Sulfur metabolism
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Sulfur Sulfur (or sulphur in British English) is a chemical element with the symbol S and atomic number 16. It is abundant, multivalent and nonmetallic. Under normal conditions, sulfur atoms form cyclic octatomic molecules with a chemical formula ...
is
metabolized Metabolism (, from el, μεταβολή ''metabolē'', "change") is the set of life-sustaining chemical reactions in organisms. The three main functions of metabolism are: the conversion of the energy in food to energy available to run cell ...
by all organisms, from
bacteria Bacteria (; singular: bacterium) are ubiquitous, mostly free-living organisms often consisting of one biological cell. They constitute a large domain of prokaryotic microorganisms. Typically a few micrometres in length, bacteria were among ...
and
archaea Archaea ( ; singular archaeon ) is a domain of single-celled organisms. These microorganisms lack cell nuclei and are therefore prokaryotes. Archaea were initially classified as bacteria, receiving the name archaebacteria (in the Archaebac ...
to
plant Plants are predominantly photosynthetic eukaryotes of the kingdom Plantae. Historically, the plant kingdom encompassed all living things that were not animals, and included algae and fungi; however, all current definitions of Plantae exclud ...
s and
animal Animals are multicellular, eukaryotic organisms in the Kingdom (biology), biological kingdom Animalia. With few exceptions, animals Heterotroph, consume organic material, Cellular respiration#Aerobic respiration, breathe oxygen, are Motilit ...
s. Sulfur is reduced or
oxidized Redox (reduction–oxidation, , ) is a type of chemical reaction in which the oxidation states of substrate change. Oxidation is the loss of electrons or an increase in the oxidation state, while reduction is the gain of electrons or a ...
by organisms in a variety of forms. The element is present in
protein Proteins are large biomolecules and macromolecules that comprise one or more long chains of amino acid residues. Proteins perform a vast array of functions within organisms, including catalysing metabolic reactions, DNA replication, respo ...
s, sulfate esters of
polysaccharide Polysaccharides (), or polycarbohydrates, are the most abundant carbohydrates found in food. They are long chain polymeric carbohydrates composed of monosaccharide units bound together by glycosidic linkages. This carbohydrate can react with wa ...
s,
steroid A steroid is a biologically active organic compound with four rings arranged in a specific molecular configuration. Steroids have two principal biological functions: as important components of cell membranes that alter membrane fluidity; and a ...
s,
phenol Phenol (also called carbolic acid) is an aromatic organic compound with the molecular formula . It is a white crystalline solid that is volatile. The molecule consists of a phenyl group () bonded to a hydroxy group (). Mildly acidic, it req ...
s, and sulfur-containing
coenzyme A cofactor is a non-protein chemical compound or metallic ion that is required for an enzyme's role as a catalyst (a catalyst is a substance that increases the rate of a chemical reaction). Cofactors can be considered "helper molecules" that ass ...
s.


Oxidation

Reduced sulfur compounds are oxidized by most organisms, including higher animals and higher plants. Some organisms can conserve energy (i.e., produce ATP) from the oxidation of sulfur. Sulfur is the sole energy source for some
lithotroph Lithotrophs are a diverse group of organisms using an inorganic substrate (usually of mineral origin) to obtain reducing equivalents for use in biosynthesis (e.g., carbon dioxide fixation) or energy conservation (i.e., ATP production) via aerobi ...
ic bacteria and archaea. Reduced sulfur compounds, such as
hydrogen sulfide Hydrogen sulfide is a chemical compound with the formula . It is a colorless chalcogen-hydride gas, and is poisonous, corrosive, and flammable, with trace amounts in ambient atmosphere having a characteristic foul odor of rotten eggs. The unde ...
, elemental sulfur,
sulfite Sulfites or sulphites are compounds that contain the sulfite ion (or the sulfate(IV) ion, from its correct systematic name), . The sulfite ion is the conjugate base of bisulfite. Although its acid ( sulfurous acid) is elusive, its salts are wide ...
,
thiosulfate Thiosulfate ( IUPAC-recommended spelling; sometimes thiosulphate in British English) is an oxyanion of sulfur with the chemical formula . Thiosulfate also refers to the compounds containing this anion, which are the salts of thiosulfuric acid, ...
, and various polythionates (e.g.,
tetrathionate The tetrathionate anion, , is a sulfur oxoanion derived from the compound tetrathionic acid, H2S4O6. Two of the sulfur atoms present in the ion are in oxidation state 0 and two are in oxidation state +5. Alternatively, the compound can be viewed ...
), are used by various lithotrophic bacteria and are all oxidized by ''
Acidithiobacillus ''Acidithiobacillus'' is a genus of the ''Acidithiobacillia'' in the " Pseudomonadota". The genus includes acidophilic organisms capable of iron and/or sulfur oxidation. Like all ''"Pseudomonadota"'', ''Acidithiobacillus'' spp. are Gram-negative ...
''. Sulfur oxidizers use enzymes such as Sulfide:quinone reductase, sulfur dioxygenase and
sulfite oxidase Sulfite oxidase () is an enzyme in the mitochondria of all eukaryotes, with exception of the yeasts. It oxidizes sulfite to sulfate and, via cytochrome c, transfers the electrons produced to the electron transport chain, allowing generation o ...
to oxidize sulfur compounds to
sulfate The sulfate or sulphate ion is a polyatomic anion with the empirical formula . Salts, acid derivatives, and peroxides of sulfate are widely used in industry. Sulfates occur widely in everyday life. Sulfates are salts of sulfuric acid and many ar ...
. Lithotrophs that can produce sugars through
chemosynthesis In biochemistry, chemosynthesis is the biological conversion of one or more carbon-containing molecules (usually carbon dioxide or methane) and nutrients into organic matter using the oxidation of inorganic compounds (e.g., hydrogen gas, hydro ...
make up the base of some
food chain A food chain is a linear network of links in a food web starting from producer organisms (such as grass or algae which produce their own food via photosynthesis) and ending at an apex predator species (like grizzly bears or killer whales), det ...
s. Food chains have formed in the absence of sunlight around
hydrothermal vent A hydrothermal vent is a fissure on the seabed from which geothermally heated water discharges. They are commonly found near volcanically active places, areas where tectonic plates are moving apart at mid-ocean ridges, ocean basins, and hotspot ...
s, which emit hydrogen sulfide and
carbon dioxide Carbon dioxide (chemical formula ) is a chemical compound made up of molecules that each have one carbon atom covalently double bonded to two oxygen atoms. It is found in the gas state at room temperature. In the air, carbon dioxide is transpar ...
. Chemosynthetic archaea use hydrogen sulfide as an energy source for
carbon fixation Biological carbon fixation or сarbon assimilation is the process by which inorganic carbon (particularly in the form of carbon dioxide) is converted to organic compounds by living organisms. The compounds are then used to store energy and as ...
, producing sugars.


Reduction

Sulfur reduction occurs in plants, fungi, and many bacteria.Pathway: sulfate reduction I (assimilatory)
" ''MetaCyc''.
Sulfate can serve as an
electron acceptor An electron acceptor is a chemical entity that accepts electrons transferred to it from another compound. It is an oxidizing agent that, by virtue of its accepting electrons, is itself reduced in the process. Electron acceptors are sometimes mista ...
in
anaerobic respiration 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. In aerobic organisms undergoing re ...
and can also be reduced for the formation of
organic compound In chemistry, organic compounds are generally any chemical compounds that contain carbon-hydrogen or carbon-carbon bonds. Due to carbon's ability to catenate (form chains with other carbon atoms), millions of organic compounds are known. The ...
s.
Sulfate-reducing bacteria 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 () as termina ...
reduce sulfate and other oxidized sulfur compounds, such as sulfite, thiosulfate, and elemental sulfur, to
sulfide Sulfide (British English also sulphide) is an inorganic anion of sulfur with the chemical formula S2− or a compound containing one or more S2− ions. Solutions of sulfide salts are corrosive. ''Sulfide'' also refers to chemical compounds lar ...
. Sulfate reduction can be dissimilatory or assimilatory. Sulfate reduction by sulfate-reducing bacteria, for example, is dissimilatory; the purpose of reducing the sulfate is to produce energy, and the sulfide is excreted.
Dissimilatory sulfate reduction 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-reduc ...
use the enzymes
ATP sulfurylase In enzymology, a sulfate adenylyltransferase () is an enzyme that catalyzes the chemical reaction :ATP + sulfate \rightleftharpoonspyrophosphate + adenylyl sulfate Thus, the two substrates of this enzyme are ATP and sulfate, whereas its two p ...
, APS reductase, and
sulfite reductase Sulfite reductases () 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 fer ...
. In assimilatory sulfate reduction, however, the sulfate is assimilated, or incorporated into organic compounds. In some organisms (e.g.,
gut flora Gut microbiota, gut microbiome, or gut flora, are the microorganisms, including bacteria, archaea, fungi, and viruses that live in the digestive tracts of animals. The gastrointestinal metagenome is the aggregate of all the genomes of the gut mi ...
,
cyanobacteria Cyanobacteria (), also known as Cyanophyta, are a phylum of gram-negative bacteria that obtain energy via photosynthesis. The name ''cyanobacteria'' refers to their color (), which similarly forms the basis of cyanobacteria's common name, blu ...
, and
yeast Yeasts are eukaryotic, single-celled microorganisms classified as members of the fungus kingdom. The first yeast originated hundreds of millions of years ago, and at least 1,500 species are currently recognized. They are estimated to constitut ...
), Pathway: sulfate reduction II (assimilatory)
" ''MetaCyc''.
assimilatory sulfate reduction is a more complex process that makes use of the enzymes ATP sulfurylase,
APS kinase APS or Aps or aps or similar may refer to: Education * Abbottabad Public School * Adarsh Public School, a public school in New Delhi, India * Alamogordo Public Schools * Albuquerque Public Schools, New Mexico, US school district * Allendale Pu ...
, PAPS reductase, and sulfite reductase.


Use by plants and animals

Plants take up sulfate in their roots and reduce it to sulfide (see
sulfur assimilation Sulfur assimilation is the process by which organisms obtain sulfur, an essential chemical element, element for growth and metabolism of most organisms. Biologically, sulfur is often encountered in its most redox, oxidized form as sulfate or the m ...
). Plants are able to reduce APS directly to sulfite (using APS reductase) without phosphorylating APS to PAPS. From the sulfide they form the amino acids
cysteine Cysteine (symbol Cys or C; ) is a semiessential proteinogenic amino acid with the formula . The thiol side chain in cysteine often participates in enzymatic reactions as a nucleophile. When present as a deprotonated catalytic residue, sometime ...
and
methionine Methionine (symbol Met or M) () is an essential amino acid in humans. As the precursor of other amino acids such as cysteine and taurine, versatile compounds such as SAM-e, and the important antioxidant glutathione, methionine plays a critical ro ...
, 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. The amino acids cysteine and methionine are used by the body to make
glutathione Glutathione (GSH, ) is an antioxidant in plants, animals, fungi, and some bacteria and archaea. Glutathione is capable of preventing damage to important cellular components caused by sources such as reactive oxygen species, free radicals, pero ...
. 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). The lack of sulfite oxidase, known as sulfite oxidase deficiency, causes physical deformities, mental retardation, and death.


See also

*
Microbial metabolism Microbial metabolism is the means by which a microbe obtains the energy and nutrients (e.g. carbon) it needs to live and reproduce. Microbes use many different types of metabolic strategies and species can often be differentiated from each other ...
* Sulfur cycle


References


External links

*{{Commons category-inline