Genetically modified animal: Difference between revisions
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'''Genetically modified animals''' are animals that have been [[genetically modified]] for a variety of purposes including producing drugs, enhancing yields, increase resistance to disease, etc. |
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The vast majority of genetically modified animals are at the research stage with the number close to entering the market remains small.<ref name=":14">{{cite journal | vauthors = Forabosco F, Löhmus M, Rydhmer L, Sundström LF | title = Genetically modified farm animals and fish in agriculture: A review. | journal = Livestock Science | date = May 2013 | volume = 153 | issue = 1–3 | pages = 1–9 | doi = 10.1016/j.livsci.2013.01.002 }}</ref> |
The vast majority of genetically modified animals are at the research stage with the number close to entering the market remains small.<ref name=":14">{{cite journal | vauthors = Forabosco F, Löhmus M, Rydhmer L, Sundström LF | title = Genetically modified farm animals and fish in agriculture: A review. | journal = Livestock Science | date = May 2013 | volume = 153 | issue = 1–3 | pages = 1–9 | doi = 10.1016/j.livsci.2013.01.002 }}</ref> |
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=== Livestock === |
=== Livestock === |
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Livestock are modified with the intention of improving economically important traits such as growth-rate, quality of meat, milk composition, disease resistance and survival. Animals have been engineered to grow faster, be healthier<ref>{{cite journal | vauthors = Lai L, Kang JX, Li R, Wang J, Witt WT, Yong HY, Hao Y, Wax DM, Murphy CN, Rieke A, Samuel M, Linville ML, Korte SW, Evans RW, Starzl TE, Prather RS, Dai Y | title = Generation of cloned transgenic pigs rich in omega-3 fatty acids | journal = Nature Biotechnology | volume = 24 | issue = 4 | pages = 435–6 | date = April 2006 | pmid = 16565727 | pmc = 2976610 | doi = 10.1038/nbt1198 |
Livestock are modified with the intention of improving economically important traits such as growth-rate, quality of meat, milk composition, disease resistance and survival. Animals have been engineered to grow faster, be healthier<ref>{{cite journal | vauthors = Lai L, Kang JX, Li R, Wang J, Witt WT, Yong HY, Hao Y, Wax DM, Murphy CN, Rieke A, Samuel M, Linville ML, Korte SW, Evans RW, Starzl TE, Prather RS, Dai Y | display-authors = 6 | title = Generation of cloned transgenic pigs rich in omega-3 fatty acids | journal = Nature Biotechnology | volume = 24 | issue = 4 | pages = 435–6 | date = April 2006 | pmid = 16565727 | pmc = 2976610 | doi = 10.1038/nbt1198 }}</ref> and resist diseases.<ref>{{cite news |url= http://www.theguardian.com/environment/2018/jun/24/genetically-engineered-animals-the-five-controversial-science |title=Genetically modified animals|last=Tucker|first=Ian | name-list-format = vanc |date=2018-06-24|work=The Guardian |access-date=2018-12-21 |issn=0261-3077}}</ref> Modifications have also improved the wool production of sheep and udder health of cows.<ref name=":14" /> Goats have been genetically engineered to produce milk with strong spiderweb-like silk proteins in their milk.<ref>{{cite web | last = Zyga | first = Lisa | name-list-format = vanc | date = 2010 | url = http://phys.org/news194539934.html/ | title = Scientist bred goats that produce spider silk | work = Phys.org | archive-url = https://web.archive.org/web/20150430100830/http://phys.org/news194539934.html/ | dead-url = yes | archive-date = 30 April 2015 }}</ref> A GM pig called [[Enviropig]] was created with the capability of digesting plant [[phosphorus]] more efficiently than conventional pigs.<ref name="Guelph">{{cite web | publisher = University of Guelph | location = Canada | date = 2010 | url = http://www.uoguelph.ca/enviropig/index.shtml | title = Enviropig | archive-url = https://web.archive.org/web/20160130104858/http://www.uoguelph.ca/enviropig/index.shtml | archive-date = 30 January 2016}}</ref><ref>{{cite web | last = Schimdt | first = Sarah | name-list-format = vanc | url = http://www.canada.com/technology/science/Genetically+engineered+pigs+killed+after+funding+ends/6819844/story.html | title = Genetically engineered pigs killed after funding ends | work = Postmedia News | date = 22 June 2012 | access-date = 31 July 2012 }}</ref> They could reduce water pollution since they excrete 30 to 70% less phosphorus in manure.<ref name="Guelph" /><ref name="Canada">{{cite web |url= http://www.uoguelph.ca/enviropig/environmental_benefits.shtml |title=Enviropig – Environmental Benefits| publisher = University of Guelph| location = Canada |archive-url = https://web.archive.org/web/20100227041057/http://www.uoguelph.ca/enviropig/environmental_benefits.shtml |archive-date = 27 February 2010 |deadurl=yes |access-date=8 March 2010 }}</ref> [[Dairy cows]] have been genetically engineered to produce milk that would be the same as human breast milk.<ref name="richardgray">{{cite web | last = Gray | first = Richard | name-list-format = vanc | date = 2011 | url = https://www.telegraph.co.uk/earth/agriculture/geneticmodification/8423536/Genetically-modified-cows-produce-human-milk.html | title = Genetically modified cows produce 'human' milk }}</ref> This could potentially benefit mothers who cannot produce breast milk but want their children to have breast milk rather than formula.<ref>{{cite web|url=http://www.classicalmedicinejournal.com/the-classical-medicine-journal/2011/4/13/genetically-modified-cows-producing-human-milk.html|title=Genetically modified cows producing human milk|author=Classical Medicine Journal|date=14 April 2010|archive-url=https://web.archive.org/web/20141106050820/http://www.classicalmedicinejournal.com/the-classical-medicine-journal/2011/4/13/genetically-modified-cows-producing-human-milk.html|archive-date=6 November 2014|deadurl=yes}}</ref><ref>{{cite news|url=https://www.telegraph.co.uk/news/worldnews/southamerica/argentina/8569687/Scientists-create-cow-that-produces-human-milk.html|title=Scientists create cow that produces 'human' milk|last=Yapp|first=Robin | name-list-format = vanc |date=11 June 2011|work=[[The Daily Telegraph]]|access-date=15 June 2012|location=London }}</ref> Researchers have also developed a genetically engineered cow that produces allergy-free milk.<ref>{{cite journal | vauthors = Jabed A, Wagner S, McCracken J, Wells DN, Laible G | title = Targeted microRNA expression in dairy cattle directs production of β-lactoglobulin-free, high-casein milk | journal = Proceedings of the National Academy of Sciences of the United States of America | volume = 109 | issue = 42 | pages = 16811–6 | date = October 2012 | pmid = 23027958 | pmc = 3479461 | doi = 10.1073/pnas.1210057109 | bibcode = 2012PNAS..10916811J }}</ref> |
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=== Research === |
=== Research === |
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== Nematodes == |
== Nematodes == |
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The [[Nematode|nemotode]] [[Caenorhabditis elegans|''Caenorhabditis elegans'']] is one of the major model organisms for researching [[molecular biology]].<ref>{{Cite web|url=http://www.wormbook.org/chapters/www_nematodeshistory/nematodeshistory.html|title=History of research on C. elegans and other free-living nematodes as model organisms|website=www.wormbook.org|access-date=2018-12-24}}</ref> [[RNA interference]] (RNAi) was discovered in ''C elegans''<ref>{{Cite journal|last=Hopkin|first=Michael|date=2006-10-02|title=RNAi scoops medical Nobel|url=http://www.nature.com/doifinder/10.1038/news061002-2|journal=news@nature|doi=10.1038/news061002-2|issn=1744-7933}}</ref> and could be induced by simply feeding them bacteria modified to express [[Double-stranded RNA|double stranded RNA]].<ref>{{ |
The [[Nematode|nemotode]] [[Caenorhabditis elegans|''Caenorhabditis elegans'']] is one of the major model organisms for researching [[molecular biology]].<ref>{{Cite web|url=http://www.wormbook.org/chapters/www_nematodeshistory/nematodeshistory.html|title=History of research on C. elegans and other free-living nematodes as model organisms|website=www.wormbook.org|access-date=2018-12-24}}</ref> [[RNA interference]] (RNAi) was discovered in ''C elegans''<ref>{{Cite journal|last=Hopkin|first=Michael|date=2006-10-02|title=RNAi scoops medical Nobel|url=http://www.nature.com/doifinder/10.1038/news061002-2|journal=news@nature|doi=10.1038/news061002-2|issn=1744-7933}}</ref> and could be induced by simply feeding them bacteria modified to express [[Double-stranded RNA|double stranded RNA]].<ref>{{cite journal | vauthors = Conte D, MacNeil LT, Walhout AJ, Mello CC | title = RNA Interference in Caenorhabditis elegans | journal = Current Protocols in Molecular Biology | volume = 109 | pages = 26.3.1-30 | date = January 2015 | pmid = 25559107 | pmc = 5396541 | doi = 10.1002/0471142727.mb2603s109 }}</ref> It is also relatively easy to produce stable transgenic nemotodes and this along with RNAi are the major tools used in studying their genes.<ref name=":16">{{cite journal | vauthors = Praitis V, Maduro MF | title = Transgenesis in C. elegans | journal = Methods in Cell Biology | volume = 106 | pages = 161–85 | date = 2011 | pmid = 22118277 | doi = 10.1016/B978-0-12-544172-8.00006-2 | url = https://www.sciencedirect.com/science/article/pii/B9780125441728000062 }}</ref> The most common use of transgenic nematodes has been studying gene expression and localisation by attaching reporter genes. Transgenes can also be combined with RNAi to rescue phenotypes, altered to study gene function, imaged in real time as the cells develop or used to control expression for different tissues or developmental stages.<ref name=":16" /> Transgenic nematodes have been used to study viruses,<ref>{{cite journal | vauthors = Diogo J, Bratanich A | title = The nematode Caenorhabditis elegans as a model to study viruses | journal = Archives of Virology | volume = 159 | issue = 11 | pages = 2843–51 | date = November 2014 | pmid = 25000902 | doi = 10.1007/s00705-014-2168-2 }}</ref> toxicology,<ref>{{cite journal | vauthors = Tejeda-Benitez L, Olivero-Verbel J | title = Caenorhabditis elegans, a Biological Model for Research in Toxicology | journal = Reviews of Environmental Contamination and Toxicology | volume = 237 | pages = 1–35 | date = 2016 | pmid = 26613986 | doi = 10.1007/978-3-319-23573-8_1 }}</ref> and diseases<ref>{{cite journal | vauthors = Schmidt J, Schmidt T | title = Animal Models of Machado-Joseph Disease | journal = Advances in Experimental Medicine and Biology | volume = 1049 | pages = 289–308 | date = 2018 | pmid = 29427110 | doi = 10.1007/978-3-319-71779-1_15 }}</ref><ref>{{cite journal | vauthors = Griffin EF, Caldwell KA, Caldwell GA | title = Genetic and Pharmacological Discovery for Alzheimer's Disease Using Caenorhabditis elegans | journal = ACS Chemical Neuroscience | volume = 8 | issue = 12 | pages = 2596–2606 | date = December 2017 | pmid = 29022701 | doi = 10.1021/acschemneuro.7b00361 }}</ref> and to detect environmental pollutants.<ref>{{cite book |last=Daniells|first=Clare |last2=Mutwakil|first2=Mohammed H. A. Z.|last3=Power|first3=Rowena S.|last4=David|first4=Helen E.|last5= De Pomerai|first5=David I. | name-list-format = vanc |chapter =Transgenic Nematodes as Biosensors of Environmental Stress |date=2002 |chapter-url=https://link.springer.com/chapter/10.1007/978-94-010-0357-5_15|title =Biotechnology for the Environment: Strategy and Fundamentals|pages=221–236|series=Focus on Biotechnology|publisher=Springer, Dordrecht |doi=10.1007/978-94-010-0357-5_15|isbn=9789401039079|access-date=2018-12-24}}</ref> |
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== Other == |
== Other == |
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The gene responsible for [[Albinism]] in [[Sea cucumber|sea cucumbers]] has been found and used to engineer [[White sea cucumber|white sea cucumbers]], a rare delicacy that is worth more per gram than gold. The technology also opens the way to investigate the genes responsible for some of the cucumbers more unusual traits, including [[Hibernation|hibernating]] in summer, [[Evisceration (autotomy)|eviscerating]] their intestines, and dissolving their bodies upon death.<ref>{{Cite web|url=https://www.scmp.com/tech/science-research/article/1846481/more-valuable-gold-not-long-genetically-modified-sea-cucumbers|title=More valuable than gold, but not for long: genetically-modified sea cucumbers headed to China's dinner tables|date=2015-08-05|website=South China Morning Post|language=en|access-date=2018-12-23}}</ref> [[Flatworm|Flatworms]] have the ability to regenerate themselves from a single cell.<ref name="pmid29906446">{{cite journal | vauthors = Zeng A, Li H, Guo L, Gao X, McKinney S, Wang Y, Yu Z, Park J, Semerad C, Ross E, Cheng LC, Davies E, Lei K, Wang W, Perera A, Hall K, Peak A, Box A, Sánchez Alvarado A |
The gene responsible for [[Albinism]] in [[Sea cucumber|sea cucumbers]] has been found and used to engineer [[White sea cucumber|white sea cucumbers]], a rare delicacy that is worth more per gram than gold. The technology also opens the way to investigate the genes responsible for some of the cucumbers more unusual traits, including [[Hibernation|hibernating]] in summer, [[Evisceration (autotomy)|eviscerating]] their intestines, and dissolving their bodies upon death.<ref>{{Cite web|url=https://www.scmp.com/tech/science-research/article/1846481/more-valuable-gold-not-long-genetically-modified-sea-cucumbers|title=More valuable than gold, but not for long: genetically-modified sea cucumbers headed to China's dinner tables|date=2015-08-05|website=South China Morning Post|language=en|access-date=2018-12-23}}</ref> [[Flatworm|Flatworms]] have the ability to regenerate themselves from a single cell.<ref name="pmid29906446">{{cite journal | vauthors = Zeng A, Li H, Guo L, Gao X, McKinney S, Wang Y, Yu Z, Park J, Semerad C, Ross E, Cheng LC, Davies E, Lei K, Wang W, Perera A, Hall K, Peak A, Box A, Sánchez Alvarado A | title = + Neoblasts Are Adult Pluripotent Stem Cells Underlying Planaria Regeneration | journal = Cell | volume = 173 | issue = 7 | pages = 1593–1608.e20 | date = June 2018 | pmid = 29906446 | doi = 10.1016/j.cell.2018.05.006 | lay-summary = http://www.nature.com/articles/d41586-018-05440-2 | lay-source = Nature }}</ref> Until 2017 there was no effective way to transform them, which hampered research. By using microinjection and radiation scientist have now created the first genetically modified flatworms.<ref>{{cite journal | vauthors = Wudarski J, Simanov D, Ustyantsev K, de Mulder K, Grelling M, Grudniewska M, Beltman F, Glazenburg L, Demircan T, Wunderer J, Qi W, Vizoso DB, Weissert PM, Olivieri D, Mouton S, Guryev V, Aboobaker A, Schärer L, Ladurner P, Berezikov E | title = Efficient transgenesis and annotated genome sequence of the regenerative flatworm model Macrostomum lignano | journal = Nature Communications | volume = 8 | issue = 1 | pages = 2120 | date = December 2017 | pmid = 29242515 | doi = 10.1038/s41467-017-02214-8 | url = https://www.nature.com/articles/s41467-017-02214-8 }}</ref> The [[bristle worm]], a marine [[annelid]], has been modified. It is of interest due to its reproductive cycle being synchronised with lunar phases, regeneration capacity and slow evolution rate.<ref>{{cite journal | vauthors = Zantke J, Bannister S, Rajan VB, Raible F, Tessmar-Raible K | title = Genetic and genomic tools for the marine annelid Platynereis dumerilii | journal = Genetics | volume = 197 | issue = 1 | pages = 19–31 | date = May 2014 | pmid = 24807110 | doi = 10.1534/genetics.112.148254 }}</ref> [[Cnidarians|Cnidaria]] such as ''[[Hydra (genus)|Hydra]]'' and the sea anemone ''[[Starlet sea anemone|Nematostella vectensis]]'' are attractive model organisms to study the [[evolution]] of [[immunity (medical)|immunity]] and certain developmental processes.<ref>{{cite journal | vauthors = Wittlieb J, Khalturin K, Lohmann JU, Anton-Erxleben F, Bosch TC | title = Transgenic Hydra allow in vivo tracking of individual stem cells during morphogenesis | journal = Proceedings of the National Academy of Sciences of the United States of America | volume = 103 | issue = 16 | pages = 6208–11 | date = April 2006 | pmid = 16556723 | pmc = 1458856 | doi = 10.1073/pnas.0510163103 | bibcode = 2006PNAS..103.6208W }}</ref> Other organisms that have been genetically modified include [[Snail|snails]],<ref name="pmid25529990">{{cite journal | vauthors = Perry KJ, Henry JQ | title = CRISPR/Cas9-mediated genome modification in the mollusc, Crepidula fornicata | journal = Genesis | volume = 53 | issue = 2 | pages = 237–44 | date = February 2015 | pmid = 25529990 | doi = 10.1002/dvg.22843 }}</ref> [[Gecko|geckos]], [[Turtle|turtles]],<ref>{{cite journal | vauthors = Nomura T, Yamashita W, Gotoh H, Ono K | title = Genetic manipulation of reptilian embryos: toward an understanding of cortical development and evolution | journal = Frontiers in Neuroscience | volume = 9 | pages = 45 | date = 2015-02-24 | pmid = 25759636 | pmc = 4338674 | doi = 10.3389/fnins.2015.00045 }}</ref> [[crayfish]], [[Oyster|oysters]], [[shrimp]], [[Clam|clams]], [[abalone]]<ref>{{Cite journal|last=Rasmussen|first=Rosalee S.|last2=Morrissey|first2=Michael T. | name-list-format = vanc |date=2007|title=Biotechnology in Aquaculture: Transgenics and Polyploidy |journal=Comprehensive Reviews in Food Science and Food Safety |volume=6|issue=1|pages=2–16|doi=10.1111/j.1541-4337.2007.00013.x }}</ref> and [[Sponge|sponges]].<ref>{{cite journal | vauthors = Ebert MS, Sharp PA | title = MicroRNA sponges: progress and possibilities | journal = Rna | volume = 16 | issue = 11 | pages = 2043–50 | date = November 2010 | pmid = 20855538 | pmc = 2957044 | doi = 10.1261/rna.2414110 }}</ref> |
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== References == |
== References == |
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Revision as of 15:21, 24 December 2018
Genetically modified animals are animals that have been genetically modified for a variety of purposes including producing drugs, enhancing yields, increase resistance to disease, etc.
The vast majority of genetically modified animals are at the research stage with the number close to entering the market remains small.[1]
Mammals
The process of genetically engineering mammals is a slow, tedious, and expensive process. However, new technologies are making genetic modifications easier and more precise.[2] The first transgenic mammals were produced by injecting viral DNA into embryos and then implanting the embryos in females.[3] The embryo would develop and it would be hoped that some of the genetic material would be incorporated into the reproductive cells. Then researchers would have to wait until the animal reached breeding age and then offspring would be screened for presence of the gene in every cell. The development of the CRISPR-Cas9 gene editing system has effectively halved the amount of time needed to develop genetically modified animals.[4] Despite the differences and difficulties in modifying them, the end aims are much the same as plants. GM mammals are created for research purposes, production of industrial or therapeutic products, agricultural uses or improving their health. There is also a market for creating genetically modified pets.[5]
Medicine
Mammals are the best models for human disease, making genetic engineered ones vital to the discovery and development of cures and treatments for many serious diseases. Knocking out genes responsible for human genetic disorders allows researchers to study the mechanism of the disease and to test possible cures. Genetically modified mice have been the most common mammals used in biomedical research, as they are cheap and easy to manipulate. Pigs are also a good target as they have a similar body size and anatomical features, physiology, pathophysiological response and diet.[6] Nonhuman primates are the most similar model organisms to humans, but there is less public acceptance towards using them as research animals.[7] In 2009, scientists 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.[8][9] Their first research target for these marmosets was Parkinson's disease, but they were also considering amyotrophic lateral sclerosis and Huntington's disease.[10]
Human proteins expressed in mammals are more likely to be similar to their natural counterparts than those expressed in plants or microorganisms. Stable expression has been accomplished in sheep, pigs, rats and other animals. In 2009, the first human biological drug produced from such an animal, a goat., was approved. The drug, ATryn, is an anticoagulant which reduces the probability of blood clots during surgery or childbirth was extracted from the goat's milk.[11] Human alpha-1-antitrypsin is another protein that is used in treating humans with this deficiency.[12] Another area is in creating pigs with greater capacity for human organ transplants (xenotransplantation). Pigs have been genetically modified so that their organs can no longer carry retroviruses[13] or have modifications to reduce the chance of rejection.[14][15] Pig lungs from genetically modified pigs are being considered for transplantation into humans.[16][17] There is even potential to create chimeric pigs that can carry human organs.[6][18]
Livestock
Livestock are modified with the intention of improving economically important traits such as growth-rate, quality of meat, milk composition, disease resistance and survival. Animals have been engineered to grow faster, be healthier[19] and resist diseases.[20] Modifications have also improved the wool production of sheep and udder health of cows.[1] Goats have been genetically engineered to produce milk with strong spiderweb-like silk proteins in their milk.[21] A GM pig called Enviropig was created with the capability of digesting plant phosphorus more efficiently than conventional pigs.[22][23] They could reduce water pollution since they excrete 30 to 70% less phosphorus in manure.[22][24] Dairy cows have been genetically engineered to produce milk that would be the same as human breast milk.[25] This could potentially benefit mothers who cannot produce breast milk but want their children to have breast milk rather than formula.[26][27] Researchers have also developed a genetically engineered cow that produces allergy-free milk.[28]
Research
Scientists have genetically engineered several organisms, including some mammals, to include green fluorescent protein (GFP), for research purposes.[29] GFP and other similar reporting genes allow easy visualisation and localisation of the products of the genetic modification.[30] Fluorescent pigs have been bred to study human organ transplants, regenerating ocular photoreceptor cells, and other topics.[31] In 2011 green-fluorescent cats were created to find therapies for HIV/AIDS and other diseases[32] as feline immunodeficiency virus (FIV) is related to HIV.[33]
Conservation
Genetically modification with a myxoma virus has been proposed to conserve European wild rabbits in the Iberian peninsula and to help regulate them in Australia. To protect the Iberian species from viral diseases, the myxoma virus was genetically modified to immunize the rabbits, while in Australia the same myxoma virus was genetically modified to lower fertility in the Australian rabbit population.[34] There have also been suggestions that genetic engineering could be used to bring animals back from extinction. It involves changing the genome of a close living relative to resemble the extinct one and is currently being attempted with the passenger pigeon.[35] Genes associated with the woolly mammoth have been added to the genome of an African Elephant, although the lead researcher says he has no intention of using live elephants.[36]
Humans
Gene therapy,[37] uses genetically modified viruses to deliver genes which 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,[38] and Leber's congenital amaurosis.[39] Treatments are also being developed for a range of other currently incurable diseases, such as cystic fibrosis,[40] sickle cell anemia,[41] Parkinson's disease,[42][43] cancer,[44][45][46] diabetes,[47] heart disease[48] and muscular dystrophy.[49] These treatments only effect somatic cells, meaning any changes would not be inheritable. Germline gene therapy results in any change being inheritable, which has raised concerns within the scientific community.[50][51] In 2015, CRISPR was used to edit the DNA of non-viable human embryos.[52][53] In November 2018, He Jiankui announced that he had edited the genomes of two human embryos, to attempt to disable the CCR5 gene, which codes for a receptor that HIV uses to enter cells. He said that twin girls, Lulu and Nana, had been born a few weeks earlier and that they carried functional copies of CCR5 along with disabled CCR5 (mosaicism) and were still vulnerable to HIV. The work was widely condemned as unethical, dangerous, and premature.[54]
Fish
Genetically modified fish are used for scientific research, as pets and as a food source. Aquaculture is a growing industry, currently providing over half the consumed fish worldwide.[55] Through genetic engineering It is possible to increase growth rates, reduce food intake, remove allergenic properties, increase cold tolerance and provide disease resistance. Fish can also be used to detect aquatic pollution or function as bioreactors.[56] Several groups have been developing zebrafish to detect pollution by attaching fluorescent proteins to genes activated by the presence of pollutants. The fish will then glow and can be used as environmental sensors.[57][58] The GloFish is a brand of genetically modified fluorescent zebrafish with bright red, green, and orange fluorescent color. It was originally developed by one of the groups to detect pollution, but is now part of the ornamental fish trade, becoming the first genetically modified animal to become publicly available as a pet when it was introduced for sale in 2003.[59]
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.[60] Zebrafish are model organisms for developmental processes, regeneration, genetics, behaviour, disease mechanisms and toxicity testing.[61] Their transparency allows researchers to observe developmental stages, intestinal functions and tumour growth.[62][63] The generation of transgenic protocols (whole organism, cell or tissue specific, tagged with reporter genes) has increased the level of information gained by studying these fish.[64]
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,[65] trout[66] and tilapia.[67] AquaBounty Technologies, a biotechnology company working on bringing a GM salmon to market, says that their GM AquAdvantage salmon can mature in half the time as wild salmon.[68] It obtained regulatory approval in 2015, the first non-plant GMO food to be commerialised.[69]
Insects
In biological research, transgenic fruit flies (Drosophila melanogaster) are model organisms used to study the effects of genetic changes on development.[70] Fruit flies are often preferred over other animals due to their short life cycle and low maintenance requirements. It also has a relatively simple genome compared to many vertebrates, with typically only one copy of each gene, making phenotypic analysis easy.[71] Drosophila have been used to study genetics and inheritance, embryonic development, learning, behavior, and aging.[72] Transposons (particularly P elements) are well developed in Drosophila and provided an early method to add transgenes to their genome, although this has been taken over by more modern gene-editing techniques.[73]
Due to their significance to human health, scientists are looking at ways to control mosquitoes through genetic engineering. Malaria-resistant mosquitoes have been developed in the laboratory.[74] by inserting a gene that reduces the development of the malaria parasite[75] and then use homing endonucleases to rapidly spread that gene throughout the male population (known as a gene drive).[76] This has been taken further by swapping it for a lethal gene.[77][78] In trials the populations of Aedes aegypti mosquitoes, the single most important carrier of dengue fever and Zika virus, were reduced by between 80% and by 90%.[79] [80][78] Another approach is to use the sterile insect technique, whereby males genetically engineered to be sterile out compete viable males, to reduce population numbers.[81]
Other insect pests that make attractive targets are moths. Diamondback moths cause US$4 to $5 billion of damage a year worldwide.[82] The approach is similar to the mosquitoes, where males transformed with a gene that prevents females from reaching maturity will be released.[83] They underwent field trials in 2017.[82] Genetically modified moths have previously been released in field trials.[84] A strain of pink bollworm that were sterilised with radiation were genetically engineered to express a red fluorescent protein making it easier for researchers to monitor them.[85]
Silkworm, the larvae stage of Bombyx mori, is an economically important insect in sericulture. Scientists are developing strategies to enhance silk quality and quantity. There is also potential to use the silk producing machinery to make other valuable proteins.[86] Proteins expressed by silkworms include; human serum albumin, human collagen α-chain, mouse monoclonal antibody and N-glycanase.[87] Silkworms have been created that produce spider silk, a stronger but extremely difficult to harvest silk,[88] and even novel silks.[89]
Birds
Systems have been developed to create transgenic organisms in a wide variety of other animals. Chickens have been genetically modified for a variety of purposes. This includes studying embryo development,[90] preventing the transmission of bird flu[91] and providing evolutionary insights using reverse engineering to recreate dinosaur-like phenotypes.[92] A GM chicken that produces the drug Kanuma, an enzyme that treats a rare condition, in its egg passed regulatory approval in 2015.[93] Genetically modified frogs, in particular Xenopus laevis and Xenopus tropicalis, are used in development biology. GM frogs can also being used as pollution sensors, especially for endocrine disrupting chemicals.[94] There are proposals to use genetic engineering to control cane toads in Australia.[95][96]
Nematodes
The nemotode Caenorhabditis elegans is one of the major model organisms for researching molecular biology.[97] RNA interference (RNAi) was discovered in C elegans[98] and could be induced by simply feeding them bacteria modified to express double stranded RNA.[99] It is also relatively easy to produce stable transgenic nemotodes and this along with RNAi are the major tools used in studying their genes.[100] The most common use of transgenic nematodes has been studying gene expression and localisation by attaching reporter genes. Transgenes can also be combined with RNAi to rescue phenotypes, altered to study gene function, imaged in real time as the cells develop or used to control expression for different tissues or developmental stages.[100] Transgenic nematodes have been used to study viruses,[101] toxicology,[102] and diseases[103][104] and to detect environmental pollutants.[105]
Other
The gene responsible for Albinism in sea cucumbers has been found and used to engineer white sea cucumbers, a rare delicacy that is worth more per gram than gold. The technology also opens the way to investigate the genes responsible for some of the cucumbers more unusual traits, including hibernating in summer, eviscerating their intestines, and dissolving their bodies upon death.[106] Flatworms have the ability to regenerate themselves from a single cell.[107] Until 2017 there was no effective way to transform them, which hampered research. By using microinjection and radiation scientist have now created the first genetically modified flatworms.[108] The bristle worm, a marine annelid, has been modified. It is of interest due to its reproductive cycle being synchronised with lunar phases, regeneration capacity and slow evolution rate.[109] Cnidaria such as Hydra and the sea anemone Nematostella vectensis are attractive model organisms to study the evolution of immunity and certain developmental processes.[110] Other organisms that have been genetically modified include snails,[111] geckos, turtles,[112] crayfish, oysters, shrimp, clams, abalone[113] and sponges.[114]
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