Big bacteria.

A small number of prokaryotic species have a unique physiology or ecology related to their development of unusually large size. The biomass of bacteria varies over more than 10 orders of magnitude, from the 0.2 microm wide nanobacteria to the largest cells of the colorless sulfur bacteria, Thiomargarita namibiensis, with a diameter of 750 microm. All bacteria, including those that swim around in the environment, obtain their food molecules by molecular diffusion. Only the fastest and largest swimmers known, Thiovulum majus, are able to significantly increase their food supply by motility and by actively creating an advective flow through the entire population. Diffusion limitation generally restricts the maximal size of prokaryotic cells and provides a selective advantage for microm-sized cells at the normally low substrate concentrations in the environment. The largest heterotrophic bacteria, the 80 x 600 microm large Epulopiscium sp. from the gut of tropical fish, are presumably living in a very nutrient-rich medium. Many large bacteria contain numerous inclusions in the cells that reduce the volume of active cytoplasm. The most striking examples of competitive advantage from large cell size are found among the colorless sulfur bacteria that oxidize hydrogen sulfide to sulfate with oxygen or nitrate. The several-cm-long filamentous species can penetrate up through the ca 500-microm-thick diffusive boundary layer and may thereby reach into water containing their electron acceptor, oxygen or nitrate. By their ability to store vast quantities of both nitrate and elemental sulfur in the cells, these bacteria have become independent of the coexistence of their substrates. In fact, a close relative, T. namibiensis, can probably respire in the sulfidic mud for several months before again filling up their large vacuoles with nitrate.

Dense populations of a giant sulfur bacterium in Namibian shelf sediments.

A previously unknown giant sulfur bacterium is abundant in sediments underlying the oxygen minimum zone of the Benguela Current upwelling system. The bacterium has a spherical cell that exceeds by up to 100-fold the biovolume of the largest known prokaryotes. On the basis of 16S ribosomal DNA sequence data, these bacteria are closely related to the marine filamentous sulfur bacteria Thioploca, abundant in the upwelling area off Chile and Peru. Similar to Thioploca, the giant bacteria oxidize sulfide with nitrate that is accumulated to

Community Structure of Filamentous, Sheath-Building Sulfur Bacteria, Thioploca spp., off the Coast of Chile.

The filamentous sulfur bacteria Thioploca spp. produce dense bacterial mats in the shelf area off the coast of Chile and Peru. The mat consists of common sheaths, shared by many filaments, that reach 5 to 10 cm down into the sediment. The structure of the Thioploca communities off the Bay of Concepcion was investigated with respect to biomass, species distribution, and three-dimensional orientation of the sheaths. Thioploca sheaths and filaments were found across the whole shelf area within the oxygen minimum zone. The maximum wet weight of sheaths, 800 g m(sup-2), was found at a depth of 90 m. The bacterial filaments within the sheaths contributed about 10% of this weight. The highest density of filaments was found within the uppermost 1 cm of the mat. On the basis of diameter classes, it was possible to distinguish populations containing only Thioploca spp. from mixed populations containing Beggiatoa spp. Three distinct size classes of Thioploca spp. were found, two of which have been described previously as Thioploca araucae and Thioploca chileae. Many Thioploca filaments did not possess a visible sheath, and about 20% of the sheaths contained more than one Thioploca species. The three-dimensional sheath structure showed that Thioploca filaments can move from the surface and deep into the sediment.