Sulfolobus Metallicus
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''Sulfolobus metallicus'' is a coccoid shaped
thermophilic A thermophile is an organism—a type of extremophile—that thrives at relatively high temperatures, between . Many thermophiles are archaea, though they can be bacteria or fungi. Thermophilic eubacteria are suggested to have been among the earl ...
archaeon 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 Archaebact ...
. It is a strict
chemolithoautotroph A lithoautotroph is an organism which derives energy from reactions of reduced compounds of mineral (inorganic) origin. Two types of lithoautotrophs are distinguished by their energy source; photolithoautotrophs derive their energy from light while ...
gaining energy by oxidation of
sulphur 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 ...
and sulphidic ores into sulfuric acid. Its type strain is Kra 23 (DSM 6482). It has many uses that take advantage of its ability to grow on metal media under acidic and hot environments.


Taxonomy

The name "metallicus" is
Latin Latin (, or , ) is a classical language belonging to the Italic branch of the Indo-European languages. Latin was originally a dialect spoken in the lower Tiber area (then known as Latium) around present-day Rome, but through the power of the ...
meaning "the miner". The use of
biochemistry Biochemistry or biological chemistry is the study of chemical processes within and relating to living organisms. A sub-discipline of both chemistry and biology, biochemistry may be divided into three fields: structural biology, enzymology and ...
and
phylogeny A phylogenetic tree (also phylogeny or evolutionary tree Felsenstein J. (2004). ''Inferring Phylogenies'' Sinauer Associates: Sunderland, MA.) is a branching diagram or a tree showing the evolutionary relationships among various biological spec ...
were originally the main source of
classification Classification is a process related to categorization, the process in which ideas and objects are recognized, differentiated and understood. Classification is the grouping of related facts into classes. It may also refer to: Business, organizat ...
, but
16S rRNA 16S rRNA may refer to: * 16S ribosomal RNA 16 S ribosomal RNA (or 16 S rRNA) is the RNA component of the 30S subunit of a prokaryotic ribosome ( SSU rRNA). It binds to the Shine-Dalgarno sequence and provides most of the SSU structure. The g ...
sequences were performed on it to elucidate the phylogeny. ''Sulfolobus sp.'' strain 7r sp. nov. is a close relative to ''Sulfolobus metallicus'' with comparable phylogenetic properties. Both ''Sulfolobus metallicus'' and ''Sulfolobus sp.'' strain 7r sp. nov. are grouped primarily based on their thermoacidophilic nature within the
Thermoproteota The Thermoproteota (also known as crenarchaea) are archaea that have been classified as a phylum of the Archaea domain. Initially, the Thermoproteota were thought to be sulfur-dependent extremophiles but recent studies have identified characteris ...
archaeal phylum.


History

''Sulfolobus metallicus'' was first isolated in 1991 by doctors Gertrud Huber and Karl O. Stetter from solfataric fields in
Iceland Iceland ( is, Ísland; ) is a Nordic island country in the North Atlantic Ocean and in the Arctic Ocean. Iceland is the most sparsely populated country in Europe. Iceland's capital and largest city is Reykjavík, which (along with its s ...
. The collected samples were grown on ore supplemented media under acidic conditions at the optimal temperature of 65°C. The archaea were routinely grown at the optimal temperature of 65°C on ore media. After ''Sulfolobus metallicus'' was shown to be able to oxidize sulfur compounds, researchers looked into its potential to
oxidize 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 d ...
reduced sulfur emissions.


Metabolism

''Sulfolobus metallicus'' was grown on media that contains reduced sulfur, and the production of sulfuric acid was tested. ''Sulfolobus metallicus'' was found to be a strict
aerobic Aerobic means "requiring air," in which "air" usually means oxygen. Aerobic may also refer to * Aerobic exercise, prolonged exercise of moderate intensity * Aerobics, a form of aerobic exercise * Aerobic respiration, the aerobic process of cellu ...
chemolithotroph 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 ...
Bandeiras, Tiago M., et al. "The Respiratory Chain Of The Thermophilic Archaeon ''Sulfolobus metallicus'': Studies On The Type-II NADH Dehydrogenase." ''BBA – Bioenergetics'' 1557.1-3 (2003): 13. because it
oxidize 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 d ...
s
sulphur 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 ...
contained in minerals such as
pyrite The mineral pyrite (), or iron pyrite, also known as fool's gold, is an iron sulfide with the chemical formula Iron, FeSulfur, S2 (iron (II) disulfide). Pyrite is the most abundant sulfide mineral. Pyrite's metallic Luster (mineralogy), lust ...
,
chalcopyrite Chalcopyrite ( ) is a copper iron sulfide mineral and the most abundant copper ore mineral. It has the chemical formula CuFeS2 and crystallizes in the tetragonal system. It has a brassy to golden yellow color and a hardness of 3.5 to 4 on the Mo ...
, elemental sulfur and
sphalerite Sphalerite (sometimes spelled sphaelerite) is a sulfide mineral with the chemical formula . It is the most important ore of zinc. Sphalerite is found in a variety of deposit types, but it is primarily in Sedimentary exhalative deposits, sedimen ...
into
sulfuric acid Sulfuric acid (American spelling and the preferred IUPAC name) or sulphuric acid ( Commonwealth spelling), known in antiquity as oil of vitriol, is a mineral acid composed of the elements sulfur, oxygen and hydrogen, with the molecular formu ...
. ''Sulfolobus metallicus'' can also oxidize
iron(II) In chemistry, iron(II) refers to the element iron in its +2 oxidation state. In ionic compounds (salts), such an atom may occur as a separate cation (positive ion) denoted by Fe2+. The adjective ferrous or the prefix ferro- is often used to spe ...
. ''Sulfolobus metallicus'' has a unique type II
NADH dehydrogenase NADH dehydrogenase is an enzyme that converts nicotinamide adenine dinucleotide (NAD) from its reduced form (NADH) to its oxidized form (NAD+). Members of the NADH dehydrogenase family and analogues are commonly systematically named using the for ...
with no
iron-sulfur clusters Iron–sulfur proteins (or iron–sulphur proteins in British spelling) are proteins characterized by the presence of iron–sulfur clusters containing sulfide-linked di-, tri-, and tetrairon centers in variable oxidation states. Iron–sulfur clu ...
that is covalently linked to a flavin molecule.


Genetics

''Sulfolobus metallicus''s
GC-content In molecular biology and genetics, GC-content (or guanine-cytosine content) is the percentage of nitrogenous bases in a DNA or RNA molecule that are either guanine (G) or cytosine (C). This measure indicates the proportion of G and C bases out o ...
is around 38 mol%.


Genes of proteins

Though its whole
genome In the fields of molecular biology and genetics, a genome is all the genetic information of an organism. It consists of nucleotide sequences of DNA (or RNA in RNA viruses). The nuclear genome includes protein-coding genes and non-coding ge ...
has not been sequenced, the coding sequences of some genes have been sequenced:


Carboxylase genes

Genes were found in this organism that encode the
biotin carboxylase In enzymology, a biotin carboxylase () is an enzyme that Catalysis, catalyzes the chemical reaction : ATP + biotin-carboxyl-carrier protein + CO2 \rightleftharpoons ADP + phosphate + carboxybiotin-carboxyl-carrier protein The three Substrate ...
, carboxyl transferase and
biotin carboxyl carrier protein Biotin Carboxyl Carrier Protein (BCCP) refers to proteins containing a biotin attachment domain that carry biotin and carboxybiotin throughout the ATP-dependent carboxylation by biotin-dependent carboxylases. In the case of '' E. coli'' Acetyl-C ...
. These genes were discovered using
Sanger sequencing Sanger sequencing is a method of DNA sequencing that involves electrophoresis and is based on the random incorporation of chain-terminating dideoxynucleotides by DNA polymerase during in vitro DNA replication. After first being developed by Frederi ...
. The amino acid residue sequence of all three proteins that are coded for by these genes have also been sequenced using microsequencing. Biotin carboxylase, carboxyl transferase and biotin carboxyl carrier protein could likely be a complex of proteins that help fix carbon dioxide for
autotrophy An autotroph or primary producer is an organism that produces complex organic compounds (such as carbohydrates, fats, and proteins) using carbon from simple substances such as carbon dioxide,Morris, J. et al. (2019). "Biology: How Life Works", ...
.


''fox'' genes

''S. metallicus'' up-regulates genes that help it grow in environments with higher sulfur and iron concentrations. It has a
cluster may refer to: Science and technology Astronomy * Cluster (spacecraft), constellation of four European Space Agency spacecraft * Asteroid cluster, a small asteroid family * Cluster II (spacecraft), a European Space Agency mission to study t ...
of genes, called ''fox'' genes, that encode membrane proteins that are similar to
cytochrome c oxidase The enzyme cytochrome c oxidase or Complex IV, (was , now reclassified as a translocasEC 7.1.1.9 is a large transmembrane protein complex found in bacteria, archaea, and mitochondria of eukaryotes. It is the last enzyme in the respiratory electr ...
. The gene that codes for sulfur oxygenase-reductase was found to be expressed when the archaea was grown on sulfur media.
cDNA In genetics, complementary DNA (cDNA) is DNA synthesized from a single-stranded RNA (e.g., messenger RNA (mRNA) or microRNA (miRNA)) template in a reaction catalyzed by the enzyme reverse transcriptase. cDNA is often used to express a speci ...
of the ''fox'' genes that were known to be in other sulfur oxidizing cells were used to test the presence of the ''fox'' genes in ''Sulfolobus metallicus''.


Phylogeny

The
16S rRNA 16S rRNA may refer to: * 16S ribosomal RNA 16 S ribosomal RNA (or 16 S rRNA) is the RNA component of the 30S subunit of a prokaryotic ribosome ( SSU rRNA). It binds to the Shine-Dalgarno sequence and provides most of the SSU structure. The g ...
DNA sequences are not always consistent with the phylogenetic analysis of the Sofolobales order of bacteria. However, a 16S rRNA analysis was performed on the members of Sulfolobales to make a phylogenetic tree. The closest member is the ''Sulfolobus'' strain LM, sharing 98.2% of the genome that codes for the 16S rRNA. Of the ''Sulfolobales'', it is least related to ''Sulfolobus hakonensis'' with a 15.7% difference in the genes that code for the 16S rRNA. ''S. metallicus'' has an 87.7% similarity to ''S. hakonensis'', an 87.6% similarity to ''A. brierleyi'', an 87.4% similarity to ''M. sedula'', an 87.5% similarity to ''M. prunae'', 88.5% similarity to ''A. ambivalens'' and an 88.8% similarity to ''A. infernus''.


Physiology

''Sulfolobus metallicus'' is a coccoid shaped
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 ...
. Its
cell envelope The cell envelope comprises the inner cell membrane and the cell wall of a bacterium. In gram-negative bacteria an outer membrane is also included. This envelope is not present in the Mollicutes where the cell wall is absent. Bacterial cell env ...
contains an
S-layer An S-layer (surface layer) is a part of the cell envelope found in almost all archaea, as well as in many types of bacteria. The S-layers of both archaea and bacteria consists of a monomolecular layer composed of only one (or, in a few cases, two) i ...
, Isopranyl ether
lipids Lipids are a broad group of naturally-occurring molecules which includes fats, waxes, sterols, fat-soluble vitamins (such as vitamins A, D, E and K), monoglycerides, diglycerides, phospholipids, and others. The functions of lipids include ...
and caldariellaquinone. The lipids protect the archaea from the acidic environments in which they live while still maintaining stability at high temperatures. It has high levels of
polyphosphate Polyphosphates are salts or esters of polymeric oxyanions formed from tetrahedral PO4 (phosphate) structural units linked together by sharing oxygen atoms. Polyphosphates can adopt linear or a cyclic ring structures. In biology, the polyphosphate e ...
.


Uses


Bioleaching

It is a key organism in
bioleaching Bioleaching is the extraction of metals from their ores through the use of living organisms. This is much cleaner than the traditional heap leaching using cyanide. Bioleaching is one of several applications within biohydrometallurgy and several ...
of
copper Copper is a chemical element with the symbol Cu (from la, cuprum) and atomic number 29. It is a soft, malleable, and ductile metal with very high thermal and electrical conductivity. A freshly exposed surface of pure copper has a pinkis ...
,
cobalt Cobalt is a chemical element with the symbol Co and atomic number 27. As with nickel, cobalt is found in the Earth's crust only in a chemically combined form, save for small deposits found in alloys of natural meteoric iron. The free element, pr ...
,
nickel Nickel is a chemical element with symbol Ni and atomic number 28. It is a silvery-white lustrous metal with a slight golden tinge. Nickel is a hard and ductile transition metal. Pure nickel is chemically reactive but large pieces are slow to ...
and
gold Gold is a chemical element with the symbol Au (from la, aurum) and atomic number 79. This makes it one of the higher atomic number elements that occur naturally. It is a bright, slightly orange-yellow, dense, soft, malleable, and ductile met ...
. Bioleaching techniques can be used to separate metal sulfides into ionic metal, which can be collected, and hydrogen sulfide. The process requires both acidic protons and oxidized iron (Fe3+). ''Sulfolobus metallicus'', being an acidophile, can not only withstand the acidic conditions necessary for bioleaching, but also produces sulfuric acid that can used for the bioleaching process and maintains the necessary levels of oxidized iron through its metabolism. Bioleaching at temperatures conducive of thermophiles like ''Sulfolobus metallicus'' has shown to be more effective than bioleaching with
mesophiles A mesophile is an organism that grows best in moderate temperature, neither too hot nor too cold, with an optimum growth range from . The optimum growth temperature for these organisms is 37°C. The term is mainly applied to microorganisms. Organi ...
. Bioleaching is adventitious to traditional metal extraction methods because it is more cost efficient and poses fewer hazards to the environment.


Oxidation of sulfur

''Sulfolobus metallicus'' could potentially be utilized to eliminate reduced sulfur compounds, such as hydrogen sulphide (H2S), near
urban Urban means "related to a city". In that sense, the term may refer to: * Urban area, geographical area distinct from rural areas * Urban culture, the culture of towns and cities Urban may also refer to: General * Urban (name), a list of people ...
areas that cause disturbing odours. Many industries have H2S gas emissions that present many environmental problems as well as a foul odour. ''S. metalllicus'' has shown that it can
oxidize 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 d ...
these compounds and potentially eliminate many of those
emissions Emission may refer to: Chemical products * Emission of air pollutants, notably: **Flue gas, gas exiting to the atmosphere via a flue ** Exhaust gas, flue gas generated by fuel combustion ** Emission of greenhouse gases, which absorb and emit rad ...
. Many of these industrial emissions occur at high temperatures and low concentrations. ''Sulfolobus metallicus'' has an advantage over other microbes for the task of sulfur oxidation in the fact that it is a
thermophile A thermophile is an organism—a type of extremophile—that thrives at relatively high temperatures, between . Many thermophiles are archaea, though they can be bacteria or fungi. Thermophilic eubacteria are suggested to have been among the earl ...
, so it can be used to treat reduced sulfur at industrial temperatures that other sulfur oxidizers could not withstand.


Archaeal phospholipids

''Sulfolobus metallicus'' could be used for mass-producing archeal phospholipids. These
lipids Lipids are a broad group of naturally-occurring molecules which includes fats, waxes, sterols, fat-soluble vitamins (such as vitamins A, D, E and K), monoglycerides, diglycerides, phospholipids, and others. The functions of lipids include ...
have promising applications in
drug delivery Drug delivery refers to approaches, formulations, manufacturing techniques, storage systems, and technologies involved in transporting a pharmaceutical compound to its target site to achieve a desired therapeutic effect. Principles related to d ...
by acting as
liposomes A liposome is a small artificial Vesicle (biology and chemistry), vesicle, spherical in shape, having at least one lipid bilayer. Due to their hydrophobicity and/or hydrophilicity, biocompatibility, particle size and many other properties, lipo ...
, or they can be used as
lubricants A lubricant (sometimes shortened to lube) is a substance that helps to reduce friction between surfaces in mutual contact, which ultimately reduces the heat generated when the surfaces move. It may also have the function of transmitting forces, t ...
but can be expensive to synthesize. ''Sulfolobus metallicus'' can potentially be used to provide a cheaper way to synthesize these lipids. If ''Sulfolobus metallicus'' is used as a bioleacher on the industrial scale, it grows in volume in tons per day. Researchers can
centrifuge A centrifuge is a device that uses centrifugal force to separate various components of a fluid. This is achieved by spinning the fluid at high speed within a container, thereby separating fluids of different densities (e.g. cream from milk) or ...
the solution and separate the
lipid Lipids are a broad group of naturally-occurring molecules which includes fats, waxes, sterols, fat-soluble vitamins (such as vitamins A, D, E and K), monoglycerides, diglycerides, phospholipids, and others. The functions of lipids include ...
without interfering with the extraction of the metal.


Zinc binding domains

''Sulfolobus metallicus'' is capable of synthesizing two isoforms of
ferredoxin Ferredoxins (from Latin ''ferrum'': iron + redox, often abbreviated "fd") are iron–sulfur proteins that mediate electron transfer in a range of metabolic reactions. The term "ferredoxin" was coined by D.C. Wharton of the DuPont Co. and applied t ...
: FdA and FdB. The tertiary structure of ferredoxin is typically stabilized by electrostatic interactions with
zinc Zinc is a chemical element with the symbol Zn and atomic number 30. Zinc is a slightly brittle metal at room temperature and has a shiny-greyish appearance when oxidation is removed. It is the first element in group 12 (IIB) of the periodi ...
ions (Zn2+). In ''S. metallicus'', FdA binds zinc ion, but FdB does not. Therefore, ''S. metallicus'' serves as a good model for studying how zinc binding affects the stability of
proteins 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 ...
like ferredoxin.


Growth and tolerance

''Sulfolobus metallicus'' can grow between 50 °C and 75 °C (meaning it is a hyperthermophilic archaea) in acidic environments between pH 1.0 and 4.5. It can grow in 0-3% NaCl.


Polyphosphate

''Sulfolobus metallicus'' can tolerate up to 200 mM
copper sulfate Copper sulfate may refer to: * Copper(II) sulfate, CuSO4, a common compound used as a fungicide and herbicide * Copper(I) sulfate Copper(I) sulfate, also known as cuprous sulfate, is an inorganic compound with the chemical formula Cu2 SO4. It ...
. It was used as a
model organism A model organism (often shortened to model) is a non-human species that is extensively studied to understand particular biological phenomena, with the expectation that discoveries made in the model organism will provide insight into the workin ...
to study the mechanism of metal transport using
polyphosphate Polyphosphates are salts or esters of polymeric oxyanions formed from tetrahedral PO4 (phosphate) structural units linked together by sharing oxygen atoms. Polyphosphates can adopt linear or a cyclic ring structures. In biology, the polyphosphate e ...
because this archaea has a higher capacity to accumulate polyphosphate than other ''Sulfolobus'' archaea. It is proposed that the accumulation of high levels of
polyphosphate Polyphosphates are salts or esters of polymeric oxyanions formed from tetrahedral PO4 (phosphate) structural units linked together by sharing oxygen atoms. Polyphosphates can adopt linear or a cyclic ring structures. In biology, the polyphosphate e ...
s contribute to the
mechanism Mechanism may refer to: *Mechanism (engineering), rigid bodies connected by joints in order to accomplish a desired force and/or motion transmission *Mechanism (biology), explaining how a feature is created *Mechanism (philosophy), a theory that a ...
of the tolerance of these copper ions. Researchers put these archaea in an increased concentration of metal and found the exopolyphosphatase activity increased as polyphosphate levels decreased. This suggests that ''Sulfolobus'' archaea can tolerate metals through a polyphosphate mechanism.


Notes and references


Further reading

*


External links

*
LPSNType strain of ''Sulfolobus metallicus'' at Bac''Dive'' - the Bacterial Diversity Metadatabase
{{Taxonbar, from=Q12603503 Thermoproteota Archaea described in 1992