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cyanobacteria
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{{redirect|Cyanobacterium|the genus|Cyanobacterium (genus)}}{{inadequate lead|date=June 2016}}{{Use dmy dates|date=January 2011}}{hide}Automatic taxobox
|fossil_range = {{long fossil range|2700|0{edih}
|image = CSIRO ScienceImage 4203 A bluegreen algae species Cylindrospermum sp under magnification.jpg
|image_caption = Microscope image of Cylindrospermum, a genus of Cyanobacteria
|display_parents = 3
|parent_authority = Stanier, 1973
|taxon = Cyanophyceae
|authority =
|subdivision_ranks = OrdersJOURNAL, Komárek J, Kaštovský J, Mareš J, Johansen JR, 2014, Taxonomic classification of cyanoprokaryotes (cyanobacterial genera) 2014, using a polyphasic approach,weblink Preslia, 86, 295–335,
|subdivision = {{as of| 2014}} the taxonomy was under revisionWEB, Cyanophyceae,weblink Access Science, 21 April 2011, JOURNAL, Oren A, A proposal for further integration of the cyanobacteria under the Bacteriological Code, International Journal of Systematic and Evolutionary Microbiology, 54, Pt 5, 1895–902, September 2004, 15388760, 10.1099/ijs.0.03008-0,
}}(File:Cyanobacteria guerrero negro.jpg|thumb|255px| Light microscope view of cyanobacteria from a microbial mat)Cyanobacteria {{IPAc-en|s|aɪ|ˌ|æ|n|oʊ|b|æ|k|ˈ|t|ɪər|i|ə}}, also known as Cyanophyta, are a phylum of bacteria that obtain their energy through photosynthesisWEB, Life History and Ecology of Cyanobacteria,weblink University of California Museum of Paleontology, 17 July 2012, and are the only photosynthetic prokaryotes able to produce oxygen.JOURNAL, Hamilton TL, Bryant DA, Macalady JL, The role of biology in planetary evolution: cyanobacterial primary production in low-oxygen Proterozoic oceans, Environmental Microbiology, 18, 2, 325–40, February 2016, 26549614, 5019231, 10.1111/1462-2920.13118, The name cyanobacteria comes from the color of the bacteria ().WEB,weblink cyan {{!, Origin and meaning of cyan by Online Etymology Dictionary|website=www.etymonline.com|access-date=2018-01-21}}WEB,weblink Henry George Liddell, Robert Scott, A Greek-English Lexicon, κύα^νος, www.perseus.tufts.edu, 2018-01-21, Cyanobacteria, which are prokaryotes, are also called "blue-green algae",WEB, Taxonomy Browser - Cyanobacteria,weblink National Center for Biotechnology Information, NCBI:txid1117, 12 April 2018, though the term algae in modern usage is restricted to eukaryotes.ENCYCLOPEDIA, 1992, Algae, The Concise Dictionary of Botany, Oxford University Press, Oxford, Allaby, M, Cyanobacteria appear to have originated in freshwater or a terrestrial environment.BOOK,weblink The Marine Microbiome: An Untapped Source of Biodiversity and Biotechnological Potential, Lucas J., Stal, Mariana Silvia, Cretoiu, 3 June 2016, Springer, Google Books, 9783319330006, Unlike heterotrophic prokaryotes, cyanobacteria have internal membranes. These are flattened sacs called thylakoids where photosynthesis is performed.BOOK, The Cyanobacteria: Molecular Biology, Genomics, and Evolution, Chapter 10. Membrane Systems in Cyanobacteria, Michelle, Liberton, Himadri B., Pakrasi, vanc, Horizon Scientific Press, 2008, 978-1-904455-15-8, Norwich, United Kingdom, 217–287, Antonia, Herrero, Enrique, Flore, JOURNAL, Liberton M, Page LE, O'Dell WB, O'Neill H, Mamontov E, Urban VS, Pakrasi HB, Organization and flexibility of cyanobacterial thylakoid membranes examined by neutron scattering, The Journal of Biological Chemistry, 288, 5, 3632–40, February 2013, 23255600, 3561581, 10.1074/jbc.M112.416933, Phototrophic eukaryotes perform photosynthesis by plastids that may have their ancestry in cyanobacteria, acquired long ago via a process called endosymbiosis. These endosymbiotic cyanobacteria in eukaryotes may have evolved or differentiated into specialized organelles such as chloroplasts, etioplasts and leucoplasts.By producing and releasing oxygen (as a byproduct of photosynthesis), cyanobacteria are thought to have converted the early oxygen-poor, reducing atmosphere into an oxidizing one, causing the Great Oxygenation Event and the "rusting of the Earth",BOOK, Whitton, Brian A., vanc, Ecology of Cyanobacteria II: Their Diversity in Space and Time, {{google books, y, 4oJ_vi27s18C, 17, |date=5 July 2012|chapter=The fossil record of cyanobacteria|publisher=Springer Science & Business Media|isbn=978-94-007-3855-3|pages=17–}} which dramatically changed the composition of the Earth's life forms and led to the near-extinction of anaerobic organisms.

Description

Cyanobacteria are a group of photosynthetic bacteria, some of which are nitrogen-fixing, that live in a wide variety of moist soils and water either freely or in a symbiotic relationship with plants or lichen-forming fungi (as in the lichen genus Peltigera).JOURNAL, Dodds WK, Gudder DA, Mollenhauer D, 1995, T he ecology of Nostoc, Journal of Phycology, 31, 2–18, 10.1111/j.0022-3646.1995.00002.x, They range from unicellular to filamentous and include colonial species. Colonies may form filaments, sheets, or even hollow spheres. Some filamentous species can differentiate into several different cell types: vegetative cells – the normal, photosynthetic cells that are formed under favorable growing conditions; akinetes – climate-resistant spores that may form when environmental conditions become harsh; and thick-walled heterocysts – which contain the enzyme nitrogenase, vital for nitrogen fixationJOURNAL, Meeks JC, Elhai J, Thiel T, Potts M, Larimer F, Lamerdin J, Predki P, Atlas R, An overview of the genome of Nostoc punctiforme, a multicellular, symbiotic cyanobacterium, Photosynthesis Research, 70, 1, 85–106, 2001, 16228364, 10.1023/A:1013840025518, JOURNAL, Golden JW, Yoon HS, Heterocyst formation in Anabaena, Current Opinion in Microbiology, 1, 6, 623–9, December 1998, 10066546, 10.1016/s1369-5274(98)80106-9, JOURNAL, Fay P, Oxygen relations of nitrogen fixation in cyanobacteria, Microbiological Reviews, 56, 2, 340–73, June 1992, 1620069, 372871, in an anaerobic environment due to its sensitivity to oxygen.

Nitrogen fixation

Some cyanobacteria can fix atmospheric nitrogen in anaerobic conditions by means of specialized cells called heterocysts. Heterocysts may also form under the appropriate environmental conditions (anoxic) when fixed nitrogen is scarce. Heterocyst-forming species are specialized for nitrogen fixation and are able to fix nitrogen gas into ammonia ({{Nh3}}), nitrites ({{No2-}}) or nitrates ({{No3-}}), which can be absorbed by plants and converted to protein and nucleic acids (atmospheric nitrogen is not bioavailable to plants, except for those having endosymbiotic nitrogen-fixing bacteria, especially the Fabaceae family, among others).Free-living cyanobacteria are present in the water of rice paddies, and cyanobacteria can be found growing as epiphytes on the surfaces of the green alga, Chara, where they may fix nitrogen.JOURNAL, Sims GK, Dunigan EP, 1984, Diurnal and seasonal variations in nitrogenase activity (C2H2 reduction) of rice roots, Soil Biology and Biochemistry, 16, 15–18, 10.1016/0038-0717(84)90118-4, Cyanobacteria such as Anabaena (a symbiont of the aquatic fern Azolla) can provide rice plantations with biofertilizer.WEB, Azolla-Anabaena as a Biofertilizer for Rice Paddy Fields in the Po Valley, a Temperate Rice Area in Northern Italy,weblink International Journal of Agronomy, 21 April 2011,

Morphology

File:CyanobacteriaColl1.jpg|thumb|left|Colonies of Nostoc pruniformeNostoc pruniformeMany cyanobacteria form motile filaments of cells, called hormogonia, that travel away from the main biomass to bud and form new colonies elsewhere.JOURNAL, Risser DD, Chew WG, Meeks JC, Genetic characterization of the hmp locus, a chemotaxis-like gene cluster that regulates hormogonium development and motility in Nostoc punctiforme, Molecular Microbiology, 92, 2, 222–33, April 2014, 24533832, 10.1111/mmi.12552, JOURNAL, Khayatan B, Bains DK, Cheng MH, Cho YW, Huynh J, Kim R, Omoruyi OH, Pantoja AP, Park JS, Peng JK, Splitt SD, Tian MY, Risser DD, A Putative O-Linked β-N-Acetylglucosamine Transferase Is Essential for Hormogonium Development and Motility in the Filamentous Cyanobacterium Nostoc punctiforme, Journal of Bacteriology, 199, 9, May 2017, 28242721, 5388816, 10.1128/JB.00075-17, e00075–17, The cells in a hormogonium are often thinner than in the vegetative state, and the cells on either end of the motile chain may be tapered. To break away from the parent colony, a hormogonium often must tear apart a weaker cell in a filament, called a necridium.Each individual cell (each single cyanobacterium) typically has a thick, gelatinous cell wall.BOOK, {{google books, y, xNQE_89dat8C, 72, |title=Text Book Of Botany Diversity Of Microbes And Cryptogams|last=Singh|publisher=Rastogi Publications|isbn=9788171338894}} They lack flagella, but hormogonia of some species can move about by gliding along surfaces.NEWS,weblink Differences between Bacteria and Cyanobacteria, 2015-10-29, Microbiology Notes, 2018-01-21, Many of the multicellular filamentous forms of Oscillatoria are capable of a waving motion; the filament oscillates back and forth. In water columns, some cyanobacteria float by forming gas vesicles, as in archaea.JOURNAL, Walsby AE, Gas vesicles, Microbiological Reviews, 58, 1, 94–144, March 1994, 8177173, 372955, These vesicles are not organelles as such. They are not bounded by lipid membranes but by a protein sheath.

Ecology

{{Further|Algal bloom}}File:2010 Filamentous Cyanobacteria Bloom near Fiji.jpg|thumb|right|upright|Cyanobacterial bloom near FijiFijiCyanobacteria can be found in almost every terrestrial and aquatic habitat—oceans, fresh water, damp soil, temporarily moistened rocks in deserts, bare rock and soil, and even Antarctic rocks. They can occur as planktonic cells or form phototrophic biofilms. They are found in endolithic ecosystem.JOURNAL, de los Ríos A, Grube M, Sancho LG, Ascaso C, Ultrastructural and genetic characteristics of endolithic cyanobacterial biofilms colonizing Antarctic granite rocks, FEMS Microbiology Ecology, 59, 2, 386–95, February 2007, 17328119, 10.1111/j.1574-6941.2006.00256.x, A few are endosymbionts in lichens, plants, various protists, or sponges and provide energy for the host. Some live in the fur of sloths, providing a form of camouflage.BOOK, Vaughan, Terry, vanc, Mammalogy, 2011, Jones and Barlett, 21, {{google books, y, LD1nDlzXYicC, 21, |isbn=9780763762995}}Aquatic cyanobacteria are known for their extensive and highly visible blooms that can form in both freshwater and marine environments. The blooms can have the appearance of blue-green paint or scum. These blooms can be toxic, and frequently lead to the closure of recreational waters when spotted. Marine bacteriophages are significant parasites of unicellular marine cyanobacteria.MAGAZINE, Schultz, Nora, 30 August 2009,weblink Photosynthetic viruses keep world's oxygen levels up, New Scientist, Cyanobacteria growth is favored in ponds and lakes where waters are calm and have less turbulent mixing.JOURNAL, Jöhnk, Klaus D., Huisman, Jef, Sharples, Jonathan, Sommeijer, Ben, Visser, Petra M., Stroom, Jasper M., vanc, Summer heatwaves promote blooms of harmful cyanobacteria, Global Change Biology, 1 March 2008, 14, 3, 495–512, 10.1111/j.1365-2486.2007.01510.x, 1365-2486, 2008GCBio..14..495J,weblink Their life cycles are disrupted when the water naturally or artificially mixes from churning currents caused by the flowing water of streams or the churning water of fountains. For this reason blooms of cyanobacteria seldom occur in rivers unless the water is flowing slowly. Growth is also favored at higher temperatures which enable Microcystis species to outcompete diatoms and green algae, and potentially allow development of toxins.Based on environmental trends, models and observations suggest cyanobacteria will likely increase their dominance in aquatic environments. This can lead to serious consequences, particularly the contamination of sources of drinking water. Cyanobacteria can interfere with water treatment in various ways, primarily by plugging filters (often large beds of sand and similar media) and by producing cyanotoxins, which have the potential to cause serious illness if consumed. Consequences may also lie within fisheries and waste management practices. Anthropogenic eutrophication, rising temperatures, vertical stratification and increased atmospheric carbon dioxide are contributors to cyanobacteria increasing dominance of aquatic ecosystems.JOURNAL, Paerl HW, Paul VJ, Climate change: links to global expansion of harmful cyanobacteria, Water Research, 46, 5, 1349–63, April 2012, 21893330, 10.1016/j.watres.2011.08.002, Cyanobacteria have been found to play an important role in terrestrial habitats. It has been widely reported that cyanobacteria soil crusts help to stabilize soil to prevent erosion and retain water.JOURNAL, Thomas AD, Dougill AJ, Spatial and temporal distribution of cyanobacterial soil crusts in the Kalahari: Implications for soil surface properties, Geomorphology, 15 March 2007, 85, 1, 17–29, 10.1016/j.geomorph.2006.03.029, 2007Geomo..85...17T, An example of a cyanobacterial species that does so is Microcoleus vaginatus. M. vaginatus stabilizes soil using a polysaccharide sheath that binds to sand particles and absorbs water.JOURNAL, Belnap, Jayne, Gardner, John S., vanc, Soil Microstructure in Soils of the Colorado Plateau: The Role of the Cyanobacterium Microcoleus Vaginatus, The Great Basin Naturalist, 1993, 53, 1, 40–47, 41712756, Some of these organisms contribute significantly to global ecology and the oxygen cycle. The tiny marine cyanobacterium Prochlorococcus was discovered in 1986 and accounts for more than half of the photosynthesis of the open ocean.JOURNAL, Nadis S, The cells that rule the seas, Scientific American, 289, 6, 52–3, December 2003, 14631732, 10.1038/scientificamerican1203-52,weblink 2003SciAm.289f..52N, 19 April 2014,weblink" title="web.archive.org/web/20140419222251weblink">weblink 19 April 2014, yes, Circadian rhythms were once thought to only exist in eukaryotic cells but many cyanobacteria display a bacterial circadian rhythm.Cyanobacteria are arguably the most successful group of microorganisms on earth. They are the most genetically diverse; they occupy a broad range of habitats across all latitudes, widespread in freshwater, marine, and terrestrial ecosystems, and they are found in the most extreme niches such as hot springs, salt works, and hypersaline bays. Photoautotrophic, oxygen-producing cyanobacteria created the conditions in the planet's early atmosphere that directed the evolution of aerobic metabolism and eukaryotic photosynthesis. Cyanobacteria fulfill vital ecological functions in the world's oceans, being important contributors to global carbon and nitrogen budgets. – Stewart and FalconerBOOK, Walsh, Patrick J., Smith, Sharon, Fleming, Lora, Helena, Solo-Gabriele, William H., Gerwick, vanc, Oceans and Human Health: Risks and Remedies from the Seas, {{google books, y, LMZPqW-PmFYC, 271, |date=2 September 2011|chapter=Cyanobacteria and cyanobacterial toxins|pp=271–296|publisher=Academic Press|isbn=978-0-08-087782-2}}

Photosynthesis

While contemporary cyanobacteria are linked to the plant kingdom as descendants of the progenitor of the endosymbiotic chloroplast, there are several features which are unique to this group.

Carbon fixation

Cyanobacteria use the energy of sunlight to drive photosynthesis, a process where the energy of light is used to synthesize organic compounds from carbon dioxide. Because they are aquatic organisms, they typically employ several strategies which are collectively known as a "carbon concentrating mechanism" to aid in the acquisition of inorganic carbon ({{CO2}} or bicarbonate). Among the more specific strategies is the widespread prevalence of the bacterial microcompartments known as carboxysomes.JOURNAL, Kerfeld CA, Heinhorst S, Cannon GC, Bacterial microcompartments, Annual Review of Microbiology, 64, 1, 391–408, 2010, 20825353, 10.1146/annurev.micro.112408.134211,weblink These icosahedral structures are composed of hexameric shell proteins that assemble into cage-like structures that can be several hundreds of nanometers in diameter. It is believed that these structures tether the {{CO2}}-fixing enzyme, RuBisCO, to the interior of the shell, as well as the enzyme carbonic anhydrase, using metabolic channeling to enhance the local {{CO2}} concentrations and thus increase the efficiency of the RuBisCO enzyme.JOURNAL, Long BM, Badger MR, Whitney SM, Price GD, Analysis of carboxysomes from Synechococcus PCC7942 reveals multiple Rubisco complexes with carboxysomal proteins CcmM and CcaA, The Journal of Biological Chemistry, 282, 40, 29323–35, October 2007, 17675289, 10.1074/jbc.M703896200,

Electron transport

In contrast to purple bacteria and other bacteria performing anoxygenic photosynthesis, thylakoid membranes of cyanobacteria are not continuous with the plasma membrane but are separate compartments.JOURNAL, Vothknecht UC, Westhoff P, Biogenesis and origin of thylakoid membranes, Biochimica et Biophysica Acta, 1541, 1–2, 91–101, December 2001, 11750665, 10.1016/S0167-4889(01)00153-7, The photosynthetic machinery is embedded in the thylakoid membranes, with phycobilisomes acting as light-harvesting antennae attached to the membrane, giving the green pigmentation observed (with wavelengths from 450 nm to 660 nm) in most cyanobacteria.JOURNAL, Sobiechowska-Sasim, Monika, StoÅ„-Egiert, Joanna, Kosakowska, Alicja, vanc, February 2014, Quantitative analysis of extracted phycobilin pigments in cyanobacteria—an assessment of spectrophotometric and spectrofluorometric methods, J Appl Phycol, 26, 5, 2065–2074, 10.1007/s10811-014-0244-3, 25346572, 4200375, (File:Cyanobacterium-inline.svg|right|440x440px|Diagram of a typical cyanobacterial cell)While most of the high-energy electrons derived from water are used by the cyanobacterial cells for their own needs, a fraction of these electrons may be donated to the external environment via electrogenic activity.JOURNAL, Pisciotta JM, Zou Y, Baskakov IV, Light-dependent electrogenic activity of cyanobacteria, PLOS ONE, 5, 5, e10821, May 2010, 20520829, 2876029, 10.1371/journal.pone.0010821, 2010PLoSO...510821P, Yang, Ching-Hong, vanc,

Respiration

Respiration in cyanobacteria can occur in the thylakoid membrane alongside photosynthesis,BOOK, Photosynthesis and Respiration in Cyanobacteria. eLS., Vermaas, Wim FJ, vanc, 2001, John Wiley & Sons, Ltd, 978-0-470-01590-2, 10.1038/npg.els.0001670, Photosynthesis and Respiration in Cyanobacteria, with their photosynthetic electron transport sharing the same compartment as the components of respiratory electron transport. While the goal of photosynthesis is to store energy by building carbohydrates from CO2, respiration is the reverse of this, with carbohydrates turned back into CO2 accompanying energy release.Cyanobacteria appear to separate these two processes with their plasma membrane containing only components of the respiratory chain, while the thylakoid membrane hosts an interlinked respiratory and photosynthetic electron transport chain. Cyanobacteria use electrons from succinate dehydrogenase rather than from NADPH for respiration.Cyanobacteria only respire during the night (or in the dark) because the facilities used for electron transport are used in reverse for photosynthesis while in the light.BOOK,weblink How the Earth Turned Green: A Brief 3.8-Billion-Year History of Plants, Armstronf, Joseph E., vanc, 2015, The University of Chicago Press, 978-0-226-06977-7,

Electron transport chain

Many cyanobacteria are able to reduce nitrogen and carbon dioxide under aerobic conditions, a fact that may be responsible for their evolutionary and ecological success. The water-oxidizing photosynthesis is accomplished by coupling the activity of photosystem (PS) II and I (Z-scheme). In contrast to green sulfur bacteria which only use one photosystem, the use of water as an electron donor is energetically demanding, requiring two photosystems.JOURNAL, Klatt JM, de Beer D, Häusler S, Polerecky L, Cyanobacteria in Sulfidic Spring Microbial Mats Can Perform Oxygenic and Anoxygenic Photosynthesis Simultaneously during an Entire Diurnal Period, Frontiers in Microbiology, 7, 1973, 2016, 28018309, 10.3389/fmicb.2016.01973, 5156726, Attached to the thylakoid membrane, phycobilisomes act as light-harvesting antennae for the photosystems.JOURNAL, Grossman AR, Schaefer MR, Chiang GG, Collier JL, The phycobilisome, a light-harvesting complex responsive to environmental conditions, Microbiological Reviews, 57, 3, 725–49, September 1993, 8246846, 372933, The phycobilisome components (phycobiliproteins) are responsible for the blue-green pigmentation of most cyanobacteria.WEB,weblink Colors from bacteria {{!, Causes of Color|website=www.webexhibits.org|access-date=2018-01-22}} The variations on this theme are due mainly to carotenoids and phycoerythrins that give the cells their red-brownish coloration. In some cyanobacteria, the color of light influences the composition of the phycobilisomes.BOOK, 10.1016/B978-012373944-5.00250-9, Cyanobacteria, Encyclopedia of Microbiology, third, 107–124, F., Garcia-Pichel, Moselio, Schaechter, vanc, 978-0-12-373944-5, 2009, JOURNAL, Kehoe DM, Chromatic adaptation and the evolution of light color sensing in cyanobacteria, Proceedings of the National Academy of Sciences of the United States of America, 107, 20, 9029–9030, May 2010, 20457899, 2889117, 10.1073/pnas.1004510107, 2010PNAS..107.9029K, In green light, the cells accumulate more phycoerythrin, whereas in red light they produce more phycocyanin. Thus, the bacteria appear green in red light and red in green light.JOURNAL, Kehoe DM, Gutu A, Responding to color: the regulation of complementary chromatic adaptation, Annual Review of Plant Biology, 57, 127–50, 2006, 16669758, 10.1146/annurev.arplant.57.032905.105215, This process of complementary chromatic adaptation is a way for the cells to maximize the use of available light for photosynthesis.A few genera lack phycobilisomes and have chlorophyll b instead (Prochloron, Prochlorococcus, Prochlorothrix). These were originally grouped together as the prochlorophytes or chloroxybacteria, but appear to have developed in several different lines of cyanobacteria. For this reason, they are now considered as part of the cyanobacterial group.JOURNAL, Palenik B, Haselkorn R, Multiple evolutionary origins of prochlorophytes, the chlorophyll b-containing prokaryotes, Nature, 355, 6357, 265–7, January 1992, 1731224, 10.1038/355265a0, 1992Natur.355..265P, JOURNAL, Urbach E, Robertson DL, Chisholm SW, Multiple evolutionary origins of prochlorophytes within the cyanobacterial radiation, Nature, 355, 6357, 267–70, January 1992, 1731225, 10.1038/355267a0, 1992Natur.355..267U,

Metabolism

In general, photosynthesis in cyanobacteria uses water as an electron donor and produces oxygen as a byproduct, though some may also use hydrogen sulfideJOURNAL, Cohen Y, Jørgensen BB, Revsbech NP, Poplawski R, Adaptation to Hydrogen Sulfide of Oxygenic and Anoxygenic Photosynthesis among Cyanobacteria, Applied and Environmental Microbiology, 51, 2, 398–407, February 1986, 16346996, 238881, a process which occurs among other photosynthetic bacteria such as the purple sulfur bacteria.Carbon dioxide is reduced to form carbohydrates via the Calvin cycle.BOOK, Molecular Mechanisms of Photosynthesis, Robert E., Blankenship, vanc, Wiley-Blackwell, 2014, 978-1-4051-8975-0, 147–173, The large amounts of oxygen in the atmosphere are considered to have been first created by the activities of ancient cyanobacteria.JOURNAL, Och, Lawrence M., Shields-Zhou, Graham A., vanc, The Neoproterozoic oxygenation event: Environmental perturbations and biogeochemical cycling, Earth-Science Reviews, 110, 1–4, 26–57, 10.1016/j.earscirev.2011.09.004, January 2012, 2012ESRv..110...26O, They are often found as symbionts with a number of other groups of organisms such as fungi (lichens), corals, pteridophytes (Azolla), angiosperms (Gunnera), etc.BOOK, The Prokaryotes, Adams, David G., Bergman, Birgitta, Nierzwicki-Bauer, Sandra A., Duggan, Paula S., Rai, Amar N., Schüßler, Arthur, vanc, 2013, Springer, Berlin, Heidelberg, 978-3-642-30193-3, 359–400, 10.1007/978-3-642-30194-0_17, There are some groups capable of heterotrophic growth,BOOK, Smith, A.J., Synthesis of metabolic intermediates, Carr, N. G., Whitton, Brian A., vanc, The Biology of Blue-green Algae, {{google books, y, fSRPg-D0Jk0C, 30, |year=1973|publisher=University of California Press|isbn=978-0-520-02344-4|pages=30–}} while others are parasitic, causing diseases in invertebrates or algae (e.g., the black band disease).JOURNAL, Jangoux, M, vanc, 1987, Diseases of Echinodermata. I. Agents microorganisms and protistans, Dis. Aquat. Org., 2, 147–162, 10.3354/dao002147, BOOK, Kinne O,weblink Diseases of Marine Animals, 1, John Wiley & Sons, Chichester, UK, 978-0-471-99584-5, 1980, JOURNAL, Kristiansen, Aase, vanc, 10.2216/i0031-8884-4-1-19.1,weblink Sarcinastrum urosporae, a Colourless Parasitic Blue-green Alga, Phycologia, 1964, 4, 1, 19–22, yes,weblink" title="web.archive.org/web/20150106045403weblink">weblink 6 January 2015, {{clear}}

Relationship to chloroplasts

{{See also|Chloroplast#Chloroplast lineages and evolution}}Primary chloroplasts are cell organelles found in some eukaryotic lineages, where they are specialized in performing the photosynthesis. They are known to have evolved from cyanobacteria through endosymbiosis, i.e. after the engulfment of a cell by another. In this case, a photosynthesizing cyanobacteria that was engulfed in some ancient eukaryotic cell.JOURNAL, Keeling PJ, The number, speed, and impact of plastid endosymbioses in eukaryotic evolution, Annual Review of Plant Biology, 64, 583–607, 2013, 23451781, 10.1146/annurev-arplant-050312-120144, After some years of debate,JOURNAL, Howe CJ, Barbrook AC, Nisbet RE, Lockhart PJ, Larkum AW, The origin of plastids, Philosophical Transactions of the Royal Society of London. Series B, Biological Sciences, 363, 1504, 2675–85, August 2008, 18468982, 2606771, 10.1098/rstb.2008.0050, it is now generally accepted that the three major groups of primary endosymbiotic eukaryotes (i.e. green plants, red algae and glaucophytes) form one large monophyletic group called Archaeplastida, which evolved after one unique endosymbiotic event.JOURNAL, Rodríguez-Ezpeleta N, Brinkmann H, Burey SC, Roure B, Burger G, Löffelhardt W, Bohnert HJ, Philippe H, Lang BF, Monophyly of primary photosynthetic eukaryotes: green plants, red algae, and glaucophytes, Current Biology, 15, 14, 1325–30, July 2005, 16051178, 10.1016/j.cub.2005.06.040, JOURNAL, Adl SM, Simpson AG, Lane CE, Lukeš J, Bass D, Bowser SS, Brown MW, Burki F, Dunthorn M, Hampl V, Heiss A, Hoppenrath M, Lara E, Le Gall L, Lynn DH, McManus H, Mitchell EA, Mozley-Stanridge SE, Parfrey LW, Pawlowski J, Rueckert S, Shadwick L, Shadwick L, Schoch CL, Smirnov A, Spiegel FW, The revised classification of eukaryotes, The Journal of Eukaryotic Microbiology, 59, 5, 429–93, September 2012, 23020233, 3483872, 10.1111/j.1550-7408.2012.00644.x, JOURNAL, Price DC, Chan CX, Yoon HS, Yang EC, Qiu H, Weber AP, Schwacke R, Gross J, Blouin NA, Lane C, Reyes-Prieto A, Durnford DG, Neilson JA, Lang BF, Burger G, Steiner JM, Löffelhardt W, Meuser JE, Posewitz MC, Ball S, Arias MC, Henrissat B, Coutinho PM, Rensing SA, Symeonidi A, Doddapaneni H, Green BR, Rajah VD, Boore J, Bhattacharya D, Cyanophora paradoxa genome elucidates origin of photosynthesis in algae and plants, Science, 335, 6070, 843–7, February 2012, 22344442, 10.1126/science.1213561, 2012Sci...335..843P, JOURNAL, Ponce-Toledo RI, Deschamps P, López-García P, Zivanovic Y, Benzerara K, Moreira D, An Early-Branching Freshwater Cyanobacterium at the Origin of Plastids, Current Biology, 27, 3, 386–391, February 2017, 28132810, 10.1016/j.cub.2016.11.056, 5650054, The morphological similarity between chloroplasts and cyanobacteria was first reported by German botanist Andreas Schimper in the 19th centuryJOURNAL, Schimper, A.F.W., Über die Entwicklung der Chlorophyllkörner und Farbkörper, About the development of the chlorophyll grains and stains, German, Bot. Zeitung, 1883, 41, 105–14, 121–31, 137–46, 153–62,weblink yes,weblink" title="web.archive.org/web/20131019121025weblink">weblink 19 October 2013, dmy-all, Chloroplasts are only found in plants and algae,BOOK, Alberts, Bruce, vanc, Molecular biology of the cell, 2002, Garland, New York [u.a.], 978-0-8153-4072-0,weblink 4., thus paving the way for Russian biologist Konstantin Mereschkowski to suggest the symbiogenic origin of the plastid in 1905.JOURNAL, Mereschkowsky C, Über Natur und Ursprung der Chromatophoren im Pflanzenreiche, About the nature and origin of chromatophores in the vegetable kingdom, German, Biol Centralbl, 1905, 25, 593–604,weblink Lynn Margulis brought this hypothesis back to attention more than 60 years laterJOURNAL, Sagan L, On the origin of mitosing cells, Journal of Theoretical Biology, 14, 3, 255–74, March 1967, 11541392, 10.1016/0022-5193(67)90079-3, but it was not until supplementary data started to accumulate that the idea became fully accepted. The cyanobacterial origin of plastids is now supported by various pieces of phylogenetic,JOURNAL, Schwartz RM, Dayhoff MO, Origins of prokaryotes, eukaryotes, mitochondria, and chloroplasts, Science, 199, 4327, 395–403, January 1978, 202030, 10.1126/science.202030, 1978Sci...199..395S, genomic,JOURNAL, Archibald JM, Genomic perspectives on the birth and spread of plastids, Proceedings of the National Academy of Sciences of the United States of America, 112, 33, 10147–53, August 2015, 25902528, 4547232, 10.1073/pnas.1421374112, 2015PNAS..11210147A, biochemicalJOURNAL, Blankenship RE, Early evolution of photosynthesis, Plant Physiology, 154, 2, 434–8, October 2010, 20921158, 2949000, 10.1104/pp.110.161687, JOURNAL, Rockwell NC, Lagarias JC, Bhattacharya D, Primary endosymbiosis and the evolution of light and oxygen sensing in photosynthetic eukaryotes, Frontiers in Ecology and Evolution, 2, 66, 2014, 25729749, 4343542, 10.3389/fevo.2014.00066, and structural evidence.Summarised in JOURNAL, Cavalier-Smith T, Membrane heredity and early chloroplast evolution, Trends in Plant Science, 5, 4, 174–82, April 2000, 10740299, 10.1016/S1360-1385(00)01598-3, The fact that another independent and more recent primary endosymbiosis event has been described between a cyanobacterium and a separate eukaryote lineage (the rhizarian Paulinella chromatophora) also gives credibility to the endosymbiotic origin of the plastid.JOURNAL, Nowack EC, Melkonian M, Glöckner G, Chromatophore genome sequence of Paulinella sheds light on acquisition of photosynthesis by eukaryotes, Current Biology, 18, 6, 410–8, March 2008, 18356055, 10.1016/j.cub.2008.02.051, In addition to this primary endosymbiosis, many eukaryotic lineages have been subject to secondary or even tertiary endosymbiotic events, that is the "Matryoshka-like" engulfment by a eukaryote of another plastid-bearing eukaryote.JOURNAL, Archibald JM, The puzzle of plastid evolution, Current Biology, 19, 2, R81–8, January 2009, 19174147, 10.1016/j.cub.2008.11.067, Within this evolutionary context, it is noteworthy that, as far as we can tell, oxygenic photosynthesis only evolved once (in cyanobacteria), and all other photosynthetic eukaryotes (including all plants and algae) have acquired this ability from them. In other words, all the oxygen that makes the atmosphere breathable for aerobic organisms originally comes from cyanobacteria or their later descendants.WEB,weblink How do plants make oxygen? Ask cyanobacteria, 30 March 2017, Phys.org, Science X, 2017-10-26,

DNA repair

Cyanobacteria are challenged by environmental stresses and internally generated reactive oxygen species that cause DNA damages. Cyanobacteria possess numerous E. coli-like DNA repair genes.JOURNAL, Cassier-Chauvat C, Veaudor T, Chauvat F, Comparative Genomics of DNA Recombination and Repair in Cyanobacteria: Biotechnological Implications, Front Microbiol, 7, 1809, 2016, 27881980, 5101192, 10.3389/fmicb.2016.01809, Several DNA repair genes are highly conserved in cyanobacteria, even in small genomes, suggesting that core DNA repair processes such as recombinational repair, nucleotide excision repair and methyl-directed DNA mismatch repair are common among cyanobacteria.

Natural genetic transformation

Cyanobacteria are capable of natural genetic transformation.JOURNAL, Orkwiszewski KG, Kaney AR, Genetic transformation of the blue-green bacterium, Anacystis nidulans, Arch Mikrobiol, 98, 1, 31–7, June 1974, 4209657, 10.1007/BF00425265, JOURNAL, Stevens SE, Porter RD, Transformation in Agmenellum quadruplicatum, Proc. Natl. Acad. Sci. U.S.A., 77, 10, 6052–6, October 1980, 16592896, 350211, 10.1073/pnas.77.10.6052, Grigorieva G, Shestakov S. Transformation in the cyanobacterium Synechocystis sp. 6803 FEMS Microbiology Letters 13 (1982) 367-370 Published by Elsevier Biomedical Pressweblink Natural genetic transformation is the genetic alteration of a cell resulting from the direct uptake and incorporation of exogenous DNA from its surroundings. For bacterial transformation to take place, the recipient bacteria must be in a state of competence, which may occur in nature as a response to conditions such as starvation, high cell density or exposure to DNA damaging agents. In chromosomal transformation, homologous transforming DNA can be integrated into the recipient genome by homologous recombination, and this process appears to be an adaptation for repairing DNA damage.JOURNAL, Bernstein H, Bernstein C, Michod RE, Sex in microbial pathogens, Infect. Genet. Evol., 57, 8–25, January 2018, 29111273, 10.1016/j.meegid.2017.10.024,

Classification

{{See also|Bacterial taxonomy}}File:Haeckel arbol bn.png|thumb|upright|Tree of Life in Generelle Morphologie der Organismen (1866). Note the location of the genusNostocNostocHistorically, bacteria were first classified as plants constituting the class Schizomycetes, which along with the Schizophyceae (blue-green algae/Cyanobacteria) formed the phylum Schizophyta,JOURNAL, Von Nägeli C, Caspary R, 1857, Bericht über die Verhandlungen der 33. Versammlung deutscher Naturforscher und Ärzte, gehalten in Bonn von 18 bis 24 September 1857, Report on the Proceedings of the 33rd Meeting of German Natural Scientists and Physicians, held in Bonn, 18 to 24 September 1857, Botanische Zeitung, 15, 749–776, then in the phylum Monera in the kingdom Protista by Haeckel in 1866, comprising Protogens, Protamaeba, Vampyrella, Protomonae, and Vibrio, but not Nostoc and other cyanobacteria, which were classified with algae,BOOK, Haeckel, Ernst, vanc, 1867, Generelle Morphologie der Organismen, Reimer, Berlin, later reclassified as the Prokaryotes by Chatton.JOURNAL, Chatton É, Pansporella perplexa. Réflexions sur la biologie et la phylogénie des protozoaires, Ann. Sci. Nat. Zool., 10-VII, 1–84, 1925, The cyanobacteria were traditionally classified by morphology into five sections, referred to by the numerals I–V. The first three – Chroococcales, Pleurocapsales, and Oscillatoriales – are not supported by phylogenetic studies. The latter two – Nostocales and Stigonematales – are monophyletic, and make up the heterocystous cyanobacteria.JOURNAL, Gugger MF, Hoffmann L, Polyphyly of true branching cyanobacteria (Stigonematales), International Journal of Systematic and Evolutionary Microbiology, 54, Pt 2, 349–357, March 2004, 15023942, 10.1099/ijs.0.02744-0, JOURNAL, Howard-Azzeh M, Shamseer L, Schellhorn HE, Gupta RS, Phylogenetic analysis and molecular signatures defining a monophyletic clade of heterocystous cyanobacteria and identifying its closest relatives, Photosynthesis Research, 122, 2, 171–185, November 2014, 24917519, 10.1007/s11120-014-0020-x, The members of Chroococales are unicellular and usually aggregate in colonies. The classic taxonomic criterion has been the cell morphology and the plane of cell division. In Pleurocapsales, the cells have the ability to form internal spores (baeocytes). The rest of the sections include filamentous species. In Oscillatoriales, the cells are uniseriately arranged and do not form specialized cells (akinetes and heterocysts).JOURNAL,weblink Taxonomic classification of cyanoprokaryotes (cyanobacterial genera) 2014, using a polyphasic approach, Komárek J, Kaštovský J, Mareš J, Johansen JR, Preslia, 86, 295–335, 2014, In Nostocales and Stigonematales, the cells have the ability to develop heterocysts in certain conditions. Stigonematales, unlike Nostocales, include species with truly branched trichomes.Most taxa included in the phylum or division Cyanobacteria have not yet been validly published{{according to whom|date=June 2016}} under the Bacteriological Code, except: The remainder are validly published under the International Code of Nomenclature for algae, fungi, and plants.Formerly, some bacteria, like Beggiatoa, were thought to be colorless Cyanobacteria.BOOK, Pringsheim, Ernst Georg, vanc, Farblose Algen: Ein Beitrag zur Evolutionsforschung, {{google books, y, RYnPAAAAMAAJ, |year=1963|publisher=Gustav Fischer Verlag}}

Earth history

Stromatolites are layered biochemical accretionary structures formed in shallow water by the trapping, binding, and cementation of sedimentary grains by biofilms (microbial mats) of microorganisms, especially cyanobacteria.JOURNAL, Riding, 2007, R., The term stromatolite: towards an essential definition, Lethaia, 32, 4, 321–330, 10.1111/j.1502-3931.1999.tb00550.x, File:Stromatolites.jpg|left|thumb|Stromatolites left behind by cyanobacteria are the oldest known fossils of life on Earth. This one-billion-year-old fossil is from Glacier National Park in Montana.]]During the Precambrian, stromatolite communities of microorganisms grew in most marine and non-marine environments in the photic zone. After the Cambrian explosion of marine animals, grazing on the stromatolite mats by herbivores greatly reduced the occurrence of the stromatolites in marine environments. Since then, they are found mostly in hypersaline conditions where grazing invertebrates cannot live (e.g. Shark Bay, Western Australia). Stromatolites provide ancient records of life on Earth by fossil remains which might date from more than 3.5 Ga ago, but this is disputed.JOURNAL, Patterns In Palaeontology: The first 3 billion years of evolution, Russell J., Garwood, 2012, Palaeontology Online, 2, 11, 1–14,weblink 25 June 2015, {{As of|2010}} the oldest undisputed evidence of cyanobacteria is from 2.1 Ga ago, but there is some evidence for them as far back as 2.7 Ga ago. Oxygen concentrations in the atmosphere remained around or below 1% of today's level until 2.4 Ga ago (the Great Oxygenation Event). The rise in oxygen may have caused a fall in the concentration of atmospheric methane, and triggered the Huronian glaciation from around 2.4 to 2.1 Ga ago. In this way, cyanobacteria may have killed off much of the other bacteria of the time.Lane, Nick (6 February 2010) "First breath: Earth's billion-year struggle for oxygen". New Scientist, pp. 36–9. See accompanying graph as well.Oncolites are sedimentary structures composed of oncoids, which are layered structures formed by cyanobacterial growth. Oncolites are similar to stromatolites, but instead of forming columns, they form approximately spherical structures that were not attached to the underlying substrate as they formed.JOURNAL, Corsetti FA, Awramik SM, Pierce D, A complex microbiota from snowball Earth times: microfossils from the Neoproterozoic Kingston Peak Formation, Death Valley, USA, Proceedings of the National Academy of Sciences of the United States of America, 100, 8, 4399–404, April 2003, 12682298, 153566, 10.1073/pnas.0730560100, 2003PNAS..100.4399C, The oncoids often form around a central nucleus, such as a shell fragment,JOURNAL, Gutschick RC, Perry TG, 1959-11-01, Sappington (Kinderhookian) sponges and their environment [Montana], Journal of Paleontology, 33, 6, 977–985,weblink 2007-06-28,
and a calcium carbonate structure is deposited by encrusting microbes. Oncolites are indicators of warm waters in the photic zone, but are also known in contemporary freshwater environments.Riding, Robert. (1991). Calcareous Algae and Stromatolites, p. 32. Springer-Verlag Press. These structures rarely exceed 10 cm in diameter.

Biotechnology and applications

The unicellular cyanobacterium Synechocystis sp. PCC6803 was the third prokaryote and first photosynthetic organism whose genome was completely sequenced.JOURNAL, 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, 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, DNA Research, 3, 3, 109–36, June 1996, 8905231, 10.1093/dnares/3.3.109, It continues to be an important model organism.JOURNAL, Tabei Y, Okada K, Tsuzuki M, Sll1330 controls the expression of glycolytic genes in Synechocystis sp. PCC 6803, Biochemical and Biophysical Research Communications, 355, 4, 1045–50, April 2007, 17331473, 10.1016/j.bbrc.2007.02.065, Cyanothece ATCC 51142 is an important diazotrophic model organism. The smallest genomes have been found in Prochlorococcus spp. (1.7 Mb)JOURNAL, 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, Genome divergence in two Prochlorococcus ecotypes reflects oceanic niche differentiation, Nature, 424, 6952, 1042–7, August 2003, 12917642, 10.1038/nature01947, 2003Natur.424.1042R, JOURNAL, 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, Genome sequence of the cyanobacterium Prochlorococcus marinus SS120, a nearly minimal oxyphototrophic genome, Proceedings of the National Academy of Sciences of the United States of America, 100, 17, 10020–5, August 2003, 12917486, 187748, 10.1073/pnas.1733211100, 2003PNAS..10010020D, and the largest in Nostoc punctiforme (9 Mb).JOURNAL, Meeks JC, Elhai J, Thiel T, Potts M, Larimer F, Lamerdin J, Predki P, Atlas R, An overview of the genome of Nostoc punctiforme, a multicellular, symbiotic cyanobacterium, Photosynthesis Research, 70, 1, 85–106, 2001, 16228364, 10.1023/A:1013840025518, Those of Calothrix spp. are estimated at 12–15 Mb,JOURNAL, 10.1099/00221287-111-1-73, Genome Size of Cyanobacteria, Journal of General Microbiology, 111, 73–85, 1979, Herdman, M., Janvier, M., Rippka, R., Stanier, R. Y., vanc, as large as yeast.File:OncolitesAlamoBreccia.jpg|thumb|left|Oncolites from the Late Devonian Alamo bolide impactAlamo bolide impactRecent research has suggested the potential application of cyanobacteria to the generation of renewable energy by converting sunlight into electricity. Internal photosynthetic pathways can be coupled to chemical mediators that transfer electrons to external electrodes.JOURNAL, Quintana N, Van der Kooy F, Van de Rhee MD, Voshol GP, Verpoorte R, Renewable energy from Cyanobacteria: energy production optimization by metabolic pathway engineering, Applied Microbiology and Biotechnology, 91, 3, 471–90, August 2011, 21691792, 3136707, 10.1007/s00253-011-3394-0, Currently, efforts are underway to commercialize algae-based fuels such as diesel, gasoline, and jet fuel.Blue green bacteria may help generate 'green' electricity, The Hindu, 21 June 2010Joule wins key patent for GMO cyanobacteria that create fuels from sunlight, CO2, and water. Biofuels Digest (2010-09-14). Retrieved on 2011-04-06.(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.)Researchers from a company called Algenol have cultured genetically modified cyanobacteria in sea water inside a clear plastic enclosure so they first make sugar (pyruvate) from {{CO2}} and the water via photosynthesis. Then, the bacteria secrete ethanol from the cell into the salt water. As the day progresses, and the solar radiation intensifies, ethanol concentrations build up and the ethanol itself evaporates onto the roof of the enclosure. As the sun recedes, evaporated ethanol and water condense into droplets, which run along the plastic walls and into ethanol collectors, from where it is extracted from the enclosure with the water and ethanol separated outside the enclosure. As of March 2013, Algenol was claiming to have tested its technology in Florida and to have achieved yields of 9,000 US gallons per acre per year.WEB, Algenol Biofuels exceeds 9,000 gallons of ethanol per year per, Algenol Biofuels, 6 March 2013,weblink yes,weblink" title="web.archive.org/web/20140420020614weblink">weblink 20 April 2014, dmy-all, This could potentially meet US demands for ethanol in gasoline in 2025, assuming a B30 blend, from an area of around half the size of California's San Bernardino County, requiring less than one-tenth of the area than ethanol from other biomass, such as corn, and only very limited amounts of fresh water.JOURNAL, Jim, Lane, vanc, 25 September 2012,weblink Take it to the Limit: Algenol and rising yields in advanced biofuels, Biofuels Digest, Cyanobacteria may possess the ability to produce substances that could one day serve as anti-inflammatory agents and combat bacterial infections in humans.JOURNAL,weblink Nuisance seaweed found to produce compounds with biomedical potential, Chemistry & Biology, 19, 5, 589–598, 10.1016/j.chembiol.2012.03.014, 22633410, 3361693, 24 May 2012, 1 June 2012, Choi, Hyukjae, Mascuch, Samantha J., Villa, Francisco A., Byrum, Tara, Teasdale, Margaret E., Smith, Jennifer E., Preskitt, Linda B., Rowley, David C., Gerwick, Lena, Gerwick, William H., Spirulina's extracted blue color is used as a natural food coloring in gum and candy.WEB,weblink Federal Register, Volume 78 Issue 156 (Tuesday, August 13, 2013), www.govinfo.gov, Researchers from several space agencies argue that cyanobacteria could be used for producing goods for human consumption in future manned outposts on Mars, by transforming materials available on this planet.JOURNAL, Sustainable life support on Mars – the potential roles of cyanobacteria, International Journal of Astrobiology, 2016, Verseux, Cyprien, Baqué, Mickael, Lehto, Kirsi, de Vera, Jean-Pierre P., Rothschild, Lynn J., Billi, Daniela, vanc, 10.1017/S147355041500021X, 1, 15, 65–92, 2016IJAsB..15...65V,

Health risks

Cyanobacteria can produce neurotoxins, cytotoxins, endotoxins, and hepatotoxins (e.g., the microcystin-producing bacteria genus microcystis), which are collectively known as cyanotoxins.Specific toxins include, anatoxin-a, anatoxin-as, aplysiatoxin, cyanopeptolin, cylindrospermopsin, domoic acid, nodularin R (from Nodularia), neosaxitoxin, and saxitoxin. Cyanobacteria reproduce explosively under certain conditions. This results in algal blooms, which can become harmful to other species, and pose a danger to humans and animals, if the cyanobacteria involved produce toxins. Several cases of human poisoning have been documented, but a lack of knowledge prevents an accurate assessment of the risks.JOURNAL, Thébault L, Lesne J, Boutin JP, [Cyanobacteria, their toxins and health risks], Medecine Tropicale, 55, 4, 375–380, 1995, 8830224, weblink" title="web.archive.org/web/20080530024705weblink">Blue-Green Algae (Cyanobacteria) and their Toxins. Hc-sc.gc.ca (2013-01-30). Retrieved on 2014-04-19.Harmful Bloom in Lake Atitlán, Guatemala from NASA Earth Observatory, retrieved on 9 January 2010.Recent studies suggest that significant exposure to high levels of cyanobacteria producing toxins such as BMAA can cause amyotrophic lateral sclerosis (ALS).People living within half a mile of cyanobacterially contaminated lakes have had a 2.3 times greater risk of developing ALS than the rest of the population; people around New Hampshire's Lake Mascoma had an up to 25 times greater risk of ALS than the expected incidence.JOURNAL, Caller TA, Doolin JW, Haney JF, Murby AJ, West KG, Farrar HE, Ball A, Harris BT, Stommel EW, A cluster of amyotrophic lateral sclerosis in New Hampshire: a possible role for toxic cyanobacteria blooms, Amyotrophic Lateral Sclerosis, 10 Suppl 2, 101–108, 2009, 19929741, 10.3109/17482960903278485, BMAA from desert crusts found throughout Qatar might have contributed to higher rates of ALS in Gulf War veterans.JOURNAL, Cox PA, Richer R, Metcalf JS, Banack SA, Codd GA, Bradley WG, Cyanobacteria and BMAA exposure from desert dust: a possible link to sporadic ALS among Gulf War veterans, Amyotrophic Lateral Sclerosis, 10 Suppl 2, 109–117, 2009, 19929742, 10.3109/17482960903286066,

Chemical control

Several chemicals can eliminate cyanobacterial blooms from water-based systems. They include: calcium hypochlorite, copper sulphate, cupricide, and simazine.WEB, Main, D.C., vanc, Toxic Algae Blooms, Veterinary Pathologist, South Perth, agric.wa.gov.au, 2006,weblink 18 November 2014, The calcium hypochlorite amount needed varies depending on the cyanobacteria bloom, and treatment is needed periodically. According to the Department of Agriculture Australia, a rate of 12 g of 70% material in 1000 l of water is often effective to treat a bloom. Copper sulfate is also used commonly, but no longer recommended by the Australian Department of Agriculture, as it kills livestock, crustaceans, and fish. Cupricide is a chelated copper product that eliminates blooms with lower toxicity risks than copper sulfate. Dosage recommendations vary from 190 ml to 4.8 l per 1000 m2. Ferric alum treatments at the rate of 50 mg/l will reduce algae blooms.JOURNAL, May V, Baker H, 1978, Reduction of toxic algae in farm dams by ferric alum, Techn. Bull., 19, 1–16, Simazine, which is also a herbicide, will continue to kill blooms for several days after an application. Simazine is marketed at different strengths (25, 50, and 90%), the recommended amount needed for one cubic meter of water per product is 25% product 8 ml; 50% product 4 ml; or 90% product 2.2 ml.

Dietary supplementation

File:Spirulina tablets.jpg|thumb|right|upright|Spirulina tablets]]Some cyanobacteria are sold as food, notably Aphanizomenon flos-aquae and Arthrospira platensis (Spirulina).JOURNAL, Spolaore P, Joannis-Cassan C, Duran E, Isambert A, Commercial applications of microalgae, Journal of Bioscience and Bioengineering, 101, 2, 87–96, February 2006, 16569602, 10.1263/jbb.101.87, Despite the associated toxins which many of the members of this phylum produce, some microalgae also contain substances of high biological value, such as polyunsaturated fatty acids, amino acids, proteins, pigments, antioxidants, vitamins, and minerals.JOURNAL, Christaki E, Florou-Paneri P, Bonos E, Microalgae: a novel ingredient in nutrition, International Journal of Food Sciences and Nutrition, 62, 8, 794–799, December 2011, 21574818, 10.3109/09637486.2011.582460, Edible blue-green algae reduce the production of pro-inflammatory cytokines by inhibiting NF-κB pathway in macrophages and splenocytes.JOURNAL, Ku CS, Pham TX, Park Y, Kim B, Shin MS, Kang I, Lee J, Edible blue-green algae reduce the production of pro-inflammatory cytokines by inhibiting NF-κB pathway in macrophages and splenocytes, Biochimica et Biophysica Acta, 1830, 4, 2981–2988, April 2013, 23357040, 3594481, 10.1016/j.bbagen.2013.01.018, Sulfate polysaccharides exhibit immunomodulatory, antitumor, antithrombotic, anticoagulant, anti-mutagenic, anti-inflammatory, antimicrobial, and even antiviral activity against HIV, herpes, and hepatitis.BOOK, Mišurcová L, Škrovánková S, Samek D, Ambrožová J, Machů L, Health benefits of algal polysaccharides in human nutrition, Advances in Food and Nutrition Research, 66, 75–145, 2012, 22909979, 10.1016/B978-0-12-394597-6.00003-3, 978-0-12-394597-6,

See also

{{div col|colwidth=18em}} {{div col end}}

References

{{Reflist|30em}}
Attribution
{{CC-notice|cc=by2.5|url=https://journals.plos.org/plosone/article?id=10.1371/journal.pone.0010821}}

Further reading

  • BOOK, Gillian, Cribbs, vanc, Nature's Superfood: the Blue-Green Algae Revolution, Newleaf, 978-0-7522-0569-4, 1997, first,
  • BOOK, Marshall, Savage, vanc, Marshall Savage, The Millennial Project: Colonizing the Galaxy in Eight Easy Steps, Little Brown & Co, 978-0-316-77163-4, 1994,
  • BOOK, Fogg GE, Stewart WD, Fay P, Walsby AE, 1973, The Blue-green Algae, Academic Press, London and New York, 978-0-12-261650-1,
  • "Architects of the earth's atmosphere", Introduction to the Cyanobacteria, University of California, Berkeley, 3 February 2006.
    • JOURNAL, 10.1080/09670262.2010.492914, Pentecost A, Franke U, 2010, Photosynthesis and calcification of the stromatolitic freshwater cyanobacterium Rivularia, Eur. J. Phycol., 45, 4, 345–353,
  • BOOK, Whitton BA, Potts M, 2000, The Ecology of Cyanobacteria: their Diversity in Time and Space, Springer, 978-0-7923-4735-4,
  • WEB,weblink From Micro-Algae to Blue Oil, ParisTech Review, December 2011, 2 March 2012,weblink" title="web.archive.org/web/20160417030653weblink">weblink 17 April 2016, yes,

External links

{{Commons category|Cyanobacteria}} {{Plankton}}{{microorganisms}}{{Bacteria classification}}{{Life on Earth}}{{Taxonbar|from=Q93315}}

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Eastern Philosophy
History of Philosophy
GETWIKI 09 MAY 2016
GETWIKI 18 OCT 2015
M.R.M. Parrott
Biographies
GETWIKI 20 AUG 2014
GETWIKI 19 AUG 2014
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