Brocadiaceae
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"''Candidatus'' Brocadia" is a ''
candidatus In prokaryote nomenclature, ''Candidatus'' (abbreviated ''Ca.''; Latin for "candidate of Roman office") is used to name prokaryotic taxa that are well characterized but yet- uncultured. Contemporary sequencing approaches, such as 16S ribosomal R ...
'' genus of
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, prokaryotic microorganisms. Typically a few micr ...
, meaning that while it is well-characterized, it has not been grown as a pure culture yet. Due to this, much of what is known about ''Candidatus'' species (including Brocadia) has been discovered using culture-independent techniques such as
metagenomic Metagenomics is the study of all genetic material from all organisms in a particular environment, providing insights into their composition, diversity, and functional potential. Metagenomics has allowed researchers to profile the microbial co ...
sequence analysis. Some notable species within this genus include the type species, ''Candidatus'' Brocadia anammoxidans, along with ''Candidatus'' Brocadia sinica and ''Candidatus'' Brocadia fulgida. Many of the species in this genus, including those already listed, are capable of anaerobic ammonium oxidation, also known as
anammox Anammox, an abbreviation for "anaerobic ammonium oxidation", is a globally important microbial process of the nitrogen cycle that takes place in many natural environments. The bacteria mediating this process were identified in 1999, and were a gr ...
, an important part of the global nitrogen cycle. Anammox works by converting fixed nitrogen back into N2 gas in the atmosphere. Anammox bacteria have a unique, membrane-bound organelle where this anammox process takes place—it is called the anammoxosome.


Phylogeny

Phylogeny 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 ...


Structure

''Candidatus'' Brocadia falls under the phylum of
Planctomycetota The Planctomycetota are a phylum of widely distributed bacteria, occurring in both aquatic and terrestrial habitats. They play a considerable role in global carbon and nitrogen cycles, with many species of this phylum capable of anaerobic ammoni ...
. These bacteria are known for a few distinguishing phenotypic features including cell surface pits (crateriform structures), no present
peptidoglycan Peptidoglycan or murein is a unique large macromolecule, a polysaccharide, consisting of sugars and amino acids that forms a mesh-like layer (sacculus) that surrounds the bacterial cytoplasmic membrane. The sugar component consists of alternating ...
in the cell wall, compartmentalization, and even budding reproduction rather than
binary fission Binary may refer to: Science and technology Mathematics * Binary number, a representation of numbers using only two values (0 and 1) for each digit * Binary function, a function that takes two arguments * Binary operation, a mathematical o ...
. Compartmentalization of ''Ca.'' Brocadia and anammox bacteria in general are of great interest. They are known to have three main compartments: (1) the paryphoplasm, (2) the riboplasm, and (3) the anammoxosome, as seen in Figure 1. The paryphoplasm is the outermost compartment of the cell, while the inner most compartment is the riboplasm, which holds the ribosomes and DNA of the cell. The anammoxosome is a membrane-bound component of the cell that does not contain DNA or ribosomes, and it is where the
anammox Anammox, an abbreviation for "anaerobic ammonium oxidation", is a globally important microbial process of the nitrogen cycle that takes place in many natural environments. The bacteria mediating this process were identified in 1999, and were a gr ...
process takes place. This is something that makes anammox bacteria unique, as prokaryotes typically do not have membrane-bound organelles. ''Ca.'' Brocadia, like other anammox bacteria, have integral ladderane lipids embedded in the membranes of their anammoxosome. This helps keep any toxic intermediates from the anammox process—such as hydrazine—from contaminating and damaging the rest of the cell.


Natural and Industrial Environments


Natural

These bacteria are found in many natural environments that the
anammox Anammox, an abbreviation for "anaerobic ammonium oxidation", is a globally important microbial process of the nitrogen cycle that takes place in many natural environments. The bacteria mediating this process were identified in 1999, and were a gr ...
process takes place in, and are a major part of the conversion of fixed nitrogen to gaseous nitrogen (N2) that is released into the atmosphere. Anammox bacteria like those falling within ''Ca.'' Brocadia were first found in the
Black Sea The Black Sea is a marginal sea, marginal Mediterranean sea (oceanography), mediterranean sea lying between Europe and Asia, east of the Balkans, south of the East European Plain, west of the Caucasus, and north of Anatolia. It is bound ...
, and subsequently the
Arabian Sea The Arabian Sea () is a region of sea in the northern Indian Ocean, bounded on the west by the Arabian Peninsula, Gulf of Aden and Guardafui Channel, on the northwest by Gulf of Oman and Iran, on the north by Pakistan, on the east by India, and ...
, both bodies of water being mostly
anoxic Anoxia means a total depletion in the level of oxygen, an extreme form of hypoxia or "low oxygen". The terms anoxia and hypoxia are used in various contexts: * Anoxic waters, sea water, fresh water or groundwater that are depleted of dissolved ox ...
and ideal conditions for the anammox process.


Industrial

The genus, ''Ca.'' Brocadia and others are used in industrial
waste water 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 ...
plants using
sequencing batch reactor Sequencing batch reactors (SBR) or sequential batch reactors are a type of activated sludge process for the wastewater treatment, treatment of wastewater. SBRs treat wastewater such as sewage or output from anaerobic digesters or mechanical biolog ...
s,
rotating biological contactor A rotating biological contactor or RBC is a biological fixed-film treatment process used in the secondary treatment of wastewater following primary treatment. The primary treatment process involves removal of grit, sand and coarse suspended mat ...
s,
membrane bioreactor Membrane bioreactors are combinations of membrane processes like microfiltration or ultrafiltration with a biological wastewater treatment process, the activated sludge process. These technologies are now widely used for municipal and industri ...
s, and many others. These reactors are ideal for industrial waste water treatment due to their high sludge retention.


Notable Species

Many studies have been performed regarding the physiological attributes of the different anammox bacteria, including their role in the
anammox Anammox, an abbreviation for "anaerobic ammonium oxidation", is a globally important microbial process of the nitrogen cycle that takes place in many natural environments. The bacteria mediating this process were identified in 1999, and were a gr ...
process, how they perform in the anammox process, and other key features that differentiate them from each other. Comparisons between important and notable species are listed here.


''Candidatus'' Brocadia anammoxidans

''Ca.'' B. anammoxidans is the
type species In International_Code_of_Zoological_Nomenclature, zoological nomenclature, a type species (''species typica'') is the species name with which the name of a genus or subgenus is considered to be permanently taxonomically associated, i.e., the spe ...
for the genera of ''Ca.'' Brocadia, and the first known anammox bacteria. Its name was chosen based on where the bacteria was discovered: 'Brocadia' for the pilot plant it was found in at Gist-brocades, and 'anammoxidans' due to its ability to perform anammox. This was determined after its
16S ribosomal DNA 16S 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 genes coding for it are referred to as 16S rRNA ...
was extracted, amplified, and sequenced, and it was seen to fall within the phylum of
Planctomycetota The Planctomycetota are a phylum of widely distributed bacteria, occurring in both aquatic and terrestrial habitats. They play a considerable role in global carbon and nitrogen cycles, with many species of this phylum capable of anaerobic ammoni ...
.


''Candidatus'' Brocadia sinica

''Ca.'' B. sinica was directly compared against its type species in both natural and man-made environments where anammox was performed. ''Ca.'' B. sinica was found to have slightly higher growth temperatures, while ''Ca.'' B. anammoxidans had slightly lower temperatures, but a wider range. Growth pHs remained similar, with ''Ca.'' B. sinica having slightly higher pH requirements. ''Ca.'' B. sinica was also shown to have higher resistance to dissolved oxygen in
growth media A growth medium or culture medium is a solid, liquid, or semi-solid designed to support the growth of a population of microorganisms or cells via the process of cell proliferation or small plants like the moss ''Physcomitrella patens''. Differe ...
, comparatively. ''Ca.'' B. sinica outcompetes ''Ca.'' B. anammoxidans in man-made environments with high
ammonium Ammonium is a modified form of ammonia that has an extra hydrogen atom. It is a positively charged (cationic) polyatomic ion, molecular ion with the chemical formula or . It is formed by the protonation, addition of a proton (a hydrogen nucleu ...
and
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 ...
, but it would be outcompeted in natural environments due to limited ammonium and nitrite availability.


''Candidatus'' Brocadia fulgida

''Ca.'' B. fulgida has been found to autofluoresce in the right concentrations and aggregations, but not as a single cell. More specifically, an autofluorescent substance has been found surrounding its cells, but has not yet been further identified. Two excitation and emission values were found associated with this autofluorescence, which could suggest more than one compound contributes to this unique trait. This species is noted to have higher
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 ...
and
acetate An acetate is a salt formed by the combination of acetic acid with a base (e.g. alkaline, earthy, metallic, nonmetallic, or radical base). "Acetate" also describes the conjugate base or ion (specifically, the negatively charged ion called ...
oxidation rates than its type species ''Ca.'' B. anammoxidans. It's proposed that ''Ca.'' B. fulgida converts acetate to CO2 first, and then incorporates the CO2 into
acetyl-CoA Acetyl-CoA (acetyl coenzyme A) is a molecule that participates in many biochemical reactions in protein, carbohydrate and lipid metabolism. Its main function is to deliver the acetyl group to the citric acid cycle (Krebs cycle) to be oxidation, o ...
. This is unique because it is much more difficult than the pathway that converts acetate to acetyl-CoA.


References

{{Taxonbar, from=Q12062662 Planctomycetota Candidatus taxa