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Polyploidy

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321: 38: 1385: 715:. 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. 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. 298:. Each of these species experienced independent autopolyploidy events (within-species polyploidy, described below), which then enabled subsequent interspecies gene flow of adaptive alleles, in this case stabilising each young polyploid lineage. Such polyploidy-enabled adaptive introgression may allow polyploids at act as 'allelic sponges', whereby they accumulate cryptic genomic variation that may be recruited upon encountering later environmental challenges. 8874: 2039: 987: 8898: 1597: 8910: 8886: 1133:, 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. 772:
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
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gametes. Natural or artificial selection for fertility can quickly stabilize meiosis in autopolyploids by restoring bivalent pairing during meiosis. Rapid adaptive evolution of the meiotic machinery, resulting in reduced levels of multivalents (and therefore stable autopolyploid meiosis) has been
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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
1354:(1956) using a cold-shock treatment of the eggs close to the time of fertilization, which produced triploid embryos that successfully matured. Cold or heat shock has also been shown to result in unreduced amphibian gametes, though this occurs more commonly in eggs than in sperm. 289:
On the other hand, polyploidization can also be a mechanism for a kind of 'reverse speciation', whereby gene flow is enabled following the polyploidy event, even between lineages that previously experienced no gene flow as diploids. This has been detailed at the genomic level in
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and eco-niche exploitation. 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
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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.
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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
76:, 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 6713:
Minato T, Yoshida S, Ishiguro T, Shimada E, Mizutani S, Kobayashi O, Yoshimoto H (March 2009). "Expression profiling of the bottom fermenting yeast Saccharomyces pastorianus orthologous genes using oligonucleotide microarrays".
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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
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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
905:(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. 150:, 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. In addition, it can be induced in plants and 1020:
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.
1749:). 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. 748:. 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 2204:
double-strand breaks. Although chromosomes are shattered into many fragments, complete chromosomes can be regenerated by making use of overlapping fragments. The mechanism employs single-stranded
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Devier B, Aguileta G, Hood ME, Giraud T (2009). "Using phylogenies of pheromone receptor genes in the Microbotryum violaceum species complex to investigate possible speciation by hybridization".
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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
6067:"Diploid/polyploid syntenic shuttle mapping and haplotype-specific chromosome walking toward a rust resistance gene (Bru1) in highly polyploid sugarcane (2n approximately 12x approximately 115)" 1376:
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.
7253: 1481:(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) 3706:
Marburger, Sarah; Monnahan, Patrick; Seear, Paul J.; Martin, Simon H.; Koch, Jordan; Paajanen, Pirita; Bohutínská, Magdalena; Higgins, James D.; Schmickl, Roswitha; Yant, Levi (2019-11-18).
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Marburger, Sarah; Monnahan, Patrick; Seear, Paul J.; Martin, Simon H.; Koch, Jordan; Paajanen, Pirita; Bohutínská, Magdalena; Higgins, James D.; Schmickl, Roswitha; Yant, Levi (2019-11-18).
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Marburger, Sarah; Monnahan, Patrick; Seear, Paul J.; Martin, Simon H.; Koch, Jordan; Paajanen, Pirita; Bohutínská, Magdalena; Higgins, James D.; Schmickl, Roswitha; Yant, Levi (2019-11-18).
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remodeling, all of which affect gene content and/or expression levels. Many of these rapid changes may contribute to reproductive isolation and speciation. However, seed generated from
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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
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This image shows haploid (single), diploid (double), triploid (triple), and tetraploid (quadruple) sets of chromosomes. Triploid and tetraploid chromosomes are examples of polyploidy.
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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.
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species diversity appear to have coincided with the timing of ancient genome duplications shared by many species. It has been established that 15% of angiosperm and 31% of fern
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can contain up to 80 chromosome copies per cell. However this is only observed in fast growing cultures, whereas cultures grown in synthetic minimal media are not polyploid.
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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
8785: 8585: 629:), 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 760:
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
6344:"Insights into the Dekkera bruxellensis genomic landscape: comparative genomics reveals variations in ploidy and nutrient utilisation potential amongst wine isolates" 1623:
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 (
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multivalent pairing is common in many recently formed allopolyploids, so it is likely that the majority of meiotic stabilization occurs gradually through selection.
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produce unreduced gametes that fuse to form allopolyploid progeny. Hybridization followed by genome duplication may be a more common path to allopolyploidy because F
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Seear, Paul J.; France, Martin G.; Gregory, Catherine L.; Heavens, Darren; Schmickl, Roswitha; Yant, Levi; Higgins, James D. (2020-07-15). Grelon, Mathilde (ed.).
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Zahradka K, Slade D, Bailone A, Sommer S, Averbeck D, Petranovic M, et al. (October 2006). "Reassembly of shattered chromosomes in Deinococcus radiodurans".
8832: 8532: 1301:) has been reported as an exception to this 'rule'. However, careful analysis using chromosome paints shows that there are only two copies of each chromosome in 1241:
contains many different species with as many as 12 sets of chromosomes (dodecaploid). Polyploid lizards are also quite common. Most are sterile and reproduce by
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Manimekalai R, Suresh G, Govinda Kurup H, Athiappan S, Kandalam M (September 2020). "Role of NGS and SNP genotyping methods in sugarcane improvement programs".
1642:) gametes. Both autopolyploids (e.g. potato) and allopolyploids (such as canola, wheat and cotton) can be found among both wild and domesticated plant species. 8837: 7722: 8822: 7700:, Schemske DW, Hancock JF, Thompson JN, Husband BC, Judd WS (2007). "Autopolyploidy in Angiosperms: Have We Grossly Underestimated the Number of Species?". 4747: 8827: 6652:
Scannell DR, Byrne KP, Gordon JL, Wong S, Wolfe KH (March 2006). "Multiple rounds of speciation associated with reciprocal gene loss in polyploid yeasts".
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Ancient genome duplications probably occurred in the evolutionary history of all life. Duplication events that occurred long ago in the history of various
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Osborn TC, Pires JC, Birchler JA, Auger DL, Chen ZJ, Lee HS, et al. (March 2003). "Understanding mechanisms of novel gene expression in polyploids".
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Bogart JP, Bi K, Fu J, Noble DW, Niedzwiecki J (February 2007). "Unisexual salamanders (genus Ambystoma) present a new reproductive mode for eukaryotes".
169:, a high frequency of polyploid cells is found in organs such as the brain, liver, heart, and bone marrow. It also occurs in the somatic cells of other 1658:, such as between polyploids and their parent species, usually have aberrant endosperm development which impairs their viability, thus contributing to 2982:
Aïnouche ML, Fortune PM, Salmon A, Parisod C, Grandbastien MA, Fukunaga K, et al. (2008). "Hybridization, polyploidy and invasion: Lessons from
2431:"Analysis of Three Sugarcane Homo/Homeologous Regions Suggests Independent Polyploidization Events of Saccharum officinarum and Saccharum spontaneum" 320: 1956:
is dotted by past and recent whole-genome duplication events (see Albertin and Marullo 2012 for review). Several examples of polyploids are known:
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Winkelmann M, Pfitzer P, Schneider W (December 1987). "Significance of polyploidy in megakaryocytes and other cells in health and tumor disease".
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Yant, Levi; Hollister, Jesse D.; Wright, Kevin M.; Arnold, Brian J.; Higgins, James D.; Franklin, F. Chris H.; Bomblies, Kirsten (November 2013).
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is disturbed, these plants are sterile, with all plants having the same genetic constitution: Among them, the exclusively vegetatively propagated
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chromosomes have been described, as well as more punctual genetic modifications such as gene loss. The homoealleles of the allotetraploid yeast
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Schematic phylogeny of the Chromalveolata. Red circles indicate polyploidy, blue squares indicate hybridization. From Albertin and Marullo, 2012
1170:. 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 691:(particularly in recently formed autopolyploids, also known as neopolyploids) and an associated decrease in fertility due to the production of 8033: 3029: 1447:, with 69 chromosomes (sometimes called 69, XXX), and tetraploidy with 92 chromosomes (sometimes called 92, XXXX). Triploidy, usually due to 2482:"Speciation by genome duplication: Repeated origins and genomic composition of the recently formed allopolyploid species Mimulus peregrinus" 5986:
Les DH, Philbrick CT (1993). "Studies of hybridization and chromosome number variation in aquatic angiosperms: Evolutionary implications".
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Seear, Paul J.; France, Martin G.; Gregory, Catherine L.; Heavens, Darren; Schmickl, Roswitha; Yant, Levi; Higgins, James D. (2020-07-15).
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Ohno S, Muramoto J, Christian L, Atkin NB (1967). "Diploid-tetraploid relationship among old-world members of the fish family Cyprinidae".
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Polyploid plants can arise spontaneously in nature by several mechanisms, including meiotic or mitotic failures, and fusion of unreduced (2
1235:). Some fish have as many as 400 chromosomes. Polyploidy also occurs commonly in amphibians; for example the biomedically important genus 718:
About half of all polyploids are thought to be the result of autopolyploidy, although many factors make this proportion hard to estimate.
7304:"Extremely radiation-resistant mutants of a halophilic archaeon with increased single-stranded DNA-binding protein (RPA) gene expression" 3391:
Cotter D, O'Donovan V, O'Maoiléidigh N, Rogan G, Roche N, Wilkins NP (June 2000). "An evaluation of the use of triploid Atlantic salmon (
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Raes J, Vandepoele K, Simillion C, Saeys Y, Van de Peer Y (2003). "Investigating ancient duplication events in the Arabidopsis genome".
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von Wangenheim KH, Peterson HP (June 2004). "Aberrant endosperm development in interploidy crosses reveals a timer of differentiation".
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have experienced one or more polyploidization events during their evolutionary history (see Albertin and Marullo, 2012 for review). The
8914: 6759:"Adaptive response to DNA-damaging agents in natural Saccharomyces cerevisiae populations from "Evolution Canyon", Mt. Carmel, Israel" 6205:"Evidence for autotetraploidy associated with reproductive isolation in Saccharomyces cerevisiae: towards a new domesticated species" 3868:
Le Comber SC, Ainouche ML, Kovarik A, Leitch AR (April 2010). "Making a functional diploid: from polysomic to disomic inheritance".
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The induction of polyploidy is a common technique to overcome the sterility of a hybrid species during plant breeding. For example,
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Most polyploids display novel variation or morphologies relative to their parental species, that may contribute to the processes of
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Chepurnov VA, Mann DG, Vyverman W, Sabbe K, Danielidis DB (2002). "Sexual Reproduction, Mating System, and Protoplast Dynamics of
6818:"Genetic characterization of the natural hybrid species Phytophthora alni as inferred from nuclear and mitochondrial DNA analyses" 2120:
underwent three successive rounds of whole-genome duplication and established itself as a major model for paleopolyploid studies.
1455:; those that do survive to term typically die shortly after birth. In some cases, survival past birth may be extended if there is 8485: 8480: 2934:"The second highest chromosome count among vertebrates is observed in cultured sturgeon and is associated with genome plasticity" 1368:
hailed the work for its potential medical applications and, in describing the results, became one of the first to use the word "
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has only one set of chromosomes, and the term is usually only applied to cells or organisms that are normally diploid. Males of
2091:, which are non-true fungi members, contain several examples of paleopolyploid and polyploid species, such as within the genus 5003:"Nobel Prize in Physiology or Medicine 2012 Awarded for Discovery That Mature Cells Can Be Reprogrammed to Become Pluripotent" 3348:
Varoquaux F, Blanvillain R, Delseny M, Gallois P (June 2000). "Less is better: new approaches for seedless fruit production".
2530: 8522: 8163: 7376: 5151: 4247: 3274:"Gametes with the somatic chromosome number: mechanisms of their formation and role in the evolution of autopolyploid plants" 3087: 2823: 1451:, occurs in about 2–3% of all human pregnancies and ~15% of miscarriages. The vast majority of triploid conceptions end as a 8670: 6025: 1364:(an extension of the work of Briggs and King in 1952) that were able to develop to the tadpole stage. The British scientist 8595: 8565: 8527: 3071: 1485:
of the egg. Digyny is most commonly caused by either failure of one meiotic division during oogenesis leading to a diploid
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and these are generally considered 'harmless'. It is not clear whether these tetraploid cells simply tend to arise during
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from diploid males of related species to trigger egg development but not incorporating the males' DNA into the offspring.
2907: 8285: 7254:"Physiological responses of the halophilic archaeon Halobacterium sp. strain NRC1 to desiccation and gamma irradiation" 5875:"Co-occurrence of related asexual, but not sexual, lineages suggests that reproductive interference limits coexistence" 2279: 2161:
is highly resistant to such exposures. The mechanism by which the genome is accurately restored involves RecA-mediated
846:) derived from hybridization among the diploid species. A similar relationship exists between three diploid species of 2051:
genomes strikingly reflect the evolutionary fate of plant polyploid ones. Large chromosomal rearrangements leading to
8026: 5742:"Numerous and rapid nonstochastic modifications of gene products in newly synthesized Brassica napus allotetraploids" 4798:"Molecular cytogenetics and allotetraploidy in the red vizcacha rat, Tympanoctomys barrerae (Rodentia, Octodontidae)" 3920: 3522: 2889: 8674: 3959:
Ramsey J, Schemske DW (January 1998). "Pathways, Mechanisms, and Rates of Polyploid Formation in Flowering Plants".
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Because pairing between homoeologous chromosomes is rare in established allopolyploids, they may benefit from fixed
8302: 1662:. Polyploids may also interbreed with diploids and produce polyploid seeds, as observed in the agamic complexes of 146:
Polyploidy is the result of whole-genome duplication during the evolution of species. It may occur due to abnormal
4417:"Little evidence for enhanced phenotypic evolution in early teleosts relative to their living fossil sister group" 2067:
diversification is also observed following polyploidization and/or hybridization in fungi, producing the fuel for
227:(six sets of chromosomes) with the common name of bread wheat. Many agriculturally important plants of the genus 3025:"Phytopharmacological potential of different species of Morus alba and their bioactive phytochemicals: A review" 1347:= 112) offspring that were, by virtue of their doubled chromosomes, reproductively isolated from their parents. 901:
Organisms in which a particular chromosome, or chromosome segment, is under- or over-represented are said to be
248: because polyploids are usually unable to interbreed with their diploid ancestors. An example is the plant 3500:
Dwivedi SL, Upadhyaya HD, Stalker HT, Blair MW, Bertioli DJ, Nielen S, Ortiz R (January 2007). Janick J (ed.).
8148: 1569:. There are, at any rate, very few clinical reports of fetuses/infants diagnosed with tetraploidy mosaicism. 7522:"Genome duplication in the teleost fish Tetraodon nigroviridis reveals the early vertebrate proto-karyotype" 6817: 158:, which can result in chromosome doubling, though its use may have other less obvious consequences as well. 8019: 6168:
Emerson R, Wilson CM (1954). "Interspecific Hybrids and the Cytogenetics and Cytotaxonomy of Euallomyces".
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Phillips N, Kapraun DF, Gómez Garreta A, Ribera Siguan MA, Rull JL, Salvador Soler N, et al. (2011).
3773:"A novel allele of ASY3 is associated with greater meiotic stability in autotetraploid Arabidopsis lyrata" 2933: 2671:"A novel allele of ASY3 is associated with greater meiotic stability in autotetraploid Arabidopsis lyrata" 652:
Rarely, autopolyploids arise from spontaneous, somatic genome doubling, which has been observed in apple (
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Dermen H (May 1951). "Tetraploid and Diploid Adventitious Shoots: From a Giant Sport of McIntosh Apple".
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a community-editable project with information, research, education, and a bibliography about polyploidy.
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Van de Peer Y, Meyer A (2005). "Large-scale gene and ancient genome duplications". In Gregory TR (ed.).
6556:"Reconstruction of the genome origins and evolution of the hybrid lager yeast Saccharomyces pastorianus" 6393:
Ma LJ, Ibrahim AS, Skory C, Grabherr MG, Burger G, Butler M, et al. (July 2009). Madhani HD (ed.).
881:). Complex patterns of allopolyploid evolution have also been observed in animals, as in the frog genus 8791: 5464:
De Bodt S, Maere S, Van de Peer Y (November 2005). "Genome duplication and the origin of angiosperms".
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Ramsey J (January 1998). "Pathways, Mechanisms, and Rates of Polyploid Formation in Flowering Plants".
3218:"Neutral markers confirm the octoploid origin and reveal spontaneous autopolyploidy in white sturgeon, 120: 17: 4837:
Svartman M, Stone G, Stanyon R (April 2005). "Molecular cytogenetics discards polyploidy in mammals".
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Havelka, Miloš; Bytyutskyy, Dmytro; Symonová, Radka; Ráb, Petr; Flajšhans, Martin (11 February 2016).
8295: 1985: 1655: 5693:"Genetic and epigenetic mechanisms for gene expression and phenotypic variation in plant polyploids" 4522: 2429:
Vilela MM, Del Bem LE, Van Sluys MA, de Setta N, Kitajima JP, Cruz GM, et al. (February 2017).
8745: 8558: 8349: 8133: 7470:"Genomic changes in resynthesized Brassica napus and their effect on gene expression and phenotype" 3830:
Parisod C, Holderegger R, Brochmann C (April 2010). "Evolutionary consequences of autopolyploidy".
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Le Cunff L, Garsmeur O, Raboin LM, Pauquet J, Telismart H, Selvi A, et al. (September 2008).
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Jaillon O, Aury JM, Brunet F, Petit JL, Stange-Thomann N, Mauceli E, et al. (October 2004).
7013:"Multiplicity of genome equivalents in the radiation-resistant bacterium Micrococcus radiodurans" 6395:"Genomic analysis of the basal lineage fungus Rhizopus oryzae reveals a whole-genome duplication" 6283:
Libkind D, Hittinger CT, Valério E, Gonçalves C, Dover J, Johnston M, et al. (August 2011).
5002: 2226: 2196: 2130: 1171: 1147: 930: 660:. This is also the most common pathway of artificially induced polyploidy, where methods such as 8850: 8806: 8739: 8708: 8626: 8600: 8255: 8228: 6285:"Microbe domestication and the identification of the wild genetic stock of lager-brewing yeast" 4517: 2173: 2031: 1321:) number is 102 and so its cells are roughly twice normal size. Its closest living relation is 865: 704:, with specific adaptive alleles of these species shared between only the evolved polyploids. 544: 195: 5020:
Velicky P, Meinhardt G, Plessl K, Vondra S, Weiss T, Haslinger P, et al. (October 2018).
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Albertin W, Marullo P, Aigle M, Bourgais A, Bely M, Dillmann C, et al. (November 2009).
5708: 4373:"Gene and genome duplications: the impact of dosage-sensitivity on the fate of nuclear genes" 3554: 2813: 1551: 1401: 712: 684: 266:
both of which have been introduced and naturalised in the United Kingdom. New populations of
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and reticulate evolution that appear to be highly prevalent in several fungal taxa. Indeed,
37: 8698: 8510: 8435: 8420: 8250: 8198: 8173: 8113: 7918: 7648: 7533: 7403: 7315: 7069: 6975: 6964:"Global trends of whole-genome duplications revealed by the ciliate Paramecium tetraurelia" 6940: 6928: 6770: 6661: 6457: 6296: 5995: 5740:
Albertin W, Balliau T, Brabant P, Chèvre AM, Eber F, Malosse C, Thiellement H (June 2006).
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5022:"Genome amplification and cellular senescence are hallmarks of human placenta development" 4162:
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weeks gestational age or those in which an embryo is present. There are also two distinct
1358:(1958) transplanted intact nuclei from somatic cells to produce diploid eggs in the frog, 1092:
probably have experienced a polyploidy event at some point in their evolutionary history.
8: 8941: 8878: 8735: 8551: 8178: 6603:
Nakao Y, Kanamori T, Itoh T, Kodama Y, Rainieri S, Nakamura N, et al. (April 2009).
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2390: 2205: 1294: 1158:, and whose relationship was completely homologous in an ancestral species. For example, 1009: 769: 641:
mating systems. In agricultural systems, autotriploidy can result in seedlessness, as in
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Polyploidy in crop plants is most commonly induced by treating seeds with the chemical
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Polyploidy occurs in some tissues of animals that are otherwise diploid, such as human
708: 423: 7879: 7852: 7780: 7755: 7679: 7636: 7186: 7161: 7137: 7112: 7037: 7012: 6962:
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of homoeologous alleles. In certain cases, such heterozygosity can have beneficial
745: 538: 532: 208: 8000: 5140: 2398: 2312: 1428:
True polyploidy rarely occurs in humans, although polyploid cells occur in highly
311:"Triploid" redirects here. For the human chromosomal disorder (69 XXX, etc.), see 27:
Condition where cells of an organism have more than two paired sets of chromosomes
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Swarup H (1956). "Production of Heteroploidy in the Three-Spined Stickleback,
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is the result of the inter-species hybridization of two diploid grass species
740:
As in autopolyploidy, this primarily occurs through the fusion of unreduced (2
649:. Triploidy is also utilized in salmon and trout farming to induce sterility. 8930: 8858: 8193: 7869: 7392:"Analysis of the genome sequence of the flowering plant Arabidopsis thaliana" 3798: 3739: 3708:"Interspecific introgression mediates adaptation to whole genome duplication" 3674: 3643:"Interspecific introgression mediates adaptation to whole genome duplication" 3609: 3486: 3334: 3255: 3056: 2755: 2696: 2637: 2606:"Interspecific introgression mediates adaptation to whole genome duplication" 2582: 2060: 2027: 2006: 1870: 1786: 1718: 1482: 1323: 1260: 1256: 1119: 1055: 818: 809: 394: 364: 147: 81: 8011: 6883: 6868:"Estimates of nuclear DNA content in 98 species of brown algae (Phaeophyta)" 6620: 6309: 6050:". In Zeder MA, Decker-Walters D, Emshwiller E, Bradley D, Smith BD (eds.). 5525: 5414: 5334: 5185: 5168: 4441: 4338: 4280: 3395:
L.) in minimising the impact of escaped farmed salmon on wild populations".
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2798: 1305:, not the four expected if it were truly a tetraploid. This rodent is not a 1129:
Although the replication and transcription of DNA is highly standardized in
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during meiosis. In this case, allopolyploidy can actually restore normal,
80:. Polyploidy is especially common in plants. Most eukaryotes have diploid 8387: 8382: 8245: 8223: 8218: 8086: 7046: 6865: 5806: 5417:"One thousand plant transcriptomes and the phylogenomics of green plants" 3390: 2446: 2150: 1864: 1498: 1452: 1355: 1066: 922: 597:. Most instances of autopolyploidy result from the fusion of unreduced (2 559: 124: 104: 7756:"Genome duplication, a trait shared by 22000 species of ray-finned fish" 7746: 7713: 7546: 7521: 7081: 6988: 6963: 6673: 5788: 5584: 5567: 5534: 812:
describes the relationships between the three common diploid Brassicas (
737:
are polyploids with chromosomes derived from two or more diverged taxa.
8716: 8631: 8574: 8407: 8397: 8392: 8377: 8307: 8183: 8157: 8108: 8098: 8046: 7738: 7670: 7451: 7434: 7222: 6479: 6269: 6189: 5659: 5084: 4936: 4745: 4183: 3202: 2380: 2357: 2138: 2115: 2072: 2064: 2023: 1917: 1904: 1892: 1806: 1651: 1646: 1632: 1628: 1600: 1572: 1539: 1490: 1456: 1448: 1437: 1433: 1351: 1314: 1310: 1267: 1159: 1142: 1130: 1081: 947: 902: 896: 848: 692: 665: 642: 418: 401: 182: 155: 132: 69: 61: 7771: 7754:
Taylor JS, Braasch I, Frickey T, Meyer A, Van de Peer Y (March 2003).
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Gaeta RT, Pires JC, Iniguez-Luy F, Leon E, Osborn TC (November 2007).
6682: 5940: 5923: 4897: 4215: 4198: 4053: 4036: 4012: 3995: 2746: 2730:"Adaptive introgression: how polyploidy reshapes gene flow landscapes" 2729: 2573: 2557:"Adaptive introgression: how polyploidy reshapes gene flow landscapes" 2556: 2497: 2038: 986: 8359: 8103: 8081: 8062: 7969: 7416: 7391: 7328: 7303: 6727: 2110: 2088: 1940:
cultivars) are available as either diploid or tetraploid; apples and
1933: 1886: 1878: 1730: 1516: 1415: 1397: 1389: 1286: 1228: 1224: 1112: 1106: 1089: 1059: 926: 789: 781: 749: 492: 457: 234: 7634: 7252:
Kottemann M, Kish A, Iloanusi C, Bjork S, DiRuggiero J (June 2005).
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Ahuja MR, Neale DB (2002). "Origins of Polyploidy in Coast Redwood (
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Xu X, Pan S, Cheng S, Zhang B, Mu D, Ni P, et al. (July 2011).
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types are labeled according to the number of chromosome sets in the
8345: 8129: 7562: 6524: 6446:"Gene order evolution and paleopolyploidy in hemiascomycete yeasts" 6052:
Documenting Domestication: New Genetic and Archaeological Paradigms
2251: 2241: 2134: 1844: 1825: 1782: 1754: 1693: 1596: 1543: 1520: 1275: 1212: 1062: 1024:
In many cases, these events can be inferred only through comparing
1017: 831: 669: 638: 637:. In fact, stable autotriploidy in plants is often associated with 578:
are polyploids with multiple chromosome sets derived from a single
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Paterson AH, Bowers JE, Van de Peer Y, Vandepoele K (March 2005).
7162:"Changes of ploidy during the Azotobacter vinelandii growth cycle" 4960:"Genome duplication in amphibians and fish: an extended synthesis" 3347: 8781: 8616: 8515: 8369: 8280: 8275: 7385: 7209:
Soppa J (January 2011). "Ploidy and gene conversion in Archaea".
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cell culture or whether they are also present in placental cells
1536: 1528: 1478: 1460: 1393: 1369: 1360: 1271: 1237: 1115: 1074: 953: 883: 688: 687:
sets, which can lead to high rates of multivalent pairing during
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Otto SP, Whitton J (2000). "Polyploid incidence and evolution".
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2026:
number) has been evidenced for some fungal species (such as the
1952:
Besides plants and animals, the evolutionary history of various
1550:
develops. These parent-of-origin effects reflect the effects of
1223:. Within vertebrates, examples of stable polyploidy include the 994:
shows the relationship between the best-documented instances of
585:
Two examples of natural autopolyploids are the piggyback plant,
352:
is used to represent the number of chromosomes in a single set:
8425: 8312: 8076: 8072: 8067: 6757:
Lidzbarsky GA, Shkolnik T, Nevo E (June 2009). Idnurm A (ed.).
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Borneman AR, Zeppel R, Chambers PJ, Curtin CD (February 2014).
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Drauch Schreier A, Gille D, Mahardja B, May B (November 2011).
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Examples in animals are more common in non-vertebrates such as
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Barker MS, Arrigo N, Baniaga AE, Li Z, Levin DA (April 2016).
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Justin R (January 2002). "Neopolyploidy in Flowering Plants".
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set. In digyny, there is typically an asymmetric poorly grown
711:
among duplicated chromosomes causes autopolyploids to display
8543: 8490: 8475: 8470: 8465: 8460: 8455: 8450: 8445: 8440: 8415: 8353: 6064: 5415:
One Thousand Plant Transcriptomes Initiative (October 2019).
2479: 2146: 2098: 2084: 1971: 1925: 1874: 1828: 1821: 1817: 1774: 1734: 1532: 1524: 1328: 1216: 1111:
A karyotype is the characteristic chromosome complement of a
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Vallejo-Marín M, Buggs RJ, Cooley AM, Puzey JR (June 2015).
792:
are examples of an allopolyploids with six chromosome sets.
7695: 3829: 1929: 1848: 1584:
A polyploidy event occurred within the stem lineage of the
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Brancati F, Mingarelli R, Dallapiccola B (December 2003).
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Types of Polyploids: Their Classification and Significance
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Mendell JE, Clements KD, Choat JH, Angert ER (May 2008).
3770: 3575: 2931: 2668: 2201: 1882: 1742: 1306: 254:. Sequencing confirmed that this species originated from 100: 6510: 5606: 4037:"Double trouble: taxonomy and definitions of polyploidy" 3022: 1270:
liver cells, are polyploid, rare instances of polyploid
8533:
International System for Human Cytogenetic Nomenclature
6815: 6651: 6602: 5568:"Genome sequence and analysis of the tuber crop potato" 5463: 4795: 4482:(6th ed.). London: Chapman & Hall. p. 28. 3907:. Advances in Genetics. Vol. 1. pp. 403–429. 7809: 7720: 7519: 6756: 6392: 5837: 2157:
into hundred of short random fragments. Nevertheless,
2014:
In addition, polyploidy is frequently associated with
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Evolution; International Journal of Organic Evolution
4957: 3993: 2905: 2486:
Evolution; International Journal of Organic Evolution
1404:, both the female (XX) and male (XY) versions of the 1118:. The preparation and study of karyotypes is part of 7900:. San Diego, California: Elsevier. pp. 329–368. 7805:. San Diego, California: Elsevier. pp. 371–426. 7515:. San Diego, California: Elsevier. pp. 427–517. 6961: 5872: 5267:"On the abundance of polyploids in flowering plants" 5216:
Wick JB, Johnson KJ, O'Brien J, Wick MJ (May 2013).
4836: 4551: 4549: 4034: 3065: 2890:"Triticum - an overview | ScienceDirect Topics" 7110: 6816:Ioos R, Andrieux A, Marçais B, Frey P (July 2006). 5921: 4796:Gallardo MH, González CA, Cebrián I (August 2006). 4414: 3271: 3267: 3265: 3114:"Definition of Eight Mulberry Species in the Genus 2811: 1255:(mostly triploids) are all female and reproduce by 1194:parent, though each chromosome pair unto itself is 1054:), and two rounds of whole genome duplication (the 921:. Species whose cells do not have nuclei, that is, 219:(four sets of chromosomes) with the common name of 7364: 5873:Whitton J, Sears CJ, Maddison WP (December 2017). 5139: 4493:Stebbins GL (1950). "Chapter XII: The Karyotype". 4475: 2531:"Make Room for a New Bloom: New Flower Discovered" 162:will also double the existing chromosome content. 7721:Soltis DE, Buggs RJ, Doyle JJ, Soltis PS (2010). 7160:Maldonado R, Jiménez J, Casadesús J (July 1994). 7111:Nagpal P, Jafri S, Reddy MA, Das HK (June 1989). 5304: 5302: 4755:Nature Reviews. Gastroenterology & Hepatology 4546: 4231: 4196: 3863: 3861: 2805: 2297:"The origins and functions of hepatic polyploidy" 2294: 1932:are commonly found as both diploid and triploid; 1470:infection, which indicates a weak immune system. 1190:to the opposite chromosome pair derived from the 1012:can be difficult to detect because of subsequent 324:Organ-specific patterns of endopolyploidy (from 2 278:via genome duplication from local populations of 211:and modification by humans, has strains that are 30:Not to be confused with "polypoid", resembling a 8928: 7895: 7796: 7113:"Multiple chromosomes of Azotobacter vinelandii" 4415:Clarke JT, Lloyd GT, Friedman M (October 2016). 3262: 3111: 744:) gametes, which can take place before or after 6443: 6244:Lu BC (1964). "Polyploidy in the Basidiomycete 6118: 6054:. Berkeley, CA: University of California Press. 5565: 5308: 5137: 5133: 5131: 4643: 4612: 4262: 3118:by Internal Transcribed Spacer-Based Phylogeny" 1924:Some crops are found in a variety of ploidies: 1615:, which self-fertilize to produce a tetraploid 127:generation is haploid, and produces gametes by 7432: 7362: 5299: 4558:"A suggestion to the medical librarians. 1920" 4410: 4408: 4074: 4072: 3958: 3858: 3536: 3534: 2295:Zhang S, Lin YH, Tarlow B, Zhu H (June 2019). 1527:that are dependent on the origin of the extra 1477:(the extra haploid set is from the mother) or 1088:) have paleopolyploidy in their ancestry. All 804:are allotetraploids with multiple origins. In 107:, for example, are monoploid. Unlike animals, 8559: 8041: 8027: 7816:Journal of Structural and Functional Genomics 7600:Journal of Structural and Functional Genomics 7295: 7245: 7202: 7053: 6167: 6114: 6112: 6110: 5368:"The evolutionary consequences of polyploidy" 4919:Swarup H (1959). "Production of triploidy in 4830: 4114: 4112: 4110: 3068:Advances in Plant Breeding Strategies: Fruits 3030:Asian Pacific Journal of Tropical Biomedicine 2728:Schmickl, Roswitha; Yant, Levi (April 2021). 2555:Schmickl, Roswitha; Yant, Levi (April 2021). 2109:) contain apparent polyploid genomes. In the 1497:. Diandry appears to predominate among early 1436:, heart muscle, placenta and in bone marrow. 601:) gametes, which results in either triploid ( 7904: 7510: 7153: 7104: 6553: 5985: 5684: 5635: 5600: 5492: 5359: 5258: 5209: 5128: 4637: 4256: 4035:Doyle JJ, Sherman-Broyles S (January 2017). 3429: 2901: 2899: 2727: 2554: 1692:There are few naturally occurring polyploid 1683:). Also, the extremely rare Tasmanian shrub 1073:lineage) and another near the origin of the 244:Polyploidization can be a mechanism of  7850: 7723:"What we still don't know about polyploidy" 7004: 5922:Halabi K, Shafir A, Mayrose I (June 2023). 5264: 5013: 4958:Mable BK, Alexandrou MA, Taylor MI (2011). 4507: 4405: 4069: 3531: 3509:. John Wiley & Sons. pp. 179–230. 3272:Bretagnolle F, Thompson JD (January 1995). 3177:Soltis DE (1984-01-01). "Autopolyploidy in 2845: 2167:synthesis-dependent strand annealing (SDSA) 1635:events are accompanied by ploidy increase. 1443:Polyploidy occurs in humans in the form of 1339:-like ancestor produced tetraploid (i.e., 2 1136: 8566: 8552: 8034: 8020: 7810:Van de Peer Y, Taylor JS, Meyer A (2003). 6107: 6045: 5962: 5117:. National Organization for Rare Disorders 4562:Journal of the Medical Library Association 4497:. New York, NY: Columbia University Press. 4370: 4107: 1960:autopolyploid: the aquatic fungi of genus 1251:, maintain sexual reproduction. Polyploid 1174:with two sets of chromosomes derived from 7930: 7878: 7868: 7779: 7678: 7660: 7580: 7545: 7493: 7450: 7415: 7327: 7185: 7136: 7036: 6987: 6891: 6792: 6782: 6681: 6628: 6579: 6487: 6469: 6420: 6410: 6369: 6359: 6318: 6308: 6220: 6144: 6090: 5939: 5898: 5814: 5765: 5716: 5667: 5583: 5542: 5524: 5440: 5383: 5342: 5282: 5241: 5184: 5047: 5037: 4997: 4995: 4975: 4813: 4722: 4712: 4581: 4521: 4450: 4440: 4388: 4347: 4337: 4314:"Extreme polyploidy in a large bacterium" 4288: 4237: 4214: 4052: 4011: 3806: 3788: 3747: 3682: 3617: 3289: 3245: 3151: 3141: 3046: 2959: 2949: 2896: 2812:Stouder DJ, Bisson PA, Naiman RJ (1997). 2776: 2745: 2704: 2686: 2645: 2572: 2505: 2454: 2320: 1944:can be diploid, triploid, or tetraploid. 1372:" in reference to animals. Later work by 1178:and two sets of chromosomes derived from 925:, may be polyploid, as seen in the large 7433:Eakin GS, Behringer RR (December 2003). 6444:Wong S, Butler G, Wolfe KH (July 2002). 5709:10.1146/annurev.arplant.58.032806.103835 5169:"Recurrent triploidy of maternal origin" 4492: 4081:Annual Review of Ecology and Systematics 3961:Annual Review of Ecology and Systematics 3943: 3902: 3555:10.1146/annurev.ecolsys.33.010802.150437 3543:Annual Review of Ecology and Systematics 3312: 3106:This review and book cite this research. 2522: 2037: 1721:, include a large number of polyploids. 1595: 1383: 1335:= 56. It was therefore surmised that an 1331:Viscacha-Rat of the same family, whose 2 1182:. Each chromosome pair derived from the 985: 319: 36: 7565:"Ancient duplication of cereal genomes" 5309:Rieseberg LH, Willis JH (August 2007). 4686: 2906:Crowhurst RN, Whittaker D, Gardner RC. 2269: 2022:(hybrid speciation without a change in 1989:, the allotriploid wine spoilage yeast 1714:= 66), although the origin is unclear. 1627:) in their genomes. Huge explosions in 1266:While some tissues of mammals, such as 14: 8929: 7010: 6941:10.1046/j.1529-8817.2002.t01-1-01233.x 6022:"Seedless Fruits Make Others Needless" 5641: 4992: 4918: 4871: 4197:Schmid M, Evans BJ, Bogart JP (2015). 4161: 4078: 3540: 3464: 3176: 2165:and a process referred to as extended 1473:Triploidy may be the result of either 1095: 683:Autopolyploids possess at least three 282:. Because of a rare genetic mutation, 199:), including both wild and cultivated 8547: 8523:List of organisms by chromosome count 8015: 8001:Polyploidy on Kimball's Biology Pages 7955: 7435:"Tetraploid development in the mouse" 7208: 4555: 4473: 4263:Parmacek MS, Epstein JA (July 2009). 4118: 3479:10.1093/oxfordjournals.jhered.a106189 2528: 1150:are those brought together following 633:. Triploids may also persist through 154:by some chemicals: the best known is 121:alternating multicellular generations 8885: 7812:"Are all fishes ancient polyploids?" 5690: 5365: 3072:Springer International Publishing AG 1274:are known, but most often result in 258:, a sterile triploid hybrid between 8909: 7390:Genome Initiative (December 2000). 6554:Dunn B, Sherlock G (October 2008). 6119:Albertin W, Marullo P (July 2012). 5924:"PloiDB: The plant ploidy database" 4269:The New England Journal of Medicine 4242:. San Francisco, CA: W.H. Freeman. 4240:An Introduction to genetic analysis 2272:Solomon/Martin/Martin/Berg, Biology 1998:paleopolyploid: the human pathogen 1696:. One example is the Coast Redwood 938:is defined with respect to a cell. 237:can have ploidy levels higher than 135:generation is diploid and produces 56:have more than two paired sets of ( 24: 7905:Wolfe KH, Shields DC (June 1997). 7355: 6243: 5284:10.1111/j.0014-3820.2006.tb01198.x 5173:European Journal of Human Genetics 5138:Baker P, Monga A, Baker P (2006). 4574:10.1023/B:RFBF.0000033049.00668.fe 3327:10.1111/j.1601-5223.1936.tb03204.x 3291:10.1111/j.1469-8137.1995.tb03005.x 1350:Polyploidy was induced in fish by 981: 969:A polyploid that is newly formed. 869:) and two allotetraploid species ( 207:, for example, after millennia of 25: 8953: 7994: 5466:Trends in Ecology & Evolution 5265:Meyers LA, Levin DA (June 2006). 4613:Cannatella DC, De Sa RO (1993). " 4495:Variation and Evolution in Plants 3946:Variation and Evolution in Plants 2908:"The genetic origin of kiwifruit" 2383:Critical Reviews in Biotechnology 2274:. Cengage Learning. p. 344. 2078: 2059:show unequal contribution to the 1983:, the allotetraploid lager yeast 972: 959: 908: 721: 570: 565: 8908: 8896: 8884: 8873: 8872: 7582:10.1111/j.1469-8137.2005.01347.x 7211:Biochemical Society Transactions 7178:10.1128/jb.176.13.3911-3919.1994 6955: 6908: 6859: 6809: 6750: 6706: 6645: 6222:10.1111/j.1420-9101.2009.01828.x 6125:Proceedings. Biological Sciences 5879:Proceedings. Biological Sciences 4977:10.1111/j.1469-7998.2011.00829.x 4619:Society of Systematic Biologists 4093:10.1146/annurev.ecolsys.29.1.467 3973:10.1146/annurev.ecolsys.29.1.467 3882:10.1111/j.1469-8137.2009.03117.x 3844:10.1111/j.1469-8137.2009.03142.x 3247:10.1111/j.1439-0426.2011.01873.x 2848:Current Opinion in Plant Biology 964: 64:. Most species whose cells have 8144:Macrochromosome/Microchromosome 7363:Snustad DP, Simmons MJ (2006). 7129:10.1128/jb.171.6.3133-3138.1989 6596: 6547: 6504: 6437: 6386: 6335: 6276: 6237: 6209:Journal of Evolutionary Biology 6196: 6161: 6058: 6039: 6014: 5979: 5956: 5915: 5866: 5831: 5782: 5733: 5457: 5408: 5160: 5107: 5064: 4951: 4912: 4865: 4789: 4739: 4680: 4606: 4501: 4486: 4467: 4364: 4305: 4203:Cytogenetic and Genome Research 4190: 4155: 4028: 3987: 3952: 3937: 3896: 3823: 3764: 3699: 3634: 3569: 3493: 3458: 3423: 3384: 3341: 3306: 3209: 3170: 3014: 2975: 2925: 2882: 2839: 2770: 2721: 1717:Aquatic plants, especially the 1689:is a triploid sterile species. 1408:(bottom right), as well as the 1392:of a human, showing the normal 452:), for example some species of 8573: 6182:10.1080/00275514.1954.12024382 5697:Annual Review of Plant Biology 4532:10.1146/annurev.genet.34.1.401 3226:Journal of Applied Ichthyology 2662: 2597: 2548: 2473: 2372: 2337: 2288: 2263: 2113:group, the remarkable species 1977:allopolyploid: the widespread 301: 13: 1: 5621:10.1016/S0168-9525(03)00015-5 3913:10.1016/s0065-2660(08)60490-3 3417:10.1016/S0044-8486(99)00367-1 3362:10.1016/s0167-7799(00)01448-7 2423:This review cites this study: 2399:10.1080/07388551.2020.1765730 2313:10.1080/15384101.2019.1618123 2257: 1767: 1671:Some plants are triploid. As 1038:), mustard weed/thale cress ( 829:) and three allotetraploids ( 473:), for example some cultured 6784:10.1371/journal.pone.0005914 6412:10.1371/journal.pgen.1000549 6361:10.1371/journal.pgen.1004161 6008:10.1016/0304-3770(93)90071-4 5642:Chen ZJ, Ni Z (March 2006). 5039:10.1371/journal.pgen.1007698 4687:Epstein CJ (February 1967). 3790:10.1371/journal.pgen.1008900 3452:10.1016/0044-8486(83)90179-5 3143:10.1371/journal.pone.0135411 2938:Genetics Selection Evolution 2818:. Springer. pp. 30–31. 2688:10.1371/journal.pgen.1008900 2435:Genome Biology and Evolution 1576:favoring the diploid cells. 1100: 941: 890: 436:), for example Kenai Birch ( 48:is a condition in which the 7: 8903:Evolutionary biology Portal 7898:The Evolution of the Genome 7803:The Evolution of the Genome 7513:The Evolution of the Genome 7029:10.1128/JB.134.1.71-75.1978 6825:Fungal Genetics and Biology 6083:10.1534/genetics.108.091355 5852:10.1016/j.ydbio.2004.03.014 5758:10.1534/genetics.106.057554 5142:Gynaecology by Ten Teachers 4851:10.1016/j.ygeno.2004.12.004 4815:10.1016/j.ygeno.2006.02.010 4371:Edger PP, Pires JC (2009). 3048:10.1016/j.apjtb.2017.09.015 2215: 2137:contains 4-8 copies of its 2123: 1816:Tetraploid crops: very few 1201: 1152:inter-species hybridization 676:are used to disrupt normal 215:(two sets of chromosomes), 10: 8958: 8184:Dinoflagellate chromosomes 6250:American Journal of Botany 5478:10.1016/j.tree.2005.07.008 5385:10.1016/j.cell.2007.10.022 4164:American Journal of Botany 3948:. Oxford University Press. 3732:10.1038/s41467-019-13159-5 3667:10.1038/s41467-019-13159-5 3183:American Journal of Botany 2630:10.1038/s41467-019-13159-5 2210:homologous recombinational 2180: 1489:or failure to extrude one 1421: 1413: 1412:(to scale at bottom left). 1206: 1140: 1104: 1001: 945: 917:tissues. This is known as 894: 530:), for example the plants 310: 29: 8868: 8815: 8764: 8707: 8655: 8609: 8581: 8528:List of sequenced genomes 8503: 8406: 8368: 8338: 8296:Chromosomal translocation 8266: 8169:A chromosome/B chromosome 8160:(or accessory chromosome) 8122: 8053: 7273:10.1007/s00792-005-0437-4 6837:10.1016/j.fgb.2006.02.006 5433:10.1038/s41586-019-1693-2 5366:Otto SP (November 2007). 4767:10.1038/s41575-020-0284-x 4556:Smith LE (October 2012). 4510:Annual Review of Genetics 4390:10.1007/s10577-009-9055-9 3602:10.1016/j.cub.2013.08.059 3515:10.1002/9780470380130.ch3 3080:10.1007/978-3-319-91944-7 3000:10.1007/s10530-008-9383-2 2951:10.1186/s12711-016-0194-0 2868:10.1016/j.pbi.2005.01.001 1986:Saccharomyces pastorianus 1702:, which is a hexaploid (6 1591: 1396:(that is, non-polyploid) 1379: 542:  or the amphibian 8746:Nonecological speciation 8350:Telomere-binding protein 8164:Supernumerary chromosome 7958:Nature Reviews. Genetics 7870:10.1186/gb-2004-5-12-250 7011:Hansen MT (April 1978). 4199:"Polyploidy in Amphibia" 4121:Nature Reviews. Genetics 2191:Halobacterium salinarium 2163:homologous recombination 2097:. Some species of brown 1967:Saccharomyces cerevisiae 1947: 1916:Dodecaploid crops: some 1724: 1706:) with 66 chromosomes (2 1546:. In diandry, a partial 1148:Homoeologous chromosomes 1137:Homoeologous chromosomes 1122:and, more specifically, 1035:Saccharomyces cerevisiae 591:and the white sturgeon, 306: 7828:10.1023/A:1022652814749 7612:10.1023/A:1022666020026 7166:Journal of Bacteriology 7117:Journal of Bacteriology 7017:Journal of Bacteriology 6310:10.1073/pnas.1105430108 5526:10.1073/pnas.0811575106 5335:10.1126/science.1137729 5186:10.1038/sj.ejhg.5201076 5073:Klinische Wochenschrift 4631:10.1093/sysbio/42.4.476 4442:10.1073/pnas.1607237113 4339:10.1073/pnas.0707522105 4281:10.1056/NEJMcibr0903347 4265:"Cardiomyocyte renewal" 3350:Trends in Biotechnology 3220:Acipenser transmontanus 2799:10.1078/0044-5231-00032 2227:Eukaryote hybrid genome 2197:Deinococcus radiodurans 2194:is polyploid and, like 2131:Deinococcus radiodurans 1813:, white pulp of coconut 1579: 931:Epulopiscium fishelsoni 594:Acipenser transmontanum 513:), for example certain 389:), for example sterile 223:or macaroni wheat, and 8851:Punctuated equilibrium 8807:Character displacement 8627:Reproductive isolation 8596:Laboratory experiments 8286:Structural alterations 7851:Van de Peer Y (2004). 7662:10.1073/pnas.212522399 7486:10.1105/tpc.107.054346 7439:Developmental Dynamics 7367:Principles of Genetics 6919:(Bacillariophyceae)". 6471:10.1073/pnas.142101099 6137:10.1098/rspb.2012.0434 5891:10.1098/rspb.2017.1579 5234:10.1055/s-0032-1331378 4921:Gasterosteus aculeatus 4874:Gasterosteus aculeatus 4617:as a Model Organism". 3503:Plant Breeding Reviews 2174:Azotobacter vinelandii 2043: 2032:Microbotryum violaceum 1620: 1607:cell undergoes failed 1432:tissue, such as liver 1419: 1402:homologous chromosomes 1317:. Its "new" diploid (2 1299:Tympanoctomys barrerae 1293:regions, known as the 999: 545:Xenopus ruwenzoriensis 338: 233:are also tetraploids. 196:Hibiscus rosa-sinensis 42: 8732:Ecological speciation 8647:Evidence of evolution 8303:Numerical alterations 8291:Chromosomal inversion 8189:Homologous chromosome 8006:The polyploidy portal 6884:10.1093/aobpla/plr001 6621:10.1093/dnares/dsp003 6572:10.1101/gr.076075.108 5840:Developmental Biology 4714:10.1073/pnas.57.2.327 4238:Griffiths AJ (1999). 3712:Nature Communications 3647:Nature Communications 2944:12. Springer Nature. 2779:Zoologischer Anzeiger 2610:Nature Communications 2041: 1773:Triploid crops: some 1599: 1422:Further information: 1414:Further information: 1387: 1010:evolutionary lineages 989: 713:polysomic inheritance 685:homologous chromosome 397:, also common in the 323: 272:the Scottish mainland 251:Erythranthe peregrina 185:. It is common among 40: 8511:Extrachromosomal DNA 8199:Satellite chromosome 8174:Lampbrush chromosome 8114:Nuclear organization 7797:Tate JA, Soltis DE, 6921:Journal of Phycology 5965:Sequoia sempervirens 3944:Stebbins GL (1950). 3903:Stebbins GL (1947). 2988:Biological Invasions 2393:(T&F): 865–880. 2391:Taylor & Francis 2247:Reciprocal silencing 2020:homoploid speciation 1992:Dekkera bruxellensis 1699:Sequoia sempervirens 1660:polyploid speciation 1611:, producing diploid 1467:Pneumocystis carinii 1410:mitochondrial genome 1248:Liolaemus chiliensis 1156:allopolyploidization 1069:(which includes the 1041:Arabidopsis thaliana 635:asexual reproduction 554:(forty-four sets; 44 552:tetratetracontaploid 395:seedless watermelons 363:), for example male 335:Dinoponera australis 246:sympatric speciation 91:(eggs and sperm) by 8736:Parallel speciation 8204:Centromere position 8179:Polytene chromosome 8149:Circular chromosome 7923:1997Natur.387..708W 7653:2002PNAS...9913627S 7647:(21): 13627–13632. 7569:The New Phytologist 7547:10.1038/nature03025 7538:2004Natur.431..946J 7408:2000Natur.408..796T 7320:2007RadR..168..507D 7082:10.1038/nature05160 7074:2006Natur.443..569Z 6989:10.1038/nature05230 6980:2006Natur.444..171A 6933:2002JPcgy..38.1004C 6775:2009PLoSO...4.5914L 6674:10.1038/nature04562 6666:2006Natur.440..341S 6462:2002PNAS...99.9272W 6301:2011PNAS..10814539L 6295:(35): 14539–14544. 6131:(1738): 2497–2509. 6000:1993AqBot..44..181L 5928:The New Phytologist 5585:10.1038/nature10158 5517:2009PNAS..10613875W 5511:(33): 13875–13879. 5327:2007Sci...317..910R 4925:Journal of Genetics 4890:1956Natur.178.1124S 4884:(4542): 1124–1125. 4705:1967PNAS...57..327E 4433:2016PNAS..11311531C 4427:(41): 11531–11536. 4377:Chromosome Research 4330:2008PNAS..105.6730M 4041:The New Phytologist 4000:The New Phytologist 3870:The New Phytologist 3832:The New Phytologist 3724:2019NatCo..10.5218M 3659:2019NatCo..10.5218M 3594:2013CBio...23.2151Y 3467:Journal of Heredity 3444:1983Aquac..30..375L 3409:2000Aquac.186...61C 3278:The New Phytologist 3238:2011JApIc..27...24D 3134:2015PLoSO..1035411Z 3074:. pp. 89–130. 2860:2005COPB....8..135A 2791:2001ZooAn.240..247B 2622:2019NatCo..10.5218M 2535:Scientific American 2206:DNA binding protein 2105:, Laminariales and 1777:varieties (such as 1656:interploidy crosses 1295:plains viscacha rat 1192:Aegilops speltoides 1180:Aegilops speltoides 1168:Aegilops speltoides 1096:Other similar terms 707:The high degree of 698:Arabidopsis arenosa 415:Plains viscacha rat 332:) in the giant ant 292:Arabidopsis arenosa 8937:Classical genetics 8816:Speciation in taxa 8751:Assortative mating 7739:10.1002/tax.595006 7452:10.1002/dvdy.10363 7308:Radiation Research 7223:10.1042/BST0390150 6246:Cyathus stercoreus 5885:(1868): 20171579. 5807:10.1093/aob/mcn017 5660:10.1002/bies.20374 5609:Trends in Genetics 5311:"Plant speciation" 5146:. London: Arnold. 5085:10.1007/BF01734832 4964:Journal of Zoology 4937:10.1007/BF02984740 3181:(Saxifragaceae)". 2447:10.1093/gbe/evw293 2358:10.1007/BF00293307 2270:Solomon E (2014). 2044: 1980:Cyathus stercoreus 1621: 1603:via polyploidy: A 1559:prenatal diagnosis 1552:genomic imprinting 1420: 1000: 702:Arabidopsis lyrata 674:mitotic inhibitors 664:or treatment with 613:) or tetraploid (2 424:Gossypium hirsutum 365:European fire ants 339: 296:Arabidopsis lyrata 111:and multicellular 43: 8924: 8923: 8802:Secondary contact 8774:Hybrid speciation 8722:Natural selection 8709:Isolating factors 8541: 8540: 8499: 8498: 8236:Centromere number 8153:Linear chromosome 7917:(6634): 708–713. 7772:10.1101/gr.640303 7532:(7011): 946–957. 7480:(11): 3403–3417. 7402:(6814): 796–815. 7378:978-0-471-69939-2 7172:(13): 3911–3919. 7068:(7111): 569–573. 6974:(7116): 171–178. 6660:(7082): 341–345. 6566:(10): 1610–1623. 6456:(14): 9272–9277. 6215:(11): 2157–2170. 5941:10.1111/nph.19057 5578:(7355): 189–195. 5427:(7780): 679–685. 5321:(5840): 910–914. 5153:978-0-340-81662-2 5079:(23): 1115–1131. 4898:10.1038/1781124a0 4474:White MJ (1973). 4324:(18): 6730–6734. 4249:978-0-7167-3520-5 4216:10.1159/000431388 4054:10.1111/nph.14276 4013:10.1111/nph.13698 3588:(21): 2151–2156. 3179:Tolmiea menziesii 3089:978-3-319-91943-0 2825:978-0-412-98691-8 2747:10.1111/nph.17204 2574:10.1111/nph.17204 2498:10.1111/evo.12678 2307:(12): 1302–1315. 2237:Polyploid complex 2083:Other eukaryotic 2069:natural selection 1891:Octaploid crops: 1869:Hexaploid crops: 1739:Triticum turgidum 1733:is the hybrid of 1686:Lomatia tasmanica 1548:hydatidiform mole 1542:and a very small 1424:Triploid syndrome 1026:sequenced genomes 992:phylogenetic tree 662:protoplast fusion 475:Siberian sturgeon 421:fish, the cotton 313:Triploid syndrome 286:is not sterile. 16:(Redirected from 8949: 8912: 8911: 8900: 8888: 8887: 8876: 8875: 8727:Sexual selection 8656:Geographic modes 8568: 8561: 8554: 8545: 8544: 8366: 8365: 8330:Polyploidization 8158:Extra chromosome 8073:Genetic material 8036: 8029: 8022: 8013: 8012: 7989: 7970:10.1038/35072009 7952: 7934: 7901: 7892: 7882: 7872: 7847: 7806: 7793: 7783: 7750: 7733:(5): 1387–1403. 7717: 7692: 7682: 7664: 7631: 7606:(1–4): 117–129. 7594: 7584: 7559: 7549: 7516: 7507: 7497: 7464: 7454: 7429: 7419: 7417:10.1038/35048692 7382: 7370: 7350: 7349: 7331: 7329:10.1667/RR0935.1 7299: 7293: 7292: 7258: 7249: 7243: 7242: 7206: 7200: 7199: 7189: 7157: 7151: 7150: 7140: 7123:(6): 3133–3138. 7108: 7102: 7101: 7057: 7051: 7050: 7040: 7008: 7002: 7001: 6991: 6959: 6953: 6952: 6927:(5): 1004–1019. 6912: 6906: 6905: 6895: 6863: 6857: 6856: 6822: 6813: 6807: 6806: 6796: 6786: 6754: 6748: 6747: 6728:10.1002/yea.1654 6710: 6704: 6703: 6685: 6649: 6643: 6642: 6632: 6600: 6594: 6593: 6583: 6551: 6545: 6544: 6508: 6502: 6501: 6491: 6473: 6441: 6435: 6434: 6424: 6414: 6390: 6384: 6383: 6373: 6363: 6339: 6333: 6332: 6322: 6312: 6280: 6274: 6273: 6241: 6235: 6234: 6224: 6200: 6194: 6193: 6165: 6159: 6158: 6148: 6116: 6105: 6104: 6094: 6062: 6056: 6055: 6043: 6037: 6036: 6034: 6033: 6024:. Archived from 6018: 6012: 6011: 5994:(2–3): 181–228. 5983: 5977: 5976: 5960: 5954: 5953: 5943: 5919: 5913: 5912: 5902: 5870: 5864: 5863: 5835: 5829: 5828: 5818: 5795:Annals of Botany 5786: 5780: 5779: 5769: 5752:(2): 1101–1113. 5737: 5731: 5730: 5720: 5691:Chen ZJ (2007). 5688: 5682: 5681: 5671: 5639: 5633: 5632: 5604: 5598: 5597: 5587: 5563: 5557: 5556: 5546: 5528: 5496: 5490: 5489: 5461: 5455: 5454: 5444: 5412: 5406: 5405: 5387: 5363: 5357: 5356: 5346: 5306: 5297: 5296: 5286: 5277:(6): 1198–1206. 5262: 5256: 5255: 5245: 5213: 5207: 5206: 5188: 5164: 5158: 5157: 5145: 5135: 5126: 5125: 5123: 5122: 5111: 5105: 5104: 5068: 5062: 5061: 5051: 5041: 5032:(10): e1007698. 5017: 5011: 5010: 4999: 4990: 4989: 4979: 4955: 4949: 4948: 4916: 4910: 4909: 4869: 4863: 4862: 4834: 4828: 4827: 4817: 4793: 4787: 4786: 4752: 4743: 4737: 4736: 4726: 4716: 4684: 4678: 4677: 4641: 4635: 4634: 4610: 4604: 4603: 4585: 4553: 4544: 4543: 4525: 4505: 4499: 4498: 4490: 4484: 4483: 4481: 4471: 4465: 4464: 4454: 4444: 4412: 4403: 4402: 4392: 4368: 4362: 4361: 4351: 4341: 4309: 4303: 4302: 4292: 4260: 4254: 4253: 4235: 4229: 4228: 4218: 4209:(3–4): 315–330. 4194: 4188: 4187: 4159: 4153: 4152: 4116: 4105: 4104: 4076: 4067: 4066: 4056: 4032: 4026: 4025: 4015: 3991: 3985: 3984: 3956: 3950: 3949: 3941: 3935: 3934: 3900: 3894: 3893: 3865: 3856: 3855: 3827: 3821: 3820: 3810: 3792: 3768: 3762: 3761: 3751: 3703: 3697: 3696: 3686: 3638: 3632: 3631: 3621: 3573: 3567: 3566: 3538: 3529: 3528: 3508: 3497: 3491: 3490: 3462: 3456: 3455: 3438:(1–4): 375–380. 3427: 3421: 3420: 3388: 3382: 3381: 3345: 3339: 3338: 3321:(2–3): 363–378. 3310: 3304: 3303: 3293: 3269: 3260: 3259: 3249: 3213: 3207: 3206: 3189:(9): 1171–1174. 3174: 3168: 3165: 3155: 3145: 3101: 3060: 3050: 3018: 3012: 3011: 2994:(5): 1159–1173. 2979: 2973: 2972: 2970: 2968: 2963: 2953: 2929: 2923: 2922: 2920: 2919: 2910:. Archived from 2903: 2894: 2893: 2886: 2880: 2879: 2843: 2837: 2836: 2834: 2832: 2809: 2803: 2802: 2785:(3–4): 247–252. 2774: 2768: 2767: 2749: 2725: 2719: 2718: 2708: 2690: 2666: 2660: 2659: 2649: 2601: 2595: 2594: 2576: 2552: 2546: 2545: 2543: 2541: 2526: 2520: 2519: 2509: 2492:(6): 1487–1500. 2477: 2471: 2468: 2458: 2418: 2376: 2370: 2369: 2341: 2335: 2334: 2324: 2292: 2286: 2285: 2267: 2153:can shatter its 2075:and speciation. 1970:strains used in 1942:kinnow mandarins 1779:Belle de Boskoop 1514: 1513: 1509: 1506: 1440:is more common. 1366:J. B. S. Haldane 1253:mole salamanders 1086:flowering plants 735:heteropolyploids 654:Malus domesticus 588:Tolmiea menzisii 539:Spartina anglica 533:Celosia argentea 526:(twelve sets; 12 486:, (eight sets; 8 413:), for example, 21: 8957: 8956: 8952: 8951: 8950: 8948: 8947: 8946: 8927: 8926: 8925: 8920: 8864: 8847:Paleopolyploidy 8811: 8766:Hybrid concepts 8760: 8703: 8651: 8621:Species complex 8605: 8577: 8572: 8542: 8537: 8495: 8402: 8364: 8334: 8323:Paleopolyploidy 8268: 8262: 8118: 8092:Heterochromatin 8055: 8049: 8040: 7997: 7992: 7760:Genome Research 7379: 7358: 7356:Further reading 7353: 7300: 7296: 7256: 7250: 7246: 7207: 7203: 7158: 7154: 7109: 7105: 7058: 7054: 7009: 7005: 6960: 6956: 6913: 6909: 6864: 6860: 6820: 6814: 6810: 6755: 6751: 6711: 6707: 6650: 6646: 6601: 6597: 6560:Genome Research 6552: 6548: 6509: 6505: 6442: 6438: 6405:(7): e1000549. 6391: 6387: 6354:(2): e1004161. 6340: 6336: 6281: 6277: 6262:10.2307/2440307 6242: 6238: 6201: 6197: 6166: 6162: 6117: 6108: 6063: 6059: 6048:Oxalis tuberosa 6044: 6040: 6031: 6029: 6020: 6019: 6015: 5984: 5980: 5969:Silvae Genetica 5961: 5957: 5920: 5916: 5871: 5867: 5836: 5832: 5787: 5783: 5738: 5734: 5689: 5685: 5640: 5636: 5605: 5601: 5564: 5560: 5497: 5493: 5472:(11): 591–597. 5462: 5458: 5413: 5409: 5364: 5360: 5307: 5300: 5263: 5259: 5214: 5210: 5179:(12): 972–974. 5165: 5161: 5154: 5136: 5129: 5120: 5118: 5113: 5112: 5108: 5069: 5065: 5018: 5014: 5001: 5000: 4993: 4956: 4952: 4917: 4913: 4870: 4866: 4835: 4831: 4794: 4790: 4750: 4744: 4740: 4685: 4681: 4658:10.1139/g06-152 4642: 4638: 4611: 4607: 4554: 4547: 4523:10.1.1.323.1059 4506: 4502: 4491: 4487: 4478:The Chromosomes 4472: 4468: 4413: 4406: 4369: 4365: 4310: 4306: 4261: 4257: 4250: 4236: 4232: 4195: 4191: 4176:10.2307/2438023 4160: 4156: 4133:10.1038/nrg1711 4127:(11): 836–846. 4117: 4108: 4077: 4070: 4033: 4029: 3992: 3988: 3957: 3953: 3942: 3938: 3923: 3901: 3897: 3866: 3859: 3828: 3824: 3783:(7): e1008900. 3769: 3765: 3704: 3700: 3639: 3635: 3582:Current Biology 3574: 3570: 3539: 3532: 3525: 3506: 3498: 3494: 3463: 3459: 3428: 3424: 3389: 3385: 3346: 3342: 3311: 3307: 3270: 3263: 3214: 3210: 3195:10.2307/2443640 3175: 3171: 3128:(8): e0135411. 3090: 3070:. Vol. 2. 3019: 3015: 2980: 2976: 2966: 2964: 2930: 2926: 2917: 2915: 2904: 2897: 2888: 2887: 2883: 2844: 2840: 2830: 2828: 2826: 2810: 2806: 2775: 2771: 2734:New Phytologist 2726: 2722: 2681:(7): e1008900. 2667: 2663: 2602: 2598: 2561:New Phytologist 2553: 2549: 2539: 2537: 2527: 2523: 2478: 2474: 2377: 2373: 2342: 2338: 2293: 2289: 2282: 2268: 2264: 2260: 2218: 2183: 2149:irradiation or 2126: 2081: 2071:and subsequent 2001:Rhizopus oryzae 1950: 1910:Oxalis tuberosa 1770: 1727: 1625:paleopolyploidy 1594: 1582: 1511: 1507: 1504: 1502: 1426: 1418: 1382: 1374:Shinya Yamanaka 1245:; others, like 1243:parthenogenesis 1209: 1204: 1184:Triticum urartu 1176:Triticum urartu 1164:Triticum urartu 1145: 1139: 1109: 1103: 1098: 1006: 1004:Paleopolyploidy 996:paleopolyploidy 984: 982:Paleopolyploidy 975: 967: 962: 950: 944: 911: 899: 893: 764:chromosomes or 759: 753: 731:amphipolyploids 724: 631:triploid bridge 573: 568: 558:), for example 490:), for example 376:), for example 316: 309: 304: 35: 28: 23: 22: 15: 12: 11: 5: 8955: 8945: 8944: 8939: 8922: 8921: 8919: 8918: 8906: 8894: 8882: 8869: 8866: 8865: 8863: 8862: 8855:Macroevolution 8840: 8835: 8830: 8825: 8819: 8817: 8813: 8812: 8810: 8809: 8804: 8799: 8789: 8770: 8768: 8762: 8761: 8759: 8758: 8756:Haldane's rule 8753: 8748: 8743: 8729: 8724: 8719: 8713: 8711: 8705: 8704: 8702: 8701: 8696: 8682: 8679:Founder effect 8659: 8657: 8653: 8652: 8650: 8649: 8644: 8639: 8634: 8629: 8624: 8613: 8611: 8610:Basic concepts 8607: 8606: 8604: 8603: 8598: 8593: 8588: 8582: 8579: 8578: 8571: 8570: 8563: 8556: 8548: 8539: 8538: 8536: 8535: 8530: 8525: 8520: 8519: 8518: 8507: 8505: 8501: 8500: 8497: 8496: 8494: 8493: 8488: 8483: 8478: 8473: 8468: 8463: 8458: 8453: 8448: 8443: 8438: 8433: 8428: 8423: 8418: 8412: 8410: 8404: 8403: 8401: 8400: 8395: 8390: 8385: 8380: 8374: 8372: 8363: 8362: 8357: 8342: 8340: 8336: 8335: 8333: 8332: 8327: 8326: 8325: 8320: 8315: 8310: 8300: 8299: 8298: 8293: 8283: 8278: 8272: 8270: 8264: 8263: 8261: 8260: 8259: 8258: 8253: 8248: 8243: 8233: 8232: 8231: 8226: 8221: 8216: 8214:Submetacentric 8211: 8201: 8196: 8191: 8186: 8181: 8176: 8171: 8166: 8161: 8155: 8146: 8141: 8140:or heterosome) 8134:Sex chromosome 8126: 8124: 8120: 8119: 8117: 8116: 8111: 8106: 8101: 8096: 8095: 8094: 8089: 8079: 8070: 8065: 8059: 8057: 8051: 8050: 8039: 8038: 8031: 8024: 8016: 8010: 8009: 8003: 7996: 7995:External links 7993: 7991: 7990: 7964:(5): 333–341. 7953: 7902: 7893: 7857:Genome Biology 7848: 7822:(1–4): 65–73. 7807: 7794: 7766:(3): 382–390. 7751: 7718: 7693: 7632: 7595: 7575:(3): 658–661. 7560: 7517: 7508: 7474:The Plant Cell 7465: 7445:(4): 751–766. 7430: 7383: 7377: 7359: 7357: 7354: 7352: 7351: 7314:(4): 507–514. 7294: 7267:(3): 219–227. 7244: 7217:(1): 150–154. 7201: 7152: 7103: 7052: 7003: 6954: 6907: 6858: 6831:(7): 511–529. 6808: 6749: 6722:(3): 147–165. 6705: 6644: 6615:(2): 115–129. 6595: 6546: 6525:10.3852/09-192 6519:(3): 689–696. 6503: 6436: 6385: 6334: 6275: 6256:(3): 343–347. 6236: 6195: 6176:(4): 393–434. 6160: 6106: 6077:(1): 649–660. 6057: 6038: 6013: 5988:Aquatic Botany 5978: 5955: 5934:(3): 918–927. 5914: 5865: 5846:(2): 277–289. 5830: 5801:(6): 833–843. 5781: 5732: 5683: 5654:(3): 240–252. 5634: 5615:(3): 141–147. 5599: 5558: 5491: 5456: 5407: 5378:(3): 452–462. 5358: 5298: 5257: 5208: 5159: 5152: 5127: 5106: 5063: 5012: 4991: 4970:(3): 151–182. 4950: 4931:(2): 129–142. 4911: 4864: 4845:(4): 425–430. 4829: 4808:(2): 214–221. 4788: 4761:(7): 391–405. 4738: 4699:(2): 327–334. 4679: 4652:(2): 119–136. 4636: 4625:(4): 476–507. 4615:Xenopus laevis 4605: 4568:(4 Suppl): B. 4545: 4516:(1): 401–437. 4500: 4485: 4466: 4404: 4383:(5): 699–717. 4363: 4304: 4255: 4248: 4230: 4189: 4170:(7): 487–499. 4154: 4106: 4087:(1): 467–501. 4068: 4047:(2): 487–493. 4027: 4006:(2): 391–398. 3986: 3967:(1): 467–501. 3951: 3936: 3921: 3895: 3876:(1): 113–122. 3857: 3822: 3763: 3698: 3633: 3568: 3549:(1): 589–639. 3530: 3523: 3492: 3473:(3): 145–149. 3457: 3422: 3403:(1–2): 61–75. 3383: 3356:(6): 233–242. 3340: 3305: 3261: 3208: 3169: 3167: 3166: 3109: 3107: 3104: 3102: 3088: 3063: 3061: 3013: 2974: 2924: 2895: 2881: 2854:(2): 135–141. 2838: 2824: 2804: 2769: 2740:(2): 457–461. 2720: 2661: 2596: 2567:(2): 457–461. 2547: 2521: 2472: 2470: 2469: 2441:(2): 266–278. 2426: 2424: 2421: 2419: 2371: 2336: 2287: 2281:978-1285423586 2280: 2261: 2259: 2256: 2255: 2254: 2249: 2244: 2239: 2234: 2229: 2224: 2222:Diploidization 2217: 2214: 2208:and is likely 2182: 2179: 2159:D. radiodurans 2143:D. radiodurans 2141:. Exposure of 2125: 2122: 2080: 2079:Chromalveolata 2077: 2057:S. pastorianus 2012: 2011: 1996: 1975: 1954:fungal species 1949: 1946: 1922: 1921: 1914: 1889: 1867: 1814: 1811:saffron crocus 1791:Ribston Pippin 1769: 1766: 1747:Secale cereale 1726: 1723: 1719:Monocotyledons 1681:Crocus sativus 1677:saffron crocus 1593: 1590: 1581: 1578: 1535:, with marked 1430:differentiated 1406:sex chromosome 1400:. It shows 22 1381: 1378: 1261:spermatophores 1208: 1205: 1203: 1200: 1141:Main article: 1138: 1135: 1105:Main article: 1102: 1099: 1097: 1094: 1058:) in an early 1014:diploidization 1002:Main article: 998:in eukaryotes. 983: 980: 974: 973:Mesopolyploidy 971: 966: 963: 961: 960:Temporal terms 958: 946:Main article: 943: 940: 919:endopolyploidy 910: 909:Endopolyploidy 907: 895:Main article: 892: 889: 866:T. porrifolius 778:heterozygosity 766:nondisjunction 757: 751: 727:Allopolyploids 723: 722:Allopolyploidy 720: 696:documented in 627:triploid block 576:Autopolyploids 572: 571:Autopolyploidy 569: 567: 566:Classification 564: 563: 562: 560:black mulberry 549: 517: 504: 477: 469:(seven sets; 7 460: 443: 439:Betula kenaica 427: 404: 391:saffron crocus 385:(three sets; 3 380: 367: 308: 305: 303: 300: 280:E. × robertsii 276:Orkney Islands 256:E. × robertsii 189:and flowering 84:, but produce 26: 9: 6: 4: 3: 2: 8954: 8943: 8940: 8938: 8935: 8934: 8932: 8917: 8916: 8907: 8905: 8904: 8899: 8895: 8893: 8892: 8883: 8881: 8880: 8871: 8870: 8867: 8860: 8859:Chronospecies 8856: 8852: 8848: 8844: 8841: 8839: 8836: 8834: 8831: 8829: 8826: 8824: 8821: 8820: 8818: 8814: 8808: 8805: 8803: 8800: 8797: 8793: 8792:Reinforcement 8790: 8787: 8786:Recombination 8783: 8779: 8775: 8772: 8771: 8769: 8767: 8763: 8757: 8754: 8752: 8749: 8747: 8744: 8741: 8737: 8733: 8730: 8728: 8725: 8723: 8720: 8718: 8715: 8714: 8712: 8710: 8706: 8700: 8697: 8694: 8690: 8686: 8683: 8680: 8676: 8672: 8668: 8664: 8661: 8660: 8658: 8654: 8648: 8645: 8643: 8640: 8638: 8635: 8633: 8630: 8628: 8625: 8622: 8618: 8615: 8614: 8612: 8608: 8602: 8599: 8597: 8594: 8592: 8589: 8587: 8584: 8583: 8580: 8576: 8569: 8564: 8562: 8557: 8555: 8550: 8549: 8546: 8534: 8531: 8529: 8526: 8524: 8521: 8517: 8514: 8513: 8512: 8509: 8508: 8506: 8502: 8492: 8489: 8487: 8484: 8482: 8479: 8477: 8474: 8472: 8469: 8467: 8464: 8462: 8459: 8457: 8454: 8452: 8449: 8447: 8444: 8442: 8439: 8437: 8434: 8432: 8429: 8427: 8424: 8422: 8419: 8417: 8414: 8413: 8411: 8409: 8405: 8399: 8396: 8394: 8391: 8389: 8386: 8384: 8381: 8379: 8376: 8375: 8373: 8371: 8367: 8361: 8358: 8355: 8351: 8347: 8344: 8343: 8341: 8337: 8331: 8328: 8324: 8321: 8319: 8316: 8314: 8311: 8309: 8306: 8305: 8304: 8301: 8297: 8294: 8292: 8289: 8288: 8287: 8284: 8282: 8279: 8277: 8274: 8273: 8271: 8269:and evolution 8265: 8257: 8254: 8252: 8249: 8247: 8244: 8242: 8239: 8238: 8237: 8234: 8230: 8227: 8225: 8222: 8220: 8217: 8215: 8212: 8210: 8207: 8206: 8205: 8202: 8200: 8197: 8195: 8194:Isochromosome 8192: 8190: 8187: 8185: 8182: 8180: 8177: 8175: 8172: 8170: 8167: 8165: 8162: 8159: 8156: 8154: 8150: 8147: 8145: 8142: 8139: 8135: 8131: 8128: 8127: 8125: 8121: 8115: 8112: 8110: 8107: 8105: 8102: 8100: 8097: 8093: 8090: 8088: 8085: 8084: 8083: 8080: 8078: 8074: 8071: 8069: 8066: 8064: 8061: 8060: 8058: 8052: 8048: 8044: 8037: 8032: 8030: 8025: 8023: 8018: 8017: 8014: 8007: 8004: 8002: 7999: 7998: 7987: 7983: 7979: 7975: 7971: 7967: 7963: 7959: 7954: 7950: 7946: 7942: 7938: 7933: 7932:10.1038/42711 7928: 7924: 7920: 7916: 7912: 7908: 7903: 7899: 7894: 7890: 7886: 7881: 7876: 7871: 7866: 7862: 7858: 7854: 7849: 7845: 7841: 7837: 7833: 7829: 7825: 7821: 7817: 7813: 7808: 7804: 7800: 7795: 7791: 7787: 7782: 7777: 7773: 7769: 7765: 7761: 7757: 7752: 7748: 7744: 7740: 7736: 7732: 7728: 7724: 7719: 7715: 7711: 7707: 7703: 7699: 7694: 7690: 7686: 7681: 7676: 7672: 7668: 7663: 7658: 7654: 7650: 7646: 7642: 7638: 7633: 7629: 7625: 7621: 7617: 7613: 7609: 7605: 7601: 7596: 7592: 7588: 7583: 7578: 7574: 7570: 7566: 7561: 7557: 7553: 7548: 7543: 7539: 7535: 7531: 7527: 7523: 7518: 7514: 7509: 7505: 7501: 7496: 7491: 7487: 7483: 7479: 7475: 7471: 7466: 7462: 7458: 7453: 7448: 7444: 7440: 7436: 7431: 7427: 7423: 7418: 7413: 7409: 7405: 7401: 7397: 7393: 7389: 7384: 7380: 7374: 7369: 7368: 7361: 7360: 7347: 7343: 7339: 7335: 7330: 7325: 7321: 7317: 7313: 7309: 7305: 7298: 7290: 7286: 7282: 7278: 7274: 7270: 7266: 7262: 7261:Extremophiles 7255: 7248: 7240: 7236: 7232: 7228: 7224: 7220: 7216: 7212: 7205: 7197: 7193: 7188: 7183: 7179: 7175: 7171: 7167: 7163: 7156: 7148: 7144: 7139: 7134: 7130: 7126: 7122: 7118: 7114: 7107: 7099: 7095: 7091: 7087: 7083: 7079: 7075: 7071: 7067: 7063: 7056: 7048: 7044: 7039: 7034: 7030: 7026: 7022: 7018: 7014: 7007: 6999: 6995: 6990: 6985: 6981: 6977: 6973: 6969: 6965: 6958: 6950: 6946: 6942: 6938: 6934: 6930: 6926: 6922: 6918: 6911: 6903: 6899: 6894: 6889: 6885: 6881: 6877: 6873: 6869: 6862: 6854: 6850: 6846: 6842: 6838: 6834: 6830: 6826: 6819: 6812: 6804: 6800: 6795: 6790: 6785: 6780: 6776: 6772: 6768: 6764: 6760: 6753: 6745: 6741: 6737: 6733: 6729: 6725: 6721: 6717: 6709: 6701: 6697: 6693: 6689: 6684: 6679: 6675: 6671: 6667: 6663: 6659: 6655: 6648: 6640: 6636: 6631: 6626: 6622: 6618: 6614: 6610: 6606: 6599: 6591: 6587: 6582: 6577: 6573: 6569: 6565: 6561: 6557: 6550: 6542: 6538: 6534: 6530: 6526: 6522: 6518: 6514: 6507: 6499: 6495: 6490: 6485: 6481: 6477: 6472: 6467: 6463: 6459: 6455: 6451: 6447: 6440: 6432: 6428: 6423: 6418: 6413: 6408: 6404: 6400: 6399:PLOS Genetics 6396: 6389: 6381: 6377: 6372: 6367: 6362: 6357: 6353: 6349: 6348:PLOS Genetics 6345: 6338: 6330: 6326: 6321: 6316: 6311: 6306: 6302: 6298: 6294: 6290: 6286: 6279: 6271: 6267: 6263: 6259: 6255: 6251: 6247: 6240: 6232: 6228: 6223: 6218: 6214: 6210: 6206: 6199: 6191: 6187: 6183: 6179: 6175: 6171: 6164: 6156: 6152: 6147: 6142: 6138: 6134: 6130: 6126: 6122: 6115: 6113: 6111: 6102: 6098: 6093: 6088: 6084: 6080: 6076: 6072: 6068: 6061: 6053: 6049: 6042: 6028:on 2019-02-03 6027: 6023: 6017: 6009: 6005: 6001: 5997: 5993: 5989: 5982: 5974: 5970: 5966: 5959: 5951: 5947: 5942: 5937: 5933: 5929: 5925: 5918: 5910: 5906: 5901: 5896: 5892: 5888: 5884: 5880: 5876: 5869: 5861: 5857: 5853: 5849: 5845: 5841: 5834: 5826: 5822: 5817: 5812: 5808: 5804: 5800: 5796: 5792: 5785: 5777: 5773: 5768: 5763: 5759: 5755: 5751: 5747: 5743: 5736: 5728: 5724: 5719: 5714: 5710: 5706: 5702: 5698: 5694: 5687: 5679: 5675: 5670: 5665: 5661: 5657: 5653: 5649: 5645: 5638: 5630: 5626: 5622: 5618: 5614: 5610: 5603: 5595: 5591: 5586: 5581: 5577: 5573: 5569: 5562: 5554: 5550: 5545: 5540: 5536: 5532: 5527: 5522: 5518: 5514: 5510: 5506: 5502: 5495: 5487: 5483: 5479: 5475: 5471: 5467: 5460: 5452: 5448: 5443: 5438: 5434: 5430: 5426: 5422: 5418: 5411: 5403: 5399: 5395: 5391: 5386: 5381: 5377: 5373: 5369: 5362: 5354: 5350: 5345: 5340: 5336: 5332: 5328: 5324: 5320: 5316: 5312: 5305: 5303: 5294: 5290: 5285: 5280: 5276: 5272: 5268: 5261: 5253: 5249: 5244: 5239: 5235: 5231: 5227: 5223: 5219: 5212: 5204: 5200: 5196: 5192: 5187: 5182: 5178: 5174: 5170: 5163: 5155: 5149: 5144: 5143: 5134: 5132: 5116: 5110: 5102: 5098: 5094: 5090: 5086: 5082: 5078: 5074: 5067: 5059: 5055: 5050: 5045: 5040: 5035: 5031: 5027: 5026:PLOS Genetics 5023: 5016: 5009:. 8 Oct 2012. 5008: 5004: 4998: 4996: 4987: 4983: 4978: 4973: 4969: 4965: 4961: 4954: 4946: 4942: 4938: 4934: 4930: 4926: 4922: 4915: 4907: 4903: 4899: 4895: 4891: 4887: 4883: 4879: 4875: 4868: 4860: 4856: 4852: 4848: 4844: 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3239: 3235: 3231: 3227: 3223: 3221: 3212: 3204: 3200: 3196: 3192: 3188: 3184: 3180: 3173: 3163: 3159: 3154: 3149: 3144: 3139: 3135: 3131: 3127: 3123: 3119: 3117: 3110: 3108: 3105: 3103: 3099: 3095: 3091: 3085: 3081: 3077: 3073: 3069: 3064: 3062: 3058: 3054: 3049: 3044: 3040: 3036: 3032: 3031: 3026: 3021: 3020: 3017: 3009: 3005: 3001: 2997: 2993: 2989: 2985: 2978: 2962: 2957: 2952: 2947: 2943: 2939: 2935: 2928: 2914:on 2021-02-25 2913: 2909: 2902: 2900: 2891: 2885: 2877: 2873: 2869: 2865: 2861: 2857: 2853: 2849: 2842: 2827: 2821: 2817: 2816: 2808: 2800: 2796: 2792: 2788: 2784: 2780: 2773: 2765: 2761: 2757: 2753: 2748: 2743: 2739: 2735: 2731: 2724: 2716: 2712: 2707: 2702: 2698: 2694: 2689: 2684: 2680: 2676: 2675:PLOS Genetics 2672: 2665: 2657: 2653: 2648: 2643: 2639: 2635: 2631: 2627: 2623: 2619: 2615: 2611: 2607: 2600: 2592: 2588: 2584: 2580: 2575: 2570: 2566: 2562: 2558: 2551: 2536: 2532: 2529:Fessenden M. 2525: 2517: 2513: 2508: 2503: 2499: 2495: 2491: 2487: 2483: 2476: 2466: 2462: 2457: 2452: 2448: 2444: 2440: 2436: 2432: 2427: 2425: 2422: 2420: 2416: 2412: 2408: 2404: 2400: 2396: 2392: 2388: 2384: 2379: 2378: 2375: 2367: 2363: 2359: 2355: 2351: 2347: 2340: 2332: 2328: 2323: 2318: 2314: 2310: 2306: 2302: 2298: 2291: 2283: 2277: 2273: 2266: 2262: 2253: 2250: 2248: 2245: 2243: 2240: 2238: 2235: 2233: 2230: 2228: 2225: 2223: 2220: 2219: 2213: 2211: 2207: 2203: 2199: 2198: 2193: 2192: 2188: 2178: 2176: 2175: 2170: 2168: 2164: 2160: 2156: 2152: 2148: 2144: 2140: 2136: 2133: 2132: 2121: 2119: 2117: 2112: 2108: 2104: 2100: 2096: 2095: 2090: 2086: 2076: 2074: 2070: 2066: 2062: 2061:transcriptome 2058: 2054: 2050: 2049:Saccharomyces 2040: 2036: 2034: 2033: 2029: 2028:basidiomycota 2025: 2021: 2017: 2016:hybridization 2009: 2008: 2007:Saccharomyces 2003: 2002: 1997: 1994: 1993: 1988: 1987: 1982: 1981: 1976: 1973: 1969: 1968: 1963: 1959: 1958: 1957: 1955: 1945: 1943: 1939: 1935: 1931: 1927: 1919: 1915: 1912: 1911: 1906: 1902: 1898: 1894: 1890: 1888: 1884: 1880: 1876: 1872: 1871:chrysanthemum 1868: 1866: 1862: 1858: 1854: 1850: 1846: 1842: 1838: 1834: 1830: 1827: 1823: 1819: 1815: 1812: 1808: 1804: 1800: 1796: 1792: 1788: 1784: 1780: 1776: 1772: 1771: 1765: 1763: 1758: 1756: 1750: 1748: 1744: 1740: 1736: 1732: 1722: 1720: 1715: 1713: 1709: 1705: 1701: 1700: 1695: 1690: 1688: 1687: 1682: 1678: 1674: 1669: 1667: 1666: 1661: 1657: 1653: 1648: 1643: 1641: 1636: 1634: 1630: 1626: 1618: 1614: 1610: 1606: 1602: 1598: 1589: 1587: 1577: 1574: 1570: 1568: 1564: 1560: 1555: 1553: 1549: 1545: 1541: 1538: 1534: 1530: 1526: 1522: 1518: 1500: 1496: 1492: 1488: 1484: 1483:fertilization 1480: 1476: 1471: 1469: 1468: 1462: 1458: 1454: 1450: 1446: 1441: 1439: 1435: 1431: 1425: 1417: 1411: 1407: 1403: 1399: 1395: 1391: 1386: 1377: 1375: 1371: 1367: 1363: 1362: 1357: 1353: 1348: 1346: 1342: 1338: 1334: 1330: 1326: 1325: 1324:Octomys mimax 1320: 1316: 1312: 1309:, but kin to 1308: 1304: 1300: 1296: 1292: 1288: 1284: 1281: 1277: 1273: 1269: 1264: 1262: 1259:, "stealing" 1258: 1257:kleptogenesis 1254: 1250: 1249: 1244: 1240: 1239: 1234: 1230: 1226: 1222: 1218: 1214: 1199: 1197: 1193: 1189: 1185: 1181: 1177: 1173: 1172:hybrid genome 1169: 1165: 1161: 1157: 1153: 1149: 1144: 1134: 1132: 1127: 1125: 1121: 1117: 1114: 1108: 1093: 1091: 1087: 1083: 1079: 1076: 1072: 1068: 1064: 1061: 1057: 1056:2R hypothesis 1053: 1052: 1047: 1043: 1042: 1037: 1036: 1031: 1030:baker's yeast 1027: 1022: 1019: 1015: 1011: 1005: 997: 993: 988: 979: 970: 965:Neopolyploidy 957: 955: 949: 939: 937: 933: 932: 928: 924: 920: 916: 906: 904: 898: 888: 887: 885: 880: 879: 874: 873: 868: 867: 862: 860: 856: 851: 850: 845: 844: 839: 837: 833: 828: 827: 822: 820: 816: 811: 810:Triangle of U 807: 806:Brassicaceous 803: 799: 795: 791: 787: 783: 779: 774: 771: 767: 763: 755: 747: 746:hybridization 743: 738: 736: 732: 728: 719: 716: 714: 710: 705: 703: 699: 694: 690: 686: 681: 679: 675: 671: 667: 663: 659: 655: 650: 648: 644: 640: 636: 632: 628: 624: 620: 616: 612: 608: 604: 600: 596: 595: 590: 589: 583: 581: 577: 561: 557: 553: 550: 547: 546: 541: 540: 535: 534: 529: 525: 521: 518: 516: 512: 509:(ten sets; 10 508: 505: 503: 499: 495: 494: 489: 485: 481: 478: 476: 472: 468: 464: 461: 459: 455: 451: 447: 444: 441: 440: 435: 432:(five sets; 5 431: 428: 426: 425: 420: 416: 412: 409:(four sets; 4 408: 405: 403: 400: 396: 392: 388: 384: 381: 379: 375: 371: 368: 366: 362: 358: 355: 354: 353: 351: 348:. The letter 347: 343: 337: 336: 331: 327: 322: 318: 314: 299: 297: 293: 287: 285: 281: 277: 273: 269: 265: 261: 257: 253: 252: 247: 242: 240: 236: 232: 231: 226: 222: 218: 214: 210: 209:hybridization 206: 202: 198: 197: 192: 188: 184: 180: 176: 172: 168: 163: 161: 157: 153: 152:cell cultures 149: 148:cell division 144: 142: 138: 134: 130: 126: 122: 118: 114: 110: 106: 102: 98: 94: 90: 87: 83: 82:somatic cells 79: 75: 71: 67: 63: 59: 55: 51: 47: 39: 33: 19: 8913: 8901: 8889: 8877: 8777: 8693:Ring species 8642:Cospeciation 8637:Cladogenesis 8586:Introduction 8317: 8235: 8203: 8043:Cytogenetics 7961: 7957: 7914: 7910: 7897: 7860: 7856: 7819: 7815: 7802: 7763: 7759: 7730: 7726: 7708:(1): 13–30. 7705: 7701: 7644: 7640: 7603: 7599: 7572: 7568: 7529: 7525: 7512: 7477: 7473: 7442: 7438: 7399: 7395: 7387: 7366: 7311: 7307: 7297: 7264: 7260: 7247: 7214: 7210: 7204: 7169: 7165: 7155: 7120: 7116: 7106: 7065: 7061: 7055: 7023:(1): 71–75. 7020: 7016: 7006: 6971: 6967: 6957: 6924: 6920: 6916: 6910: 6875: 6871: 6861: 6828: 6824: 6811: 6769:(6): e5914. 6766: 6762: 6752: 6719: 6715: 6708: 6657: 6653: 6647: 6612: 6609:DNA Research 6608: 6598: 6563: 6559: 6549: 6516: 6512: 6506: 6453: 6449: 6439: 6402: 6398: 6388: 6351: 6347: 6337: 6292: 6288: 6278: 6253: 6249: 6245: 6239: 6212: 6208: 6198: 6173: 6169: 6163: 6128: 6124: 6074: 6070: 6060: 6051: 6047: 6041: 6030:. 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Retrieved 5109: 5076: 5072: 5066: 5029: 5025: 5015: 5007:ScienceDaily 5006: 4967: 4963: 4953: 4928: 4924: 4920: 4914: 4881: 4877: 4873: 4867: 4842: 4838: 4832: 4805: 4801: 4791: 4758: 4754: 4741: 4696: 4692: 4682: 4649: 4645: 4639: 4622: 4618: 4614: 4608: 4565: 4561: 4513: 4509: 4503: 4494: 4488: 4477: 4469: 4424: 4420: 4380: 4376: 4366: 4321: 4317: 4307: 4275:(1): 86–88. 4272: 4268: 4258: 4239: 4233: 4206: 4202: 4192: 4167: 4163: 4157: 4124: 4120: 4084: 4080: 4044: 4040: 4030: 4003: 3999: 3989: 3964: 3960: 3954: 3945: 3939: 3904: 3898: 3873: 3869: 3835: 3831: 3825: 3780: 3776: 3766: 3715: 3711: 3701: 3650: 3646: 3636: 3585: 3581: 3571: 3546: 3542: 3502: 3495: 3470: 3466: 3460: 3435: 3431: 3425: 3400: 3396: 3392: 3386: 3353: 3349: 3343: 3318: 3314: 3308: 3281: 3277: 3229: 3225: 3219: 3211: 3186: 3182: 3178: 3172: 3125: 3121: 3115: 3067: 3034: 3028: 3016: 2991: 2987: 2986:(Poaceae)". 2983: 2977: 2967:15 September 2965:. Retrieved 2941: 2937: 2927: 2916:. 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