Thiomargarita Namibiensis
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''Thiomargarita namibiensis'' is a
Gram-negative Gram-negative bacteria are bacteria that do not retain the crystal violet stain used in the Gram staining method of bacterial differentiation. They are characterized by their cell envelopes, which are composed of a thin peptidoglycan cell wall ...
coccoid
bacterium Bacteria (; singular: bacterium) are ubiquitous, mostly free-living organisms often consisting of one biological cell. They constitute a large domain of prokaryotic microorganisms. Typically a few micrometres in length, bacteria were among ...
, found in the ocean sediments of the
continental shelf A continental shelf is a portion of a continent that is submerged under an area of relatively shallow water, known as a shelf sea. Much of these shelves were exposed by drops in sea level during glacial periods. The shelf surrounding an island ...
of
Namibia Namibia (, ), officially the Republic of Namibia, is a country in Southern Africa. Its western border is the Atlantic Ocean. It shares land borders with Zambia and Angola to the north, Botswana to the east and South Africa to the south and ea ...
. It is the second largest
bacterium Bacteria (; singular: bacterium) are ubiquitous, mostly free-living organisms often consisting of one biological cell. They constitute a large domain of prokaryotic microorganisms. Typically a few micrometres in length, bacteria were among ...
ever discovered, as a rule in diameter, but sometimes attaining . Cells of ''Thiomargarita namibiensis'' are large enough to be visible to the naked eye. Although the species held the record for the largest known bacterium, ''
Epulopiscium fishelsoni "''Candidatus'' Epulonipiscium" is a genus of Gram-positive bacteria that have a symbiotic relationship with surgeonfish. These bacteria are known for their unusually large size, many ranging from 200–700 μm in length. Until the discovery of ...
'' – previously discovered in the gut of
surgeonfish Acanthuridae are the family of surgeonfishes, tangs, and unicornfishes. The family includes about 86 extant species of marine fish living in tropical seas, usually around coral reefs. Many of the species are brightly colored and popular in ...
– grows slightly longer, but narrower. ''Thiomargarita'' means "sulfur pearl". This refers to the appearance of the cells; they contain microscopic sulfur granules that scatter incident light, lending the cell a pearly lustre. Like many coccoid bacteria such as ''
Streptococcus ''Streptococcus'' is a genus of gram-positive ' (plural ) or spherical bacteria that belongs to the family Streptococcaceae, within the order Lactobacillales (lactic acid bacteria), in the phylum Bacillota. Cell division in streptococci occurs ...
'', their cellular division tends to occur along a single axis, causing their cells to form chains, rather like strings of pearls. The species name ''namibiensis'' means "of Namibia".


Occurrence

The species was discovered by Heide N. Schulz and others in 1997, in the coastal
seafloor sediments Marine sediment, or ocean sediment, or seafloor sediment, are deposits of insoluble particles that have accumulated on the seafloor. These particles have their origins in soil and rocks and have been transported from the land to the sea, mainly ...
of
Walvis Bay Walvis Bay ( en, lit. Whale Bay; af, Walvisbaai; ger, Walfischbucht or Walfischbai) is a city in Namibia and the name of the bay on which it lies. It is the second largest city in Namibia and the largest coastal city in the country. The c ...
(Namibia). Schulz and her colleagues, from the
Max Planck Institute for Marine Microbiology __NOTOC__ The Max Planck Institute for Marine Microbiology is located in Bremen, Germany. It was founded in 1992, almost a year after the foundation of its sister institute, the Max Planck Institute for Terrestrial Microbiology at Marburg. In 1996 ...
, were on a Russian research vessel, the ''Petr Kottsov'', when the white color of this microbe caught their interest. They were actually looking for other recently found sulfide-eating marine bacteria, ''
Thioploca ''Thioploca '' is a genus of filamentous sulphur-oxidizing bacteria which occurs along of coast off the west of South America. Was discovered in 1907 by R. Lauterborn classified as belonging to the order Thiotrichales, part of the Gammaproteobac ...
'' and ''
Beggiatoa ''Beggiatoa'' is a genus of ''Gammaproteobacteria'' belonging the order ''Thiotrichales,'' in the ''Pseudomonadota'' phylum. This genus was one of the first bacteria discovered by Ukrainian botanist Sergei Sergei Winogradsky, Winogradsky. During ...
''. They ended up with an entire new discovery, of a much larger cousin strain of the two other bacteria. In 2005, a closely related strain was discovered in the
Gulf of Mexico The Gulf of Mexico ( es, Golfo de México) is an oceanic basin, ocean basin and a marginal sea of the Atlantic Ocean, largely surrounded by the North American continent. It is bounded on the northeast, north and northwest by the Gulf Coast of ...
. Among other differences from the Namibian strain, the Mexican strain does not seem to divide along a single axis and accordingly does not form chains. The previously largest known bacterium was ''
Epulopiscium fishelsoni "''Candidatus'' Epulonipiscium" is a genus of Gram-positive bacteria that have a symbiotic relationship with surgeonfish. These bacteria are known for their unusually large size, many ranging from 200–700 μm in length. Until the discovery of ...
'', at 0.5 mm long. ''
Thiomargarita magnifica ''Thiomargarita magnifica'' is a species of sulfur-oxidizing gammaproteobacteria, found growing underwater on the detached leaves of red mangroves from the Guadeloupe archipelago in the Lesser Antilles. This filament-shaped bacteria is the large ...
'' described in 2022 is larger.


Structure

Although ''Thiomargarita'' is closely related to ''Thioploca'' and ''Beggiatoa'' in function, their structures proved to be vastly different. ''Thioploca'' and ''Beggiatoa'' cells are much smaller and grow tightly stacked on each other in long filaments. Their shape is necessary for them to shuttle down into the ocean sediments to find more sulfide and nitrate. In contrast, ''Thiomargarita'' grow in rows of separate single ball-shaped cells, not allowing them to have the range of mobility that ''Thioploca'' and ''Beggiota'' have. With their lack of movement, ''
Thiomargarita ''Thiomargarita'' is a genus (family Thiotrichaceae) which includes the vacuolate sulfur bacteria species ''Thiomargarita namibiensis'', ''Candidatus Thiomargarita nelsonii'', and ''Ca. Thiomargarita joergensii''. In 2022, scientists working ...
'' have adapted by evolving very large nitrate-storing bubbles, called
vacuole A vacuole () is a membrane-bound organelle which is present in plant and fungal cells and some protist, animal, and bacterial cells. Vacuoles are essentially enclosed compartments which are filled with water containing inorganic and organic mo ...
s, allowing them to survive long periods of nitrate and sulfide starvation. The vacuoles give them the ability to stay immobile, just waiting for nitrate-rich waters to sweep over them once again. These vacuoles are what account for the size that scientists had previously thought impossible. Scientists disregarded large bacterium, because bacteria rely on chemiosmosis across their membranes to make ATP, as the cells become larger, they make proportionately less ATP, thus energy production limits their size. ''Thiomargarita'' are an exception to this size constraint, as their cytoplasm forms along the periphery of the cell, while the nitrate-storing vacuoles occupy the center of the cell. As these vacuoles swell, they greatly contribute to the record-holding size. It holds the record for the world's largest bacteria, with a volume three million times more than that of the average bacteria.


Metabolism

The bacterium is
chemolithotroph Lithotrophs are a diverse group of organisms using an inorganic substrate (usually of mineral origin) to obtain reducing equivalents for use in biosynthesis (e.g., carbon dioxide fixation) or energy conservation (i.e., ATP production) via aerobi ...
ic and is capable of using
nitrate Nitrate is a polyatomic ion A polyatomic ion, also known as a molecular ion, is a covalent bonded set of two or more atoms, or of a metal complex, that can be considered to behave as a single unit and that has a net charge that is not zer ...
as the terminal electron acceptor in the
electron transport chain An electron transport chain (ETC) is a series of protein complexes and other molecules that transfer electrons from electron donors to electron acceptors via redox reactions (both reduction and oxidation occurring simultaneously) and couples th ...
. The organism will oxidize
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 ...
(H2S) into elemental
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 ...
(S). This is deposited as granules in its periplasm and is highly refractile and opalescent, making the organism look like a pearl. While the sulfide is available in the surrounding sediment, produced by other bacteria from dead
microalgae Microalgae or microphytes are microscopic algae invisible to the naked eye. They are phytoplankton typically found in freshwater and marine systems, living in both the water column and sediment. They are unicellular species which exist indiv ...
that sank down to the sea bottom, the nitrate comes from the above seawater. Since the bacterium is
sessile Sessility, or sessile, may refer to: * Sessility (motility), organisms which are not able to move about * Sessility (botany), flowers or leaves that grow directly from the stem or peduncle of a plant * Sessility (medicine), tumors and polyps that ...
, and the concentration of available nitrate fluctuates considerably over time, it stores nitrate at high concentration (up to 0.8 molar) in a large
vacuole A vacuole () is a membrane-bound organelle which is present in plant and fungal cells and some protist, animal, and bacterial cells. Vacuoles are essentially enclosed compartments which are filled with water containing inorganic and organic mo ...
like an inflated balloon, which is responsible for about 80% of its size. When nitrate concentrations in the environment are low, the bacterium uses the contents of its vacuole for respiration. Thus, the presence of a central vacuole in its cells enables a prolonged survival in sulfidic sediments. The non-motility of ''Thiomargarita'' cells is compensated by its large cellular size. Recent research has also indicated that the bacterium may be
facultatively anaerobic A facultative anaerobic organism is an organism that makes ATP by aerobic respiration if oxygen is present, but is capable of switching to fermentation if oxygen is absent. Some examples of facultatively anaerobic bacteria are ''Staphylococcus' ...
rather than obligately anaerobic, and thus capable of respiring with oxygen if it is plentiful.


Significance

Gigantism Gigantism ( el, γίγας, ''gígas'', "giant", plural γίγαντες, ''gígantes''), also known as giantism, is a condition characterized by excessive growth and height significantly above average. In humans, this condition is caused by ove ...
is usually a disadvantage for bacteria. Bacteria obtain their nutrients via simple diffusion process across their cell membrane, as they lack the sophisticated nutrient uptake mechanism found in
eukaryote Eukaryotes () are organisms whose cells have a nucleus. All animals, plants, fungi, and many unicellular organisms, are Eukaryotes. They belong to the group of organisms Eukaryota or Eukarya, which is one of the three domains of life. Bacte ...
s. A bacterium of large size would imply a lower ratio of cell membrane surface area to cell volume. This would limit the rate of uptake of nutrients to threshold levels. Large bacteria might starve easily unless they have a different backup mechanism. ''T. namibiensis'' overcomes this problem by harboring large vacuoles that can be filled up with life-supporting nitrates.


See also

*
Valonia ventricosa ''Valonia ventricosa'', also known as bubble algae, sea grape or sailor's eyeballs is a species of algae found in oceans throughout the world in tropical and subtropical regions, within the phylum Chlorophyta. It is one of the largest known unice ...
- a large, 5 centimetre-wide unicellular species of algae


References


External links


Macrophotos of ''Thiomargarita namibiensis''
{{Taxonbar, from=Q132118 Thiotrichales Bacteria described in 1999