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Parthenogenesis (; from the Greek + q.v..) is a natural form of asexual reproduction in which the develops directly from an without need for . In , parthenogenesis means the development of an embryo from an unfertilized . In , parthenogenesis is a component process of . In , parthenogenesis can mean the development of an embryo from either an individual sperm or an individual egg.

Parthenogenesis occurs naturally in some plants, , animal (including , some , , some , , some mites, some , some , and ), and a few , such as some , , and . This type of reproduction has been induced artificially in animal species that naturally reproduce through sex, including fish, amphibians, and mice.

Normal egg cells form in the process of and are , with half as many as their mother's body cells. Haploid individuals, however, are usually non-viable, and parthenogenetic offspring usually have the chromosome number. Depending on the mechanism involved in restoring the diploid number of chromosomes, parthenogenetic offspring may have anywhere between all and half of the mother's . In some types of parthenogenesis, the offspring that have all of the mother's genetic material are called full and those having only half are called half clones. Full clones are usually formed without meiosis. If meiosis occurs, the offspring get only a fraction of the mother's alleles since crossing over of takes place during meiosis, creating variation.

Parthenogenetic offspring in species that use either the XY or the X0 sex-determination system have two X chromosomes and are female. In species that use the ZW sex-determination system, they have either two Z chromosomes (male) or two W chromosomes (mostly non-viable but rarely a female), or they could have one Z and one W chromosome (female).


Life history types
Parthenogenesis is a form of asexual reproduction in which the develops directly from an without need for . It occurs naturally in some plants, , animal species (including , some , , some , , some mites, some , some , and ), and a few , such as some , , ,
(1986). 9780920269817, Torstar Books.
and . This type of reproduction has been induced artificially in several animal species that naturally reproduce through sex, including fish, amphibians, and mice.

Some species reproduce exclusively by parthenogenesis (such as the ), while others can switch between sexual reproduction and parthenogenesis. This is called facultative parthenogenesis (other terms are cyclical parthenogenesis, heterogamy

(1996). 9783110106619, Walter de Gruyter. .
or heterogony). The switch between sexuality and parthenogenesis in such species may be triggered by the season (, some ), or by a lack of males or by conditions that favour rapid population growth ( and like ). In these species, asexual reproduction occurs either in summer (aphids) or as long as conditions are favourable. This is because in asexual reproduction, a successful genotype can spread quickly without being modified by sex or wasting resources on male offspring who will not give birth. Some species can produce both sexually and through parthenogenesis, and offspring in the same clutch of a species of tropical lizard can be a mix of sexually produced offspring and parthenogenically produced offspring. In California condors, facultative parthenogenesis can occur even when a male is present and available for a female to breed with. In times of stress, offspring produced by sexual reproduction may be fitter as they have new, possibly beneficial gene combinations. In addition, sexual reproduction provides the benefit of meiotic recombination between non-sister chromosomes, a process associated with repair of double-strand breaks and other DNA damages that may be induced by stressful conditions.
(1987). 9780120176243

Many taxa with heterogony have within them species that have lost the sexual phase and are now completely asexual. Many other cases of obligate parthenogenesis (or gynogenesis) are found among polyploids and hybrids where the chromosomes cannot pair for meiosis.

(2025). 9780190688554 .

The production of female offspring by parthenogenesis is referred to as (e.g., aphids) while the production of males by parthenogenesis is referred to as (e.g., bees). When unfertilized eggs develop into both males and females, the phenomenon is called deuterotoky.


Types and mechanisms
Parthenogenesis can occur without meiosis through mitotic oogenesis. This is called apomictic parthenogenesis. Mature egg cells are produced by mitotic divisions, and these cells directly develop into embryos. In flowering plants, cells of the can undergo this process. The offspring produced by apomictic parthenogenesis are full clones of their mother, as in aphids.

Parthenogenesis involving is more complicated. In some cases, the offspring are haploid (e.g., male ). In other cases, collectively called automictic parthenogenesis, the ploidy is restored to diploidy by various means. This is because haploid individuals are not viable in most species. In automictic parthenogenesis, the offspring differ from one another and their mother. They are called half clones of their mother.


Automixis
Automixis includes several reproductive mechanisms, some of which are parthenogenetic.

Diploidy can be restored by the doubling of the chromosomes without cell division before meiosis begins or after meiosis is completed. This is an endomitotic cycle. Diploidy can also be restored by fusion of the first two , or by fusion of the meiotic products. The chromosomes may not separate at one of the two anaphases (restitutional meiosis)l or the nuclei produced may fuse, or one of the polar bodies may fuse with the egg cell at some stage during its maturation.

Some authors consider all forms of automixis sexual as they involve recombination. Many others classify the endomitotic variants as asexual and consider the resulting embryos parthenogenetic. Among these authors, the threshold for classifying automixis as a sexual process depends on when the products of anaphase I or of anaphase II are joined. The criterion for sexuality varies from all cases of restitutional meiosis, to those where the nuclei fuse or to only those where gametes are mature at the time of fusion. Those cases of automixis that are classified as sexual reproduction are compared to self-fertilization in their mechanism and consequences.

The genetic composition of the offspring depends on what type of automixis takes place. When endomitosis occurs before meiosisCosín, Darío J. Díaz, Marta Novo, and Rosa Fernández. "Reproduction of Earthworms: Sexual Selection and Parthenogenesis". In Biology of Earthworms Https://doi.org/10.1007%2F978-3-642-14636-7_5.< /ref> or when central fusion occurs (restitutional meiosis of anaphase I or the fusion of its products), the offspring get all to more than half of the mother's genetic material and heterozygosity is mostly preserved (if the mother has two alleles for a locus, the offspring will likely get both). This is because in the homologous chromosomes are separated. Heterozygosity is not completely preserved when crossing over occurs in central fusion. In the case of pre-meiotic doubling, recombination, if it happens, occurs between identical sister chromatids.

If terminal fusion (restitutional meiosis of anaphase II or the fusion of its products) occurs, a little over half of the mother's genetic material is present in the offspring, and the offspring are mostly homozygous. This is because at anaphase II the sister chromatids are separated and whatever heterozygosity is present is due to crossing over. In the case of endomitosis after meiosis, the offspring is completely homozygous and has only half the mother's genetic material. This can result in parthenogenetic offspring being unique from each other and from their mother.


Sex of the offspring
In apomictic parthenogenesis, the offspring are clones of the mother and hence (except for aphids) are usually female. In the case of aphids, parthenogenetically produced males and females are clones of their mother except that the males lack one of the X chromosomes (XO).

When meiosis is involved, the sex of the offspring depends on the type of sex determination system and the type of apomixis. In species that use the XY sex-determination system, parthenogenetic offspring have two X chromosomes and are female. In species that use the ZW sex-determination system the offspring genotype may be one of ZW (female), ZZ (male), or WW (non-viable in most species, but a fertile, viable female in a few, e.g., ). ZW offspring are produced by before meiosis or by central fusion. ZZ and WW offspring occur either by terminal fusion or by endomitosis in the egg cell.

In polyploid obligate parthenogens, like the whiptail lizard, all the offspring are female.

In many hymenopteran insects, such as honeybees, female eggs are produced sexually, using sperm from a drone father, while the production of further drones (males) depends on the queen (and occasionally workers) producing unfertilized eggs. This means that females (workers and queens) are always diploid, while males (drones) are always haploid and are produced parthenogenetically.


Facultative
Facultative parthenogenesis occurs when a female can produce offspring either sexually or via asexual reproduction.Bell, G. (1982). The Masterpiece of Nature: The Evolution and Genetics of Sexuality, University of California Press, Berkeley, pp. 1–635 (see p. 295). Facultative parthenogenesis is extremely rare in nature, with only a few examples of animal taxa capable of facultative parthenogenesis. One of the best-known examples of taxa exhibiting facultative parthenogenesis are ; presumably, this is the default reproductive mode of all species in this insect order. Facultative parthenogenesis has generally been believed to be a response to a lack of a viable male. A female may undergo facultative parthenogenesis if a male is absent from the habitat or if it is unable to produce viable offspring. However, California condors and the tropical lizard Lepidophyma smithii both can produce parthenogenic offspring in the presence of males, indicating that facultative parthenogenesis may be more common than previously thought and is not simply a response to a lack of males.

In , a generation sexually conceived by a male and a female produces only females. The reason for this is the non-random segregation of the 'X' and 'O' during .

Facultative parthenogenesis is often used to describe cases of spontaneous parthenogenesis in normally sexual animals. For example, many cases of spontaneous parthenogenesis in , some , , and a variety of domesticated birds were widely attributed to facultative parthenogenesis. These cases are examples of spontaneous parthenogenesis. The occurrence of such asexually produced eggs in sexual animals can be explained by a meiotic error, leading to eggs produced via .


Obligate
Obligate parthenogenesis is the process in which organisms exclusively reproduce through asexual means. Many species have transitioned to obligate parthenogenesis over evolutionary time. Well-documented transitions to obligate parthenogenesis have been found in numerous metazoan taxa, albeit through highly diverse mechanisms. These transitions often occur as a result of inbreeding or mutation within large populations. web. Some documented species, specifically salamanders and geckos, rely on obligate parthenogenesis as their major method of reproduction. As such, there are over 80 species of unisex reptiles (mostly lizards but including a single snake species), amphibians, and fishes in nature for which males are no longer a part of the reproductive process. A female produces an ovum with a full set (two sets of genes) provided solely by the mother. Thus, a male is not needed to provide sperm to fertilize the egg. This form of asexual reproduction is thought in some cases to be a serious threat to biodiversity due to the subsequent lack of gene variation and potentially decreased fitness of the offspring.

Some invertebrate species that feature (partial) sexual reproduction in their native range are found to reproduce solely by parthenogenesis in areas to which they have been . Relying solely on parthenogenetic reproduction has several advantages for an : it obviates the need for individuals in a very sparse initial population to search for mates; and an exclusively female sex distribution allows a population to multiply and invade more rapidly (potentially twice as fast). Examples include several species and the willow sawfly, Nematus oligospilus, which is sexual in its native habitat but parthenogenetic where it has been introduced into the Southern Hemisphere.


Natural occurrence
Parthenogenesis does not apply to species.
(1978). 9780521218870, CUP Archive. .
Parthenogenesis occurs naturally in , , , , and some other invertebrates, as well as in many plants. Among , strict parthenogenesis is only known to occur in lizards, snakes, birds, and sharks. Fish, amphibians, and reptiles make use of various forms of gynogenesis and hybridogenesis (an incomplete form of parthenogenesis). The first all-female (unisexual) reproduction in was described in the fish in 1932. Since then, at least 50 species of unisexual vertebrates have been described, including at least 20 fish, 25 lizards, a single snake species, frogs, and salamanders.Vrijenhoek, R.C., R.M. Dawley, C.J. Cole, and J.P. Bogart. 1989. "A list of the known unisexual vertebrates", pp. 19–23 in: Evolution and Ecology of Unisexual Vertebrates. R.M. Dawley and J.P. Bogart (eds.) Bulletin 466, New York State Museum, Albany


Artificial induction
Use of an electrical or chemical stimulus can produce the beginning of the process of parthenogenesis in the asexual development of viable offspring.

During oocyte development, high metaphase-promoting factor (MPF) activity causes mammalian oocytes to arrest at the metaphase II stage until fertilization by a sperm. The fertilization event causes intracellular calcium oscillations and targeted degradation of cyclin B, a regulatory subunit of MPF, thus permitting the MII-arrested oocyte to proceed through meiosis.

To initiate unfertilised development of swine oocytes, various methods exist to induce an artificial activation that mimics sperm entry, such as calcium ionophore treatment, microinjection of calcium ions, or electrical stimulation. Treatment with cycloheximide, a non-specific protein synthesis inhibitor, enhances the development of unfertilised eggs in swine presumably by continual inhibition of MPF/cyclin B. As meiosis proceeds, extrusion of the second polar is blocked by exposure to cytochalasin B. This treatment results in a diploid (2 maternal genomes) parthenote The resulting embryos can be surgically transferred to a recipient oviduct for further development, but will succumb to developmental failure after ≈30 days of gestation. The swine placenta in these cases often appears hypo-vascular: see free image (Figure 1) in linked reference.

Induced parthenogenesis of this type in and results in abnormal development. This is because mammals have imprinted genetic regions, where either the maternal or the paternal chromosome is inactivated in the offspring for development to proceed normally. A mammal developing from parthenogenesis would have double doses of maternally imprinted genes and lack paternally imprinted genes, leading to developmental abnormalities. It has been suggested that defects in folding or interdigitation are one cause of swine abortive development. As a consequence, research on the induced development of unfertilised eggs in humans is focused on the production of embryonic stem cells for use in medical treatment, not as a reproductive strategy.

In 2022, researchers reported that they had produced viable offspring born from unfertilized eggs in mice, addressing the problems of genomic imprinting by "targeted DNA methylation rewriting of seven imprinting control regions".


In humans
In 1955, , a geneticist specializing in the reproductive biology of the ( Lebistes reticulatus), claimed that parthenogenesis may occur (though very rarely) in humans, leading to so-called "virgin births". This created some sensation among her colleagues and the lay public alike. Time, 28 November 1955; Editorial in The Lancet, 2: 967 (1955) Sometimes an embryo may begin to divide without fertilization, but it cannot fully develop on its own; so while it may create some skin and nerve cells, it cannot create others (such as skeletal muscle) and becomes a type of benign tumor called an ovarian . Spontaneous ovarian activation is not rare and has been known about since the 19th century. Some teratomas can even become primitive fetuses (fetiform teratoma) with imperfect heads, limbs, and other structures, but are non-viable.

In 1995, there was a reported case of partial human parthenogenesis; a boy was found to have some of his cells (such as white blood cells) lacking any genetic content from his father. Scientists believe that an unfertilized egg began to self-divide but then had some (but not all) of its cells fertilized by a sperm cell; this must have happened early in development, as self-activated eggs quickly lose their ability to be fertilized. The unfertilized cells eventually duplicated their DNA, boosting their chromosomes to 46. When the unfertilized cells hit a developmental block, the fertilized cells took over and developed that tissue. The boy had asymmetrical facial features and learning difficulties but was otherwise healthy. This would make him a parthenogenetic chimera (a child with two cell lineages in his body).Philip Cohen, "The boy whose blood has no father", New Scientist, 7.10.1995 While over a dozen similar cases have been reported since then (usually discovered after the patient demonstrated clinical abnormalities), there have been no scientifically confirmed reports of a non-chimeric, clinically healthy human parthenote (i.e. produced from a single, parthenogenetic-activated oocyte).

In 2007, the International Stem Cell Corporation of California announced that Elena Revazova had intentionally created human stem cells from unfertilized human eggs using parthenogenesis. The process may offer a way to create stem cells genetically matched to a particular female to treat degenerative diseases. The same year, Revazova and ISCC published an article describing how to produce human stem cells that are homozygous in the HLA region of DNA. These stem cells are called HLA homozygous parthenogenetic human stem cells (hpSC-Hhom) and would allow derivatives of these cells to be implanted without immune rejection. With the selection of oocyte donors according to HLA , it would be possible to generate a bank of cell lines whose tissue derivatives, collectively, could be MHC-matched with a significant number of individuals within the human population.

After an independent investigation, it was revealed that the discredited South Korean scientist unknowingly produced the first human embryos resulting from parthenogenesis. Initially, Hwang claimed he and his team had extracted stem cells from cloned human embryos, a result later found to be fabricated. Further examination of the chromosomes of these cells shows indicators of parthenogenesis in those extracted stem cells, similar to those found in the mice created by Tokyo scientists in 2004. Although Hwang deceived the world about being the first to create artificially cloned human embryos, he contributed a breakthrough to stem cell research by creating human embryos using parthenogenesis.Williams, Chris. "Stem cell fraudster made 'virgin birth' breakthrough: Silver lining for Korean science scandal", The Register, 3 August 2007.


Similar phenomena

Gynogenesis
A form of asexual reproduction related to parthenogenesis is gynogenesis. Here, offspring are produced by the same mechanism as in parthenogenesis, but with the requirement that the egg merely be stimulated by the presence of to develop. However, the sperm cell does not contribute any genetic material to the offspring. Since gynogenetic species are all female, activation of their eggs requires mating with males of a closely related species for the needed stimulus. Some of the genus are gynogenetic and appear to have been so for over a million years. The success of those salamanders may be due to rare fertilization of eggs by males, introducing new material to the gene pool, which may result from perhaps only one mating out of a million. In addition, the is known to reproduce by gynogenesis.


Hybridogenesis
Hybridogenesis is a mode of reproduction of hybrids. Hybridogenetic hybrids (for example AB ), usually females, during exclude one of parental genomes (A) and produce with unrecombined of second parental species (B), instead of containing mixed recombined parental genomes. First genome (A) is restored by of these gametes with gametes from the first species (AA, sexual host, usually male). Hybridogenesis is not completely asexual, but hemiclonal: half the genome is passed to the next generation clonally, unrecombined, intact (B), other half sexually, recombined (A). This process continues, so that each generation is half (or hemi-) clonal on the mother's side and has half new genetic material from the father's side.

This form of reproduction is seen in some live-bearing fish of the genus as well as in some of the spp. ("green frogs" or "waterfrogs"):

  • P. kl. esculentus (edible frog): P. lessonae × P. ridibundus,
  • P. kl. grafi (Graf's hybrid frog): P. perezi × P. ridibundus
  • P. kl. hispanicus (Italian edible frog) – unknown origin: P. bergeri × P. ridibundus or P. kl. esculentus

Other examples where hybridogenesis is at least one of the modes of reproduction include i.e.,

  • Iberian minnow Tropidophoxinellus alburnoides ( Squalius pyrenaicus × hypothetical ancestor related with Anaecypris hispanica)
  • spined loaches hankugensis × C. longicorpus
  • Bacillus stick insects × Bacillus grandii benazzii


In human culture
Parthenogenesis, in the form of reproduction from a single individual (typically a god), is common in mythology, religion, and folklore around the world, including in ancient ; for example, was born from the head of .
(2025). 9780230618862, Palgrave Macmillan.
In Christianity, Islam, and the Bahá'í Faith, there is the virgin birth of Jesus, and stories of miraculous births also appear in other global religions.
(2025). 9789053565032, Eerdmans.
The theme is one of several aspects of reproductive biology explored in science fiction.


See also
  • - a form of quasi-sexual reproduction in which a male is the sole source of the nuclear genetic material in the embryo
  • Telescoping generations
  • – conducted experiments that established what is now termed parthenogenesis in aphids
  • – Polish and a pioneer of parthenogenesis among
  • – caused the eggs of sea urchins to begin embryonic development without sperm
  • – plants with seedless fruit


Further reading
  • Dawley, Robert M. & Bogart, James P. (1989). Evolution and Ecology of Unisexual Vertebrates. Albany: New York State Museum.
  • Futuyma, Douglas J. & Slatkin, Montgomery. (1983). Coevolution. Sunderland, Mass: Sinauer Associates.
  • Maynard Smith, John. (1978). The Evolution of Sex. Cambridge: Cambridge University Press.
  • Michod, Richard E. & Levin, Bruce R. (1988). The Evolution of Sex. Sunderland, Mass: Sinauer Associates.
  • Stearns, Stephan C. (1988). The Evolution of Sex and Its Consequences (Experientia Supplementum, Vol. 55). Boston: Birkhauser.


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