|Stained micrograph of Thiomargarita namibiensis|
Schulz et al., 1999
Thiomargarita namibiensis is a gram-negative coccoid Proteobacterium, found in the ocean sediments of the continental shelf of Namibia. It is one of the largest bacteria ever discovered, as a rule 0.1–0.3 mm (100–300 µm) in diameter, but sometimes attaining 0.75 mm (750 µm). Cells of Thiomargarita namibiensis are large enough to be visible to the naked eye. Although the species holds the record for the most massive bacterium, Epulopiscium fishelsoni – previously discovered in the gut of surgeonfish – grows slightly longer, but narrower.
Thiomargarita means "sulfur pearl". This refers to the appearance of the cells; they contain microscopic sulfur granules that scatter incandescent light, lending the cell a pearly lustre. Like many coccoid bacteria such as Streptococcus 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".
The species was discovered by Heide N. Schulz and others in 1997, in the coastal seafloor sediments of Walvis Bay (Namibia). Schulz and her colleagues, from the Max Planck Institute of Marine Biology, 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 and Beggiatoa. In the end 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. 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. No other species in the genus Thiomargarita are known at present.
Although Thiomargarita namibiensis 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 they have adapted by evolving very large nitrate-storing bubbles, called vacuoles, allowing them to survive long periods of nitrate and sulfide starvation. The vacuoles give Thiomargarita 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 diffusion to move chemicals around, a process that works only over tiny distances. This implies that the cytoplasm has to be close to the cell wall, greatly limiting their size. But Thiomargarita are an exception to this size constraint as their cytoplasms form along the peripheral cell membrane, while the nitrate-storing vacuoles occupy the cells of the Thiomargarita. 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.
The bacterium is chemolithotrophic, and is capable of using nitrate as the terminal electron acceptor in the electron transport chain. The organism will oxidize hydrogen sulfide (H2S) into elemental sulfur (S). This is deposited as granules in its cytoplasm 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 that sank down to the sea bottom, the nitrate comes from the above seawater. Since the bacterium is sessile, 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, 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.
Gigantism 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 eukaryotes. 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.
- "The largest Bacterium: Scientist discovers new bacterial life form off the African coast", Max Planck Institute for Marine Microbiology, 8 April 1999[dead link]
- List of Prokaryotic names with Standing in Nomenclature - Genus Thiomargarita
- Amsden, Brandi, Thiomargarita namibiensis
- Karen M. Kalanetra, Samantha B. Joye, Nicole R. Sunseri, Douglas C. Nelson. Novel vacuolate sulfur bacteria from the Gulf of Mexico reproduce by reductive division in three dimensions. Environmental Microbiology (2005) 7 (9), 1451–1460 doi:10.1111/j.1462-2920.2005.00832.x
- Randerson, James (8 June 2002), "Record Breaker", New Scientist
- The World's Largest Bacteria, October 2001
- Schulz HN, Brinkhoff T, Ferdelman TG, Mariné MH, Teske A, Jorgensen BB (April 1999), "Dense populations of a giant sulfur bacterium in Namibian shelf sediments", Science 284 (5413): 493–5, doi:10.1126/science.284.5413.493, PMID 10205058.
- Kalanetra KM, Joye SB, Sunseri NR, Nelson DC (September 2005), "Novel vacuolate sulfur bacteria from the Gulf of Mexico reproduce by reductive division in three dimensions", Environ. Microbiol. 7 (9): 1451–60, doi:10.1111/j.1462-2920.2005.00832.x, PMID 16104867.
- The genus Thiomargarita. Heide Schulz. The Prokaryotes 2006, part 3, section 3.3, 1156-1163
- Schulz HN, de Beer D (November 2002), "Uptake Rates of Oxygen and Sulfide Measured with Individual Thiomargarita namibiensis Cells by Using Microelectrodes", Applied and Environmental Microbiology 68 (11): 5746–9, doi:10.1128/AEM.68.11.5746-5749.2002.
- Giant bacterium carries thousands of genomes. Nature News, 8 May 2008.
- Extreme polyploidy in a large bacterium. Proc Natl Acad Sci USA 2008, 105:6730-6734.