Hybrid (biology)

In biology and specifically, genetics, the term hybrid has several meanings, all referring to the offspring of sexual reproduction.[1]

  1. In general usage, hybrid is synonymous with heterozygous: any offspring resulting from the mating of two distinctly homozygous individuals
  2. a genetic hybrid carries two different alleles of the same gene
  3. a structural hybrid results from the fusion of gametes that have differing structure in at least one chromosome, as a result of structural abnormalities
  4. a numerical hybrid results from the fusion of gametes having different haploid numbers of chromosomes
  5. a permanent hybrid is a situation where only the heterozygous genotype occurs, because all homozygous combinations are lethal.

From a taxonomic perspective, hybrid refers to offspring resulting from the interbreeding between two animals or plants of different taxa.[2]

  1. Hybrids between different subspecies within a species (such as between the Bengal tiger and Siberian tiger) are known as intra-specific hybrids. Hybrids between different species within the same genus (such as between lions and tigers) are sometimes known as interspecific hybrids or crosses. Hybrids between different genera (such as between sheep and goats) are known as intergeneric hybrids. Extremely rare interfamilial hybrids have been known to occur (such as the guineafowl hybrids).[3] No interordinal (between different orders) animal hybrids are known.
  2. The second type of hybrid consists of crosses between populations, breeds or cultivars within a single species. This meaning is often used in plant and animal breeding, where hybrids are commonly produced and selected because they have desirable characteristics not found or inconsistently present in the parent individuals or populations. This flow of genetic material between populations or races is often called hybridization.

Contents

Etymology

According to the Oxford English Dictionary, the word is derived from Latin hybrida, meaning the "offspring of a tame sow and a wild boar", "child of a freeman and slave", etc.[4] The term entered into popular use in English in the 19th century, though examples of its use have been found from the early 17th century.[5]

Types of hybrids

Depending on the parents, there are a number of different types of hybrids;[6]

Interspecific hybrids

Interspecific hybrids are bred by mating two species, normally from within the same genus. The offspring display traits and characteristics of both parents. The offspring of an interspecific cross are very often sterile; thus, hybrid sterility prevents the movement of genes from one species to the other, keeping both species distinct.[10] Sterility is often attributed to the different number of chromosomes the two species have, for example donkeys have 62 chromosomes, while horses have 64 chromosomes, and mules and hinnies have 63 chromosomes. Mules, hinnies, and other normally sterile interspecific hybrids cannot produce viable gametes, because differences in chromosome structure prevent appropriate pairing and segregation during meiosis, meiosis is disrupted, and viable sperm and eggs are not formed. However, fertility in female mules has been reported with a donkey as the father.[11]

Most often other processes occurring in plants and animals keep gametic isolation and species distinction. Species often have different mating or courtship patterns or behaviors, the breeding seasons may be distinct and even if mating does occur antigenic reactions to the sperm of other species prevent fertilization or embryo development. Hybridisation is much more common among that spawn indiscriminately, like soft corals and among plants.

While it is possible to predict the genetic composition of a backcross on average, it is not possible to accurately predict the composition of a particular backcrossed individual, due to random segregation of chromosomes. In a species with two pairs of chromosomes, a twice backcrossed individual would be predicted to contain 12.5% of one species' genome (say, species A). However, it may, in fact, still be a 50% hybrid if the chromosomes from species A were lucky in two successive segregations, and meiotic crossovers happened near the telomeres. The chance of this is fairly high: \left(\frac{1}{2}\right)^{(2 \times 2)} = \frac{1}{16} (where the "two times two" comes about from two rounds of meiosis with two chromosomes); however, this probability declines markedly with chromosome number and so the actual composition of a hybrid will be increasingly closer to the predicted composition.

Hybrids are often named by the portmanteau method, combining the names of the two parent species. For example, a zeedonk is a cross between a zebra and a donkey. Since the traits of hybrid offspring often vary depending on which species was mother and which was father, it is traditional to use the father's species as the first half of the portmanteau. For example, a liger is a cross between a male lion and a female tiger, while a tiglon is a cross between a male tiger and a female lion.

Domestic and wild hybrids

Hybrids between domesticated and wild animals in particular may be problematic. Breeders of domesticated species discourage crossbreeding with wild species, unless a deliberate decision is made to incorporate a trait of a wild ancestor back into a given breed or strain. Wild populations of animals and plants have evolved naturally over millions of years through a process of natural selection in contrast to human controlled selective breeding or artificial selection for desirable traits from the human point of view. Normally, these two methods of reproduction operate independently of one another. However, an intermediate form of selective breeding, wherein animals or plants are bred by humans, but with an eye to adaptation to natural region-specific conditions and an acceptance of natural selection to weed out undesirable traits, created many ancient domesticated breeds or types now known as landraces.

Many times, domesticated species live in or near areas which also still hold naturally evolved, region-specific wild ancestor species and subspecies. In some cases, a domesticated species of plant or animal may become feral, living wild. Other times, a wild species will come into an area inhabited by a domesticated species. Some of these situations lead to the creation of hybridized plants or animals, a cross between the native species and a domesticated one. This type of crossbreeding, termed genetic pollution by those who are concerned about preserving the genetic base of the wild species, has become a major concern. Hybridization is also a concern to the breeders of purebred species as well, particularly if the gene pool is small and if such crossbreeding or hybridization threatens the genetic base of the domesticated purebred population.

The concern with genetic pollution of a wild population is that hybridized animals and plants may not be as genetically strong as naturally evolved region specific wild ancestors wildlife which can survive without human husbandry and have high immunity to natural diseases. The concern of purebred breeders with wildlife hybridizing a domesticated species is that it can coarsen or degrade the specific qualities of a breed developed for a specific purpose, sometimes over many generations. Thus, both purebred breeders and wildlife biologists share a common interest in preventing accidental hybridization.

Hybrid species

Main article: Hybrid speciation

While not very common, a few animal species has be recognized as being the result of hybridization. The Lonicera fly is an example of a novel animal species that resulted from natural hybridization. The American red wolf too appear to be a hybrid species between gray wolf and coyote.[12] The European edible frog appear a species, but is actually semi-permanent hybrids between pool frogs and marsh frogs, the edible frog population being dependent on the presence of at least one of the parents species to be maintained.[13]

Hybrid species of plants are a lot more common than animals. Many of the crop species are hybrids, and hybridization appear to be an important factor in speciation in some plant groups.

Examples of hybrid animals

Hybrids should not be confused with genetic chimeras such as that between sheep and goat known as the geep. Wider interspecific hybrids can be made via in vitro fertilization or somatic hybridization, however the resulting cells are not able to develop into a full organism. An example of interspecific hybrid cell lines is humster (hamster x human) cells.

Hybrid plants

Plant species hybridize more readily than animal species, and the resulting hybrids are more often fertile hybrids and may reproduce, though there still exist sterile hybrids and selective hybrid elimination where the offspring are less able to survive and are thus eliminated before they can reproduce. A number of plant species are the result of hybridization and polyploidy with many plant species easily cross pollinating and producing viable seeds, the distinction between each species is often maintained by geographical isolation or differences in the flowering period. Since plants hybridize frequently without much work, they are often created by humans in order to produce improved plants. These improvements can include the production of more or improved; seeds, fruits or other plant parts for consumption, or to make a plant more winter or heat hardy or improve its growth and/or appearance for use in horticulture. Much work is now being done with hybrids to produce more disease resistant plants for both agricultural and horticultural crops. In many groups of plants hybridization has been used to produce larger and more showy flowers and new flower colors.

Many plant genera and species have their origins in polyploidy. Autopolyploidy resulting from the sudden multiplication in the number of chromosomes in typical normal populations caused by unsuccessful separation of the chromosomes during meiosis. Tetraploids or plants with four sets of chromosomes are common in a number of different groups of plants and over time these plants can differentiate into distinct species from the normal diploid line. In Oenothera lamarchiana the diploid species has 14 chromosomes, this species has spontaneously given rise to plants with 28 chromosomes that have been given the name Oenthera gigas. Tetraploids can develop into a breeding population within the diploid population and when hybrids are formed with the diploid population the resulting offspring tend to be sterile triploids, thus effectively stopping the intermixing of genes between the two groups of plants (unless the diploids, in rare cases, produce unreduced gametes).

Another form of polyploidy called allopolyploidy occurs when two different species mate and produce hybrids. Usually the typical chromosome number is doubled in successful allopolyploid species, with four sets of chromosomes the genotypes can sort out to form a complete diploid set from the parent species, thus they can produce fertile offspring that can mate and reproduce with each other but can not back-cross with the parent species. Allopolyploidy in plants often gives them a condition called hybrid vigour, which results in plants that are larger and stronger growing than either of the two parent species. Allopolyploids are often more aggressive growing and can be invaders of new habitats.

Sterility in a hybrid is often a result of chromosome number; if parents are of differing chromosome pair number, the offspring will have an odd number of chromosomes, leaving them unable to produce chromosomally balanced gametes.[16] While this is a negative in a crop such as wheat, when growing a crop which produces no seeds would be pointless, it is an attractive attribute in some fruits. Bananas and seedless watermelon, for instance, are intentionally bred to be triploid, so that they will produce no seeds. Many hybrids are created by humans, but natural hybrids occur as well.

Heterosis

Main article: heterosis

Hybrids are sometimes stronger than either parent variety, a phenomenon most common with plant hybrids, which when present is known as hybrid vigor (heterosis) or heterozygote advantage.[17] A transgressive phenotype is a phenotype displaying more extreme characteristics than either of the parent lines.[18] Plant breeders make use of a number of techniques to produce hybrids, including line breeding and the formation of complex hybrids. An economically important example is hybrid maize (corn), which provides a considerable seed yield advantage over open pollinated varieties. Hybrid seed dominates the commercial maize seed market in the United States, Canada and many other major maize producing countries.[19]

Examples of species hybrids

Some plant hybrids include:

Some natural hybrids are:

Some horticultural hybrids:

Hybrids in nature

Hybridisation between two closely related species is actually a common occurrence in nature but is also being greatly influenced by anthropogenic changes as well.[20] Hybridization is a naturally occurring genetic process where individuals from two genetically distinct populations mate.[21] As stated above, it can occur both intraspecifically, between different distinct populations within the same species, and interspecifically, between two different species. Hybrids can be either sterile/not viable or viable/fertile. This affects the kind of effect that this hybrid will have on its and other populations that it interacts with.[22] Many hybrid zones are known where the ranges of two species meet, and hybrids are continually produced in great numbers. These hybrid zones are useful as biological model systems for studying the mechanisms of speciation (Hybrid speciation). Recently DNA analysis of a bear shot by a hunter in the North West Territories confirmed the existence of naturally-occurring and fertile grizzly–polar bear hybrids.[23] There have been reports of similar supposed hybrids, but this is the first to be confirmed by DNA analysis. In 1943, Clara Helgason described a male bear shot by hunters during her childhood. It was large and off-white with hair all over its paws. The presence of hair on the bottom of the feet suggests it was a natural hybrid of Kodiak and Polar bear.

Anthropogenic hybridization

Anthropogenic caused changes to the environment such as fragmentation and Introduced species are becoming more widespread.[24] This is allowing another kind of hybridization that is more of the focus of conservation genetics to occur: anthropogenic hybridization. This anthropogenically caused hybridization increases the challenges in managing certain populations that are experiencing introgression.

Introduced species and habitat fragmentation

Humans have been introducing species world wide to environments for a long time both directly such as establishing a population to be used as a biological control and indirectly such as accidental escapes of individuals out of agriculture. This causes drastic global affects on various populations with hybridization being one of the reasons introduced species can be so detrimental.[22][25] When habitats become broken apart, one of two things can occur, genetically speaking. The first is that populations that were once connected can be cut off from one another, preventing their genes from interacting. Occasionally, this will result in a population of one species breeding with a population of another species as a means of surviving such as the case with the red wolves. Their population numbers being so small, they needed another means of survival. Habitat fragmentation also led to the influx of generalist species into areas where they would not have been, leading to competition and in some cases interbreeding/incorporation of a population into another. In this way, habitat fragmentation is essentially an indirect method of introducing species to an area.

The hybridization continuum

There is a kind of continuum with three semi-distinct categories dealing with anthropogenic hybridization: hybridization without Introgression, hybridization with widespread introgression, and essentially a Hybrid swarm.[20] Depending on where a population falls along this continuum, the management plans for that population will change. Hybridization is currently an area of great discussion within Wildlife management and habitat management fields. Global climate change, which is also anthropogenically caused, is creating other changes such as difference in population distributions which are indirect causes for an increase in anthropogenic hybridization.

Consequences

Hybridization can be a less discussed way toward Extinction then within detection of where a population lies along the hybrid continuum. The dispute of hybridization is how to manage the resulting hybrids. When a population experiences hybridization with substantial introgression, there still exists parent types of each set of individuals. When a complete hybrid swarm is created, all the individuals are hybrids.

Management of hybrids

Conservationists disagree on when is the proper time to give up on a population that is becoming a hybrid swarm or to try and save the still existing pure individuals. Once it becomes a complete mixture, we should look to conserve those hybrids to avoid their loss.[20] Most leave it as a case-by-case basis, depending on detecting of hybrids within the group. It is nearly impossible to regulate hybridization via policy because hybridization can occur beneficially when it occurs "naturally" and there is the matter of protecting those previously mentioned hybrid swarms because if they are the only remaining evidence of prior species, they need to be conserved as well.[20]

In some species, hybridisation plays an important role in evolutionary biology. While most hybrids are disadvantaged as a result of genetic incompatibility, the fittest survive, regardless of species boundaries. They may have a beneficial combination of traits allowing them to exploit new habitats or to succeed in a marginal habitat where the two parent species are disadvantaged. This has been seen in experiments on sunflower species. Unlike mutation, which affects only one gene, hybridisation creates multiple variations across genes or gene combinations simultaneously. Successful hybrids could evolve into new species within 50-60 generations. This leads some scientists to speculate that life is a genetic continuum rather than a series of self-contained species.

Where there are two closely related species living in the same area, less than 1 in 1000 individuals are likely to be hybrids because animals rarely choose a mate from a different species (otherwise species boundaries would completely break down). In some closely related species there are recognized "hybrid zones".

Some species of Heliconius butterflies exhibit dramatic geographical polymorphism of their wing patterns, which act as aposematic signals advertising their unpalatability to potential predators. Where different-looking geographical races abut, inter-racial hybrids are common, healthy and fertile. Heliconius hybrids can breed with other hybrid individuals and with individuals of either parental race. These hybrid backcrosses are disadvantaged by natural selection because they lack the parental form's warning coloration, and are therefore not avoided by predators.

A similar case in mammals is hybrid White-Tail/Mule Deer. The hybrids don't inherit either parent's escape strategy. White-tail Deer dash while Mule Deer bound. The hybrids are easier prey than the parent species.

In birds, healthy Galapagos Finch hybrids are relatively common, but their beaks are intermediate in shape and less efficient feeding tools than the specialised beaks of the parental species so they lose out in the competition for food. Following a major storm in 1983, the local habitat changed so that new types of plants began to flourish, and in this changed habitat, the hybrids had an advantage over the birds with specialised beaks - demonstrating the role of hybridization in exploiting new ecological niches. If the change in environmental conditions is permanent or is radical enough that the parental species cannot survive, the hybrids become the dominant form. Otherwise, the parental species will re-establish themselves when the environmental change is reversed, and hybrids will remain in the minority.

Natural hybrids may occur when a species is introduced into a new habitat. In Britain, there is hybridisation of native European Red Deer and introduced Chinese Sika Deer. Conservationists want to protect the Red Deer, but the environment favors the Sika Deer genes. There is a similar situation with White-headed Ducks and Ruddy Ducks.

Expression of parental traits in hybrids

When two distinct types of organisms breed with each other, the resulting hybrids typically have intermediate traits (e.g., one parent has red flowers, the other has white, and the hybrid, pink flowers).[26] Commonly, hybrids also combine traits seen only separately in one parent or the other (e.g., a bird hybrid might combine the yellow head of one parent with the orange belly of the other).[26] Most characteristics of the typical hybrid are of one of these two types, and so, in a strict sense, are not really new. However, an intermediate trait does differ from those seen in the parents (e.g., the pink flowers of the intermediate hybrid just mentioned are not seen in either of its parents). Likewise, combined traits are new when viewed as a combination.

In a hybrid, any trait that falls outside the range of parental variation is termed heterotic. Heterotic hybrids do have new traits, that is, they are not intermediate. Positive heterosis produces more robust hybrids, they might be stronger or bigger; while the term negative heterosis refers to weaker or smaller hybrids.[27] Heterosis is common in both animal and plant hybrids. For example, hybrids between a lion and a tigress ("ligers") are much larger than either of the two progenitors, while a tigon (lioness × tiger) is smaller. Also the hybrids between the Common Pheasant (Phasianus colchicus) and domestic fowl (Gallus gallus) are larger than either of their parents, as are those produced between the Common Pheasant and hen Golden Pheasant (Chrysolophus pictus).[28] Spurs are absent in hybrids of the former type, although present in both parents.[29]

When populations hybridize, often the first generation (F1) hybrids are very uniform. Typically, however, the individual members of subsequent hybrid generations are quite variable. High levels of variability in a natural population, then, are indicative of hybridity. Researchers use this fact to ascertain whether a population is of hybrid origin. Since such variability generally occurs only in later hybrid generations, the existence of variable hybrids is also an indication that the hybrids in question are fertile.

Genetic mixing and extinction

Main article: Genetic pollution

Regionally developed ecotypes can be threatened with extinction when new alleles or genes are introduced that alter that ecotype. This is sometimes called genetic mixing.[30] Hybridization and introgression of new genetic material can lead to the replacement of local genotypes if the hybrids are more fit and have breeding advantages over the indigenous ecotype or species. These hybridization events can result from the introduction of non native genotypes by humans or through habitat modification, bringing previously isolated species into contact. Genetic mixing can be especially detrimental for rare species in isolated habitats, ultimately affecting the population to such a degree that none of the originally genetically distinct population remains.[31][32]

Effect on biodiversity and food security

Main articles: biodiversity and food security

In agriculture and animal husbandry, the Green Revolution's use of conventional hybridization increased yields by breeding "high-yielding varieties". The replacement of locally indigenous breeds, compounded with unintentional cross-pollination and crossbreeding (genetic mixing), has reduced the gene pools of various wild and indigenous breeds resulting in the loss of genetic diversity.[33] Since the indigenous breeds are often well-adapted to local extremes in climate and have immunity to local pathogens this can be a significant genetic erosion of the gene pool for future breeding. Therefore, commercial plant geneticists strive to breed "widely adapted" cultivars to counteract this tendency.[34]

Limiting factors

A number of conditions exist that limit the success of hybridization, the most obvious is great genetic diversity between most species. But in animals and plants that are more closely related hybridization barriers can include morphological differences, differing times of fertility, mating behaviors and cues, physiological rejection of sperm cells or the developing embryo.

In plants, barriers to hybridization include blooming period differences, different pollinator vectors, inhibition of pollen tube growth, somatoplastic sterility, cytoplasmic-genic male sterility and structural differences of the chromosomes.[35]

Mythical and legendary hybrids

Ancient folktales often contain mythological creatures, sometimes these are described as hybrids (e.g., Hippogriff as the offspring of a griffin and a horse, and the Minotaur which is the offspring of Pasiphaë and a white bull). More often they are kind of chimera, i.e., a composite of the physical attributes of two or more kinds of animals, mythical beasts, and often humans, with no suggestion that they are the result of interbreeding, e.g., Harpies, mermaids, and centaurs.

See also

References

  1. ^ Rieger, R.; Michaelis A.; Green, M. M. (1991). Glossary of Genetics (5th ed.). Springer-Verlag. ISBN 0-387-52054-6 page 256
  2. ^ Keeton, William T. 1980. Biological science. New York: Norton. ISBN 0-393-95021-2 page A9.
  3. ^ Ghigi A. 1936. "Galline di faraone e tacchini" Milano (Ulrico Hoepli)
  4. ^ askoxford.com
  5. ^ Oxford English Dictionary Online, Oxford University Press 2007.
  6. ^ Wricke, Gunter, and Eberhard Weber. 1986. Quantitative genetics and selection in plant breeding. Berlin: W. de Gruyter. Page 257.
  7. ^ J. O. Rawlings, C. Clark Cockerham Analysis of Double Cross Hybrid Populations. J. O. Rawlings, C. Clark Cockerham Biometrics, Vol. 18, No. 2 (Jun., 1962), pp. 229-244 doi:10.2307/2527461
  8. ^ Roy, Darbeshwar. 2000. Plant breeding analysis and exploitation of variation. Pangbourne, UK: Alpha Science International. Page 446.
  9. ^ Arnold, M.L. (1996). Natural Hybridization and Evolution. New York: Oxford University Press. pp. 232. ISBN 978-0-19-509975-1. 
  10. ^ Keeton, William T. 1980. Biological science. New York: Norton. ISBN 0-393-95021-2 Page 800
  11. ^ "McBeath S, Tan PP, Bai Q, Speed RM.". 1988. http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&list_uids=3378453&dopt=Abstract. 
  12. ^ Esch, Mary. "Study: Eastern wolves are hybrids with coyotes". AP News. http://www.google.com/hostednews/ap/article/ALeqM5hIGn3MyVkj3dkd4W2UljCK0y4sVg?docId=787311a8475a4d3aae6e8498aa2edd80. Retrieved 1 June 201`. 
  13. ^ Frost, Grant, Faivovich, Bain, Haas, Haddad, de Sá, Channing, Wilkinson, Donnellan, Raxworthy, Campbell, Blotto, Moler, Drewes, Nussbaum, Lynch, Green, and Wheeler 2006. The amphibian tree of life. Bulletin of the American Museum of Natural History. Number 297. New York. Issued March 15, 2006.
  14. ^ http://www.savingwildplaces.com/swp-home/swp-crocodile/8287793?preview=&psid=&ph=class%2525253dawc-148772
  15. ^ http://www.pacificbulbsociety.org/pbswiki/index.php/LiliumHybrids
  16. ^ University of Colorado Principles of Genetics (MCDB 2150) Lecture 33: Chromosomal changes: Monosomy, Trisomy, Polyploidy, Structural Changes
  17. ^ Evaluating the utility of Arabidopsis thaliana as a model for understanding heterosis in hybrid crops Journal Euphytica Publisher Springer Netherlands ISSN 0014-2336 (Print) 1573-5060 (Online) Issue Volume 156, Numbers 1-2 / July, 2007 DOI 10.1007/s10681-007-9362-1 Pages 157-171
  18. ^ Rieseberg, Loren H.; Margaret A. Archer and Robert K. Wayne (1999-07). "Transgressive segregation, adaptation and speciation". Heredity 83 (4): 363–372. doi:10.1038/sj.hdy.6886170. PMID 10583537. 
  19. ^ Smith C. Wayne. Corn: Origin, History, Technology, and Production. Wiley Series in Crop Science, 2004, p. 332.
  20. ^ a b c d Allendorf, Fred W.; R.F. Leary, P. Spruell, & J.K. Wenburg (November 2001). "The problems with hybrids: setting conservation guidelines". TRENDS in Ecology & Evolution 16 (11): 613–622. doi:10.1016/S0169-5347(01)02290-X. 
  21. ^ Allendorf, Fred (2007). Conservation and the Genetics of Populations. Malden, MA: Blackwell Publishing. pp. 534. 
  22. ^ a b Allendorf, Fred (2007). Conservation and the Genetics of Populations. Malden, MA: Blackwell Publishing. pp. 421–448. 
  23. ^ "Hybrid bear shot dead in Canada". BBC News. 13 May 2006. http://news.bbc.co.uk/2/hi/science/nature/4766217.stm. 
  24. ^ Ehrlich, Paul; John Holdren (26). "Impact of population Growth". Science 171 (3977): 1212–1216. doi:10.1126/science.171.3977.1212. 
  25. ^ Vitousek, Peter; Carla M. D'Antonio, Lloyd L. Loope, Marcel Rejmánek, & Randy Westbrooks (1997). "Introduced Species: A Significant Component of Human-cause Global Change". New Zealand Journal of Ecology 21 (1): 1–16. 
  26. ^ a b McCarthy, Eugene M. 2006. Handbook of Avian Hybrids of the World. Oxford: Oxford University Press. Pp. 16-17.
  27. ^ McCarthy, Eugene M. 2006. Handbook of Avian Hybrids of the World. Oxford: Oxford University Press. P. 17.
  28. ^ Darwin, C. 1868. Variation of Animals and Plants under Domestication, vol. II, p. 125
  29. ^ Spicer, J. W. G. 1854. Note on hybrid gallinaceous birds. The Zoologist, 12: 4294-4296 (see p. 4295).
  30. ^ Mooney, H. A.; Cleland, E. E. (2001). "The evolutionary impact of invasive species". Proc Natl Acad Sci U S A. 98 (10): 5446–5451. doi:10.1073/pnas.091093398. PMC 33232. PMID 11344292. http://www.pubmedcentral.nih.gov/articlerender.fcgi?artid=33232. 
  31. ^ Rhymer JM and Simberloff, D. (1996) Extinction by Hybridization and Introgression. Annual Review of Ecology and Systematics 27: 83-109 (doi:10.1146/annurev.ecolsys.27.1.83), [1]
  32. ^ Brad M. Potts, Robert C. Barbour, Andrew B. Hingston (2001) Genetic Pollution from Farm Forestry using eucalypt species and hybrids; A report for the RIRDC/L&WA/FWPRDC; Joint Venture Agroforestry Program; RIRDC Publication No 01/114; RIRDC Project No CPF - 3A; ISBN 0 642 58336 6; ISSN 1440-6845; Australian Government, Rural Industrial Research and Development Corporation
  33. ^ Devinder Sharma “Genetic Pollution: The Great Genetic Scandal”; Bulletin 28. hosted by www.farmedia.org
  34. ^ Troyer, A. Forrest. Breeding Widely Adapted Cultivars: Examples from Maize. Encyclopedia of Plant and Crop Science, 27 February 2004.
  35. ^ Barriers to hybridization of Solanum bulbocastanumDun. and S. VerrucosumSchlechtd. and structural hybridity in their F1 plants Journal Euphytica [googilygoogilygoo..bwahahaa] Publisher Springer Netherlands ISSN 0014-2336 (Print) 1573-5060 (Online) Issue Volume 25, Number 1 / January, 1976 Category Articles DOI 10.1007/BF00041523 Pages 1-10

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