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 photosynthesis. They live in anaerobic aquatic environments. In contrast to plants, green sulfur bacteria mainly use sulfide ions as electron donors. They are autotrophs that utilize the reverse tricarboxylic acid cycle to perform carbon fixation. They are also mixotrophs and reduce nitrogen. Characteristics. Green sulfur bacteria are gram-negative rod or spherical shaped bacteria. Some types of green sulfur bacteria have gas vacuoles that allow for movement . They are photolithoautotrophs, and use light energy and reduced sulfur compounds as the electron source. Electron donors include H, HS, S. The major photosynthetic pigment in these bacteria is Bacteriochlorophylls "c" or "d" in green species and "e" in brown species, and is located in the chlorosomes and plasma membranes. Chlorosomes are a unique feature that allow them to capture light in low-light conditions. Habitat. The majority of green sulfur bacteria are mesophilic, preferring moderate temperatures, and all live in aquatic environments. They require anaerobic conditions and reduced sulfur; they are usually found in the top millimeters of sediment . They are capable of photosynthesis in low light conditions. The Black Sea, an extremely anoxic environment, was found to house a large population of green sulfur bacteria at about 100 m depth. Due to the lack of light available in this region of the sea, most bacteria were photosynthetically inactive. The photosynthetic activity detected in the sulfide chemocline suggests that the bacteria need very little energy for cellular maintenance. A species of green sulfur bacteria has been found living near a black smoker off the coast of Mexico at a depth of 2,500 m in the Pacific Ocean. At this  depth, the bacterium, designated GSB1, lives off the dim glow of the thermal vent since no sunlight can penetrate to that depth. Green sulfur bacteria has also been found living on coral reef colonies in Taiwan, they make up the majority of a "green layer" on these colonies. They likely play a role in the coral system, and there could be a symbiotic relationship between the bacteria and the coral host. The coral could provide an anaerobic environment and  a source of carbon for the bacteria. The bacteria can provide nutrients and detoxify the coral by oxidizing sulfide. One type  of green sulfur bacteria, Chlorobium tepidum, has been found in sulfur springs. These organisms are thermophilic, unlike most other green sulfur bacteria. Phylogeny. The currently accepted phylogeny is based on 16S rRNA-based LTP release 123 by The All-Species Living Tree Project. Specific characteristics of genera. Green sulfur bacteria are family Chlorobiaceae. There are four genera; "Chloroherpeton", "Prosthecochloris", "Chlorobium" and "Chlorobaculum". Characteristics used to distinguish between these genera include some metabolic properties, pigments, cell morphology and absorption spectra. However, it is difficult to distinguish these properties and therefore the taxonomic division is sometimes unclear. Generally, "Chlorobium" are rod or  vibroid shaped and some species contain gas vesicles. They can develop as single or aggregate cells. They can be green or dark brown. The green strains use photosynthetic pigments Bchl "c" or "d" with chlorobactene carotenoids and the brown strains use photosynthetic pigment  Bchl "e" with isorenieratene carotenoids. Low amounts of salt are required for growth. "Prosthecochlori"s are made up of vibroid, ovid or rod shaped cells. They start as single cells that form appendages that do not branch, referred to as non-branching prosthecae. They can also form gas vesicles. The photosynthetic pigments present include Bchl "c, d" or  e. Furthermore, salt is necessary for growth. "Chlorobaculum" develop as single cells and are generally vibroid or rod-shaped. Some of these can form gas vesicles. The photosynthetic pigments in this genus are Bchl "c, d" or "e". Some species require NaCl (sodium chloride) for growth. Members of this genus used to be a part of the genus Chlorobium, but have formed a separate lineage. The genus "Chloroherpeton" is unique because members of this genus are motile. They are flexing long rods, and can move by gliding. They are green in color and contain the photosynthetic pigment Bchl "c" as  well as "γ-carotene". Salt is required for growth. Metabolism. Photosynthesis. The green sulfur bacteria use a Type I reaction center for photosynthesis. Type I reaction centers are the bacterial homologue of photosystem I (PSI) in plants and cyanobacteria. The GSB reaction centers contain bacteriochlorophyll a and are known as P840 reaction centers due to the excitation wavelength of 840 nm that powers the flow of electrons. In green sulfur bacteria the reaction center is associated with a large antena complex called the chlorosome that captures and funnels light energy to the reaction center. The chlorosomes have a peak absorption  in the far red region of the spectrum between 720-750 nm because they contain bacteriochlorophyll c, d and e. A protein complex called the Fenna-Matthews-Olson complex (FMO) is physically located between the chlorosomes and the P840 RC. The FMO complex helps efficiently transfer the energy absorbed by the antena to the reaction center. PSI and Type I reaction centers are able to reduce ferredoxin (Fd), a strong reductant that can be used to fix and reduce NADPH. Once the reaction center (RC) has given an electron to Fd it becomes an oxidizing agent (P840) with a reduction  potential of around +300 mV. While this is not positive enough to strip electrons from water to synthesize ( = +820 mV), it can accept electrons from other sources like , thiosulphate or ions. This transport of electrons from donors like to the acceptor Fd is called linear electron flow or linear electron transport. The oxidation of sulfide ions leads to the production of sulfur as a waste product that accumulates as globules on the extracellular side of the membrane. These globules of sulfur give green sulfur bacteria their name. When sulfide is depleted, the sulfur globules are consumed and further oxidized  to sulfate. However, the pathway of sulfur oxidation is not well-understood. Instead of passing the electrons onto Fd, the Fe-S clusters in the P840 reaction center can transfer the electrons to menaquinone (MQ:) which returns the electrons to the P840 via an electron transport chain (ETC). On the way back to the RC the electrons from MQH2 pass through a cytochrome bc complex (similar to the complex III of mitochondria) that pumps ions across the membrane. The electrochemical potential of the protons across the membrane is used to synthesize ATP by the FF ATP synthase. This cyclic electron  transport is responsible for converting light energy into cellular energy in the form of ATP. Sulfur metabolism. Green sulfur bacteria oxidize inorganic sulfur compounds to use as electron donors for anaerobic photosynthesis, specifically in carbon dioxide fixation. They usually prefer to utilize sulfide over other sulfur compounds as an electron donor, however they can utilize thiosulfate or H. The intermediate is usually sulfur, which is deposited outside of the cell, and the end product is sulfate. The sulfur, which is deposited extracellularly, is in the form of sulfur globules, which can be later oxidized completely. The mechanisms of sulfur oxidation  in green sulfur bacteria are not well characterized. Some enzymes thought to be involved in sulfide oxidation include flavocytochrome c, sulfide:quinone oxidoreductase and the Sox system. Flavocytochrome can catalyze the transfer of electrons to cytochromes from sulfide, and these cytochromes could then move the electrons to the photosynthetic reaction center. However, not all green sulfur bacteria produce this enzyme, demonstrating that it is not needed for the oxidation of sulfide. Sulfide:quinone oxidoreductase (SQR) also helps with electron transport, but, when alone, has been found to produce decreased rates of sulfide oxidation in green sulfur bacteria, suggesting that there  is a different, more effective mechanism. However, most green sulfur bacteria contain a homolog of the SQR gene. The oxidation of thiosulfate to sulfate could be catalyzed by the enzymes in the Sox system. It is thought that the enzymes and genes related to sulfur metabolism were obtained via horizontal gene transfer during the evolution of green sulfur bacteria. Carbon fixation. Green sulfur bacteria are photoautotrophs: they not only get energy from light, they can grow using carbon dioxide as their sole source of carbon. They fix carbon dioxide using the reverse tricarboxylic acid cycle (rTCA) cycle where energy is  consumed to reduce carbon dioxide, rather than oxidize as seen in the forward TCA cycle, in order to synthesize pyruvate and acetate. These molecules are used as the raw materials to synthesize all the building blocks a cell needs to generate macromolecules. The rTCA cycle is highly energy efficient enabling the bacteria to grow under low light conditions. However it has several oxygen sensitive enzymes that limits its efficiency in aerobic conditions. The reactions of reversal of the oxidative tricarboxylic acid cycle are catalyzed by four enzymes: However, the oxidative TCA cycle (OTCA) still is present in green sulfur bacteria . The OTCA can assimilate acetate, however the OTCA appears to be incomplete in green sulfur bacteria due to the location and down regulation of the gene during phototrophic growth. Mixotrophy. Green sulfur bacteria are obligate photoautotrophs: they cannot grow in the absence of light even if they are provided with organic matter. However they exhibit a form of mixotrophy where they can consume simple organic compounds in the presence of light and CO. Nitrogen fixation. The majority of green sulfur bacteria are diazotrophs: they can reduce nitrogen to ammonia which is then used to synthesize amino acids. Green sulfur