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Cyanobacteria
Cyanobacteria /saɪˌænoʊbækˈtɪəriə/, also known as
Cyanophyta, are a phylum consisting of free-living Cyanobacteria
photosynthetic bacteria and the endosymbiotic plastids, a Temporal range: 2100–0 Ma
sister group to Gloeomargarita, that are present in some Had'n Archean Proterozoic Pha.
eukaryotes. They commonly obtain their energy through
oxygenic photosynthesis.[4] The oxygen gas in the atmosphere
of earth is produced by cyanobacteria of this phylum, either as
free-living bacteria or as the endosymbiotic plastids.[5] The
name cyanobacteria comes from the color of the bacteria
(Greek: κυανός, romanized: kyanós, lit. 'blue').[6][7]
Cyanobacteria, which are prokaryotes, are also called "blue-
green algae",[4][8] though some modern botanists restrict
the term algae to eukaryotes.[9] Cyanobacteria appear to have Microscope image of Cylindrospermum,
originated in freshwater or a terrestrial environment.[10]
a genus of Cyanobacteria
Unlike heterotrophic prokaryotes, cyanobacteria have internal Scientific classification
membranes. These are flattened sacs called thylakoids where
photosynthesis is performed.[11][12] Domain: Bacteria
(unranked): Terrabacteria
Phototrophic eukaryotes such as green plants perform
photosynthesis in plastids that are thought to have their (unranked): Cyanobacteria-
ancestry in cyanobacteria, acquired long ago via a process Melainabacteria
called endosymbiosis. These endosymbiotic cyanobacteria in group
eukaryotes then evolved and differentiated into specialized
organelles such as chloroplasts, etioplasts and leucoplasts. Phylum: Cyanobacteria
Stanier, 1973
By producing and releasing oxygen (as a byproduct of
photosynthesis), cyanobacteria are thought to have converted Class: Cyanophyceae
the early oxygen-poor, reducing atmosphere into an oxidizing
Orders[3]
one, causing the Great Oxygenation Event and the "rusting of
the Earth",[13] which dramatically changed the composition of
the Earth's life forms and led to the near-extinction of As of 2014 the taxonomy was
anaerobic organisms.[14] under revision[1][2]
Contents Synechococcales
Incertae sedis
Description
Nitrogen fixation †Gunflintia
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Morphology †Ozarkcollenia
Ecology Plastids (endosymbiotic)
Photosynthesis
Synonyms
Carbon fixation
Electron transport
Myxophyceae Wallroth, 1833
Respiration
Electron transport chain Phycochromaceae Rabenhorst,
Metabolism 1865
Description
Light microscope view of cyanobacteria
Cyanobacteria are a group of photosynthetic bacteria, some from a microbial mat
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).[15] 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 fixation[16][17][18] in an anaerobic environment due to its sensitivity to oxygen.[18]
Nitrogen fixation
Some cyanobacteria can fix atmospheric nitrogen in anaerobic conditions by means of specialized
cells called heterocysts.[17][18] 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 family Fabaceae, among others).
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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.[19]
Cyanobacteria such as Anabaena (a symbiont of the aquatic fern Azolla) can provide rice plantations
with biofertilizer.[20]
Morphology
Colonies of Nostoc pruniforme Each individual cell (each single cyanobacterium) typically has a
thick, gelatinous cell wall.[23] They lack flagella, but hormogonia
of some species can move about by gliding along surfaces.[24]
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.[25] These vesicles are not organelles as such. They are not bounded by lipid
membranes but by a protein sheath.
Ecology
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 in endolithic
ecosystem.[26] 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.[27]
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.[28]
Cyanobacterial growth is favored in ponds and lakes where waters are calm and have little turbulent
mixing.[30] 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.[30]
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.[31]
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"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
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photosynthesis.
Cyanobacteria fulfill
vital ecological
functions in the
world's oceans, being
important
contributors to global
carbon and nitrogen
budgets." – Stewart
and Falconer[35]
Photosynthesis
Cyanobacteria have several unique features. As the endosymbiotic plastids are endosymbiotic
cyanobacteria, they share these features insofar as they have not lost them.
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.[36] 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.[37]
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.[38] 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.[39]
Respiration
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.[41] Cyanobacteria use electrons from succinate
dehydrogenase rather than from NADPH for respiration.[41]
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.[42]
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.[43]
Attached to the thylakoid membrane, phycobilisomes act as light-harvesting antennae for the
photosystems.[44] The phycobilisome components (phycobiliproteins) are responsible for the blue-
green pigmentation of most cyanobacteria.[45] 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.[46][47] 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.[48] 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.[49][50]
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[51] 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.[52] The large amounts of
oxygen in the atmosphere are considered to have been first created by the activities of ancient
cyanobacteria.[53] They are often found as symbionts with a number of other groups of organisms
such as fungi (lichens), corals, pteridophytes (Azolla), angiosperms (Gunnera), etc.[54]
There are some groups capable of heterotrophic growth,[55] while others are parasitic, causing
diseases in invertebrates or algae (e.g., the black band disease).[56][57][58]
Relationship to chloroplasts
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Primary chloroplasts are cell organelles found in some eukaryotic lineages, where they are specialized
in performing the photosynthesis. They are considered to have evolved from endosymbiotic
cyanobacteria.[59][60] After some years of debate,[61] 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.[62][63][64][65]
The morphological similarity between chloroplasts and cyanobacteria was first reported by German
botanist Andreas Schimper in the 19th century[66] Chloroplasts are only found in plants and algae,[67]
thus paving the way for Russian biologist Konstantin Mereschkowski to suggest the symbiogenic
origin of the plastid in 1905.[68] Lynn Margulis brought this hypothesis back to attention more than
60 years later[69] 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,[70][62][65] genomic,[71] biochemical[72][73] and structural evidence.[74] 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.[75]
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.[76][59]
Within this evolutionary context, it is noteworthy that, as far as we can tell, oxygenic photosynthesis
only evolved once (in prokaryotic cyanobacteria), and all 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.[77]
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.[78]
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.[78]
Classification
Historically, bacteria were first classified as plants constituting the class Schizomycetes, which along
with the Schizophyceae (blue-green algae/Cyanobacteria) formed the phylum Schizophyta,[83] then
in the phylum Monera in the kingdom Protista by Haeckel in 1866, comprising Protogens,
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The remainder are validly published under the International Code of Nomenclature for algae, fungi,
and plants.
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.[90]
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 date from 3.5 Ga ago.[91] 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.[92]
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Cyanobacteria may possess the ability to produce substances that could one day serve as anti-
inflammatory agents and combat bacterial infections in humans.[107]
Spirulina's extracted blue color is used as a natural food coloring in gum and candy.[108]
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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.[109]
Health risks
Some cyanobacteria can produce neurotoxins, cytotoxins, endotoxins, and hepatotoxins (e.g., the
microcystin-producing bacteria genus microcystis), which are collectively known as cyanotoxins.
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.[113] BMAA from desert crusts found throughout Qatar might
have contributed to higher rates of ALS in Gulf War veterans.[111][114]
Chemical control
Several chemicals can eliminate cyanobacterial blooms from smaller water-based systems such as
swimming pools. They include: calcium hypochlorite, copper sulphate, cupricide, and simazine.[115]
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.[115] Copper sulfate is also used
commonly, but no longer recommended by the Australian Department of Agriculture, as it kills
livestock, crustaceans, and fish.[115] 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.[115] Ferric alum treatments at the rate of 50 mg/l will reduce algae blooms.[115][116]
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.[115]
Dietary supplementation
Some cyanobacteria are sold as food, notably Aphanizomenon flos-
aquae and Arthrospira platensis (Spirulina).[117]
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Gallery
See also
Archean Eon Geological history of oxygen
Bacterial phyla, other major lineages of Hypolith
Bacteria Microbial mats
Biodiesel Microalgae
Cyanobiont Phytoplankton
Endosymbiotic theory Proterozoic Eon
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Attribution
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Further reading
Cribbs G (1997). Nature's Superfood: the Blue-Green Algae Revolution (first ed.). Newleaf.
ISBN 978-0-7522-0569-4.
Savage M (1994). The Millennial Project: Colonizing the Galaxy in Eight Easy Steps (https://archi
ve.org/details/millennialprojec00sava). Little Brown & Co. ISBN 978-0-316-77163-4.
Fogg GE, Stewart WD, Fay P, Walsby AE (1973). The Blue-green Algae. London and New York:
Academic Press. ISBN 978-0-12-261650-1.
"Architects of the earth's atmosphere (http://www.ucmp.berkeley.edu/bacteria/cyanointro.html)",
Introduction to the Cyanobacteria, University of California, Berkeley, 3 February 2006.
Whitton BA (25 April 2002). "Phylum Cyanophyta (Cyanobacteria)". The Freshwater Algal Flora of
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External links
What are Cyanobacteria and What are its Types? (http://www.thebigger.com/biology/monera/what
-are-cyanobacteria-and-what-are-its-types/)
Webserver for Cyanobacteria Research (https://web.archive.org/web/20190916232500/http://ww
w-cyanosite.bio.purdue.edu/)
Diving an Antarctic Time Capsule Filled With Primordial Life (http://scientistatwork.blogs.nytimes.c
om/2013/01/31/diving-an-antarctic-time-capsule-filled-with-primordial-life)
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