Genetically modified organism
A genetically modified organism (GMO), also known as a transgenic organism, is any organism whose genetic material has been altered using genetic engineering techniques. GMOs are the source of medicines and genetically modified foods and are also widely used in scientific research and to produce 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").
- 1 Production
- 2 History
- 3 Uses
- 3.1 Microbes
- 3.2 Plants
- 3.3 Mammals
- 3.4 Fish
- 3.5 Invertebrates
- 4 Regulation
- 5 Controversy
- 6 See also
- 7 References
- 8 External links
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. To do this artificially may require:
- attaching the genes to a virus
- physically inserting the extra DNA into the nucleus of the intended host with a very small syringe
- with the use of electroporation (that is, introducing DNA from one organism into the cell of another by use of an electric pulse)
- with very small particles fired 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 When nucleic acid sequences are combined in a laboratory, the resulting DNA is called recombinant DNA. Recombinant DNA may contain oligonucleotides from the same or similar species, in which case it is called "cisgenic", or may contain oligonucleotides from different organisms that could not naturally interbreed, in which case it is called "transgenic". Recombinant DNA may also contain synthetic sequences.
The first recombinant DNA molecules were produced by Paul Berg in 1972. Genetic engineering, the direct manipulation of genes using biotechnology, was first accomplished by Herbert Boyer and Stanley Cohen in 1973. Whereas selective breeding depends on naturally occurring genetic variation within a population or species, genetic engineering can involve the intentional introduction of genes from different species. Advances have allowed scientists to manipulate, remove, and add genes to a variety of different organisms to induce a range of different traits. From 1976 the technology became commercialized, with companies producing and selling genetically modified foods and medicines.
GMOs are used in biological and medical research, production of pharmaceutical drugs, experimental medicine (e.g. gene therapy), and agriculture (e.g. golden rice, resistance to herbicides). 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 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.
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 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 has applied for regulatory approval to market their GM salmon in the US. As of December 2012 the application was still pending. 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.
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. Aedes aegypti mosquitoes, the single most important carrier of dengue fever, were reduced by 80% in a 2010 trial of these GM mosquitoes in the Cayman Islands. Between 50 and 100 million people are affected by dengue fever every year and 40,000 people die from it.
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.
Cnidarians such as Hydra and the sea anemone Nematostella vectensis have become 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 stably 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 coexistence of GM and nonGM crops. 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, non-governmental 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 which had been labeled as non-GMO grew 30 percent to $1.1 billion.
There is a general scientific agreement that food from genetically modified crops is not inherently riskier to human health than conventional food. 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 those being considered by Vermont's governor, Peter Shumlin, to require labeling of foods containing genetically modified ingredients, could have the unintended consequence of interrupting the benign process of spreading GM technologies to impoverished countries that suffer with food security problems.
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.
- 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
- Reading Scientific Services (detecting GMOs)
- SMART breeding
- Sperm-mediated gene transfer
- Timeline of genetically modified organisms
- Cornell Chronicle, 14 May 1987, page 3. Biologists invent gun for shooting cells with DNA
- 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. doi:10.1038/327070a0.
- Lee LY, Gelvin SB (February 2008). "T-DNA binary vectors and systems". Plant Physiol. 146 (2): 325–332. doi:10.1104/pp.107.113001. OCLC 1642351. PMC 2245830. PMID 18250230.
- Park F (October 2007). "Lentiviral vectors: are they the future of animal transgenesis?". Physiol. Genomics 31 (2): 159–173. doi:10.1152/physiolgenomics.00069.2007. OCLC 37367250. PMID 17684037.
- 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.
- Nature Education, 2014. Definition: recombinant DNA technology / DNA cloning; gene cloning; cloning
- Special Topics in Modern Genetics 5: Genetically modified foods
- 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. doi:10.1073/pnas.69.10.2904. PMC 389671. PMID 4342968.
- 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.
- Melo, Eduardo O.; Canavessi, Aurea M. O.; Franco, Mauricio M.; Rumpf, Rodolpho (2007). "Animal transgenesis: state of the art and applications". J. Appl. Genet. 48 (1): 47–61. doi:10.1007/BF03194657. PMID 17272861. Archived from the original (PDF) on 26 September 2009.
- Arpino, JA; et al. (Jul 2013). "Tuning the dials of Synthetic Biology". Microbiology 159 (7): 1236–53. doi:10.1099/mic.0.067975-0. PMC 3749727. PMID 23704788.
- 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 (journal) 509: 385–388. 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. doi:10.1038/nrd2399. PMID 18097458.
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. doi:10.1007/s00253-004-1809-x. PMID 15580495.
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. doi:10.1160/TH07-10-0593. PMID 18449413.
- 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. doi:10.1002/14651858.CD004440.pub2. PMID 17636758.
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. doi:10.1002/14651858.CD003887.pub2. PMID 17253498.
- 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 553867. 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 
- (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. 
- Baur, A., R. Reski, G. Gorr (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, 331–340 
- Protalix website – technology platform
- Gali Weinreb and Koby Yeshayahou for Globes 2 May 2012. "FDA approves Protalix Gaucher treatment"
- 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. doi:10.1038/nbt0303-227. PMID 12610561.
- 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. doi:10.1038/sj.embor.7400769. PMC 1525145. PMID 16880817.
- 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.
- EFSA (2012). Genetically modified animals Europe: EFSA
- Murray, Joo (20). Genetically modified animals. Canada: Brainwaving
- 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. Sci. 71 (4): 1250–1254. Bibcode:1974PNAS...71.1250J. doi:10.1073/pnas.71.4.1250. PMC 388203. PMID 4364530.
- 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. doi:10.1098/rstb.1999.0447. PMC 1692600. PMID 10434294.
- Spencer, L; Humphries, J; Brantly, M. (12 May 2005). "Antibody Response to Aerosolized Transgenic Human Alpha1-Antitrypsin". New England Journal of Medicine 352: 19. 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" 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. doi:10.1038/nmeth.1703. PMID 21909101.
- Staff (3 April 2012) Biology of HIV 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. doi:10.1038/nature08090. PMID 19478777.
- Schatten, G.; Mitalipov, S. (2009). "Developmental biology: Transgenic primate offspring". Nature 459 (7246): 515–516. Bibcode:2009Natur.459..515S. doi:10.1038/459515a. PMC 2777739. PMID 19478771.
- Cyranoski, D. (2009). "Marmoset model takes centre stage". Nature 459 (7246): 492–492. doi:10.1038/459492a. PMID 19478751.
- Louis-Marie Houdebine (2009) "Production of Pharmaceutical by transgenic animals". Comparative Immunology, Microbiology & Infectious Diseases 32(2): 107–121 
- 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. doi:10.1038/nbt1198. PMC 2976610. PMID 16565727. 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.".
- 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. doi:10.1073/pnas.1210057109.
- Zyga, Lisa(2010). "Scientist bred goats that produce spider silk".
- Selkirk SM (October 2004). "Gene therapy in clinical medicine". Postgrad Med J 80 (948): 560–70. doi:10.1136/pgmj.2003.017764. PMC 1743106. PMID 15466989.
- Cavazzana-Calvo M, Fischer A (June 2007). "Gene therapy for severe combined immunodeficiency: are we there yet?". J. Clin. Invest. 117 (6): 1456–65. doi:10.1172/JCI30953. PMC 1878528. PMID 17549248.
- 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. doi:10.1016/S1474-4422(11)70039-4. PMID 21419704.
- 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. doi:10.1038/sj.gt.3302877. PMID 17066097.
- 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.
- Shao Jun Du 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 
- Devlin RF et al. (2001) "Growth of domesticated transgenic fish". Nature 409, 781–782 
- 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 
- Pollack, Andrew (December 21, 2012). "Engineered Fish Moves a Step Closer to Approval". The New York Times.
- Staff (26 December 2012) Draft Environmental Assessment and Preliminary Finding of No Significant Impact Concerning a Genetically Engineered Atlantic Salmon; Availability Federal Register / Vol. 77, No. 247 / Wednesday, 26 December 2012 / Notices, Retrieved 2 January 2013
- 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 July 5, 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. 2000;919:133–47 
- 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 | January 2002 
- Mattingly CJ et al. (2001) "Green fluorescent protein (GFP) as a marker of aryl hydrocarbon receptor (AhR) function in developing zebrafish (Danio rerio)". Environ Health Perspect. 2001 Aug;109(8):845–9 
- 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.
- "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. doi:10.1371/journal.ppat.1001003. PMC 2904800. PMID 20664791.
- 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. doi:10.1038/nature09937. PMC 3093433. PMID 21508956.
- 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. doi:10.1073/pnas.1019295108.
- 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. doi:10.1038/nbt.2019. PMID 22037376.
- Staff (March 2011) "Cayman demonstrates RIDL potential" Oxitec Newsletter, March 2011. Retrieved 20 September 2011
- 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. doi:10.1371/journal.pone.0038547. PMID 22693645.
- Wittlieb J, Khalturin K, Lohmann JU, Anton-Erxleben F and Bosch TCG (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. doi:10.1073/pnas.0510163103. PMC 1458856. PMID 16556723.
- 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. doi:10.1126/science.285.5426.384. PMID 10411496.
- "The History and Future of GM Potatoes". PotatoPro.com.
- 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
- 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".
- "Vermont v science", The Economist (Montpelier) 411 (8886), 10 May 2014: 25–26
- American Association for the Advancement of Science (AAAS), Board of Directors (2012). Statement by the AAAS Board of Directors On Labeling of Genetically Modified Foods, and associated Press release: Legally Mandating GM Food Labels Could Mislead and Falsely Alarm Consumers
- A decade of EU-funded GMO research (2001–2010) (PDF). Directorate-General for Research and Innovation. Biotechnologies, Agriculture, Food. European Union. 2010. doi:10.2777/97784. ISBN 978-92-79-16344-9.
"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." (p. 16)
- Ronald, Pamela (2011). "Plant genetics, sustainable agriculture and global food security". Genetics 188 (1): 11–20. doi:10.1534/genetics.111.128553. PMC 3120150. PMID 21546547.
- 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. doi:10.1258/jrsm.2008.070372. PMC 2408621. PMID 18515776.
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."
- 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
- 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
- Everything you wanted to know about GM organisms — Provided by New Scientist.
- Transgenic Organism Research
- International Society for Transgenic Technologies (ISTT)
- GMO-Compass: Information on genetically modified organisms
- Co-Extra: Research on co-existence and traceability of GM and non-GM supply chains
- ISAAA Knowledge Center: Information on genetically modified organisms