Nitrospira Defluvii
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''Nitrospira'' (from Latin: nitro, meaning "nitrate" and Greek: spira, meaning "spiral") is a genus of bacteria within the monophyletic clade of the
Nitrospirota The Nitrospirota are a phylum of bacteria. They include multiple genera such as '' Nitrospira'', the largest. History of knowledge The first member of this phylum, '' Nitrospira marina'', was discovered in 1985. The second member, '' Nitrospi ...
phylum. The first member of this genus was described 1986 by Watson et al., isolated from the
Gulf of Maine The Gulf of Maine is a large gulf of the Atlantic Ocean on the east coast of North America. It is bounded by Cape Cod at the eastern tip of Massachusetts in the southwest and by Cape Sable Island at the southern tip of Nova Scotia in the northea ...
. The bacterium was named '' Nitrospira marina''. Populations were initially thought to be limited to
marine ecosystem Marine ecosystems are the largest of Earth's aquatic ecosystems and exist in Saline water, waters that have a high salt content. These systems contrast with freshwater ecosystems, which have a lower salt content. Marine waters cover more than 7 ...
s, but it was later discovered to be well-suited for numerous habitats, including
activated sludge The activated sludge process is a type of biological wastewater treatment process for treating sewage or Industrial wastewater treatment, industrial wastewaters using aeration and a biological floc (biofilm), floc composed of bacteria and protozoa ...
of
wastewater treatment Wastewater treatment is a process which removes and eliminates contaminants from wastewater. It thus converts it into an effluent that can be returned to the water cycle. Once back in the water cycle, the effluent creates an acceptable impact on ...
systems, natural biological marine settings (such as the
Seine River The Seine ( , ) is a river in northern France. Its drainage basin is in the Paris Basin (a geological relative lowland) covering most of northern France. It rises at Source-Seine, northwest of Dijon in northeastern France in the Langres p ...
in France and beaches in
Cape Cod Cape Cod is a peninsula extending into the Atlantic Ocean from the southeastern corner of Massachusetts, in the northeastern United States. Its historic, maritime character and ample beaches attract heavy tourism during the summer months. The ...
in the United States), water circulation
biofilter Biofiltration is a pollution control technique using a bioreactor containing living material to capture and biologically degrade pollutants. Common uses include processing waste water, capturing harmful chemicals or silt from surface runoff, a ...
s in
aquarium An aquarium (: aquariums or aquaria) is a vivarium of any size having at least one transparent side in which aquatic plants or animals are kept and displayed. fishkeeping, Fishkeepers use aquaria to keep fish, invertebrates, amphibians, aquati ...
tanks, terrestrial systems, fresh and salt water ecosystems, agricultural lands and
hot spring A hot spring, hydrothermal spring, or geothermal spring is a Spring (hydrology), spring produced by the emergence of Geothermal activity, geothermally heated groundwater onto the surface of the Earth. The groundwater is heated either by shallow ...
s. ''Nitrospira'' is a ubiquitous bacterium that plays a role in the nitrogen cycle by performing nitrite oxidation in the second step of nitrification. ''Nitrospira'' live in a wide array of environments including but not limited to, drinking water systems, waste treatment plants, rice paddies, forest soils, geothermal springs, and sponge tissue. Despite being abundant in many natural and engineered ecosystems ''Nitrospira'' are difficult to culture, so most knowledge of them is from molecular and genomic data. However, due to their difficulty to be cultivated in laboratory settings, the entire genome was only sequenced in one species, '' Nitrospira defluvii''. In addition, ''Nitrospira'' bacteria's
16S rRNA 16S ribosomal RNA (or 16Svedberg, 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 genes coding for it are referred to as ...
sequences are too dissimilar to use for PCR primers, thus some members go unnoticed. In addition, members of ''Nitrospira'' with the capabilities to perform complete
nitrification ''Nitrification'' is the biological oxidation of ammonia to nitrate via the intermediary nitrite. Nitrification is an important step in the nitrogen cycle in soil. The process of complete nitrification may occur through separate organisms or ent ...
(
comammox Comammox (COMplete AMMonia OXidation) is the name attributed to an organism that can convert ammonia into nitrite and then into nitrate through the process of nitrification. Nitrification has traditionally been thought to be a two-step process, wher ...
bacteria) has also been discovered and cultivated.


Morphology

For the following description, ''
Nitrospira moscoviensis ''Nitrospira moscoviensis'' was the second bacterium classified under the most diverse nitrite-oxidizing bacteria phylum, ''Nitrospirae''. It is a gram-negative, non-motile, facultative lithoauthotropic bacterium that was discovered in Moscow, R ...
'' will be representative of the ''Nitrospira'' genus. ''Nitrospira'' is a
Gram-negative Gram-negative bacteria are bacteria that, unlike gram-positive bacteria, do not retain the crystal violet stain used in the Gram staining method of bacterial differentiation. Their defining characteristic is that their cell envelope consists ...
nitrite The nitrite polyatomic ion, ion has the chemical formula . Nitrite (mostly sodium nitrite) is widely used throughout chemical and pharmaceutical industries. The nitrite anion is a pervasive intermediate in the nitrogen cycle in nature. The name ...
-oxidizing organism with a helical to vibroid morphology (0.9–2.2 × 0.2–0.4
micrometre The micrometre (English in the Commonwealth of Nations, Commonwealth English as used by the International Bureau of Weights and Measures; SI symbol: μm) or micrometer (American English), also commonly known by the non-SI term micron, is a uni ...
s in size). They are non-
planktonic Plankton are the diverse collection of organisms that drift in water (or air) but are unable to actively propel themselves against currents (or wind). The individual organisms constituting plankton are called plankters. In the ocean, they pro ...
organisms that reside as clumps, known as aggregates, in
biofilm A biofilm is a Syntrophy, syntrophic Microbial consortium, community of microorganisms in which cell (biology), cells cell adhesion, stick to each other and often also to a surface. These adherent cells become embedded within a slimy ext ...
s. Visualization using
transmission electron microscopy Transmission electron microscopy (TEM) is a microscopy technique in which a beam of electrons is transmitted through a specimen to form an image. The specimen is most often an ultrathin section less than 100 nm thick or a suspension on a g ...
(TEM) confirms star-like protrusions on the outer membrane (6–8 nm thick). The
periplasmic space The periplasm is a concentrated gel-like matrix in the space between the inner cytoplasmic membrane and the bacterial outer membrane called the ''periplasmic space'' in Gram-negative (more accurately "diderm") bacteria. Using cryo-electron microsco ...
is exceptionally wide (34–41 nm thick), which provides space to accommodate electron-rich molecules. Electron-deprived structures are located in the
cytosol The cytosol, also known as cytoplasmic matrix or groundplasm, is one of the liquids found inside cells ( intracellular fluid (ICF)). It is separated into compartments by membranes. For example, the mitochondrial matrix separates the mitochondri ...
and are believed to be
glycogen Glycogen is a multibranched polysaccharide of glucose that serves as a form of energy storage in animals, fungi, and bacteria. It is the main storage form of glucose in the human body. Glycogen functions as one of three regularly used forms ...
storage vesicles;
polyhydroxybutyrate Polyhydroxybutyrate (PHB) is a Polyhydroxyalkanoates, polyhydroxyalkanoate (PHA), a polymer belonging to the polyesters class that are of interest as bio-derived and biodegradable plastics. The poly-3-hydroxybutyrate (P3HB) form of PHB is probabl ...
and
polyphosphate A polyphosphate is a Salt (chemistry), salt or ester of polymeric oxyanions formed from tetrahedral PO4 (phosphate) structural units linked together by sharing oxygen atoms. Polyphosphates can adopt linear or a cyclic (also called, ring) structure ...
granules are also identified in the cytoplasm. DNA analysis determined 56.9 +/- 0.4 mol% of the DNA to be
guanine Guanine () (symbol G or Gua) is one of the four main nucleotide bases found in the nucleic acids DNA and RNA, the others being adenine, cytosine, and thymine ( uracil in RNA). In DNA, guanine is paired with cytosine. The guanine nucleoside ...
and
cytosine Cytosine () (symbol C or Cyt) is one of the four nucleotide bases found in DNA and RNA, along with adenine, guanine, and thymine ( uracil in RNA). It is a pyrimidine derivative, with a heterocyclic aromatic ring and two substituents attac ...
base pairs.


General metabolism

''Nitrospira'' are capable of aerobic hydrogen oxidation and nitrite oxidation to obtain electrons, but high concentrations of nitrite have shown to inhibit their growth. The optimal temperature for nitrite oxidation and growth in ''
Nitrospira moscoviensis ''Nitrospira moscoviensis'' was the second bacterium classified under the most diverse nitrite-oxidizing bacteria phylum, ''Nitrospirae''. It is a gram-negative, non-motile, facultative lithoauthotropic bacterium that was discovered in Moscow, R ...
'' is 39 °C (can range from 33–44 °C) at a pH range of 7.6–8.0 Despite being commonly classified as obligate
chemolithotrophs 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 aerob ...
, some are capable of
mixotroph A mixotroph is an organism that uses a mix of different sources of energy and carbon, instead of having a single trophic mode, on the continuum from complete autotrophy to complete heterotrophy. It is estimated that mixotrophs comprise more than ...
y. For instance, under different environments, ''Nitrospira'' can choose to assimilate carbon by
carbon fixation Biological carbon fixation, or сarbon assimilation, is the Biological process, process by which living organisms convert Total inorganic carbon, inorganic carbon (particularly carbon dioxide, ) to Organic compound, organic compounds. These o ...
or by consuming organic molecules (
glycerol Glycerol () is a simple triol compound. It is a colorless, odorless, sweet-tasting, viscous liquid. The glycerol backbone is found in lipids known as glycerides. It is also widely used as a sweetener in the food industry and as a humectant in pha ...
,
pyruvate Pyruvic acid (CH3COCOOH) is the simplest of the alpha-keto acids, with a carboxylic acid and a ketone functional group. Pyruvate, the conjugate base, CH3COCOO−, is an intermediate in several metabolic pathways throughout the cell. Pyruvic ...
, or
formate Formate (IUPAC name: methanoate) is the conjugate base of formic acid. Formate is an anion () or its derivatives such as ester of formic acid. The salts and esters are generally colorless. Fundamentals When dissolved in water, formic acid co ...
). New studies also show that ''Nitrospira'' can use
urea Urea, also called carbamide (because it is a diamide of carbonic acid), is an organic compound with chemical formula . This amide has two Amine, amino groups (–) joined by a carbonyl functional group (–C(=O)–). It is thus the simplest am ...
as a source of nutrients.
Urease Ureases (), functionally, belong to the superfamily of amidohydrolases and phosphotriesterases. Ureases are found in numerous Bacteria, Archaea, fungi, algae, plants, and some invertebrates. Ureases are nickel-containing metalloenzymes of high ...
encoded within their genome can break urea down to and
ammonia Ammonia is an inorganic chemical compound of nitrogen and hydrogen with the chemical formula, formula . A Binary compounds of hydrogen, stable binary hydride and the simplest pnictogen hydride, ammonia is a colourless gas with a distinctive pu ...
. The can be assimilated by
anabolism Anabolism () is the set of metabolic pathways that construct macromolecules like DNA or RNA from smaller units. These reactions require energy, known also as an Endergonic reaction, endergonic process. Anabolism is the building-up aspect of metabo ...
while the ammonia and organic by-product released by ''Nitrospira'' allow ammonium oxidizers and other microbes to co-exist in the same microenvironment.


Nitrification

All members of this genus have the nitrite oxidoreductase genes, and thus are all thought to be nitrite-oxidizers. Ever since
nitrifying bacteria Nitrifying bacteria are chemolithotrophic organisms that include species of genera such as '' Nitrosomonas'', '' Nitrosococcus'', '' Nitrobacter'', '' Nitrospina'', '' Nitrospira'' and '' Nitrococcus''. These bacteria get their energy from the oxi ...
were discovered it was accepted that nitrification occurred in two steps, although it would be energetically favourable for one organism to do both steps. Recently ''Nitrospira'' members with the abilities to perform complete
nitrification ''Nitrification'' is the biological oxidation of ammonia to nitrate via the intermediary nitrite. Nitrification is an important step in the nitrogen cycle in soil. The process of complete nitrification may occur through separate organisms or ent ...
(
comammox Comammox (COMplete AMMonia OXidation) is the name attributed to an organism that can convert ammonia into nitrite and then into nitrate through the process of nitrification. Nitrification has traditionally been thought to be a two-step process, wher ...
bacteria) have also been discovered and cultivated as in the case of '' Nitrospira inopinata''. The discovery of commamox organisms within ''Nitrospira'' redefine the way bacteria contribute to the
Nitrogen cycle The nitrogen cycle is the biogeochemical cycle by which nitrogen is converted into multiple chemical forms as it circulates among atmosphere, atmospheric, terrestrial ecosystem, terrestrial, and marine ecosystems. The conversion of nitrogen can ...
and thus a lot of future studies will be dedicated to it. With these new findings there's now a possibility to mainly use complete nitrification instead of partial nitrification in engineered systems like
wastewater treatment plants Wastewater treatment is a process which removes and eliminates contaminants from wastewater. It thus converts it into an effluent that can be returned to the water cycle. Once back in the water cycle, the effluent creates an acceptable impact on ...
because complete nitrification results in lower emissions of the
greenhouse gas Greenhouse gases (GHGs) are the gases in the atmosphere that raise the surface temperature of planets such as the Earth. Unlike other gases, greenhouse gases absorb the radiations that a planet emits, resulting in the greenhouse effect. T ...
es:
nitrous oxide Nitrous oxide (dinitrogen oxide or dinitrogen monoxide), commonly known as laughing gas, nitrous, or factitious air, among others, is a chemical compound, an Nitrogen oxide, oxide of nitrogen with the Chemical formula, formula . At room te ...
and
nitric oxide Nitric oxide (nitrogen oxide, nitrogen monooxide, or nitrogen monoxide) is a colorless gas with the formula . It is one of the principal oxides of nitrogen. Nitric oxide is a free radical: it has an unpaired electron, which is sometimes den ...
, into the atmosphere.


Genome

After sequencing and analyzing the DNA of ''Nitrospira'' members, researchers discovered both species had genes encoding ammonia monooxygenase (Amo) and hydroxlyamine dehydrogenase (hao), enzymes that ammonia-oxidizing bacteria (AOB) use to convert
ammonia Ammonia is an inorganic chemical compound of nitrogen and hydrogen with the chemical formula, formula . A Binary compounds of hydrogen, stable binary hydride and the simplest pnictogen hydride, ammonia is a colourless gas with a distinctive pu ...
into nitrite. The bacteria possess all necessary sub-units for both enzymes as well as the necessary cell membrane associated proteins and transporters to carry out the first step of nitrification. Origins of the Amo gene are debatable as one study found that it is similar to other AOB while another study found the Amo gene to be genetically distinct from other lineages. Current findings indicate that the hao gene is phylogenetically distinct from the hao gene present in other AOB, meaning that they acquired them long ago, likely by
horizontal gene transfer Horizontal gene transfer (HGT) or lateral gene transfer (LGT) is the movement of genetic material between organisms other than by the ("vertical") transmission of DNA from parent to offspring (reproduction). HGT is an important factor in the e ...
. ''Nitrospira'' also carry the genes encoding for all the sub-units of nitrite oxidoreductase (nxr), the enzyme that catalyzes the second step of nitrification.


Phylogeny

The currently accepted taxonomy is based on the
List of Prokaryotic names with Standing in Nomenclature List of Prokaryotic names with Standing in Nomenclature (LPSN) is an online database that maintains information on the naming and taxonomy of prokaryotes, following the taxonomy requirements and rulings of the International Code of Nomenclatu ...
(LSPN) and the
National Center for Biotechnology Information The National Center for Biotechnology Information (NCBI) is part of the National Library of Medicine (NLM), a branch of the National Institutes of Health (NIH). It is approved and funded by the government of the United States. The NCBI is lo ...
(NCBI). Phylogeny is based on GTDB 09-RS220 by
Genome Taxonomy Database The Genome Taxonomy Database (GTDB) is an online database that maintains information on a proposed nomenclature of prokaryotes, following a phylogenomic approach based on a set of conserved single-copy proteins. In addition to resolving parap ...
Species incertae sedis: * "''Ca.'' N. alkalitolerans" Daebeler et al. 2020 * "''Ca.'' N. bockiana" Lebedeva et al. 2008 * "''N. calida''" Lebedeva et al. 2011 * ''N. marina'' Watson et al. 1986 * "''Ca.'' N. salsa" Haaijer et al. 2013


See also

*
Nitrogen cycle The nitrogen cycle is the biogeochemical cycle by which nitrogen is converted into multiple chemical forms as it circulates among atmosphere, atmospheric, terrestrial ecosystem, terrestrial, and marine ecosystems. The conversion of nitrogen can ...
*
List of bacterial orders This article lists the orders of the Bacteria. The currently accepted taxonomy is based on the List of Prokaryotic names with Standing in Nomenclature (LPSN) and National Center for Biotechnology Information (NCBI) and the phylogeny is based on 16 ...
*
List of bacteria genera This article lists the genera of the bacteria Bacteria (; : bacterium) are ubiquitous, mostly free-living organisms often consisting of one Cell (biology), biological cell. They constitute a large domain (biology), domain of Prokaryote, ...


References


External links


MicrobeWiki -- Nitrospira
{{Taxonbar, from=Q3024630 Nitrospirota Bacteria genera