As of 2014 the taxonomy was under revision
Incertae sedis _
* Myxophyceae Wallroth, 1833
* Phycochromaceae Rabenhorst, 1865
* Cyanophyceae Sachs, 1874
* Schizophyceae Cohn, 1879
* Cyanophyta Steinecke, 1931
* Oxyphotobacteria Gibbons padding:0.5em 0 0 0; border-style:solid;
float:right;clear:right; padding:0 0.5em; margin:0.3em 0 0.8em 1.4em;
overflow: hidden;"> Life timeline view • discuss • edit -4500
— – -4000 — – -3500 — – -3000 — – -2500 — – -2000
— – -1500 — – -1000 — – -500 — – 0 — _WATER _
life _PHOTOSYNTHESIS _ EUKARYOTES Multicellular
life LAND LIFE DINOSAURS MAMMALS FLOWERS ←
Earliest Earth (−4540 ) ← Earliest water ← Earliest
life ← LHB meteorites ← Earliest oxygen ←
Atmospheric oxygen ←
Oxygen crisis ← Earliest sexual
reproduction ← Ediacara biota ←
← Earliest humans P
n Pongola Huronian
Cryogenian Andean Karoo Quaternary
Axis scale : millions of years .
Orange labels: known _ICE AGES_.
Also see: _
Human timeline _ and _Nature timeline _
Cyanobacteria are a group of photosynthetic, nitrogen-fixing bacteria
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 _). 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 .
Cyanobacteria can fix atmospheric nitrogen in anaerobic conditions by
means of specialized cells called heterocysts .
Heterocysts may also
form under the appropriate environmental conditions (anoxic) when
fixed nitrogen is scarce. Heterocyst-forming species are specialized
for nitrogen fixation and are able to fix nitrogen gas into ammonia
(NH3 ), nitrites (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,
Fabaceae family, among others).
Free-living cyanobacteria are present in the water of rice paddies ,
and cyanobacteria can be found growing as epiphytes on the surfaces of
the green alga, _Chara _, where they may fix nitrogen. Cyanobacteria
such as _
Anabaena _ (a symbiont of the aquatic fern _
Azolla _), can
provide rice plantations with biofertilizer.
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_ Colonies of
Nostoc pruniforme _ _
Cylindrospermum _ sp.
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 a weaker cell in a filament, called a necridium.
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 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.
Cyanobacterial bloom near
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 almost every endolithic ecosystem . 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
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.
Cyanobacteria prefer calm waters, such as those provided by ponds and
lakes. 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. When the bacteria are found in rivers, they have usually come
from the outfall of lakes upstream from the sampling point.
Cyanobacteria are a growing concern for drinking water utilities who
use lakes and rivers as their source water. The bacteria can interfere
with treatment in various ways, primarily by plugging filters (often
large beds of sand and similar media), and by producing cyanotoxins ,
which have the potential of causing serious illness if consumed.
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. Many cyanobacteria even display the
circadian rhythms that were once thought to exist only in eukaryotic
cells (see bacterial circadian rhythms ).
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
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
While contemporary cyanobacteria are linked to the plant kingdom as
descendants of the endosymbiotic progenitor of the chloroplast , there
are several features which are unique to this group.
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 . 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
RuBisCO , to the interior of the shell, as well as the enzyme
carbonic anhydrase , using the paradigm of metabolic channeling to
enhance the local CO2 concentrations and thus increase the efficiency
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.
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
Metabolism And Organelles
As prokaryotes, cyanobacteria do not have nuclei. In most forms, the
photosynthetic machinery is embedded into internal membrane structures
called thylakoids .
Cyanobacteria get their colour from the bluish
pigment phycocyanin , which assists chlorophyll in photosynthesis. 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 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
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 anaerobic conditions, they are able to
use only PS I—cyclic photophosphorylation —with electron donors
other than water (hydrogen sulfide , thiosulphate, or even molecular
hydrogen ) just like purple photosynthetic bacteria. Furthermore, they
share an archaeal property, the ability to reduce elemental sulfur by
anaerobic respiration in the dark. Their photosynthetic electron
transport shares the same compartment as the components of respiratory
electron transport. Their plasma membrane contains only components of
the respiratory chain, while the thylakoid membrane hosts an
interlinked respiratory and photosynthetic electron transport chain.
The terminal oxidases in the thylakoid membrane
respiratory/photosynthetic electron transport chain are essential for
survival to rapid light changes, although not for dark maintenance
under conditions where cells are not light stressed.
Attached to the thylakoid membrane, phycobilisomes act as
light-harvesting antennae for 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 phycobilisomes. 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. This process of complementary chromatic adaptation is
a way for the cells to maximize the use of available light for
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.
There are some groups capable of heterotrophic growth, while others
are parasitic , causing diseases in invertebrates or eukaryotic algae
(e.g., the black band disease ).
RELATIONSHIP TO CHLOROPLASTS
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all other cyanobacteria
Cladogram showing plastids (chloroplasts
and similar) and basal cyanobacteria
Chloroplasts found in eukaryotes (algae and plants) appear to have
evolved from an endosymbiotic relation with cyanobacteria. This
endosymbiotic theory is supported by various structural and genetic
similarities. Primary chloroplasts are found among the "true plants "
or green plants – species ranging from sea lettuce to evergreens and
flowers that contain chlorophyll _b_ – as well as among the red
algae and glaucophytes , marine species that contain phycobilins. It
now appears that these chloroplasts probably had a single origin, in
an ancestor of the clade called
Archaeplastida . However this does not
necessitate origin from cyanobacteria themselves; microbiology is
still undergoing profound classification changes and entire domains
Archaea ) are poorly mapped and understood. Other algae
likely took their chloroplasts from these forms by secondary
endosymbiosis or ingestion.
_ Tree of Life in Generelle Morphologie der Organismen_ (1866).
Note the location of the genus _
Nostoc _ with algae and not with
bacteria (kingdom "Monera") See also:
Historically, bacteria were first classified as plants constituting
the class Schizomycetes, which along with the Schizophyceae
(blue-green algae/Cyanobacteria) formed the phylum Schizophyta, then
in the phylum
Monera in the kingdom
Haeckel in 1866,
comprising _Protogens, Protamaeba, Vampyrella, Protomonae_, and
_Vibrio_, but not _Nostoc_ and other cyanobacteria, which were
classified with algae, later reclassified as the _
Prokaryotes _ by
The cyanobacteria were traditionally classified by morphology into
five sections, referred to by the numerals I-V. The first three –
Pleurocapsales , and
Oscillatoriales – are not
supported by phylogenetic studies. The latter two –
Stigonematales – are monophyletic, and make up the heterocystous
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
Nostocales and Stigonematales, the cells have the
ability to develop heterocysts in certain conditions. Stigonematales,
unlike Nostocales, include species with truly branched trichomes.
Most taxa included in the phylum or division
Cyanobacteria have not
yet been validly published under the Bacteriological Code , except:
* The classes Chroobacteria ,
Hormogoneae , and
* The orders
Pleurocapsales , and
* The families
Prochloraceae and Prochlorotrichaceae
* The genera _Halospirulina , Planktothricoides ,
Prochloron _, and _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
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 ,
Stromatolites left behind by
cyanobacteria are the oldest known fossils of life on Earth. This
one-billion-year-old fossil is from Glacier National Park in Montana.
Precambrian , stromatolite communities of microorganisms
grew in most marine and non-marine environments in the photic zone.
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
Shark Bay , Western Australia).
ancient records of life on Earth by fossil remains which might date
from more than 3.5 Ga ago, but this is disputed. 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
levels 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 methane levels, 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.
Oncolites are sedimentary structures composed of oncoids, which are
layered structures formed by cyanobacterial growth. Oncolites are
similar to stromatolites, but instead of forming columns, they form
approximately spherical structures that were not attached to the
underlying substrate as they formed. The oncoids often form around a
central nucleus, such as a shell fragment, and a calcium carbonate
structure is deposited by encrusting microbes . Oncolites are
indicators of warm waters in the photic zone , but are also known in
contemporary freshwater environments. These structures rarely exceed
10 cm in diameter.
BIOTECHNOLOGY AND APPLICATIONS
The unicellular cyanobacterium _
Synechocystis _ sp. PCC6803 was the
third prokaryote and first photosynthetic organism whose genome was
completely sequenced . It continues to be an important model
Cyanothece _ ATCC 51142 is an important diazotrophic model
organism. The smallest genomes have been found in _
spp. (1.7 Mb) and the largest in _
Nostoc punctiforme _ (9 Mb).
Those of _Calothrix _ spp. are estimated at 12–15 Mb, as large as
yeast . Oncolites from the
Alamo bolide impact in
Recent research has suggested the potential application of
cyanobacteria to the generation of renewable energy by converting
sunlight into electricity. Internal photosynthetic pathways can be
coupled to chemical mediators that transfer electrons to external
electrodes. Currently, efforts are underway to commercialize algae
-based fuels such as diesel , gasoline , and jet fuel .
Cyanobacteria cultured in specific media:
Cyanobacteria can be helpful
in agriculture as they have the ability to fix atmospheric nitrogen in
Researchers from a company called
Algenol have cultured genetically
modified cyanobacteria in sea water inside a clear plastic enclosure
so they first make sugar (pyruvate) from CO2 and the water via
photosynthesis. Then, the bacteria secrete ethanol from the cell into
the salt water. As the day progresses, and the solar radiation
intensifies, ethanol concentrations build up and the ethanol itself
evaporates onto the roof of the enclosure. As the sun recedes,
evaporated ethanol and water condense into droplets, which run along
the plastic walls and into ethanol collectors, from where it is
extracted from the enclosure with the water and ethanol separated
outside the enclosure. As of March 2013,
Algenol was claiming to have
tested its technology in Florida and to have achieved yields of 9,000
US gallons per acre per year. This could potentially meet US demands
for ethanol in gasoline in 2025, assuming a B30 blend, from an area of
around half the size of California’s San Bernardino County,
requiring less than one-tenth of the area than ethanol from other
biomass, such as corn, and only very limited amounts of fresh water.
Cyanobacteria may possess the ability to produce substances that
could one day serve as anti-inflammatory agents and combat bacterial
infections in humans.
Spirulina's extracted blue color is used as a natural food coloring
in gum and candy.
Researchers from several space agencies argue that cyanobacteria
could be used for producing goods for human consumption (food,
oxygen...) in future manned outposts on Mars, by transforming
materials available on this planet.
Cyanobacteria can produce neurotoxins , cytotoxins , endotoxins , and
hepatotoxins (e.g., the microcystin -producing bacteria genus
microcystis ), and collectively known as cyanotoxins .
Specific toxins include, anatoxin-a , anatoxin-as , aplysiatoxin ,
cyanopeptolin, cylindrospermopsin , domoic acid , nodularin R (from
Nodularia _), neosaxitoxin , and saxitoxin .
explosively under certain conditions. This results in algal blooms ,
which can become harmful to other species , and pose a danger to
humans and animals, if the cyanobacteria involved produce toxins.
Several cases of human poisoning have been documented, but a lack of
knowledge prevents an accurate assessment of the risks.
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
Lake Mascoma had an up to 25 times greater risk of ALS
than the expected incidence. BMAA from desert crusts found throughout
Qatar might have contributed to higher rates of ALS in Gulf War
Several chemicals can eliminate cyanobacterial blooms from
water-based systems. They include: calcium hypochlorite , copper
sulphate , cupricide, and simazine . The calcium hypochlorite amount
needed varies depending on the cyanobacteria bloom, and treatment is
needed periodically. According to the Department of Agriculture
Australia, a rate of 12 g of 70% material in 1000 l of water is often
effective to treat a bloom. Copper sulfate is also used commonly, but
no longer recommended by the Australian Department of Agriculture, as
it kills livestock, crustaceans, and fish. Culpricide is a chelated
copper product that eliminates blooms with lower toxicity risks than
copper sulfate. Dosage recommendations vary from 190 ml to 4.8 l per
1000 m2. Ferric alum treatments at the rate of 50 mg/l will reduce
algae blooms. Simazine, which is also a herbicide, will continue to
kill blooms for several days after an application.
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.
Some cyanobacteria are sold as food, notably _Aphanizomenon
flos-aquae _ and _Arthrospira platensis_ (Spirulina ).
Despite the associated toxins which many of the members of this
phylum produce, some microalgae also contain substances of high
biological value, such as polyunsaturated fatty acids, amino acids
(proteins), pigments, antioxidants, vitamins, and minerals. Edible
blue-green algae reduce the production of pro-inflammatory cytokines
by inhibiting NF-κB pathway in macrophages and splenocytes. Sulfate
polysaccharides exhibit immunomodulatory, antitumor, antithrombotic,
anticoagulant, anti-mutagenic, anti-inflammatory, antimicrobial, and
even antiviral activity against HIV, herpes, and hepatitis.
Bacterial phyla , other major lineages of Bacteria
Geological history of oxygen
Great Oxygenation Event
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* High lipid content microalgae
Milky seas effect
Paradox of the plankton
Paradox of the plankton
* Thin layers
* Eukaryotic picoplankton
Aeromonas salmonicida _
Enteric redmouth disease
Flavobacterium columnare _
Pelagibacter ubique _
Streptococcus iniae _
Emiliania huxleyi _
Thalassiosira pseudonana _
Pfiesteria piscicida _
Velvet (fish disease)
* Marine larvae
Algal nutrient solutions
Diel vertical migration
Fish diseases and parasites
Microbial population biology
Microbial food web
Host microbe interactions in Caenorhabditis elegans
Dark field microscopy
Marine microbial symbiosis
* International Census of Marine
Microbes in human culture
Microbial symbiosis and immunity
Human Microbiome Project
* in pregnancy
* Lines on the Antiquity of
Microbially induced sedimentary structure
Microbial dark matter
Physical factors affecting microbial life
Bacteria classification (phyla and orders )
Eukaryota (Supergroup Plant Hacrobia
Heterokont Alveolata Rhizaria Excavata Amoebozoa Opisthokonta
* _thermophiles _
FCB group )
PVC group )
Source: _Bergey\'s Manual _ (2001–2012). Alternative views:
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