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''Chlorobium'' is a
genus Genus ( plural genera ) is a taxonomic rank used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In the hierarchy of biological classification, genus com ...
of
green sulfur bacteria The green sulfur bacteria are a phylum of obligately anaerobic photoautotrophic bacteria that metabolize sulfur. Green sulfur bacteria are nonmotile (except ''Chloroherpeton thalassium'', which may glide) and capable of anoxygenic photosynthe ...
. They are photolithotrophic
oxidizer An oxidizing agent (also known as an oxidant, oxidizer, electron recipient, or electron acceptor) is a substance in a redox chemical reaction that gains or " accepts"/"receives" an electron from a (called the , , or ). In other words, an oxid ...
s of
sulfur Sulfur (or sulphur in British English) is a chemical element with the symbol S and atomic number 16. It is abundant, multivalent and nonmetallic. Under normal conditions, sulfur atoms form cyclic octatomic molecules with a chemical formula ...
and most notably utilise a noncyclic electron transport chain to reduce NAD+.
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 ...
is achieved using a Type 1 Reaction Centre using
bacteriochlorophyll Bacteriochlorophylls (BChl) are photosynthetic pigments that occur in various phototrophic bacteria. They were discovered by C. B. van Niel in 1932. They are related to chlorophylls, which are the primary pigments in plants, algae, and cyanoba ...
(BChl) ''a''. Two photosynthetic antenna complexes aid in light absorption: the Fenna-Matthews-Olson complex ("FMO", also containing BChl ''a''), and the
chlorosome A chlorosome is a photosynthetic antenna complex found in green sulfur bacteria (GSB) and some green filamentous anoxygenic phototrophs (FAP) ( Chloroflexaceae, Oscillochloridaceae; both members of Chloroflexia). They differ from other antenna ...
s which employ mostly BChl ''c'', ''d'', or ''e''.
Hydrogen sulfide Hydrogen sulfide is a chemical compound with the formula . It is a colorless chalcogen-hydride gas, and is poisonous, corrosive, and flammable, with trace amounts in ambient atmosphere having a characteristic foul odor of rotten eggs. The unde ...
is used as an electron source and
carbon dioxide Carbon dioxide (chemical formula ) is a chemical compound made up of molecules that each have one carbon atom covalently double bonded to two oxygen atoms. It is found in the gas state at room temperature. In the air, carbon dioxide is transpar ...
its carbon source.Prescott, Harley, Klein. (2005). ''Microbiology'' pp. 195, 493, 597, 618-619, 339. ''Chlorobium''
species In biology, a species is the basic unit of classification and a taxonomic rank of an organism, as well as a unit of biodiversity. A species is often defined as the largest group of organisms in which any two individuals of the appropriate s ...
exhibit a dark green color; in a
Winogradsky column The Winogradsky column is a simple device for culturing a large diversity of microorganisms. Invented in the 1880s by Sergei Winogradsky, the device is a column of pond mud and water mixed with a carbon source such as newspaper (containing cellul ...
, the green layer often observed is composed of ''Chlorobium''. This genus lives in strictly anaerobic conditions below the surface of a body of water, commonly 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 ...
zone of a
eutrophic lake The Trophic State Index (TSI) is a classification system designed to rate water bodies based on the amount of biological productivity they sustain. Although the term "trophic index" is commonly applied to lakes, any surface water body may be ind ...
. '' Chlorobium aggregatum'' is a species which exists in a symbiotic relationship with a colorless, nonphotosynthetic bacteria. This species looks like a bundle of green bacteria, attached to a central rod-like cell which can move around with a
flagellum A flagellum (; ) is a hairlike appendage that protrudes from certain plant and animal sperm cells, and from a wide range of microorganisms to provide motility. Many protists with flagella are termed as flagellates. A microorganism may have f ...
. The green, outer bacteria use light to oxidize
sulfide Sulfide (British English also sulphide) is an inorganic anion of sulfur with the chemical formula S2− or a compound containing one or more S2− ions. Solutions of sulfide salts are corrosive. ''Sulfide'' also refers to chemical compounds lar ...
into
sulfate The sulfate or sulphate ion is a polyatomic anion with the empirical formula . Salts, acid derivatives, and peroxides of sulfate are widely used in industry. Sulfates occur widely in everyday life. Sulfates are salts of sulfuric acid and many ar ...
. The inner cell, which is not able to perform
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 ...
, reduces the sulfate into sulfide. These bacteria divide in unison, giving the structure a
multicellular A multicellular organism is an organism that consists of more than one cell, in contrast to unicellular organism. All species of animals, land plants and most fungi are multicellular, as are many algae, whereas a few organisms are partially uni- ...
appearance which is highly unusual in bacteria. ''Chlorobium'' species are thought to have played an important part in
mass extinction An extinction event (also known as a mass extinction or biotic crisis) is a widespread and rapid decrease in the biodiversity on Earth. Such an event is identified by a sharp change in the diversity and abundance of multicellular organisms. It ...
events on Earth. If the oceans turn anoxic (due to the shutdown of ocean circulation) then ''Chlorobium'' would be able to out compete other photosynthetic life. They would produce huge quantities of
methane Methane ( , ) is a chemical compound with the chemical formula (one carbon atom bonded to four hydrogen atoms). It is a group-14 hydride, the simplest alkane, and the main constituent of natural gas. The relative abundance of methane on Eart ...
and
hydrogen sulfide Hydrogen sulfide is a chemical compound with the formula . It is a colorless chalcogen-hydride gas, and is poisonous, corrosive, and flammable, with trace amounts in ambient atmosphere having a characteristic foul odor of rotten eggs. The unde ...
which would cause
global warming In common usage, climate change describes global warming—the ongoing increase in global average temperature—and its effects on Earth's climate system. Climate change in a broader sense also includes previous long-term changes to E ...
and
acid rain Acid rain is rain or any other form of precipitation that is unusually acidic, meaning that it has elevated levels of hydrogen ions (low pH). Most water, including drinking water, has a neutral pH that exists between 6.5 and 8.5, but acid ...
. This would have huge consequences for other oceanic organisms and also for terrestrial organisms. Evidence for abundant ''Chlorobium'' populations is provided by chemical fossils found in sediments deposited at the
Cretaceous mass extinction The Cretaceous ( ) is a geological period that lasted from about 145 to 66 million years ago (Mya). It is the third and final period of the Mesozoic Era, as well as the longest. At around 79 million years, it is the longest geological period of th ...
.


Molecular signatures for ''Chlorobiota''

Comparative genomic analysis has led to the identification of 2
conserved signature indels Conserved signature inserts and deletions (CSIs) in protein sequences provide an important category of molecular markers for understanding phylogenetic relationships. CSIs, brought about by rare genetic changes, provide useful phylogenetic markers ...
which are uniquely found in members of the phylum ''Chlorobiota'' (formerly Chlorobi) and are thus characteristic of the phylum. The first indel is a 28-amino-acid insertion in
DNA polymerase III DNA polymerase III holoenzyme is the primary enzyme complex involved in prokaryotic DNA replication. It was discovered by Thomas Kornberg (son of Arthur Kornberg) and Malcolm Gefter in 1970. The complex has high processivity (i.e. the number of ...
and the second is a 12 to 14 amino acid insertion in alanyl-tRNA synthetase. These indels are not found in any other bacteria and thus serve as molecular markers for the phylum. In addition to the conserved signature indels, 51 proteins which are uniquely found in members of the phylum ''Chlorobiota''. 65 other proteins have been identified which are unique to the ''Chlorobiota'' phylum, however these proteins are missing in several ''Chlorobiota'' species and are not distributed throughout the phylum with any clear pattern. This means that significant gene loss may have occurred, or the presence of these proteins may be a result of
horizontal gene transfer Horizontal gene transfer (HGT) or lateral gene transfer (LGT) is the movement of genetic material between Unicellular organism, unicellular and/or multicellular organisms other than by the ("vertical") transmission of DNA from parent to offsprin ...
. Of these 65 proteins, 8 are found only in ''Chlorobium luteolum'' and ''Chlorobium phaeovibrioides''. These two species form a strongly supported clade in phylogenetic trees and a close relationship between these species is further supported by the unique sharing of these 8 proteins.


Relatedness of ''Chlorobiota'' to ''Bacteroidota'' and ''Fibrobacterota'' phyla

Species from the ''Bacteroidota'' (formerly Bacteroidetes) and ''Chlorobiota'' phyla branch very closely together in phylogenetic trees, indicating a close relationship. Through the use of comparative genomic analysis, 3 proteins have been identified which are uniquely shared by virtually all members of the ''Bacteroidota'' and ''Chlorobiota'' phyla. The sharing of these 3 proteins is significant because other than these 3 proteins, no proteins from either the'' Bacteroidota'' or ''Chlorobiota'' phyla are shared by any other groups of bacteria. Several conserved signature indels have also been identified which are uniquely shared by members of the ''Bacteroidota'' and ''Chlorobiota'' phyla. The presence of these molecular signatures supports the close relationship of the ''Bacteroidota'' and ''Chlorobiota'' phyla. Additionally, the phylum ''
Fibrobacterota Fibrobacterota is a small bacterial phylum which includes many of the major rumen bacteria, allowing for the degradation of plant-based cellulose in ruminant animals. Members of this phylum were categorized in other phyla. The genus '' Fibrobac ...
'' (formerly Fibrobacteres) is indicated to be specifically related to these two phyla. A clade consisting of these three phyla is strongly supported by phylogenetic analyses based upon a number of different proteins These phyla also branch in the same position based upon conserved signature indels in a number of important proteins. Lastly and most importantly, two conserved signature indels (in the RpoC protein and in
serine hydroxymethyltransferase Serine hydroxymethyltransferase (SHMT) is a pyridoxal phosphate (PLP) (Vitamin B6) dependent enzyme () which plays an important role in cellular one-carbon pathways by catalyzing the reversible, simultaneous conversions of L-serine to glycine ...
) and one signature protein PG00081 have been identified that are uniquely shared by all of the species from these three phyla. All of these results provide compelling evidence that the species from these three phyla shared a common ancestor exclusive of all other bacteria and it has been proposed that they should all recognized as part of a single “FCB”superphylum.


Phylogeny

The currently accepted taxonomy is based on the
List of Prokaryotic names with Standing in Nomenclature List of Prokaryotic names with Standing in Nomenclature (LPSN) is an online database that maintains information on the naming and taxonomy of prokaryotes, following the taxonomy requirements and rulings of the International Code of Nomenclature ...
(LPSN) and
National Center for Biotechnology Information The National Center for Biotechnology Information (NCBI) is part of the United States National Library of Medicine (NLM), a branch of the National Institutes of Health (NIH). It is approved and funded by the government of the United States. The ...
(NCBI)


See also

*
List of bacterial orders This article lists the orders of the Bacteria. The currently accepted taxonomy is based on the List of Prokaryotic names with Standing in Nomenclature (LPSN) and National Center for Biotechnology Information (NCBI) and the phylogeny is based on 16S ...
*
List of bacteria genera This article lists the genera of the bacteria. The currently accepted taxonomy is based on the List of Prokaryotic names with Standing in Nomenclature (LPSN) and National Center for Biotechnology Information (NCBI). However many taxonomic names are ...


References


External links


''Chlorobium'' at Bac''Dive'' - the Bacterial Diversity Metadatabase
{{Taxonbar, from=Q2725095 Phototrophic bacteria Chlorobiota Bacteria genera