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=== Biotechnology === [[File:Blue-green algae cultured in specific media.jpg|thumb|right|Cyanobacteria cultured in specific media: Cyanobacteria can be helpful in agriculture as they have the ability to fix atmospheric nitrogen in soil.]] The unicellular cyanobacterium ''[[Synechocystis]]'' sp. PCC6803 was the third prokaryote and first photosynthetic organism whose [[genome]] was completely [[DNA sequencing|sequenced]].<ref>{{cite journal | vauthors = Kaneko T, Sato S, Kotani H, Tanaka A, Asamizu E, Nakamura Y, Miyajima N, Hirosawa M, Sugiura M, Sasamoto S, Kimura T, Hosouchi T, Matsuno A, Muraki A, Nakazaki N, Naruo K, Okumura S, Shimpo S, Takeuchi C, Wada T, Watanabe A, Yamada M, Yasuda M, Tabata S | display-authors = 6 | title = Sequence analysis of the genome of the unicellular cyanobacterium Synechocystis sp. strain PCC6803. II. Sequence determination of the entire genome and assignment of potential protein-coding regions | journal = DNA Research | volume = 3 | issue = 3 | pages = 109–136 | date = June 1996 | pmid = 8905231 | doi = 10.1093/dnares/3.3.109 | doi-access = free }}</ref> It continues to be an important model organism.<ref>{{cite journal | vauthors = Tabei Y, Okada K, Tsuzuki M | title = Sll1330 controls the expression of glycolytic genes in Synechocystis sp. PCC 6803 | journal = Biochemical and Biophysical Research Communications | volume = 355 | issue = 4 | pages = 1045–1050 | date = April 2007 | pmid = 17331473 | doi = 10.1016/j.bbrc.2007.02.065 }}</ref> ''[[Crocosphaera|Crocosphaera subtropica]]'' ATCC 51142 is an important [[diazotroph]]ic model organism.<ref>{{cite journal |last1=Mareš |first1=Jan |last2=Johansen |first2=Jeffrey R. |last3=Hauer |first3=Tomáš |last4=Zima |first4=Jan |last5=Ventura |first5=Stefano |last6=Cuzman |first6=Oana |last7=Tiribilli |first7=Bruno |last8=Kaštovský |first8=Jan |title=Taxonomic resolution of the genus Cyanothece (Chroococcales, Cyanobacteria), with a treatment on Gloeothece and three new genera, Crocosphaera, Rippkaea , and Zehria |journal=Journal of Phycology |date=June 2019 |volume=55 |issue=3 |pages=578–610 |doi=10.1111/jpy.12853 |pmid=30830691 |bibcode=2019JPcgy..55..578M }}</ref> The smallest genomes of a photosynthetic organism have been found in ''Prochlorococcus'' spp. (1.7 [[Genome size|Mb]])<ref>{{cite journal | vauthors = Rocap G, Larimer FW, Lamerdin J, Malfatti S, Chain P, Ahlgren NA, Arellano A, Coleman M, Hauser L, Hess WR, Johnson ZI, Land M, Lindell D, Post AF, Regala W, Shah M, Shaw SL, Steglich C, Sullivan MB, Ting CS, Tolonen A, Webb EA, Zinser ER, Chisholm SW | display-authors = 6 | title = Genome divergence in two Prochlorococcus ecotypes reflects oceanic niche differentiation | journal = Nature | volume = 424 | issue = 6952 | pages = 1042–1047 | date = August 2003 | pmid = 12917642 | doi = 10.1038/nature01947 | doi-access = free | bibcode = 2003Natur.424.1042R }}</ref><ref>{{cite journal | vauthors = Dufresne A, Salanoubat M, Partensky F, Artiguenave F, Axmann IM, Barbe V, Duprat S, Galperin MY, Koonin EV, Le Gall F, Makarova KS, Ostrowski M, Oztas S, Robert C, Rogozin IB, Scanlan DJ, Tandeau de Marsac N, Weissenbach J, Wincker P, Wolf YI, Hess WR | display-authors = 6 | title = Genome sequence of the cyanobacterium Prochlorococcus marinus SS120, a nearly minimal oxyphototrophic genome | journal = Proceedings of the National Academy of Sciences of the United States of America | volume = 100 | issue = 17 | pages = 10020–10025 | date = August 2003 | pmid = 12917486 | pmc = 187748 | doi = 10.1073/pnas.1733211100 | doi-access = free | bibcode = 2003PNAS..10010020D }}</ref> and the largest in ''[[Nostoc punctiforme]]'' (9 Mb).<ref name="Meeks-2001"/> Those of ''[[Calothrix]]'' spp. are estimated at 12–15 Mb,<ref>{{Cite journal |doi=10.1099/00221287-111-1-73 |title=Genome Size of Cyanobacteria |journal=Journal of General Microbiology |volume=111 |pages=73–85 |year=1979 |vauthors=Herdman M, Janvier M, Rippka R, Stanier RY |issue=1 |doi-access=free}}</ref> as large as [[yeast]]. Recent research has suggested the potential application of cyanobacteria to the generation of [[renewable energy]] by directly converting sunlight into electricity. Internal photosynthetic pathways can be coupled to chemical mediators that transfer electrons to external [[electrodes]].<ref>{{cite journal | vauthors = Quintana N, Van der Kooy F, Van de Rhee MD, Voshol GP, Verpoorte R | title = Renewable energy from Cyanobacteria: energy production optimization by metabolic pathway engineering | journal = Applied Microbiology and Biotechnology | volume = 91 | issue = 3 | pages = 471–490 | date = August 2011 | pmid = 21691792 | pmc = 3136707 | doi = 10.1007/s00253-011-3394-0 }}</ref><ref>{{cite journal | vauthors = Chen X, Lawrence JM, Wey LT, Schertel L, Jing Q, Vignolini S, Howe CJ, Kar-Narayan S, Zhang JZ | display-authors = 6 | title = 3D-printed hierarchical pillar array electrodes for high-performance semi-artificial photosynthesis | journal = Nature Materials | volume = 21 | issue = 7 | pages = 811–818 | date = July 2022 | pmid = 35256790 | doi = 10.1038/s41563-022-01205-5 | bibcode = 2022NatMa..21..811C | url = https://www.repository.cam.ac.uk/handle/1810/334792 }}</ref> In the shorter term, efforts are underway to commercialize [[algae-based fuels]] such as [[diesel fuel|diesel]], [[gasoline]], and [[jet fuel]].<ref name="Pisciotta JM, Zou Y, Baskakov IV 2010 e108212"/><ref>[http://www.thehindu.com/sci-tech/science/article477049.ece "Blue green bacteria may help generate 'green' electricity"], ''The Hindu'', 21 June 2010</ref><ref>{{Cite web |date=2010-09-14 |title=Joule wins key patent for GMO cyanobacteria that create fuels from sunlight, CO2 and water : Biofuels Digest |url=https://www.biofuelsdigest.com/bdigest/2010/09/14/joule-wins-key-patent-for-gmo-cyanobacteria-that-create-fuels-from-sunlight-co2-and-water/ |access-date=2022-08-10 |language=en-US}}</ref> Cyanobacteria have been also engineered to produce ethanol<ref>{{cite journal | vauthors = Deng MD, Coleman JR | title = Ethanol synthesis by genetic engineering in cyanobacteria | journal = Applied and Environmental Microbiology | volume = 65 | issue = 2 | pages = 523–528 | date = February 1999 | pmid = 9925577 | pmc = 91056 | doi = 10.1128/AEM.65.2.523-528.1999 | bibcode = 1999ApEnM..65..523D }}</ref> and experiments have shown that when one or two CBB genes are being over expressed, the yield can be even higher.<ref>{{Cite journal |title=Engineering photoautotrophic carbon fixation for enhanced growth and productivity | vauthors = Liang F, Lindberg P, Lindblad P |date=20 November 2018 |journal=Sustainable Energy & Fuels |volume=2 |issue=12 |pages=2583–2600 |doi=10.1039/C8SE00281A|doi-access=free }}</ref><ref>{{cite journal | vauthors = Roussou S, Albergati A, Liang F, Lindblad P | title = Engineered cyanobacteria with additional overexpression of selected Calvin-Benson-Bassham enzymes show further increased ethanol production | journal = Metabolic Engineering Communications | volume = 12 | pages = e00161 | date = June 2021 | pmid = 33520653 | pmc = 7820548 | doi = 10.1016/j.mec.2021.e00161 }}</ref> Cyanobacteria may possess the ability to produce substances that could one day serve as anti-inflammatory agents and combat bacterial infections in humans.<ref>{{cite journal | vauthors = Choi H, Mascuch SJ, Villa FA, Byrum T, Teasdale ME, Smith JE, Preskitt LB, Rowley DC, Gerwick L, Gerwick WH | display-authors = 6 | title = Honaucins A-C, potent inhibitors of inflammation and bacterial quorum sensing: synthetic derivatives and structure-activity relationships | journal = Chemistry & Biology | volume = 19 | issue = 5 | pages = 589–598 | date = May 2012 | pmid = 22633410 | pmc = 3361693 | doi = 10.1016/j.chembiol.2012.03.014 }}</ref> Cyanobacteria's photosynthetic output of sugar and oxygen has been demonstrated to have therapeutic value in rats with heart attacks.<ref>{{cite news | vauthors = Frischkorn K |title=Need to Fix a Heart Attack? Try Photosynthesis |url=https://www.smithsonianmag.com/science-nature/how-light-activated-bacteria-could-help-heal-heart-attack-180963756/ |access-date=20 May 2021 |work=[[Smithsonian (magazine)|Smithsonian]] |date=19 June 2017}}</ref> While cyanobacteria can naturally produce various secondary metabolites, they can serve as advantageous hosts for plant-derived metabolites production owing to biotechnological advances in systems biology and synthetic biology.<ref>{{cite journal | vauthors = Jeong Y, Cho SH, Lee H, Choi HK, Kim DM, Lee CG, Cho S, Cho BK | display-authors = 6 | title = Current Status and Future Strategies to Increase Secondary Metabolite Production from Cyanobacteria | journal = Microorganisms | volume = 8 | issue = 12 | pages = 1849 | date = November 2020 | pmid = 33255283 | pmc = 7761380 | doi = 10.3390/microorganisms8121849 | doi-access = free }}</ref> Spirulina's extracted blue color is used as a natural food coloring.<ref>{{cite journal | vauthors = Newsome AG, Culver CA, van Breemen RB | title = Nature's palette: the search for natural blue colorants | journal = Journal of Agricultural and Food Chemistry | volume = 62 | issue = 28 | pages = 6498–6511 | date = July 2014 | pmid = 24930897 | doi = 10.1021/jf501419q | bibcode = 2014JAFC...62.6498N }}</ref> Researchers from several space agencies argue that cyanobacteria could be used for producing goods for human consumption in future crewed outposts on Mars, by transforming materials available on this planet.<ref name="Verseux2015">{{cite journal | vauthors = Verseux C, Baqué M, Lehto K, de Vera JP, Rothschild LJ, Billi D |author5-link=Lynn J. Rothschild |title=Sustainable life support on Mars – the potential roles of cyanobacteria |journal=International Journal of Astrobiology |volume=15 |issue=1 |pages=65–92 |date=2016 |doi=10.1017/S147355041500021X |bibcode=2016IJAsB..15...65V |doi-access=free}}</ref>
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