- Transgenic plant
Transgenic plants possess a
geneor genes that have been transferred from a different species. Although DNA of another species can be integrated in a plant genome by natural processes, the term "transgenic plants" refers to plants created in a laboratory using recombinant DNAtechnology. The aim is to design plants with specific characteristics by artificial insertion of genes from other species or sometimes entirely different kingdoms. "See also" Genetics, List of genetic engineering topics.
Varieties containing genes of two distinct plant species are frequently created by classical breeders who deliberately force hybridization between distinct plant species when carrying out interspecific or intergeneric "wide crosses" with the intention of developing disease resistant crop varieties. Classical plant breeders use a number of "in vitro" techniques such as
protoplastfusion, embryorescue or mutagenesis to generate diversity and produce plants that would not exist in nature ("see also Plant breeding, Heterosis, New Rice for Africa").
Such traditional techniques (used since about 1930 on) have never been controversial, or been given wide publicity except among professional biologists, and have allowed crop breeders to develop varieties of basic food crop, wheat in particular, which resist devastating plant diseases such as rusts. "Hope" is one such wheat variety bred by E. S. McFadden with a gene from a wild grass. "Hope" saved American wheat growers from devastating stem rust outbreaks in the 1930s.
Methods used in traditional breeding that generate plants with DNA from two species by non-recombinant methods are widely familiar to professional plant scientists, and serve important roles in securing a sustainable future for agriculture by protecting crops from pests and helping land and water to be used more efficiently.fact|date=October 2007 ("see also"
Food security, International Fund for Agricultural Development, International development)
Natural movements of genes between species
Natural movement of genes between species, often called
horizontal gene transferor lateral gene transfer, can occur because of gene transfer mediated by natural processes.
This natural gene movement between species has been widely detected during genetic investigation of various natural mobile genetic elements, such as
transposons, and retrotransposons that naturally translocate to new sites in a genome, and often move to new species over an evolutionary time scale. There are many types of natural mobile DNAs, and they have been detected abundantly in food crops such as rice [DNA-binding specificity of rice mariner-like transposases and interactions with Stowaway MITEs, C. Feschotte et al, Nucleic Acids Research 2005 33(7):2153-2165; [http://nar.oxfordjournals.org/cgi/content/full/33/7/2153 ] ] .
These various mobile genes play a major role in dynamic changes to chromosomes during evolution [ [http://www.pnas.org/cgi/content/full/103/21/8101 Birth of a chimeric primate gene by capture of the transposase gene from a mobile element — PNAS:1. Richard Cordaux*,2. Swalpa Udit†,3. Mark A. Batzer*, and 4. Cédric Feschotte†,‡(+Author Affiliations)1.*Department of Biological Sciences, Biological Computation and Visualization Center, Center for BioModular Multi-Scale Systems, Louisiana State University, 202 Life Sciences Building, Baton Rouge, LA 70803; and 2.†Department of Biology, University of Texas, Arlington, TX 76019 1. Edited by Susan R. Wessler, University of Georgia, Athens, GA, and approved March 27, 2006 (received for review February 10, 2006)] ] , [ [http://www.nature.com/nrg/journal/v4/n11/abs/nrg1204_fs.html November 2003 Vol 4 No 11 Nature Reviews Genetics 4, 865-875 (2003); doi:10.1038/nrg1204 THE ORIGIN OF NEW GENES: GLIMPSES FROM THE YOUNG AND OLD (By Manyuan Long, Esther Betrán, Kevin Thornton & Wen Wang ] ] , and have often been given whimsical names, such as Mariner, Hobo, Trans-Siberian Express (Transib), Osmar, Helitron, Sleeping Princess, MITE and MULE, to emphasize their mobile and transient behavior.
Genetically mobile DNA contitututes a major fraction of the DNA of many plants, and the natural dynamic changes to crop plant chromosomes caused by this natural transgenic DNA mimics many of the features of plant genetic engineering currently pursued in the laboratory, such as using
transposons as a genetic tool, and molecular cloning. "See also" transposon, retrotransposon, integron, provirus, endogenous retrovirus, heterosis, [http://www.nature.com/ng/journal/v37/n9/abs/ng1615.html;jsessionid=367F14297326E4C7BF28B89F461CDB46 Gene duplication and exon shuffling by helitron-like transposons generate intraspecies diversity in maize.]
There is new scientific literature about natural transgenic events in plants, through movement of natural mobile DNAs called MULEs between rice and Setaria millet [ [http://biology.plosjournals.org/perlserv/?request=get-document&doi=10.1371/journal.pbio.0040035 PLoS Biology - (2006) Jumping Genes Cross Plant Species Boundaries. PLoS Biol 4(1): e35 doi:10.1371/journal.pbio.0040035 Published: December 20, 2005 Copyright: © 2005 Public Library of Science.] ] .
It is becoming clear that natural rearrangements of DNA and horizontal gene transfer play a pervasive role in natural evolution. Importantly many, if not most, flowering plants evolved by transgenesis - that is, the creation of natural interspecies hybrids in which chromosome sets from different plant species were added together. There is also the long and rich history of interspecies cross-breeding with traditional methods.
Deliberate creation of transgenic plants during breeding
Production of transgenic plants in wide-crosses by plant breeders has been a vital aspect of conventional
plant breedingfor about a century. Without it, security of our food supply against losses caused by crop pests such as rusts and mildews would be severely compromised fact|date=October 2007. The first historically recorded interspecies transgenic cereal hybrid was actually between wheat and rye (Wilson, 1876).
In the 20th century, the introduction of alien probing into common foods was repeatedly achieved by traditional crop breeders by artificially overcoming fertility barriers. Novel genetic rearrangements of plant chromosomes, such as insertion of large blocks of
rye(Secale) genes into wheat chromosomes ('translocations'), has also been exploited widely for many decades [ [http://www.pnas.org/cgi/content/abstract/96/11/5937 Plant genetic resources: What can they contribute toward increased crop productivity? — PNAS: 1. David Hoisington*, 2. Mireille Khairallah, 3. Timothy Reeves, 4. Jean-Marcel Ribaut, 5. Bent Skovmand, 6. Suketoshi Taba, and 7. Marilyn Warburton] ] .
By the late 1930s with the introduction of
colchicine, perennial grasses were being hybridized with wheat with the aim of transferring aids resistance and perenniality into annual crops, and large-scale practical use of hybrids was well established, leading on to development of Triticosecale and other new transgenic cereal crops. In 1985 Plant Genetic Systems( Ghent, Belgium), founded by Marc Van Montaguand Jeff Schell, was the first company to develop genetically engineered ( tobacco) plants with insect tolerance by expressing genes encoding for insecticidal proteins from Bacillus thuringiensis(Bt). [Vaeck, M., A. Reynaerts, H. Hofte, S. Jansens, M. De Beuckeleer, C. Dean, M. Zabeau, M. Van Montagu & J. Leemans. 1987, Transgenic plants protected from insect attack, Nature 328: 33-37.]
Transgenic resistance traits in bread wheat varieties
Important transgenic pathogen and parasite resistance traits in current bread wheat varieties (gene, eg "Lr9" followed by the source species) are:
**Lr9 (from "Aegilops umbellulata")
**Lr18 "Triticum timopheevi"
**Lr23 "T. turgidum"
**Lr24 "Ag. elongatum"
**Lr25 "Secale cereale"
**Lr29 "Ag. elongatum"
**Lr32 "T. tauschii"
**Sr2 "T. turgidum" ("Hope" ) McFadden, E. S. (1930) J. Am. Soc. Agron. 22, 1020-1031 .
**Sr22 "Triticum monococcum"
**Sr36 "Triticum timopheevii"
**Yr15 "Triticum dicoccoides"
**Pm21 "Haynaldia villosa"
**Pm25 "T. monococcum"
*Wheat streak mosaic virus
**Wsm1 "Ag. elongatum"Pest resistance
**H21 "S. cereale" H23,
**H24 "T. tauschii"
**H27 "Aegilops ventricosa"
*Cereal cyst nematode
**Cre3 (Ccn-D1) "T. tauschii"
Genetically engineered plants
The intentional creation of transgenic plants by laboratory based recombinant DNA methods is more recent (from the mid-70s on) and has been a controversial development in the field of
biotechnologyopposed vigorously by many NGOs, and several governments, particularly within the European Community. These transgenic recombinant plants (biotech crops, modern transgenics) are transforming agriculture in those regions that have allowed farmers to adopt them, and the area sown to these crops has continued to grow globally in every years since their first introduction in 1996. Fact|date=October 2007
Transgenic recombinant plants are generated in a laboratory by adding one or more
genes to a plant's genome,and the techniques frequently called transformation. Transformation is usually achieved using gold particle bombardment or through the process of Horizontal gene transferusing a soil bacterium, " Agrobacterium tumefaciens", carrying an engineered plasmid vector, or carrier of selected extra genes.
Transgenic recombinant plants are identified as a class of
genetically modified organism(GMO); usually only transgenic plants created by direct DNA manipulation are given much attention in public discussions.
Transgenic plants have been deliberately developed for a variety of reasons: longer shelf life, disease resistance, herbicide resistance, pest resistance, non-biological stress resistances, such as to drought or nitrogen starvation, and nutritional improvement ("see
Golden rice"). The first modern recombinant crop approved for sale in the US, in 1994, was the FlavrSavrtomato, which was intended to have a longer shelf life. The first conventional transgenic cereal created by scientific breeders was actually a hybrid between wheat and rye in 1876 (Wilson, 1876). The first transgenic cereal may have been wheat, which itself is a natural transgenic plant derived from at least three different parenteral species.
Genetically modified organisms were prior to the coming of the commercially viable crops as the
FlavrSavrtomato, only strictly grown indoors (in laboratories). However, after the introduction of the Flavr Savrtomato, certain GMO-crops as GMO-soy and GMO-corn where in the USA being grown outdoors on large scales.
Commercial factors, especially high regulatory and research costs, have so far restricted modern transgenic crop varieties to major traded commodity crops, but recently R&D projects to enhance crops that are locally important in developing counties are being pursued, such as insect protected cow-pea for Africa. [http://www.pi.csiro.au/enewsletter/PDF/PI_info_Cowpeas.pdf] , and insect protected Brinjal eggplant for India [http://www.fbae.org/Channels/Views/indian_bt_brinjal_in_public.htm] .
Transgenic plants have been used for
bioremediationof contaminated soils. Mercury, seleniumand organic pollutants such as polychlorinated biphenyls (PCBs) have been removed from soils by transgenic plants containing genes for bacterial enzymes [cite journal | author=Meagher, RB | title=Phytoremediation of toxic elemental and organic pollutants | journal=Current Opinion In Plant Biology | volume=3 | issue=2 | year=2000 | pages=153–162 | pmid=10712958 | doi=10.1016/S1369-5266(99)00054-0] .
Regulation of transgenic plants
United Statesthe [http://usbiotechreg.nbii.gov Coordinated Framework for Regulation of Biotechnology] governs the regulation of transgenic organisms, including plants. The three agencies involved are:
Animal and Plant Health Inspection Service- who state that
The Biotechnology Regulatory Services (BRS) program of the U.S. Department of Agriculture’s (USDA) Animal and Plant Health Inspection Service (APHIS) is responsible for regulating the introduction (importation, interstate movement, and field release) of genetically engineered (GE) organisms that may pose a plant pest risk. BRS exercises this authority through APHIS regulations in Title 7, Code of Federal Regulations, Part 340 under the Plant Protection Act of 2000.
APHIS protects agriculture and the environment by ensuring that biotechnology is developed and used in a safe manner. Through a strong regulatory framework, BRS ensures the safe and confined introduction of new GE plants with significant safeguards to prevent the accidental release of any GE material.
APHIS has regulated the biotechnology industry since 1987 and has authorized more than 10,000 field tests of GE organisms. In order to emphasize the importance of the program, APHIS established BRS in August 2002 by combining units within the agency that dealt with the regulation of biotechnology. [http://www.aphis.usda.gov/publications/biotechnology/content/printable_version/BRS_FS_FedReg_02-06.pdf Biotechnology, Federal Regulation, and the U.S. Department of Agriculture, February 2006, USDA-APHIS Fact Sheet]
EPA- evaluates potential environmental impacts, especially for genes which encode for pesticide production
DHHS, Food and Drug Administration(FDA) - evaluates human health risk if the plant is intended for human consumption
The term cisgenic is now being introduced by some plant producers to refer to artificial genetic transfers that could theoretically have been replicated by conventional crossbreeding methods. Producers argue that "cisgenically" produced organisms do not have the same degree of novelty as "transgenic" organisms, and involve no environmental issues that are not already present in conventional crossbreeding. It is argued [Deborah MacKenzie, "Let them eat spuds" (cover story) New Scientist No2667 2 August 2008 30-33] that "cisgenic" modification is useful for plants that are difficult to crossbreed predictably by conventional means (such as potatoes), and that plants in the "cisgenic" category should not require the same level of legal regulation as other genetically-modified organisms.
The potential impact on nearby ecosystems is one of the greatest concerns associated with transgenic plants.
Transgenes have the potential for significant ecological impact if the plants can increase in frequency and persist in natural populations. These concerns are similar to those surrounding conventionally bred plant breeds. Several risk factors should be considered:
* Is the transgenic plant capable of growing outside a cultivated area?
* Can the transgenic plant pass its genes to a local wild species, and are the offspring also fertile?
* Does the introduction of the transgene confer a selective advantage to the plant or to hybrids in the wild?
Many domesticated plants can mate and hybridise with wild relatives when they are grown in proximity, and whatever genes the cultivated plant had can then be passed to the hybrid. This applies equally to transgenic plants and conventionally bred plants, as in either case there are advantageous genes that may have negative consequences to an ecosystem upon release. This is normally not a significant concern, despite fears over 'mutant superweeds' overgrowing local wildlife: although hybrid plants are far from uncommon, in most cases these hybrids are not fertile due to
polyploidy, and will not multiply or persist long after the original domestic plant is removed from the environment. However, this does not negate the possibility of a negative impact.
In some cases, the pollen from a domestic plant may travel many miles on the wind before fertilising another plant. This can make it difficult to assess the potential harm of crossbreeding; many of the relevant hybrids are far away from the test site. Among the solutions under study for this concern are systems designed to prevent transfer of transgenes, such as
Terminator Technology, and the genetic transformation of the chloroplastonly, so that only the seed of the transgenic plant would bear the transgene. With regard to the former, there is some controversy that the technologies may be inequitable and might force dependence upon producers for valid seed in the case of poor farmers, whereas the latter has no such concern but has technical constraints that still need to be overcome. Solutions are being developed by EU funded research programmes such as Co-Extraand Transcontainer.
There are at least three possible avenues of hybridization leading to escape of a transgene:
*Hybridization with non-transgenic crop plants of the same species and variety.
*Hybridization with wild plants of the same species.
*Hybridization with wild plants of closely related species, usually of the same genus.
However, there are a number of factors which must be present for hybrids to be created.
*The transgenic plants must be close enough to the wild species for the pollen to reach the wild plants.
*The wild and transgenic plants must flower at the same time.
*The wild and transgenic plants must be genetically compatible.
In order to persist, these hybrid offspring:
*Must be viable, and fertile.
*Must carry the transgene.
Studies suggest that a possible escape route for transgenic plants will be through hybridization with wild plants of related species.
#It is known that some crop plants have been found to hybridize with wild counterparts.
#It is understood, as a basic part of population genetics, that the spread of a transgene in a wild population will be directly related to the fitness effects of the gene in addition to the rate of influx of the
geneto the population. Advantageous genes will spread rapidly, neutral genes will spread with genetic drift, and disadvantageous genes will only spread if there is a constant influx.
#The ecological effects of transgenes are not known, but it is generally accepted that only genes which improve fitness in relation to
abioticfactors would give hybrid plants sufficient advantages to become weedy or invasive. Abiotic factors are parts of the ecosystem which are not alive, such as climate, salt and mineral content, and temperature. Genes improving fitness in relation to biotic factors could disturb the (sometimes fragile) balance of an ecosystem. For instance, a wild plant receiving a pest resistance gene from a transgenic plant might become resistant to one of its natural pests, say, a beetle. This could allow the plant to increase in frequency, while at the same time animals higher up in the food chain, which are at least partly dependent on that beetle as food source, might decrease in abundance. However, the exact consequences of a transgene with a selective advantage in the natural environment are almost impossible to predict reliably.
It is also important to refer to the demanding actions that government of developing countries had been building up among the last decades.
Agricultural impact of transgenic plants
Outcrossing of transgenic plants not only poses potential environmental risks, it may also trouble farmers and food producers. Many countries have different legislations for transgenic and conventional plants as well as the derived food and feed, and consumers demand the freedom of choice to buy GM-derived or conventional products. Therefore, farmers and producers must separate both production chains. This requires coexistence measures on the field level as well as traceability measures throughout the whole food and feed processing chain. Research projects such as
Co-Extra, SIGMEA and Transcontainer investigate how farmers can avoid outcrossing and mixing of transgenic and non-transgenic crops, and how processors can ensure and verify the separation of both production chains.
*Anderson, K. and Lee Ann Jackson. 2005. Some Implications of GM Food Technology Policies for Sub-Saharan Africa. Journal of African Economies 14(3):385-410; doi:10.1093/jae/eji013
*Heong, KL, YH Chen, DE Johnson, GC Jahn, M Hossain, RS Hamilton. 2005. Debate Over a GM Rice Trial in China. Letters. Science, Vol 310, Issue 5746, 231-233 , 14 October 2005.
*Huang, J., Ruifa Hu, Scott Rozelle, Carl Pray. 2005. Insect-Resistant GM Rice in Farmers' Fields: Assessing Productivity and Health Effects in China. Science (29 April 2005) Vol. 308. no. 5722, pp. 688 – 690. DOI: 10.1126/science.1108972
*Syvanen, M. and Kado, C. I. Horizontal Gene Transfer. Second Edition. Academic Press 2002.
*Chrispeels, M.J. and Sadova, D.E. Plants, Genes, and Crop Biotechnology. Second Edition. James and Bartlett 2003.
* [http://www.pnas.org/cgi/content/abstract/96/11/5937 Plant genetic resources: What can they contribute toward increased crop productivity? David Hoisington*, Mireille Khairallah, Timothy Reeves, Jean-Marcel Ribaut, Bent Skovmand, Suketoshi Taba, and Marilyn Warburton, Proc. Natl. Acad Sci USA. Vol. 96, Issue 11, 5937-5943, May 25, 1999. (This paper was presented at the National Academy of Sciences colloquium "Plants and Population: Is There Time?" held December 5-6, 1998, at the Arnold and Mabel Beckman Center in Irvine, CA).] (See also
* [http://www.aphis.usda.gov/publications/biotechnology/index.shtml U.S. Department of Agriculture Animal and Plant Health Inspection Service (USDA-APHIS) Publications Biotechnology.]
* [http://www.aphis.usda.gov/publications/biotechnology/content/printable_version/BRS_FS_FedReg_02-06.pdf Biotechnology, Federal Regulation, and the U.S. Department of Agriculture, February 2006, USDA-APHIS Fact Sheet]
* [http://www.aphis.usda.gov/publications/biotechnology/content/printable_version/BRS_CoordFrameBro.pdf Biotechnology Regulatory Services, Coordinated Framework for the Regulation of Biotechnology, USDA-APHIS Outreach Material]
* [http://www.aphis.usda.gov/publications/biotechnology/content/printable_version/BRS_QA_biotechandusda.pdf Questions and Answers About Biotechnology and the USDA, August 2006, USDA-APHIS Fact Sheet]
* [http://www.aphis.usda.gov/publications/biotechnology/content/printable_version/BRS_FS_pharmaceutical_02-06.pdf Permitting Genetically Engineered Plants That Produce Pharmaceutical Compounds, February 2006, USDA-APHIS Fact Sheet]
Food and Agriculture Organization
Foreign Agricultural Service
GM food controversy
International Fund for Agricultural Development
Mobile genetic elements
Genetically modified organism
Transposons as a genetic tool
United States Department of Agriculture
* [http://www.merid.org/fs-agbiotech/ Food Security and Ag-Biotech News] — balanced news on the debate over transgenic crops
* [http://www.pnas.org/cgi/content/abstract/96/11/5937 Plant genetic resources and transgenics contributions towards increased crop productivity] — Context of transgenics for food security
* [http://www.isb.vt.edu/ Information Systems for Biotechnology (ISB) based at Virginia Tech] — Authoritative information in the form of readable up-to-date articles to support the environmentally responsible use of agricultural biotechnology products, including transgenic plants.
* [http://www.fbae.org/index.htm Foundation for Biotechnology Awareness and Education]
* [http://www.pgeconomics.co.uk/who.htm PG Economics] - Reports on agricultural economic benefits.
* [http://www.agbioforum.org/ AgBioForum Journal] - Professional economics papers on crop biotechnology
* [http://www.ifpri.org/ Internation Food Policy Research Institute] The food security context.
* [http://www.truthabouttrade.org/ Truth About Trade] - Forthright support of agricultural technology for farmers and the value of free trade.
* [http://www.agbioworld.org/ AgbioWorld] - Comprehensive scientifically sound resource that supports the use on transgenic crops and provides news from around the world on agriculture.
* [http://www.gmo-compass.org/eng/home/ GMO-Compass] - Information on the use of genetic engineering in the agri-food industry.
* [http://www.gmo-safety.eu/en/ GMO Safety] - Information about research projects funded by the German Ministry of Education and Research (BMBF) on the biological safety of genetically modified plants.
* [http://www.co-extra.eu Co-Extra: Research on co-existence and traceability of transgenic and conventional plants]
* [http://sigmea.dyndns.org SIGMEA: Research on sustainable introduction of transgenic crops into European agriculture]
* [http://www.transcontainer.org/UK Transcontainer: Research on biological transgene containment]
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