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Genetic engineering
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==History== {{main|History of genetic engineering}} Humans have altered the genomes of species for thousands of years through [[selective breeding]], or artificial selection<ref name=Root>{{cite book|title=Domestication|url={{google books |plainurl=y |id=WGDYHvOHwmwC}}|first=Clive |last=Root| name-list-style = vanc |year=2007|publisher=Greenwood Publishing Groups}}</ref>{{rp|1}}<ref name=Zohary>{{cite book |title=Domestication of Plants in the Old World: The origin and spread of plants in the old world|url={{google books |plainurl=y |id=tc6vr0qzk_4C}}|first1=Daniel |last1=Zohary |first2=Maria |last2=Hopf |first3=Ehud |last3=Weiss | name-list-style = vanc |year=2012|publisher=Oxford University Press}}</ref>{{rp|1}} as contrasted with [[natural selection]]. More recently, [[mutation breeding]] has used exposure to chemicals or radiation to produce a high frequency of random mutations, for selective breeding purposes. Genetic engineering as the direct manipulation of DNA by humans outside breeding and mutations has only existed since the 1970s. The term "genetic engineering" was coined by the Russian-born geneticist [[Nikolay Timofeev-Ressovsky]] in his 1934 paper "The Experimental Production of Mutations", published in the British journal Biological Reviews.<ref>{{cite journal | vauthors = Timofeev-Ressovsky NW | title = The Experimental Production of Mutations | journal = Biological Reviews | volume = 9 | issue = 4 | pages = 411β457 | date = October 1934 | doi = 10.1111/j.1469-185X.1934.tb01255.x | s2cid = 86396986 | url = https://archive.org/details/in.ernet.dli.2015.26044/page/n449}}</ref> [[Jack Williamson]] used the term in his science fiction novel Dragon's Island, published in 1951<ref>{{cite book|last=Stableford|first=Brian M.| name-list-style = vanc |title=Historical dictionary of science fiction literature|page=133|year=2004|publisher=Scarecrow Press |isbn=978-0-8108-4938-9|url={{google books |plainurl=y |id=nzmIPZg5xicC|page=9}}}}</ref> β one year before DNA's role in [[heredity]] was confirmed by [[Alfred Hershey]] and [[Martha Chase]],<ref>{{cite journal | vauthors = Hershey AD, Chase M | title = Independent functions of viral protein and nucleic acid in growth of bacteriophage | journal = The Journal of General Physiology | volume = 36 | issue = 1 | pages = 39β56 | date = May 1952 | pmid = 12981234 | pmc = 2147348 | doi = 10.1085/jgp.36.1.39 }}</ref> and two years before [[James D. Watson|James Watson]] and [[Francis Crick]] showed that the [[DNA]] molecule has a double-helix structure β though the general concept of direct genetic manipulation was explored in rudimentary form in [[Stanley G. Weinbaum]]'s 1936 science fiction story ''Proteus Island''.<ref>{{cite encyclopedia|url=http://www.sf-encyclopedia.com/entry/genetic_engineering|title=Genetic Engineering|date=2 April 2015|encyclopedia=Encyclopedia of Science Fiction}}</ref><ref>{{Cite book|url=https://books.google.com/books?id=WJrvj0pOJx0C&pg=PA164 |title=Modern Concepts in Nanotechnology, Volume 5|year=2008|isbn=978-81-8356-296-6|publisher=Discovery Publishing House |author=Shiv Kant Prasad |author2=Ajay Dash}}</ref> [[File:Jaenisch 2003 by Sam Ogden.jpg|thumb|upright|In 1974 [[Rudolf Jaenisch]] created a [[genetically modified mouse]], the first GM animal.]] In 1972, [[Paul Berg]] created the first [[recombinant DNA]] molecules by combining DNA from the monkey virus [[SV40]] with that of the [[Lambda phage|lambda virus]].<ref>{{cite journal | vauthors = Jackson DA, Symons RH, Berg P | title = 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 | journal = Proceedings of the National Academy of Sciences of the United States of America | volume = 69 | issue = 10 | pages = 2904β9 | date = October 1972 | pmid = 4342968 | pmc = 389671 | doi = 10.1073/pnas.69.10.2904 | first8 = David A. Jackson Robert H. Symons and Paul Berg | bibcode = 1972PNAS...69.2904J | doi-access = free }}</ref> In 1973 [[Herbert Boyer]] and [[Stanley Norman Cohen|Stanley Cohen]] created the first [[transgenic organism]] by inserting [[Antibiotic resistance#Applications|antibiotic resistance genes]] into the [[plasmid]] of an ''[[Escherichia coli]]'' bacterium.<ref>{{cite web|title=History of Genetics: Genetic Engineering Timeline|last=Arnold|first=Paul| name-list-style = vanc |year=2009|url=http://www.brighthub.com/science/genetics/articles/21983.aspx}}</ref><ref>{{cite journal | vauthors = Gutschi S, Hermann W, Stenzl W, Tscheliessnigg KH | title = [Displacement of electrodes in pacemaker patients (author's transl)] | journal = Zentralblatt fΓΌr Chirurgie | volume = 104 | issue = 2 | pages = 100β4 | date = 1 May 1973 | pmid = 433482}}</ref> A year later [[Rudolf Jaenisch]] created a [[transgenic mouse]] by introducing foreign DNA into its embryo, making it the world's first [[transgenic animal]]<ref name="pmid4364530">{{cite journal | vauthors = Jaenisch R, Mintz B | title = Simian virus 40 DNA sequences in DNA of healthy adult mice derived from preimplantation blastocysts injected with viral DNA | journal = Proceedings of the National Academy of Sciences of the United States of America | volume = 71 | issue = 4 | pages = 1250β4 | date = April 1974 | pmid = 4364530 | pmc = 388203 | doi = 10.1073/pnas.71.4.1250| bibcode = 1974PNAS...71.1250J | doi-access = free }}</ref> These achievements led to concerns in the scientific community about potential risks from genetic engineering, which were first discussed in depth at the [[Asilomar Conference on Recombinant DNA|Asilomar Conference]] in 1975. One of the main recommendations from this meeting was that government oversight of recombinant DNA research should be established until the technology was deemed safe.<ref name="BergBaltimoreBrennerRoblinSinger1975">{{cite journal | vauthors = Berg P, Baltimore D, Brenner S, Roblin RO, Singer MF | title = Summary statement of the Asilomar conference on recombinant DNA molecules | journal = Proceedings of the National Academy of Sciences of the United States of America | volume = 72 | issue = 6 | pages = 1981β4 | date = June 1975 | pmid = 806076 | pmc = 432675 | doi = 10.1073/pnas.72.6.1981 | bibcode = 1975PNAS...72.1981B | doi-access = free }}</ref><ref>{{cite web | title = NIH Guidelines for research involving recombinant DNA molecules | work = Office of Biotechnology Activities | publisher = U.S. Department of Health and Human Services | url = http://oba.od.nih.gov/rdna/nih_guidelines_oba.html | archive-url = https://web.archive.org/web/20120910070047/http://oba.od.nih.gov/rdna/nih_guidelines_oba.html | archive-date = 10 September 2012 }}</ref> In 1976 [[Genentech]], the first genetic engineering company, was founded by Herbert Boyer and [[Robert A. Swanson|Robert Swanson]] and a year later the company produced a human protein ([[somatostatin]]) in ''E. coli''. Genentech announced the production of genetically engineered human [[insulin]] in 1978.<ref name="GoeddelKleidBolivarHeynekerYansuraCreaHiroseKraszewskiItakuraRiggs1979">{{cite journal | vauthors = Goeddel DV, Kleid DG, Bolivar F, Heyneker HL, Yansura DG, Crea R, Hirose T, Kraszewski A, Itakura K, Riggs AD | title = Expression in Escherichia coli of chemically synthesized genes for human insulin | journal = Proceedings of the National Academy of Sciences of the United States of America | volume = 76 | issue = 1 | pages = 106β10 | date = January 1979 | pmid = 85300 | pmc = 382885 | doi = 10.1073/pnas.76.1.106 | bibcode = 1979PNAS...76..106G | doi-access = free }}</ref> In 1980, the [[U.S. Supreme Court]] in the ''[[Diamond v. Chakrabarty]]'' case ruled that genetically altered life could be patented.<ref>{{cite web |url=http://supreme.justia.com/us/447/303/case.html |title=Diamond V Chakrabarty|volume=447|author=US Supreme Court Cases from Justia & Oyez|website=Justia |date=16 June 1980 |access-date=17 July 2010 |issue=303}}</ref> The [[Insulin (medication)|insulin]] produced by bacteria was approved for release by the [[Food and Drug Administration]] (FDA) in 1982.<ref>{{cite magazine|url=http://www.time.com/time/magazine/article/0,9171,949646-1,00.html |archive-url=https://web.archive.org/web/20111027011602/http://www.time.com/time/magazine/article/0,9171,949646-1,00.html |archive-date=27 October 2011 |title=Artificial Genes |magazine=[[Time (magazine)|Time]] |date=15 November 1982 |access-date=17 July 2010}}</ref> In 1983, a biotech company, Advanced Genetic Sciences (AGS) applied for U.S. government authorisation to perform field tests with the [[Ice-minus bacteria|ice-minus strain]] of ''[[Pseudomonas syringae]]'' to protect crops from frost, but environmental groups and protestors delayed the field tests for four years with legal challenges.<ref>{{cite journal | last1 = Bratspies | first1 = Rebecca | name-list-style = vanc | year = 2007 | title = Some Thoughts on the American Approach to Regulating Genetically Modified Organisms | ssrn = 1017832 | journal = Kansas Journal of Law & Public Policy | volume = 16 | issue = 3 | pages = 101β31 }}</ref> In 1987, the ice-minus strain of ''P. syringae'' became the first [[genetically modified organism]] (GMO) to be released into the environment<ref name=BBC2002>{{Cite news|date=2002-06-14|title=GM crops: A bitter harvest?|language=en-GB|url=http://news.bbc.co.uk/2/hi/science/nature/2045286.stm|access-date=2023-03-30}}</ref> when a strawberry field and a potato field in California were sprayed with it.<ref>{{cite news |last=Maugh |first=Thomas H. II |date=9 June 1987 |url=https://www.latimes.com/archives/la-xpm-1987-06-09-mn-6024-story.html |title=Altered Bacterium Does Its Job: Frost Failed to Damage Sprayed Test Crop, Company Says |work=[[Los Angeles Times]]}}</ref> Both test fields were attacked by activist groups the night before the tests occurred: "The world's first trial site attracted the world's first field trasher".<ref name=BBC2002 /> The first field trials of [[transgenic plants|genetically engineered plants]] occurred in France and the US in 1986, tobacco plants were engineered to be resistant to [[herbicide]]s.<ref>{{cite web |last=James |first=Clive | name-list-style = vanc |title=Global Review of the Field Testing and Commercialization of Transgenic Plants: 1986 to 1995 |url=http://www.isaaa.org/kc/Publications/pdfs/isaaabriefs/Briefs%201.pdf |archive-url=https://web.archive.org/web/20100616175626/http://isaaa.org/kc/Publications/pdfs/isaaabriefs/Briefs%201.pdf |archive-date=2010-06-16 |url-status=live |publisher=The International Service for the Acquisition of Agri-biotech Applications|access-date=17 July 2010 |year=1996}}</ref> The People's Republic of China was the first country to commercialise transgenic plants, introducing a virus-resistant tobacco in 1992.<ref name="James1997">{{cite journal|last=James |first=Clive | name-list-style = vanc |year=1997 |title=Global Status of Transgenic Crops in 1997 |journal=ISAAA Briefs No. 5. |page=31 |url=http://www.isaaa.org/resources/publications/briefs/05/download/isaaa-brief-05-1997.pdf |archive-url=https://web.archive.org/web/20090116155014/http://www.isaaa.org/Resources/Publications/briefs/05/download/isaaa-brief-05-1997.pdf |archive-date=2009-01-16 |url-status=live}}</ref> In 1994 [[Monsanto#Spin-offs and mergers|Calgene]] attained approval to commercially release the first [[genetically modified food]], the [[Flavr Savr]], a tomato engineered to have a longer shelf life.<ref>{{cite journal |doi=10.3733/ca.v054n04p6 |title=The case of the FLAVR SAVR tomato |year=2000 | vauthors = Bruening G, Lyons JM |journal=California Agriculture |volume=54 |issue=4 |pages=6β7|doi-broken-date=1 November 2024 |doi-access=free }}</ref> In 1994, the European Union approved tobacco engineered to be resistant to the herbicide [[bromoxynil]], making it the first genetically engineered crop commercialised in Europe.<ref>{{cite magazine|title=Transgenic tobacco is European first|date=18 June 1994 |last=MacKenzie |first=Debora | name-list-style = vanc |url=https://www.newscientist.com/article/mg14219301.100-transgenic-tobacco-is-european-first.html|magazine=[[New Scientist]]}}</ref> In 1995, [[Genetically modified potato#NewLeaf|Bt potato]] was approved safe by the [[United States Environmental Protection Agency|Environmental Protection Agency]], after having been approved by the FDA, making it the first pesticide producing crop to be approved in the US.<ref>{{Cite web|title=Lawrence Journal-World - Google News Archive Search|url=https://news.google.com/newspapers?nid=2199&dat=19950506&id=A0YyAAAAIBAJ&sjid=jOYFAAAAIBAJ&pg=4631,1776980&hl=en|access-date=2023-03-30|website=news.google.com}}</ref> In 2009 11 transgenic crops were grown commercially in 25 countries, the largest of which by area grown were the US, Brazil, Argentina, India, Canada, China, Paraguay and South Africa.<ref>{{Cite web|title=Executive Summary: Global Status of Commercialized Biotech/GM Crops: 2009 - ISAAA Brief 41-2009 | ISAAA.org|url=https://www.isaaa.org/resources/publications/briefs/41/executivesummary/default.asp|access-date=2023-03-30|website=www.isaaa.org}}</ref> In 2010, scientists at the [[J. Craig Venter Institute]] created the first [[synthetic genome]] and inserted it into an empty bacterial cell. The resulting bacterium, named [[Mycoplasma laboratorium]], could [[replicate (biology)|replicate]] and produce proteins.<ref>{{cite journal | vauthors = Pennisi E | author-link = Elizabeth Pennisi | title = Genomics. Synthetic genome brings new life to bacterium | journal = Science | volume = 328 | issue = 5981 | pages = 958β9 | date = May 2010 | pmid = 20488994 | doi = 10.1126/science.328.5981.958 | doi-access = free }}</ref><ref>{{cite journal | vauthors = Gibson DG, Glass JI, Lartigue C, Noskov VN, Chuang RY, Algire MA, Benders GA, Montague MG, Ma L, Moodie MM, Merryman C, Vashee S, Krishnakumar R, Assad-Garcia N, Andrews-Pfannkoch C, Denisova EA, Young L, Qi ZQ, Segall-Shapiro TH, Calvey CH, Parmar PP, Hutchison CA, Smith HO, Venter JC | display-authors = 6 | title = Creation of a bacterial cell controlled by a chemically synthesized genome | journal = Science | volume = 329 | issue = 5987 | pages = 52β6 | date = July 2010 | pmid = 20488990 | doi = 10.1126/science.1190719 | bibcode = 2010Sci...329...52G | citeseerx = 10.1.1.167.1455 | s2cid = 7320517 }}</ref> Four years later this was taken a step further when a bacterium was developed that replicated a [[plasmid]] containing a unique [[base pair]], creating the first organism engineered to use an expanded genetic alphabet.<ref>{{cite journal | vauthors = Malyshev DA, Dhami K, Lavergne T, Chen T, Dai N, Foster JM, CorrΓͺa IR, Romesberg FE | title = A semi-synthetic organism with an expanded genetic alphabet | journal = Nature | volume = 509 | issue = 7500 | pages = 385β8 | date = May 2014 | pmid = 24805238 | pmc = 4058825 | doi = 10.1038/nature13314 | bibcode = 2014Natur.509..385M }}</ref><ref>{{cite journal | vauthors = Thyer R, Ellefson J | s2cid = 4399670 | title = Synthetic biology: New letters for life's alphabet | journal = Nature | volume = 509 | issue = 7500 | pages = 291β2 | date = May 2014 | pmid = 24805244 | doi = 10.1038/nature13335 | bibcode = 2014Natur.509..291T | doi-access = free }}</ref> In 2012, [[Jennifer Doudna]] and [[Emmanuelle Charpentier]] collaborated to develop the [[CRISPR|CRISPR/Cas9]] system,<ref>{{Cite news |url=https://www.nytimes.com/2015/05/12/science/jennifer-doudna-crispr-cas9-genetic-engineering.html |archive-url=https://ghostarchive.org/archive/20220102/https://www.nytimes.com/2015/05/12/science/jennifer-doudna-crispr-cas9-genetic-engineering.html |archive-date=2022-01-02 |url-access=limited |url-status=live |title=Jennifer Doudna, a Pioneer Who Helped Simplify Genome Editing|last=Pollack|first=Andrew | name-list-style = vanc |date=2015-05-11 |work=[[The New York Times]] |access-date=2017-11-15}}{{cbignore}}</ref><ref>{{cite journal | vauthors = Jinek M, Chylinski K, Fonfara I, Hauer M, Doudna JA, Charpentier E | title = A programmable dual-RNA-guided DNA endonuclease in adaptive bacterial immunity | journal = Science | volume = 337 | issue = 6096 | pages = 816β21 | date = August 2012 | pmid = 22745249 | pmc = 6286148 | doi = 10.1126/science.1225829 | bibcode = 2012Sci...337..816J }}</ref> a technique which can be used to easily and specifically alter the genome of almost any organism.<ref>{{cite journal | vauthors = Ledford H | title = CRISPR: gene editing is just the beginning | journal = Nature | volume = 531 | issue = 7593 | pages = 156β9 | date = March 2016 | pmid = 26961639 | doi = 10.1038/531156a | bibcode = 2016Natur.531..156L | doi-access = free }}</ref>
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