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EspañolPOLYPLOIDY
(Redirected from Triploid)
'Polyploidy' is the condition of some biological cells and organisms manifested by the presence of more than two homologous sets of chromosomes. 'Polyploid' types are termed according to the number of chromosome sets in the nucleus: 'triploid' (three sets; 3x), 'tetraploid' (four sets; 4x), 'pentaploid' (five sets; 5x), 'hexaploid' (six sets; 6x) and so on.
A haploid has only one set of chromosomes. Haploidy may also occur as a normal stage in an organism's life cycle as in ferns and fungi. In some instances not all the chromosomes are duplicated and the condition is called aneuploidy. Where an organism is normally diploid, some spontaneous aberrations may occur which are usually caused by a hampered cell division.
Polyploidy occurs in some animals, such as goldfish, salmon, and salamanders, but is especially common among ferns and flowering plants, including both wild and cultivated species. Wheat, for example, after millennia of hybridization and modification by humans, has strains that are 'diploid' (two sets of chromosomes), 'tetraploid' (four sets of chromosomes) with the common name of durum or macaroni wheat, and 'hexaploid' (six sets of chromosomes) with the common name of bread wheat. Many agriculturally important plants of the genus ''Brassica'' are also tetraploids; their relationship is described by the Triangle of U.
The occurrence of polyploidy is a mechanism of speciation and is known to have resulted in new species of the plant Salsify (also known as "goatsbeard").
Examples in animals are more common in the ‘lower’ forms such as flatworms, leeches, and brine shrimp. Polyploid animals are often sterile, so they often reproduce by parthenogenesis. Polyploid salamanders and lizards are also quite common and parthenogenetic. While mammalian liver cells are polyploid, rare instances of polyploid mammals are known, but most often result in prenatal death.
The only known exception to this rule is an octodontid rodent of Argentina's harsh desert regions, known as the Red Viscacha-Rat (''Tympanoctomys barrerae''), discovered by Milton Gallardo Narcisi, member of Universidad Austral de Chile. This rodent is not a rat, but kin to guinea pigs and chinchillas. Its "new" diploid [2n] number is 102 and so its cells are roughly twice normal size. Its closest living relation is ''Octomys mimax'', the Andean Viscacha-Rat of the same family, whose 2n=56. It is surmised that an ''Octomys''-like ancestor produced tetraploid (i.e., 4n=112) offspring that were, by virtue of their doubled chromosomes, reproductively isolated from their parents; but that these likely survived the ordinarily catastrophic effects of polyploidy in mammals by shedding (via translocation or some similar mechanism) the "extra" set of sex chromosomes gained at this doubling.
Polyploidy can be induced in cell culture by some chemicals: the best known is colchicine, which can result in chromosome doubling, though its use may have other less obvious consequences as well.
There are large number of polyploid crop varieties - See Polyploid Crops below.
True polyploidy rarely occurs in humans, although it occurs in some tissues (especially in the liver). Polyploidy refers to a numerical change in a whole set of chromosomes. Organisms in which a particular chromosome, or chromosome segment, is under- or overrepresented are said to be aneuploid (from the Greek words meaning "not," "good," and "fold"). Therefore the distinction between aneuploidy and polyploidy is that aneuploidy refers to a numerical change in part of the chromosome, whereas polyploidy refers to a numerical change in the whole set of chromosomes. [1: Cytogenetic Variation (p109)]
Aneuploidy occurs in humans in the form of triploidy (69,XXX) and tetraploidy (92,XXXX), not to be confused with 47,XXX or 48, XXXX aneuploidy. Triploidy, usually due to polyspermy, occurs in about 2-3% of all human pregnancies and ~15% of miscarriages. The vast majority of triploid conceptions end as miscarriage and those that do survive to term typically die shortly after birth. In some cases survival past birth may occur longer if there is mixoploidy with both a diploid and a triploid cell population present.
Triploidy may be the result of either digyny (the extra haploid set is from the mother) or diandry (the extra haploid set is from the father). Diandry is almost always caused by the fertilization of an egg by two sperm (dispermy). Digyny is most commonly caused by either failure of one meiotic division during oogenesis leading to a diploid oocyte or failure to extrude one polar body from the oocyte. Diandry appears to predominate among early miscarriages while digyny predominates among triploidy that survives into the fetal period. However, among early miscarriages, digyny is also more common in those cases <8.5 weeks gestational age or those in which an embryo is present. There are also two distinct phenotypes in triploid placentas and fetuses that are dependent on the origin of the extra haploid set. In digyny there is typically an asymmetric poorly grown fetus, with marked adrenal hypoplasia and a very small placenta. In diandry, the fetus (when present) is typically normally grown or symmetrically growth restricted, with normal adrenal glands and an abnormally large cystic placenta that is called a partial hydatidiform mole. These parent-of-origin effects reflect the effects of genomic imprinting.
Complete tetraploidy is more rarely diagnosed than triploidy, but is observed in 1-2% of early miscarriages. However, some tetraploid cells are not uncommonly found in chromosome analysis at prenatal diagnosis and these are generally considered ‘harmless’. It is not clear whether these tetraploid cells simply tend to arise during ''in vitro'' cell culture or whether they are also present in placental cells ''in vivo''. There are, at any rate, very few clinical reports of fetuses/infants diagnosed with tetraploidy mosaicism.
Mixoploidy is quite commonly observed in human preimplantation embryos and includes haploid/diploid as well as diploid/tetraploid mixed cell populations. It is unknown whether these embryos fail to implant and are therefore rarely detected in ongoing pregnancies or if there is simply a selective process favoring the diploid cells.
Polyploid plants tend to be larger and better at flourishing in early succession habitats such as farm fields. In the breeding of crops, the tallest and best thriving plants are selected for. Thus, many crops (and agricultural weeds) may have unintentionally been bred to a higher level of ploidy.
The induction of polyploids is a common technique to overcome the sterility of a hybrid species during plant breeding. For example, Triticale is the hybrid of wheat (''Triticum turgidum'') and rye (''Secale cereale''). It combines sought-after characteristics of the parents, but the initial hybrids are sterile. After polyploidization, the hybrid becomes fertile and can thus be further propagated to become triticale.
In some situations polyploid crops are preferred because they are sterile. For example many seedless fruit varieties are seedless as a result of polyploidy. Such crops are propagated using asexual techniques such as grafting.
Polyploidy in crop plants is most commonly induced by treating seeds with the chemical colchicine.
★ Triploid crops: banana, apple, ginger, citrus [1]
★ Tetraploid crops: durum or macaroni wheat, maize, cotton, potato, cabbage, leek, tobacco, peanut, kinnow, Pelargonium
★ Hexaploid crops: chrysanthemum, bread wheat, triticale, oat
★ Octaploid crops: strawberry, dahlia, pansies, sugar cane
Some crops are found in a variety of ploidy. Apples, tulips and lilies are commonly found as both diploid and as triploid. Daylilies (Hemerocallis) cultivars are available as either diploid or tetraploid. Kinnows can be tetraploid, diploid, or triploid.
Refers to genes duplicated by polyploidy.
Autopolyploids are polyploids with chromosomes derived from a single species. Autopolyploids can arise from a spontaneous, naturally-occurring genome doubling (for example, the potato). Bananas and apples can be found as triploid autopolyploids.
Allopolyploids are polyploids with chromosomes derived from different species. Triticale is an example of an allopolyploid, having six chromosome sets, four from wheat (Triticum turgidum) and two from rye (Secale cereale). Cabbage is a very interesting example of a fertile allotetraploid crop (see Triangle of U). 'Amphidiploid' is another word for an allopolyploid.
The giant tree ''Sequoia sempervirens'' or Coast Redwood has a hexaploid (6n) genome, and is also thought to be autoallopolyploid (AAAABB). It is the only polyploid conifer species known.
Main articles: Paleopolyploidy
Ancient genome duplications probably characterize all life. Duplication events that occurred long ago in the history of various evolutionary lineages can be difficult to detect because of subsequent diploidization (such that a polyploid starts to behave cytogenetically as a diploid over time) as mutations and gene translations gradually make one copy of each chromosome unlike its other copy.
In many cases, these events can be inferred only through comparing sequenced genomes. Examples of unexpected but recently confirmed ancient genome duplications include the baker's yeast (''Saccharomyces cerevisiae''), mustard weed/thale cress (''Arabidopsis thaliana''), rice (''Oryza sativa''), and an early evolutionary ancestor of the vertebrates (which includes the human lineage) and another near the origin of the teleost fishes. Angiosperms (flowering plants) have paleopolyploidy in their ancestry. All eukaryotes probably have experienced a polyploidy event at some point in their evolutionary history.
★ Speciation
★ Polyploid complex
★ Paleopolyploidy
[1] Griffiths, A. J. et al. 2000. An introduction to genetic analysis, 7th ed. W. H. Freeman, New York ISBN 0-7167-3520-2
[2] Snustad, P. et al. 2006. Principles of Genetics, 4th ed. John Wiley & Sons, Inc. Hoboken, NJ ISBN 10 0-471-69939-X
★ Arabidopsis Genome Initiative (2000). Analysis of the genome sequence of the flowering plant ''Arabidopsis thaliana''. ''Nature'' 408: 796-815.
★ Eakin, G.S. & Behringer, R.R. (2003). Tetraploid development in the mouse. ''Developmental Dynamics'' 228: 751-766.
★ Gregory, T.R. & Mable, B.K. (2005). Polyploidy in animals. In The Evolution of the Genome (edited by T.R. Gregory). Elsevier, San Diego, pp. 427-517.
★ Jaillon, O. et al. (2004). Genome duplication in the teleost fish ''Tetraodon nigroviridis'' reveals the early vertebrate proto-karyotype. ''Nature'' 431: 946-957.
★ Paterson, A.H., Bowers, J. E., Van de Peer, Y. & Vandepoele, K. (2005). Ancient duplication of cereal genomes. ''New Phytologist'' 165: 658-661.
★ Raes, J., Vandepoele, K., Saeys, Y., Simillion, C. & Van de Peer, Y. (2003). Investigating ancient duplication events in the ''Arabidopsis'' genome. ''Journal of Structural and Functional Genomics'' 3: 117-129.
★ Simillion, C., Vandepoele, K., Van Montagu, M., Zabeau, M. & Van de Peer, Y. (2002). The hidden duplication past of ''Arabidopsis thaliana''. ''Proceedings of the National Academy of Science of the USA'' 99: 13627-13632.
★ Soltis, D. E.; Soltis, P. S.; Schemske, D. W.; Hancock, J. F.; Thompson, J. N.; Husband, B. C. & Judd, W. S. (2007).Autopolyploidy in angiosperms: have we grossly underestimated the number of species? ''Taxon'' 56 (1):13-30.
★ Taylor, J.S., Braasch, I., Frickey, T., Meyer, A. & Van de Peer, Y. (2003). Genome duplication, a trait shared by 22,000 species of ray-finned fish. ''Genome Research'' 13: 382-390.
★ Tate, J.A., Soltis, D.E., & Soltis, P.S. (2005). Polyploidy in plants. In The Evolution of the Genome (edited by T.R. Gregory). Elsevier, San Diego, pp.371-426.
★ Van de Peer, Y., Taylor, J.S. & Meyer, A. (2003). Are all fishes ancient polyploids? ''Journal of Structural and Functional Genomics'' 3: 65-73.
★ Van de Peer, Y. (2004). ''Tetraodon'' genome confirms ''Takifugu'' findings: most fish are ancient polyploids. ''Genome Biology'' 5(12):250.
★ Van de Peer, Y. and Meyer, A. (2005). Large-scale gene and ancient genome duplications. In The Evolution of the Genome (edited by T.R. Gregory). Elsevier, San Diego, pp.329-368
★ Wolfe, K.H. & Shields, D.C. (1997). Molecular evidence for an ancient duplication of the entire yeast genome. ''Nature'' 387: 708-713.
★ Wolfe, K.H. (2001). Yesterday's polyploids and the mystery of diploidization. ''Nature Reviews Genetics'' 2: 333-341.
★ Polyploidy on Kimball's Biology Pages
★
★ The polyploidy portal a community-editable project with information, research, education, and a bibliography about polyploidy.
'Polyploidy' is the condition of some biological cells and organisms manifested by the presence of more than two homologous sets of chromosomes. 'Polyploid' types are termed according to the number of chromosome sets in the nucleus: 'triploid' (three sets; 3x), 'tetraploid' (four sets; 4x), 'pentaploid' (five sets; 5x), 'hexaploid' (six sets; 6x) and so on.
A diagram that summarizes all well-known polyploidization events. See Paleopolyploidy for more detail
A haploid has only one set of chromosomes. Haploidy may also occur as a normal stage in an organism's life cycle as in ferns and fungi. In some instances not all the chromosomes are duplicated and the condition is called aneuploidy. Where an organism is normally diploid, some spontaneous aberrations may occur which are usually caused by a hampered cell division.
Examples
Polyploidy occurs in some animals, such as goldfish, salmon, and salamanders, but is especially common among ferns and flowering plants, including both wild and cultivated species. Wheat, for example, after millennia of hybridization and modification by humans, has strains that are 'diploid' (two sets of chromosomes), 'tetraploid' (four sets of chromosomes) with the common name of durum or macaroni wheat, and 'hexaploid' (six sets of chromosomes) with the common name of bread wheat. Many agriculturally important plants of the genus ''Brassica'' are also tetraploids; their relationship is described by the Triangle of U.
The occurrence of polyploidy is a mechanism of speciation and is known to have resulted in new species of the plant Salsify (also known as "goatsbeard").
Examples in animals are more common in the ‘lower’ forms such as flatworms, leeches, and brine shrimp. Polyploid animals are often sterile, so they often reproduce by parthenogenesis. Polyploid salamanders and lizards are also quite common and parthenogenetic. While mammalian liver cells are polyploid, rare instances of polyploid mammals are known, but most often result in prenatal death.
The only known exception to this rule is an octodontid rodent of Argentina's harsh desert regions, known as the Red Viscacha-Rat (''Tympanoctomys barrerae''), discovered by Milton Gallardo Narcisi, member of Universidad Austral de Chile. This rodent is not a rat, but kin to guinea pigs and chinchillas. Its "new" diploid [2n] number is 102 and so its cells are roughly twice normal size. Its closest living relation is ''Octomys mimax'', the Andean Viscacha-Rat of the same family, whose 2n=56. It is surmised that an ''Octomys''-like ancestor produced tetraploid (i.e., 4n=112) offspring that were, by virtue of their doubled chromosomes, reproductively isolated from their parents; but that these likely survived the ordinarily catastrophic effects of polyploidy in mammals by shedding (via translocation or some similar mechanism) the "extra" set of sex chromosomes gained at this doubling.
Polyploidy can be induced in cell culture by some chemicals: the best known is colchicine, which can result in chromosome doubling, though its use may have other less obvious consequences as well.
There are large number of polyploid crop varieties - See Polyploid Crops below.
Polyploidy in humans (Aneuploidy)
True polyploidy rarely occurs in humans, although it occurs in some tissues (especially in the liver). Polyploidy refers to a numerical change in a whole set of chromosomes. Organisms in which a particular chromosome, or chromosome segment, is under- or overrepresented are said to be aneuploid (from the Greek words meaning "not," "good," and "fold"). Therefore the distinction between aneuploidy and polyploidy is that aneuploidy refers to a numerical change in part of the chromosome, whereas polyploidy refers to a numerical change in the whole set of chromosomes. [1: Cytogenetic Variation (p109)]
Aneuploidy occurs in humans in the form of triploidy (69,XXX) and tetraploidy (92,XXXX), not to be confused with 47,XXX or 48, XXXX aneuploidy. Triploidy, usually due to polyspermy, occurs in about 2-3% of all human pregnancies and ~15% of miscarriages. The vast majority of triploid conceptions end as miscarriage and those that do survive to term typically die shortly after birth. In some cases survival past birth may occur longer if there is mixoploidy with both a diploid and a triploid cell population present.
Triploidy may be the result of either digyny (the extra haploid set is from the mother) or diandry (the extra haploid set is from the father). Diandry is almost always caused by the fertilization of an egg by two sperm (dispermy). Digyny is most commonly caused by either failure of one meiotic division during oogenesis leading to a diploid oocyte or failure to extrude one polar body from the oocyte. Diandry appears to predominate among early miscarriages while digyny predominates among triploidy that survives into the fetal period. However, among early miscarriages, digyny is also more common in those cases <8.5 weeks gestational age or those in which an embryo is present. There are also two distinct phenotypes in triploid placentas and fetuses that are dependent on the origin of the extra haploid set. In digyny there is typically an asymmetric poorly grown fetus, with marked adrenal hypoplasia and a very small placenta. In diandry, the fetus (when present) is typically normally grown or symmetrically growth restricted, with normal adrenal glands and an abnormally large cystic placenta that is called a partial hydatidiform mole. These parent-of-origin effects reflect the effects of genomic imprinting.
Complete tetraploidy is more rarely diagnosed than triploidy, but is observed in 1-2% of early miscarriages. However, some tetraploid cells are not uncommonly found in chromosome analysis at prenatal diagnosis and these are generally considered ‘harmless’. It is not clear whether these tetraploid cells simply tend to arise during ''in vitro'' cell culture or whether they are also present in placental cells ''in vivo''. There are, at any rate, very few clinical reports of fetuses/infants diagnosed with tetraploidy mosaicism.
Mixoploidy is quite commonly observed in human preimplantation embryos and includes haploid/diploid as well as diploid/tetraploid mixed cell populations. It is unknown whether these embryos fail to implant and are therefore rarely detected in ongoing pregnancies or if there is simply a selective process favoring the diploid cells.
Polyploid crops
Polyploid plants tend to be larger and better at flourishing in early succession habitats such as farm fields. In the breeding of crops, the tallest and best thriving plants are selected for. Thus, many crops (and agricultural weeds) may have unintentionally been bred to a higher level of ploidy.
The induction of polyploids is a common technique to overcome the sterility of a hybrid species during plant breeding. For example, Triticale is the hybrid of wheat (''Triticum turgidum'') and rye (''Secale cereale''). It combines sought-after characteristics of the parents, but the initial hybrids are sterile. After polyploidization, the hybrid becomes fertile and can thus be further propagated to become triticale.
In some situations polyploid crops are preferred because they are sterile. For example many seedless fruit varieties are seedless as a result of polyploidy. Such crops are propagated using asexual techniques such as grafting.
Polyploidy in crop plants is most commonly induced by treating seeds with the chemical colchicine.
Examples of Polyploid Crops
★ Triploid crops: banana, apple, ginger, citrus [1]
★ Tetraploid crops: durum or macaroni wheat, maize, cotton, potato, cabbage, leek, tobacco, peanut, kinnow, Pelargonium
★ Hexaploid crops: chrysanthemum, bread wheat, triticale, oat
★ Octaploid crops: strawberry, dahlia, pansies, sugar cane
Some crops are found in a variety of ploidy. Apples, tulips and lilies are commonly found as both diploid and as triploid. Daylilies (Hemerocallis) cultivars are available as either diploid or tetraploid. Kinnows can be tetraploid, diploid, or triploid.
Terminology
Homeolog
Refers to genes duplicated by polyploidy.
Autopolyploidy
Autopolyploids are polyploids with chromosomes derived from a single species. Autopolyploids can arise from a spontaneous, naturally-occurring genome doubling (for example, the potato). Bananas and apples can be found as triploid autopolyploids.
Allopolyploidy
Allopolyploids are polyploids with chromosomes derived from different species. Triticale is an example of an allopolyploid, having six chromosome sets, four from wheat (Triticum turgidum) and two from rye (Secale cereale). Cabbage is a very interesting example of a fertile allotetraploid crop (see Triangle of U). 'Amphidiploid' is another word for an allopolyploid.
The giant tree ''Sequoia sempervirens'' or Coast Redwood has a hexaploid (6n) genome, and is also thought to be autoallopolyploid (AAAABB). It is the only polyploid conifer species known.
Paleopolyploidy
Main articles: Paleopolyploidy
Ancient genome duplications probably characterize all life. Duplication events that occurred long ago in the history of various evolutionary lineages can be difficult to detect because of subsequent diploidization (such that a polyploid starts to behave cytogenetically as a diploid over time) as mutations and gene translations gradually make one copy of each chromosome unlike its other copy.
In many cases, these events can be inferred only through comparing sequenced genomes. Examples of unexpected but recently confirmed ancient genome duplications include the baker's yeast (''Saccharomyces cerevisiae''), mustard weed/thale cress (''Arabidopsis thaliana''), rice (''Oryza sativa''), and an early evolutionary ancestor of the vertebrates (which includes the human lineage) and another near the origin of the teleost fishes. Angiosperms (flowering plants) have paleopolyploidy in their ancestry. All eukaryotes probably have experienced a polyploidy event at some point in their evolutionary history.
See also
★ Speciation
★ Polyploid complex
★ Paleopolyploidy
References
[1] Griffiths, A. J. et al. 2000. An introduction to genetic analysis, 7th ed. W. H. Freeman, New York ISBN 0-7167-3520-2
[2] Snustad, P. et al. 2006. Principles of Genetics, 4th ed. John Wiley & Sons, Inc. Hoboken, NJ ISBN 10 0-471-69939-X
Further reading
★ Arabidopsis Genome Initiative (2000). Analysis of the genome sequence of the flowering plant ''Arabidopsis thaliana''. ''Nature'' 408: 796-815.
★ Eakin, G.S. & Behringer, R.R. (2003). Tetraploid development in the mouse. ''Developmental Dynamics'' 228: 751-766.
★ Gregory, T.R. & Mable, B.K. (2005). Polyploidy in animals. In The Evolution of the Genome (edited by T.R. Gregory). Elsevier, San Diego, pp. 427-517.
★ Jaillon, O. et al. (2004). Genome duplication in the teleost fish ''Tetraodon nigroviridis'' reveals the early vertebrate proto-karyotype. ''Nature'' 431: 946-957.
★ Paterson, A.H., Bowers, J. E., Van de Peer, Y. & Vandepoele, K. (2005). Ancient duplication of cereal genomes. ''New Phytologist'' 165: 658-661.
★ Raes, J., Vandepoele, K., Saeys, Y., Simillion, C. & Van de Peer, Y. (2003). Investigating ancient duplication events in the ''Arabidopsis'' genome. ''Journal of Structural and Functional Genomics'' 3: 117-129.
★ Simillion, C., Vandepoele, K., Van Montagu, M., Zabeau, M. & Van de Peer, Y. (2002). The hidden duplication past of ''Arabidopsis thaliana''. ''Proceedings of the National Academy of Science of the USA'' 99: 13627-13632.
★ Soltis, D. E.; Soltis, P. S.; Schemske, D. W.; Hancock, J. F.; Thompson, J. N.; Husband, B. C. & Judd, W. S. (2007).Autopolyploidy in angiosperms: have we grossly underestimated the number of species? ''Taxon'' 56 (1):13-30.
★ Taylor, J.S., Braasch, I., Frickey, T., Meyer, A. & Van de Peer, Y. (2003). Genome duplication, a trait shared by 22,000 species of ray-finned fish. ''Genome Research'' 13: 382-390.
★ Tate, J.A., Soltis, D.E., & Soltis, P.S. (2005). Polyploidy in plants. In The Evolution of the Genome (edited by T.R. Gregory). Elsevier, San Diego, pp.371-426.
★ Van de Peer, Y., Taylor, J.S. & Meyer, A. (2003). Are all fishes ancient polyploids? ''Journal of Structural and Functional Genomics'' 3: 65-73.
★ Van de Peer, Y. (2004). ''Tetraodon'' genome confirms ''Takifugu'' findings: most fish are ancient polyploids. ''Genome Biology'' 5(12):250.
★ Van de Peer, Y. and Meyer, A. (2005). Large-scale gene and ancient genome duplications. In The Evolution of the Genome (edited by T.R. Gregory). Elsevier, San Diego, pp.329-368
★ Wolfe, K.H. & Shields, D.C. (1997). Molecular evidence for an ancient duplication of the entire yeast genome. ''Nature'' 387: 708-713.
★ Wolfe, K.H. (2001). Yesterday's polyploids and the mystery of diploidization. ''Nature Reviews Genetics'' 2: 333-341.
External links
★ Polyploidy on Kimball's Biology Pages
★
★ The polyploidy portal a community-editable project with information, research, education, and a bibliography about polyploidy.
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