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Polyploidy

February 26, 2023

Not to be confused with "polypoid", resembling a polyp.

Polyploidy is a condition in which the cells of an organism have more than one pair of (homologous) chromosomes. Most species whose cells have nuclei (eukaryotes) are diploid, meaning they have two complete sets of chromosomes, one from each of two parents; each set contains the same number of chromosomes, and the chromosomes are joined in pairs of homologous chromosomes. However, some organisms are polyploid. Polyploidy is especially common in plants. Most eukaryotes have diploid somatic cells, but produce haploid gametes (eggs and sperm) by meiosis. A monoploid has only one set of chromosomes, and the term is usually only applied to cells or organisms that are normally diploid. Males of bees and other Hymenoptera, for example, are monoploid. Unlike animals, plants and multicellular algae have life cycles with two alternating multicellular generations. The gametophyte generation is haploid, and produces gametes by mitosis; the sporophyte generation is diploid and produces spores by meiosis.

This image shows haploid (single), diploid (double), triploid (triple), and tetraploid (quadruple) sets of chromosomes. Triploid and tetraploid chromosomes are examples of polyploidy.

Polyploidy may occur due to abnormal cell division, either during mitosis, or more commonly from the failure of chromosomes to separate during meiosis or from the fertilization of an egg by more than one sperm.[1] In addition, it can be induced in plants and cell cultures by some chemicals: the best known is colchicine, which can result in chromosome doubling, though its use may have other less obvious consequences as well. Oryzalin will also double the existing chromosome content.

Polyploidy occurs in highly differentiated human tissues in the liver, heart muscle, bone marrow and the placenta.[2][failed verification] It occurs in the somatic cells of some animals, such as goldfish,[3] salmon, and salamanders, but is especially common among ferns and flowering plants (see Hibiscus rosa-sinensis), including both wild and cultivated species. Wheat, for example, after millennia of hybridization and modification by humans, has strains that are diploid (two sets of chromosomes), tetraploid (four sets of chromosomes) with the common name of durum or macaroni wheat, and hexaploid (six sets of chromosomes) with the common name of bread wheat. Many agriculturally important plants of the genus Brassica are also tetraploids. Sugarcane can have ploidy levels higher than octaploid.[4]

Polyploidization can be a mechanism of sympatric speciation because polyploids are usually unable to interbreed with their diploid ancestors. An example is the plant Erythranthe peregrina. Sequencing confirmed that this species originated from E. × robertsii, a sterile triploid hybrid between E. guttata and E. lutea, both of which have been introduced and naturalised in the United Kingdom. New populations of E. peregrina arose on the Scottish mainland and the Orkney Islands via genome duplication from local populations of E. × robertsii.[5] Because of a rare genetic mutation, E. peregrina is not sterile.[6]

Terminology

Types

"Triploid" redirects here. For the human chromosomal disorder (69 XXX, etc.), see Triploid syndrome.

 
Organ-specific patterns of endopolyploidy (from 2x to 64x) in the giant ant Dinoponera australis

Polyploid types are labeled according to the number of chromosome sets in the nucleus. The letter x is used to represent the number of chromosomes in a single set:

Classification

Autopolyploidy

Autopolyploids are polyploids with multiple chromosome sets derived from a single taxon.

Two examples of natural autopolyploids are the piggyback plant, Tolmiea menzisii[14] and the white sturgeon, Acipenser transmontanum.[15] Most instances of autopolyploidy result from the fusion of unreduced (2n) gametes, which results in either triploid (n + 2n = 3n) or tetraploid (2n + 2n = 4n) offspring.[16] Triploid offspring are typically sterile (as in the phenomenon of triploid block), but in some cases they may produce high proportions of unreduced gametes and thus aid the formation of tetraploids. This pathway to tetraploidy is referred to as the triploid bridge.[16] Triploids may also persist through asexual reproduction. In fact, stable autotriploidy in plants is often associated with apomictic mating systems.[17] In agricultural systems, autotriploidy can result in seedlessness, as in watermelons and bananas.[18] Triploidy is also utilized in salmon and trout farming to induce sterility.[19][20]

Rarely, autopolyploids arise from spontaneous, somatic genome doubling, which has been observed in apple (Malus domesticus) bud sports.[21] This is also the most common pathway of artificially induced polyploidy, where methods such as protoplast fusion or treatment with colchicine, oryzalin or mitotic inhibitors are used to disrupt normal mitotic division, which results in the production of polyploid cells. This process can be useful in plant breeding, especially when attempting to introgress germplasm across ploidal levels.[22]

Autopolyploids possess at least three homologous chromosome sets, which can lead to high rates of multivalent pairing during meiosis (particularly in recently formed autopolyploids, also known as neopolyploids) and an associated decrease in fertility due to the production of aneuploid gametes.[23] Natural or artificial selection for fertility can quickly stabilize meiosis in autopolyploids by restoring bivalent pairing during meiosis, but the high degree of homology among duplicated chromosomes causes autopolyploids to display polysomic inheritance.[24] This trait is often used as a diagnostic criterion to distinguish autopolyploids from allopolyploids, which commonly display disomic inheritance after they progress past the neopolyploid stage.[25] While most polyploid species are unambiguously characterized as either autopolyploid or allopolyploid, these categories represent the ends of a spectrum of divergence between parental subgenomes. Polyploids that fall between these two extremes, which are often referred to as segmental allopolyploids, may display intermediate levels of polysomic inheritance that vary by locus.[26][27]

About half of all polyploids are thought to be the result of autopolyploidy,[28][29] although many factors make this proportion hard to estimate.[30]

Allopolyploidy

Allopolyploids or amphipolyploids or heteropolyploids are polyploids with chromosomes derived from two or more diverged taxa.

As in autopolyploidy, this primarily occurs through the fusion of unreduced (2n) gametes, which can take place before or after hybridization. In the former case, unreduced gametes from each diploid taxon – or reduced gametes from two autotetraploid taxa – combine to form allopolyploid offspring. In the latter case, one or more diploid F1 hybrids produce unreduced gametes that fuse to form allopolyploid progeny.[31] Hybridization followed by genome duplication may be a more common path to allopolyploidy because F1 hybrids between taxa often have relatively high rates of unreduced gamete formation – divergence between the genomes of the two taxa result in abnormal pairing between homoeologous chromosomes or nondisjunction during meiosis.[31] In this case, allopolyploidy can actually restore normal, bivalent meiotic pairing by providing each homoeologous chromosome with its own homologue. If divergence between homoeologous chromosomes is even across the two subgenomes, this can theoretically result in rapid restoration of bivalent pairing and disomic inheritance following allopolyploidization. However multivalent pairing is common in many recently formed allopolyploids, so it is likely that the majority of meiotic stabilization occurs gradually through selection.[23][25]

Because pairing between homoeologous chromosomes is rare in established allopolyploids, they may benefit from fixed heterozygosity of homoeologous alleles.[32] In certain cases, such heterozygosity can have beneficial heterotic effects, either in terms of fitness in natural contexts or desirable traits in agricultural contexts. This could partially explain the prevalence of allopolyploidy among crop species. Both bread wheat and Triticale are examples of an allopolyploids with six chromosome sets. Cotton, peanut, or quinoa are allotetraploids with multiple origins. In Brassicaceous crops, the Triangle of U describes the relationships between the three common diploid Brassicas (B. oleracea, B. rapa, and B. nigra) and three allotetraploids (B. napus, B. juncea, and B. carinata) derived from hybridization among the diploid species. A similar relationship exists between three diploid species of Tragopogon (T. dubius, T. pratensis, and T. porrifolius) and two allotetraploid species (T. mirus and T. miscellus).[33] Complex patterns of allopolyploid evolution have also been observed in animals, as in the frog genus Xenopus.[34]

Aneuploid

Main article: Aneuploidy

Organisms in which a particular chromosome, or chromosome segment, is under- or over-represented are said to be aneuploid (from the Greek words meaning "not", "good", and "fold"). Aneuploidy refers to a numerical change in part of the chromosome set, whereas polyploidy refers to a numerical change in the whole set of chromosomes.[35]

Endopolyploidy

Polyploidy occurs in some tissues of animals that are otherwise diploid, such as human muscle tissues.[36] This is known as endopolyploidy. Species whose cells do not have nuclei, that is, prokaryotes, may be polyploid, as seen in the large bacterium Epulopiscium fishelsoni.[37] Hence ploidy is defined with respect to a cell.

Monoploid

Main article: Monoploidy

A monoploid has only one set of chromosomes and the term is usually only applied to cells or organisms that are normally diploid. The more general term for such organisms is haploid.

Temporal terms

Neopolyploidy

A polyploid that is newly formed.

Mesopolyploidy

That has become polyploid in more recent history; it is not as new as a neopolyploid and not as old as a paleopolyploid. It is a middle aged polyploid. Often this refers to whole genome duplication followed by intermediate levels of diploidization.

Paleopolyploidy

 
This phylogenetic tree shows the relationship between the best-documented instances of paleopolyploidy in eukaryotes.
Main article: Paleopolyploidy

Ancient genome duplications probably occurred in the evolutionary history of all life. Duplication events that occurred long ago in the history of various evolutionary lineages can be difficult to detect because of subsequent diploidization (such that a polyploid starts to behave cytogenetically as a diploid over time) as mutations and gene translations gradually make one copy of each chromosome unlike the other copy. Over time, it is also common for duplicated copies of genes to accumulate mutations and become inactive pseudogenes.[38]

In many cases, these events can be inferred only through comparing sequenced genomes. Examples of unexpected but recently confirmed ancient genome duplications include baker's yeast (Saccharomyces cerevisiae), mustard weed/thale cress (Arabidopsis thaliana), rice (Oryza sativa), and an early evolutionary ancestor of the vertebrates (which includes the human lineage) and another near the origin of the teleost fishes.[39] Angiosperms (flowering plants) have paleopolyploidy in their ancestry. All eukaryotes probably have experienced a polyploidy event at some point in their evolutionary history.

Other similar terms

Karyotype

Main article: Karyotype

A karyotype is the characteristic chromosome complement of a eukaryote species.[40][41] The preparation and study of karyotypes is part of cytology and, more specifically, cytogenetics.

Although the replication and transcription of DNA is highly standardized in eukaryotes, the same cannot be said for their karyotypes, which are highly variable between species in chromosome number and in detailed organization despite being constructed out of the same macromolecules. In some cases, there is even significant variation within species. This variation provides the basis for a range of studies in what might be called evolutionary cytology.

Homoeologous chromosomes

Main article: Homoeology

Homoeologous chromosomes are those brought together following inter-species hybridization and allopolyploidization, and whose relationship was completely homologous in an ancestral species. For example, durum wheat is the result of the inter-species hybridization of two diploid grass species Triticum urartu and Aegilops speltoides. Both diploid ancestors had two sets of 7 chromosomes, which were similar in terms of size and genes contained on them. Durum wheat contains a hybrid genome with two sets of chromosomes derived from Triticum urartu and two sets of chromosomes derived from Aegilops speltoides. Each chromosome pair derived from the Triticum urartu parent is homoeologous to the opposite chromosome pair derived from the Aegilops speltoides parent, though each chromosome pair unto itself is homologous.

Examples

Animals

Examples in animals are more common in non-vertebrates[42] such as flatworms, leeches, and brine shrimp. Within vertebrates, examples of stable polyploidy include the salmonids and many cyprinids (i.e. carp).[43] Some fish have as many as 400 chromosomes.[43] Polyploidy also occurs commonly in amphibians; for example the biomedically important genus Xenopus contains many different species with as many as 12 sets of chromosomes (dodecaploid).[44] Polyploid lizards are also quite common. Most are sterile and reproduce by parthenogenesis;[citation needed] others, like Liolaemus chiliensis, maintain sexual reproduction. Polyploid mole salamanders (mostly triploids) are all female and reproduce by kleptogenesis,[45] "stealing" spermatophores from diploid males of related species to trigger egg development but not incorporating the males' DNA into the offspring.

While mammalian liver cells are polyploid, rare instances of polyploid mammals are known, but most often result in prenatal death. An octodontid rodent of Argentina's harsh desert regions, known as the plains viscacha rat (Tympanoctomys barrerae) has been reported as an exception to this 'rule'.[46] However, careful analysis using chromosome paints shows that there are only two copies of each chromosome in T. barrerae, not the four expected if it were truly a tetraploid.[47] This rodent is not a rat, but kin to guinea pigs and chinchillas. Its "new" diploid (2n) number is 102 and so its cells are roughly twice normal size. Its closest living relation is Octomys mimax, the Andean Viscacha-Rat of the same family, whose 2n = 56. It was therefore surmised that an Octomys-like ancestor produced tetraploid (i.e., 2n = 4x = 112) offspring that were, by virtue of their doubled chromosomes, reproductively isolated from their parents.

Polyploidy was induced in fish by Har Swarup (1956) using a cold-shock treatment of the eggs close to the time of fertilization, which produced triploid embryos that successfully matured.[48][49] Cold or heat shock has also been shown to result in unreduced amphibian gametes, though this occurs more commonly in eggs than in sperm.[50] John Gurdon (1958) transplanted intact nuclei from somatic cells to produce diploid eggs in the frog, Xenopus (an extension of the work of Briggs and King in 1952) that were able to develop to the tadpole stage.[51] The British scientist J. B. S. Haldane hailed the work for its potential medical applications and, in describing the results, became one of the first to use the word "clone" in reference to animals. Later work by Shinya Yamanaka showed how mature cells can be reprogrammed to become pluripotent, extending the possibilities to non-stem cells. Gurdon and Yamanaka were jointly awarded the Nobel Prize in 2012 for this work.[51]

Humans

 
Schematic karyogram of a human, showing the normal diploid (that is, non-polyploid) karyotype. It shows 22 homologous chromosomes, both the female (XX) and male (XY) versions of the sex chromosome (bottom right), as well as the mitochondrial genome (to scale at bottom left).
Further information: Karyotype
Further information: Triploid syndrome

True polyploidy rarely occurs in humans, although polyploid cells occur in highly differentiated tissue, such as liver parenchyma, heart muscle, placenta and in bone marrow.[2][52] Aneuploidy is more common.

Polyploidy occurs in humans in the form of triploidy, with 69 chromosomes (sometimes called 69, XXX), and tetraploidy with 92 chromosomes (sometimes called 92, XXXX). Triploidy, usually due to polyspermy, occurs in about 2–3% of all human pregnancies and ~15% of miscarriages.[citation needed] The vast majority of triploid conceptions end as a miscarriage; those that do survive to term typically die shortly after birth. In some cases, survival past birth may be extended if there is mixoploidy with both a diploid and a triploid cell population present. There has been one report of a child surviving to the age of seven months with complete triploidy syndrome. He failed to exhibit normal mental or physical neonatal development, and died from a Pneumocystis carinii infection, which indicates a weak immune system.[53]

Triploidy may be the result of either digyny (the extra haploid set is from the mother) or diandry (the extra haploid set is from the father). Diandry is mostly caused by reduplication of the paternal haploid set from a single sperm, but may also be the consequence of dispermic (two sperm) fertilization of the egg.[54] Digyny is most commonly caused by either failure of one meiotic division during oogenesis leading to a diploid oocyte or failure to extrude one polar body from the oocyte. Diandry appears to predominate among early miscarriages, while digyny predominates among triploid zygotes that survive into the fetal period.[55] However, among early miscarriages, digyny is also more common in those cases less than 8+1⁄2 weeks gestational age or those in which an embryo is present. There are also two distinct phenotypes in triploid placentas and fetuses that are dependent on the origin of the extra haploid set. In digyny, there is typically an asymmetric poorly grown fetus, with marked adrenal hypoplasia and a very small placenta.[citation needed] In diandry, a partial hydatidiform mole develops.[54] These parent-of-origin effects reflect the effects of genomic imprinting.[citation needed]

Complete tetraploidy is more rarely diagnosed than triploidy, but is observed in 1–2% of early miscarriages. However, some tetraploid cells are commonly found in chromosome analysis at prenatal diagnosis and these are generally considered 'harmless'. It is not clear whether these tetraploid cells simply tend to arise during in vitro cell culture or whether they are also present in placental cells in vivo. There are, at any rate, very few clinical reports of fetuses/infants diagnosed with tetraploidy mosaicism.

Mixoploidy is quite commonly observed in human preimplantation embryos and includes haploid/diploid as well as diploid/tetraploid mixed cell populations. It is unknown whether these embryos fail to implant and are therefore rarely detected in ongoing pregnancies or if there is simply a selective process favoring the diploid cells.

Fish

A polyploidy event occurred within the stem lineage of the teleost fish.[39]

Plants

 
Speciation via polyploidy: A diploid cell undergoes failed meiosis, producing diploid gametes, which self-fertilize to produce a tetraploid zygote.

Polyploidy is frequent in plants, some estimates suggesting that 30–80% of living plant species are polyploid, and many lineages show evidence of ancient polyploidy (paleopolyploidy) in their genomes.[56][57][58][59] Huge explosions in angiosperm species diversity appear to have coincided with the timing of ancient genome duplications shared by many species.[60] It has been established that 15% of angiosperm and 31% of fern speciation events are accompanied by ploidy increase.[61]

Polyploid plants can arise spontaneously in nature by several mechanisms, including meiotic or mitotic failures, and fusion of unreduced (2n) gametes.[62] Both autopolyploids (e.g. potato[63]) and allopolyploids (such as canola, wheat and cotton) can be found among both wild and domesticated plant species.

Most polyploids display novel variation or morphologies relative to their parental species, that may contribute to the processes of speciation and eco-niche exploitation.[57][62] The mechanisms leading to novel variation in newly formed allopolyploids may include gene dosage effects (resulting from more numerous copies of genome content), the reunion of divergent gene regulatory hierarchies, chromosomal rearrangements, and epigenetic remodeling, all of which affect gene content and/or expression levels.[64][65][66][67] Many of these rapid changes may contribute to reproductive isolation and speciation. However seed generated from interploidy crosses, such as between polyploids and their parent species, usually have aberrant endosperm development which impairs their viability,[68][69] thus contributing to polyploid speciation.

Some plants are triploid. As meiosis is disturbed, these plants are sterile, with all plants having the same genetic constitution: Among them, the exclusively vegetatively propagated saffron crocus (Crocus sativus). Also, the extremely rare Tasmanian shrub Lomatia tasmanica is a triploid sterile species.

There are few naturally occurring polyploid conifers. One example is the Coast Redwood Sequoia sempervirens, which is a hexaploid (6x) with 66 chromosomes (2n = 6x = 66), although the origin is unclear.[70]

Aquatic plants, especially the Monocotyledons, include a large number of polyploids.[71]

Crops

The induction of polyploidy is a common technique to overcome the sterility of a hybrid species during plant breeding. For example, triticale is the hybrid of wheat (Triticum turgidum) and rye (Secale cereale). It combines sought-after characteristics of the parents, but the initial hybrids are sterile. After polyploidization, the hybrid becomes fertile and can thus be further propagated to become triticale.

In some situations, polyploid crops are preferred because they are sterile. For example, many seedless fruit varieties are seedless as a result of polyploidy. Such crops are propagated using asexual techniques, such as grafting.

Polyploidy in crop plants is most commonly induced by treating seeds with the chemical colchicine.

Examples

Some crops are found in a variety of ploidies: tulips and lilies are commonly found as both diploid and triploid; daylilies (Hemerocallis cultivars) are available as either diploid or tetraploid; apples and kinnow mandarins can be diploid, triploid, or tetraploid.

Fungi

Besides plants and animals, the evolutionary history of various fungal species is dotted by past and recent whole-genome duplication events (see Albertin and Marullo 2012[75] for review). Several examples of polyploids are known:

In addition, polyploidy is frequently associated with hybridization and reticulate evolution that appear to be highly prevalent in several fungal taxa. Indeed, homoploid speciation (hybrid speciation without a change in chromosome number) has been evidenced for some fungal species (such as the basidiomycota Microbotryum violaceum[83]).

 
Schematic phylogeny of the Chromalveolata. Red circles indicate polyploidy, blue squares indicate hybridization. From Albertin and Marullo, 2012[75]

As for plants and animals, fungal hybrids and polyploids display structural and functional modifications compared to their progenitors and diploid counterparts. In particular, the structural and functional outcomes of polyploid Saccharomyces genomes strikingly reflect the evolutionary fate of plant polyploid ones. Large chromosomal rearrangements[84] leading to chimeric chromosomes[85] have been described, as well as more punctual genetic modifications such as gene loss.[86] The homoealleles of the allotetraploid yeast S. pastorianus show unequal contribution to the transcriptome.[87] Phenotypic diversification is also observed following polyploidization and/or hybridization in fungi,[88] producing the fuel for natural selection and subsequent adaptation and speciation.

Chromalveolata

Other eukaryotic taxa have experienced one or more polyploidization events during their evolutionary history (see Albertin and Marullo, 2012[75] for review). The oomycetes, which are non-true fungi members, contain several examples of paleopolyploid and polyploid species, such as within the genus Phytophthora.[89] Some species of brown algae (Fucales, Laminariales[90] and diatoms[91]) contain apparent polyploid genomes. In the Alveolata group, the remarkable species Paramecium tetraurelia underwent three successive rounds of whole-genome duplication[92] and established itself as a major model for paleopolyploid studies.

Bacteria

Each Deinococcus radiodurans bacterium contains 4-8 copies of its chromosome.[93] Exposure of D. radiodurans to X-ray irradiation or desiccation can shatter its genomes into hundred of short random fragments. Nevertheless, D. radiodurans is highly resistant to such exposures. The mechanism by which the genome is accurately restored involves RecA-mediated homologous recombination and a process referred to as extended synthesis-dependent strand annealing (SDSA).[94]

Azotobacter vinelandii can contain up to 80 chromosome copies per cell.[95] However this is only observed in fast growing cultures, whereas cultures grown in synthetic minimal media are not polyploid.[96]

Archaea

The archaeon Halobacterium salinarium is polyploid[97] and, like Deinococcus radiodurans, is highly resistant to X-ray irradiation and desiccation, conditions that induce DNA double-strand breaks.[98] Although chromosomes are shattered into many fragments, complete chromosomes can be regenerated by making use of overlapping fragments. The mechanism employs single-stranded DNA binding protein and is likely homologous recombinational repair.[99]

See also

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Further reading

  • Snustad, D. P.; et al. (2006). Principles of Genetics (4th ed.). Hoboken, New Jersey: John Wiley & Sons. ISBN 978-0-471-69939-2.
  • The Arabidopsis Genome Initiative (2000). "Analysis of the genome sequence of the flowering plant Arabidopsis thaliana". Nature. 408 (6814): 796–815. Bibcode:2000Natur.408..796T. doi:10.1038/35048692. PMID 11130711.
  • Eakin, G. S.; Behringer, R. R. (2003). "Tetraploid development in the mouse". Developmental Dynamics. 228 (4): 751–766. doi:10.1002/dvdy.10363. PMID 14648853.
  • Gaeta, R. T.; Pires, J. C.; Iniguez-Luy, F.; Leon, E.; Osborn, T. C. (2007). "Genomic Changes in Resynthesized Brassica napus and Their Effect on Gene Expression and Phenotype". The Plant Cell Online. 19 (11): 3403–3417. doi:10.1105/tpc.107.054346. PMC 2174891. PMID 18024568.
  • Gregory, T. R.; Mable, B. K. (2005). "Polyploidy in animals". In Gregory, T. R. (ed.). The Evolution of the Genome. San Diego, California: Elsevier. pp. 427–517.
  • Jaillon, O.; Aury, J.-M.; Brunet, F.; Petit, J.-L.; Stange-Thomann, N.; Mauceli, E.; Bouneau, L.; Fischer, C.; Ozouf-Costaz, C.; Bernot, A.; Nicaud, S.; Jaffe, D.; Fisher, S.; Lutfalla, G.; Dossat, C.; Segurens, B.; Dasilva, C.; Salanoubat, M.; Levy, M.; Boudet, N.; Castellano, S.; Anthouard, V.; Jubin, C.; Castelli, V.; Katinka, M.; Vacherie, B.; Biémont, C.; Skalli, Z.; Cattolico, L.; Poulain, J.; et al. (2004). "Genome duplication in the teleost fish Tetraodon nigroviridis reveals the early vertebrate proto-karyotype". Nature. 431 (7011): 946–957. Bibcode:2004Natur.431..946J. doi:10.1038/nature03025. PMID 15496914.
  • Paterson, A. H.; Bowers, J. E.; Van De Peer, Y.; Vandepoele, K. (2005). "Ancient duplication of cereal genomes". New Phytologist. 165 (3): 658–661. doi:10.1111/j.1469-8137.2005.01347.x. PMID 15720677.
  • Raes, J.; Vandepoele, K.; Simillion, C.; Saeys, Y.; Van De Peer, Y. (2003). "Investigating ancient duplication events in the Arabidopsis genome". Journal of Structural and Functional Genomics. 3 (1–4): 117–129. doi:10.1023/A:1022666020026. PMID 12836691. S2CID 9666357.
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External links

 
Look up mesopolyploid in Wiktionary, the free dictionary.
 
Look up neopolyploid in Wiktionary, the free dictionary.
  • Polyploidy on Kimball's Biology Pages
  • a community-editable project with information, research, education, and a bibliography about polyploidy.

February 26, 2023
polyploidy, confused, with, polypoid, resembling, polyp, condition, which, cells, organism, have, more, than, pair, homologous, chromosomes, most, species, whose, cells, have, nuclei, eukaryotes, diploid, meaning, they, have, complete, sets, chromosomes, from,. Not to be confused with polypoid resembling a polyp Polyploidy is a condition in which the cells of an organism have more than one pair of homologous chromosomes Most species whose cells have nuclei eukaryotes are diploid meaning they have two complete sets of chromosomes one from each of two parents each set contains the same number of chromosomes and the chromosomes are joined in pairs of homologous chromosomes However some organisms are polyploid Polyploidy is especially common in plants Most eukaryotes have diploid somatic cells but produce haploid gametes eggs and sperm by meiosis A monoploid has only one set of chromosomes and the term is usually only applied to cells or organisms that are normally diploid Males of bees and other Hymenoptera for example are monoploid Unlike animals plants and multicellular algae have life cycles with two alternating multicellular generations The gametophyte generation is haploid and produces gametes by mitosis the sporophyte generation is diploid and produces spores by meiosis This image shows haploid single diploid double triploid triple and tetraploid quadruple sets of chromosomes Triploid and tetraploid chromosomes are examples of polyploidy Polyploidy may occur due to abnormal cell division either during mitosis or more commonly from the failure of chromosomes to separate during meiosis or from the fertilization of an egg by more than one sperm 1 In addition it can be induced in plants and cell cultures by some chemicals the best known is colchicine which can result in chromosome doubling though its use may have other less obvious consequences as well Oryzalin will also double the existing chromosome content Polyploidy occurs in highly differentiated human tissues in the liver heart muscle bone marrow and the placenta 2 failed verification It occurs in the somatic cells of some animals such as goldfish 3 salmon and salamanders but is especially common among ferns and flowering plants see Hibiscus rosa sinensis including both wild and cultivated species Wheat for example after millennia of hybridization and modification by humans has strains that are diploid two sets of chromosomes tetraploid four sets of chromosomes with the common name of durum or macaroni wheat and hexaploid six sets of chromosomes with the common name of bread wheat Many agriculturally important plants of the genus Brassica are also tetraploids Sugarcane can have ploidy levels higher than octaploid 4 Polyploidization can be a mechanism of sympatric speciation because polyploids are usually unable to interbreed with their diploid ancestors An example is the plant Erythranthe peregrina Sequencing confirmed that this species originated from E robertsii a sterile triploid hybrid between E guttata and E lutea both of which have been introduced and naturalised in the United Kingdom New populations of E peregrina arose on the Scottish mainland and the Orkney Islands via genome duplication from local populations of E robertsii 5 Because of a rare genetic mutation E peregrina is not sterile 6 Contents 1 Terminology 1 1 Types 1 2 Classification 1 2 1 Autopolyploidy 1 2 2 Allopolyploidy 1 2 3 Aneuploid 1 2 4 Endopolyploidy 1 2 5 Monoploid 1 3 Temporal terms 1 3 1 Neopolyploidy 1 3 2 Mesopolyploidy 1 3 3 Paleopolyploidy 1 4 Other similar terms 1 4 1 Karyotype 1 4 2 Homoeologous chromosomes 2 Examples 2 1 Animals 2 1 1 Humans 2 1 2 Fish 2 2 Plants 2 2 1 Crops 2 2 1 1 Examples 2 3 Fungi 2 4 Chromalveolata 2 5 Bacteria 2 6 Archaea 3 See also 4 References 5 Further reading 6 External linksTerminology EditTypes Edit Triploid redirects here For the human chromosomal disorder 69 XXX etc see Triploid syndrome Organ specific patterns of endopolyploidy from 2x to 64x in the giant ant Dinoponera australisPolyploid types are labeled according to the number of chromosome sets in the nucleus The letter x is used to represent the number of chromosomes in a single set haploid one set 1x diploid two sets 2x triploid three sets 3x for example sterile saffron crocus or seedless watermelons also common in the phylum Tardigrada 7 tetraploid four sets 4x for example Plains viscacha rat Salmonidae fish 8 the cotton Gossypium hirsutum 9 pentaploid five sets 5x for example Kenai Birch Betula kenaica hexaploid six sets 6x for example some species of wheat 10 kiwifruit 11 heptaploid or septaploid seven sets 7x octaploid or octoploid eight sets 8x for example Acipenser genus of sturgeon fish dahlias decaploid ten sets 10x for example certain strawberries dodecaploid or duodecaploid twelve sets 12x for example the plants Celosia argentea and Spartina anglica 12 or the amphibian Xenopus ruwenzoriensis tetratetracontaploid fortyfour sets 44x for example black mulberry 13 Classification Edit Autopolyploidy Edit Autopolyploids are polyploids with multiple chromosome sets derived from a single taxon Two examples of natural autopolyploids are the piggyback plant Tolmiea menzisii 14 and the white sturgeon Acipenser transmontanum 15 Most instances of autopolyploidy result from the fusion of unreduced 2n gametes which results in either triploid n 2n 3n or tetraploid 2n 2n 4n offspring 16 Triploid offspring are typically sterile as in the phenomenon of triploid block but in some cases they may produce high proportions of unreduced gametes and thus aid the formation of tetraploids This pathway to tetraploidy is referred to as the triploid bridge 16 Triploids may also persist through asexual reproduction In fact stable autotriploidy in plants is often associated with apomictic mating systems 17 In agricultural systems autotriploidy can result in seedlessness as in watermelons and bananas 18 Triploidy is also utilized in salmon and trout farming to induce sterility 19 20 Rarely autopolyploids arise from spontaneous somatic genome doubling which has been observed in apple Malus domesticus bud sports 21 This is also the most common pathway of artificially induced polyploidy where methods such as protoplast fusion or treatment with colchicine oryzalin or mitotic inhibitors are used to disrupt normal mitotic division which results in the production of polyploid cells This process can be useful in plant breeding especially when attempting to introgress germplasm across ploidal levels 22 Autopolyploids possess at least three homologous chromosome sets which can lead to high rates of multivalent pairing during meiosis particularly in recently formed autopolyploids also known as neopolyploids and an associated decrease in fertility due to the production of aneuploid gametes 23 Natural or artificial selection for fertility can quickly stabilize meiosis in autopolyploids by restoring bivalent pairing during meiosis but the high degree of homology among duplicated chromosomes causes autopolyploids to display polysomic inheritance 24 This trait is often used as a diagnostic criterion to distinguish autopolyploids from allopolyploids which commonly display disomic inheritance after they progress past the neopolyploid stage 25 While most polyploid species are unambiguously characterized as either autopolyploid or allopolyploid these categories represent the ends of a spectrum of divergence between parental subgenomes Polyploids that fall between these two extremes which are often referred to as segmental allopolyploids may display intermediate levels of polysomic inheritance that vary by locus 26 27 About half of all polyploids are thought to be the result of autopolyploidy 28 29 although many factors make this proportion hard to estimate 30 Allopolyploidy Edit Allopolyploids or amphipolyploids or heteropolyploids are polyploids with chromosomes derived from two or more diverged taxa As in autopolyploidy this primarily occurs through the fusion of unreduced 2n gametes which can take place before or after hybridization In the former case unreduced gametes from each diploid taxon or reduced gametes from two autotetraploid taxa combine to form allopolyploid offspring In the latter case one or more diploid F1 hybrids produce unreduced gametes that fuse to form allopolyploid progeny 31 Hybridization followed by genome duplication may be a more common path to allopolyploidy because F1 hybrids between taxa often have relatively high rates of unreduced gamete formation divergence between the genomes of the two taxa result in abnormal pairing between homoeologous chromosomes or nondisjunction during meiosis 31 In this case allopolyploidy can actually restore normal bivalent meiotic pairing by providing each homoeologous chromosome with its own homologue If divergence between homoeologous chromosomes is even across the two subgenomes this can theoretically result in rapid restoration of bivalent pairing and disomic inheritance following allopolyploidization However multivalent pairing is common in many recently formed allopolyploids so it is likely that the majority of meiotic stabilization occurs gradually through selection 23 25 Because pairing between homoeologous chromosomes is rare in established allopolyploids they may benefit from fixed heterozygosity of homoeologous alleles 32 In certain cases such heterozygosity can have beneficial heterotic effects either in terms of fitness in natural contexts or desirable traits in agricultural contexts This could partially explain the prevalence of allopolyploidy among crop species Both bread wheat and Triticale are examples of an allopolyploids with six chromosome sets Cotton peanut or quinoa are allotetraploids with multiple origins In Brassicaceous crops the Triangle of U describes the relationships between the three common diploid Brassicas B oleracea B rapa and B nigra and three allotetraploids B napus B juncea and B carinata derived from hybridization among the diploid species A similar relationship exists between three diploid species of Tragopogon T dubius T pratensis and T porrifolius and two allotetraploid species T mirus and T miscellus 33 Complex patterns of allopolyploid evolution have also been observed in animals as in the frog genus Xenopus 34 Aneuploid Edit Main article Aneuploidy Organisms in which a particular chromosome or chromosome segment is under or over represented are said to be aneuploid from the Greek words meaning not good and fold Aneuploidy refers to a numerical change in part of the chromosome set whereas polyploidy refers to a numerical change in the whole set of chromosomes 35 Endopolyploidy Edit Polyploidy occurs in some tissues of animals that are otherwise diploid such as human muscle tissues 36 This is known as endopolyploidy Species whose cells do not have nuclei that is prokaryotes may be polyploid as seen in the large bacterium Epulopiscium fishelsoni 37 Hence ploidy is defined with respect to a cell Monoploid Edit Main article Monoploidy A monoploid has only one set of chromosomes and the term is usually only applied to cells or organisms that are normally diploid The more general term for such organisms is haploid Temporal terms Edit Neopolyploidy Edit A polyploid that is newly formed Mesopolyploidy Edit That has become polyploid in more recent history it is not as new as a neopolyploid and not as old as a paleopolyploid It is a middle aged polyploid Often this refers to whole genome duplication followed by intermediate levels of diploidization Paleopolyploidy Edit This phylogenetic tree shows the relationship between the best documented instances of paleopolyploidy in eukaryotes Main article Paleopolyploidy Ancient genome duplications probably occurred in the evolutionary history of all life Duplication events that occurred long ago in the history of various evolutionary lineages can be difficult to detect because of subsequent diploidization such that a polyploid starts to behave cytogenetically as a diploid over time as mutations and gene translations gradually make one copy of each chromosome unlike the other copy Over time it is also common for duplicated copies of genes to accumulate mutations and become inactive pseudogenes 38 In many cases these events can be inferred only through comparing sequenced genomes Examples of unexpected but recently confirmed ancient genome duplications include baker s yeast Saccharomyces cerevisiae mustard weed thale cress Arabidopsis thaliana rice Oryza sativa and an early evolutionary ancestor of the vertebrates which includes the human lineage and another near the origin of the teleost fishes 39 Angiosperms flowering plants have paleopolyploidy in their ancestry All eukaryotes probably have experienced a polyploidy event at some point in their evolutionary history Other similar terms Edit Karyotype Edit Main article Karyotype A karyotype is the characteristic chromosome complement of a eukaryote species 40 41 The preparation and study of karyotypes is part of cytology and more specifically cytogenetics Although the replication and transcription of DNA is highly standardized in eukaryotes the same cannot be said for their karyotypes which are highly variable between species in chromosome number and in detailed organization despite being constructed out of the same macromolecules In some cases there is even significant variation within species This variation provides the basis for a range of studies in what might be called evolutionary cytology Homoeologous chromosomes Edit Main article Homoeology Homoeologous chromosomes are those brought together following inter species hybridization and allopolyploidization and whose relationship was completely homologous in an ancestral species For example durum wheat is the result of the inter species hybridization of two diploid grass species Triticum urartu and Aegilops speltoides Both diploid ancestors had two sets of 7 chromosomes which were similar in terms of size and genes contained on them Durum wheat contains a hybrid genome with two sets of chromosomes derived from Triticum urartu and two sets of chromosomes derived from Aegilops speltoides Each chromosome pair derived from the Triticum urartu parent is homoeologous to the opposite chromosome pair derived from the Aegilops speltoides parent though each chromosome pair unto itself is homologous Examples EditAnimals Edit Examples in animals are more common in non vertebrates 42 such as flatworms leeches and brine shrimp Within vertebrates examples of stable polyploidy include the salmonids and many cyprinids i e carp 43 Some fish have as many as 400 chromosomes 43 Polyploidy also occurs commonly in amphibians for example the biomedically important genus Xenopus contains many different species with as many as 12 sets of chromosomes dodecaploid 44 Polyploid lizards are also quite common Most are sterile and reproduce by parthenogenesis citation needed others like Liolaemus chiliensis maintain sexual reproduction Polyploid mole salamanders mostly triploids are all female and reproduce by kleptogenesis 45 stealing spermatophores from diploid males of related species to trigger egg development but not incorporating the males DNA into the offspring While mammalian liver cells are polyploid rare instances of polyploid mammals are known but most often result in prenatal death An octodontid rodent of Argentina s harsh desert regions known as the plains viscacha rat Tympanoctomys barrerae has been reported as an exception to this rule 46 However careful analysis using chromosome paints shows that there are only two copies of each chromosome in T barrerae not the four expected if it were truly a tetraploid 47 This rodent is not a rat but kin to guinea pigs and chinchillas Its new diploid 2n number is 102 and so its cells are roughly twice normal size Its closest living relation is Octomys mimax the Andean Viscacha Rat of the same family whose 2n 56 It was therefore surmised that an Octomys like ancestor produced tetraploid i e 2n 4x 112 offspring that were by virtue of their doubled chromosomes reproductively isolated from their parents Polyploidy was induced in fish by Har Swarup 1956 using a cold shock treatment of the eggs close to the time of fertilization which produced triploid embryos that successfully matured 48 49 Cold or heat shock has also been shown to result in unreduced amphibian gametes though this occurs more commonly in eggs than in sperm 50 John Gurdon 1958 transplanted intact nuclei from somatic cells to produce diploid eggs in the frog Xenopus an extension of the work of Briggs and King in 1952 that were able to develop to the tadpole stage 51 The British scientist J B S Haldane hailed the work for its potential medical applications and in describing the results became one of the first to use the word clone in reference to animals Later work by Shinya Yamanaka showed how mature cells can be reprogrammed to become pluripotent extending the possibilities to non stem cells Gurdon and Yamanaka were jointly awarded the Nobel Prize in 2012 for this work 51 Humans Edit Schematic karyogram of a human showing the normal diploid that is non polyploid karyotype It shows 22 homologous chromosomes both the female XX and male XY versions of the sex chromosome bottom right as well as the mitochondrial genome to scale at bottom left Further information Karyotype Further information Triploid syndrome True polyploidy rarely occurs in humans although polyploid cells occur in highly differentiated tissue such as liver parenchyma heart muscle placenta and in bone marrow 2 52 Aneuploidy is more common Polyploidy occurs in humans in the form of triploidy with 69 chromosomes sometimes called 69 XXX and tetraploidy with 92 chromosomes sometimes called 92 XXXX Triploidy usually due to polyspermy occurs in about 2 3 of all human pregnancies and 15 of miscarriages citation needed The vast majority of triploid conceptions end as a miscarriage those that do survive to term typically die shortly after birth In some cases survival past birth may be extended if there is mixoploidy with both a diploid and a triploid cell population present There has been one report of a child surviving to the age of seven months with complete triploidy syndrome He failed to exhibit normal mental or physical neonatal development and died from a Pneumocystis carinii infection which indicates a weak immune system 53 Triploidy may be the result of either digyny the extra haploid set is from the mother or diandry the extra haploid set is from the father Diandry is mostly caused by reduplication of the paternal haploid set from a single sperm but may also be the consequence of dispermic two sperm fertilization of the egg 54 Digyny is most commonly caused by either failure of one meiotic division during oogenesis leading to a diploid oocyte or failure to extrude one polar body from the oocyte Diandry appears to predominate among early miscarriages while digyny predominates among triploid zygotes that survive into the fetal period 55 However among early miscarriages digyny is also more common in those cases less than 8 1 2 weeks gestational age or those in which an embryo is present There are also two distinct phenotypes in triploid placentas and fetuses that are dependent on the origin of the extra haploid set In digyny there is typically an asymmetric poorly grown fetus with marked adrenal hypoplasia and a very small placenta citation needed In diandry a partial hydatidiform mole develops 54 These parent of origin effects reflect the effects of genomic imprinting citation needed Complete tetraploidy is more rarely diagnosed than triploidy but is observed in 1 2 of early miscarriages However some tetraploid cells are commonly found in chromosome analysis at prenatal diagnosis and these are generally considered harmless It is not clear whether these tetraploid cells simply tend to arise during in vitro cell culture or whether they are also present in placental cells in vivo There are at any rate very few clinical reports of fetuses infants diagnosed with tetraploidy mosaicism Mixoploidy is quite commonly observed in human preimplantation embryos and includes haploid diploid as well as diploid tetraploid mixed cell populations It is unknown whether these embryos fail to implant and are therefore rarely detected in ongoing pregnancies or if there is simply a selective process favoring the diploid cells Fish Edit A polyploidy event occurred within the stem lineage of the teleost fish 39 Plants Edit Speciation via polyploidy A diploid cell undergoes failed meiosis producing diploid gametes which self fertilize to produce a tetraploid zygote Polyploidy is frequent in plants some estimates suggesting that 30 80 of living plant species are polyploid and many lineages show evidence of ancient polyploidy paleopolyploidy in their genomes 56 57 58 59 Huge explosions in angiosperm species diversity appear to have coincided with the timing of ancient genome duplications shared by many species 60 It has been established that 15 of angiosperm and 31 of fern speciation events are accompanied by ploidy increase 61 Polyploid plants can arise spontaneously in nature by several mechanisms including meiotic or mitotic failures and fusion of unreduced 2n gametes 62 Both autopolyploids e g potato 63 and allopolyploids such as canola wheat and cotton can be found among both wild and domesticated plant species Most polyploids display novel variation or morphologies relative to their parental species that may contribute to the processes of speciation and eco niche exploitation 57 62 The mechanisms leading to novel variation in newly formed allopolyploids may include gene dosage effects resulting from more numerous copies of genome content the reunion of divergent gene regulatory hierarchies chromosomal rearrangements and epigenetic remodeling all of which affect gene content and or expression levels 64 65 66 67 Many of these rapid changes may contribute to reproductive isolation and speciation However seed generated from interploidy crosses such as between polyploids and their parent species usually have aberrant endosperm development which impairs their viability 68 69 thus contributing to polyploid speciation Some plants are triploid As meiosis is disturbed these plants are sterile with all plants having the same genetic constitution Among them the exclusively vegetatively propagated saffron crocus Crocus sativus Also the extremely rare Tasmanian shrub Lomatia tasmanica is a triploid sterile species There are few naturally occurring polyploid conifers One example is the Coast Redwood Sequoia sempervirens which is a hexaploid 6x with 66 chromosomes 2n 6x 66 although the origin is unclear 70 Aquatic plants especially the Monocotyledons include a large number of polyploids 71 Crops Edit The induction of polyploidy is a common technique to overcome the sterility of a hybrid species during plant breeding For example triticale is the hybrid of wheat Triticum turgidum and rye Secale cereale It combines sought after characteristics of the parents but the initial hybrids are sterile After polyploidization the hybrid becomes fertile and can thus be further propagated to become triticale In some situations polyploid crops are preferred because they are sterile For example many seedless fruit varieties are seedless as a result of polyploidy Such crops are propagated using asexual techniques such as grafting Polyploidy in crop plants is most commonly induced by treating seeds with the chemical colchicine Examples Edit Triploid crops some apple varieties such as Belle de Boskoop Jonagold Mutsu Ribston Pippin banana citrus ginger watermelon 72 saffron crocus white pulp of coconut Tetraploid crops very few apple varieties durum or macaroni wheat cotton potato canola rapeseed leek tobacco peanut kinnow Pelargonium Hexaploid crops chrysanthemum bread wheat triticale oat kiwifruit 11 Octaploid crops strawberry dahlia pansies sugar cane oca Oxalis tuberosa 73 Dodecaploid crops some sugar cane hybrids 74 Some crops are found in a variety of ploidies tulips and lilies are commonly found as both diploid and triploid daylilies Hemerocallis cultivars are available as either diploid or tetraploid apples and kinnow mandarins can be diploid triploid or tetraploid Fungi Edit Besides plants and animals the evolutionary history of various fungal species is dotted by past and recent whole genome duplication events see Albertin and Marullo 2012 75 for review Several examples of polyploids are known autopolyploid the aquatic fungi of genus Allomyces 76 some Saccharomyces cerevisiae strains used in bakery 77 etc allopolyploid the widespread Cyathus stercoreus 78 the allotetraploid lager yeast Saccharomyces pastorianus 79 the allotriploid wine spoilage yeast Dekkera bruxellensis 80 etc paleopolyploid the human pathogen Rhizopus oryzae 81 the genus Saccharomyces 82 etc In addition polyploidy is frequently associated with hybridization and reticulate evolution that appear to be highly prevalent in several fungal taxa Indeed homoploid speciation hybrid speciation without a change in chromosome number has been evidenced for some fungal species such as the basidiomycota Microbotryum violaceum 83 Schematic phylogeny of the Chromalveolata Red circles indicate polyploidy blue squares indicate hybridization From Albertin and Marullo 2012 75 As for plants and animals fungal hybrids and polyploids display structural and functional modifications compared to their progenitors and diploid counterparts In particular the structural and functional outcomes of polyploid Saccharomyces genomes strikingly reflect the evolutionary fate of plant polyploid ones Large chromosomal rearrangements 84 leading to chimeric chromosomes 85 have been described as well as more punctual genetic modifications such as gene loss 86 The homoealleles of the allotetraploid yeast S pastorianus show unequal contribution to the transcriptome 87 Phenotypic diversification is also observed following polyploidization and or hybridization in fungi 88 producing the fuel for natural selection and subsequent adaptation and speciation Chromalveolata Edit Other eukaryotic taxa have experienced one or more polyploidization events during their evolutionary history see Albertin and Marullo 2012 75 for review The oomycetes which are non true fungi members contain several examples of paleopolyploid and polyploid species such as within the genus Phytophthora 89 Some species of brown algae Fucales Laminariales 90 and diatoms 91 contain apparent polyploid genomes In the Alveolata group the remarkable species Paramecium tetraurelia underwent three successive rounds of whole genome duplication 92 and established itself as a major model for paleopolyploid studies Bacteria Edit Each Deinococcus radiodurans bacterium contains 4 8 copies of its chromosome 93 Exposure of D radiodurans to X ray irradiation or desiccation can shatter its genomes into hundred of short random fragments Nevertheless D radiodurans is highly resistant to such exposures The mechanism by which the genome is accurately restored involves RecA mediated homologous recombination and a process referred to as extended synthesis dependent strand annealing SDSA 94 Azotobacter vinelandii can contain up to 80 chromosome copies per cell 95 However this is only observed in fast growing cultures whereas cultures grown in synthetic minimal media are not polyploid 96 Archaea Edit The archaeon Halobacterium salinarium is polyploid 97 and like Deinococcus radiodurans is highly resistant to X ray irradiation and desiccation conditions that induce DNA double strand breaks 98 Although chromosomes are shattered into many fragments complete chromosomes can be regenerated by making use of overlapping fragments The mechanism employs single stranded DNA binding protein and is likely homologous recombinational repair 99 See also EditDiploidization Eukaryote hybrid genome Ploidy Polyploid complex Polysomy Reciprocal silencing SympatryReferences Edit Solomon Eldra 2014 Solomon Martin Martin Berg BIOLOGY Cengage Learning p 344 ISBN 978 1285423586 a b Velicky P Meinhardt 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Van de Peer Y Meyer A 2005 Large scale gene and ancient genome duplications In Gregory T R ed The Evolution of the Genome San Diego California Elsevier pp 329 368 Wolfe K H Shields D C 1997 Molecular evidence for an ancient duplication of the entire yeast genome Nature 387 6634 708 713 Bibcode 1997Natur 387 708W doi 10 1038 42711 PMID 9192896 S2CID 4307263 Wolfe K H 2001 Yesterday s polyploids and the mystery of diploidization Nature Reviews Genetics 2 5 333 341 doi 10 1038 35072009 PMID 11331899 S2CID 20796914 External links Edit Look up mesopolyploid in Wiktionary the free dictionary Look up neopolyploid in Wiktionary the free dictionary Polyploidy on Kimball s Biology Pages The polyploidy portal a community editable project with information research education and a bibliography about polyploidy Retrieved from https en wikipedia org w index php title Polyploidy amp oldid 1139689569, wikipedia, wiki, book, books, library,

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