Borg (microbiology)
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Borgs are large (up to ~1 Mbp) extrachromosomal linear
DNA Deoxyribonucleic acid (; DNA) is a polymer composed of two polynucleotide chains that coil around each other to form a double helix. The polymer carries genetic instructions for the development, functioning, growth and reproduction of al ...
elements found in
methanotrophic Methanotrophs (sometimes called methanophiles) are prokaryotes that metabolize methane as their source of carbon and chemical energy. They are bacteria or archaea, can grow aerobically or anaerobically, and require single-carbon compounds to ...
archaea Archaea ( ) is a Domain (biology), domain of organisms. Traditionally, Archaea only included its Prokaryote, prokaryotic members, but this has since been found to be paraphyletic, as eukaryotes are known to have evolved from archaea. Even thou ...
(specifically '' Methanoperedens'' spp.) that live in oxygen-starved environments such as deep mud. They have been found in organisms isolated from wetland, aquifer, and riverbed environments, as well as a deserted mercury mine, in
California California () is a U.S. state, state in the Western United States that lies on the West Coast of the United States, Pacific Coast. It borders Oregon to the north, Nevada and Arizona to the east, and shares Mexico–United States border, an ...
and
Colorado Colorado is a U.S. state, state in the Western United States. It is one of the Mountain states, sharing the Four Corners region with Arizona, New Mexico, and Utah. It is also bordered by Wyoming to the north, Nebraska to the northeast, Kansas ...
. They were first described by Basem Al-Shayeb and Jill Banfield in 2022. The nature of Borgs remains unclear; they are thought to be "giant linear
plasmid A plasmid is a small, extrachromosomal DNA molecule within a cell that is physically separated from chromosomal DNA and can replicate independently. They are most commonly found as small circular, double-stranded DNA molecules in bacteria and ...
s" or
giant virus A giant virus, sometimes referred to as a girus, is a very large virus, some of which are larger than typical bacteria. All known giant viruses belong to the phylum ''Nucleocytoviricota''. Description While the exact criteria as defined in the sc ...
es. At least 19 different types have been identified, all of which co-occur within ''Methanoperedens'', which shares many of their genes. ''Methanoperedens''' main chromosome is only about three times larger than the Borgs it hosts. It is speculated that Borgs may augment ''Methanoperedens capacity for
anaerobic oxidation of methane Anaerobic oxidation of methane (AOM) is a methane-consuming microbial process occurring in anoxic marine and freshwater sediments. AOM is known to occur among mesophiles, but also in psychrophiles, thermophiles, halophiles, acidophiles, and al ...
and protein production.


Discovery

Borgs were discovered on March 8, 2020 by Jill Banfield and her research group at the
University of California, Berkeley The University of California, Berkeley (UC Berkeley, Berkeley, Cal, or California), is a Public university, public Land-grant university, land-grant research university in Berkeley, California, United States. Founded in 1868 and named after t ...
. The name "Borg" was chosen as a reference to the Star Trek faction of the same name, due to the novel genetic elements' apparent propensity to assimilate genes from microbes, most notably ''Methanoperedens''; the name was originally suggested by Banfield's son.


Features

The structures of Borg genomes are conserved and distinct from the plasmids and chromosomes of ''Methanoperedens'', as well as other archaeal genomes''.'' Borgs do not contain protein-coding genes that are associated with plasmids or viruses; they also lack
rRNA Ribosomal ribonucleic acid (rRNA) is a type of non-coding RNA which is the primary component of ribosomes, essential to all cells. rRNA is a ribozyme which carries out protein synthesis in ribosomes. Ribosomal RNA is transcribed from ribosomal ...
genes,
origins of replication The origin of replication (also called the replication origin) is a particular sequence in a genome at which replication is initiated. Propagation of the genetic material between generations requires timely and accurate duplication of DNA by semi ...
, or other vital genes and features that are commonly found within minichromosomes (also known as megaplasmids) found in archaea. Borgs range from 0.66-0.92 Mbp in length, larger than the genomes of any known archaeal viruses. The sizes of Borg genomes are more characteristic of
eukaryote The eukaryotes ( ) constitute the Domain (biology), domain of Eukaryota or Eukarya, organisms whose Cell (biology), cells have a membrane-bound cell nucleus, nucleus. All animals, plants, Fungus, fungi, seaweeds, and many unicellular organisms ...
-specific double-stranded DNA viruses from the phylum ''
Nucleocytoviricota ''Nucleocytoviricota'' is a phylum of viruses. Members of the phylum are also known as the nucleocytoplasmic large DNA viruses (NCLDV), which serves as the basis of the name of the phylum with the suffix - for virus phylum. These viruses are refe ...
'', also known as nucleocytoplasmic large DNA viruses (NCLDV), which can surpass 2.5 Mbp. Tandem
direct repeat Direct repeats are a type of genetic sequence that consists of two or more repeats of a specific sequence. In other words, the direct repeats are nucleotide sequences present in multiple copies in the genome. Generally, a direct repeat occurs when a ...
sequences are prevalent throughout Borg genomes, and they are terminated by
inverted repeats An inverted repeat (or IR) is a single stranded sequence of nucleotides followed downstream by its reverse complement. The intervening sequence of nucleotides between the initial sequence and the reverse complement can be any length including zero ...
. This differs from the megaplasmids of some bacteria, which carry interspaced repeats and usually are not responsible for encoding necessary genes. Below is a list of genes known to be encoded by Borgs (note that not every Borg subtype contains the same genes): * Mobile or
transposable element A transposable element (TE), also transposon, or jumping gene, is a type of mobile genetic element, a nucleic acid sequence in DNA that can change its position within a genome. The discovery of mobile genetic elements earned Barbara McClinto ...
defense systems * Genes involved with the anaerobic oxidation of methane (AOM) * Type III-A CRISPR-Cas system (No spacer acquisition machinery, primarily targets RNA) *
RPL11 60S ribosomal protein L11 is a protein that in humans is encoded by the ''RPL11'' gene. Function Ribosomes, the organelles that catalyze protein synthesis, consist of a small 40S subunit and a large 60S subunit. Together these subunits are ...
(Ribosomal protein L11) *
Glycosyltransferase Glycosyltransferases (GTFs, Gtfs) are enzymes ( EC 2.4) that establish natural glycosidic linkages. They catalyze the transfer of saccharide moieties from an activated nucleotide sugar (also known as the "glycosyl donor") to a nucleophilic gl ...
* Genes involved in DNA and RNA manipulation * Transport genes * Genes involved in energy metabolism * Genes encoding cell surface proteins ** PEGA ** S-layer * Genes encoding membrane-associated proteins (unknown function, possibly affects host's membrane properties) * ''nifHDK'' operon (involved in
nitrogen fixation Nitrogen fixation is a chemical process by which molecular dinitrogen () is converted into ammonia (). It occurs both biologically and abiological nitrogen fixation, abiologically in chemical industry, chemical industries. Biological nitrogen ...
) * Genes involved in
polyhydroxyalkanoate Polyhydroxyalkanoates or PHAs are polyesters produced in nature by numerous microorganisms, including through bacterial fermentation of sugars or lipids. When produced by bacteria they serve as both a source of energy and as a carbon store. More ...
synthesis (Carbon storage; possibly used for when resources are low) * Genes encoding
Tellurium Tellurium is a chemical element; it has symbol Te and atomic number 52. It is a brittle, mildly toxic, rare, silver-white metalloid. Tellurium is chemically related to selenium and sulfur, all three of which are chalcogens. It is occasionally fou ...
resistance proteins * ''
FtsZ FtsZ is a protein encoded by the ''ftsZ'' gene that assembles into a ring at the future site of bacterial cell division (also called the Z ring). FtsZ is a prokaryotic homologue of the eukaryotic protein tubulin. The initials FtsZ mean "Filame ...
'' (tubulin homolog, involved in bactrial cell division) *
Major Vault Protein Major vault protein, also known as lung resistance-related protein is a protein that in humans is encoded by the ''MVP'' gene. 78 copies of the protein assemble into the large compartments called vaults. Function This gene encodes the major ...
homologs *
TCA cycle The citric acid cycle—also known as the Krebs cycle, Szent–Györgyi–Krebs cycle, or TCA cycle (tricarboxylic acid cycle)—is a series of chemical reaction, biochemical reactions that release the energy stored in nutrients through acetyl-Co ...
genes **
citrate synthase Citrate synthase ( E.C. 2.3.3.1 (previously 4.1.3.7)) is an enzyme that exists in nearly all living cells. It functions as a pace-making enzyme in the first step of the citric acid cycle (or Krebs cycle). Citrate synthase is located within euka ...
**
aconitase Aconitase (aconitate hydratase; ) is an enzyme that catalyses the stereochemistry, stereo-specific isomerization of citrate to isocitrate via ''cis''-aconitate in the tricarboxylic acid cycle, a non-redox-active process. Image:Citrate wpmp.p ...
*''cfbB'' and ''cfbC'' * Coenzyme F420:L-glutamate ligase - ''cofE'' * Electron bifurcating complexes * D-Lactate dehydrogenase * Genes involved in
Tetrahydromethanopterin Tetrahydromethanopterin (THMPT, ) is a coenzyme in methanogenesis. It is the carrier of the C1 group as it is reduced to the methyl level, before transferring to the coenzyme M. Tetrahydrosarcinapterin (THSPT, ) is a modified form of THMPT, wher ...
synthesis * Genes encoding
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 ...
proteins * Genes encoding 5,6,7,8-tetrahydromethanopterin hydro-lyase (Fae) (involved in formaldehyde detoxification) * Genes encoding
Carbon monoxide dehydrogenase In enzymology, carbon monoxide dehydrogenase (CODH) () is an enzyme that catalyzes the chemical reaction :CO + H2O + A \rightleftharpoons CO2 + AH2 The chemical process catalyzed by carbon monoxide dehydrogenase is similar to the water-gas shi ...
(CODH) * Genes encoding
plastocyanin Plastocyanin is a copper-containing protein that mediates electron-transfer. It is found in a variety of plants, where it participates in photosynthesis. The protein is a prototype of the blue copper proteins, a family of intensely blue-colored ...
* Genes encoding cupredoxins * Genes encoding multiheme cytochromes (MHC)


References

{{reflist Microbiology DNA