Mutation breeding

Last updated

Mutation breeding, sometimes referred to as "variation breeding", is the process of exposing seeds to chemicals, radiation, or enzymes [1] [2] in order to generate mutants with desirable traits to be bred with other cultivars. Plants created using mutagenesis are sometimes called mutagenic plants or mutagenic seeds.

Contents

From 1930 to 2014 more than 3200 mutagenic plant varieties were released [3] [4] that have been derived either as direct mutants (70%) or from their progeny (30%). [5] Crop plants account for 75% of released mutagenic species with the remaining 25% ornamentals or decorative plants. [6] However, although the FAO/IAEA reported in 2014 that over 1,000 mutant varieties of major staple crops were being grown worldwide, [3] it is unclear how many of these varieties are currently used in agriculture or horticulture around the world, as these seeds are not always identified or labeled as having a mutagenic provenance. [7]

History

According to garden historian Paige Johnson:

After WWII, there was a concerted effort to find 'peaceful' uses for atomic energy. One of the ideas was to bombard plants with radiation and produce lots of mutations, some of which, it was hoped, would lead to plants that bore more heavily or were disease or cold-resistant or just had unusual colors. The experiments were mostly conducted in giant gamma gardens on the grounds of national laboratories in the US but also in Europe and countries of the [then-]USSR. [8]

Processes

There are different kinds of mutagenic breeding such as using chemical mutagens like ethyl methanesulfonate and dimethyl sulfate, radiation or transposons to generate mutants. Mutation breeding is commonly used to produce traits in crops such as larger seeds, new colors, or sweeter fruits, that either cannot be found in nature or have been lost during evolution. [9]

Radiation

Exposing plants to radiation is sometimes called radiation breeding and is a sub class of mutagenic breeding. Radiation breeding was discovered in the 1920s when Lewis Stadler of the University of Missouri used X-rays on maize and barley. In the case of barley, the resulting plants were white, yellow, pale yellow and some had white stripes. [10] In 1928, Stadler first published his findings on radiation-induced mutagenesis in plants. [11] During the period 1930–2024, radiation-induced mutant varieties were developed primarily using gamma rays (64%) and X-rays (22%), [12] [13] [14] [6] : 187  although other sources of radiation such as microwaves or high-energy photons and high-energy electrons can be applied. [15] [16]

Radiation breeding may take place in atomic gardens; [11] and seeds have been sent into orbit in order to expose them to more cosmic radiation. [17]

Ultraviolet has been used, for example to produce knockouts for the investigation of virulence mechanisms of plant pathogens. [18]

Chemicals

High rates of chromosome aberrations resulting from ionizing radiation and the accompanied detrimental effects made researchers look for alternate sources for inducing mutations. As a result, an array of chemical mutagens has been discovered. The most widely used chemical mutagens are alkylating agents. Ethyl methanesulfonate (EMS) is the most popular because of its effectiveness and ease of handling, especially its detoxification through hydrolysis for disposal. Nitroso compounds are the other alkylating agents widely used, but they are light-sensitive and more precautions need to be taken because of their higher volatility. EMS has become a commonly used mutagen for developing large numbers of mutants for screening such as in developing TILLING populations. [19] Although many chemicals are mutagens, only few have been used in practical breeding as the doses need to be optimised and also because the effectiveness is not high in plants for many.[ citation needed ]

Restriction endonucleases

Interest in the use of bacterial restriction endonucleases (RE) – for example Fok1 [2] and CRISPR/Cas9 [1] [2] – to study double-stranded breaks in plant DNA began in the mid-nineties. These breaks in DNA, otherwise known as DSBs, were found to be the source of much chromosomal damage in eukaryotes, causing mutations in plant varieties. REs induce a result on plant DNA similar to that of ionizing radiation or radiomimetic chemicals. Blunt ended breaks in the DNA, unlike sticky ended breaks, were found to produce more variations in chromosomal damage, making them the more useful type of break for mutation breeding. While the connection of REs to chromosomal aberrations is mostly limited to research on mammalian DNA, success in mammalian studies caused scientists to conduct more studies of RE-induced chromosomal and DNA damage on barley genomes. Due to restriction endonucleases' ability to facilitate damage in chromosomes and DNA, REs have the capability of being used as a new method of mutagenesis to promote the proliferation of mutated plant varieties. [20] [1] [2]

Space-breeding

The ability of plants to develop and thrive is dependent on conditions such as microgravity and cosmic radiation in space. China has been experimenting with this theory by sending seeds into space, testing to see if space flights will cause genetic mutations. Since 1987, China has cultivated 66 mutant varieties from space through their space-breeding program. Chromosomal aberrations greatly increased when seeds were sent into aerospace compared to their earth-bound counterparts. The effect of space flight on seeds depends on their species and variety. For example, space-bred wheat saw a large growth in seed germination in compared to its Earth-bound control, but space-bred rice had no visible advantage compared to its control. For the varieties that were positively mutated by space flight, their growth potential exceeded that of not only their Earth-grown counterparts, but also their irradiated counterparts on Earth. Compared to traditional mutagenic techniques, space-bred mutations have greater efficacy in that they experience positive effects on their first generation of mutation, whereas irradiated crops often see no advantageous mutations in their first generations. Though multiple experiments have shown the positive effects of space flight on seed mutation, there is no clear connection as to what aspect of aerospace has produced such advantageous mutations. There is much speculation around cosmic radiation being the source of chromosomal aberrations, but so far, there has been no concrete evidence of such connection. Though China's space-breeding program has been shown to be very successful, the program requires a large budget and technological support that many other countries are either unwilling or unable to provide, meaning this program is unfeasible outside of China. Due to such restraints, scientists have been trying to replicate space condition on Earth in order to promote the same expedient space-born mutations on Earth. One such replication is a magnetic field-free space (MF), which produces an area with a weaker magnetic field than that of Earth. MF treatment produced mutagenic results, and has been used to cultivate new mutant varieties of rice and alfalfa. Other replications of space conditions include irradiation of seeds by a heavy 7 Li-ion beam or mixed high-energy particles. [21] These space-bred varieties are already being introduced to the public. In 2011, during the National Lotus Flowers Exhibition in China, a mutant lotus, called the "Outer Space Sun", was shown at the flower show. [22]

Ion beam technology

Ion beams mutate DNA by deleting multiple bases from the genome. Compared to traditional sources of radiation, like gamma rays and X-rays, ion beams have been shown to cause more severe breaks in DNA that are more difficult to weave back together, causing the change in DNA to be more drastic than changes caused by traditional irradiation. Ion beams change DNA in a manner that makes it look vastly different than its original makeup, more so than when traditional irradiation techniques are used. Most experimentation, using ion beam technology, has been conducted in Japan. Notable facilities using this technology are TIARA of the Japan Atomic Energy Agency, RIKEN Accelerator Research Facility, and various other Japanese institutions. During the process of ion beam radiation, seeds are wedged between two kapton films and irradiated for roughly two minutes. Mutation frequencies are notably higher for ion beam radiation compared to electron radiation, and the mutation spectrum is broader for ion beam radiation compared to gamma ray radiation. The broader mutation spectrum was revealed through the largely varied amount of flower phenotypes produced by ion beams. Flowers mutated by the ion beams exhibited a variety of colors, patterns, and shapes. Through ion beam radiation, new varieties of plants have been cultivated. These plants had the characteristics of being ultraviolet light-B resistant, disease resistant, and chlorophyll-deficient. Ion beam technology has been used in the discovery of new genes responsible for the creation of more robust plants, but its most prevalent use is commercially for producing new flower phenotypes, like striped chrysanthemums. [23]

Mature pollen treated with gamma radiation

Gamma radiation is used on mature rice pollen to produce parent plants used for crossing. The mutated traits in the parent plants are able to be inherited by their offspring plants. Because rice pollen has a very short lifespan, researchers had to blast gamma rays at cultured spikes from rice plants. Through experimentation, it was revealed that there was a greater variety of mutation in irradiated pollen rather than irradiated dry seeds. Pollen treated with 46Gy of gamma radiation showed an increase in grain size overall and other useful variations. Typically, the length of each grain was longer after the crossing of irradiated parent rice plants. The rice progeny also exhibited a less chalky visage, improving on the appearance of the parent rice plants. This technique was used to develop two new rice cultivars, Jiaohezaozhan and Jiafuzhan, in China. Along with facilitating the creation of these two rice cultivars, the irradiation of mature rice pollen has produced roughly two hundred mutant rice lines. Each of these lines produce rice grains of both a higher quality and larger size. The mutations produced by this technique vary with each generation, meaning further breeding of these mutated plants could produce new mutations. Traditionally, gamma radiation is used on solely adult plants, and not on pollen. The irradiation of mature pollen allows mutant plants to grow without being in direct contact with gamma radiation. This discovery is in contrast to what was previously believed about gamma radiation: that it could only elicit mutations in plants and not pollen. [24]

Comparison to other techniques

In the debate over genetically modified foods, the use of transgenic processes is often compared and contrasted with mutagenic processes. [25] While the abundance and variation of transgenic organisms in human food systems, and their effect on agricultural biodiversity, ecosystem health and human health is somewhat well documented, mutagenic plants and their role on human food systems is less well known, with one journalist writing "Though poorly known, radiation breeding has produced thousands of useful mutants and a sizable fraction of the world's crops...including varieties of rice, wheat, barley, pears, peas, cotton, peppermint, sunflowers, peanuts, grapefruit, sesame, bananas, cassava and sorghum." [10] In Canada crops generated by mutation breeding face the same regulations and testing as crops obtained by genetic engineering. [26] [27] [28] [29] Mutagenic varieties tend to be made freely available for plant breeding, in contrast to many commercial plant varieties or germplasm that increasingly have restrictions on their use [6] : 187  such as terms of use, patents and proposed genetic user restriction technologies and other intellectual property regimes and modes of enforcement.

Unlike genetically modified crops, which typically involve the insertion of one or two target genes, plants developed via mutagenic processes with random, multiple and unspecific genetic changes [30] have been discussed as a concern [31] but are not prohibited by any nation's organic standards. Reports from the US National Academy of Sciences state that there is no scientific justification for regulating genetic engineered crops while not doing so for mutation breeding crops. [7]

Several organic food and seed companies promote and sell certified organic products that were developed using both chemical and nuclear mutagenesis. [32] Several certified organic brands, whose companies support strict labeling or outright bans on GMO-crops, market their use of branded wheat and other varietal strains which were derived from mutagenic processes without any reference to this genetic manipulation. [32] These organic products range from mutagenic barley and wheat ingredient used in organic beers [33] to mutagenic varieties of grapefruits sold directly to consumers as organic. [34]

Release by nation

As of 2011 the percentage of all mutagenic varieties released globally, by country, were: [6] : 187  [35]

Notable varieties per country include:

Flag of Argentina.svg  Argentina
Flag of Australia (converted).svg  Australia
Flag of Bangladesh.svg  Bangladesh
Flag of Cuba.svg  Cuba
Flag of the People's Republic of China.svg  People's Republic of China

Flag of the Czech Republic.svg  Czech Republic

Flag of Egypt.svg  Egypt
Flag of Finland.svg  Finland
Flag of France.svg  France
Flag of Germany.svg  Germany
Flag of Ghana.svg  Ghana
Flag of India.svg  India
Flag of Italy.svg  Italy
Flag of Japan.svg  Japan
Flag of Myanmar.svg  Myanmar
Flag of Pakistan.svg  Pakistan
Flag of Peru.svg  Peru
Flag of Sudan.svg  Sudan
Flag of Thailand.svg  Thailand
Flag of the United Kingdom.svg  United Kingdom
Flag of the United States (23px).png  United States
Flag of Vietnam.svg  Vietnam

In 2014, it was reported that 17 rice mutant varieties, 10 soybean, two maize and one chrysanthemum mutant varieties had been officially released to Vietnamese farmers. 15% of rice and 50% of soybean was produced from mutant varieties. [50]

See also

Related Research Articles

<span class="mw-page-title-main">Cereal</span> Grass that has edible grain

A cereal is a grass cultivated for its edible grain. Cereals are the world's largest crops, and are therefore staple foods. They include rice, wheat, rye, oats, barley, millet, and maize. Edible grains from other plant families, such as buckwheat and quinoa, are pseudocereals. Most cereals are annuals, producing one crop from each planting, though rice is sometimes grown as a perennial. Winter varieties are hardy enough to be planted in the autumn, becoming dormant in the winter, and harvested in spring or early summer; spring varieties are planted in spring and harvested in late summer. The term cereal is derived from the name of the Roman goddess of grain crops and fertility, Ceres.

<span class="mw-page-title-main">Food irradiation</span> Sterilization of food with ionizing radiations for enhanced preservation and longer shelflife

Food irradiation is the process of exposing food and food packaging to ionizing radiation, such as from gamma rays, x-rays, or electron beams. Food irradiation improves food safety and extends product shelf life (preservation) by effectively destroying organisms responsible for spoilage and foodborne illness, inhibits sprouting or ripening, and is a means of controlling insects and invasive pests.

<span class="mw-page-title-main">Wheat</span> Genus of grass cultivated for grain

Wheat is a grass widely cultivated for its seed, a cereal grain that is a staple food around the world. The many species of wheat together make up the genus Triticum ; the most widely grown is common wheat. The archaeological record suggests that wheat was first cultivated in the regions of the Fertile Crescent around 9600 BC. Botanically, the wheat kernel is a caryopsis, a type of fruit.

Agricultural biotechnology, also known as agritech, is an area of agricultural science involving the use of scientific tools and techniques, including genetic engineering, molecular markers, molecular diagnostics, vaccines, and tissue culture, to modify living organisms: plants, animals, and microorganisms. Crop biotechnology is one aspect of agricultural biotechnology which has been greatly developed upon in recent times. Desired trait are exported from a particular species of Crop to an entirely different species. These transgene crops possess desirable characteristics in terms of flavor, color of flowers, growth rate, size of harvested products and resistance to diseases and pests.

<span class="mw-page-title-main">ENU</span> Chemical compound

ENU, also known as N-ethyl-N-nitrosourea (chemical formula C3H7N3O2), is a highly potent mutagen. For a given gene in mice, ENU can induce 1 new mutation in every 700 loci. It is also toxic at high doses.

<span class="mw-page-title-main">Sterile insect technique</span> Method of biological control for insect populations

The sterile insect technique (SIT) is a method of biological insect control, whereby overwhelming numbers of sterile insects are released into the wild. The released insects are preferably male, as this is more cost-effective and the females may in some situations cause damage by laying eggs in the crop, or, in the case of mosquitoes, taking blood from humans. The sterile males compete with fertile males to mate with the females. Females that mate with a sterile male produce no offspring, thus reducing the next generation's population. Sterile insects are not self-replicating and, therefore, cannot become established in the environment. Repeated release of sterile males over low population densities can further reduce and in cases of isolation eliminate pest populations, although cost-effective control with dense target populations is subjected to population suppression prior to the release of the sterile males.

Irradiation is the process by which an object is exposed to radiation. An irradiator is a device used to expose an object to radiation, notably gamma radiation, for a variety of purposes. Irradiators may be used for sterilizing medical and pharmaceutical supplies, preserving foodstuffs, alteration of gemstone colors, studying radiation effects, eradicating insects through sterile male release programs, or calibrating thermoluminescent dosimeters (TLDs).

Plant breeders use different methods depending on the mode of reproduction of crops, which include:

<span class="mw-page-title-main">Waxy corn</span> Type of field corn

Waxy corn or glutinous corn is a type of field corn characterized by its sticky texture when cooked as a result of larger amounts of amylopectin. The corn was first described from a specimen from China in 1909. As this plant showed many peculiar traits, the American breeders long used it as a genetic marker to tag the existence of hidden genes in other maize breeding programs. In 1922 a researcher found that the endosperm of waxy maize contained only amylopectin and no amylose starch molecule in opposition to normal dent corn varieties that contain both. Until World War II, the main source of starch in the United States was tapioca, but when Japan severed the supply lines of the U.S., they forced processors to turn to waxy maize. Amylopectin or waxy starch is now used mainly in food products, but also in the textile, adhesive, corrugating and paper industry.

<span class="mw-page-title-main">Hybrid rice</span> Rice crossbred from different varieties

Hybrid rice is a type of Asian rice that has been crossbred from two very different parent varieties. As with other types of hybrids, hybrid rice typically displays heterosis or "hybrid vigor", so when grown under the same conditions as comparable purebred rice varieties, it can produce up to 30% more yield. To produce hybrid seeds in large quantity, a purebred sterile rice variety is fertilized with fertile pollen from a different variety. High-yield crops, including hybrid rice, are one of the most important tools for combatting worldwide food crises.

Intensive crop farming is a modern industrialized form of crop farming. Intensive crop farming's methods include innovation in agricultural machinery, farming methods, genetic engineering technology, techniques for achieving economies of scale in production, the creation of new markets for consumption, patent protection of genetic information, and global trade. These methods are widespread in developed nations.

<span class="mw-page-title-main">Atomic gardening</span> Form of mutation breeding in plants

Atomic gardening is a form of mutation breeding where plants are exposed to radiation. Some of the mutations produced thereby have turned out to be useful. Typically this is gamma radiation – in which case it is a gamma garden – produced by cobalt-60.

The Institute of Plant Breeding (IPB) is a research institute of the University of the Philippines Los Baños. It is the national biotechnology research center and repository for all crops other than rice, which is handled by the Philippine Rice Research Institute.

<span class="mw-page-title-main">Biofortification</span> Breeding crops for higher nutritional value

Biofortification is the idea of breeding crops to increase their nutritional value. This can be done either through conventional selective breeding, or through genetic engineering. Biofortification differs from ordinary fortification because it focuses on making plant foods more nutritious as the plants are growing, rather than having nutrients added to the foods when they are being processed. This is an important improvement on ordinary fortification when it comes to providing nutrients for the rural poor, who rarely have access to commercially fortified foods. As such, biofortification is seen as an upcoming strategy for dealing with deficiencies of micronutrients in low and middle-income countries. In the case of iron, the WHO estimated that biofortification could help cure the 2 billion people suffering from iron deficiency-induced anemia.

<span class="mw-page-title-main">Plant breeding</span> Humans changing traits, ornamental/crops

Plant breeding is the science of changing the traits of plants in order to produce desired characteristics. It has been used to improve the quality of nutrition in products for humans and animals. The goals of plant breeding are to produce crop varieties that boast unique and superior traits for a variety of applications. The most frequently addressed agricultural traits are those related to biotic and abiotic stress tolerance, grain or biomass yield, end-use quality characteristics such as taste or the concentrations of specific biological molecules and ease of processing.

<span class="mw-page-title-main">Kalanamak rice</span> Indian rice variety

Kalanamak is a scented rice of Nepal and India. Its name means black husk. This variety has been in cultivation since the original Buddhist period. It is popular in Himalayan Tarai of Nepal i.e., Kapilvastu, and eastern Uttar Pradesh, where it is known as the scented black pearl. It was featured in the book Speciality rices of the world by Food and Agriculture Organization of the United Nations.

<span class="mw-page-title-main">Philippine Nuclear Research Institute</span> Agency of the Philippine government

The Philippine Nuclear Research Institute (PNRI) is a government agency under the Department of Science and Technology mandated to undertake research and development activities in the peaceful uses of nuclear energy, institute regulations on the said uses, and carry out the enforcement of said regulations to protect the health and safety of radiation workers and the general public.

Golden Promise is a variety of spring-sown two-row barley. It was developed in the 1950s by exposing an existing variety to gamma radiation with the aim of producing a semi-dwarf variety of barley that had good malting characteristics. It became very popular with farmers during the 1970s but began to be replaced by higher yielding varieties in the 1980s. Due to its perceived better flavour than these more modern varieties, a niche market was maintained and it is still favoured by craft brewers. It has been used by researchers to study the genetics of barley and to investigate the effect of barley on the flavour of beer.

Starmaya is an F1 hybrid coffee tree that can be propagated by seed rather than through somatic embryogenesis (SE). It was propagated from a parent plant that is male-sterile. This facilitates controlled pollination because breeders do not have to manually castrate each individual flower of the autogamous coffee tree.

<span class="mw-page-title-main">Shoba Sivasankar</span> A leading geneticist for the United Nations

Shoba Sivasankar is a Geneticist who leads the Plant Breeding and Genetics group of both the International Atomic Energy Agency (IAEA) and the Food and Agriculture Organization of the United Nations.

References

  1. 1 2 3 Chen, Kunling; Wang, Yanpeng; Zhang, Rui; Zhang, Huawei; Gao, Caixia (2019-04-29). "CRISPR/Cas Genome Editing and Precision Plant Breeding in Agriculture". Annual Review of Plant Biology . 70 (1). Annual Reviews: 667–697. doi: 10.1146/annurev-arplant-050718-100049 . ISSN   1543-5008. PMID   30835493. S2CID   73471425.
  2. 1 2 3 4 Mackelprang, Rebecca; Lemaux, Peggy G. (2020-04-29). "Genetic Engineering and Editing of Plants: An Analysis of New and Persisting Questions". Annual Review of Plant Biology . 71 (1). Annual Reviews: 659–687. doi: 10.1146/annurev-arplant-081519-035916 . ISSN   1543-5008. PMID   32023090. S2CID   211049671.
  3. 1 2 (2014) Plant Breeding and Genetics Archived 2018-07-27 at the Wayback Machine Joint FAO/IAEA Division of Nuclear Techniques in Food and Agriculture, Retrieved 31 July 2014
  4. ↑ Schouten, H. J.; Jacobsen, E. (2007). "Are Mutations in Genetically Modified Plants Dangerous?". Journal of Biomedicine and Biotechnology. 2007 (7): 1–2. doi: 10.1155/2007/82612 . PMC   2218926 . PMID   18273413.
  5. ↑ M.K. Maluszynsk; K. Nichterlein; L. van Zanten; B.S. Ahloowalia (2000). "Officially released mutant varieties – the FAO/IAEA Database". Mutation Breeding Review (12): 1–84.
  6. 1 2 3 4 5 6 7 8 9 10 11 Ahloowali, B.S. (2004). "Global impact of mutation-derived varieties". Euphytica. 135 (2): 187–204. doi:10.1023/b:euph.0000014914.85465.4f. S2CID   34494057 . Retrieved 20 April 2011.
  7. 1 2 Kaskey, Jack (21 November 2013) The Scariest Veggies of Them All Bloomberg Business Week, Retrieved 31 July 2014
  8. ↑ Johnson, Paige. "Atomic Gardens" . Retrieved 20 April 2011.
  9. ↑ "New Citrus Variety Released by UC Riverside is Very Sweet, Juicy and Low-seeded".
  10. 1 2 Broad, William J. (28 August 2007). "Useful Mutants, Bred With Radiation". New York Times. Retrieved 20 April 2011.
  11. 1 2 Atomic Gardens: Public Perceptions & Public Policy Archived 2013-06-30 at the Wayback Machine , Life Sciences Foundation Magazine, Spring 2012.
  12. ↑ "EJPAU 2010. Zalewska M. , Miler N. , Tymoszuk A. , Drzewiecka B. , Winiecki J. RESULTS OF MUTATION BREEDING ACTIVITY ON Chrysanthemum × grandiflorum (Ramat.) Kitam. IN POLAND". www.ejpau.media.pl. Retrieved 2024-09-12.
  13. ↑ Miler, Natalia; Kulus, Dariusz; Sliwinska, Elwira (2020-11-01). "Nuclear DNA content as an indicator of inflorescence colour stability of in vitro propagated solid and chimera mutants of chrysanthemum". Plant Cell, Tissue and Organ Culture. 143 (2): 421–430. doi: 10.1007/s11240-020-01929-9 . ISSN   1573-5044.
  14. ↑ Kulus, Dariusz; Tymoszuk, Alicja; Jedrzejczyk, Iwona; Winiecki, Janusz (2022-06-01). "Gold nanoparticles and electromagnetic irradiation in tissue culture systems of bleeding heart: biochemical, physiological, and (cyto)genetic effects". Plant Cell, Tissue and Organ Culture. 149 (3): 715–734. doi: 10.1007/s11240-022-02236-1 . ISSN   1573-5044.
  15. ↑ Miler, Natalia; Kulus, Dariusz (2018-01-03). "Microwave treatment can induce chrysanthemum phenotypic and genetic changes". Scientia Horticulturae. 227: 223–233. Bibcode:2018ScHor.227..223M. doi:10.1016/j.scienta.2017.09.047. ISSN   0304-4238.
  16. ↑ Miler, Natalia; Jedrzejczyk, Iwona; Jakubowski, Seweryn; Winiecki, Janusz (2021-06-01). "Ovaries of Chrysanthemum Irradiated with High-Energy Photons and High-Energy Electrons Can Regenerate Plants with Novel Traits". Agronomy. 11 (6): 1111. doi: 10.3390/agronomy11061111 . ISSN   2073-4395.
  17. ↑ Smith, Peter (2011-04-12). "How Radiation is Changing the Foods that You Eat". GOOD. GOOD Worldwide, Inc. Retrieved 2011-07-16.
  18. ↑ Xu, Liangsheng; Li, Guoqing; Jiang, Daohong; Chen, Weidong (2018-08-25). "Sclerotinia sclerotiorum: An Evaluation of Virulence Theories". Annual Review of Phytopathology . 56 (1). Annual Reviews: 311–338. doi: 10.1146/annurev-phyto-080417-050052 . ISSN   0066-4286. PMID   29958073. S2CID   49615444.
  19. ↑ Pathirana, R. Plant Mutation Breeding in Agriculture. CAB Reviews: Perspectives in Agriculture, Veterinary Science, Nutrition and Natural Resources. 2011 6 No 032
  20. ↑ Stoilov, L.; Gecheff, K. (2009). Shu, Q.Y. (ed.). "Restriction Endonucleases as a Tool for In Vivo Induction of Chromosomal and DNA Damage in Barley Genome" (PDF). Induced Plant Mutations in the Genomics Era– via Food and Agriculture Organization of the United Nations.
  21. ↑ Liu, L.X.; Guo, H.J.; Zhao, L.S.; Wang, J.; Zhao, S.R. (2009). Shu, Q.Y. (ed.). "Achievements and Perspectives of Crop Space Breeding in China" (PDF). Induced Plant Mutations in the Genomics Era– via Food and Agriculture Organization of the United Nations.
  22. ↑ "Mutant Lotus Bred in Space on Display in Chongqing, China".
  23. ↑ Tanaka, A. (2009). Shu, Q.Y. (ed.). "Establishment of Ion Beam Technology for Breeding" (PDF). Induced Plant Mutations in the Genomics Era– via Food and Agriculture Organization of the United Nations.
  24. ↑ Wang, H.; Qiu, S.; Zheng, J.; Jiang, L.; Huang, H.; Huang, Y. (2009). Shu, Q.Y. (ed.). "Generation of New Rice Cultivars from Mature Pollen Treated with Gamma-Radiation" (PDF). Induced Plant Mutations in the Genomics Era– via Food and Agriculture Organization of the United Nations.
  25. ↑ UK Government Science Review First Report, Prepared by the GM Science Review panel (July 2003). Chairman Professor Sir David King, Chief Scientific Advisor to the UK Government, P 9: "...it is necessary to produce about 100 GM plants to obtain one that has the desirable characters for its use as a basis of a new GM crop variety. ... Most of these so-called conventional plant breeding methods (such as gene transfer by pollination, mutation breeding, cell selection and induced polyploidy) have a substantially greater discard rate. Mutation breeding, for instance, involves the production of unpredictable and undirected genetic changes and many thousands, even millions, of undesirable plants are discarded in order to identify plants with suitable qualities for further breeding."
  26. ↑ The Canadian regulatory system is based on whether a product has novel features regardless of method of origin. In other words, a product is regulated as genetically modified if it carries some trait not previously found in the species whether it was generated using mutation breeding or genetic engineering (or any other method including selective breeding).
  27. ↑ Evans, Brent and Lupescu, Mihai (15 July 2012) Canada – Agricultural Biotechnology Annual – 2012 Archived 2013-12-15 at the Wayback Machine GAIN (Global Agricultural Information Network) report CA12029, United States Department of Agriculture, Foreifn Agricultural Service, Retrieved 7 August 2014
  28. ↑ McHugen, Alan (September 14, 2000). "Chapter 1: Hors-d'oeuvres and entrees/What is genetic modification? What are GMOs?". Pandora's Picnic Basket . Oxford University Press. ISBN   978-0198506744.
  29. ↑ Rowland, G.G. (2009). "Chapter 110: The Effect of Plants With Novel Traits (PNT) Regulation on Mutation Breeding in Canada". In Shu, Q. Y. (ed.). Induced Plant Mutations in the Genomics Era. Plant Breeding Section, Joint FAO/IAEA Division of Nuclear Techniques in Food and Agriculture, International Atomic Energy Agency, Vienna, Austria. pp. 423–424. ISBN   978-92-5-106324-8.
  30. ↑ Useful Mutants, Bred With Radiation, by William J. Broad, New York Times, August 28, 2007.
  31. ↑ Discussion Document Excluded Methods Terminology Archived 2013-06-24 at the Wayback Machine , National Organic Standards Board GMO ad hoc Subcommittee paper, U.S. Agricultural Marketing Service, published February 6, 2013.
  32. 1 2 Mendel in the Kitchen: A Scientist's View of Genetically Modified Foods, By Nina V. Fedoroff and Nancy Marie Brow, pg. 17, Joseph Henry Press, 2004.
  33. ↑ Golden Promise Organic Ale
  34. ↑ Wasatch Organic Rio Red Grapefruit
  35. ↑ Pathirana, Ranjith (September 6, 2011) Plant mutation breeding in agriculture CAB Reviews: Perspectives in Agriculture, Veterinary Science, Nutrition and Natural Resources (CAB International); 20116 (032): 1 – 20; doi:10.1079/PAVSNNR20116032; ISSN 1749-8848; Retrieved August 6, 2014
  36. 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 Kharkwal, M. C.; Shu, Q. Y. (2008). "The role of induced mutations in world food security" (PDF). Induced Plant Mutations in the Genomics Era. Proceedings of an International Joint FAO/IAEA Symposium, 2008 2009. Vienna, Austria: Joint FAO/IAEA Division of Nuclear Techniques in Food and Agriculture, International Atomic Energy Agency: 33–38. ISBN   9789251063248.
  37. 1 2 3 4 5 6 Shu, Qing-Yao (2012). Forster, B. P.; Nakagawa, H. (eds.). Plant Mutation Breeding and Biotechnology. CABI Publishing. p. 17. ISBN   978-1780640853.
  38. ↑ "Lift-off for Chinese space potato". BBC News. 12 February 2007.
  39. 1 2 3 4 Ahloowalia, B. S.; Maluszynski, M. (2001). "Production Process in Old and Modern Spring Barley Varieties". Euphytica. 118 (2): 167. doi:10.1023/A:1004162323428. S2CID   36088943.
  40. ↑ Lipavsky, J. Petr, J. and Hradecká, D, (2002) "Production Process in Old and Modern Spring Barley Varieties" Die Bodenkultur, 53 (1) 2, Page 19
  41. ↑ Rossi, Luigi (2010). "Il miglioramento genetico del grano duro in Casaccia. Il caso CRESO" [Genetic Improvement of Durum Wheat in Casaccia. The Creso Case](PDF). Energia, Ambiente e Innovazione. ENEA. Archived from the original (PDF) on 2016-11-30. Retrieved 2016-11-29.
  42. ↑ van Harten, A. M. (1998). Mutation Breeding: Theory and Practical Applications. U. K.: Cambridge University Press. p. 239. ISBN   978-0521470742.
  43. ↑ Kotobuki, Kazuo. "Japanese pear tree named 'Osa Gold'" . Retrieved 20 April 2011.
  44. 1 2 3 (2008) NIAB – Plant Breeding & Genetics Division, Achievements Nuclear Institute for Agriculture and Biology, Faisalabad, Pakistan, Retrieved 16 May 2013
  45. 1 2 3 (2012) Improved barley varieties – Feeding people from the equator to the arctic Archived 2014-08-29 at the Wayback Machine Joint FAO/IAEAProgramme, Nuclear Techniques in Food and Agriculture, Retrieved 25 October 2013
  46. ↑ Forster, B. P. (2001). "Mutation genetics of salt tolerance in barley: An assessment of Golden Promise and other semi-dwarf mutants". Euphytica. 120 (3): 317–328. doi:10.1023/A:1017592618298. S2CID   22320510.
  47. ↑ Broad, William (2007-08-28). "Useful Mutants, Bred With Radiation". New York Times. Retrieved 2013-06-19.
  48. ↑ (2012) Successful Mutation Breeding Programmes in Vietnam Archived 2013-10-29 at the Wayback Machine Joint FAO/IAEAProgramme, Nuclear Techniques in Food and Agriculture, Retrieved 25 October 2013
  49. 1 2 Vinh, M.Q. et al (2009) Current Status and Research Directions of Induced Mutation Application to Seeds Program in Vietnam in Induced Plant Mutations in the Genomics Era, FAO of the UN, Rome, Pp 341–345, Web page version retrieved 25 October 2013
  50. ↑ (2014) Successful Mutation Breeding Programmes in Vietnam Archived 2013-10-29 at the Wayback Machine Joint FAO/IAEA Division of Nuclear Techniques in Food and Agriculture, Retrieved 31 July 2014
This page is based on this Wikipedia article
Text is available under the CC BY-SA 4.0 license; additional terms may apply.
Images, videos and audio are available under their respective licenses.