Sulfur metabolism

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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. [1] The element is present in proteins, sulfate esters of polysaccharides, steroids, phenols, and sulfur-containing coenzymes. [2]

Contents

Oxidation

Reduced sulfur compounds are oxidized by most organisms, including higher animals and higher plants. [2] Some organisms can conserve energy (i.e., produce ATP) from the oxidation of sulfur and it can serve as the sole energy source for some lithotrophic bacteria and archaea. [3] Sulfur oxidizers use enzymes such as Sulfide:quinone reductase, sulfur dioxygenase and sulfite oxidase to oxidize sulfur compounds to sulfate.

Sulfur-oxidizing microorganisms

Reduced sulfur compounds, such as hydrogen sulfide, elemental sulfur, sulfite, thiosulfate, and various polythionates (e.g., tetrathionate), are oxidized by chemotrophic, phototrophic, and mixotrophic bacteria for energy. [1] Some chemosynthetic archaea use hydrogen sulfide as an energy source for carbon fixation, producing sugars.

Chemotrophic sulfur-oxidizing bacteria

In order to have sufficient redox potential, microorganisms that use sulfur as an electron donor often use oxygen or nitrate as terminal electron acceptors. [4] Members of the chemotrophic Acidithiobacillus genus are able to oxidize a vast range of reduced sulfur compounds, but are restricted to acidic environments. [5] Chemotrophs that can produce sugars through chemosynthesis make up the base of some food chains. Food chains have formed in the absence of sunlight around hydrothermal vents, which emit hydrogen sulfide and carbon dioxide.

Phototrophic sulfur-oxidizing bacteria

Microbial sulfur cycle Microbial sulfur cycle.png
Microbial sulfur cycle

Some bacteria use light energy to couple sulfur oxidation to carbon dioxide (CO2) fixation for growth. These fall into two general groups: green sulfur bacteria (GSB) and purple sulfur bacteria (PSB). [6] However, some Cyanobacteria are also able to use hydrogen sulfide as an electron donor during anoxygenic photosynthesis. [7] All PSB are part of the class Gammaproteobacteria and are found in two families: Chromatiaceae and Ectothiorhodospiraceae. Typically, sulfur globules accumulate intracellularly in Chromatiaceae and extracellularly in Ectothiorhodospiraceae, which is one distinguishing feature between these two groups of PSB. [8] GSB are found within the family Chlorobiaceae generally oxidize sulfide or elemental sulfur, but some members are able to utilize thiosulfate. [9]

Reduction

Sulfur reduction occurs in plants, fungi, and many bacteria. [10] Sulfate can serve as an electron acceptor in anaerobic respiration and can also be reduced for the formation of organic compounds. Sulfate-reducing bacteria reduce sulfate and other oxidized sulfur compounds, such as sulfite, thiosulfate, and elemental sulfur, to sulfide.

Dissimilatory sulfur reduction

Some microorganisms are capable of reducing sulfate and elemental sulfur for energy by coupling sulfur reduction with the oxidation of molecular hydrogen or organic compounds such as acetate in anaerobic respiration. [11] These processes typically produce hydrogen sulfide as a byproduct, which can go on to serve as an electron donor in sulfur oxidation. [11] Sulfate reduction by sulfate-reducing bacteria is dissimilatory; the purpose of reducing the sulfate is to produce energy, and the sulfide is excreted. Dissimilatory sulfate reduction use the enzymes ATP sulfurylase, APS reductase, and sulfite reductase. [12]

Assimilatory sulfur reduction

In assimilatory sulfate reduction the sulfate is assimilated, or incorporated into organic compounds such as cysteine, methionine, or iron-sulfur clusters and enzyme cofactors. [13] In bacteria, sulfate and thiosulfate are transported into the cell by sulfate permeases where it can then be reduced and incorporated into biomolecules. [14] In some organisms (e.g., gut flora, cyanobacteria, and yeast), [15] assimilatory sulfate reduction is a more complex process that makes use of the enzymes ATP sulfurylase, APS kinase, PAPS reductase, and sulfite reductase. [10]

Disproportionation

Sulfur can also serve as both an electron donor and electron acceptor by microorganisms is disproportionation reactions. For example, Acidianus ambivalens uses sulfur oxygenase reductase (SOR) to convert elemental sulfur to sulfate, thiosulfate, and hydrogen sulfide through disproportionation. [16] Elemental sulfur disproportionation is restricted to environments where the concentration of the sulfide products are kept low, which typically happens in the presence of scavenging minerals that contain iron or manganese. [17] Disproportionation of thiosulfate often occurs in anoxic layers of marine and freshwater sediments. [18] [19]

Use by plants and animals

Plants take up sulfate in their roots and reduce it to sulfide (see Sulfur assimilation). However, some Brassica species are able to assimilate atmospheric sources of sulfur in the absence of other sources. [20] Plants reduce APS directly to sulfite (using APS reductase) without phosphorylating APS to PAPS. From the sulfide they form the amino acids cysteine and methionine, sulfolipids, and other sulfur compounds. Animals obtain sulfur from cysteine and methionine in the protein that they consume.

Sulfur is the third most abundant mineral element in the body. [21] The amino acids cysteine and methionine are used by the body to make glutathione. Excess cysteine and methionine are oxidized to sulfate by sulfite oxidase, eliminated in the urine, or stored as glutathione (which can serve as a store for sulfur). [21] The lack of sulfite oxidase, known as sulfite oxidase deficiency, causes physical deformities, mental retardation, and death.

See also

Related Research Articles

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Sulfur (also spelled sulphur in British English) is a chemical element; it has symbol S and atomic number 16. It is abundant, multivalent and nonmetallic. Under normal conditions, sulfur atoms form cyclic octatomic molecules with the chemical formula S8. Elemental sulfur is a bright yellow, crystalline solid at room temperature.

Anaerobic respiration is respiration using electron acceptors other than molecular oxygen (O2). Although oxygen is not the final electron acceptor, the process still uses a respiratory electron transport chain.

<span class="mw-page-title-main">Sulfate-reducing microorganism</span> Microorganisms that "breathe" sulfates

Sulfate-reducing microorganisms (SRM) or sulfate-reducing prokaryotes (SRP) are a group composed of sulfate-reducing bacteria (SRB) and sulfate-reducing archaea (SRA), both of which can perform anaerobic respiration utilizing sulfate (SO2−
4) as terminal electron acceptor, reducing it to hydrogen sulfide (H2S). Therefore, these sulfidogenic microorganisms "breathe" sulfate rather than molecular oxygen (O2), which is the terminal electron acceptor reduced to water (H2O) in aerobic respiration.

In chemistry, disproportionation, sometimes called dismutation, is a redox reaction in which one compound of intermediate oxidation state converts to two compounds, one of higher and one of lower oxidation states. The reverse of disproportionation, such as when a compound in an intermediate oxidation state is formed from precursors of lower and higher oxidation states, is called comproportionation, also known as synproportionation.

Sulfur-reducing bacteria are microorganisms able to reduce elemental sulfur (S0) to hydrogen sulfide (H2S). These microbes use inorganic sulfur compounds as electron acceptors to sustain several activities such as respiration, conserving energy and growth, in absence of oxygen. The final product of these processes, sulfide, has a considerable influence on the chemistry of the environment and, in addition, is used as electron donor for a large variety of microbial metabolisms. Several types of bacteria and many non-methanogenic archaea can reduce sulfur. Microbial sulfur reduction was already shown in early studies, which highlighted the first proof of S0 reduction in a vibrioid bacterium from mud, with sulfur as electron acceptor and H
2 as electron donor. The first pure cultured species of sulfur-reducing bacteria, Desulfuromonas acetoxidans, was discovered in 1976 and described by Pfennig Norbert and Biebel Hanno as an anaerobic sulfur-reducing and acetate-oxidizing bacterium, not able to reduce sulfate. Only few taxa are true sulfur-reducing bacteria, using sulfur reduction as the only or main catabolic reaction. Normally, they couple this reaction with the oxidation of acetate, succinate or other organic compounds. In general, sulfate-reducing bacteria are able to use both sulfate and elemental sulfur as electron acceptors. Thanks to its abundancy and thermodynamic stability, sulfate is the most studied electron acceptor for anaerobic respiration that involves sulfur compounds. Elemental sulfur, however, is very abundant and important, especially in deep-sea hydrothermal vents, hot springs and other extreme environments, making its isolation more difficult. Some bacteria – such as Proteus, Campylobacter, Pseudomonas and Salmonella – have the ability to reduce sulfur, but can also use oxygen and other terminal electron acceptors.

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.

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<span class="mw-page-title-main">Sulfur assimilation</span> Incorporation of sulfur into living organisms

Sulfur assimilation is the process by which living organisms incorporate sulfur into their biological molecules. In plants, sulfate is absorbed by the roots and then be transported to the chloroplasts by the transipration stream where the sulfur are reduced to sulfide with the help of a series of enzymatic reactions. Furthermore, the reduced sulfur is incorporated into cysteine, an amino acid that is a precursor to many other sulfur-containing compounds. In animals, sulfur assimilation occurs primarily through the diet, as animals cannot produce sulfur-containing compounds directly. Sulfur is incorporated into amino acids such as cysteine and methionine, which are used to build proteins and other important molecules. Besides, With the rapid development of economy, the increase emission of sulfur results in environmental issues, such as acid rain and hydrogen sulfilde.

<span class="mw-page-title-main">3'-Phosphoadenosine-5'-phosphosulfate</span> Chemical compound

3′-Phosphoadenosine-5′-phosphosulfate (PAPS) is a derivative of adenosine monophosphate (AMP) that is phosphorylated at the 3′ position and has a sulfate group attached to the 5′ phosphate. It is the most common coenzyme in sulfotransferase reactions and hence part of sulfation pathways. It is endogenously synthesized by organisms via the phosphorylation of adenosine 5′-phosphosulfate (APS), an intermediary metabolite. In humans such reaction is performed by bifunctional 3′-phosphoadenosine 5′-phosphosulfate synthases using ATP as the phosphate donor.

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<span class="mw-page-title-main">Adenylyl-sulfate reductase</span> Class of enzymes

Adenylyl-sulfate reductase is an enzyme that catalyzes the chemical reaction of the reduction of adenylyl-sulfate/adenosine-5'-phosphosulfate (APS) to sulfite through the use of an electron donor cofactor. The products of the reaction are AMP and sulfite, as well as an oxidized electron donor cofactor.

<span class="mw-page-title-main">Sulfite reductase</span> Enzyme family

Sulfite reductases (EC 1.8.99.1) are enzymes that participate in sulfur metabolism. They catalyze the reduction of sulfite to hydrogen sulfide and water. Electrons for the reaction are provided by a dissociable molecule of either NADPH, bound flavins, or ferredoxins.

<span class="mw-page-title-main">Thiosulfate dehydrogenase</span>

Thiosulfate dehydrogenase is an enzyme that catalyzes the chemical reaction:

Sulfurimonas is a bacterial genus within the class of Campylobacterota, known for reducing nitrate, oxidizing both sulfur and hydrogen, and containing Group IV hydrogenases. This genus consists of four species: Sulfurimonas autorophica, Sulfurimonas denitrificans, Sulfurimonas gotlandica, and Sulfurimonas paralvinellae. The genus' name is derived from "sulfur" in Latin and "monas" from Greek, together meaning a “sulfur-oxidizing rod”. The size of the bacteria varies between about 1.5-2.5 μm in length and 0.5-1.0 μm in width. Members of the genus Sulfurimonas are found in a variety of different environments which include deep sea-vents, marine sediments, and terrestrial habitats. Their ability to survive in extreme conditions is attributed to multiple copies of one enzyme. Phylogenetic analysis suggests that members of the genus Sulfurimonas have limited dispersal ability and its speciation was affected by geographical isolation rather than hydrothermal composition. Deep ocean currents affect the dispersal of Sulfurimonas spp., influencing its speciation. As shown in the MLSA report of deep-sea hydrothermal vents Campylobacterota, Sulfurimonas has a higher dispersal capability compared with deep sea hydrothermal vent thermophiles, indicating allopatric speciation.

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<span class="mw-page-title-main">Dissimilatory sulfate reduction</span>

Dissimilatory sulfate reduction is a form of anaerobic respiration that uses sulfate as the terminal electron acceptor to produce hydrogen sulfide. This metabolism is found in some types of bacteria and archaea which are often termed sulfate-reducing organisms. The term "dissimilatory" is used when hydrogen sulfide is produced in an anaerobic respiration process. By contrast, the term "assimilatory" would be used in relation to the biosynthesis of organosulfur compounds, even though hydrogen sulfide may be an intermediate.

<span class="mw-page-title-main">Microbial oxidation of sulfur</span>

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).

<span class="mw-page-title-main">Hydrothermal vent microbial communities</span> Undersea unicellular organisms

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.

Dissimilatory sulfite reductase is an enzyme that participates in sulfur metabolism in dissimilatory sulfate reduction.

Sulfur isotope biogeochemistry is the study of the distribution of sulfur isotopes in biological and geological materials. In addition to its common isotope, 32S, sulfur has three rare stable isotopes: 34S, 36S, and 33S. The distribution of these isotopes in the environment is controlled by many biochemical and physical processes, including biological metabolisms, mineral formation processes, and atmospheric chemistry. Measuring the abundance of sulfur stable isotopes in natural materials, like bacterial cultures, minerals, or seawater, can reveal information about these processes both in the modern environment and over Earth history.

References

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