Genetic pollution is a controversial term for uncontrolled gene flow into wild populations. It is defined as “the dispersal of contaminated altered genes from genetically engineered organisms to natural organisms, esp. by cross-pollination”, but has come to be used in some broader ways. It is related to the population genetics concept of gene flow, and genetic rescue, which is genetic material intentionally introduced to increase the fitness of a population. It is called genetic pollution when it negatively impacts on the fitness of a population, such as through outbreeding depression and the introduction of unwanted phenotypes which can lead to extinction.
Conservation biologists and conservationists have used the term to describe gene flow from domestic, feral, and non-native species into wild indigenous species, which they consider undesirable. They promote awareness of the effects of introduced invasive species that may "hybridize with native species, causing genetic pollution". In the fields of agriculture, agroforestry and animal husbandry, genetic pollution is used to describe gene flows between genetically engineered species and wild relatives. The use of the word “pollution” is meant to convey the idea that mixing genetic information is bad for the environment, but because the mixing of genetic information can lead to a variety of outcomes, “pollution” may not always be the most accurate descriptor.
- 1 Gene flow to wild population
- 2 Controversy
- 3 See also
- 4 References
Gene flow to wild population
Some conservation biologists and conservationists have used genetic pollution for a number of years as a term to describe gene flow from a non-native, invasive subspecies, domestic, or genetically-engineered population to a wild indigenous population.
The introduction of genetic material into the gene pool of a population by human intervention can have both positive and negative effects on populations. When genetic material is intentionally introduced to increase the fitness of a population, this is called genetic rescue. When genetic material is unintentionally introduced to a population, this is called genetic pollution and can negatively affect the fitness of a population (primarily through outbreeding depression), introduce other unwanted phenotypes, or theoretically lead to extinction.
An introduced species is one that is not native to a given population that is either intentionally or accidentally brought into a given ecosystem. Effects of introduction are highly variable, but if an introduced species has a major negative impact on its new environment, it can be considered an invasive species. One such example is the introduction of the Asian Longhorned beetle in North America, which was first detected in 1996 in Brooklyn, New York. It is believed that these beetles were introduced through cargo at trade ports. The beetles are highly damaging to the environment, and are estimated to cause risk to 35% of urban trees, excluding natural forests. These beetles cause severe damage to the wood of trees by larval funneling. Their presence in the ecosystem destabilizes community structure, having a negative influence on many species in the system. Introduced species are not always disruptive to an environment, however. Tomás Carlo and Jason Gleditch of Penn State University found that the number of invasive honeysuckle plants in the area correlated with the number and diversity of the birds in the Happy Valley Region of Pennsylvania, suggesting introduced honeysuckle plants and birds formed a mutually beneficial relationship. Presence of introduced honeysuckle was associated with higher diversity of the bird populations in that area, demonstrating that introduced species are not always detrimental to a given environment and it is completely context dependent.
Conservation biologists and conservationists have, for a number of years, used the term to describe gene flow from domestic, feral, and non-native species into wild indigenous species, which they consider undesirable. For example, TRAFFIC is the international wildlife trade monitoring network that works to limit trade in wild plants and animals so that it is not a threat to conservationist goals. They promote awareness of the effects of introduced invasive species that may "hybridize with native species, causing genetic pollution". Furthermore, the Joint Nature Conservation Committee, the statutory adviser to the UK government, has stated that invasive species "will alter the genetic pool (a process called genetic pollution), which is an irreversible change."
Invasive species can invade both large and small native populations and have a profound effect. Upon invasion, invasive species interbreed with native species to form sterile or more evolutionarily fit hybrids that can outcompete the native populations. Invasive species can cause extinctions of small populations on islands that are particularly vulnerable due to their smaller amounts of genetic diversity. In these populations, local adaptations can be disrupted by the introduction of new genes that may not be as suitable for the small island environments. For example, the Cercocarpus traskiae of the Catalina Island off the coast of California has faced near extinction with only a single population remaining due to the hybridization of its offspring with Cercocarpus betuloides.
Increased contact between wild and domesticated populations of organisms can lead to reproductive interactions that are detrimental to the wild population's ability to survive. A wild population is one that lives in natural areas and is not regularly looked after by humans. This contrast with domesticated populations that live in human controlled areas and are regularly, and historically, in contact with humans. Genes from domesticated populations are added to wild populations as a result of reproduction. In many crop populations this can be the result of pollen traveling from farmed crops to neighboring wild plants of the same species. For farmed animals, this reproduction may happen as the result of escaped or released animals.
Aquaculture is the practice of farming aquatic animals or plants for the purpose of consumption. This practice is becoming increasingly common for the production of salmon. This is specifically termed aquaculture of salmonoids. One of the dangers of this practice is the possibility of domesticated salmon breaking free from their containment. The occurrence of escaping incidents is becoming increasingly common as aquaculture gains popularity. Farming structures may be ineffective at holding the vast number of fast growing animals they house. Natural disasters, high tides, and other environmental occurrences can also trigger aquatic animal escapes. The reason these escapes are considered dangers is the impact they pose for the wild population they reproduce with after escaping. In many instances the wild population experiences a decreased likelihood of survival after reproducing with domesticated populations of salmon.
The Washington Department of Fish and Wildlife cites that "commonly expressed concerns surrounding escaped Atlantic salmon include competition with native salmon, predation, disease transfer, hybridization, and colonization" A report done by that organization in 1999 did not find that escaped salmon posed a significant risk to wild populations.
Crops refer to groups of plants grown for consumption. Despite domestication over many years, these plants are not so far removed from their wild relatives that they could reproduce if brought together. Many crops are still grown in the areas they originated and gene flow between crops and wild relatives impacts the evolution of wild populations. Farmers can avoid reproduction between the different populations by timing their planting of crops so that crops are not flowering when wild relatives would be. Domesticated crops have been changed through artificial selection and genetic engineering. The genetic make up of many crops is different than that of its wild relatives, but the closer they grow to one another the more likely they are to share genes through pollen. Gene flow persists between crops and wild counterparts.
Genetically engineered organisms
Genetically engineered organisms are genetically modified in a laboratory, and therefore distinct from those that were bred through artificial selection. In the fields of agriculture, agroforestry and animal husbandry, genetic pollution is being used to describe gene flows between GE species and wild relatives. An early use of the term "genetic pollution" in this later sense appears in a wide-ranging review of the potential ecological effects of genetic engineering in The Ecologist magazine in July 1989. It was also popularized by environmentalist Jeremy Rifkin in his 1998 book The Biotech Century. While intentional crossbreeding between two genetically distinct varieties is described as hybridization with the subsequent introgression of genes, Rifkin, who had played a leading role in the ethical debate for over a decade before, used genetic pollution to describe what he considered to be problems that might occur due the unintentional process of (modernly) genetically modified organisms (GMOs) dispersing their genes into the natural environment by breeding with wild plants or animals.
Concerns about negative consequences from gene flow between genetically engineered organisms and wild populations are valid. Most corn and soybean crops grown in the midwestern USA are genetically modified. There are corn and soybean varieties that are resistant to herbicides like glyphosate and corn that produces neonicotinoid pesticide within all of its tissues. These genetic modifications are meant to increase yields of crops but there is little evidence that yields actually increase. While scientists are concerned genetically engineered organisms can have negative effects on surrounding plant and animal communities, the risk of gene flow between genetically engineered organisms and wild populations is yet another concern. Many farmed crops may be weed resistant and reproduce with wild relatives. More research is necessary to understand how much gene flow between genetically engineered crops and wild populations occurs, and the impacts of genetic mixing.
Mutations within organisms can be executed through the process of exposing the organism to chemicals or radiation in order to generate mutations. This has been done in plants in order to create mutants that have a desired trait. These mutants can then be bred with other mutants or individuals that are not mutated in order to maintain the mutant trait. However, similar to the risks associated with introducing individuals to a certain environment, the variation created by mutated individuals could have a negative impact on native populations as well.
Since 2005 there has existed a GM Contamination Register, launched for GeneWatch UK and Greenpeace International that records all incidents of intentional or accidental release of organisms genetically modified using modern techniques.
Genetic use restriction technologies (GURTs) were developed for the purpose of property protection, but could be beneficial in preventing the dispersal of transgenes. GeneSafe technologies introduced a method that became known as “Terminator.” This method is based on seeds that produce sterile plants. This would prevent movement of transgenes into wild populations as hybridization would not be possible. However, this technology has never been deployed as it disproportionately negatively affects farmers in developing countries, who save seeds to use each year (whereas in developed countries, farmers generally buy seeds from seed production companies).
Physical containment has also been utilized to prevent the escape of transgenes. Physical containment includes barriers such as filters in labs, screens in greenhouses, and isolation distances in the field. Isolation distances have not always been successful, such as transgene escape from an isolated field into the wild in herbicide-resistant bentgrass Agrostis stolonifera.
Another suggested method that applies specifically to protection traits (e.g. pathogen resistance) is mitigation. Mitigation involves linking the positive trait (beneficial to fitness) to a trait that is negative (harmful to fitness) to wild but not domesticated individuals. In this case, if the protection trait was introduced to a weed, the negative trait would also be introduced in order to decrease overall fitness of the weed and decrease possibility of the individual’s reproduction and thus propagation of the transgene.
Not all genetically engineered organisms cause genetic pollution. Genetic engineering has a variety of uses and is specifically defined as a direct manipulation of the genome of an organism. Genetic pollution can occur in response to the introduction of a species that is not native to a particular environment, and genetically engineered organisms are examples of individuals that could cause genetic pollution following introduction. Due to these risks, studies have been done in order to assess the risks of genetic pollution associated with organisms that have been genetically engineered:
- Genetic In a 10-year study of four different crops, none of the genetically engineered plants were found to be more invasive or more persistent than their conventional counterparts. An often cited claimed example of genetic pollution is the reputed discovery of transgenes from GE maize in landraces of maize in Oaxaca, Mexico. The report from Quist and Chapela, has since been discredited on methodological grounds. The scientific journal that originally published the study concluded that "the evidence available is not sufficient to justify the publication of the original paper."  More recent attempts to replicate the original studies have concluded that genetically modified corn is absent from southern Mexico in 2003 and 2004.
- A 2009 study verified the original findings of the controversial 2001 study, by finding transgenes in about 1% of 2000 samples of wild maize in Oaxaca, Mexico, despite Nature retracting the 2001 study and a second study failing to back up the findings of the initial study. The study found that the transgenes are common in some fields, but non-existent in others, hence explaining why a previous study failed to find them. Furthermore, not every laboratory method managed to find the transgenes.
- A 2004 study performed near an Oregon field trial for a genetically modified variety of creeping bentgrass (Agrostis stolonifera) revealed that the transgene and its associate trait (resistance to the glyphosate herbicide) could be transmitted by wind pollination to resident plants of different Agrostis species, up to 14 km from the test field. In 2007, the Scotts Company, producer of the genetically modified bentgrass, agreed to pay a civil penalty of $500,000 to the United States Department of Agriculture (USDA). The USDA alleged that Scotts "failed to conduct a 2003 Oregon field trial in a manner which ensured that neither glyphosate-tolerant creeping bentgrass nor its offspring would persist in the environment".
Not only are there risks in terms of genetic engineering, but there are risks that emerge from species hybridization In Czechoslovakia, ibex were introduced from Turkey and Sinai to help promote the ibex population there, which caused hybrids that produced offspring too early, which caused the overall population to disappear completely. The genes of each population of the ibex in Turkey and Sinai were locally adapted to their environments so when placed in a new environmental context did not flourish. Additionally, the environmental toll that may arise from the introduction of a new species may be so disruptive that the ecosystem is no longer able to sustain certain populations.
The use of the word “pollution” in the term genetic pollution has a deliberate negative connotation and is meant to convey the idea that mixing genetic information is bad for the environment. However, because the mixing of genetic information can lead to a variety of outcomes, “pollution” may not be the most accurate descriptor. Gene flow is undesirable according to some environmentalists and conservationists, including groups such as Greenpeace, TRAFFIC, and GeneWatch UK 
"Invasive species have been a major cause of extinction throughout the world in the past few hundred years. Some of them prey on native wildlife, compete with it for resources, or spread disease, while others may hybridize with native species, causing "genetic pollution". In these ways, invasive species are as big a threat to the balance of nature as the direct overexploitation by humans of some species."</ref>.
It can also be considered undesirable if it leads to a loss of fitness in the wild populations. The term can be associated with the gene flow from a mutation bred, synthetic organism or genetically engineered organism to a non GE organism, by those who consider such gene flow detrimental. These environmentalist groups stand in complete opposition to the development and production of genetically engineered organisms.
Seeds of Destruction: Hidden Agenda of Genetic Manipulation is a book by American writer, Frederick William Engdahl. In his book, Engdahl explains how the (oil-rich) Rockefeller family is planing to control world farming by the Green Revolution plan. Rockefeller family is trying to finish other family farms in the United States and also is trying do same as this plan to other countries and finally take the whole world food security on their hands. Engdahl believes that green revolution is happening by the new world empire. He also sees the Rockefeller family hand behind the great Genetically modified organism (GMO) project to control world population.
From a governmental perspective, genetic pollution is defined as follows by the Food and Agriculture Organization of the United Nations:
"Uncontrolled spread of genetic information (frequently referring to transgenes) into the genomes of organisms in which such genes are not present in nature."
Use of the term 'genetic pollution' and similar phrases such as genetic deterioration, genetic swamping, genetic takeover, and genetic aggression, are being debated by scientists as many do not find it scientifically appropriate. Rhymer and Simberloff argue that these types of terms:
"...imply either that hybrids are less fit than the parentals, which need not be the case, or that there is an inherent value in "pure" gene pools."
They recommend that gene flow from invasive species be termed genetic mixing since:
"Mixing" need not be value-laden, and we use it here to denote mixing of gene pools whether or not associated with a decline in fitness.
Patrick Moore has questioned whether the term "genetic pollution" is more political than scientific. The term is considered to arouse emotional feelings towards the subject matter. In an interview he comments:
"If you take a term used quite frequently these days, the term "genetic pollution," otherwise referred to as genetic contamination, it is a propaganda term, not a technical or scientific term. Pollution and contamination are both value judgments. By using the word "genetic" it gives the public the impression that they are talking about something scientific or technical--as if there were such a thing as genes that amount to pollution.
Thus, using the term “genetic pollution” is inherently political. A scientific approach to discussing gene flow between introduced and native species would be to use terms like genetic mixing or gene flow. Such mixing can definitely have negative consequences on the fitness of native populations, so it is important not to minimize the risk. However, because genetic mixing can also lead to fitness recovery in cases that could be described as “genetic rescue”, it is important to distinguish that just mixing genes from introduced into native populations can lead to variable outcomes for the fitness of native populations.
- Rhymer JM, 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.
- Competitive Enterprise Institute staff (2004). "What's Wrong with the Environmental Movement: an interview with Patrick Moore". Environment News. The Heartland Institute. Archived from the original on 24 November 2006.
- Boffey PM (December 13, 1983). "Italy's Wild Dogs Winning Darwinian Battle". The New York Times.
Although wolves and dogs have always lived in close contact in Italy and have presumably mated in the past, the newly worrisome element, in Dr. Boitani's opinion, is the increasing disparity in numbers, which suggests that interbreeding will become fairly common. As a result, genetic pollution of the wolf gene pool might reach irreversible levels, he warned. By hybridization, dogs can easily absorb the wolf genes and destroy the wolf, as it is, he said. The wolf might survive as a more doglike animal, better adapted to living close to people, he said, but it would not be what we today call a wolf.
- Ellstrand NC (2001). "When Transgenes Wander, Should We Worry?" (PDF). Plant Physiol. 125 (4): 1543–1545. doi:10.1104/pp.125.4.1543.
- "the definition of genetic pollution". Dictionary.com. Retrieved 2018-04-30.
- Waller, Donald M. (June 2015). "Genetic rescue: a safe or risky bet?". Molecular Ecology. 24 (11): 2595–2597. doi:10.1111/mec.13220. ISSN 1365-294X. PMID 26013990.
- Butler D (18 August 1994). "Bid to protect wolves from genetic pollution". Nature. 370 (6490): 497. doi:10.1038/370497a0.
- Potts BM, Barbour RC, Hingston AB, Vaillancourt RE (2003). "Corrigendum to: TURNER REVIEW No. 6 Genetic pollution of native eucalypt gene pools—identifying the risks". Australian Journal of Botany. 51 (3): 333. doi:10.1071/BT02035_CO.
- Haack, Robert A., et al. Managing Invasive Populations of Asian Longhorned Beetle and Citrus Longhorned Beetle: A Worldwide Perspective. vol. 55, Annual Review of Entomology, 2010, Managing Invasive Populations of Asian Longhorned Beetle and Citrus Longhorned Beetle: A Worldwide Perspective.
- 2011, Invasive Plants Can Create Positive Ecological Change.
- "When is wildlife trade a problem?". TRAFFIC.org, the wildlife trade monitoring network, a joint programme of WWF and IUCN. The World Conservation Union. Archived from the original on 24 December 2007.
- Effects of the introduction of invasive/non-native species - Joint Nature Conservation Committee (JNCC), a statutory adviser to Government on UK and international nature conservation. Accessed on November 25, 2007. : "Occasionally non-native species can reproduce with native species and produce hybrids, which will alter the genetic pool (a process called genetic pollution), which is an irreversible change."
- Levin DA, Francisco-Ortega J, Jansen RK (1996-02-01). "Hybridization and the Extinction of Rare Plant Species". Conservation Biology. 10 (1): 10–16. doi:10.1046/j.1523-1739.1996.10010010.x. ISSN 1523-1739.
- Anderson R. "More than 160,000 non-native Atlantic salmon escaped into Washington waters in fish farm accident". latimes.com. Retrieved 2018-04-30.
- "'Environmental Nightmare' After Thousands Of Atlantic Salmon Escape Fish Farm". NPR.org. Retrieved 2018-04-30.
- Scotti A. "Thousands of salmon escape from fish farm, and no one knows what will happen next". nydailynews.com. Retrieved 2018-04-30.
- "Escapes: Net-pens are poor containment structures and escaped farmed salmon can compete with wild salmon for food and spawning habitat". Living Oceans. 2013-03-12. Retrieved 2018-04-30.
- Montanari S. "How Did The Eclipse Let Thousands Of Farmed Salmon Escape?". Forbes. Retrieved 2018-04-30.
- "Spill of farmed Atlantic salmon near San Juan Islands much bigger than first estimates". The Seattle Times. 2017-08-24. Retrieved 2018-04-30.
- Braun, Ashley. "Farmed and Dangerous? Pacific Salmon Confront Rogue Atlantic Cousins". Scientific American. Retrieved 2018-05-01.
- video, tronc. "Farmed salmon escape into Washington state waters". chicagotribune.com. Retrieved 2018-05-01.
- "Atlantic Salmon (Salmo salar) - Aquatic Invasive Species | Washington Department of Fish & Wildlife". wdfw.wa.gov. Retrieved 2018-05-01.
- Appleby, Kevin H. Amos and Andrew. "Atlantic Salmon in Washington State: A Fish Management Perspective - WDFW Publications | Washington Department of Fish & Wildlife". wdfw.wa.gov. Retrieved 2018-05-01.
- Ellstrand, Norman C.; Prentice, Honor C.; Hancock, James F. (1999). "Gene Flow and Introgression from Domesticated Plants into Their Wild Relatives". Annual Review of Ecology and Systematics. 30 (1): 539–563. doi:10.1146/annurev.ecolsys.30.1.539.
- Carroll, Sean B. (2010-05-24). "Tracking the Ancestry of Corn Back 9,000 Years". The New York Times. ISSN 0362-4331. Retrieved 2018-05-01.
- "Gene flow from GM to non-GM populations in the crop, forestry, animal and fishery sectors". Background document to Conference 7: May 31 - July 6, 2002; Electronic Forum on Biotechnology in Food and Agriculture. Food and Agriculture Organization of the United Nations (FAO).
- Rifkin J (1998). The Biotech Century: Harnessing the Gene and Remaking the World. J P Tarcher. ISBN 978-0-87477-909-7.
- Quinion M. "Genetic Pollution". World Wide Words.
- Otchet A (1998). "Jeremy Rifkin: fears of a brave new world". an interview hosted by The United Nations Educational, Scientific and Cultural Organization (UNESCO).
- Waltz, Emily (June 2010). "Glyphosate resistance threatens Roundup hegemony". Nature Biotechnology. 28 (6): 537–538. doi:10.1038/nbt0610-537. ISSN 1087-0156. PMID 20531318.
- Krupke, C. H.; Holland, J. D.; Long, E. Y.; Eitzer, B. D. (2017-05-22). "Planting of neonicotinoid-treated maize poses risks for honey bees and other non-target organisms over a wide area without consistent crop yield benefit". Journal of Applied Ecology. 54 (5): 1449–1458. doi:10.1111/1365-2664.12924. ISSN 0021-8901.
- Brown, Paul (2005-07-25). "GM crops created superweed, say scientists". the Guardian. Retrieved 2018-05-01.
- "Illegal Genetically Engineered Corn from Monsanto Detected in Argentina". GM Contamination Register. Archived from the original on 2011-06-22. Retrieved 2010-07-08.
- "Brazil – Illegal Roundup Ready cotton grown on 16,000 hectares". GM Contamination Register.
- "GM Contamination Register".
- Sang, Yi; Millwood, Reginald J.; Neal Stewart Jr, C. (2013-06-04). "Gene use restriction technologies for transgenic plant bioconfinement". Plant Biotechnology Journal. 11 (6): 649–658. doi:10.1111/pbi.12084. ISSN 1467-7644. PMID 23730743.
- Gressel, Jonathan (2014-08-15). "Dealing with transgene flow of crop protection traits from crops to their relatives". Pest Management Science. 71 (5): 658–667. doi:10.1002/ps.3850. ISSN 1526-498X. PMID 24977384.
- Crawley MJ, Brown SL, Hails RS, Kohn D, Rees M (8 February 2001). "Biotechnology: Transgenic crops in natural habitats". Nature. 409 (6821): 682–683. doi:10.1038/35055621. PMID 11217848.
- Quist D, Chapela IH (November 2001). "Transgenic DNA introgressed into traditional maize landraces in Oaxaca, Mexico". Nature. 414 (6863): 541–3. doi:10.1038/35107068. PMID 11734853.
- Christou P (2002). "No Credible Scientific Evidence is Presented to Support Claims that Transgenic DNA was Introgressed into Traditional Maize Landraces in Oaxaca, Mexico". Transgenic Research. 11 (1): 3–5. doi:10.1023/A:1013903300469.
- Metz M, Fütterer J (April 2002). "Biodiversity (Communications arising): suspect evidence of transgenic contamination". Nature. 416 (6881): 600–1, discussion 600, 602. doi:10.1038/nature738. PMID 11935144. Archived from the original (– Scholar search) on October 31, 2008.
- Ortiz-García S, Ezcurra E, Schoel B, Acevedo F, Soberón J, Snow AA (August 2005). "Absence of detectable transgenes in local landraces of maize in Oaxaca, Mexico (2003-2004)". Proceedings of the National Academy of Sciences of the United States of America. 102 (35): 12338–43. doi:10.1073/pnas.0503356102. PMC 1184035. PMID 16093316.
- "'Alien' genes escape into wild corn". New Scientist. 18 February 2009.
- Watrud LS, Lee EH, Fairbrother A, Burdick C, Reichman JR, Bollman M, Storm M, King G, Van de Water PK (October 2004). "Evidence for landscape-level, pollen-mediated gene flow from genetically modified creeping bentgrass with CP4 EPSPS as a marker". Proceedings of the National Academy of Sciences of the United States of America. 101 (40): 14533–8. doi:10.1073/pnas.0405154101. PMC 521937. PMID 15448206.
- "USDA Concludes Genetically Engineered Creeping Bentgrass Investigation".
- Genetic Rescue: a Safe or Risky Bet?
- GE agriculture and genetic pollution web article hosted by Greenpeace.org
- "Say no to genetic pollution". Greenpeace.
- Greenpeace. "Genetic Pollution a Multiplying Nightmare" (PDF).
- Milot E, Perrier C, Papillon L, Dodson JJ, Bernatchez L (April 2013). "Reduced fitness of Atlantic salmon released in the wild after one generation of captive breeding". Evolutionary Applications. 6 (3): 472–85. doi:10.1111/eva.12028. PMC 3673475. PMID 23745139.
- Todhunter, Colin. "On Independence Day, India's new rulers are the World Bank, IMF, WTO and Monsanto". The Ecologist. theecologist.org. Retrieved 29 April 2019.
- Todhunter, Colin. "Poisoned Agriculture: Depopulation and Human Extinction". Counter Punch. counterpunch.org. Retrieved 29 April 2019.
- Zaid A, Hughes HG, Porceddu E, Nicholas F (26 October 2007). Glossary of Biotechnology for Food and Agriculture - A Revised and Augmented Edition of the Glossary of Biotechnology and Genetic Engineering. A FAO Research and Technology Paper. Food and Agriculture Organization of the United Nations. ISBN 978-92-5-104683-8. Archived from the original on 30 November 2011.