Cyanobacteria
Cyanobacteria Temporal range:
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Microscope image of Cylindrospermum, a filamentous genus of cyanobacteria | |
Scientific classification | |
Domain: | Bacteria |
Clade: | Terrabacteria |
Clade: | Cyanobacteria-Melainabacteria group |
Phylum: | Cyanobacteria Stanier, 1973 |
Class: | Cyanophyceae |
Orders[3] | |
As of 2014[update] the taxonomy was under revision[1][2]
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Synonyms | |
List
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Cyanobacteria (
Cyanobacteria are probably the most numerous taxon to have ever existed on Earth and the first organisms known to have produced oxygen.[11] 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 Oxidation Event and the "rusting of the Earth",[12] which dramatically changed the composition of life forms on Earth.[13]
Cyanobacteria use
Sericytochromatia, the proposed name of the paraphyletic and most basal group, is the ancestor of both the non-photosynthetic group Melainabacteria and the photosynthetic cyanobacteria, also called Oxyphotobacteria.[16]
The cyanobacteria
Overview
Cyanobacteria are a very large and diverse phylum of
Cyanobacteria are globally widespread photosynthetic prokaryotes and are major contributors to global biogeochemical cycles.[25] They are the only oxygenic photosynthetic prokaryotes, and prosper in diverse and extreme habitats.[26] They are among the oldest organisms on Earth with fossil records dating back at least 2.1 billion years.[27] Since then, cyanobacteria have been essential players in the Earth's ecosystems. Planktonic cyanobacteria are a fundamental component of marine food webs and are major contributors to global carbon and nitrogen fluxes.[28][29] Some cyanobacteria form harmful algal blooms causing the disruption of aquatic ecosystem services and intoxication of wildlife and humans by the production of powerful toxins (cyanotoxins) such as microcystins, saxitoxin, and cylindrospermopsin.[30][31] Nowadays, cyanobacterial blooms pose a serious threat to aquatic environments and public health, and are increasing in frequency and magnitude globally.[32][25]
Cyanobacteria are ubiquitous in marine environments and play important roles as
Within the cyanobacteria, only a few lineages colonized the open ocean:
Marine cyanobacteria include the smallest known photosynthetic organisms. The smallest of all,
Morphology
Cyanobacteria are variable in morphology, ranging from
Many cyanobacteria form motile filaments of cells, called hormogonia, that travel away from the main biomass to bud and form new colonies elsewhere.[51][52] 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.
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
- Thick-walled heterocysts – which contain the enzyme nitrogenase vital for nitrogen fixation[54][55][56] in an anaerobic environment due to its sensitivity to oxygen.[56]
Each individual cell (each single cyanobacterium) typically has a thick, gelatinous
Nitrogen fixation
Some cyanobacteria can fix atmospheric
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.[60] Cyanobacteria such as Anabaena (a symbiont of the aquatic fern Azolla) can provide rice plantations with biofertilizer.[61]
Photosynthesis
Carbon fixation
Cyanobacteria use the energy of
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.[66] 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.[67]
While most of the high-energy
Respiration
Respiration in cyanobacteria can occur in the thylakoid membrane alongside photosynthesis,[69] 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.[69] Cyanobacteria use electrons from succinate dehydrogenase rather than from NADPH for respiration.[69]
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.[70]
Electron transport chain
Many cyanobacteria are able to reduce nitrogen and carbon dioxide under
Attached to the thylakoid membrane, phycobilisomes act as light-harvesting antennae for the photosystems.[72] The phycobilisome components (phycobiliproteins) are responsible for the blue-green pigmentation of most cyanobacteria.[73] 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.[74][75] In green light, the cells accumulate more phycoerythrin, which absorbs green light, whereas in red light they produce more phycocyanin which absorbs red. Thus, these bacteria can change from brick-red to bright blue-green depending on whether they are exposed to green light or to red light.[76] 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.[77][78]
Metabolism
In general, photosynthesis in cyanobacteria uses water as an electron donor and produces oxygen as a byproduct, though some may also use hydrogen sulfide[79] a process which occurs among other photosynthetic bacteria such as the purple sulfur bacteria.
There are some groups capable of
Ecology
Cyanobacteria can be found in almost every terrestrial and
Aquatic cyanobacteria are known for their extensive and highly visible
Cyanobacterial growth is favoured in ponds and lakes where waters are calm and have little turbulent mixing.
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
Cyanobacteria have been found to play an important role in terrestrial habitats and organism communities. It has been widely reported that cyanobacteria soil crusts help to stabilize soil to prevent erosion and retain water.[96] 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.[97] M. vaginatus also makes a significant contribution to the cohesion of biological soil crust.[98]
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.[99] 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
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 Falconer[100]
Cyanobionts
Some cyanobacteria, the so-called
The relationships between
Collective behaviour
Some cyanobacteria – even single-celled ones – show striking collective behaviours and form colonies (or blooms) that can float on water and have important ecological roles. For instance, billions of years ago, communities of marine Paleoproterozoic cyanobacteria could have helped create the biosphere as we know it by burying carbon compounds and allowing the initial build-up of oxygen in the atmosphere.[124] On the other hand, toxic cyanobacterial blooms are an increasing issue for society, as their toxins can be harmful to animals.[32] Extreme blooms can also deplete water of oxygen and reduce the penetration of sunlight and visibility, thereby compromising the feeding and mating behaviour of light-reliant species.[123]
As shown in the diagram on the right, bacteria can stay in suspension as individual cells, adhere collectively to surfaces to form biofilms, passively sediment, or flocculate to form suspended aggregates. Cyanobacteria are able to produce sulphated polysaccharides (yellow haze surrounding clumps of cells) that enable them to form floating aggregates. In 2021, Maeda et al. discovered that oxygen produced by cyanobacteria becomes trapped in the network of polysaccharides and cells, enabling the microorganisms to form buoyant blooms.[125] It is thought that specific protein fibres known as pili (represented as lines radiating from the cells) may act as an additional way to link cells to each other or onto surfaces. Some cyanobacteria also use sophisticated intracellular gas vesicles as floatation aids.[123]
The diagram on the left above shows a proposed model of microbial distribution, spatial organization, carbon and O2 cycling in clumps and adjacent areas. (a) Clumps contain denser cyanobacterial filaments and heterotrophic microbes. The initial differences in density depend on cyanobacterial motility and can be established over short timescales. Darker blue color outside of the clump indicates higher oxygen concentrations in areas adjacent to clumps. Oxic media increase the reversal frequencies of any filaments that begin to leave the clumps, thereby reducing the net migration away from the clump. This enables the persistence of the initial clumps over short timescales; (b) Spatial coupling between photosynthesis and respiration in clumps. Oxygen produced by cyanobacteria diffuses into the overlying medium or is used for aerobic respiration.
It has been unclear why and how cyanobacteria form communities. Aggregation must divert resources away from the core business of making more cyanobacteria, as it generally involves the production of copious quantities of extracellular material. In addition, cells in the centre of dense aggregates can also suffer from both shading and shortage of nutrients.[127][128] So, what advantage does this communal life bring for cyanobacteria?[123]
New insights into how cyanobacteria form blooms have come from a 2021 study on the cyanobacterium
Previous studies on Synechocystis have shown
The bubble flotation mechanism identified by Maeda et al. joins a range of known strategies that enable cyanobacteria to control their buoyancy, such as using gas vesicles or accumulating carbohydrate ballasts.[134] Type IV pili on their own could also control the position of marine cyanobacteria in the water column by regulating viscous drag.[135] Extracellular polysaccharide appears to be a multipurpose asset for cyanobacteria, from floatation device to food storage, defence mechanism and mobility aid.[132][123]
Cellular death
One of the most critical processes determining cyanobacterial eco-physiology is
Cyanophages
Cyanophages are viruses that infect cyanobacteria. Cyanophages can be found in both freshwater and marine environments.[145] Marine and freshwater cyanophages have icosahedral heads, which contain double-stranded DNA, attached to a tail by connector proteins.[146] The size of the head and tail vary among species of cyanophages. Cyanophages, like other bacteriophages, rely on Brownian motion to collide with bacteria, and then use receptor binding proteins to recognize cell surface proteins, which leads to adherence. Viruses with contractile tails then rely on receptors found on their tails to recognize highly conserved proteins on the surface of the host cell.[147]
Cyanophages infect a wide range of cyanobacteria and are key regulators of the cyanobacterial populations in aquatic environments, and may aid in the prevention of cyanobacterial blooms in freshwater and marine ecosystems. These blooms can pose a danger to humans and other animals, particularly in
The first cyanophage, LPP-1, was discovered in 1963.[149] Cyanophages are classified within the bacteriophage families Myoviridae (e.g. AS-1, N-1), Podoviridae (e.g. LPP-1) and Siphoviridae (e.g. S-1).[149]
Movement
It has long been known that
that does not involve flagellar motion.Many species of cyanobacteria are capable of gliding.
Cyanobacteria have strict light requirements. Too little light can result in insufficient energy production, and in some species may cause the cells to resort to heterotrophic respiration.[21] Too much light can inhibit the cells, decrease photosynthesis efficiency and cause damage by bleaching. UV radiation is especially deadly for cyanobacteria, with normal solar levels being significantly detrimental for these microorganisms in some cases.[20][162][22]
Filamentous cyanobacteria that live in microbial mats often migrate vertically and horizontally within the mat in order to find an optimal niche that balances their light requirements for photosynthesis against their sensitivity to photodamage. For example, the filamentous cyanobacteria Oscillatoria sp. and Spirulina subsalsa found in the hypersaline benthic mats of Guerrero Negro, Mexico migrate downwards into the lower layers during the day in order to escape the intense sunlight and then rise to the surface at dusk.[163] In contrast, the population of Microcoleus chthonoplastes found in hypersaline mats in Camargue, France migrate to the upper layer of the mat during the day and are spread homogeneously through the mat at night.[164] An in vitro experiment using Phormidium uncinatum also demonstrated this species' tendency to migrate in order to avoid damaging radiation.[20][162] These migrations are usually the result of some sort of photomovement, although other forms of taxis can also play a role.[165][22]
Photomovement – the modulation of cell movement as a function of the incident light – is employed by the cyanobacteria as a means to find optimal light conditions in their environment. There are three types of photomovement: photokinesis, phototaxis and photophobic responses.[166][167][168][22]
Photokinetic microorganisms modulate their gliding speed according to the incident light intensity. For example, the speed with which Phormidium autumnale glides increases linearly with the incident light intensity.[169][22]
Phototactic microorganisms move according to the direction of the light within the environment, such that positively phototactic species will tend to move roughly parallel to the light and towards the light source. Species such as Phormidium uncinatum cannot steer directly towards the light, but rely on random collisions to orient themselves in the right direction, after which they tend to move more towards the light source. Others, such as Anabaena variabilis, can steer by bending the trichome.[170][22]
Finally, photophobic microorganisms respond to spatial and temporal light gradients. A step-up photophobic reaction occurs when an organism enters a brighter area field from a darker one and then reverses direction, thus avoiding the bright light. The opposite reaction, called a step-down reaction, occurs when an organism enters a dark area from a bright area and then reverses direction, thus remaining in the light.[22]
Evolution
Earth history
million years ago) |
During the
One former classification scheme of cyanobacterial fossils divided them into the
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Stromatolitesleft behind by cyanobacteria are the oldest known fossils of life on Earth. This fossil is one billion years old.
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Oncolitic limestone formed from successive layers of calcium carbonate precipitated by cyanobacteria
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Late Devonian Alamo bolide impactin Nevada
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Cyanobacterial remains of an annulated tubular microfossil Oscillatoriopsis longa [180]
Scale bar: 100 μm
Origin of photosynthesis
As far as we can tell,
Cyanobacteria remained the principal
Origin of chloroplasts
Primary chloroplasts are cell organelles found in some
The morphological similarity between chloroplasts and cyanobacteria was first reported by German botanist Andreas Franz Wilhelm Schimper in the 19th century[192] Chloroplasts are only found in plants and algae,[193] thus paving the way for Russian biologist Konstantin Mereschkowski to suggest in 1905 the symbiogenic origin of the plastid.[194] Lynn Margulis brought this hypothesis back to attention more than 60 years later[195] but the idea did not become fully accepted until supplementary data started to accumulate. The cyanobacterial origin of plastids is now supported by various pieces of phylogenetic,[196][188][191] genomic,[197] biochemical[198][199] and structural evidence.[200] The description of another independent and more recent primary endosymbiosis event between a cyanobacterium and a separate eukaryote lineage (the rhizarian Paulinella chromatophora) also gives credibility to the endosymbiotic origin of the plastids.[201]
In addition to this primary endosymbiosis, many eukaryotic lineages have been subject to
Marine origins
Part of a series on |
Plankton |
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Cyanobacteria have fundamentally transformed the geochemistry of the planet.[210][207] Multiple lines of geochemical evidence support the occurrence of intervals of profound global environmental change at the beginning and end of the Proterozoic (2,500–542 Mya).[211] [212][213] While it is widely accepted that the presence of molecular oxygen in the early fossil record was the result of cyanobacteria activity, little is known about how cyanobacteria evolution (e.g., habitat preference) may have contributed to changes in biogeochemical cycles through Earth history. Geochemical evidence has indicated that there was a first step-increase in the oxygenation of the Earth's surface, which is known as the Great Oxidation Event (GOE), in the early Paleoproterozoic (2,500–1,600 Mya).[210][207] A second but much steeper increase in oxygen levels, known as the Neoproterozoic Oxygenation Event (NOE),[212][81][214] occurred at around 800 to 500 Mya.[213][215] Recent chromium isotope data point to low levels of atmospheric oxygen in the Earth's surface during the mid-Proterozoic,[211] which is consistent with the late evolution of marine planktonic cyanobacteria during the Cryogenian;[216] both types of evidence help explain the late emergence and diversification of animals.[217][43]
Understanding the evolution of planktonic cyanobacteria is important because their origin fundamentally transformed the
Genetics
Cyanobacteria are capable of natural genetic transformation.[220][221][222] 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.[223]
DNA repair
Cyanobacteria are challenged by environmental stresses and internally generated reactive oxygen species that cause DNA damage. Cyanobacteria possess numerous E. coli-like DNA repair genes.[224] 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.[224]
Classification
Phylogeny
16S rRNA based | GTDB 08-RS214 by Genome Taxonomy Database[228][229][230] | ||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||
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Taxonomy
Historically, bacteria were first classified as plants constituting the class Schizomycetes, which along with the Schizophyceae (blue-green algae/Cyanobacteria) formed the phylum Schizophyta,
The cyanobacteria were traditionally classified by morphology into five sections, referred to by the numerals I–V. The first three –
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).[236] In Nostocales and Stigonematales, the cells have the ability to develop heterocysts in certain conditions. Stigonematales, unlike Nostocales, include species with truly branched trichomes.[234]
Most taxa included in the phylum or division Cyanobacteria have not yet been validly published under The International Code of Nomenclature of Prokaryotes (ICNP) except:
- The classes Gloeobacteria
- The orders Stigonematales
- The families Prochloraceae and Prochlorotrichaceae
- The genera Prochlorothrix
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.[237]
The currently accepted taxonomy is based on the List of Prokaryotic names with Standing in Nomenclature (LPSN)[238] and National Center for Biotechnology Information (NCBI).[239] Class "Cyanobacteriia"
- Subclass "Gloeobacteria" Cavalier-Smith 2002
- GloeobacteralesCavalier-Smith 2002
- Subclass "Phycobacteria" Cavalier-Smith 2002
- Acaryochloridales Miyashita et al. 2003 ex Strunecký & Mareš 2022 [incl. Thermosynechococcales]
- Aegeococcales Strunecký & Mareš 2022
- "Elainellales"
- "Eurycoccales"
- Geitlerinematales Strunecký & Mareš 2022
- Gloeoemargaritales Moreira et al. 2016
- "Leptolyngbyales" Strunecký & Mareš 2022
- Nodosilineales Strunecký & Mareš 2022
- Oculatellales Strunecký & Mareš 2022
- "Phormidesmiales"
- Prochlorococcaceae Komárek & Strunecky 2020 {"PCC-6307"}
- Pseudanabaenales Hoffmann, Komárek & Kastovsky 2005
- "Pseudophormidiales"
- Thermostichales Komárek & Strunecký 2020
- Synechococcophycidae Hoffmann, Komárek & Kastovsky 2005
- "Limnotrichales"
- Prochlorotrichales Strunecký & Mareš 2022 (PCC-9006)
- Synechococcales Hoffmann, Komárek & Kastovsky 2005
- Nostocophycidae Hoffmann, Komárek & Kastovsky 2005
- Stigonematales)
Relation to humans
Biotechnology
The unicellular cyanobacterium Synechocystis sp. PCC6803 was the third prokaryote and first photosynthetic organism whose genome was completely sequenced.[240] It continues to be an important model organism.[241] Cyanothece ATCC 51142 is an important diazotrophic model organism. The smallest genomes have been found in Prochlorococcus spp. (1.7 Mb)[242][243] and the largest in Nostoc punctiforme (9 Mb).[144] Those of Calothrix spp. are estimated at 12–15 Mb,[244] as large as yeast.
Recent research has suggested the potential application of cyanobacteria to the generation of
Cyanobacteria may possess the ability to produce substances that could one day serve as anti-inflammatory agents and combat bacterial infections in humans.[252] Cyanobacteria's photosynthetic output of sugar and oxygen has been demonstrated to have therapeutic value in rats with heart attacks.[253] 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.[254]
Spirulina's extracted blue color is used as a natural food coloring.[255]
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.[256]
Human nutrition
Some cyanobacteria are sold as food, notably Arthrospira platensis (Spirulina) and others (Aphanizomenon flos-aquae).[257]
Some microalgae contain substances of high biological value, such as polyunsaturated fatty acids, amino acids, proteins, pigments, antioxidants, vitamins, and minerals.[258] Edible blue-green algae reduce the production of pro-inflammatory cytokines by inhibiting NF-κB pathway in macrophages and splenocytes.[259] Sulfate polysaccharides exhibit immunomodulatory, antitumor, antithrombotic, anticoagulant, anti-mutagenic, anti-inflammatory, antimicrobial, and even antiviral activity against HIV, herpes, and hepatitis.[260]
Health risks
Some cyanobacteria can produce
Specific toxins include
Recent studies suggest that significant exposure to high levels of cyanobacteria producing toxins such as
Chemical control
Several chemicals can eliminate cyanobacterial blooms from smaller water-based systems such as swimming pools. They include calcium hypochlorite, copper sulphate, Cupricide (chelated copper), and simazine.[268] 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.[268] Copper sulfate is also used commonly, but no longer recommended by the Australian Department of Agriculture, as it kills livestock, crustaceans, and fish.[268] 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.[268] Ferric alum treatments at the rate of 50 mg/L will reduce algae blooms.[268][269] 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.[268]
Climate change
The capacity of the harmful cyanobacterial genus
Gallery
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Cyanobacteria activity turns Coatepeque Caldera lake a turquoise color
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Cyanobacterial bloom near Fiji
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Cyanobacteria inLake Köyliö.
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Video – Oscillatoria and Gleocapsa – with oscillatory movement as filaments of Oscillatoria orient towards light
See also
- Archean Eon
- Bacterial phyla, other major lineages of Bacteria
- Biodiesel
- Cyanobiont
- Endosymbiotic theory
- Geological history of oxygen
- Hypolith
Notes
- ^ Botanists restrict the name algae to protist eukaryotes, which does not extend to cyanobacteria, which are prokaryotes. However, the common name blue-green algae continues to be used synonymously with cyanobacteria outside of the biological sciences.
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Attribution
This article incorporates text available under the CC BY 2.5 license.
Further reading
- Gillian C (1997). Nature's Superfood: the Blue-Green Algae Revolution (first ed.). Newleaf. ISBN 978-0-7522-0569-4.
- ISBN 978-0-316-77163-4.
- Fogg GE, Stewart WD, Fay P, Walsby AE (1973). The Blue-green Algae. London and New York: ISBN 978-0-12-261650-1.
- "Architects of the earth's atmosphere, Introduction to the Cyanobacteria". University of California, Berkeley. 3 February 2006.
- Whitton BA (2002). "Phylum Cyanophyta (Cyanobacteria)". The Freshwater Algal Flora of the British Isles. Cambridge: ISBN 978-0-521-77051-4.
- Pentecost A, Franke U (2010). "Photosynthesis and calcification of the stromatolitic freshwater cyanobacterium Rivularia". European Journal of Phycology. 45 (4): 345–53. .
- Whitton BA, Potts M, eds. (2000). The Ecology of Cyanobacteria: their Diversity in Time and Space. Springer. ISBN 978-0-7923-4735-4.
- "From Micro-Algae to Blue Oil". ParisTech Review. December 2011. Archived from the original on 17 April 2016. Retrieved 2 March 2012.