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Autopolyploidy

Autopolyploids are polyploids with multiple chromosome sets derived from a single taxon. 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.[1] 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”.[1] Triploids may also persist through asexual reproduction. In fact, stable autotriploidy in plants is often associated with apomictic mating systems.[2] In agricultural systems, autotriploidy can result in seedlessness, as in watermelons and bananas.[3] Triploidy is also utilized in salmon and trout farming to induce sterility.[4][5]

Rarely, autopolyploids arise from spontaneous, somatic genome doubling, which has been observed in apple (Malus domesticus) bud sports.[6] 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.[7]

Autopolyploids possess at least three homologous chromosome sets, which can lead to high rates of multivalent pairing during meiosis (particularly in recently formed autopolyploids, a.k.a. neopolyploids) and an associated decrease in fertility due to the production of aneuploid gametes.[8] 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.[9] 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.[10]

Allopolyploidy

Allopolyploids 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 taxa – 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.[11] 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[11]. 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.[8][10]

Because pairing between homoeologous chromosomes is rare in established allopolyploids, they may benefit from fixed heterozygosity of homoeologous alleles.[12] 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 is an allotetraploid 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).[13] Complex patterns of allopolyploid evolution have also been observed in animals, as in the frog genus Xenopus.[14]

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Autopolyploidy

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Autopolyploids are polyploids with multiple chromosome sets derived from a single species. Autopolyploids can arise from a spontaneous, naturally occurring genome doubling, like the potato.[15] Others might form following fusion of 2n gametes (unreduced gametes). Bananas and apples can be found as autotriploids. Autopolyploid plants typically display polysomic inheritance, and therefore have low fertility, but may be propagated clonally.

Allopolyploidy

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Allopolyploids are polyploids with chromosomes derived from different species. Precisely it is the result of multiplying the chromosome number in an F1 hybrid. Triticale is an example of an allopolyploid, having six chromosome sets, allohexaploid, four from wheat (Triticum turgidum) and two from rye (Secale cereale). Amphidiploids are a type of allopolyploids (they are allotetraploid, containing the diploid chromosome sets of both parents[16]). Some of the best examples of allopolyploids come from the Brassicas, and 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 diploids.

  1. ^ a b Bretagnolle, F.; Thompson, J. D. (1995-01-01). "Gametes with the somatic chromosome number: mechanisms of their formation and role in the evolution of autopolyploid plants". New Phytologist. 129 (1): 1–22. doi:10.1111/j.1469-8137.1995.tb03005.x. ISSN 1469-8137.
  2. ^ Müntzing, Arne (1936-03-01). "The Evolutionary Significance of Autopolyploidy". Hereditas. 21 (2–3): 363–378. doi:10.1111/j.1601-5223.1936.tb03204.x. ISSN 1601-5223.
  3. ^ Varoquaux, F.; Blanvillain, R.; Delseny, M.; Gallois, P. (2000-06-01). "Less is better: new approaches for seedless fruit production". Trends in Biotechnology. 18 (6): 233–242. ISSN 0167-7799. PMID 10802558.
  4. ^ Cotter, D.; O'Donovan, V.; O'Maoiléidigh, N.; Rogan, G.; Roche, N.; Wilkins, N. P. (2000-06-01). "An evaluation of the use of triploid Atlantic salmon (Salmo salar L.) in minimising the impact of escaped farmed salmon on wild populations". Aquaculture. 186 (1–2): 61–75. doi:10.1016/S0044-8486(99)00367-1.
  5. ^ "Production of all-female triploid rainbow trout - ScienceDirect". www.sciencedirect.com. Retrieved 2017-03-27.
  6. ^ Dermen, Haig (1951-05-01). "TETRAPLOID AND DIPLOID ADVENTITIOUS SHOOTSFrom a Giant Sport of McIntosh Apple". Journal of Heredity. 42 (3): 145–149. doi:10.1093/oxfordjournals.jhered.a106189. ISSN 0022-1503.
  7. ^ Dwivedi, Sangam L.; Upadhyaya, Hari D.; Stalker, H. Thomas; Blair, Matthew W.; Bertioli, David J.; Nielen, Stephan; Ortiz, Rodomiro (2007-01-01). Janick, Jules (ed.). Plant Breeding Reviews. John Wiley & Sons, Inc. pp. 179–230. doi:10.1002/9780470380130.ch3. ISBN 9780470380130.
  8. ^ a b "Neopolyploidy in Flowering Plants". Annual Review of Ecology and Systematics. 33 (1): 589–639. 2002-01-01. doi:10.1146/annurev.ecolsys.33.010802.150437.
  9. ^ Parisod, Christian; Holderegger, Rolf; Brochmann, Christian (2010-04-01). "Evolutionary consequences of autopolyploidy". The New Phytologist. 186 (1): 5–17. doi:10.1111/j.1469-8137.2009.03142.x. ISSN 1469-8137. PMID 20070540.
  10. ^ a b Le Comber, S. C.; Ainouche, M. L.; Kovarik, A.; Leitch, A. R. (2010-04-01). "Making a functional diploid: from polysomic to disomic inheritance". The New Phytologist. 186 (1): 113–122. doi:10.1111/j.1469-8137.2009.03117.x. ISSN 1469-8137. PMID 20028473.
  11. ^ a b "Pathways, Mechanisms, and Rates of Polyploid Formation in Flowering Plants". Annual Review of Ecology and Systematics. 29 (1): 467–501. 1998-01-01. doi:10.1146/annurev.ecolsys.29.1.467.
  12. ^ Comai, Luca (2005-11-01). "The advantages and disadvantages of being polyploid". Nature Reviews. Genetics. 6 (11): 836–846. doi:10.1038/nrg1711. ISSN 1471-0056. PMID 16304599.
  13. ^ Ownbey, Marion (1950-01-01). "Natural Hybridization and Amphiploidy in the Genus Tragopogon". American Journal of Botany. 37 (7): 487–499. doi:10.2307/2438023.
  14. ^ Schmid, Michael; Evans, Ben J.; Bogart, James P. "Polyploidy in Amphibia". Cytogenetic and Genome Research. 145 (3–4): 315–330. doi:10.1159/000431388.
  15. ^ Xu, Xun; Xu, Shengkai; Pan, Shifeng; Cheng, Bo; Zhang, Desheng; Mu, Peixiang; Ni, Gengyun; Zhang, Shuang; Yang, Ruiqiang; Li, Jun; Wang, Gisella; Orjeda, Frank; Guzman, Michael; Torres, Roberto; Lozano, Olga; Ponce, Diana; Martinez, Germán; De La Cruz, S. K.; Chakrabarti, Virupaksh U.; Patil, Konstantin G.; Skryabin, Boris B.; Kuznetsov, Nikolai V.; Ravin, Tatjana V.; Kolganova, Alexey V.; Beletsky, Andrei V.; Mardanov, Alex; Di Genova, Daniel M.; Bolser, David M. A.; Martin, Guangcun; Li, Yu (2011). "Genome sequence and analysis of the tuber crop potato". Nature. 475 (7355): 189–95. doi:10.1038/nature10158. PMID 21743474.
  16. ^ Rieger, R.; Michaelis, A.; Green, M.M. (1968). A glossary of genetics and cytogenetics: Classical and molecular.[page needed]
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