Genetically modified organism

Part of a series on
Genetic engineering
Genetically modified organisms
Bacteria Viruses Animals Mammals Fish Insects Plants Maize/corn Rice Soybean Potato
History and regulation
History Regulation Substantial equivalence Cartagena Protocol on Biosafety
Process
Techniques Molecular cloning Recombinant DNA Gene delivery Transformation Transfection Transduction Genome editing TALEN CRISPR
Applications
Genetically modified crops food Gene therapy Designer baby
Controversies
Genetically modified food controversies GMO conspiracy theories Pusztai affair Séralini affair StarLink corn recall He Jiankui affair
v t e

A genetically modified organism (GMO) is any

Genetically engineered organism (GEO) can be considered a more precise term compared to GMO when describing organisms' genomes that have been directly manipulated with biotechnology.[16]^[8] The Cartagena Protocol on Biosafety used the synonym living modified organism (LMO) in 2000 and defined it as "any living organism that possesses a novel combination of genetic material obtained through the use of modern biotechnology."^[17] Modern biotechnology is further defined as "In vitro nucleic acid techniques, including recombinant deoxyribonucleic acid (DNA) and direct injection of nucleic acid into cells or organelles, or fusion of cells beyond the taxonomic family."^[18]

Originally, the term GMO was not commonly used by scientists to describe genetically engineered organisms until after usage of GMO became common in popular media.

The definitions focus on the process more than the product, which means there could be GMOS and non-GMOs with very similar genotypes and phenotypes.[21]^[22] This has led scientists to label it as a scientifically meaningless category,^[23] saying that it is impossible to group all the different types of GMOs under one common definition.^[24] It has also caused issues for organic institutions and groups looking to ban GMOs.^[25][26] It also poses problems as new processes are developed. The current definitions came in before genome editing became popular and there is some confusion as to whether they are GMOs. The EU has adjudged that they are^[27] changing their GMO definition to include "organisms obtained by mutagenesis", but has excluded them from regulation based on their "long safety record" and that they have been "conventionally been used in a number of applications".^[9] In contrast the USDA has ruled that gene edited organisms are not considered GMOs.^[28]

Even greater inconsistency and confusion is associated with various "Non-GMO" or "GMO-free" labeling schemes in food marketing, where even products such as water or salt, which do not contain any organic substances and genetic material (and thus cannot be genetically modified by definition), are being labeled to create an impression of being "more healthy".^[29][30][31]

Production

Creating a genetically modified organism (GMO) is a multi-step process. Genetic engineers must isolate the gene they wish to insert into the host organism. This gene can be taken from a

Humans have

In 1974,

Left: Bacteria transformed with pGLO under ambient light
Right: Bacteria transformed with pGLO visualized under ultraviolet light

Bacteria were the first organisms to be genetically modified in the laboratory, due to the relative ease of modifying their chromosomes.^[88] This ease made them important tools for the creation of other GMOs. Genes and other genetic information from a wide range of organisms can be added to a plasmid and inserted into bacteria for storage and modification. Bacteria are cheap, easy to grow, clonal, multiply quickly and can be stored at −80 °C almost indefinitely. Once a gene is isolated it can be stored inside the bacteria, providing an unlimited supply for research.^[89] A large number of custom plasmids make manipulating DNA extracted from bacteria relatively easy.^[90]

Their ease of use has made them great tools for scientists looking to study gene function and

Other uses for genetically modified bacteria include bioremediation, where the bacteria are used to convert pollutants into a less toxic form. Genetic engineering can increase the levels of the enzymes used to degrade a toxin or to make the bacteria more stable under environmental conditions.^[109] Bioart has also been created using genetically modified bacteria. In the 1980s artist Jon Davis and geneticist Dana Boyd converted the Germanic symbol for femininity (ᛉ) into binary code and then into a DNA sequence, which was then expressed in Escherichia coli.^[110] This was taken a step further in 2012, when a whole book was encoded onto DNA.^[111] Paintings have also been produced using bacteria transformed with fluorescent proteins.^[110]

Viruses

Viruses are often modified so they can be used as vectors for inserting genetic information into other organisms. This process is called transduction and if successful the recipient of the introduced DNA becomes a GMO. Different viruses have different efficiencies and capabilities. Researchers can use this to control for various factors; including the target location, insert size, and duration of gene expression. Any dangerous sequences inherent in the virus must be removed, while those that allow the gene to be delivered effectively are retained.^[112]

While viral vectors can be used to insert DNA into almost any organism it is especially relevant for its potential in treating human disease. Although primarily still at trial stages,

Another potential use of genetically modified viruses is to alter them so they can directly treat diseases. This can be through expression of protective proteins or by directly targeting infected cells. In 2004, researchers reported that a genetically modified virus that exploits the selfish behavior of cancer cells might offer an alternative way of killing tumours.[122]^[123] Since then, several researchers have developed genetically modified oncolytic viruses that show promise as treatments for various types of cancer.^{[124][125][126][127][128]} In 2017, researchers genetically modified a virus to express spinach defensin proteins. The virus was injected into orange trees to combat citrus greening disease that had reduced orange production by 70% since 2005.^[129]

Natural viral diseases, such as myxomatosis and rabbit hemorrhagic disease, have been used to help control pest populations. Over time the surviving pests become resistant, leading researchers to look at alternative methods. Genetically modified viruses that make the target animals infertile through immunocontraception have been created in the laboratory^[130] as well as others that target the developmental stage of the animal.^[131] There are concerns with using this approach regarding virus containment^[130] and cross species infection.^[132] Sometimes the same virus can be modified for contrasting purposes. Genetic modification of the myxoma virus has been proposed to conserve European wild rabbits in the Iberian peninsula and to help regulate them in Australia. To protect the Iberian species from viral diseases, the myxoma virus was genetically modified to immunize the rabbits, while in Australia the same myxoma virus was genetically modified to lower fertility in the Australian rabbit population.^[133]

Outside of biology scientists have used a genetically modified virus to construct a

Fungi can be used for many of the same processes as bacteria. For industrial applications, yeasts combine the bacterial advantages of being a single-celled organism that is easy to manipulate and grow with the advanced protein modifications found in eukaryotes. They can be used to produce large complex molecules for use in food, pharmaceuticals, hormones, and steroids.^[136] Yeast is important for wine production and as of 2016 two genetically modified yeasts involved in the fermentation of wine have been commercialized in the United States and Canada. One has increased malolactic fermentation efficiency, while the other prevents the production of dangerous ethyl carbamate compounds during fermentation.^[96] There have also been advances in the production of biofuel from genetically modified fungi.^[137]

Fungi, being the most common

Agaricus bisporus the common white button mushroom, has been gene edited to resist browning, giving it a longer shelf life. The process used CRISPR to knock out a gene that encodes polyphenol oxidase. As it didn't introduce any foreign DNA into the organism it was not deemed to be regulated under existing GMO frameworks and as such is the first CRISPR-edited organism to be approved for release.^[142] This has intensified debates as to whether gene-edited organisms should be considered genetically modified organisms^[143] and how they should be regulated.^[144]

Plants

Plants have been engineered for scientific research, to display new flower colors, deliver vaccines, and to create enhanced crops. Many plants are

Much of the advances in the field of genetic engineering has come from experimentation with tobacco. Major advances in tissue culture and plant cellular mechanisms for a wide range of plants has originated from systems developed in tobacco.^[146] It was the first plant to be altered using genetic engineering and is considered a model organism for not only genetic engineering, but a range of other fields.^[147] As such the transgenic tools and procedures are well established making tobacco one of the easiest plants to transform.^[148] Another major model organism relevant to genetic engineering is Arabidopsis thaliana. Its small genome and short life cycle makes it easy to manipulate and it contains many homologs to important crop species.^[149] It was the first plant sequenced, has a host of online resources available and can be transformed by simply dipping a flower in a transformed Agrobacterium solution.^[150]

In research, plants are engineered to help discover the functions of certain genes. The simplest way to do this is to remove the gene and see what

Some genetically modified plants are purely

It has been proposed to genetically modify some plant species threatened by extinction to be resistant to invasive plants and diseases, such as the emerald ash borer in North American and the fungal disease, Ceratocystis platani, in European plane trees.^[158] The papaya ringspot virus devastated papaya trees in Hawaii in the twentieth century until transgenic papaya plants were given pathogen-derived resistance.^[159] However, genetic modification for conservation in plants remains mainly speculative. A unique concern is that a transgenic species may no longer bear enough resemblance to the original species to truly claim that the original species is being conserved. Instead, the transgenic species may be genetically different enough to be considered a new species, thus diminishing the conservation worth of genetic modification.^[158]

Crops

Genetically modified crops are genetically modified plants that are used in

There are three main aims to agricultural advancement; increased production, improved conditions for agricultural workers and sustainability. GM crops contribute by improving harvests through reducing insect pressure, increasing nutrient value and tolerating different abiotic stresses. Despite this potential, as of 2018, the commercialized crops are limited mostly to cash crops like cotton, soybean, maize and canola and the vast majority of the introduced traits provide either herbicide tolerance or insect resistance.^[160] Soybeans accounted for half of all genetically modified crops planted in 2014.^[161] Adoption by farmers has been rapid, between 1996 and 2013, the total surface area of land cultivated with GM crops increased by a factor of 100.^[162] Geographically though the spread has been uneven, with strong growth in the Americas and parts of Asia and little in Europe and Africa.^[160] Its socioeconomic spread has been more even, with approximately 54% of worldwide GM crops grown in developing countries in 2013.^[162] Although doubts have been raised,^[163] most studies have found growing GM crops to be beneficial to farmers through decreased pesticide use as well as increased crop yield and farm profit.^{[164][165][166]}

The majority of GM crops have been modified to be resistant to selected herbicides, usually a

Vaccine production and storage has great potential in transgenic plants. Vaccines are expensive to produce, transport, and administer, so having a system that could produce them locally would allow greater access to poorer and developing areas.[181] As well as purifying vaccines expressed in plants it is also possible to produce edible vaccines in plants. Edible vaccines stimulate the immune system when ingested to protect against certain diseases. Being stored in plants reduces the long-term cost as they can be disseminated without the need for cold storage, don't need to be purified, and have long term stability. Also being housed within plant cells provides some protection from the gut acids upon digestion. However the cost of developing, regulating, and containing transgenic plants is high, leading to most current plant-based vaccine development being applied to veterinary medicine, where the controls are not as strict.^[187]

Genetically modified crops have been proposed as one of the ways to reduce farming-related CO₂ emissions due to higher yield, reduced use of pesticides, reduced use of tractor fuel and no tillage. According to a 2021 study, in EU alone widespread adoption of GE crops would reduce greenhouse gas emissions by 33 million tons of CO₂ equivalent or 7.5% of total farming-related emissions.^[188]

Animals

The vast majority of genetically modified animals are at the research stage with the number close to entering the market remaining small.^[189] As of 2018 only three genetically modified animals have been approved, all in the USA. A goat and a chicken have been engineered to produce medicines and a salmon has increased its own growth.^[190] Despite the differences and difficulties in modifying them, the end aims are much the same as for plants. GM animals are created for research purposes, production of industrial or therapeutic products, agricultural uses, or improving their health. There is also a market for creating genetically modified pets.^[191]

Mammals

The process of genetically engineering mammals is slow, tedious, and expensive. However, new technologies are making genetic modifications easier and more precise.^[192] The first transgenic mammals were produced by injecting viral DNA into embryos and then implanting the embryos in females.^[60] The embryo would develop and it would be hoped that some of the genetic material would be incorporated into the reproductive cells. Then researchers would have to wait until the animal reached breeding age and then offspring would be screened for the presence of the gene in every cell. The development of the CRISPR-Cas9 gene editing system as a cheap and fast way of directly modifying germ cells, effectively halving the amount of time needed to develop genetically modified mammals.^[193]

Scientists have genetically engineered several organisms, including some mammals, to include green fluorescent protein (GFP), for research purposes.^[218] GFP and other similar reporting genes allow easy visualization and localization of the products of the genetic modification.^[219] Fluorescent pigs have been bred to study human organ transplants, regenerating ocular photoreceptor cells, and other topics.^[220] In 2011, green-fluorescent cats were created to help find therapies for HIV/AIDS and other diseases^[221] as feline immunodeficiency virus is related to HIV.^[222]

There have been suggestions that genetic engineering could be used to bring animals back from extinction. It involves changing the genome of a close living relative to resemble the extinct one and is currently being attempted with the passenger pigeon.^[223] Genes associated with the woolly mammoth have been added to the genome of an African Elephant, although the lead researcher says he has no intention of creating live elephants and transferring all the genes and reversing years of genetic evolution is a long way from being feasible.^[224][225] It is more likely that scientists could use this technology to conserve endangered animals by bringing back lost diversity or transferring evolved genetic advantages from adapted organisms to those that are struggling.^[226]

Humans

Genetically modified fish are used for scientific research, as pets and as a food source. Aquaculture is a growing industry, currently providing over half the consumed fish worldwide.^[246] Through genetic engineering it is possible to increase growth rates, reduce food intake, remove allergenic properties, increase cold tolerance and provide disease resistance. Fish can also be used to detect aquatic pollution or function as bioreactors.^[247]

Several groups have been developing zebrafish to detect pollution by attaching fluorescent proteins to genes activated by the presence of pollutants. The fish will then glow and can be used as environmental sensors.^[248][249] The GloFish is a brand of genetically modified fluorescent zebrafish with bright red, green, and orange fluorescent color. It was originally developed by one of the groups to detect pollution, but is now part of the ornamental fish trade, becoming the first genetically modified animal to become publicly available as a pet when in 2003 it was introduced for sale in the USA.^[250]

GM fish are widely used in basic research in genetics and development. Two species of fish, zebrafish and

GM fish have been developed with promoters driving an over-production of growth hormone for use in the aquaculture industry to increase the speed of development and potentially reduce fishing pressure on wild stocks. This has resulted in dramatic growth enhancement in several species, including salmon,^[256] trout^[257] and tilapia.^[258] AquaBounty Technologies, a biotechnology company, have produced a salmon (called AquAdvantage salmon) that can mature in half the time as wild salmon.^[259] It obtained regulatory approval in 2015, the first non-plant GMO food to be commercialized.^[260] As of August 2017, GMO salmon is being sold in Canada.^[261] Sales in the US started in May 2021.^[262]

Insects

Due to their significance to human health, scientists are looking at ways to control mosquitoes through genetic engineering. Malaria-resistant mosquitoes have been developed in the laboratory by inserting a gene that reduces the development of the malaria parasite[268] and then use homing endonucleases to rapidly spread that gene throughout the male population (known as a gene drive).^[269][270] This approach has been taken further by using the gene drive to spread a lethal gene.^[271][272] In trials the populations of Aedes aegypti mosquitoes, the single most important carrier of dengue fever and Zika virus, were reduced by between 80% and by 90%.^{[273][274][272]} Another approach is to use a sterile insect technique, whereby males genetically engineered to be sterile out compete viable males, to reduce population numbers.^[275]

Other insect pests that make attractive targets are moths. Diamondback moths cause US$4 to $5 billion of damage each year worldwide.^[276] The approach is similar to the sterile technique tested on mosquitoes, where males are transformed with a gene that prevents any females born from reaching maturity.^[277] They underwent field trials in 2017.^[276] Genetically modified moths have previously been released in field trials.^[278] In this case a strain of pink bollworm that were sterilized with radiation were genetically engineered to express a red fluorescent protein making it easier for researchers to monitor them.^[279]

Silkworm, the larvae stage of

human collagen α-chain, mouse monoclonal antibody and N-glycanase.^[281] Silkworms have been created that produce spider silk, a stronger but extremely difficult to harvest silk,^[282] and even novel silks.^[283]

Other

Systems have been developed to create transgenic organisms in a wide variety of other animals. Chickens have been genetically modified for a variety of purposes. This includes studying

developmental biology research. GM frogs can also be used as pollution sensors, especially for endocrine disrupting chemicals.^[288] There are proposals to use genetic engineering to control cane toads in Australia.^[289][290]

The

double stranded RNA.^[293] It is also relatively easy to produce stable transgenic nematodes and this along with RNAi are the major tools used in studying their genes.^[294] The most common use of transgenic nematodes has been studying gene expression and localization by attaching reporter genes. Transgenes can also be combined with RNAi techniques to rescue phenotypes, study gene function, image cell development in real time or control expression for different tissues or developmental stages.^[294] Transgenic nematodes have been used to study viruses,^[295] toxicology,^[296] diseases,^[297][298] and to detect environmental pollutants.^[299]

The gene responsible for

immunity and certain developmental processes.^[304] Other animals that have been genetically modified include snails,^[305] geckos, turtles,^[306] crayfish, oysters, shrimp, clams, abalone^[307] and sponges.^[308]

Regulation

Main article: Regulation of genetic engineering

Genetically modified organisms are regulated by government agencies. This applies to research as well as the release of genetically modified organisms, including crops and food. The development of a regulatory framework concerning genetic engineering began in 1975, at

Asilomar meeting recommended a set of guidelines regarding the cautious use of recombinant technology and any products resulting from that technology.^[309] The Cartagena Protocol on Biosafety was adopted on 29 January 2000 and entered into force on 11 September 2003.^[310] It is an international treaty that governs the transfer, handling, and use of genetically modified organisms.^[311] One hundred and fifty-seven countries are members of the Protocol and many use it as a reference point for their own regulations.^[312]

Universities and research institutes generally have a special committee that is responsible for approving any experiments that involve genetic engineering. Many experiments also need permission from a national regulatory group or legislation. All staff must be trained in the use of GMOs and all laboratories must gain approval from their regulatory agency to work with GMOs.[313] The legislation covering GMOs are often derived from regulations and guidelines in place for the non-GMO version of the organism, although they are more severe.^[314] There is a near-universal system for assessing the relative risks associated with GMOs and other agents to laboratory staff and the community. They are assigned to one of four risk categories based on their virulence, the severity of the disease, the mode of transmission, and the availability of preventive measures or treatments. There are four biosafety levels that a laboratory can fall into, ranging from level 1 (which is suitable for working with agents not associated with disease) to level 4 (working with life-threatening agents). Different countries use different nomenclature to describe the levels and can have different requirements for what can be done at each level.^[314]

There are differences in the regulation for the release of GMOs between countries, with some of the most marked differences occurring between the US and Europe.^[315] Regulation varies in a given country depending on the intended use of the products of the genetic engineering. For example, a crop not intended for food use is generally not reviewed by authorities responsible for food safety.^[316] Some nations have banned the release of GMOs or restricted their use, and others permit them with widely differing degrees of regulation.^{[317][318][319][320]} In 2016, thirty eight countries officially ban or prohibit the cultivation of GMOs and nine (Algeria, Bhutan, Kenya, Kyrgyzstan, Madagascar, Peru, Russia, Venezuela and Zimbabwe) ban their importation.^[321] Most countries that do not allow GMO cultivation do permit research using GMOs.^[322] Despite regulation, illegal releases have sometimes occurred, due to weakness of enforcement.^[8]

The European Union (EU) differentiates between approval for cultivation within the EU and approval for import and processing.^[323] While only a few GMOs have been approved for cultivation in the EU a number of GMOs have been approved for import and processing.^[324] The cultivation of GMOs has triggered a debate about the market for GMOs in Europe.^[325] Depending on the coexistence regulations, incentives for cultivation of GM crops differ.^[326] The US policy does not focus on the process as much as other countries, looks at verifiable scientific risks and uses the concept of substantial equivalence.^[327] Whether gene edited organisms should be regulated the same as genetically modified organism is debated. USA regulations sees them as separate and does not regulate them under the same conditions, while in Europe a GMO is any organism created using genetic engineering techniques.^[28]

One of the key issues concerning regulators is whether GM products should be labeled. The

feed which contains greater than 0.9% of approved GMOs must be labeled.^[334] In 2014, sales of products that had been labeled as non-GMO grew 30 percent to $1.1 billion.^[335]

Controversy

See also: Genetically modified food controversies

There is controversy over GMOs, especially with regard to their release outside laboratory environments. The dispute involves consumers, producers, biotechnology companies, governmental regulators, non-governmental organizations, and scientists. Many of these concerns involve GM crops and whether food produced from them is safe and what impact growing them will have on the environment. These controversies have led to litigation, international trade disputes, and protests, and to restrictive regulation of commercial products in some countries.

food supply.^[337]

There is a scientific consensus^{[338][339][340][341]} that currently available food derived from GM crops poses no greater risk to human health than conventional food,^{[342][343][344][345][346]} but that each GM food needs to be tested on a case-by-case basis before introduction.^{[347][348][349]} Nonetheless, members of the public are much less likely than scientists to perceive GM foods as safe.^{[350][351][352][353]} The legal and regulatory status of GM foods varies by country, with some nations banning or restricting them, and others permitting them with widely differing degrees of regulation.^{[354][355][356][357]}

As late as the 1990s

triploid, 99% are reproductively sterile, and raised in areas where escaped salmon could not survive.^[364][365] Bacteria have also been modified to depend on nutrients that cannot be found in nature,^[366] and genetic use restriction technology has been developed, though not yet marketed, that causes the second generation of GM plants to be sterile.^[367]

Other environmental and

monarch butterflies. Follow up studies have since shown that the toxicity levels encountered in the field were not high enough to harm the larvae.^[372]

Accusations that scientists are "playing God" and other religious issues have been ascribed to the technology from the beginning.^[373] With the ability to genetically engineer humans now possible there are ethical concerns over how far this technology should go, or if it should be used at all.^[374] Much debate revolves around where the line between treatment and enhancement is and whether the modifications should be inheritable.^[375] Other concerns include contamination of the non-genetically modified food supply,^[376][377] the rigor of the regulatory process,^[378][379] consolidation of control of the food supply in companies that make and sell GMOs,^[380] exaggeration of the benefits of genetic modification,^[381] or concerns over the use of herbicides with glyphosate.^[382] Other issues raised include the patenting of life^[383] and the use of intellectual property rights.^[384]

There are large differences in consumer acceptance of GMOs, with Europeans more likely to view GM food negatively than North Americans.^[385] GMOs arrived on the scene as the public confidence in food safety, attributed to recent food scares such as Bovine spongiform encephalopathy and other scandals involving government regulation of products in Europe, was low.^[386] This along with campaigns run by various non-governmental organizations (NGO) have been very successful in blocking or limiting the use of GM crops.^[387] NGOs like the Organic Consumers Association, the Union of Concerned Scientists,^{[388][389][390]} Greenpeace and other groups have said that risks have not been adequately identified and managed^[391] and that there are unanswered questions regarding the potential long-term impact on human health from food derived from GMOs. They propose mandatory labeling^[392][393] or a moratorium on such products.^{[380][378][394]}

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341. ^ But see also:
     
     • Domingo, José L.; Bordonaba, Jordi Giné (2011). "A literature review on the safety assessment of genetically modified plants" (PDF). Environment International. 37 (4): 734–742.
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       (PDF) from the original on 31 August 2019. I began this article with the testimonials from respected scientists that there is literally no scientific controversy over the health effects of GMOs. My investigation into the scientific literature tells another story.

     And contrast:
     

     • Panchin, Alexander Y.; Tuzhikov, Alexander I. (14 January 2016). "Published GMO studies find no evidence of harm when corrected for multiple comparisons". Critical Reviews in Biotechnology. 37 (2): 213–217.
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       Despite various concerns, today, the American Association for the Advancement of Science, the World Health Organization, and many independent international science organizations agree that GMOs are just as safe as other foods. Compared with conventional breeding techniques, genetic engineering is far more precise and, in most cases, less likely to create an unexpected outcome.
342. ^ "Statement by the AAAS Board of Directors on Labeling of Genetically Modified Foods" (PDF). American Association for the Advancement of Science. 20 October 2012. Retrieved 30 August 2019. The EU, for example, has invested more than €300 million in research on the biosafety of GMOs. Its recent report states: 'The main conclusion to be drawn from the efforts of more than 130 research projects, covering a period of more than 25 years of research and involving more than 500 independent research groups, is that biotechnology, and in particular GMOs, are not per se riskier than e.g. conventional plant breeding technologies.' The World Health Organization, the American Medical Association, the U.S. National Academy of Sciences, the British Royal Society, and every other respected organization that has examined the evidence has come to the same conclusion: consuming foods containing ingredients derived from GM crops is no riskier than consuming the same foods containing ingredients from crop plants modified by conventional plant improvement techniques.
     
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347. ^ "Frequently asked questions on genetically modified foods". World Health Organization. Retrieved 30 August 2019. Different GM organisms include different genes inserted in different ways. This means that individual GM foods and their safety should be assessed on a case-by-case basis and that it is not possible to make general statements on the safety of all GM foods.
     
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External links

• ISAAA database
• GMO-Compass: Information on genetically modified organisms