From Wikipedia, the free encyclopedia
Cyanobacteria
Cyanobacteria
Scientific classification
Domain: Bacteria
Phylum: Cyanobacteria
Orders
The taxonomy is currently under revision.[1]
Cyanobacteria, also known as blue-green algae, blue-
green bacteria or Cyanophyta, is a phylum of bacteria
that obtain their energy through photosynthesis. The
name "cyanobacteria" comes from the color of the bac-
teria (Greek: κυανός (kyanós) = blue). They are a signific-
ant component of the marine nitrogen cycle and an im-
portant primary producer in many areas of the ocean,
but are also found in habitats other than the marine en-
vironment; in particular cyanobacteria are known to oc-
cur in both freshwater,[2] hypersaline inland lakes[3] and
in arid areas where they are a major component of bio-
logical soil crusts.
Stromatolites of fossilized oxygen-producing cy-
anobacteria have been found from 2.8 billion years
ago.[4] The ability of cyanobacteria to perform oxygenic
photosynthesis is thought to have converted the early
reducing atmosphere into an oxidizing one, which dra-
matically changed the composition of life forms on Earth
by provoking an explosion of biodiversity and leading to
the near-extinction of oxygen-intolerant organisms.
Chloroplasts in plants and eukaryotic algae have evolved
from cyanobacteria via endosymbiosis.
Forms
Cyanobacteria are found in almost every conceivable en-
vironment, from oceans to fresh water to bare rock to
soil. They can occur as planktonic cells or form photo-
trophic biofilms in fresh water and marine environ-
ments, they occur in damp soil, or even temporarily
moistened rocks in deserts. 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.
1
Cyanobacteria
Colonies of Nostoc pruniforme.
Cyanobacteria include unicellular and colonial spe-
cies. Colonies may form filaments, sheets or even hollow
balls. Some filamentous colonies show the ability to dif-
ferentiate into several different cell types: vegetative
cells, the normal, photosynthetic cells that are formed
under favorable growing conditions; akinetes, the
climate-resistant spores that may form when environ-
mental conditions become harsh; and thick-walled het-
erocysts, which contain the enzyme nitrogenase, vital
for nitrogen fixation. Heterocysts may also form under
the appropriate environmental conditions (anoxic)
wherever nitrogen is necessary. 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 ni-
trogen cannot be used by plants directly]. The rice pad-
dies of Asia, which produce about 75% of the world’s
rice[5], could not do so were it not for healthy popula-
tions of nitrogen-fixing cyanobacteria in the rice paddy
fertilizer.
Many cyanobacteria also form motile filaments,
called hormogonia, that travel away from the main bio-
mass to bud and form new colonies elsewhere. The cells
in a hormogonium are often thinner than in the vegetat-
ive state, and the cells on either end of the motile chain
may be tapered. In order 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 of a cyanobacterium typically
has a thick, gelatinous cell wall. They differ from other
gram-negative bacteria in that the quorum sensing mo-
lecules autoinducer-2[6] and acyl-homoserine lactones[7]
are absent. They lack flagella, but hormogonia and some
From Wikipedia, the free encyclopedia
unicellular species may move about by gliding along sur-
faces. In water columns some cyanobacteria float by
forming gas vesicles, like in archaea.
Some of these organisms contribute significantly to
global ecology and the oxygen cycle. The tiny marine cy-
anobacterium Prochlorococcus was discovered in 1986 and
accounts for more than half of the photosynthesis of the
open ocean.[8] Many cyanobacteria even display the cir-
cadian rhythms that were once thought to exist only in
eukaryotic cells (see bacterial circadian rhythms).
Photosynthesis
Cyanobacteria have an elaborate and highly organized
system of internal membranes which function in photo-
synthesis. Photosynthesis in cyanobacteria generally
uses water as an electron donor and produces oxygen as
a by-product, though some may also use hydrogen sulf-
ide as occurs among other photosynthetic bacteria. Car-
bon dioxide is reduced to form carbohydrates via the
Calvin cycle. In most forms the photosynthetic ma-
chinery is embedded into folds of the cell membrane,
called thylakoids. The large amounts of oxygen in the at-
mosphere are considered to have been first created by
the activities of ancient cyanobacteria. Due to their abil-
ity to fix nitrogen in aerobic conditions they are often
found as symbionts with a number of other groups of or-
ganisms such as fungi (lichens), corals, pteridophytes
(Azolla), angiosperms (Gunnera) etc.
Cyanobacteria are the only group of organisms that
are able to reduce nitrogen and carbon in aerobic condi-
tions, a fact that may be responsible for their evolution-
ary and ecological success. The water-oxidizing photo-
synthesis is accomplished by coupling the activity of
photosystem (PS) II and I (Z-scheme). In anaerobic con-
ditions, they are also able to use only PS I — cyclic pho-
tophosphorylation — with electron donors other than
water (hydrogen sulfide, thiosulphate, or even molecu-
lar hydrogen) just like purple photosynthetic bacteria.
Furthermore, they share an archaeal property, the abil-
ity to reduce elemental sulfur by anaerobic respiration
in the dark. Their photosynthetic electron transport
shares the same compartment as the components of res-
piratory electron transport. Actually, their plasma mem-
brane contains only components of the respiratory
chain, while the thylakoid membrane hosts both respir-
atory and photosynthetic electron transport.
Attached to thylakoid membrane, phycobilisomes
act as light harvesting antennae for the photosystems .
The phycobilisome components (phycobiliproteins) are
responsible for the blue-green pigmentation of most cy-
anobacteria. The variations to this theme is mainly due
to carotenoids and phycoerythrins which give the cells
the red-brownish coloration. In some cyanobacteria, the
color of light influences the composition of phycobili-
somes. In green light, the cells accumulate more
2
Cyanobacteria
phycoerythrin, whereas in red light they produce more
phycocyanin. Thus the bacteria appear green in red light
and red in green light. This process is known as comple-
mentary chromatic adaptation and is a way for the cells
to maximize the use of available light for
photosynthesis.
A few genera, however, 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 cyanobac-
teria. For this reason they are now considered as part of
the cyanobacterial group.
Relationship to chloroplasts
Gloeobacter
Prochlorococcus
Synechococcus
plastids
all other cyanobacteria
Cladogram showing plastids (chloroplasts
and similar) and basal cyanobacteria.[9]
Chloroplasts found in eukaryotes (algae and plants)
likely evolved from an endosymbiotic relation with cy-
anobacteria. This endosymbiotic theory is supported by
various structural and genetic similarities. Primary
chloroplasts are found among the green plants, where
they contain chlorophyll b, and among the red algae and
glaucophytes, where they contain phycobilins. It now
appears that these chloroplasts probably had a single
origin, in an ancestor of the clade called Primoplantae.
Other algae likely took their chloroplasts from these
forms by secondary endosymbiosis or ingestion.
It was once thought that the mitochondria in euka-
ryotes also developed from an endosymbiotic relation-
ship with cyanobacteria; however, it is now suspected
that this evolutionary event occurred when aerobic bac-
teria were engulfed by anaerobic host cells. Mitochon-
dria are believed to have originated not from cyanobac-
teria but from an ancestor of Rickettsia.
Relationship to Earth history
The biochemical capacity to use water as the source for
electrons in photosynthesis evolved once, in a common
ancestor of extant cyanobacteria. The geologic record
From Wikipedia, the free encyclopedia
Oncolites; Guilmette Formation (Late Devonian) near Hancock
Summit, Pahranagat Range, Nevada.
indicates that this transforming event took place early
in our planet’s history, at least 2450-2320 million years
ago (mya), and probably much earlier. Geobiological in-
terpretation of Archean (>2500 mya) sedimentary rocks
remains a challenge; available evidence indicates that
life existed 3500 mya, but the question of when oxygenic
photosynthesis evolved continues to engender debate
and research. A clear paleontological window on cy-
anobacterial evolution opened about 2000 mya, reveal-
ing an already diverse biota of blue-greens. Cyanobac-
teria remained principal primary producers throughout
the Proterozoic Eon (2500-543 mya), in part because the
redox structure of the oceans favored photautotrophs
capable of nitrogen fixation. Green algae joined blue-
greens as major primary producers on continental
shelves near the end of the Proterozoic, but only with
the Mesozoic (251-65 mya) radiations of dinoflagellates,
coccolithophorids, and diatoms did primary production
in marine shelf waters take modern form. The most
common cyanobacterial structures in the fossil record
include stromatolites and oncolites. Cyanobacteria re-
main critical to marine ecosystems as primary producers
in oceanic gyres, as agents of biological nitrogen fixa-
tion, and, in modified form, as the plastids of marine al-
gae.[10]
Classification
The cyanobacteria were traditionally classified by mor-
phology into five sections, referred to by the numerals I-
V. The first three - Chroococcales, Pleurocapsales, and
Oscillatoriales - are not supported by phylogenetic stud-
ies. However, the latter two - Nostocales and Sti-
gonematales - are monophyletic, and make up the het-
erocystous cyanobacteria. The members of Chroococales
are unicellular and usually aggregate in colonies. The
classic taxonomic criterion has been the cell morpho-
logy and the plane of cell division. In Pleurocapsales, the
cells have the ability to form internal spores (baeocytes).
3
Cyanobacteria
The rest of the sections include filamentous species. In
Oscillatoriales, the cells are uniseriately arranged and do
not form specialized cells (akinetes and heterocysts). In
Nostocales and Stigonematales the cells have the ability
to develop heterocysts in certain conditions. Sti-
gonematales, unlike Nostocales, includes species with
truly branched trichomes. Most taxa included in the
phylum or division Cyanobacteria have not yet been val-
idly published under the Bacteriological Code. Except:
• The classes Chroobacteria, Hormogoneae and
Gloeobacteria
• The orders Chroococcales, Gloeobacterales,
Nostocales, Oscillatoriales, Pleurocapsales and
Stigonematales
• The families Prochloraceae and Prochlorotrichaceae
• The genera Halospirulina, Planktothricoides,
Prochlorococcus, Prochloron, Prochlorothrix.
Biotechnology and applications
The unicellular cyanobacterium Synechocystis sp.
PCC6803 was the third prokaryote and first photosyn-
thetic organism whose genome was completely se-
quenced.[11] It continues to be an important model or-
ganism.[12] The smallest genomes have been found in
Prochlorococcus spp. (1.7 Mb)[13][14] and the largest in
Nostoc punctiforme (9 Mb)[15]. Those of Calothrix spp. are
estimated at 12-15 Mb,[16] as large as yeast.
Some cyanobacteria are sold as food, notably Aph-
anizomenon flos-aquae and Arthrospira platensis
(Spirulina).[17]
Health risks
Certain cyanobacteria produce cyanotoxins like
anatoxin-a, anatoxin-as, aplysiatoxin, cylindrospermop-
sin, domoic acid, microcystin LR, nodularin R (from Nod-
ularia), or saxitoxin. Sometimes a mass-reproduction of
cyanobacteria results in algal blooms.
These toxins can be neurotoxins, hepatotoxins, cyto-
toxins, and endotoxins, and can be dangerous to animals
and humans. Several cases of human poisoning have
been documented but a lack of knowledge prevents an
accurate assessment of the risks.[18][19]
See also
• Archean
• Cyanobiont
• Hypolith
• Oxygen Catastrophe
• Proterozoic
• Bacterial Circadian Rhythms
From Wikipedia, the free encyclopedia
References
[1] Ahoren Oren (2004). "A proposal for further integration
of the cyanobacteria under the Bacteriological Code". Int.
J. Syst. Evol. Microbiol. 54: 1895–1902. doi:10.1099/
ijs.0.03008-0. PMID 15388760.
[2] Betsey Betsey Dexter Dyer, 2003 A Field Guide to
Bacteria, Cornell University Press ISBN 0801488540
[3] C. Michael Hogan (2008) Makgadikgadi, The
Megalithic Portal, ed. A. Burnham
[4] Olson JM (2006). "Photosynthesis in the Archean era".
Photosyn. Res. 88 (2): 109–17. doi:10.1007/
s11120-006-9040-5. PMID 16453059.
[5] United Nations Conference on Trade and
Development
[6] J. Sun, et al. (2004). "Is autoinducer-2 a universal signal
for interspecies communication? A comparative genomic
and phylogenetic analysis of the synthesis and signal
transduction pathways". BMC Evol. Biol. 4: 36.
doi:10.1186/1471-2148-4-36.
[7] E. Dittmann, et al. (2001). "Altered expression of two
light-dependent genes in a microcystin-lacking mutant
of Microcystis aeruginosa PCC7806". Microbiology 147:
3113–3119.
[8] Steve Nadis, The Cells That Rule the Seas, Scientific
American, Nov. 2003 [1]
[9] Enrique Flores AH (2008). The Cyanobacteria: Molecular
Biology, Genomics and Evolution. Horizon. pp. 3. ISBN
1904455158. http://books.google.com/
Further reading
books?hl=sv&lr=&id=xgMahO1BXrQC&oi=fnd&pg=PA1&ots=m58kg-1Qno&sig=mt9OxD-__GmdITpqr11TtyLfzkM#PPA3,M1.
[10] Herrero A and Flores E (editor). (2008). The
Cyanobacteria: Molecular Biology, Genomics and
Evolution (1st ed.). Caister Academic Press. ISBN
978-1-904455-15-8 . http://www.horizonpress.com/cyan.
[11] T. Kaneko, et al. (1996). "Kaneko, T. et al. (1996) Sequence
analysis of the genome of the unicellular cyanobacterium
Synechocystis sp. strain PCC6803. II. Sequence
determination of the entire genome and assignment of
potential protein-coding regions". DNA Res. 3: 109–136.
doi:10.1093/dnares/3.3.109. PMID 8905231.
[12] Tabei Y, Okada K, Tsuzuki M (2007). "Sll1330 controls the
expression of glycolytic genes in Synechocystis sp. PCC
6803". Biochem. Biophys. Res. Commun. 355 (4): 1045–50.
doi:10.1016/j.bbrc.2007.02.065. PMID 17331473.
[13] G. Rocap, et al. (2003). "Genome divergence in two
Prochlorococcus ecotypes reflects oceanic niche
differentiation". Nature 424: 1042–1047. doi:10.1038/
nature01947.
External links
[14] A. Dufresne, et al. (2003). "Genome sequence of the
cyanobacterium Prochlorococcus marinus SS120, a
nearly minimal oxyphototrophic genome.". Proc. Natl
Retrieved from "http://en.wikipedia.org/wiki/Cyanobacteria"
Categories: Phototrophic bacteria, Cyanobacteria, Photosynthesis
4
Cyanobacteria
Acad. Sci. USA 100: 10020–10025. doi:10.1073/
pnas.1733211100. PMID 12917486.
[15]J.C. Meeks, et al. (2001). "An overview of the genome of
Nostoc punctiforme, a multicellular, symbiotic
cyanobacterium". Photosynth. Res. 70: 85–106 A number
of important advances have occurred in cyanobacterial
biotechnology in the recent years. World wide attention
is drawn towards cyanobacteria for their possible use in
mariculture, food, feed, fuel, fertilizer, colourant,
production of various secondary metabolites including
vitamins, toxins, enzymes, pharmaceuticals,
pharmacological probes and pollution abatement. Only a
few cyanobacterial strains (including Spirulina) have
been well-characterized or exploited commercially
(Thajuddin and Subramanian. Cyanobacterial
biodiversity and potential applications in biotechnology.
CURRENT SCIENCE, VOL. 89, NO. 1, 10 JULY 2005).
doi:10.1023/A:1013840025518.
[16]M. Herdman, et al. (1979). "Genome size of
cyanobacteria". J. Gen. Microbiol. 111: 73–85.
[17]Spolaore P, Joannis-Cassan C, Duran E, Isambert A (2006).
"Commercial applications of microalgae". J. Biosci.
Bioeng. 101 (2): 87–96. doi:10.1263/jbb.101.87. PMID
16569602. http://www.jstage.jst.go.jp/article/jbb/101/2/
101_87/_article.
[18]Cyanobacteria, their toxins and health risks
[19]Blue-Green Algae (Cyanobacteria) and their Toxins
• Gillian Cribbs (1997) Nature’s Superfood, the Blue-Green
Algae Revolution. Newleaf. ISBN 0-7522-0569-2
• Marshall Savage, (1992, 1994) The Millennial Project:
Colonizing the Galaxy in Eight Easy Steps. Little,
Brown. ISBN 0-316-77163-5
• Fogg, G.E., Stewart, W.D.P., Fay, P.and Walsby, A.E.
1973. The Blue-green Algae. Academic Press, London
and New York. ISBN 0-12-261650-2
• "Architects of the earth’s atmosphere." Introduction
to the Cyanobacteria. University of California,
Berkeley. 03 Feb. 2006.
• Whitton, B.A. Phylum Cyanophyta (Cyanobacteria).
in The Freshwater Algal Flora of the British Isles.
Cambridge University Press, Cambridge ISBN 0 521
77051 3
• Overview of cyanobacteria
• Webserver for Cyanobacteria Research
From Wikipedia, the free encyclopedia Cyanobacteria

This page was last modified on 15 May 2009, at 12:29 (UTC). All text is available under the terms of the GNU Free
Documentation License. (See Copyrights for details.) Wikipedia® is a registered trademark of the Wikimedia
Foundation, Inc., a U.S. registered 501(c)(3) tax-deductible nonprofit charity. Privacy policy About Wikipedia
Disclaimers