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Cyanobacteria

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91-446: As of 2014 the taxonomy was under revision Cyanobacteria ( / s aɪ ˌ æ n oʊ b æ k ˈ t ɪər i . ə / ), also called Cyanobacteriota or Cyanophyta , are a phylum of autotrophic gram-negative bacteria that can obtain biological energy via oxygenic photosynthesis . The name "cyanobacteria" (from Ancient Greek κύανος ( kúanos )  'blue') refers to their bluish green ( cyan ) color, which forms

182-433: A 2021 study on the cyanobacterium Synechocystis . These use a set of genes that regulate the production and export of sulphated polysaccharides , chains of sugar molecules modified with sulphate groups that can often be found in marine algae and animal tissue. Many bacteria generate extracellular polysaccharides, but sulphated ones have only been seen in cyanobacteria. In Synechocystis these sulphated polysaccharide help

273-453: A certain degree of evolutionary relatedness (the phylogenetic definition). Attempting to define a level of the Linnean hierarchy without referring to (evolutionary) relatedness is unsatisfactory, but a phenetic definition is useful when addressing questions of a morphological nature—such as how successful different body plans were. The most important objective measure in the above definitions

364-407: A character unique to a sub-set of the crown group. Furthermore, organisms in the stem group of a phylum can possess the "body plan" of the phylum without all the characteristics necessary to fall within it. This weakens the idea that each of the phyla represents a distinct body plan. A classification using this definition may be strongly affected by the chance survival of rare groups, which can make

455-415: 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. M. vaginatus also makes a significant contribution to the cohesion of biological soil crust . Some of these organisms contribute significantly to global ecology and the oxygen cycle . The tiny marine cyanobacterium Prochlorococcus

546-457: 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. Attached to the thylakoid membrane, phycobilisomes act as light-harvesting antennae for

637-596: A group containing Viridiplantae and the algal Rhodophyta and Glaucophyta divisions. The definition and classification of plants at the division level also varies from source to source, and has changed progressively in recent years. Thus some sources place horsetails in division Arthrophyta and ferns in division Monilophyta, while others place them both in Monilophyta, as shown below. The division Pinophyta may be used for all gymnosperms (i.e. including cycads, ginkgos and gnetophytes), or for conifers alone as below. Since

728-404: A key role in developmental processes, such as akinete and heterocyst differentiation, as well as strategy for population survival. Cyanophages are viruses that infect cyanobacteria. Cyanophages can be found in both freshwater and marine environments. Marine and freshwater cyanophages have icosahedral heads, which contain double-stranded DNA, attached to a tail by connector proteins. The size of

819-400: A phylum based on body plan has been proposed by paleontologists Graham Budd and Sören Jensen (as Haeckel had done a century earlier). The definition was posited because extinct organisms are hardest to classify: they can be offshoots that diverged from a phylum's line before the characters that define the modern phylum were all acquired. By Budd and Jensen's definition, a phylum is defined by

910-471: A phylum much more diverse than it would be otherwise. Total numbers are estimates; figures from different authors vary wildly, not least because some are based on described species, some on extrapolations to numbers of undescribed species. For instance, around 25,000–27,000 species of nematodes have been described, while published estimates of the total number of nematode species include 10,000–20,000; 500,000; 10 million; and 100 million. The kingdom Plantae

1001-689: A phylum, other phylum-level ranks appear, such as the case of Bacillariophyta (diatoms) within Ochrophyta . These differences became irrelevant after the adoption of a cladistic approach by the ISP, where taxonomic ranks are excluded from the classifications after being considered superfluous and unstable. Many authors prefer this usage, which lead to the Chromista-Protozoa scheme becoming obsolete. Currently there are 40 bacterial phyla (not including " Cyanobacteria ") that have been validly published according to

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1092-551: 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 "CO 2 concentrating mechanism" to aid in the acquisition of inorganic carbon (CO 2 or bicarbonate ). Among the more specific strategies is the widespread prevalence of the bacterial microcompartments known as carboxysomes , which co-operate with active transporters of CO 2 and bicarbonate, in order to accumulate bicarbonate into

1183-442: A proposed model of microbial distribution, spatial organization, carbon and O 2 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

1274-492: A range of environments, including soil, water, and animal habitats. They can be often be found in the gut of humans and in the respiratory tract, oral environments, and skin surface, though rarely. Melainabacteria is often found in natural environments such as groundwater aquifers and lake sediment, as well as soil and bioreactors. Melainabacteria are also found in the aphotic zone of aquatic environments such as lake sediment and aquifers. Cyanobacteria bloom in freshwater systems as

1365-410: A range of toxins known as cyanotoxins that can cause harmful health effects in humans and animals. Cyanobacteria are a very large and diverse phylum of photosynthetic prokaryotes . They are defined by their unique combination of pigments and their ability to perform oxygenic photosynthesis . They often live in colonial aggregates that can take on a multitude of forms. Of particular interest are

1456-552: A response to biotic and abiotic stresses. However, cell death research in cyanobacteria is a relatively young field and understanding of the underlying mechanisms and molecular machinery underpinning this fundamental process remains largely elusive. However, reports on cell death of marine and freshwater cyanobacteria indicate this process has major implications for the ecology of microbial communities/ Different forms of cell demise have been observed in cyanobacteria under several stressful conditions, and cell death has been suggested to play

1547-495: A result of excess nutrients and high temperatures, resulting in a scum on the water surface that resembles spilled paint. Because Melainabacteria and Cyanobacteria are related, it has raised concern because Melainabacteria thrive in groundwater systems. The genomes of Melainabacteria were found to be bigger when found in aquifer systems and algal cultivation ponds than when in the mammalian gut environment. The Great Oxygenation Event (GOE) that occurred 2.4 billion years ago altered

1638-484: A serious threat to aquatic environments and public health, and are increasing in frequency and magnitude globally. Cyanobacteria are ubiquitous in marine environments and play important roles as primary producers . They are part of the marine phytoplankton , which currently contributes almost half of the Earth's total primary production. About 25% of the global marine primary production is contributed by cyanobacteria. Within

1729-401: A set of characters shared by all its living representatives. This approach brings some small problems—for instance, ancestral characters common to most members of a phylum may have been lost by some members. Also, this definition is based on an arbitrary point of time: the present. However, as it is character based, it is easy to apply to the fossil record. A greater problem is that it relies on

1820-401: A subjective decision about which groups of organisms should be considered as phyla. The approach is useful because it makes it easy to classify extinct organisms as " stem groups " to the phyla with which they bear the most resemblance, based only on the taxonomically important similarities. However, proving that a fossil belongs to the crown group of a phylum is difficult, as it must display

1911-496: A symbiotic relationship with plants or lichen -forming fungi (as in the lichen genus Peltigera ). Cyanobacteria are globally widespread photosynthetic prokaryotes and are major contributors to global biogeochemical cycles . They are the only oxygenic photosynthetic prokaryotes, and prosper in diverse and extreme habitats. They are among the oldest organisms on Earth with fossil records dating back at least 2.1 billion years. Since then, cyanobacteria have been essential players in

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2002-416: A waving motion; the filament oscillates back and forth. In water columns, some cyanobacteria float by forming gas vesicles , as in archaea . These vesicles are not organelles as such. They are not bounded by lipid membranes , but by a protein sheath. Some cyanobacteria can fix atmospheric nitrogen in anaerobic conditions by means of specialized cells called heterocysts . Heterocysts may also form under

2093-406: A weaker cell in a filament, called a necridium. Some filamentous species can differentiate into several different cell types: Each individual cell (each single cyanobacterium) typically has a thick, gelatinous cell wall . They lack flagella , but hormogonia of some species can move about by gliding along surfaces. Many of the multicellular filamentous forms of Oscillatoria are capable of

2184-577: 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 eutrophic freshwater lakes. Infection by these viruses is highly prevalent in cells belonging to Synechococcus spp. in marine environments, where up to 5% of cells belonging to marine cyanobacterial cells have been reported to contain mature phage particles. The first cyanophage, LPP-1 ,

2275-467: Is a paraphyletic taxon, which is less acceptable to present-day biologists than in the past. Proposals have been made to divide it among several new kingdoms, such as Protozoa and Chromista in the Cavalier-Smith system . Protist taxonomy has long been unstable, with different approaches and definitions resulting in many competing classification schemes. Many of the phyla listed below are used by

2366-417: 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

2457-405: Is defined in various ways by different biologists (see Current definitions of Plantae ). All definitions include the living embryophytes (land plants), to which may be added the two green algae divisions, Chlorophyta and Charophyta , to form the clade Viridiplantae . The table below follows the influential (though contentious) Cavalier-Smith system in equating "Plantae" with Archaeplastida ,

2548-726: Is favoured in ponds and lakes where waters are calm and have little turbulent mixing. Their lifecycles 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 favoured 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

2639-472: Is generally included in kingdom Fungi, though its exact relations remain uncertain, and it is considered a protozoan by the International Society of Protistologists (see Protista , below). Molecular analysis of Zygomycota has found it to be polyphyletic (its members do not share an immediate ancestor), which is considered undesirable by many biologists. Accordingly, there is a proposal to abolish

2730-486: Is known regarding the nature (e.g., genetic diversity, host or cyanobiont specificity, and cyanobiont seasonality) of the symbiosis involved, particularly in relation to dinoflagellate host. 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

2821-481: Is later used to make amino acids and proteins. Marine picocyanobacteria ( Prochlorococcus and Synechococcus ) numerically dominate most phytoplankton assemblages in modern oceans, contributing importantly to primary productivity. While some planktonic cyanobacteria are unicellular and free living cells (e.g., Crocosphaera , Prochlorococcus , Synechococcus ); others have established symbiotic relationships with haptophyte algae , such as coccolithophores . Amongst

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2912-416: Is possibly the most plentiful genus on Earth: a single millilitre of surface seawater can contain 100,000 cells of this genus or more. Worldwide there are estimated to be several octillion (10, a billion billion billion) individuals. Prochlorococcus is ubiquitous between latitudes 40°N and 40°S, and dominates in the oligotrophic (nutrient-poor) regions of the oceans. The bacterium accounts for about 20% of

3003-436: Is the "certain degree" that defines how different organisms need to be members of different phyla. The minimal requirement is that all organisms in a phylum should be clearly more closely related to one another than to any other group. Even this is problematic because the requirement depends on knowledge of organisms' relationships: as more data become available, particularly from molecular studies, we are better able to determine

3094-454: Is the aggregate of all species which have gradually evolved from one and the same common original form, as, for example, all vertebrates. We name this aggregate [a] Stamm [i.e., stock] ( Phylon )." In plant taxonomy , August W. Eichler (1883) classified plants into five groups named divisions, a term that remains in use today for groups of plants, algae and fungi. The definitions of zoological phyla have changed from their origins in

3185-643: The Bacteriological Code Currently there are 2 phyla that have been validly published according to the Bacteriological Code Other phyla that have been proposed, but not validly named, include: Melainabacteria Scale bar, 5.0 μm. Melainabacteria is a phylum related to Cyanobacteria . Organisms belonging to this phylum have been found in the human gut and various aquatic habitats such as groundwater. By analyzing genomes of Melainabacteria, predictions are possible about

3276-552: The Catalogue of Life , and correspond to the Protozoa-Chromista scheme, with updates from the latest (2022) publication by Cavalier-Smith . Other phyla are used commonly by other authors, and are adapted from the system used by the International Society of Protistologists (ISP). Some of the descriptions are based on the 2019 revision of eukaryotes by the ISP. The number of protist phyla varies greatly from one classification to

3367-511: The International Code of Nomenclature for algae, fungi, and plants accepts the terms as equivalent. Depending on definitions, the animal kingdom Animalia contains about 31 phyla, the plant kingdom Plantae contains about 14 phyla, and the fungus kingdom Fungi contains about 8 phyla. Current research in phylogenetics is uncovering the relationships among phyla within larger clades like Ecdysozoa and Embryophyta . The term phylum

3458-446: The biosphere as we know it by burying carbon compounds and allowing the initial build-up of oxygen in the atmosphere. On the other hand, toxic cyanobacterial blooms are an increasing issue for society, as their toxins can be harmful to animals. 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. As shown in

3549-482: The black band disease ). Cyanobacteria 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 inside stones and shells (in endolithic ecosystems ). A few are endosymbionts in lichens , plants, various protists , or sponges and provide energy for

3640-618: The early Earth 's anoxic, weakly reducing prebiotic atmosphere , into an oxidizing one with free gaseous oxygen (which previously would have been immediately removed by various surface reductants ), resulting in the Great Oxidation Event and the " rusting of the Earth " during the early Proterozoic , dramatically changing the composition of life forms on Earth. The subsequent adaptation of early single-celled organisms to survive in oxygenous environments likely had led to endosymbiosis between anaerobes and aerobes , and hence

3731-445: The filamentous species , which often dominate the upper layers of microbial mats found in extreme environments such as hot springs , hypersaline water , deserts and the polar regions, but are also widely distributed in more mundane environments as well. They are evolutionarily optimized for environmental conditions of low oxygen. Some species are nitrogen-fixing and live in a wide variety of moist soils and water, either freely or in

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3822-499: The host . Some live in the fur of sloths , providing a form of camouflage . 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. Cyanobacterial growth

3913-460: The Earth's ecosystems. Planktonic cyanobacteria are a fundamental component of marine food webs and are major contributors to global carbon and nitrogen fluxes . 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 . Nowadays, cyanobacterial blooms pose

4004-491: The Zygomycota phylum. Its members would be divided between phylum Glomeromycota and four new subphyla incertae sedis (of uncertain placement): Entomophthoromycotina , Kickxellomycotina , Mucoromycotina , and Zoopagomycotina . Kingdom Protista (or Protoctista) is included in the traditional five- or six-kingdom model, where it can be defined as containing all eukaryotes that are not plants, animals, or fungi. Protista

4095-598: The accumulation of particulate organic carbon (cells, sheaths and heterotrophic organisms) in clumps. 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. So, what advantage does this communal life bring for cyanobacteria? New insights into how cyanobacteria form blooms have come from

4186-485: The activities of ancient cyanobacteria. They are often found as symbionts with a number of other groups of organisms such as fungi (lichens), corals , pteridophytes ( Azolla ), angiosperms ( Gunnera ), etc. The carbon metabolism of cyanobacteria include the incomplete Krebs cycle , the pentose phosphate pathway , and glycolysis . There are some groups capable of heterotrophic growth, while others are parasitic , causing diseases in invertebrates or algae (e.g.,

4277-480: 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 ( NH 3 ), nitrites ( NO − 2 ) or nitrates ( NO − 3 ), 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

4368-442: The basis of cyanobacteria's informal common name , blue-green algae , although as prokaryotes they are not scientifically classified as algae . Cyanobacteria are probably the most numerous taxon to have ever existed on Earth and the first organisms known to have produced oxygen , having appeared in the middle Archean eon and apparently originated in a freshwater or terrestrial environment . Their photopigments can absorb

4459-826: The cell structure and metabolic abilities. The bacterial cell is similar to cyanobacteria in being surrounded by two membranes. It differs from cyanobacteria in its ability to move by flagella (like gram-negative flagella), though some members (e.g. Gastranaerophilales ) lack flagella. Melainabacteria are not able to perform photosynthesis , but obtain energy by fermentation . " Cyanobacteriota " Vampirovibrio "Margulisbacteria" " Sericytochromatia " Cyanobacteria " Ca. Caenarcanum " " Ca. Obscuribacter " Vampirovibrio " Ca. Adamsella " " Ca. Galligastranaerophilus " Ca. Avigastranaerophilus " Ca. Limenecus " Ca. Spyradomonas " Ca. Gastranaerophilus " Ca. Scatousia " Ca. Scatenecus " Ca. Stercorousia " Melainabacteria can be found in

4550-432: The clump from the overlying medium and is also produced within the clump by respiration. In oxic solutions, high O 2 concentrations reduce the efficiency of CO 2 fixation and result in the excretion of glycolate. Under these conditions, clumping can be beneficial to cyanobacteria if it stimulates the retention of carbon and the assimilation of inorganic carbon by cyanobacteria within clumps. This effect appears to promote

4641-915: The contamination of sources of drinking water . Researchers including Linda Lawton at Robert Gordon University , have developed techniques to study these. 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. 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. An example of

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4732-460: The course of life on Earth forever by increasing the abundance of oxygen in the atmosphere. Bacteria that existed before the GEO did not rely on the presence of oxygen as a source for metabolism, such as the billion-year-old Cyanobacteria. Melainabacteria is a close relative to Cyanobacteria, though Melainabacteria diverged and do not photosynthesize. Cyanobacteria produced atmospheric oxygen and supported

4823-429: The cyanobacteria, only a few lineages colonized the open ocean: Crocosphaera and relatives, cyanobacterium UCYN-A , Trichodesmium , as well as Prochlorococcus and Synechococcus . From these lineages, nitrogen-fixing cyanobacteria are particularly important because they exert a control on primary productivity and the export of organic carbon to the deep ocean, by converting nitrogen gas into ammonium, which

4914-501: The cyanobacterial group. In general, photosynthesis in cyanobacteria uses water as an electron donor and produces oxygen as a byproduct, though some may also use hydrogen sulfide 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 . The large amounts of oxygen in the atmosphere are considered to have been first created by

5005-457: The cyanobacterium form buoyant aggregates by trapping oxygen bubbles in the slimy web of cells and polysaccharides. Previous studies on Synechocystis have shown type IV pili , which decorate the surface of cyanobacteria, also play a role in forming blooms. These retractable and adhesive protein fibres are important for motility, adhesion to substrates and DNA uptake. The formation of blooms may require both type IV pili and Synechan – for example,

5096-493: The cytoplasm of the cell. Carboxysomes are icosahedral structures composed of hexameric shell proteins that assemble into cage-like structures that can be several hundreds of nanometres in diameter. It is believed that these structures tether the CO 2 -fixing enzyme, RuBisCO , to the interior of the shell, as well as the enzyme carbonic anhydrase , using metabolic channeling to enhance the local CO 2 concentrations and thus increase

5187-471: The development of early plant cells. The genomes of Melainabacteria organisms isolated from ground water indicate that the organism has the capacity to fix nitrogen. Melainabacteria lack linked electron transport chains but have multiple methods to generate a membrane potential which can then produce ATP via ATP synthase . They are able to use Fe hydrogenases for H 2 production that can be consumed by other microorganisms. Melainabacteria from

5278-432: 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

5369-457: The efficiency of the RuBisCO enzyme. 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. The photosynthetic machinery is embedded in the thylakoid membranes, with phycobilisomes acting as light-harvesting antennae attached to the membrane, giving

5460-920: The evolution of eukaryotes during the Paleoproterozoic . Cyanobacteria use photosynthetic pigments such as various forms of chlorophyll , carotenoids , phycobilins to convert the photonic energy in sunlight to chemical energy . Unlike heterotrophic prokaryotes, cyanobacteria have internal membranes . These are flattened sacs called thylakoids where photosynthesis is performed. Photoautotrophic eukaryotes such as red algae , green algae and plants perform photosynthesis in chlorophyllic organelles that are thought to have their ancestry in cyanobacteria, acquired long ago via endosymbiosis. These endosymbiont cyanobacteria in eukaryotes then evolved and differentiated into specialized organelles such as chloroplasts , chromoplasts , etioplasts , and leucoplasts , collectively known as plastids . Sericytochromatia,

5551-441: The family Fabaceae , 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. Cyanobacteria such as Anabaena (a symbiont of the aquatic fern Azolla ) can provide rice plantations with biofertilizer . Cyanobacteria use the energy of sunlight to drive photosynthesis ,

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5642-472: The filamentous forms, Trichodesmium are free-living and form aggregates. However, filamentous heterocyst-forming cyanobacteria (e.g., Richelia , Calothrix ) are found in association with diatoms such as Hemiaulus , Rhizosolenia and Chaetoceros . Marine cyanobacteria include the smallest known photosynthetic organisms. The smallest of all, Prochlorococcus , is just 0.5 to 0.8 micrometres across. In terms of numbers of individuals, Prochlorococcus

5733-504: The first publication of the APG system in 1998, which proposed a classification of angiosperms up to the level of orders , many sources have preferred to treat ranks higher than orders as informal clades. Where formal ranks have been provided, the traditional divisions listed below have been reduced to a very much lower level, e.g. subclasses . Wolf plants Hepatophyta Liver plants Coniferophyta Cone-bearing plant Phylum Microsporidia

5824-402: The first signs of multicellularity. Many cyanobacteria form motile filaments of cells, called hormogonia , that travel away from the main biomass to bud and form new colonies elsewhere. 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

5915-478: The green pigmentation observed (with wavelengths from 450 nm to 660 nm) in most cyanobacteria. 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. Respiration in cyanobacteria can occur in the thylakoid membrane alongside photosynthesis, with their photosynthetic electron transport sharing

6006-563: The gut and how they got there. Ongoing studies such as, "The human gut and groundwater harbor non-photosynthetic bacteria belonging to a new candidate phylum sibling to Cyanobacteria," funded by various organizations such as the National Institutes of Health , the David and Lucile Packard Foundation , The Hartwell Foundation, the Arnold and Mabel Beckman Foundation , the U.S. Department of Energy ,

6097-416: 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. Cyanophages infect

6188-519: The human gut also synthesize several B and K vitamins , which suggests that these bacteria are beneficial to their host because they are consumed along with plant fibers. Melainabacteria have been found to potentially play a role in digesting fiber in the human gut, and are more commonly in herbivorous mammals and those with plant-rich diets. Because plant diets require more fiber break-down, Melainabacteria may aid in this digestive function. However, scientists are unsure of why these microbes are in

6279-537: The most abundant photosynthetic organisms on Earth, accounting for a quarter of all carbon fixed in marine ecosystems. In contrast to free-living marine cyanobacteria, some cyanobionts are known to be responsible for nitrogen fixation rather than carbon fixation in the host. However, the physiological functions of most cyanobionts remain unknown. Cyanobionts have been found in numerous protist groups, including dinoflagellates , tintinnids , radiolarians , amoebae , diatoms , and haptophytes . Among these cyanobionts, little

6370-452: 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 Falconer Some cyanobacteria,

6461-404: The network of polysaccharides and cells, enabling the microorganisms to form buoyant blooms. 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. The diagram on the left above shows

6552-411: The next. The Catalogue of Life includes Rhodophyta and Glaucophyta in kingdom Plantae, but other systems consider these phyla part of Protista. In addition, less popular classification schemes unite Ochrophyta and Pseudofungi under one phylum, Gyrista , and all alveolates except ciliates in one phylum Myzozoa , later lowered in rank and included in a paraphyletic phylum Miozoa . Even within

6643-528: The other hand, the highly parasitic phylum Mesozoa was divided into two phyla ( Orthonectida and Rhombozoa ) when it was discovered the Orthonectida are probably deuterostomes and the Rhombozoa protostomes . This changeability of phyla has led some biologists to call for the concept of a phylum to be abandoned in favour of placing taxa in clades without any formal ranking of group size. A definition of

6734-417: The oxygen in the Earth's atmosphere. Cyanobacteria are variable in morphology, ranging from unicellular and filamentous to colonial forms . Filamentous forms exhibit functional cell differentiation such as heterocysts (for nitrogen fixation), akinetes (resting stage cells), and hormogonia (reproductive, motile filaments). These, together with the intercellular connections they possess, are considered

6825-711: The photosystems. The phycobilisome components ( phycobiliproteins ) are responsible for the blue-green pigmentation of most cyanobacteria. 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. 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. This process of "complementary chromatic adaptation"

6916-441: The pili may allow cyanobacteria from the same species to recognise each other and make initial contacts, which are then stabilised by building a mass of extracellular polysaccharide. 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. Type IV pili on their own could also control

7007-411: The pili may help to export the polysaccharide outside the cell. Indeed, the activity of these protein fibres may be connected to the production of extracellular polysaccharides in filamentous cyanobacteria. A more obvious answer would be that pili help to build the aggregates by binding the cells with each other or with the extracellular polysaccharide. As with other kinds of bacteria, certain components of

7098-478: The position of marine cyanobacteria in the water column by regulating viscous drag. Extracellular polysaccharide appears to be a multipurpose asset for cyanobacteria, from floatation device to food storage, defence mechanism and mobility aid. One of the most critical processes determining cyanobacterial eco-physiology is cellular death . Evidence supports the existence of controlled cellular demise in cyanobacteria, and various forms of cell death have been described as

7189-476: 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. The cyanobacteria Synechocystis and Cyanothece are important model organisms with potential applications in biotechnology for bioethanol production, food colorings, as a source of human and animal food, dietary supplements and raw materials. Cyanobacteria produce

7280-468: The red- and blue-spectrum frequencies of sunlight (thus reflecting a greenish color) to split water molecules into hydrogen ions and oxygen. The hydrogen ions are used to react with carbon dioxide to produce complex organic compounds such as carbohydrates (a process known as carbon fixation ), and the oxygen is released as a byproduct . By continuously producing and releasing oxygen over billions of years, cyanobacteria are thought to have converted

7371-469: The relationships between groups. So phyla can be merged or split if it becomes apparent that they are related to one another or not. For example, the bearded worms were described as a new phylum (the Pogonophora) in the middle of the 20th century, but molecular work almost half a century later found them to be a group of annelids , so the phyla were merged (the bearded worms are now an annelid family ). On

7462-427: 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. Dissolved inorganic carbon (DIC) diffuses into

7553-443: The root surface within a restricted zone by Nostoc . The relationships between cyanobionts (cyanobacterial symbionts) and protistan hosts are particularly noteworthy, as some nitrogen-fixing cyanobacteria ( diazotrophs ) play an important role in primary production , especially in nitrogen-limited oligotrophic oceans. Cyanobacteria, mostly pico-sized Synechococcus and Prochlorococcus , are ubiquitously distributed and are

7644-505: The roots of wheat and cotton plants. Calothrix sp. has also been found on the root system of wheat. Monocots , such as wheat and rice, have been colonised by Nostoc spp., In 1991, Ganther and others isolated diverse heterocystous nitrogen-fixing cyanobacteria, including Nostoc , Anabaena and Cylindrospermum , from plant root and soil. Assessment of wheat seedling roots revealed two types of association patterns: loose colonization of root hair by Anabaena and tight colonization of

7735-415: The same compartment as the components of respiratory electron transport. While the goal of photosynthesis is to store energy by building carbohydrates from CO 2 , respiration is the reverse of this, with carbohydrates turned back into CO 2 accompanying energy release. Cyanobacteria appear to separate these two processes with their plasma membrane containing only components of the respiratory chain, while

7826-399: The six Linnaean classes and the four embranchements of Georges Cuvier . Informally, phyla can be thought of as groupings of organisms based on general specialization of body plan . At its most basic, a phylum can be defined in two ways: as a group of organisms with a certain degree of morphological or developmental similarity (the phenetic definition), or a group of organisms with

7917-423: The so-called cyanobionts (cyanobacterial symbionts), have a symbiotic relationship with other organisms, both unicellular and multicellular. As illustrated on the right, there are many examples of cyanobacteria interacting symbiotically with land plants . Cyanobacteria can enter the plant through the stomata and colonize the intercellular space, forming loops and intracellular coils. Anabaena spp. colonize

8008-470: 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. Many cyanobacteria are able to reduce nitrogen and carbon dioxide under aerobic conditions,

8099-464: Was coined in 1866 by Ernst Haeckel from the Greek phylon ( φῦλον , "race, stock"), related to phyle ( φυλή , "tribe, clan"). Haeckel noted that species constantly evolved into new species that seemed to retain few consistent features among themselves and therefore few features that distinguished them as a group ("a self-contained unity"): "perhaps such a real and completely self-contained unity

8190-460: Was discovered in 1963. 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 ). Phylum (biology) In biology , a phylum ( / ˈ f aɪ l əm / ; pl. : phyla ) is a level of classification or taxonomic rank below kingdom and above class . Traditionally, in botany the term division has been used instead of phylum, although

8281-499: Was discovered in 1986 and accounts for more than half of the photosynthesis of the open ocean. 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

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