Genetically modified organism
A genetically modified organism (GMO) is any organism whose genetic material has been altered using genetic engineering techniques (i.e., a genetically engineered organism). GMOs are used to produce many medications and genetically modified foods and are widely used in scientific research and the production of other goods. The term GMO is very close to the technical legal term, 'living modified organism', defined in the Cartagena Protocol on Biosafety, which regulates international trade in living GMOs (specifically, "any living organism that possesses a novel combination of genetic material obtained through the use of modern biotechnology").
A more specifically defined type of GMO is a "transgenic organism." This is an organism whose genetic makeup has been altered by the addition of genetic material from an unrelated organism. This should not be confused with the more general way in which "GMO" is used to classify genetically altered organisms, as typically GMOs are organisms whose genetic makeup has been altered without the addition of genetic material from an unrelated organism.
Genetic modification involves the mutation, insertion, or deletion of genes. Inserted genes usually come from a different species in a form of horizontal gene-transfer. In nature this can occur when exogenous DNA penetrates the cell membrane for any reason. This can be accomplished artificially by:
- attaching the genes to a virus.
- physically inserting the extra DNA into the nucleus of the intended host with a very small syringe.
- using electroporation (that is, introducing DNA from one organism into the cell of another by use of an electric pulse).
- firing small particles from a gene gun.
Other methods exploit natural forms of gene transfer, such as the ability of Agrobacterium to transfer genetic material to plants, or the ability of lentiviruses to transfer genes to animal cells.
Humans have domesticated plants and animals since around 12,000 BCE, using selective breeding or artificial selection (as contrasted with natural selection).:25 The process of selective breeding, in which organisms with desired traits (and thus with the desired genes) are used to breed the next generation and organisms lacking the trait are not bred, is a precursor to the modern concept of genetic modification.:1:1 Various advancements in genetics allowed humans to directly alter the DNA and therefore genes of organisms. In 1972 Paul Berg created the first recombinant DNA molecule when he combined DNA from a monkey virus with that of the lambda virus.
Herbert Boyer and Stanley Cohen made the first genetically modified organism (GMO) in 1973. They took a gene from a bacterium that provided resistance to the antibiotic kanamycin, inserted it into a plasmid and then induced another bacteria to uptake the plasmid. The bacteria was then able to survive in the presence of kanamycin. Boyer and Cohen expressed other genes in bacteria. This included genes from the toad Xenopus laevis in 1974, creating the first GMO expressing a gene from an organism from different kingdom.
In 1974 Rudolf Jaenisch created a transgenic mouse by introducing foreign DNA into its embryo, making it the world’s first transgenic animal. However it took another eight years before transgenic mice were developed that passed the transgene to their offspring. Genetically modified mice were created in 1984 that carried cloned oncogenes, predisposed them to developing cancer. Mice with genes knocked out (knockout mouse) were created in 1989. The first transgenic livestock were produced in 1985 and the first animal to synthesise transgenic proteins in their milk were mice, engineered to produce human tissue plasminogen activator in 1987.
In 1983 the first genetically engineered plant was developed by Michael W. Bevan, Richard B. Flavell and Mary-Dell Chilton. They infected tobacco with Agrobacterium transformed with an antibiotic resistance gene and through tissue culture techniques were able to grow a new plant containing the resistance gene. The gene gun was invented in 1987, allowing transformation of plants not susceptible to Agrobacterium infection. In 2000, Vitamin A-enriched golden rice, was the first plant developed with increased nutrient value.
In 1976 Genentech, the first genetic engineering company was founded by Herbert Boyer and Robert Swanson; a year later, the company produced a human protein (somatostatin) in E.coli. Genentech announced the production of genetically engineered human insulin in 1978. The insulin produced by bacteria, branded humulin, was approved for release by the Food and Drug Administration in 1982. In 1988 the first human antibodies were produced in plants. In 1987, the ice-minus strain of P. syringae became the first genetically modified organism to be released into the environment when a strawberry field and a potato field in California were sprayed with it.
The first genetically modified crop, an antibiotic-resistant tobacco plant, was produced in 1982. China was the first country to commercialize transgenic plants, introducing a virus-resistant tobacco in 1992. In 1994 Calgene attained approval to commercially release the Flavr Savr tomato, the first genetically modified food. Also in 1994, the European Union approved tobacco engineered to be resistant to the herbicide bromoxynil, making it the first genetically engineered crop commercialized in Europe. An insect resistant Potato was approved for release in the USA in 1995, and by 1996 approval had been granted to commercially grow 8 transgenic crops and one flower crop (carnation) in 6 countries plus the EU.
In 2010, scientists at the J. Craig Venter Institute, announced that they had created the first synthetic bacterial genome. They named it Synthia and it was the world's first synthetic life form.
The first genetically modified animal to be commercialised was the GloFish, a Zebra fish with a fluorescent gene added that allows it to glow in the dark under ultraviolet light. The first genetically modified animal to be approved for food use was AquAdvantage salmon in 2015. The salmon were transformed with a growth hormone-regulating gene from a Pacific Chinook salmon and a promoter from an ocean pout enabling it to grow year-round instead of only during spring and summer.
GMOs are used in biological and medical research, production of pharmaceutical drugs, experimental medicine (e.g. gene therapy and vaccines against the Ebola virus), and agriculture (e.g. golden rice, resistance to herbicides), with developing uses in conservation. The term "genetically modified organism" does not always imply, but can include, targeted insertions of genes from one species into another. For example, a gene from a jellyfish, encoding a fluorescent protein called GFP, or green fluorescent protein, can be physically linked and thus co-expressed with mammalian genes to identify the location of the protein encoded by the GFP-tagged gene in the mammalian cell. Such methods are useful tools for biologists in many areas of research, including those who study the mechanisms of human and other diseases or fundamental biological processes in eukaryotic or prokaryotic cells.
They continue to be important model organisms for experiments in genetic engineering. In the field of synthetic biology, they have been used to test various synthetic approaches, from synthesizing genomes to creating novel nucleotides.
Genetically modified bacteria are used to produce the protein insulin to treat diabetes. Similar bacteria have been used to produce biofuels, clotting factors to treat haemophilia, and human growth hormone to treat various forms of dwarfism.
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 70% since 2005.
In addition, various genetically engineered micro-organisms are routinely used as sources of enzymes for the manufacture of a variety of processed foods. These include alpha-amylase from bacteria, which converts starch to simple sugars, chymosin from bacteria or fungi, which clots milk protein for cheese making, and pectinesterase from fungi, which improves fruit juice clarity.
Transgenic plants have been engineered for scientific research, to create new colours in plants, and to create different crops.
In research, plants are engineered to help discover the functions of certain genes. One way to do this is to knock out the gene of interest and see what phenotype develops. Another strategy is to attach the gene to a strong promoter and see what happens when it is over expressed. A common technique used to find out where the gene is expressed is to attach it to GUS or a similar reporter gene that allows visualisation of the location.'
After thirteen years of collaborative research, an Australian company – Florigene, and a Japanese company – Suntory, created a blue rose (actually lavender or mauve) in 2004. The genetic engineering involved three alterations – adding two genes, and interfering with another. One of the added genes was for the blue plant pigment delphinidin cloned from the pansy. The researchers then used RNA interference (RNAi) technology to depress all color production by endogenous genes by blocking a crucial protein in color production, called dihydroflavonol 4-reductase) (DFR), and adding a variant of that protein that would not be blocked by the RNAi but that would allow the delphinidin to work. The roses are sold in Japan, the United States, and Canada. Florigene has also created and sells lavender-colored carnations that are genetically engineered in a similar way.
Simple plants and plant cells have been genetically engineered for production of biopharmaceuticals in bioreactors as opposed to cultivating plants in open fields. Work has been done with duckweed Lemna minor, the algae Chlamydomonas reinhardtii and the moss Physcomitrella patens. An Israeli company, Protalix, has developed a method to produce therapeutics in cultured transgenic carrot and tobacco cells. Protalix and its partner, Pfizer, received FDA approval to market its drug Elelyso, a treatment for Gaucher's disease, in 2012.
Genetically modified crops
Genetically modified crops (GM crops, or biotech crops) are plants used in agriculture, the DNA of which has been modified using genetic engineering techniques. In most cases the aim is to introduce a new trait to the plant which does not occur naturally in the species. Examples in food crops include resistance to certain pests, diseases, or environmental conditions, reduction of spoilage, or resistance to chemical treatments (e.g. resistance to a herbicide), or improving the nutrient profile of the crop. Examples in non-food crops include production of pharmaceutical agents, biofuels, and other industrially useful goods, as well as for bioremediation.
Farmers have widely adopted GM technology. Between 1996 and 2013, the total surface area of land cultivated with GM crops increased by a factor of 100, from 17,000 square kilometers (4,200,000 acres) to 1,750,000 km2 (432 million acres). 10% of the world's croplands were planted with GM crops in 2010. In the US, by 2014, 94% of the planted area of soybeans, 96% of cotton and 93% of corn were genetically modified varieties. In recent years GM crops expanded rapidly in developing countries. In 2013 approximately 18 million farmers grew 54% of worldwide GM crops in developing countries.
For discussions of issues about GM crops and GM food, see the Controversies section below and the article on genetically modified food controversies.
Cisgenesis, sometimes also called intragenesis, is a product designation for a category of genetically engineered plants. A variety of classification schemes have been proposed that order genetically modified organisms based on the nature of introduced genotypical changes rather than the process of genetic engineering.
While some genetically modified plants are developed by the introduction of a gene originating from distant, sexually incompatible species into the host genome, cisgenic plants contain genes that have been isolated either directly from the host species or from sexually compatible species. The new genes are introduced using recombinant DNA methods and gene transfer. Some scientists hope that the approval process of cisgenic plants might be simpler than that of proper transgenics, but it remains to be seen.
Conservation in plants
Genetically modified organisms have been proposed to aid conservation of plant species threatened by extinction. Many trees face the threat of invasive plants and diseases, such as the emerald ash borer in North American and the fungal disease, Ceratocystis platani, in European plane trees. A suggested solution to increase the resilience of threatened tree species is to genetically modify individuals by transferring resistant genes. Papaya trees are an example of a species that was successfully conserved using genetic modification. The papaya ringspot virus (PRSV) devastated papaya trees in Hawaii in the twentieth century until transgenic papaya plants were given pathogen-derived resistance.
However, genetic modification for conservation in plants remains mainly speculative and further experimentation is needed before the technique can be widely implemented. A main concern with using genetic modification for conservation purposes 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.
Genetically modified mammals are an important category of genetically modified organisms. Ralph L. Brinster and Richard Palmiter developed the techniques responsible for transgenic mice, rats, rabbits, sheep, and pigs in the early 1980s, and established many of the first transgenic models of human disease, including the first carcinoma caused by a transgene. The process of genetically engineering animals is a slow, tedious, and expensive process. However, new technologies are making genetic modifications easier and more precise.
The first transgenic (genetically modified) animal was produced by injecting DNA into mouse embryos then implanting the embryos in female mice.
Genetically modified animals currently being developed can be placed into six different broad classes based on the intended purpose of the genetic modification:
- to research human diseases (for example, to develop animal models for these diseases);
- to produce industrial or consumer products (fibres for multiple uses);
- to produce products intended for human therapeutic use (pharmaceutical products or tissue for implantation);
- to enrich or enhance the animals' interactions with humans (hypo-allergenic pets);
- to enhance production or food quality traits (faster growing fish, pigs that digest food more efficiently);
- to improve animal health (disease resistance)
Genetically modified (genetically engineered) animals are becoming more vital to the discovery and development of cures and treatments for many serious diseases. By altering the DNA or transferring DNA to an animal, we can develop certain proteins that may be used in medical treatment. Stable expressions of human proteins have been developed in many animals, including sheep, pigs, and rats. Human-alpha-1-antitrypsin, which has been tested in sheep and is used in treating humans with this deficiency and transgenic pigs with human-histo-compatibility have been studied in the hopes that the organs will be suitable for transplant with less chances of rejection.
Scientists have genetically engineered several organisms, including some mammals, to include green fluorescent protein (GFP), first observed in the jellyfish, Aequorea victoria in 1962, for medical research purposes (Chalfie, Shimoura, and Tsien were awarded the Nobel prize in Chemistry in 2008 for the discovery and development of GFP ). For example, fluorescent pigs have been bred to study human organ transplants (xenotransplantation), regenerating ocular photoreceptor cells, and other topics. In 2011 a Japanese-American team created green-fluorescent cats to find therapies for HIV/AIDS and other diseases as feline immunodeficiency virus (FIV) is related to HIV.
In 2009, scientists in Japan announced that they had successfully transferred a gene into a primate species (marmosets) and produced a stable line of breeding transgenic primates for the first time. Their first research target for these marmosets was Parkinson's disease, but they were also considering amyotrophic lateral sclerosis and Huntington's disease.
Producing human therapeutics
Within the field known as pharming, intensive research has been conducted to develop transgenic animals that produce biotherapeutics. On 6 February 2009, the U.S. Food and Drug Administration approved the first human biological drug produced from such an animal, a goat. The drug, ATryn, is an anticoagulant which reduces the probability of blood clots during surgery or childbirth. It is extracted from the goat's milk.
Production or food quality traits
Enviropig was a genetically enhanced line of Yorkshire pigs in Canada created with the capability of digesting plant phosphorus more efficiently than conventional Yorkshire pigs. The project ended in 2012. These pigs produced the enzyme phytase, which breaks down the indigestible phosphorus, in their saliva. The enzyme was introduced into the pig chromosome by pronuclear microinjection. With this enzyme, the animal is able to digest cereal grain phosphorus. The use of these pigs would reduce the potential of water pollution since they excrete from 30 to 70.7% less phosphorus in manure depending upon the age and diet. The lower concentrations of phosphorus in surface runoff reduces algal growth, because phosphorus is the limiting nutrient for algae. Because algae consume large amounts of oxygen, it can result in dead zones for fish.
In 2011, Chinese scientists generated dairy cows genetically engineered with genes from human beings to produce milk that would be the same as human breast milk. This could potentially benefit mothers who cannot produce breast milk but want their children to have breast milk rather than formula. Aside from milk production, the researchers claim these transgenic cows to be identical to regular cows. Two months later scientists from Argentina presented Rosita, a transgenic cow incorporating two human genes, to produce milk with similar properties as human breast milk. In 2012, researchers from New Zealand also developed a genetically engineered cow that produced allergy-free milk.
Goats have been genetically engineered to produce milk with strong spiderweb-like silk proteins in their milk.
Human gene therapy
Gene therapy, uses genetically modified viruses to deliver genes that can cure disease in humans. Although gene therapy is still relatively new, it has had some successes. It has been used to treat genetic disorders such as severe combined immunodeficiency, and Leber's congenital amaurosis. Treatments are also being developed for a range of other currently incurable diseases, such as cystic fibrosis, sickle cell anemia, Parkinson's disease, cancer, diabetes, heart disease and muscular dystrophy.
Genetically modified organisms have been used to conserve European wild rabbits in the Iberian peninsula and Australia. In both cases, the genetically modified organism used was a myxoma virus, but for opposite purposes: to protect the endangered population in Europe with immunizations and to regulate the overabundant population in Australia with contraceptives.
In the Iberian peninsula, the European wild rabbit population has experienced a sharp decline from viral diseases and overhunting. To protect the species from viral diseases, the myxoma virus was genetically modified to immunize the rabbits. The European wild rabbit population in Australia faces the opposite problem: lack of natural predators has made the introduced species invasive. The same myxoma virus was genetically modified to lower fertility in the Australian rabbit population.
Genetically modified fish are used for scientific research and as pets, and are being considered for use as food and as aquatic pollution sensors.
GM fish are widely used in basic research in genetics and development. Two species of fish, zebrafish and medaka, are most commonly modified because they have optically clear chorions (membranes in the egg), rapidly develop, and the 1-cell embryo is easy to see and microinject with transgenic DNA.
The GloFish is a patented brand of genetically modified (GM) fluorescent zebrafish with bright red, green, and orange fluorescent color. Although not originally developed for the ornamental fish trade, it became the first genetically modified animal to become publicly available as a pet when it was introduced for sale in 2003. They were quickly banned for sale in California.
GM fish have been developed with promoters driving an over-production of "all fish" 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, trout and tilapia. AquaBounty Technologies, a biotechnology company working on bringing a GM salmon to market, claims that their GM AquAdvantage salmon can mature in half the time as wild salmon. AquaBounty applied for regulatory approval to market their GM salmon in the US, and was approved in November 2015. On 25 November 2013 Canada approved commercial scale production and export of GM Salmon eggs but they are not approved for human consumption in Canada.
Several academic groups have been developing GM zebrafish to detect aquatic pollution. The lab that originated the GloFish discussed above originally developed them to change color in the presence of pollutants, to be used as environmental sensors. A lab at University of Cincinnati has been developing GM zebrafish for the same purpose, as has a lab at Tulane University.
Genetically modified frogs are used for scientific research and are widely used in basic research including genetics and early development. Two species of frog, Xenopus laevis and Xenopus tropicalis, are most commonly used.
In biological research, transgenic fruit flies (Drosophila melanogaster) are model organisms used to study the effects of genetic changes on development. Fruit flies are often preferred over other animals due to their short life cycle, low maintenance requirements, and relatively simple genome compared to many vertebrates.
In 2010, scientists created "malaria-resistant mosquitoes" in the laboratory. The World Health Organization estimated that malaria killed almost one million people in 2008. Genetically modified male mosquitoes containing a lethal gene have been developed to combat the spread of dengue fever and the Zika virus. Aedes aegypti mosquitoes, the single most important carrier of dengue fever and the Zika virus, were reduced by 80% in a 2010 trial of these GM mosquitoes in the Cayman Islands and by 90% in a 2015 trial in Bahia, Brazil. In comparison, the Florida Keys Mosquito Control District has achieved only 30%-60% population reduction with traps and pesticide spraying. In 2016 FDA approved a genetically modified mosquito intervention for Key West, Florida. UK firm Oxitec proposed the release of millions of modified male (non-biting) mosquitoes to compete with wild males for mates. The males are engineered so that their offspring die before maturing, helping to eradicate mosquito-borne disease. Final approval was to be based on a local referendum to be held in November. Andrea Crisanti, a molecular biologist at Imperial College in London is working on ways to stop the A. gambiae mosquito from transmitting disease.
A strain of Pectinophora gossypiella (Pink bollworm) has been genetically engineered to express a red fluorescent protein. This allows researchers to monitor bollworms that have been sterilized by radiation and released to reduce bollworm infestation. The strain has been field tested for over three years and has been approved for release.
Cnidaria such as Hydra and the sea anemone Nematostella vectensis are attractive model organisms to study the evolution of immunity and certain developmental processes. An important technical breakthrough was the development of procedures for generation of stable transgenic hydras and sea anemones by embryo microinjection.
The regulation of genetic engineering concerns the approaches taken by governments to assess and manage the risks associated with the use of genetic engineering technology and the development and release of genetically modified organisms (GMO), including genetically modified crops and genetically modified fish. There are differences in the regulation of GMOs between countries, with some of the most marked differences occurring between the USA and Europe. 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. The European Union differentiates between approval for cultivation within the EU and approval for import and processing. While only a few GMOs have been approved for cultivation in the EU a number of GMOs have been approved for import and processing. The cultivation of GMOs has triggered a debate about the market for GMOs in Europe. Depending on the coexistence regulations, incentives for cultivation of GM crops differ.
There is controversy over GMOs, especially with regard to their use in producing food. The dispute involves buyers, biotechnology companies, governmental regulators, nongovernmental organizations, and scientists. The key areas of controversy related to GMO food are whether GM food should be labeled, the role of government regulators, the effect of GM crops on health and the environment, the effect on pesticide resistance, the impact of GM crops for farmers, and the role of GM crops in feeding the world population. In 2014, sales of products that had been labeled as non-GMO grew 30 percent to $1.1 billion.
There is a scientific consensus that currently available food derived from GM crops poses no greater risk to human health than conventional food, but that each GM food needs to be tested on a case-by-case basis before introduction. Nonetheless, members of the public are much less likely than scientists to perceive GM foods as safe. 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.
No reports of ill effects have been proven in the human population from ingesting GM food. Although labeling of GMO products in the marketplace is required in many countries, it is not required in the United States and no distinction between marketed GMO and non-GMO foods is recognized by the US FDA. In a May 2014 article in The Economist it was argued that, while GM foods could potentially help feed 842 million malnourished people globally, laws such as the one passed in Vermont, to require labeling of foods containing genetically modified ingredients, could have the unintended consequence of interrupting the process of spreading GM technologies to impoverished countries that suffer with food security problems.
A 2014 critical review focused on histopathology studies on rat digestive tracts (eating approved widely eaten GM crops) found significant flaws, inadequacies, and a lack of transparency in methodology and results. Published studies could be found for only 19% of these widely eaten crops. Most of reviewed studies were performed after the approval of crop. The authors conclude that more research is needed, including long-term animal feeding studies and thorough histopathological investigations.
The Organic Consumers Association, and the Union of Concerned Scientists, and Greenpeace stated that risks have not been adequately identified and managed, and they have questioned the objectivity of regulatory authorities. Some health groups say there are unanswered questions regarding the potential long-term impact on human health from food derived from GMOs, and propose mandatory labeling or a moratorium on such products. Concerns include contamination of the non-genetically modified food supply, effects of GMOs on the environment and nature, the rigor of the regulatory process, and consolidation of control of the food supply in companies that make and sell GMOs, or concerns over the use of herbicides with glyphosate.
- Chimera (genetics)
- Council for Responsible Genetics
- Detection of genetically modified organisms
- Gene flow
- Gene pool
- Genetic erosion
- Horizontal gene transfer
- Non-GMO Project
- Organic farming
- SMART breeding
- Sperm-mediated gene transfer
- Timeline of genetically modified organisms
- StarLink corn recall
- "History of Genetically Modified Foods". umich.edu. Archived from the original on 21 October 2015.
- Cornell Chronicle, 14 May 1987, page 3. Biologists invent gun for shooting cells with DNA Archived 29 October 2013 at the Wayback Machine.
- Sanford, JC; et al. (1987). "Delivery of substances into cells and tissues using a particle bombardment process". Journal of Particulate Science and Technology. 5: 27–37. doi:10.1080/02726358708904533.
- Klein, TM; et al. (1987). "High-velocity microprojectiles for delivering nucleic acids into living cells". Nature. 327 (6117): 70–73. Bibcode:1987Natur.327...70K. doi:10.1038/327070a0.
- Lee LY, Gelvin SB (February 2008). "T-DNA binary vectors and systems". Plant Physiol. 146 (2): 325–332. OCLC 1642351. PMC . PMID 18250230. doi:10.1104/pp.107.113001.
- Park F (October 2007). "Lentiviral vectors: are they the future of animal transgenesis?". Physiol. Genomics. 31 (2): 159–173. OCLC 37367250. PMID 17684037. doi:10.1152/physiolgenomics.00069.2007.
- Noel Kingsbury. Hybrid: The History and Science of Plant Breeding University of Chicago Press, Oct 15, 2009
- Clive Root (2007). Domestication. Greenwood Publishing Groups.
- Daniel Zohary; Maria Hopf; Ehud Weiss (2012). Domestication of Plants in the Old World: The Origin and Spread of Plants in the Old World. Oxford University Press.
- Jackson, DA; Symons, RH; Berg, P (1 October 1972). "Biochemical Method for Inserting New Genetic Information into DNA of Simian Virus 40: Circular SV40 DNA Molecules Containing Lambda Phage Genes and the Galactose Operon of Escherichia coli". PNAS. 69 (10): 2904–2909. Bibcode:1972PNAS...69.2904J. PMC . PMID 4342968. doi:10.1073/pnas.69.10.2904.
- M. K. Sateesh (25 August 2008). Bioethics And Biosafety. I. K. International Pvt Ltd. pp. 456–. ISBN 978-81-906757-0-3. Retrieved 27 March 2013.
- "Genome and genetics timeline – 1973". Genome news network.
- Morrow, J. F.; Cohen, S. N.; Chang, A. C.; Boyer, H. W.; Goodman, H. M.; Helling, R. B. (1974-05-01). "Replication and transcription of eukaryotic DNA in Escherichia coli". Proceedings of the National Academy of Sciences of the United States of America. 71 (5): 1743–1747. Bibcode:1974PNAS...71.1743M. ISSN 0027-8424. PMC . PMID 4600264. doi:10.1073/pnas.71.5.1743.
- Jaenisch, R. and Mintz, B. (1974 ) Simian virus 40 DNA sequences in DNA of healthy adult mice derived from preimplantation blastocysts injected with viral DNA. Proc. Natl. Acad. 71(4):1250–1254 
- "'Any idiot can do it.' Genome editor CRISPR could put mutant mice in everyone's reach". Science | AAAS. 2016-11-02. Retrieved 2016-12-02.
- Gordon, J.; Ruddle, F. (1981). "Integration and stable germ line transmission of genes injected into mouse pronuclei". Science. 214 (4526): 1244–6. Bibcode:1981Sci...214.1244G. PMID 6272397. doi:10.1126/science.6272397.
- Costantini, F.; Lacy, E. (1981). "Introduction of a rabbit β-globin gene into the mouse germ line". Nature. 294 (5836): 92–4. Bibcode:1981Natur.294...92C. PMID 6945481. doi:10.1038/294092a0.
- Hanahan, D.; Wagner, E. F.; Palmiter, R. D. (2007). "The origins of oncomice: A history of the first transgenic mice genetically engineered to develop cancer". Genes & Development. 21 (18): 2258–2270. PMID 17875663. doi:10.1101/gad.1583307.
- Brophy, B.; Smolenski, G.; Wheeler, T.; Wells, D.; l'Huillier, P.; Laible, G. T. (2003). "Cloned transgenic cattle produce milk with higher levels of β-casein and κ-casein". Nature Biotechnology. 21 (2): 157–162. PMID 12548290. doi:10.1038/nbt783.
- A. John Clark (1998). "The Mammary Gland as a Bioreactor: Expression, Processing, and Production of Recombinant Proteins". Journal of Mammary Gland Biology and Neoplasia. 3 (3): 337–350. PMID 10819519. doi:10.1023/a:1018723712996.
- K. Gordon; E. Lee; J. Vitale; A. Smith; H. Westphal; L. Hennighausen (1987). "Production of human tissue plasmnogen activator in transgenic mouse milk". Biotechnology. 5 (11): 1183±1187. doi:10.1038/nbt1187-1183.
- Bevan, M. W.; Flavell, R. B.; Chilton, M. D. (1983). "A chimaeric antibiotic resistance gene as a selectable marker for plant cell transformation". Nature. 304 (5922): 184–187. Bibcode:1983Natur.304..184B. doi:10.1038/304184a0.
- Roger Segelken for the Cornell Chronicle. Mary 14, 1987. Biologists Invent Gun for Shooting Cells with DNA Issue available as pdf download here , page 3
- Ye, Xudong; Al-Babili, Salim; Klöti, Andreas; Zhang, Jing; Lucca, Paola; Beyer, Peter; Potrykus, Ingo (2000-01-14). "Engineering the Provitamin A (β-Carotene) Biosynthetic Pathway into (Carotenoid-Free) Rice Endosperm". Science. 287 (5451): 303–305. Bibcode:2000Sci...287..303Y. ISSN 0036-8075. PMID 10634784. doi:10.1126/science.287.5451.303.
- Goeddel, D. V.; Kleid, D. G.; Bolivar, F.; Heyneker, H. L.; Yansura, D. G.; Crea, R.; Hirose, T.; Kraszewski, A.; Itakura, K.; Riggs, A. D. (1979). "Expression in Escherichia coli of chemically synthesized genes for human insulin". Proceedings of the National Academy of Sciences. 76 (1): 106–10. Bibcode:1979PNAS...76..106G. PMC . PMID 85300. doi:10.1073/pnas.76.1.106.
- "Artificial Genes". TIME. 15 November 1982. Retrieved 17 July 2010.
- Woodard, S. L.; Woodard, J. A.; Howard, M. E. (2004). "Plant molecular farming: Systems and products". Plant Cell Reports. 22 (10): 711–720. PMID 14997337. doi:10.1007/s00299-004-0767-1.
- BBC News 14 June 2002 GM crops: A bitter harvest?
- Thomas H. Maugh II for the Los Angeles Times. June 09, 1987. Altered Bacterium Does Its Job : Frost Failed to Damage Sprayed Test Crop, Company Says
- Fraley, RT; et al. (1983). "Expression of bacterial genes in plant cells" (PDF). Proc. Natl. Acad. Sci. USA. 80 (15): 4803–4807. Bibcode:1983PNAS...80.4803F. PMC . PMID 6308651. doi:10.1073/pnas.80.15.4803.
- James, Clive (1997). "Global Status of Transgenic Crops in 1997" (PDF). ISAAA Briefs No. 5.: 31.
- Bruening, G.; Lyons, J. M. (2000). "The case of the FLAVR SAVR tomato". California Agriculture. 54 (4): 6–7. doi:10.3733/ca.v054n04p6.
- Debora MacKenzie (18 June 1994). "Transgenic tobacco is European first". New Scientist.
- Genetically Altered Potato Ok'd For Crops Lawrence Journal-World - 6 May 1995
- James, Clive (1996). "Global Review of the Field Testing and Commercialization of Transgenic Plants: 1986 to 1995" (PDF). The International Service for the Acquisition of Agri-biotech Applications. Retrieved 17 July 2010.
- Gibson, D. G.; Glass, J. I.; Lartigue, C.; Noskov, V. N.; Chuang, R.-Y.; Algire, M. A.; Benders, G. A.; Montague, M. G.; Ma, L.; Moodie, M. M.; Merryman, C.; Vashee, S.; Krishnakumar, R.; Assad-Garcia, N.; Andrews-Pfannkoch, C.; Denisova, E. A.; Young, L.; Qi, Z.-Q.; Segall-Shapiro, T. H.; Calvey, C. H.; Parmar, P. P.; Hutchison Ca, C. A.; Smith, H. O.; Venter, J. C. (2010). "Creation of a Bacterial Cell Controlled by a Chemically Synthesized Genome". Science. 329 (5987): 52–6. Bibcode:2010Sci...329...52G. PMID 20488990. doi:10.1126/science.1190719.
- Sample, Ian (20 May 2010). "Craig Venter creates synthetic life form". London: guardian.co.uk.
- Vàzquez-Salat, Núria; Salter, Brian; Smets, Greet; Houdebine, Louis-Marie (2012-11-01). "The current state of GMO governance: Are we ready for GM animals?". Biotechnology Advances. Special issue on ACB 2011. 30 (6): 1336–1343. PMID 22361646. doi:10.1016/j.biotechadv.2012.02.006.
- "AQUABOUNTY CLEARED TO SELL SALMON IN USA FOR COMMERCIAL PURPOSES".
- Bodnar, Anastasia (October 2010). "Risk Assessment and Mitigation of AquAdvantage Salmon" (PDF). ISB News Report.
- Whipple, Tom (2016-05-14). "Trailblazing GM vaccines ‘are held back by red tape’". www.thetimes.co.uk. The Times. Retrieved 2016-05-14.
- Thomas, Michael A.; Roemer, Gary W.; Donlan, C. Josh; Dickson, Brett G.; Matocq, Marjorie; Malaney, Jason (2013-09-26). "Ecology: Gene tweaking for conservation". Nature. 501 (7468): 485–486. PMID 24073449. doi:10.1038/501485a.
- Melo, Eduardo O.; Canavessi, Aurea M. O.; Franco, Mauricio M.; Rumpf, Rodolpho (2007). "Animal transgenesis: state of the art and applications" (PDF). J. Appl. Genet. 48 (1): 47–61. PMID 17272861. doi:10.1007/BF03194657. Archived from the original (PDF) on 27 September 2009.
- Arpino, JA; et al. (Jul 2013). "Tuning the dials of Synthetic Biology". Microbiology. 159 (7): 1236–53. PMC . PMID 23704788. doi:10.1099/mic.0.067975-0.
- Pollack, Andrew (7 May 2014). "Researchers Report Breakthrough in Creating Artificial Genetic Code". New York Times. Retrieved 7 May 2014.
- Malyshev, Denis A.; Dhami, Kirandeep; Lavergne, Thomas; Chen, Tingjian; Dai, Nan; Foster, Jeremy M.; Corrêa, Ivan R.; Romesberg, Floyd E. (7 May 2014). "A semi-synthetic organism with an expanded genetic alphabet". Nature. 509 (7500): 385–388. Bibcode:2014Natur.509..385M. PMC . PMID 24805238. doi:10.1038/nature13314. Retrieved 7 May 2014.
- Leader, Benjamin; Baca, Qentin J.; Golan, David E. (January 2008). "Protein therapeutics: a summary and pharmacological classification". Nature Reviews Drug Discovery. A guide to drug discovery. 7 (1): 21–39. PMID 18097458. doi:10.1038/nrd2399.
Leader 2008 — Fee required for access to full text.
- Walsh, Gary (April 2005). "Therapeutic insulins and their large-scale manufacture". Appl. Microbiol. Biotechnol. 67 (2): 151–159. PMID 15580495. doi:10.1007/s00253-004-1809-x.
Walsh 2005 — Fee required for access to full text.
- Summers, Rebecca (24 April 2013) "Bacteria churn out first ever petrol-like biofuel" New Scientist, Retrieved 27 April 2013
- Pipe, Steven W. (May 2008). "Recombinant clotting factors". Thromb. Haemost. 99 (5): 840–850. PMID 18449413. doi:10.1160/TH07-10-0593.
- Bryant, Jackie; Baxter, Louise; Cave, Carolyn B.; Milne, Ruairidh; Bryant, Jackie (2007). Bryant, Jackie, ed. "Recombinant growth hormone for idiopathic short stature in children and adolescents". Cochrane Database Syst Rev (3): CD004440. PMID 17636758. doi:10.1002/14651858.CD004440.pub2.
Bryant 2007 — Fee required for access to full text.
- Baxter L, Bryant J, Cave CB, Milne R (2007). Bryant, Jackie, ed. "Recombinant growth hormone for children and adolescents with Turner syndrome". Cochrane Database Syst Rev (1): CD003887. PMID 17253498. doi:10.1002/14651858.CD003887.pub2.
- Molteni, Megan (2017-04-12). "Florida’s Orange Trees Are Dying, But a Weaponized Virus Could Save Them". WIRED. Retrieved 2017-04-17.
- Panesar, Pamit et al. (2010) Enzymes in Food Processing: Fundamentals and Potential Applications, Chapter 10, I K International Publishing House, ISBN 978-93-80026-33-6
- Jefferson R. A. Kavanagh T. A. Bevan M. W. (1987). "GUS fusions: beta-glucuronidase as a sensitive and versatile gene fusion marker in higher plants". The EMBO Journal. 6 (13): 3901–3907. ISSN 0261-4189. PMC . PMID 3327686.
- Nosowitz, Dan (15 September 2011) "Suntory Creates Mythical Blue (Or, Um, Lavender-ish) Rose" Popular Science, Retrieved 30 August 2012
- Phys.Org website. 4 April 2005 Plant gene replacement results in the world's only blue rose
- Kyodo (11 September 2011 "Suntory to sell blue roses overseas" The Japan Times, Retrieved 30 August 2012
- "World's First 'Blue' Rose Soon Available in U.S.". WIRED. 14 September 2011.
- Gasdaska JR et al. (2003) "Advantages of Therapeutic Protein Production in the Aquatic Plant Lemna". BioProcessing Journal Mar/Apr 2003 pp 49–56 [dead link]
- (10 December 2012) "Engineering algae to make complex anti-cancer 'designer' drug" PhysOrg, Retrieved 15 April 2013
- Büttner-Mainik, A.; et al. (2011). "Production of biologically active recombinant human factor H in Physcomitrella". Plant Biotechnology Journal. 9: 373–383. PMID 20723134. doi:10.1111/j.1467-7652.2010.00552.x.
- Baur, A.; Reski, R.; Gorr, G. (2005). "Enhanced recovery of a secreted recombinant human growth factor using stabilizing additives and by co-expression of human serum albumin in the moss Physcomitrella patens". Plant Biotech. J. 3 (3): 331–340. PMID 17129315. doi:10.1111/j.1467-7652.2005.00127.x.
- Protalix technology platform Archived 27 October 2012 at the Wayback Machine.
- Gali Weinreb and Koby Yeshayahou for Globes 2 May 2012. "FDA approves Protalix Gaucher treatment Archived 29 May 2013 at the Wayback Machine."
- ISAAA 2013 Annual Report Executive Summary, Global Status of Commercialized Biotech/GM Crops: 2013 ISAAA Brief 46-2013, Retrieved 6 August 2014
- James, C (2011). "ISAAA Brief 43, Global Status of Commercialized Biotech/GM Crops: 2011". ISAAA Briefs. Ithaca, New York: International Service for the Acquisition of Agri-biotech Applications (ISAAA). Retrieved 2012-06-02.
- Jorge Fernandez-Cornejo; Seth James Wechsler. "USDA ERS - Adoption of Genetically Engineered Crops in the U.S.". usda.gov.
- Nielsen, K. M. (2003). "Transgenic organisms—time for conceptual diversification?". Nature Biotechnology. 21 (3): 227–228. PMID 12610561. doi:10.1038/nbt0303-227.
- Schouten, H.; Krens, F.; Jacobsen, E. (2006). "Cisgenic plants are similar to traditionally bred plants: international regulations for genetically modified organisms should be altered to exempt cisgenesis". EMBO Reports. 7 (8): 750–753. PMC . PMID 16880817. doi:10.1038/sj.embor.7400769.
- Prins, T. W. and Kok, E. J. (2010) Food and feed safety aspects of cisgenic crop plant varieties Report 2010.001, Project number: 120.72.667.01, RIKILT – Institute of Food Safety, Netherlands. Retrieved 6 September 2010.
- Adams, Jonathan M.; Piovesan, Gianluca; Strauss, Steve; Brown, Sandra (2002-08-01). "The Case for Genetic Engineering of Native and Landscape Trees against Introduced Pests and Diseases". Conservation Biology. 16 (4): 874–879. ISSN 1523-1739. doi:10.1046/j.1523-1739.2002.00523.x.
- Tripathi, Savarni (2007). "Development of Genetically Engineered Resistant Papaya for papaya ringspot virus in a Timely Manner". Methods in Molecular Biology.
- EFSA (2012). Genetically modified animals Europe: EFSA
- Murray, Joo (20). Genetically modified animals. Canada: Brainwaving
- Jaenisch, R.; Mintz, B. (1974). "Simian virus 40 DNA sequences in DNA of healthy adult mice derived from preimplantation blastocysts injected with viral DNA.". Proc. Natl. Acad. Sci. 71 (4): 1250–1254. Bibcode:1974PNAS...71.1250J. PMC . PMID 4364530. doi:10.1073/pnas.71.4.1250.
- Rudinko, Larisa (20). Guidance for industry. USA: Center for veterinary medicine Link.
- Sathasivam K, Hobbs C, Mangiarini L, et al. (June 1999). "Transgenic models of Huntington's disease". Philosophical Transactions of the Royal Society B. 354 (1386): 963–9. PMC . PMID 10434294. doi:10.1098/rstb.1999.0447.[permanent dead link]
- Spencer, L; Humphries, J; Brantly, M. (12 May 2005). "Antibody Response to Aerosolized Transgenic Human Alpha1-Antitrypsin". New England Journal of Medicine. 352 (19): 19. PMID 15888711. doi:10.1056/nejm200505123521923. Retrieved 28 April 2011.
- "Green fluorescent protein takes Nobel prize". Lewis Brindley. Retrieved 2015-05-31.
- Randall S. et al. (2008) "Genetically Modified Pigs for Medicine and Agriculture Archived 26 March 2014 at the Wayback Machine." Biotechnology and Genetic Engineering Reviews – Vol. 25, 245–266, Retrieved 31 August 2012
- Wongsrikeao P, Saenz D, Rinkoski T, Otoi T, Poeschla E (2011). "Antiviral restriction factor transgenesis in the domestic cat". Nature Methods. 8 (10): 853–9. PMC . PMID 21909101. doi:10.1038/nmeth.1703.
- Staff (3 April 2012) Biology of HIV Archived 11 April 2014 at the Wayback Machine. National Institute of Allergy and Infectious Diseases, Retrieved 31 August 2012.
- Sasaki, E.; Suemizu, H.; Shimada, A.; Hanazawa, K.; Oiwa, R.; Kamioka, M.; Tomioka, I.; Sotomaru, Y.; Hirakawa, R.; Eto, T.; Shiozawa, S.; Maeda, T.; Ito, M.; Ito, R.; Kito, C.; Yagihashi, C.; Kawai, K.; Miyoshi, H.; Tanioka, Y.; Tamaoki, N.; Habu, S.; Okano, H.; Nomura, T. (2009). "Generation of transgenic non-human primates with germline transmission". Nature. 459 (7246): 523–527. Bibcode:2009Natur.459..523S. PMID 19478777. doi:10.1038/nature08090.
- Schatten, G.; Mitalipov, S. (2009). "Developmental biology: Transgenic primate offspring". Nature. 459 (7246): 515–516. Bibcode:2009Natur.459..515S. PMC . PMID 19478771. doi:10.1038/459515a.
- Cyranoski, D. (2009). "Marmoset model takes centre stage". Nature. 459 (7246): 492–492. PMID 19478751. doi:10.1038/459492a.
- Houdebine, Louis-Marie (2009). "Production of Pharmaceutical by transgenic animals" (PDF). Comparative Immunology, Microbiology & Infectious Diseases. 32 (2): 107–121. PMID 18243312. doi:10.1016/j.cimid.2007.11.005.
- Britt Erickson, 10 February 2009, for Chemical & Engineering News. FDA Approves Drug From Transgenic Goat Milk Accessed 6 October 2012
- Lai L, et al. (2006). "Generation of cloned transgenic pigs rich in omega-3 fatty acids" (PDF). Nature Biotechnology. 24 (4): 435–436. PMC . PMID 16565727. doi:10.1038/nbt1198. Archived from the original (PDF) on 16 August 2009. Retrieved 2009-03-29.
- Guelph(2010). Enviropig. Canada:
- Schimdt, Sarah. "Genetically engineered pigs killed after funding ends", Postmedia News, 22 June 2012. Accessed 31 July 2012.
- Canada. "Enviropig — Environmental Benefits | University of Guelph". Uoguelph.ca. Retrieved 8 March 2010.
- Gray,Richard(2011). "Genetically modified cows produce 'human' milk"
- Classical Medicine Journal (14 April 2010). "Genetically modified cows producing human milk.". Archived from the original on 6 November 2014.
- Yapp, Robin (11 June 2011). "Scientists create cow that produces 'human' milk". The Daily Telegraph. London. Retrieved 15 June 2012.
- Jabed, A.; Wagner, S.; McCracken, J.; Wells, D. N.; Laible, G. (2012). "Targeted microRNA expression in dairy cattle directs production of -lactoglobulin-free, high-casein milk". Proceedings of the National Academy of Sciences. 109 (42): 16811–16816. Bibcode:2012PNAS..10916811J. doi:10.1073/pnas.1210057109.
- Zyga, Lisa(2010). "Scientist bred goats that produce spider silk Archived 30 April 2015 at the Wayback Machine.".
- Selkirk SM (October 2004). "Gene therapy in clinical medicine". Postgrad Med J. 80 (948): 560–70. PMC . PMID 15466989. doi:10.1136/pgmj.2003.017764.
- Cavazzana-Calvo M, Fischer A (June 2007). "Gene therapy for severe combined immunodeficiency: are we there yet?". J. Clin. Invest. 117 (6): 1456–65. PMC . PMID 17549248. doi:10.1172/JCI30953.
- Richards, Sabrina (6 November 2012) "Gene therapy arrives in Europe" The Scientist, Retrieved 15 April 2013
- Rosenecker J, Huth S, Rudolph C (October 2006). "Gene therapy for cystic fibrosis lung disease: current status and future perspectives". Current Opinion in Molecular Therapeutics. 8 (5): 439–45. PMID 17078386.
- Persons DA, Nienhuis AW (July 2003). "Gene therapy for the hemoglobin disorders". Curr. Hematol. Rep. 2 (4): 348–55. PMID 12901333.
- Lewitt, P. A.; Rezai, A. R.; Leehey, M. A.; Ojemann, S. G.; Flaherty, A. W.; Eskandar, E. N.; Kostyk, S. K.; Thomas, K.; Sarkar, A.; Siddiqui, M. S.; Tatter, S. B.; Schwalb, J. M.; Poston, K. L.; Henderson, J. M.; Kurlan, R. M.; Richard, I. H.; Van Meter, L.; Sapan, C. V.; During, M. J.; Kaplitt, M. G.; Feigin, A. (2011). "AAV2-GAD gene therapy for advanced Parkinson's disease: A double-blind, sham-surgery controlled, randomised trial". The Lancet Neurology. 10 (4): 309–319. PMID 21419704. doi:10.1016/S1474-4422(11)70039-4.
- Gallaher, James "Gene therapy 'treats' Parkinson's disease" BBC News Health, 17 March 2011. Retrieved 24 April 2011
- Urbina, Zachary (12 February 2013) "Genetically Engineered Virus Fights Liver Cancer" United Academics, Retrieved 15 February 2013
- "Treatment for Leukemia Is Showing Early Promise". The New York Times. Associated Press. 11 August 2011. p. A15. Retrieved 21 January 2013.
- Coghlan, Andy (26 March 2013) "Gene therapy cures leukaemia in eight days" The New Scientist, Retrieved 15 April 2013
- Staff (13 February 2013) "Gene therapy cures diabetic dogs" New Scientist, Retrieved 15 February 2013
- (30 April 2013) "New gene therapy trial gives hope to people with heart failure" British Heart Foundation, Retrieved 5 May 2013
- Foster K, Foster H, Dickson JG (December 2006). "Gene therapy progress and prospects: Duchenne muscular dystrophy". Gene Ther. 13 (24): 1677–85. PMID 17066097. doi:10.1038/sj.gt.3302877.
- Moreno, Sacramento (18 December 2007). "Long-term decline of the European wild rabbit (Oryctolagus cuniculus) in south-western Spain". Wildlife Research.
- Angulo, E.; Cooke, B. (2002-12-01). "First synthesize new viruses then regulate their release? The case of the wild rabbit". Molecular Ecology. 11 (12): 2703–2709. ISSN 1365-294X. PMID 12453252. doi:10.1046/j.1365-294X.2002.01635.x.
- Hackett, P. B., Ekker, S. E. and Essner, J. J. (2004) Applications of transposable elements in fish for transgenesis and functional genomics. Fish Development and Genetics (Z. Gong and V. Korzh, eds.) World Scientific, Inc., Chapter 16, 532–580.
- Published PCT Application WO2000049150 "Chimeric Gene Constructs for Generation of Fluorescent Transgenic Ornamental Fish". National University of Singapore 
- Eric Hallerman "Glofish, The First GM Animal Commercialized: Profits amid Controversy". June, 2004. Accessed 3 September 2012.
- Schuchat, S. (17 December 2003). "Why GloFish won't glow in California". San Francisco Chronicle.
- Jun Du, Shao; et al. (1992). "Growth Enhancement in Transgenic Atlantic Salmon by the Use of an 'All Fish' Chimeric Growth Hormone Gene Construct". Nature Biotechnology. 10: 176–181. doi:10.1038/nbt0292-176.
- Devlin, RF; et al. (2001). "Growth of domesticated transgenic fish". Nature. 409: 781–782. PMID 11236982. doi:10.1038/35057314.
- Rahman, MA; et al. (2001). "Growth and nutritional trials on transgenic Nile tilapia containing an exogenous fish growth hormone gene". Journal of Fish Biology. 59 (1): 62–78. doi:10.1111/j.1095-8649.2001.tb02338.x.
- Pollack, Andrew (December 21, 2012). "Engineered Fish Moves a Step Closer to Approval". The New York Times.
- http://www.fda.gov/ForConsumers/ConsumerUpdates/ucm472487.htm. Missing or empty
- Goldenberg, Suzanne (25 November 2013). "Canada approves production of GM salmon eggs on commercial scale". The Guardian. Retrieved 26 November 2013.
- National University of Singapore Enterprise webpage Archived 9 May 2014 at the Wayback Machine.
- "Zebra Fish as Pollution Indicators" Page last modified on 31 July 2001. Accessed October 2012
- Carvan, MJ; et al. (2000). "Transgenic zebrafish as sentinels for aquatic pollution". Ann N Y Acad Sci. 919: 133–47. Bibcode:2000NYASA.919..133C. PMID 11083105. doi:10.1111/j.1749-6632.2000.tb06875.x.
- Nebert, DW; et al. (2002). "Use of Reporter Genes and Vertebrate DNA Motifs in Transgenic Zebrafish as Sentinels for Assessing Aquatic Pollution". Environmental Health Perspectives. 110 (1): A15. PMC . PMID 11813700. doi:10.1289/ehp.110-a15.
- Mattingly, CJ; et al. (Aug 2001). "Green fluorescent protein (GFP) as a marker of aryl hydrocarbon receptor (AhR) function in developing zebrafish (Danio rerio)". Environ Health Perspect. 109 (8): 845–9. PMC . PMID 11564622. doi:10.1289/ehp.01109845.
- Huntingford, F.A., Adams, C., Braithwaite, V.A., Kadri, S., Pottinger, T.G., Sandøe, P. and Turnbull, J.F. (2006). "Review paper: Current issues in fish welfare" (PDF). Journal of Fish Biology. 68 (2): 332–372. doi:10.1111/j.0022-1112.2006.001046.x.
- Fini, Jean-Baptiste; Le Mevel, Sebastien; Turque, Nathalie; Palmier, Karima; Zalko, Daniel; Cravedi, Jean-Pierre; Demeneix, Barbara A. (2007-08-15). "An in vivo multiwell-based fluorescent screen for monitoring vertebrate thyroid hormone disruption". Environmental Science & Technology. 41 (16): 5908–5914. Bibcode:2007EnST...41.5908F. ISSN 0013-936X. PMID 17874805. doi:10.1021/es0704129.
- "Online Education Kit: 1981-82: First Transgenic Mice and Fruit Flies". genome.gov.
- Gallagher, James "GM mosquitoes offer malaria hope" BBC News, Health, 20 April 2011. Retrieved 22 April 2011
- Corby-Harris, V.; Drexler, A.; Watkins De Jong, L.; Antonova, Y.; Pakpour, N.; Ziegler, R.; Ramberg, F.; Lewis, E. E.; Brown, J. M.; Luckhart, S.; Riehle, M. A. (2010). Vernick, Kenneth D., ed. "Activation of Akt Signaling Reduces the Prevalence and Intensity of Malaria Parasite Infection and Lifespan in Anopheles stephensi Mosquitoes". PLoS Pathogens. 6 (7): e1001003. PMC . PMID 20664791. doi:10.1371/journal.ppat.1001003.
- Windbichler, N.; Menichelli, M.; Papathanos, P. A.; Thyme, S. B.; Li, H.; Ulge, U. Y.; Hovde, B. T.; Baker, D.; Monnat Jr, R. J.; Burt, A.; Crisanti, A. (2011). "A synthetic homing endonuclease-based gene drive system in the human malaria mosquito". Nature. 473 (7346): 212–215. Bibcode:2011Natur.473..212W. PMC . PMID 21508956. doi:10.1038/nature09937.
- World Health Organization, Malaria, Key Facts Retrieved 22 April 2011
- Wise De Valdez, M. R.; Nimmo, D.; Betz, J.; Gong, H. -F.; James, A. A.; Alphey, L.; Black, W. C. (2011). "Genetic elimination of dengue vector mosquitoes". Proceedings of the National Academy of Sciences. 108 (12): 4772–4775. Bibcode:2011PNAS..108.4772W. doi:10.1073/pnas.1019295108.
- Knapton, Sarah (6 February 2016). "Releasing millions of GM mosquitoes ‘could solve zika crisis’". The Telegraph. Retrieved 14 March 2016.
- Harris, A. F.; Nimmo, D.; McKemey, A. R.; Kelly, N.; Scaife, S.; Donnelly, C. A.; Beech, C.; Petrie, W. D.; Alphey, L. (2011). "Field performance of engineered male mosquitoes". Nature Biotechnology. 29 (11): 1034–1037. PMID 22037376. doi:10.1038/nbt.2019.
- Staff (March 2011) "Cayman demonstrates RIDL potential" Oxitec Newsletter, March 2011. Retrieved 20 September 2011
- Page, Michael Le. "GM mosquito trial in Florida given the go-ahead by regulator". Retrieved 2016-08-09.
- Adler. "A World Without Mosquitoes". Smithsonian. 47 (3).
- Nicholls, Henry (14 September 2011) "Swarm troopers: Mutant armies waging war in the wild" The New Scientist. Retrieved 20 September 2011
- Staff Pink Bollworm Oxitec, Retrieved 17 August 2014
- Walters, M.; et al. (2012). "Field longevity of a fluorescent protein marker in an engineered strain of the pink bollworm, Pectinophora gossypiella (Saunders)". PLoS ONE. 7 (6): e38547. Bibcode:2012PLoSO...738547W. PMC . PMID 22693645. doi:10.1371/journal.pone.0038547.
- Wittlieb J, Khalturin K, Lohmann JU, Anton-Erxleben F, Bosch TC (2006). "Transgenic Hydra allow in vivo tracking of individual stem cells during morphogenesis". Proc. Natl. Acad. Sci. U.S.A. 103 (16): 6208–6211. Bibcode:2006PNAS..103.6208W. PMC . PMID 16556723. doi:10.1073/pnas.0510163103.
- Gaskell, G.; Bauer, M. W.; Durant, J.; Allum, N. C. (1999). "Worlds Apart? The Reception of Genetically Modified Foods in Europe and the U.S". Science. 285 (5426): 384–387. PMID 10411496. doi:10.1126/science.285.5426.384.
- "The History and Future of GM Potatoes". PotatoPro.com.
- Purnhagen, Wesseler (2016): "The "Honey" Judgment of Bablok and Others Versus Freistaat Bayern in the Court of Justice of the European Union: Implications for Co-existence". In N. Kalaitzandonakes et al. (eds.), The Coexistence of Genetically Modified, Organic and Conventional Foods., pp. 149–165 – 150-153. New York: Springer Science
- Wesseler, J. and N. Kalaitzandonakes (2011): "Present and Future EU GMO policy". In Arie Oskam, Gerrit Meesters and Huib Silvis (eds.), EU Policy for Agriculture, Food and Rural Areas. Second Edition, pp. 23–323 – 23-332. Wageningen: Wageningen Academic Publishers
- Purnhagen, Wesseler (2016): Social, Economic and Legal Avenues". In N. Kalaitzandonakes et al. (eds.), The Coexistence of Genetically Modified, Organic and Conventional Foods., pp. 71–85. New York: Springer Science
- Beckmann, V., C. Soregaroli, J. Wesseler (2011): "Coexistence of genetically modified (GM) and non-modified (non GM) crops: Are the two main property rights regimes equivalent with respect to the coexistence value?" In Genetically modified food and global welfare edited by Colin Carter, GianCarlo Moschini and Ian Sheldon, pp 201–224. Volume 10 in Frontiers of Economics and Globalization Series. Bingley, UK: Emerald Group Publishing
- Smithonian (2015). "Some Brands Are Labeling Products "GMO-free" Even if They Don’t Have Genes".
- Nicolia, Alessandro; Manzo, Alberto; Veronesi, Fabio; Rosellini, Daniele (2013). "An overview of the last 10 years of genetically engineered crop safety research" (PDF). Critical Reviews in Biotechnology. 34: 1–12. PMID 24041244. doi:10.3109/07388551.2013.823595.
We have reviewed the scientific literature on GE crop safety for the last 10 years that catches the scientific consensus matured since GE plants became widely cultivated worldwide, and we can conclude that the scientific research conducted so far has not detected any significant hazard directly connected with the use of GM crops.
The literature about Biodiversity and the GE food/feed consumption has sometimes resulted in animated debate regarding the suitability of the experimental designs, the choice of the statistical methods or the public accessibility of data. Such debate, even if positive and part of the natural process of review by the scientific community, has frequently been distorted by the media and often used politically and inappropriately in anti-GE crops campaigns.
- "State of Food and Agriculture 2003–2004. Agricultural Biotechnology: Meeting the Needs of the Poor. Health and environmental impacts of transgenic crops". Food and Agriculture Organization of the United Nations. Retrieved February 8, 2016.
Currently available transgenic crops and foods derived from them have been judged safe to eat and the methods used to test their safety have been deemed appropriate. These conclusions represent the consensus of the scientific evidence surveyed by the ICSU (2003) and they are consistent with the views of the World Health Organization (WHO, 2002). These foods have been assessed for increased risks to human health by several national regulatory authorities (inter alia, Argentina, Brazil, Canada, China, the United Kingdom and the United States) using their national food safety procedures (ICSU). To date no verifiable untoward toxic or nutritionally deleterious effects resulting from the consumption of foods derived from genetically modified crops have been discovered anywhere in the world (GM Science Review Panel). Many millions of people have consumed foods derived from GM plants - mainly maize, soybean and oilseed rape - without any observed adverse effects (ICSU).
- Ronald, Pamela (May 5, 2011). "Plant Genetics, Sustainable Agriculture and Global Food Security". Genetics. 188: 11–20. PMC . PMID 21546547. doi:10.1534/genetics.111.128553.
There is broad scientific consensus that genetically engineered crops currently on the market are safe to eat. After 14 years of cultivation and a cumulative total of 2 billion acres planted, no adverse health or environmental effects have resulted from commercialization of genetically engineered crops (Board on Agriculture and Natural Resources, Committee on Environmental Impacts Associated with Commercialization of Transgenic Plants, National Research Council and Division on Earth and Life Studies 2002). Both the U.S. National Research Council and the Joint Research Centre (the European Union's scientific and technical research laboratory and an integral part of the European Commission) have concluded that there is a comprehensive body of knowledge that adequately addresses the food safety issue of genetically engineered crops (Committee on Identifying and Assessing Unintended Effects of Genetically Engineered Foods on Human Health and National Research Council 2004; European Commission Joint Research Centre 2008). These and other recent reports conclude that the processes of genetic engineering and conventional breeding are no different in terms of unintended consequences to human health and the environment (European Commission Directorate-General for Research and Innovation 2010).
- 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: 734–742. PMID 21296423. doi:10.1016/j.envint.2011.01.003.
In spite of this, the number of studies specifically focused on safety assessment of GM plants is still limited. However, it is important to remark that for the first time, a certain equilibrium in the number of research groups suggesting, on the basis of their studies, that a number of varieties of GM products (mainly maize and soybeans) are as safe and nutritious as the respective conventional non-GM plant, and those raising still serious concerns, was observed. Moreover, it is worth mentioning that most of the studies demonstrating that GM foods are as nutritional and safe as those obtained by conventional breeding, have been performed by biotechnology companies or associates, which are also responsible of commercializing these GM plants. Anyhow, this represents a notable advance in comparison with the lack of studies published in recent years in scientific journals by those companies.
Krimsky, Sheldon (2015). "An Illusory Consensus behind GMO Health Assessment" (PDF). Science, Technology, & Human Values. 40: 1–32. doi:10.1177/0162243915598381.
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.
Panchin, Alexander Y.; Tuzhikov, Alexander I. (January 14, 2016). "Published GMO studies find no evidence of harm when corrected for multiple comparisons". Critical Reviews in Biotechnology: 1–5. ISSN 0738-8551. PMID 26767435. doi:10.3109/07388551.2015.1130684.
Here, we show that a number of articles some of which have strongly and negatively influenced the public opinion on GM crops and even provoked political actions, such as GMO embargo, share common flaws in the statistical evaluation of the data. Having accounted for these flaws, we conclude that the data presented in these articles does not provide any substantial evidence of GMO harm.
The presented articles suggesting possible harm of GMOs received high public attention. However, despite their claims, they actually weaken the evidence for the harm and lack of substantial equivalency of studied GMOs. We emphasize that with over 1783 published articles on GMOs over the last 10 years it is expected that some of them should have reported undesired differences between GMOs and conventional crops even if no such differences exist in reality.
Yang, Y.T.; Chen, B. (2016). "Governing GMOs in the USA: science, law and public health". Journal of the Science of Food and Agriculture. 96: 1851–1855. PMID 26536836. doi:10.1002/jsfa.7523.
It is therefore not surprising that efforts to require labeling and to ban GMOs have been a growing political issue in the USA (citing Domingo and Bordonaba, 2011).
Overall, a broad scientific consensus holds that currently marketed GM food poses no greater risk than conventional food... Major national and international science and medical associations have stated that no adverse human health effects related to GMO food have been reported or substantiated in peer-reviewed literature to date.
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.
- "Statement by the AAAS Board of Directors On Labeling of Genetically Modified Foods" (PDF). American Association for the Advancement of Science. October 20, 2012. Retrieved February 8, 2016.
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 more risky 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.
Pinholster, Ginger (October 25, 2012). "AAAS Board of Directors: Legally Mandating GM Food Labels Could "Mislead and Falsely Alarm Consumers"". American Association for the Advancement of Science. Retrieved February 8, 2016.
- "A decade of EU-funded GMO research (2001–2010)" (PDF). Directorate-General for Research and Innovation. Biotechnologies, Agriculture, Food. European Commission, European Union. 2010. ISBN 978-92-79-16344-9. doi:10.2777/97784. Retrieved February 8, 2016.
- "AMA Report on Genetically Modified Crops and Foods (online summary)". American Medical Association. January 2001. Retrieved March 19, 2016.
A report issued by the scientific council of the American Medical Association (AMA) says that no long-term health effects have been detected from the use of transgenic crops and genetically modified foods, and that these foods are substantially equivalent to their conventional counterparts. (from online summary prepared by ISAAA)" "Crops and foods produced using recombinant DNA techniques have been available for fewer than 10 years and no long-term effects have been detected to date. These foods are substantially equivalent to their conventional counterparts. (from original report by AMA: )
"REPORT 2 OF THE COUNCIL ON SCIENCE AND PUBLIC HEALTH (A-12): Labeling of Bioengineered Foods" (PDF). American Medical Association. 2012. Archived from the original on 7 September 2012. Retrieved March 19, 2016.
Bioengineered foods have been consumed for close to 20 years, and during that time, no overt consequences on human health have been reported and/or substantiated in the peer-reviewed literature.
- "Restrictions on Genetically Modified Organisms: United States. Public and Scholarly Opinion". Library of Congress. June 9, 2015. Retrieved February 8, 2016.
Several scientific organizations in the US have issued studies or statements regarding the safety of GMOs indicating that there is no evidence that GMOs present unique safety risks compared to conventionally bred products. These include the National Research Council, the American Association for the Advancement of Science, and the American Medical Association. Groups in the US opposed to GMOs include some environmental organizations, organic farming organizations, and consumer organizations. A substantial number of legal academics have criticized the US's approach to regulating GMOs.
- "Genetically Engineered Crops: Experiences and Prospects". The National Academies of Sciences, Engineering, and Medicine (US). 2016. p. 149. Retrieved May 19, 2016.
Overall finding on purported adverse effects on human health of foods derived from GE crops: On the basis of detailed examination of comparisons of currently commercialized GE with non-GE foods in compositional analysis, acute and chronic animal toxicity tests, long-term data on health of livestock fed GE foods, and human epidemiological data, the committee found no differences that implicate a higher risk to human health from GE foods than from their non-GE counterparts.
- "Frequently asked questions on genetically modified foods". World Health Organization. Retrieved February 8, 2016.
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.
GM foods currently available on the international market have passed safety assessments and are not likely to present risks for human health. In addition, no effects on human health have been shown as a result of the consumption of such foods by the general population in the countries where they have been approved. Continuous application of safety assessments based on the Codex Alimentarius principles and, where appropriate, adequate post market monitoring, should form the basis for ensuring the safety of GM foods.
- Haslberger, Alexander G. (2003). "Codex guidelines for GM foods include the analysis of unintended effects". Nature Biotechnology. 21: 739–741. PMID 12833088. doi:10.1038/nbt0703-739.
These principles dictate a case-by-case premarket assessment that includes an evaluation of both direct and unintended effects.
- Some medical organizations, including the British Medical Association, advocate further caution based upon the precautionary principle:
"Genetically modified foods and health: a second interim statement" (PDF). British Medical Association. March 2004. Retrieved March 21, 2016.
In our view, the potential for GM foods to cause harmful health effects is very small and many of the concerns expressed apply with equal vigour to conventionally derived foods. However, safety concerns cannot, as yet, be dismissed completely on the basis of information currently available.
When seeking to optimise the balance between benefits and risks, it is prudent to err on the side of caution and, above all, learn from accumulating knowledge and experience. Any new technology such as genetic modification must be examined for possible benefits and risks to human health and the environment. As with all novel foods, safety assessments in relation to GM foods must be made on a case-by-case basis.
Members of the GM jury project were briefed on various aspects of genetic modification by a diverse group of acknowledged experts in the relevant subjects. The GM jury reached the conclusion that the sale of GM foods currently available should be halted and the moratorium on commercial growth of GM crops should be continued. These conclusions were based on the precautionary principle and lack of evidence of any benefit. The Jury expressed concern over the impact of GM crops on farming, the environment, food safety and other potential health effects.
The Royal Society review (2002) concluded that the risks to human health associated with the use of specific viral DNA sequences in GM plants are negligible, and while calling for caution in the introduction of potential allergens into food crops, stressed the absence of evidence that commercially available GM foods cause clinical allergic manifestations. The BMA shares the view that that there is no robust evidence to prove that GM foods are unsafe but we endorse the call for further research and surveillance to provide convincing evidence of safety and benefit.
- Funk, Cary; Rainie, Lee (January 29, 2015). "Public and Scientists' Views on Science and Society". Pew Research Center. Retrieved February 24, 2016.
The largest differences between the public and the AAAS scientists are found in beliefs about the safety of eating genetically modified (GM) foods. Nearly nine-in-ten (88%) scientists say it is generally safe to eat GM foods compared with 37% of the general public, a difference of 51 percentage points.
- Marris, Claire (2001). "Public views on GMOs: deconstructing the myths" (PDF). EMBO Reports. 2: 545–548. PMC . PMID 11463731. doi:10.1093/embo-reports/kve142.
- Final Report of the PABE research project (December 2001). "Public Perceptions of Agricultural Biotechnologies in Europe". Commission of European Communities. Retrieved February 24, 2016.
- Scott, Sydney E.; Inbar, Yoel; Rozin, Paul (2016). "Evidence for Absolute Moral Opposition to Genetically Modified Food in the United States" (PDF). Perspectives on Psychological Science. 11 (3): 315–324. PMID 27217243. doi:10.1177/1745691615621275.
- "Restrictions on Genetically Modified Organisms". Library of Congress. June 9, 2015. Retrieved February 24, 2016.
- Bashshur, Ramona (February 2013). "FDA and Regulation of GMOs". American Bar Association. Retrieved February 24, 2016.
- Sifferlin, Alexandra (October 3, 2015). "Over Half of E.U. Countries Are Opting Out of GMOs". Time.
- Lynch, Diahanna; Vogel, David (April 5, 2001). "The Regulation of GMOs in Europe and the United States: A Case-Study of Contemporary European Regulatory Politics". Council on Foreign Relations. Retrieved February 24, 2016.
- American Medical Association (2012). "Report 2 of the Council on Science and Public Health: Labeling of Bioengineered Foods" "Bioengineered foods have been consumed for close to 20 years, and during that time, no overt consequences on human health have been reported and/or substantiated in the peer-reviewed literature." (first page)
- United States Institute of Medicine and National Research Council (2004). "Safety of Genetically Engineered Foods: Approaches to Assessing Unintended Health Effects". National Academies Press. Free full-text. National Academies Press. pp R9-10: "In contrast to adverse health effects that have been associated with some traditional food production methods, similar serious health effects have not been identified as a result of genetic engineering techniques used in food production. This may be because developers of bioengineered organisms perform extensive compositional analyses to determine that each phenotype is desirable and to ensure that unintended changes have not occurred in key components of food."
- Key S, Ma JK, Drake PM (June 2008). "Genetically modified plants and human health". J R Soc Med. 101 (6): 290–8. PMC . PMID 18515776. doi:10.1258/jrsm.2008.070372.
pp 292-293. "Foods derived from GM crops have been consumed by hundreds of millions of people across the world for more than 15 years, with no reported ill effects (or legal cases related to human health), despite many of the consumers coming from that most litigious of countries, the USA."
- "Vermont v science", The Economist, Montpelier, 411 (8886), pp. 25–26, 10 May 2014
- Dan Charles, Allison Aubrey (March 27, 2016). "How Little Vermont Got Big Food Companies To Label GMOs". FOOD FOR THOUGHT. the salt. Retrieved 2017-01-11.
- Zdziarski, I.M.; Edwards, J.W.; Carman, J.A.; Haynes, J.I. (December 2014). "GM crops and the rat digestive tract: A critical review". Environment International. 73: 423–433. PMID 25244705. doi:10.1016/j.envint.2014.08.018.
- José L. Domingo. Safety assessment of GM plants: An updated review of the scientific literature
- Nathanael Johnson for Grist. Jul 8, 2013 The genetically modified food debate: Where do we begin?
- JoAnna Wendel for the Genetic Literacy Project. 10 September 2013 Scientists, journalists and farmers join lively GMO forum
- Keith Kloor for Discover Magazine's CollideAScape 22 August 2014 On Double Standards and the Union of Concerned Scientists
- Union of Concerned Scientists. Alternatives to Genetic Engineering. Page source description: "Biotechnology companies produce genetically engineered crops to control insects and weeds and to manufacture pharmaceuticals and other chemicals. The Union of Concerned Scientists works to strengthen the federal oversight needed to prevent such products from contaminating our food supply."
- Emily Marden, Risk and Regulation: U.S. Regulatory Policy on Genetically Modified Food and Agriculture 44 B.C.L. Rev. 733 (2003). Quote: "By the late 1990s, public awareness of GM foods reached a critical level and a number of public interest groups emerged to focus on the issue. One of the early groups to focus on the issue was Mothers for Natural Law ("MFNL"), an Iowa based organization that aimed to ban GM foods from the market.... The Union of Concerned Scientists ("UCS"), an alliance of 50,000 citizens and scientists, has been another prominent voice on the issue.... As the pace of GM products entering the market increased in the 1990s, UCS became a vocal critic of what it saw as the agency’s collusion with industry and failure to fully take account of allergenicity and other safety issues."
- British Medical Association Board of Science and Education (2004). "Genetically modified food and health: A second interim statement". March.
- Public Health Association of Australia (2007) "Genetically Modified Foods" PHAA AGM 2007 Archived 20 January 2014 at the Wayback Machine.
- Canadian Association of Physicians for the Environment (2013) "Statement on Genetically Modified Organisms in the Environment and the Marketplace". October 2013
- Irish Doctors' Environmental Association "IDEA Position on Genetically Modified Foods". Retrieved 3/25/14
- PR Newswire "Genetically Modified Maize: Doctors' Chamber Warns of 'Unpredictable Results' to Humans". 11 November 2013
- Chartered Institute of Environmental Health (2006) "Proposals for managing the coexistence of GM, conventional and organic crops Response to the Department for Environment, Food and Rural Affairs consultation paper". October 2006
- Paull, John (2015) GMOs and organic agriculture: Six lessons from Australia, Agriculture & Forestry, 61(1): 7-14.
- American Medical Association (2012). "Report 2 of the Council on Science and Public Health: Labeling of Bioengineered Foods". "To better detect potential harms of bioengineered foods, the Council believes that pre-market safety assessment should shift from a voluntary notification process to a mandatory requirement." page 7
- Landrigan, Philip J.; Benbrook, Charles (2015). "GMOs, Herbicides, and Public Health". New England Journal of Medicine. New England Journal of Medicine. 373 (8): 693–5. PMID 26287848. doi:10.1056/NEJMp1505660.