''Cyanothece'' is a genus of
unicellular
A unicellular organism, also known as a single-celled organism, is an organism that consists of a single cell, unlike a multicellular organism that consists of multiple cells. Organisms fall into two general categories: prokaryotic organisms and ...
,
diazotroph Diazotrophs are bacteria and archaea that fix gaseous nitrogen in the atmosphere into a more usable form such as ammonia.
A diazotroph is a microorganism that is able to grow without external sources of fixed nitrogen. Examples of organisms that ...
ic, oxygenic photosynthesizing
cyanobacteria.
Modern organisms and cellular organization
In 1976, Jiří Komárek defined the
prokaryotic
A prokaryote () is a single-celled organism that lacks a nucleus and other membrane-bound organelles. The word ''prokaryote'' comes from the Greek πρό (, 'before') and κάρυον (, 'nut' or 'kernel').Campbell, N. "Biology:Concepts & Connec ...
cyanobacteria genus ''Cyanothece'' as distinct from ''
Synechococcus
''Synechococcus'' (from the Greek ''synechos'', in succession, and the Greek ''kokkos'', granule) is a unicellular cyanobacterium that is very widespread in the marine environment. Its size varies from 0.8 to 1.5 µm. The photosynthetic c ...
'' NAG 1949.
Organisms in both genera share characteristics in addition to being oxygenic
phototrophs
Phototrophs () are organisms that carry out photon capture to produce complex organic compounds (e.g. carbohydrates) and acquire energy. They use the energy from light to carry out various cellular metabolic processes. It is a common misconcep ...
. They are both unicellular, forming aggregates, but not found in mucilaginous colonies.
They may have a thin
mucilage
Mucilage is a thick, gluey substance produced by nearly all plants and some microorganisms. These microorganisms include protists which use it for their locomotion. The direction of their movement is always opposite to that of the secretion of m ...
layer around each cell.
Both genera also divide by
binary fission
Binary may refer to:
Science and technology Mathematics
* Binary number, a representation of numbers using only two digits (0 and 1)
* Binary function, a function that takes two arguments
* Binary operation, a mathematical operation that ta ...
along an axis perpendicular to the cell's longitudinal axis.
A handful of characteristics distinguish the two genera. While ''Synechococcus'' species are usually cylindrical, ''Cyanothece'' species are normally oval and longer than 3 μm.,
''Cyanothece’s'' outer
cell wall layer is relatively thick and contains spherical, glassy
vesicles
Vesicle may refer to:
; In cellular biology or chemistry
* Vesicle (biology and chemistry), a supramolecular assembly of lipid molecules, like a cell membrane
* Synaptic vesicle
; In human embryology
* Vesicle (embryology), bulge-like features o ...
whose function has yet to be defined.
''Cyanothece’s'' nucleoids are spread loosely throughout the cell, with a net-like appearance.
Instead of
concentric thylakoid membranes that share a center or axis, ''Cyanothece’s'' exhibit short, wavy and
radially arranged.,
All ''Cyanothece'' had
nitrogenase
Nitrogenases are enzymes () that are produced by certain bacteria, such as cyanobacteria (blue-green bacteria) and rhizobacteria. These enzymes are responsible for the reduction of nitrogen (N2) to ammonia (NH3). Nitrogenases are the only fa ...
activity at one time; although some strains have lost the necessary genes.
During
nitrogen-fixing
Nitrogen fixation is a chemical process by which molecular nitrogen (), with a strong triple covalent bond, in the air is converted into ammonia () or related nitrogenous compounds, typically in soil or aquatic systems but also in industry. Atm ...
conditions, ''Cyanothece'' creates
inclusion storage bodies under the control of a
circadian rhythm.
Evolutionary history
Between 2.5 and 3.0 billion years ago, cyanobacteria started using the energy from light to split water, releasing
oxygen
Oxygen is the chemical element with the symbol O and atomic number 8. It is a member of the chalcogen group in the periodic table, a highly reactive nonmetal, and an oxidizing agent that readily forms oxides with most elements as ...
into the
anaerobic
Anaerobic means "living, active, occurring, or existing in the absence of free oxygen", as opposed to aerobic which means "living, active, or occurring only in the presence of oxygen." Anaerobic may also refer to:
* Anaerobic adhesive, a bonding a ...
,
reducing environment
A reducing atmosphere is an atmospheric condition in which oxidation is prevented by removal of oxygen and other oxidizing gases or vapours, and which may contain actively reducing gases such as hydrogen, carbon monoxide, and gases such as hydro ...
.
Parts of this ancient cyanobacterial
metabolism
Metabolism (, from el, μεταβολή ''metabolē'', "change") is the set of life-sustaining chemical reactions in organisms. The three main functions of metabolism are: the conversion of the energy in food to energy available to run c ...
are still maintained today.
Bandyopadhyay ''et al.'' 2011 created a
phylogenic tree for cyanobacteria using 226
homolog
In biology, homology is similarity due to shared ancestry between a pair of structures or genes in different taxa. A common example of homologous structures is the forelimbs of vertebrates, where the wings of bats and birds, the arms of prima ...
protein groups. They grouped five of the six major ''Cyanothece'' strains (PCC 7424, PCC 7822, ATCC 51142, PCC 8801, PCC 8802) as belonging to a single
clade, but had ''Cyanothece'' sp PCC 7425 branched off earlier. PCC 7425's
nitrogenase
Nitrogenases are enzymes () that are produced by certain bacteria, such as cyanobacteria (blue-green bacteria) and rhizobacteria. These enzymes are responsible for the reduction of nitrogen (N2) to ammonia (NH3). Nitrogenases are the only fa ...
cluster is arranged differently from the other five strains and can only fix nitrogen anaerobically.
Most other cyanobacteria may have lost their ability to fix nitrogen. As Earth's climate became more
oxidated, the process of fixing nitrogen became unfavorable, and natural selection eliminated some of the necessary genes for the
nitrogenase
Nitrogenases are enzymes () that are produced by certain bacteria, such as cyanobacteria (blue-green bacteria) and rhizobacteria. These enzymes are responsible for the reduction of nitrogen (N2) to ammonia (NH3). Nitrogenases are the only fa ...
protein complex to increase
evolutionary fitness
Fitness (often denoted w or ω in population genetics models) is the quantitative representation of individual reproductive success. It is also equal to the average contribution to the gene pool of the next generation, made by the same individua ...
.
Photosynthesis/pigments
Cyanobacteria turn energy from the sun into chemical energy through oxygenic
photosynthesis
Photosynthesis is a process used by plants and other organisms to convert light energy into chemical energy that, through cellular respiration, can later be released to fuel the organism's activities. Some of this chemical energy is stored i ...
. Their
light-harvesting complex
A light-harvesting complex consists of a number of chromophores which are complex subunit proteins that may be part of a larger super complex of a photosystem, the functional unit in photosynthesis. It is used by plants and photosynthetic bacteri ...
that captures the photons usually includes the pigments
chlorophyll a and
phycocyanin
Phycocyanin is a pigment-protein complex from the light-harvesting phycobiliprotein family, along with allophycocyanin and phycoerythrin. It is an accessory pigment to chlorophyll. All phycobiliproteins are water-soluble, so they cannot exist w ...
. A cyanobacterium's typical blue-green color is a result of the combination of these two pigments. Three ''Cyanothece'' strains, sp. PCC 7424, 7822 and 8801, have the additional pigment
phycoerythrin, which expands the wavelengths of light these species use for energy. Phycoerythrin also gives these three species a brownish-green color.
The rate of oxygen created by
photosystem II is much higher when ''Cyanothece'' does not fix nitrogen (when the medium is nitrogen-replete).
The genera's
circadian rhythm controls photosynthetic oxygen generation by regulating when the proteins for their photosynthetic machinery are produced.
This
diurnal oscillation occurs even when the organisms are kept in the light continuously
or in the dark continuously.
Photosynthesis is
downregulated
In the biological context of organisms' production of gene products, downregulation is the process by which a cell decreases the quantity of a cellular component, such as RNA or protein, in response to an external stimulus. The complementary proc ...
when the nitrogen-fixing enzyme, nitrogenase, is
upregulated
In the biological context of organisms' production of gene products, downregulation is the process by which a cell decreases the quantity of a cellular component, such as RNA or protein, in response to an external stimulus. The complementary pro ...
. Decreasing the oxygen in the cell allows the oxygen-sensitive nitrogenase to fix nitrogen from the air for the organism's needs.
Metabolism, biosynthesis, symbiosis
''Cyanothece'' balances the production of oxygen through photosynthesis and oxygen-sensitive nitrogen fixation and fermentation all in one cell. They accomplish this by separating the two processes in time under the control of their circadian rhythm.
During the day, they use the energy harnessed from photosynthesis to produce the carbohydrate
glycogen, which is stored in
granules.
At night, the organisms break down the glycogen, providing the energy for nitrogen fixation.
In a very energy-intensive process, nitrogenase is first synthesized
and then takes N
2 from the air, combining it with
protons
A proton is a stable subatomic particle, symbol , H+, or 1H+ with a positive electric charge of +1 ''e'' elementary charge. Its mass is slightly less than that of a neutron and 1,836 times the mass of an electron (the proton–electron mas ...
and
electrons
The electron ( or ) is a subatomic particle with a negative one elementary electric charge. Electrons belong to the first generation of the lepton particle family,
and are generally thought to be elementary particles because they have no ...
to produce
ammonia
Ammonia is an inorganic compound of nitrogen and hydrogen with the formula . A stable binary hydride, and the simplest pnictogen hydride, ammonia is a colourless gas with a distinct pungent smell. Biologically, it is a common nitrogenous wa ...
and
hydrogen
Hydrogen is the chemical element with the symbol H and atomic number 1. Hydrogen is the lightest element. At standard conditions hydrogen is a gas of diatomic molecules having the formula . It is colorless, odorless, tasteless, non-toxic ...
gas. The organisms also store
cyanophycin
Cyanophycin, also known as CGP (cyanophycin granule polypeptide) or multi-L-arginyl-poly (L-aspartic acid), is a non-protein, non-ribosomally produced amino acid polymer composed of an aspartic acid backbone and arginine side groups.
Cyanophycin ...
, a nitrogen-reserve molecule which is a polymer of
arginine and
asparagine
Asparagine (symbol Asn or N) is an α-amino acid that is used in the biosynthesis of proteins. It contains an α-amino group (which is in the protonated −NH form under biological conditions), an α-carboxylic acid group (which is in the depro ...
, for use by the organism during the day.
Different ''Cyanothece'' species metabolize nitrogen-containing compounds through a variety of pathways; all have an
arginine decarboxylase, but vary after that point.
To provide the
anoxic environment needed by nitrogenase, ''Cyanothece'' boosts its
respiration
Respiration may refer to:
Biology
* Cellular respiration, the process in which nutrients are converted into useful energy in a cell
** Anaerobic respiration, cellular respiration without oxygen
** Maintenance respiration, the amount of cellul ...
as night begins by using its glycogen stores
while turning off photosynthesis.
In addition, the organisms produce
peroxidases
Peroxidases or peroxide reductases ( EC numberbr>1.11.1.x are a large group of enzymes which play a role in various biological processes. They are named after the fact that they commonly break up peroxides.
Functionality
Peroxidases typically ca ...
and
catalases which help scavenge any oxygen left in the cell.
The circadian rhythm ensures that this occurs even when the organism is growing in continuous light
or continuous darkness.
In the dark, the cyanobacteria act as
heterotroph
A heterotroph (; ) is an organism that cannot produce its own food, instead taking nutrition from other sources of organic carbon, mainly plant or animal matter. In the food chain, heterotrophs are primary, secondary and tertiary consumers, but ...
s, getting their energy and carbon from the
medium
Medium may refer to:
Science and technology
Aviation
*Medium bomber, a class of war plane
*Tecma Medium, a French hang glider design
Communication
* Media (communication), tools used to store and deliver information or data
* Medium of ...
. ''Cyanothece'' has the genes for the use of a variety of
sugar molecules;
although
glycerol
Glycerol (), also called glycerine in British English and glycerin in American English, is a simple triol compound. It is a colorless, odorless, viscous liquid that is sweet-tasting and non-toxic. The glycerol backbone is found in lipids known ...
is the only one that has been used successfully to grow ''Cyanothece'' in the dark.
Many of the genes that are unique to the genera have
homologs
A couple of homologous chromosomes, or homologs, are a set of one maternal and one paternal chromosome that pair up with each other inside a cell during fertilization. Homologs have the same genes in the same loci where they provide points alon ...
in
anaerobic
Anaerobic means "living, active, occurring, or existing in the absence of free oxygen", as opposed to aerobic which means "living, active, or occurring only in the presence of oxygen." Anaerobic may also refer to:
* Anaerobic adhesive, a bonding a ...
bacteria, including those responsible for
formate production through mixed-acid fermentation and also
fermentative lactate production.
Some ''Cyanothece'' species also are capable of
tryptophan
Tryptophan (symbol Trp or W)
is an α-amino acid that is used in the biosynthesis of proteins. Tryptophan contains an α-amino group, an α-carboxylic acid group, and a side chain indole, making it a polar molecule with a non-polar aromatic ...
degradation,
methionine salvage, conversion of stored
lipids
Lipids are a broad group of naturally-occurring molecules which includes fats, waxes, sterols, fat-soluble vitamins (such as vitamins A, D, E and K), monoglycerides, diglycerides, phospholipids, and others. The functions of lipids include ...
into
carbohydrates,
alkane and higher
alcohol synthesis, and
phosphonate
In organic chemistry, phosphonates or phosphonic acids are organophosphorus compounds containing groups (where R = alkyl, aryl, or just hydrogen). Phosphonic acids, typically handled as salts, are generally nonvolatile solids that are poorly ...
metabolism.
They can switch between a
photoautotrophic Photoautotrophs are organisms that use light energy and inorganic carbon to produce organic materials. Eukaryotic photoautotrophs absorb energy through the chlorophyll molecules in their chloroplasts while prokaryotic photoautotrophs use chlorophyll ...
and
photoheterotrophic metabolism depending on the environmental conditions that maximize their growth, employing the pathways that use the least amount of energy.
Genome size, organization, and ploidy options
The
genomes
In the fields of molecular biology and genetics, a genome is all the genetic information of an organism. It consists of nucleotide sequences of DNA (or RNA in RNA viruses). The nuclear genome includes protein-coding genes and non-coding g ...
of many of ''Cyanothece'' species have been sequenced, ranging from 4.79 to 7.84 Mbp. Between 4367 and 6642 coding sequences are an amalgamation of genes encoding capabilities for
fermentation and aerobic nitrogen fixation (like
filamentous cyanobacteria).
Unusually, the genes for nitrogen fixation are in a large, contiguous
cluster
may refer to:
Science and technology Astronomy
* Cluster (spacecraft), constellation of four European Space Agency spacecraft
* Asteroid cluster, a small asteroid family
* Cluster II (spacecraft), a European Space Agency mission to study t ...
(under the control of multiple
promoters),
including genes for the
uptake hydrogenase,
regulators
Regulator may refer to:
Technology
* Regulator (automatic control), a device that maintains a designated characteristic, as in:
** Battery regulator
** Pressure regulator
** Diving regulator
** Voltage regulator
* Regulator (sewer), a control de ...
, and
transporters.
The organism's robust circadian rhythm is apparent in the co-ordination of
transcription
Transcription refers to the process of converting sounds (voice, music etc.) into letters or musical notes, or producing a copy of something in another medium, including:
Genetics
* Transcription (biology), the copying of DNA into RNA, the fir ...
of correlated processes.
Using
microarrays
A microarray is a multiplex lab-on-a-chip. Its purpose is to simultaneously detect the expression of thousands of genes from a sample (e.g. from a tissue). It is a two-dimensional array on a solid substrate—usually a glass slide or silicon ...
, about 30% of 5000 genes tested exhibited
diurnal oscillations in 12-hour light/dark conditions, while 10% continued the behavior in continuous light.
About 1,705 of the gene groups are >99.5%
homologous with other cyanobacteria genera, largely ''
Microcystis
''Microcystis'' is a genus of freshwater cyanobacteria that includes the harmful algal bloom-forming ''Microcystis aeruginosa''. Many members of a ''Microcystis'' community can produce neurotoxins and hepatotoxins, such as microcystin and cyan ...
'' and filamentous, nitrogen-fixing strains. Typical
GC content
In molecular biology and genetics, GC-content (or guanine-cytosine content) is the percentage of nitrogenous bases in a DNA or RNA molecule that are either guanine (G) or cytosine (C). This measure indicates the proportion of G and C bases out ...
is about 40%.
''Cyanothece'' species also have three to six
plasmids ranging between 10 and 330 kb.
Unique to some species of this genus is one to three linear pieces of
DNA.
The linear DNA encodes enzymes for glucose and
pyruvate metabolism
(recall that
glycerol
Glycerol (), also called glycerine in British English and glycerin in American English, is a simple triol compound. It is a colorless, odorless, viscous liquid that is sweet-tasting and non-toxic. The glycerol backbone is found in lipids known ...
is the only organic carbon source on which ''Cyanothece'' has grown successfully
),
lactate fermentation,
transposons
A transposable element (TE, transposon, or jumping gene) is a nucleic acid sequence in DNA that can change its position within a genome, sometimes creating or reversing mutations and altering the cell's genetic identity and genome size. Tran ...
, and
CRISPR proteins.
''Cyanothece'' species do not typically use
homologous recombination
Homologous recombination is a type of genetic recombination in which genetic information is exchanged between two similar or identical molecules of double-stranded or single-stranded nucleic acids (usually DNA as in cellular organisms but may ...
, which greatly hinders genetic manipulation; an exception is ''Cyanothece'' sp. PCC 7822 in which
gene knock-outs can be generated.
Cell size, growth patterns, sex
''Cyanothece'' species are normally
oval
An oval () is a closed curve in a plane which resembles the outline of an egg. The term is not very specific, but in some areas (projective geometry, technical drawing, etc.) it is given a more precise definition, which may include either one ...
and longer than 3 μm.
They
double
A double is a look-alike or doppelgänger; one person or being that resembles another.
Double, The Double or Dubble may also refer to:
Film and television
* Double (filmmaking), someone who substitutes for the credited actor of a character
* ...
in 10 to 14 hours in the presence of
nitrate, when they do not need to use energy to fix nitrogen, and 16 to 20 hours when fixing nitrogen.
They divide by
binary fission
Binary may refer to:
Science and technology Mathematics
* Binary number, a representation of numbers using only two digits (0 and 1)
* Binary function, a function that takes two arguments
* Binary operation, a mathematical operation that ta ...
in one plane that is perpendicular to their longitudinal axis.
Daughter cells remain joined for just a short time after division.
Cell division
Cell division is the process by which a parent cell divides into two daughter cells. Cell division usually occurs as part of a larger cell cycle in which the cell grows and replicates its chromosome(s) before dividing. In eukaryotes, there ar ...
does not proceed until the daughter cells reach their mature size and original shape.
Habitat range
''Cyanothece'' has been found in a variety of environments all over the world. One point in common is that the
pH is usually lower than 7.
Typically they are associated with
water
Water (chemical formula ) is an Inorganic compound, inorganic, transparent, tasteless, odorless, and Color of water, nearly colorless chemical substance, which is the main constituent of Earth's hydrosphere and the fluids of all known living ...
in
benthic marine environments,
rice fields
A paddy field is a flooded field of arable land used for growing semiaquatic crops, most notably rice and taro. It originates from the Neolithic rice-farming cultures of the Yangtze River basin in southern China, associated with pre-Aust ...
,
acidic marshes,
peaty bogs,
intertidal zones,
moors
The term Moor, derived from the ancient Mauri, is an exonym first used by Christian Europeans to designate the Muslim inhabitants of the Maghreb, the Iberian Peninsula, Sicily and Malta during the Middle Ages.
Moors are not a distinct or ...
and clear
lake
A lake is an area filled with water, localized in a basin, surrounded by land, and distinct from any river or other outlet that serves to feed or drain the lake. Lakes lie on land and are not part of the ocean, although, like the much large ...
s,
but sometimes are found in
mountain
A mountain is an elevated portion of the Earth's crust, generally with steep sides that show significant exposed bedrock. Although definitions vary, a mountain may differ from a plateau in having a limited summit area, and is usually highe ...
soils
Soil, also commonly referred to as earth or dirt, is a mixture of organic matter, minerals, gases, liquids, and organisms that together support life. Some scientific definitions distinguish ''dirt'' from ''soil'' by restricting the former ter ...
.
Walls and resting cysts
''Cyanothece'' species have a thin mucilaginous layer around a thick