). Social Behaviour: Genes, Ecology and Evolution. Cambridge University Press. p. 231. ISBN   978-0-521-88317-7.
  4. 1 2 Royle NJ, Smiseth PT, Kölliker M (9 August 2012). Kokko H, Jennions M (eds.). The Evolution of Parental Care. Oxford University Press. p. 103. ISBN   978-0-19-969257-6. The answer is that there is an agreement by convention: individuals producing the smaller of the two gamete types-sperm or pollen- are males, and those producing larger gametes-eggs or ovules- are females.
  5. 1 2 Avise JC (18 March 2011). Hermaphroditism: A Primer on the Biology, Ecology, and Evolution of Dual Sexuality. Columbia University Press. pp. 1–7. ISBN   978-0-231-52715-6. Archived from the original on 11 October 2020. Retrieved 18 September 2020.
  6. 1 2 Moore D, Robson JD, Trinci AP (2020). 21st Century guidebook to fungi (2 ed.). Cambridge University press. pp. 211–228. ISBN   978-1-108-74568-0.
  7. 1 2 Kumar R, Meena M, Swapnil P (2019). "Anisogamy". In Vonk J, [[Todd K. Shackelford ]|Shackelford T]] (eds.). Encyclopedia of Animal Cognition and Behavior. Cham: Springer International Publishing. pp. 1–5. doi:10.1007/978-3-319-47829-6_340-1. ISBN   978-3-319-47829-6. Archived from the original on 4 November 2020. Anisogamy can be defined as a mode of sexual reproduction in which fusing gametes, formed by participating parents, are dissimilar in size.
  8. 1 2 3 4 Hake L, O'Connor C. "Genetic Mechanisms of Sex Determination | Learn Science at Scitable". Retrieved 13 April 2021.
  9. Allaby M (29 March 2012). A Dictionary of Plant Sciences. OUP Oxford. p. 350. ISBN   978-0-19-960057-1.
  10. Pitnick SS, Hosken DJ, Birkhead TR (21 November 2008). Sperm Biology: An Evolutionary Perspective. Academic Press. pp. 43–44. ISBN   978-0-08-091987-4.
  11. Low BS (4 January 2015). Why Sex Matters: A Darwinian Look at Human Behavior - Revised Edition. Princeton University Press. p. 32. ISBN   978-1-4008-5235-2. This disruptive selection leads to anisogamy (unlike gametes) and a bimodal size distribution: small gametes (sperm) and large gametes (egg).
  12. Pitnick SS, Hosken DJ, Birkhead TR (21 November 2008). Sperm Biology: An Evolutionary Perspective. Academic Press. p. 60. ISBN   978-0-08-091987-4. In anisogamous species, a third gamete size is either unable to invade or to persist, so two genders are evolutionarily stable
  13. 1 2 Lehtonen J, Parker GA (2014). "Gamete competition, gamete limitation, and the evolution of the two sexes". Molecular Human Reproduction. 20 (12): 1161–1168. doi:10.1093/molehr/gau068. PMID   25323972.
  14. 1 2 Dusenbery DB (2009). Living at Micro Scale: The Unexpected Physics of Being Small. Harvard University Press. pp. 308–326. ISBN   978-0-674-03116-6.
  15. 1 2 3 Lehtonen J, Kokko H, Parker GA (October 2016). "What do isogamous organisms teach us about sex and the two sexes?". Philosophical Transactions of the Royal Society of London. Series B, Biological Sciences. 371 (1706). doi:10.1098/rstb.2015.0532. PMC   5031617 . PMID   27619696.
  16. Stiglec R, Ezaz T, Graves JA (2007). "A new look at the evolution of avian sex chromosomes". Cytogenetic and Genome Research. 117 (1–4): 103–9. doi:10.1159/000103170. PMID   17675850. S2CID   12932564.
  17. Veyrunes F, Waters PD, Miethke P, Rens W, McMillan D, Alsop AE, et al. (June 2008). "Bird-like sex chromosomes of platypus imply recent origin of mammal sex chromosomes". Genome Research. 18 (6): 965–73. doi:10.1101/gr.7101908. PMC   2413164 . PMID   18463302.
  18. Muyle A, Bachtrog D, Marais GA, Turner JM (June 2021). "Epigenetics drive the evolution of sex chromosomes in animals and plants". Philosophical Transactions of the Royal Society of London. Series B, Biological Sciences. 376 (1826): 20200124. doi:10.1098/rstb.2020.0124. PMC  8059572. PMID   33866802. Archived from the original on June 2021.
  19. 1 2 3 4 5 6 Bachtrog D, Mank JE, Peichel CL, Kirkpatrick M, Otto SP, Ashman TL, et al. (July 2014). "Sex determination: why so many ways of doing it?". PLOS Biology. 12 (7): e1001899. doi:10.1371/journal.pbio.1001899. PMC   4077654 . PMID   24983465.
  20. 1 2 Kliman, Richard (2016). Encyclopedia of Evolutionary Biology. 2. Academic Press. pp. 212–224. ISBN   978-0-12-800426-5. Archived from the original on 2016.
  21. West S (28 September 2009). Sex Allocation. Princeton University Press. p. 1-2. ISBN   978-1-4008-3201-9.
  22. Kuwamura T, Sunobe T, Sakai Y, Kadota T, Sawada K (1 July 2020). "Hermaphroditism in fishes: an annotated list of species, phylogeny, and mating system". Ichthyological Research. 67 (3): 341–360. doi: 10.1007/s10228-020-00754-6 . ISSN   1616-3915. S2CID   218527927.
  23. Kobayashi K, Kitano T, Iwao Y, Kondo M (1 June 2018). Reproductive and Developmental Strategies: The Continuity of Life. Springer. p. 290. ISBN   978-4-431-56609-0.
  24. Renner, Susanne S. (2014). "The relative and absolute frequencies of angiosperm sexual systems: dioecy, monoecy, gynodioecy, and an updated online database". American Journal of Botany. 101 (10): 1588–1596. doi: 10.3732/ajb.1400196 . PMID   25326608.
  25. 1 2 Sabath N, Goldberg EE, Glick L, Einhorn M, Ashman TL, Ming R, et al. (February 2016). "Dioecy does not consistently accelerate or slow lineage diversification across multiple genera of angiosperms". The New Phytologist. 209 (3): 1290–300. doi:10.1111/nph.13696. PMID   26467174.
  26. Beentje H (2016). The Kew plant glossary (2 ed.). Royal Botanic Gardens, Kew: Kew Publishing. ISBN   978-1-84246-604-9.
  27. Leite Montalvão, Ana Paula; Kersten, Birgit; Fladung, Matthias; Müller, Niels Andreas (2021). "The Diversity and Dynamics of Sex Determination in Dioecious Plants". Frontiers in Plant Science. 11: 580488. doi: 10.3389/fpls.2020.580488 . ISSN   1664-462X. PMC   7843427 . PMID   33519840.
  28. 1 2 Walas Ł, Mandryk W, Thomas PA, Tyrała-Wierucka Ż, Iszkuło G (2018). "Sexual systems in gymnosperms: A review" (PDF). Basic and Applied Ecology. 31: 1–9. doi:10.1016/j.baae.2018.05.009.
  29. Minelli A, Fusco G (10 October 2019). The Biology of Reproduction. Cambridge University Press. p. 134. ISBN   978-1-108-49985-9.
  30. 1 2 3 4 Choe J (21 January 2019). "Body Size and Sexual Dimorphism". In Cox R (ed.). Encyclopedia of Animal Behavior. Volume 2. Academic Press. pp. 7–11. ISBN   978-0-12-813252-4.|volume= has extra text (help)
  31. Zhang B, Claßen-Bockhoff R (August 2019). "Sex-differential reproduction success and selection on floral traits in gynodioecious Salvia pratensis". BMC Plant Biology. 19 (1): 375. doi:10.1186/s12870-019-1972-y. PMC   6712674 . PMID   31455268.
  32. Perry LE, Pannell JR, Dorken ME (19 April 2012). "Two's company, three's a crowd: experimental evaluation of the evolutionary maintenance of trioecy in Mercurialis annua (Euphorbiaceae)". PLOS ONE. 7 (4): e35597. Bibcode:2012PLoSO...735597P. doi:10.1371/journal.pone.0035597. PMC   3330815 . PMID   22532862.
  33. Albert B, Morand-Prieur MÉ, Brachet S, Gouyon PH, Frascaria-Lacoste N, Raquin C (October 2013). "Sex expression and reproductive biology in a tree species, Fraxinus excelsior L". Comptes Rendus Biologies. 336 (10): 479–85. doi:10.1016/j.crvi.2013.08.004. PMID   24246889.
  34. Alberts B, Johnson A, Lewis J, Raff M, Roberts K, Walter P (2002). "The Benefits of Sex". Molecular Biology of the Cell (4th ed.). New York: Garland Science. ISBN   978-0-8153-3218-3.
  35. Alberts B, Johnson A, Lewis J, Raff M, Roberts K, Walter P (2002). "Meiosis". Molecular Biology of the Cell (4th ed.). New York: Garland Science. ISBN   978-0-8153-3218-3.
  36. Hine R, Martin E (2015). A Dictionary of Biology. Oxford University Press. p. 542. ISBN   978-0-19-871437-8.
  37. Gilbert SF (2000). "1.2. Multicellularity: Evolution of Differentiation". Developmental Biology (6th ed.). Sunderland (MA): Sinauer Associates. ISBN   978-0-87893-243-6.
  38. Gee H (22 November 1999). "Size and the single sex cell". Nature. Retrieved 4 June 2018.
  39. Fusco G, Minelli A (10 October 2019). The Biology of Reproduction. Cambridge University Press. pp. 111–113. ISBN   978-1-108-49985-9. Archived from the original on 1 April 2021. Retrieved 29 March 2021.
  40. Alberts B, Johnson A, Lewis J, Raff M, Roberts K, Walter P (2002). "Caenorhabditis Elegans: Development from the Perspective of the Individual Cell". Molecular Biology of the Cell (4th ed.). New York: Garland Science. ISBN   978-0-8153-3218-3.
  41. Smith JM, Smith RT (24 August 1978). The Evolution of Sex. CUP Archive. p. 125. ISBN   978-0-521-29302-0.
  42. Alberts B, Johnson A, Lewis J, Raff M, Roberts K, Walter P (2002). "Mendelian genetics in eukaryotic life cycles". Molecular Biology of the Cell (4th ed.). New York: Garland Science. ISBN   978-0-8153-3218-3.
  43. Judson, Olivia (14 August 2002). Dr. Tatiana's Sex Advice to All Creation: The Definitive Guide to the Evolutionary Biology of Sex. Henry Holt and Company. p. 88. ISBN   978-0-8050-6331-8.
  44. Alberts B, Johnson A, Lewis J, Raff M, Roberts K, Walter P (2002). "Sperm". Molecular Biology of the Cell (4th ed.). New York: Garland Science. ISBN   978-0-8153-3218-3.
  45. Alberts B, Johnson A, Lewis J, Raff M, Roberts K, Walter P (2002). "Eggs". Molecular Biology of the Cell (4th ed.). New York: Garland Science. ISBN   978-0-8153-3218-3.
  46. 1 2 Yoshida M, Asturiano JF (25 March 2020). Reproduction in Aquatic Animals: From Basic Biology to Aquaculture Technology. Springer Nature. p. 17. ISBN   978-981-15-2290-1.
  47. Sadava DE, Heller HC, Purves WK, Orians GH, Hillis DM (2008). Life: The Science of Biology. W. H. Freeman. p. 905. ISBN   978-0-7167-7671-0.
  48. 1 2 Stewart-Williams, Steve (17 September 2018). The Ape that Understood the Universe: How the Mind and Culture Evolve. Cambridge University Press. pp. 109–110. ISBN   978-1-108-57752-6.
  49. Ritchison G. "Avian Reproduction". Eastern Kentucky University. Archived from the original on 12 April 2008. Retrieved 3 April 2008.
  50. Alberts B, Johnson A, Lewis J, Raff M, Roberts K, Walter P (2002). "Fertilization". Molecular Biology of the Cell (4th ed.). New York: Garland Science. ISBN   978-0-8153-3218-3.
  51. Jones AG, Avise JC (October 2003). "Male pregnancy". Current Biology. 13 (20): R791. doi:10.1016/j.cub.2003.09.045. PMID   14561416. S2CID   5282823.
  52. Klowden MJ (15 May 2013). Physiological Systems in Insects. Academic Press. p. 228. ISBN   978-0-12-415970-9.
  53. Peinert M, Wipfler B, Jetschke G, Kleinteich T, Gorb SN, Beutel RG, Pohl H (April 2016). "Traumatic insemination and female counter-adaptation in Strepsiptera (Insecta)". Scientific Reports. 6 (1): 25052. Bibcode:2016NatSR...625052P. doi:10.1038/srep25052. PMC   4850473 . PMID   27125507.
  54. Schaffer A (27 September 2007). "Pas de Deux: Why Are There Only Two Sexes?". Slate . Archived from the original on 14 December 2007. Retrieved 30 November 2007.
  55. Smith GM (1947). "On the reproduction of some Pacific coast species of Ulva". American Journal of Botany. 34 (2): 80–87. doi:10.1002/j.1537-2197.1947.tb12961.x. JSTOR   2437232. PMID   20286318.
  56. Gilbert SF (2000). "Gamete Production in Angiosperms". Developmental Biology (6th ed.). Sunderland (MA): Sinauer Associates. ISBN   978-0-87893-243-6.
  57. Micheneau C, Johnson SD, Fay MF (2009). "Orchid pollination: from Darwin to the present day". Botanical Journal of the Linnean Society. 161 (1): 1–19. doi:10.1111/j.1095-8339.2009.00995.x.
  58. Judd, Walter S.; Campbell, Christopher S.; Kellogg, Elizabeth A.; Stevens, Peter F.; Donoghue, Michael J. (2002). Plant systematics, a phylogenetic approach (2 ed.). Sunderland MA, USA: Sinauer Associates Inc. ISBN   0-87893-403-0.
  59. Farjon A (27 April 2010). A Handbook of the World's Conifers: Revised and Updated Edition. BRILL. p. 14. ISBN   978-90-474-3062-9.
  60. Farjon A (2008). The natural history of conifers. Portland, Oregon, US: Timber Press, Inc. pp. 16–21. ISBN   978-0-88192-869-3.
  61. Renner SS, Ricklefs RE (1995). "Dioecy and its correlates in the flowering plants". American Journal of Botany. 82 (5): 596–606. doi:10.2307/2445418. JSTOR   2445418.
  62. Stace CA (2019). New Flora of the British Isles (Fourth ed.). Middlewood Green, Suffolk, U.K.: C & M Floristics. p. 398. ISBN   978-1-5272-2630-2.
  63. Raven PH, Evert RF, Eichhorn SE (2005). Biology of Plants. Macmillan. pp. 436–451. ISBN   978-0-7167-1007-3.
  64. Avise J (18 March 2011). Hermaphroditism: A Primer on the Biology, Ecology, and Evolution of Dual Sexuality. Columbia University Press. pp. 43–46. ISBN   978-0-231-52715-6. Archived from the original on 11 October 2020. Retrieved 18 September 2020.
  65. Bell PR, Hemsley AR (2000). Green plants, their origin and diversity (2 ed.). Cambridge, UK: Cambridge University Press. p. 294. ISBN   0-521-64673-1.
  66. Raven PH, Evert RF, Eichhorn SE (2005). Biology of Plants. Macmillan. pp. 460–463. ISBN   978-0-7167-1007-3.
  67. 1 2 Heitman J, Howlett BJ, Crous PW, Stukenbrock EH, James TY, Gow NR (10 July 2020). Coelho M, Bakkeren G, Sun S, Hood M, Giraud T (eds.). The Fungal Kingdom. John Wiley & Sons. pp. 147–163. ISBN   978-1-55581-958-3.
  68. James T (1 December 2015). "Why mushrooms have evolved to be so promiscuous: Insights from evolutionary and ecological patterns". Fungal Biology Reviews. 29 (3–4): 167–178. doi:10.1016/j.fbr.2015.10.002. ISSN   1749-4613.
  69. Heitman, Joseph; Howlett, Barbara J.; Crous, Pedro W.; Stukenbrock, Eva H.; James, Timothy Yong; Gow, Neil A. R. (10 July 2020). The Fungal Kingdom. John Wiley & Sons. pp. 177–191. ISBN   978-1-55581-958-3.
  70. Watkinson SC, Boddy L, Money N (2015). The Fungi. Elsevier Science. p. 115. ISBN   978-0-12-382035-8. Archived from the original on 26 February 2020. Retrieved 18 February 2018.
  71. Lane N (13 October 2005). Power, Sex, Suicide: Mitochondria and the Meaning of Life. Oxford University Press, UK. p. 236. ISBN   978-0-19-280481-5.
  72. Lodish H, Berk A, Zipursky SL, Matsudaira P, Baltimore D, Darnell J (2000). "Section 14.1: Cell-Type Specification and Mating-Type Conversion in Yeast". Molecular Cell Biology (Fourth ed.). WH Freeman and Co. ISBN   978-0-7167-4366-8.
  73. Laybourn-Parry J (8 March 2013). A Functional Biology of Free-Living Protozoa. Springer Science & Business Media. pp. 86–88. ISBN   978-1-4684-7316-2.
  74. Wang G, Chen K, Zhang J, Deng S, Xiong J, He X, et al. (December 2020). "Drivers of Mating Type Composition in Tetrahymena thermophila". Genome Biology and Evolution. 12 (12): 2328–2343. doi:10.1093/gbe/evaa197. PMC   7846192 . PMID   32946549.
  75. Tanurdzic M, Banks JA (2004). "Sex-determining mechanisms in land plants". The Plant Cell. 16 Suppl: S61-71. doi:10.1105/tpc.016667. PMC   2643385 . PMID   15084718.
  76. Warner DA, Shine R (January 2008). "The adaptive significance of temperature-dependent sex determination in a reptile". Nature. 451 (7178): 566–8. Bibcode:2008Natur.451..566W. doi:10.1038/nature06519. PMID   18204437. S2CID   967516.
  77. Beukeboom LW, Perrin N (2014). The Evolution of Sex Determination. Oxford University Press. p. 16. ISBN   978-0-19-965714-8.
  78. Wallis MC, Waters PD, Graves JA (October 2008). "Sex determination in mammals--before and after the evolution of SRY". Cellular and Molecular Life Sciences. 65 (20): 3182–95. doi:10.1007/s00018-008-8109-z. PMID   18581056. S2CID   31675679.
  79. Kaiser VB, Bachtrog D (2010). "Evolution of sex chromosomes in insects". Annual Review of Genetics. 44: 91–112. doi:10.1146/annurev-genet-102209-163600. PMC   4105922 . PMID   21047257.
  80. Dellaporta SL, Calderon-Urrea A (October 1993). "Sex determination in flowering plants". The Plant Cell. 5 (10): 1241–51. doi:10.1105/tpc.5.10.1241. JSTOR   3869777. PMC   160357 . PMID   8281039.
  81. Fusco G, Minelli A (10 October 2019). The Biology of Reproduction. Cambridge University Press. pp. 306–308. ISBN   978-1-108-49985-9.
  82. Smith CA, Katz M, Sinclair AH (February 2003). "DMRT1 is upregulated in the gonads during female-to-male sex reversal in ZW chicken embryos". Biology of Reproduction. 68 (2): 560–70. doi: 10.1095/biolreprod.102.007294 . PMID   12533420.
  83. Majerus ME (2003). Sex Wars: Genes, Bacteria, and Biased Sex Ratios. Princeton University Press. p. 59. ISBN   978-0-691-00981-0.
  84. "Evolution of the Y Chromosome". Annenberg Media. Annenberg Media. Archived from the original on 4 November 2004. Retrieved 1 April 2008.
  85. Bull JJ (1983). Evolution of sex determining mechanisms. p. 17. ISBN   0-8053-0400-2.
  86. Thiriot-Quiévreux C (2003). "Advances in chromosomal studies of gastropod molluscs". Journal of Molluscan Studies. 69 (3): 187–202. doi: 10.1093/mollus/69.3.187 .
  87. Yoshimura A (2005). "Karyotypes of two American field crickets: Gryllus rubens and Gryllus sp. (Orthoptera: Gryllidae)". Entomological Science. 8 (3): 219–222. doi:10.1111/j.1479-8298.2005.00118.x. S2CID   84908090.
  88. Meyer BJ (1997). "Sex Determination and X Chromosome Dosage Compensation: Sexual Dimorphism". In Riddle DL, Blumenthal T, Meyer BJ, Priess JR (eds.). C. elegans II. Cold Spring Harbor Laboratory Press. ISBN   978-0-87969-532-3.
  89. Handbuch Der Zoologie / Handbook of Zoology. Walter de Gruyter. 1925. ISBN   978-3-11-016210-3. Archived from the original on 11 October 2020. Retrieved 29 September 2020 via Google Books.
  90. Tanurdzic M, Banks JA (2004). "Sex-determining mechanisms in land plants". The Plant Cell. 16 Suppl (Suppl): S61-71. doi:10.1105/tpc.016667. PMC   2643385 . PMID   15084718.
  91. Fusco G, Minelli A (10 October 2019). The Biology of Reproduction. Cambridge University Press. p. 124. ISBN   978-1-108-49985-9.
  92. Todd EV, Liu H, Muncaster S, Gemmell NJ (2016). "Bending Genders: The Biology of Natural Sex Change in Fish". Sexual Development. 10 (5–6): 223–241. doi: 10.1159/000449297 . PMID   27820936. S2CID   41652893.
  93. Gilbert SF (2000). "Environmental Sex Determination". Developmental Biology. 6th Edition.
  94. Charlesworth B (August 2003). "Sex determination in the honeybee". Cell. 114 (4): 397–8. doi: 10.1016/S0092-8674(03)00610-X . PMID   12941267.
  95. de la Filia A, Bain S, Ross L (June 2015). "Haplodiploidy and the reproductive ecology of Arthropods". Current Opinion in Insect Science. 9: 36–43. doi:10.1016/j.cois.2015.04.018. PMID   32846706.
  96. Hamilton, W.D. (1967). "Extraordinary sex ratios". Science. 156 (3774): 477–488. Bibcode:1967Sci...156..477H. doi:10.1126/science.156.3774.477. PMID   6021675.
  97. Whitfield J (June 2004). "Everything you always wanted to know about sexes". PLOS Biology. 2 (6): e183. doi:10.1371/journal.pbio.0020183. PMC   423151 . PMID   15208728. One thing biologists do agree on is that males and females count as different sexes. And they also agree that the main difference between the two is gamete size: males make lots of small gametes—sperm in animals, pollen in plants—and females produce a few big eggs.
  98. Pierce BA (2012). Genetics: A Conceptual Approach. W. H. Freeman. p. 74. ISBN   978-1-4292-3252-4.
  99. Dawkins, Richard (2016). The Selfish Gene. Oxford University Press. pp. 183–184. ISBN   978-0-19-878860-7. However, there is one fundamental feature of the sexes which can be used to label males as males, and females as females, throughout animals and plants. This is that the sex cells or ‘gametes' of males are much smaller and more numerous than the gametes of females. This is true whether we are dealing with animals or plants. One group of individuals has large sex cells, and it is convenient to use the word female for them. The other group, which it is convenient to call male, has small sex cells. The difference is especially pronounced in reptiles and in birds, where a single egg cell is big enough and nutritious enough to feed a developing baby for. Even in humans, where the egg is microscopic, it is still many times larger than the sperm. As we shall see, it is possible to interpret all the other differences between the sexes as stemming from this one basic difference.
  100. 1 2 Mori, Emiliano; Mazza, Giuseppe; Lovari, Sandro (2017). "Sexual Dimorphism". In Vonk, Jennifer; Shackelford, Todd (eds.). Encyclopedia of Animal Cognition and Behavior. Cham: Springer International Publishing. pp. 1–7. doi:10.1007/978-3-319-47829-6_433-1. ISBN   978-3-319-47829-6 . Retrieved 5 June 2021.
  101. Mealey, Linda (13 April 2000). Sex Differences: Developmental and Evolutionary Strategies. Academic Press. pp. 87–88. ISBN   978-0-08-054113-6.
  102. Buss, David M. (27 December 2016). The Evolution of Desire: Strategies of Human Mating. Basic Books. pp. 31–32. ISBN   978-0-465-09776-0.
  103. Pack PE (20 December 2016). CliffsNotes AP Biology, 5th Edition. Houghton Mifflin Harcourt. p. 219. ISBN   978-0-544-78417-8.
  104. Minelli A, Fusco G (10 October 2019). The Biology of Reproduction. Cambridge University Press. pp. 116–117. ISBN   9781108499859. Archived from the original on 11 October 2020. Retrieved 11 October 2020. However, species are also considered gonochoric if (as develop mental abnormalities[ clarification needed ] or because of genetic mutations) intersex individuals accidentally occur, i.e. ones with a mix of both male and female phenotypic characters ( Section 6.1.1 ).
  105. 1 2 "intersex | Definition & Facts". Encyclopedia Britannica. Archived from the original on 25 July 2020. Retrieved 11 October 2020. The [intersex] condition usually results from extra chromosomes or a hormonal abnormality during embryological development.
  106. Farrell A (1 June 2011). Encyclopedia of Fish Physiology: From Genome to Environment. Academic Press. ISBN   978-0-08-092323-9. Thus, strictly speaking, all hermaphrodites are intersex at one time point, but not all intersexes are hermaphrodites. This definition is usually applied to gonochoristic species to describe those individuals that are not normal for the species.
  107. Wilkinson GS, Reillo PR (22 January 1994). "Female choice response to artificial selection on an exaggerated male trait in a stalk-eyed fly" (PDF). Proceedings of the Royal Society B. 225 (1342): 1–6. Bibcode:1994RSPSB.255....1W. CiteSeerX . doi:10.1098/rspb.1994.0001. S2CID   5769457. Archived from the original (PDF) on 10 September 2006.
  108. Drees BM, Jackman J (1999). "Southern black widow spider". Field Guide to Texas Insects. Houston, Texas: Gulf Publishing Company. Archived from the original on 31 August 2003. Retrieved 8 August 2012 via Extension Entomology,, Texas A&M University.
  109. Stuart-Smith J, Swain R, Stuart-Smith R, Wapstra E (2007). "Is fecundity the ultimate cause of female-biased size dimorphism in a dragon lizard?". Journal of Zoology. 273 (3): 266–272. doi:10.1111/j.1469-7998.2007.00324.x.
  110. Shaw AJ (2000). "Population ecology, population genetics, and microevolution". In Shaw AJ, Goffinet B (eds.). Bryophyte Biology. Cambridge: Cambridge University Press. pp. 379–380. ISBN   978-0-521-66097-6.
  111. 1 2 Schuster RM (1984). "Comparative Anatomy and Morphology of the Hepaticae". New Manual of Bryology. 2. Nichinan, Miyazaki, Japan: The Hattori botanical Laboratory. p. 891.
  112. Crum HA, Anderson LE (1980). Mosses of Eastern North America. 1. New York: Columbia University Press. p. 196. ISBN   978-0-231-04516-2.
  113. Briggs DA (1965). "Experimental taxonomy of some British species of genus Dicranum". New Phytologist. 64 (3): 366–386. doi: 10.1111/j.1469-8137.1965.tb07546.x .
  114. Zahavi A, Zahavi A (1997). The handicap principle: a missing piece of Darwin's puzzle. Oxford University Press. ISBN   978-0-19-510035-8.

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