Green sulfur bacteria | |
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Green sulfur bacteria in a Winogradsky column | |
Scientific classification | |
Domain: | Bacteria |
(unranked): | Bacteroidota–Chlorobiota group |
Phylum: | Chlorobiota Iino et al. 2021 [1] |
Class: | "Chlorobia" Garrity and Holt 2001 [2] |
Order: | Chlorobiales Gibbons and Murray 1978 (Approved Lists 1980) [3] |
Families and Genera | |
Synonyms | |
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The green sulfur bacteria are a phylum, Chlorobiota, [4] of obligately anaerobic photoautotrophic bacteria that metabolize sulfur. [5]
Green sulfur bacteria are nonmotile (except Chloroherpeton thalassium, which may glide) and capable of anoxygenic photosynthesis. [5] [6] They live in anaerobic aquatic environments. [7] In contrast to plants, green sulfur bacteria mainly use sulfide ions as electron donors. [8] They are autotrophs that utilize the reverse tricarboxylic acid cycle to perform carbon fixation. [9] They are also mixotrophs and reduce nitrogen. [10] [11]
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. [12] Electron donors include H2, H2S, 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. [7] Chlorosomes are a unique feature that allow them to capture light in low-light conditions. [13]
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. [7]
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. [14]
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. [15]
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. [16] 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. [17]
One type of green sulfur bacteria, Chlorobaculum tepidum , has been found in sulfur springs. These organisms are thermophilic, unlike most other green sulfur bacteria. [7]
16S rRNA based LTP_08_2023 [18] [19] [20] | 120 marker proteins based GTDB 08-RS214 [21] [22] [23] | ||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||
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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. [24]
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. [24]
Prosthecochloris 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. [24]
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. [24]
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. [24]
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 and 750 nm because they contain bacteriochlorophyll c, d and e. [25] 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 CO
2 and reduce NAD+. 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 O
2 (E
0 = +820 mV), it can accept electrons from other sources like H
2S, thiosulphate or Fe2+
ions. [26] This transport of electrons from donors like H
2S 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. [8]
Instead of passing the electrons onto Fd, the Fe-S clusters in the P840 reaction center can transfer the electrons to menaquinone (MQ:MQH
2) 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 bc1 complex (similar to the complex III of mitochondria) that pumps H+
ions across the membrane. The electrochemical potential of the protons across the membrane is used to synthesize ATP by the FoF1 ATP synthase. This cyclic electron transport is responsible for converting light energy into cellular energy in the form of ATP. [25]
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 H2. [27] The intermediate is usually sulfur, which is deposited outside of the cell, [28] 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. [27]
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 SO
x 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. [27] However, most green sulfur bacteria contain a homolog of the SQR gene. [29] The oxidation of thiosulfate to sulfate could be catalyzed by the enzymes in the SO
x system. [27]
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. [29]
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 [9] where energy is consumed to reduce carbon dioxide, rather than oxidize as seen in the forward TCA cycle, [9] 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. [30] However it has several oxygen sensitive enzymes that limits its efficiency in aerobic conditions. [30]
The reactions of reversal of the oxidative tricarboxylic acid cycle are catalyzed by four enzymes: [9]
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. [9]
Green sulfur bacteria are often referred to as obligate photoautotrophs as they cannot grow in the absence of light even if they are provided with organic matter. [9] [26] However they exhibit a form of mixotrophy where they can consume simple organic compounds in the presence of light and CO2. [9] In the presence of CO2 or HCO3−, some green sulfur bacteria can utilize acetate or pyruvate. [9]
Mixotrophy in green sulfur bacteria is best modeled by the representative green sulfur bacterium Chlorobaculum tepidum. [31] Mixotrophy occurs during amino acid biosynthesis/carbon utilization and energy metabolism. [32] The bacterium uses electrons, generated from the oxidation of sulfur, and the energy it captures from light to run the rTCA. C. tepidum also exhibits use of both pyruvate and acetate as an organic carbon source. [32]
An example of mixotrophy in C. tepidum that combines autotrophy and heterotrophy is in its synthesis of acetyl-CoA. C. tepidum can autotrophically generate acetyl-CoA through the rTCA cycle, or it can heterotrophically generate it from the uptake of acetate. Similar mixotrophic activity occurs when pyruvate is used for amino acid biosynthesis, but mixotrophic growth using acetate yields higher growth rates. [31] [32]
In energy metabolism, C. tepidum relies on light reactions to produce energy (NADPH and NADH) because the pathways typically responsible for energy production (oxidative pentose phosphate pathway and normal TCA cycle) are only partly functional. [32] Photons absorbed from the light are used to produce NADPH and NADH, the cofactors of energy metabolism. C. tepidum also generates energy in the form of ATP using the proton motive force derived from sulfide oxidation. [31] Energy production from both sulfide oxidation and photon absorption via bacteriochlorophylls. [32]
The majority of green sulfur bacteria are diazotrophs: they can reduce nitrogen to ammonia which is then used to synthesize amino acids. [33] Nitrogen fixation among green sulfur bacteria is generally typical of an anoxygenic phototroph, and requires the presence of light. Green sulfur bacteria exhibit activity from a Type-1 secretion system and a ferredoxin-NADP+ oxidoreductase to generate reduced iron, a trait that evolved to support nitrogen fixation. [34] Like purple sulfur bacteria, they can regulate the activity of nitrogenase post-translationally in response to ammonia concentrations. Their possession of nif genes, even though evolutionarily distinct, may suggest their nitrogen fixation abilities arose in two different events or through a shared very distant ancestor. [33]
Examples of green sulfur bacteria capable of nitrogen fixation include the genus Chlorobium and Pelodictyon, excluding P. phaeoclathratiforme. Prosthecochloris aestuarii and Chloroherpeton thalassium also fall into this category. [33] Their N2 fixation is widespread and plays an important role in overall nitrogen availability for ecosystems. Green sulfur bacteria living in coral reefs, such as Prosthecochloris, are crucial in generating available nitrogen in the already nutrient-limited environment. [16]
Photosynthesis is a system of biological processes by which photosynthetic organisms, such as most plants, algae, and cyanobacteria, convert light energy, typically from sunlight, into the chemical energy necessary to fuel their metabolism. Photosynthesis usually refers to oxygenic photosynthesis, a process that produces oxygen. Photosynthetic organisms store the chemical energy so produced within intracellular organic compounds like sugars, glycogen, cellulose and starches. To use this stored chemical energy, an organism's cells metabolize the organic compounds through cellular respiration. Photosynthesis plays a critical role in producing and maintaining the oxygen content of the Earth's atmosphere, and it supplies most of the biological energy necessary for complex life on Earth.
Chloroflexus aurantiacus is a photosynthetic bacterium isolated from hot springs, belonging to the green non-sulfur bacteria. This organism is thermophilic and can grow at temperatures from 35 to 70 °C. Chloroflexus aurantiacus can survive in the dark if oxygen is available. When grown in the dark, Chloroflexus aurantiacus has a dark orange color. When grown in sunlight it is dark green. The individual bacteria tend to form filamentous colonies enclosed in sheaths, which are known as trichomes.
Biological carbon fixation, or сarbon assimilation, is the process by which living organisms convert inorganic carbon to organic compounds. These organic compounds are then used to store energy and as structures for other biomolecules. Carbon is primarily fixed through photosynthesis, but some organisms use chemosynthesis in the absence of sunlight. Chemosynthesis is carbon fixation driven by chemical energy rather than from sunlight.
Chlorophyll a is a specific form of chlorophyll used in oxygenic photosynthesis. It absorbs most energy from wavelengths of violet-blue and orange-red light, and it is a poor absorber of green and near-green portions of the spectrum. Chlorophyll does not reflect light but chlorophyll-containing tissues appear green because green light is diffusively reflected by structures like cell walls. This photosynthetic pigment is essential for photosynthesis in eukaryotes, cyanobacteria and prochlorophytes because of its role as primary electron donor in the electron transport chain. Chlorophyll a also transfers resonance energy in the antenna complex, ending in the reaction center where specific chlorophylls P680 and P700 are located.
Purple bacteria or purple photosynthetic bacteria are Gram-negative proteobacteria that are phototrophic, capable of producing their own food via photosynthesis. They are pigmented with bacteriochlorophyll a or b, together with various carotenoids, which give them colours ranging between purple, red, brown, and orange. They may be divided into two groups – purple sulfur bacteria and purple non-sulfur bacteria. Purple bacteria are anoxygenic phototrophs widely spread in nature, but especially in aquatic environments, where there are anoxic conditions that favor the synthesis of their pigments.
The Chromatiaceae are one of the two families of purple sulfur bacteria, together with the Ectothiorhodospiraceae. They belong to the order Chromatiales of the class Gammaproteobacteria, which is composed by unicellular Gram-negative organisms. Most of the species are photolithoautotrophs and conduct an anoxygenic photosynthesis, but there are also representatives capable of growing under dark and/or microaerobic conditions as either chemolithoautotrophs or chemoorganoheterotrophs.
Photoheterotrophs are heterotrophic phototrophs—that is, they are organisms that use light for energy, but cannot use carbon dioxide as their sole carbon source. Consequently, they use organic compounds from the environment to satisfy their carbon requirements; these compounds include carbohydrates, fatty acids, and alcohols. Examples of photoheterotrophic organisms include purple non-sulfur bacteria, green non-sulfur bacteria, and heliobacteria. These microorganisms are ubiquitous in aquatic habitats, occupy unique niche-spaces, and contribute to global biogeochemical cycling. Recent research has also indicated that the oriental hornet and some aphids may be able to use light to supplement their energy supply.
Lithotrophs are a diverse group of organisms using an inorganic substrate to obtain reducing equivalents for use in biosynthesis or energy conservation via aerobic or anaerobic respiration. While lithotrophs in the broader sense include photolithotrophs like plants, chemolithotrophs are exclusively microorganisms; no known macrofauna possesses the ability to use inorganic compounds as electron sources. Macrofauna and lithotrophs can form symbiotic relationships, in which case the lithotrophs are called "prokaryotic symbionts". An example of this is chemolithotrophic bacteria in giant tube worms or plastids, which are organelles within plant cells that may have evolved from photolithotrophic cyanobacteria-like organisms. Chemolithotrophs belong to the domains Bacteria and Archaea. The term "lithotroph" was created from the Greek terms 'lithos' (rock) and 'troph' (consumer), meaning "eaters of rock". Many but not all lithoautotrophs are extremophiles.
A chlorosome is a photosynthetic antenna complex found in green sulfur bacteria (GSB) and many green non-sulfur bacteria (GNsB), together known as green bacteria. They differ from other antenna complexes by their large size and lack of protein matrix supporting the photosynthetic pigments. Green sulfur bacteria are a group of organisms that generally live in extremely low-light environments, such as at depths of 100 metres in the Black Sea. The ability to capture light energy and rapidly deliver it to where it needs to go is essential to these bacteria, some of which see only a few photons of light per chlorophyll per day. To achieve this, the bacteria contain chlorosome structures, which contain up to 250,000 chlorophyll molecules. Chlorosomes are ellipsoidal bodies, in GSB their length varies from 100 to 200 nm, width of 50-100 nm and height of 15 – 30 nm, in GNsB the chlorosomes are somewhat smaller.
Microbial metabolism is the means by which a microbe obtains the energy and nutrients it needs to live and reproduce. Microbes use many different types of metabolic strategies and species can often be differentiated from each other based on metabolic characteristics. The specific metabolic properties of a microbe are the major factors in determining that microbe's ecological niche, and often allow for that microbe to be useful in industrial processes or responsible for biogeochemical cycles.
The reverse Krebs cycle is a sequence of chemical reactions that are used by some bacteria and archaea to produce carbon compounds from carbon dioxide and water by the use of energy-rich reducing agents as electron donors.
The Fenna–Matthews–Olson (FMO) complex is a water-soluble complex and was the first pigment-protein complex (PPC) to be structure analyzed by x-ray spectroscopy. It appears in green sulfur bacteria and mediates the excitation energy transfer from light-harvesting chlorosomes to the membrane-embedded bacterial reaction center (bRC). Its structure is trimeric (C3-symmetry). Each of the three monomers contains eight bacteriochlorophyll a molecules. They are bound to the protein scaffold via chelation of their central magnesium atom either to amino acids of the protein or water-bridged oxygen atoms.
Sulfur is metabolized by all organisms, from bacteria and archaea to plants and animals. Sulfur can have an oxidation state from -2 to +6 and is reduced or oxidized by a diverse range of organisms. The element is present in proteins, sulfate esters of polysaccharides, steroids, phenols, and sulfur-containing coenzymes.
Anoxygenic photosynthesis is a special form of photosynthesis used by some bacteria and archaea, which differs from the better known oxygenic photosynthesis in plants in the reductant used and the byproduct generated.
Chlorobaculum tepidum, previously known as Chlorobium tepidum, is an anaerobic, thermophilic green sulfur bacteria first isolated from New Zealand. Its cells are gram-negative and non-motile rods of variable length. They contain chlorosomes and bacteriochlorophyll a and c.
Chlorobium chlorochromatii, originally known as Chlorobium aggregatum, is a symbiotic green sulfur bacteria that performs anoxygenic photosynthesis and functions as an obligate photoautotroph using reduced sulfur species as electron donors. Chlorobium chlorochromatii can be found in stratified freshwater lakes.
Microbial oxidation of sulfur is the oxidation of sulfur by microorganisms to build their structural components. The oxidation of inorganic compounds is the strategy primarily used by chemolithotrophic microorganisms to obtain energy to survive, grow and reproduce. Some inorganic forms of reduced sulfur, mainly sulfide (H2S/HS−) and elemental sulfur (S0), can be oxidized by chemolithotrophic sulfur-oxidizing prokaryotes, usually coupled to the reduction of oxygen (O2) or nitrate (NO3−). Anaerobic sulfur oxidizers include photolithoautotrophs that obtain their energy from sunlight, hydrogen from sulfide, and carbon from carbon dioxide (CO2).
The hydrothermal vent microbial community includes all unicellular organisms that live and reproduce in a chemically distinct area around hydrothermal vents. These include organisms in the microbial mat, free floating cells, or bacteria in an endosymbiotic relationship with animals. Chemolithoautotrophic bacteria derive nutrients and energy from the geological activity at Hydrothermal vents to fix carbon into organic forms. Viruses are also a part of the hydrothermal vent microbial community and their influence on the microbial ecology in these ecosystems is a burgeoning field of research.
Chlorobium limicola is a gram negative bacterial member of green sulfur bacteria genus found in freshwater hot springs. C. limicola is a non-motile mesophile, photoautotrophic / photosynthetic strict anaerobe important to carbon, nitrogen and sulfur cycles in anoxic freshwater environments. Strain DSMZ 245 T was isolated from Gilroy Hot Springs in California, and whole genome sequencing was accomplished. Believed to be morphologically diverse, it was determined that culturing techniques determine some characteristics like slime production and morphology. As a green sulfur bacteria, C. limicola fixes carbon via reverse TCA cycle reactions.
Prosthecochloris aestuarii is a green sulfur bacterium in the genus Prosthecochloris. This organism was originally isolated from brackish lagoons located in Sasyk-Sivash and Sivash. They are characterized by the presence of "prosthecae" on their cell surface; the inner part of these appendages house the photosynthetic machinery within chlorosomes, which are characteristic structures of green sulfur bacteria. Additionally, like other green sulfur bacteria, they are Gram-negative, non-motile, and non-spore forming. Of the four major groups of green sulfur bacteria, P. aestuarii serves as the type species for Group 4.