Terrabacteria | |
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Scanning electron micrograph of Actinomyces israelii (Actinomycetota) | |
Scientific classification | |
Domain: | Bacteria |
Clade: | Terrabacteria Battistuzzi et al., 2004, Battistuzzi & Hedges, 2009 |
Phyla | |
Synonyms | |
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Terrabacteria is a taxon containing approximately two-thirds of prokaryote species, including those in the gram positive phyla (Actinomycetota and Bacillota) as well as the phyla "Cyanobacteria", Chloroflexota, and Deinococcota. [1] [2]
It derives its name (terra = "land") from the evolutionary pressures of life on land. Terrabacteria possess important adaptations such as resistance to environmental hazards (e.g., desiccation, ultraviolet radiation, and high salinity) and oxygenic photosynthesis. Also, the unique properties of the cell wall in gram-positive taxa, which likely evolved in response to terrestrial conditions, have contributed toward pathogenicity in many species. [2] These results now leave open the possibility that terrestrial adaptations may have played a larger role in prokaryote evolution than currently understood. [1] [2]
Terrabacteria was proposed in 2004 for Actinomycetota, "Cyanobacteria", and Deinococcota [1] and was expanded later to include Bacillota and Chloroflexota. [2] Other phylogenetic analyses [3] [4] [5] have supported the close relationships of these phyla. Most species of prokaryotes not placed in Terrabacteria were assigned to the taxon Hydrobacteria, [2] in reference to the moist environment inferred for the common ancestor of those species. Some molecular phylogenetic analyses [6] [7] have not supported this dichotomy of Terrabacteria and Hydrobacteria, but the most recent genomic analyses, [4] [5] including those that have focused on rooting the tree, [4] have found these two groups to be monophyletic. [4]
Terrabacteria and Hydrobacteria were inferred to have diverged approximately 3 billion years ago, suggesting that land (continents) had been colonized by prokaryotes at that time. [2] Together, Terrabacteria and Hydrobacteria form a large group containing 97% of prokaryotes and 99% of all species of Bacteria known by 2009, and placed in the taxon Selabacteria, in allusion to their phototrophic abilities (selas = light). [8] Currently, the bacterial phyla that are outside of Terrabacteria + Hydrobacteria, and thus justifying the taxon Selabacteria, are debated and may or may not include Fusobacteria. [2] [4]
The name “Glidobacteria” [9] included some members of Terrabacteria but excluded the large gram positive groups, Bacillota and Actinomycetota, and is not supported by molecular phylogenetic data. [1] [2] [3] [6] [7] [4] [5] Moreover, the article naming Glidobacteria [9] did not include a molecular phylogeny or statistical analyses and did not follow the widely used three-domain system. For example, it claimed that eukaryotes split from Archaea very recently (~900 Mya), which is contradicted by the fossil record, [10] and that lineage of eukaryotes + Archaea was nested within Bacteria as a close relative of Actinomycetota.
In 2022, new rules were introduced for kingdom-level taxa of prokaryotes, and the same two authors who proposed those new rules, proposed new names in 2024. [11] They concluded that “the taxonomically preferable solution for bacterial kingdoms seems to be to accept the subdivision apparent in the study by Battistuzzi and Hedges,” with refinement. [2] The new (and only valid) name is Bacillati. [11]
The phylogenetic tree according to the phylogenetic analyses of Battistuzzi and Hedges (2009) is the following and with a molecular clock calibration. [1] [2]
Recent molecular analyses have found roughly the following relationships including other phyla, whose relationships were uncertain. [12] [13] [14] [15] [16] [17]
Terrabacteria |
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On the other hand, Coleman et al. [4] named the clade composed of Thermotogota, Deinococcota, Synergistota and related as DST and furthermore the analysis suggests that ultra-small bacteria (CPR group) may belong to Terrabacteria being more closely related to Chloroflexota. According to this study the phylum Aquificota sometimes included belongs to Hydrobacteria and that the phylum Fusobacteriota can belong to both Terrabacteria and Hydrobacteria. The result was the following: [4]
Terrabacteria | |
In bacteriology, gram-positive bacteria are bacteria that give a positive result in the Gram stain test, which is traditionally used to quickly classify bacteria into two broad categories according to their type of cell wall.
Bacillota is a phylum of bacteria, most of which have gram-positive cell wall structure. The renaming of phyla such as Firmicutes in 2021 remains controversial among microbiologists, many of whom continue to use the earlier names of long standing in the literature.
The Actinomycetota are a diverse phylum of Gram-positive bacteria with high GC content. They can be terrestrial or aquatic. They are of great importance to land flora because of their contributions to soil systems. In soil they help to decompose the organic matter of dead organisms so the molecules can be taken up anew by plants. While this role is also played by fungi, Actinomycetota are much smaller and likely do not occupy the same ecological niche. In this role the colonies often grow extensive mycelia, as fungi do, and the name of an important order of the phylum, Actinomycetales, reflects that they were long believed to be fungi. Some soil actinomycetota live symbiotically with the plants whose roots pervade the soil, fixing nitrogen for the plants in exchange for access to some of the plant's saccharides. Other species, such as many members of the genus Mycobacterium, are important pathogens.
The Chloroflexia are a class of bacteria in the phylum Chloroflexota. Chloroflexia are typically filamentous, and can move about through bacterial gliding. It is named after the order Chloroflexales.
Deinococcota is a phylum of bacteria with a single class, Deinococci, that are highly resistant to environmental hazards, also known as extremophiles. These bacteria have thick cell walls that give them gram-positive stains, but they include a second membrane and so are closer in structure to those of gram-negative bacteria.
The phylum Bacteroidota is composed of three large classes of Gram-negative, nonsporeforming, anaerobic or aerobic, and rod-shaped bacteria that are widely distributed in the environment, including in soil, sediments, and sea water, as well as in the guts and on the skin of animals.
Mollicutes is a class of bacteria distinguished by the absence of a cell wall. The word "Mollicutes" is derived from the Latin mollis, and cutis. Individuals are very small, typically only 0.2–0.3 μm in size and have a very small genome size. They vary in form, although most have sterols that make the cell membrane somewhat more rigid. Many are able to move about through gliding, but members of the genus Spiroplasma are helical and move by twisting. The best-known genus in the Mollicutes is Mycoplasma. Colonies show the typical "fried-egg" appearance.
The Thermotogota are a phylum of the domain Bacteria. The phylum contains a single class, Thermotogae. The phylum Thermotogota is composed of Gram-negative staining, anaerobic, and mostly thermophilic and hyperthermophilic bacteria. It's the sole phylum in the kingdom Thermotogati.
Gracilicutes is a clade in bacterial phylogeny.
The Chloroflexota are a phylum of bacteria containing isolates with a diversity of phenotypes, including members that are aerobic thermophiles, which use oxygen and grow well in high temperatures; anoxygenic phototrophs, which use light for photosynthesis ; and anaerobic halorespirers, which uses halogenated organics as electron acceptors.
In biology, a phylum is a level of classification or taxonomic rank below kingdom and above class. Traditionally, in botany the term division has been used instead of phylum, although the International Code of Nomenclature for algae, fungi, and plants accepts the terms as equivalent. Depending on definitions, the animal kingdom Animalia contains about 31 phyla, the plant kingdom Plantae contains about 14 phyla, and the fungus kingdom Fungi contains about 8 phyla. Current research in phylogenetics is uncovering the relationships among phyla within larger clades like Ecdysozoa and Embryophyta.
Bacterial phyla constitute the major lineages of the domain Bacteria. While the exact definition of a bacterial phylum is debated, a popular definition is that a bacterial phylum is a monophyletic lineage of bacteria whose 16S rRNA genes share a pairwise sequence identity of ~75% or less with those of the members of other bacterial phyla.
Bacterial taxonomy is subfield of taxonomy devoted to the classification of bacteria specimens into taxonomic ranks.
There are several models of the Branching order of bacterial phyla, one of these was proposed in 2004 by Battistuzzi and Hedges, note the coinage of the taxa Terrabacteria and Hydrobacteria.
Conserved signature inserts and deletions (CSIs) in protein sequences provide an important category of molecular markers for understanding phylogenetic relationships. CSIs, brought about by rare genetic changes, provide useful phylogenetic markers that are generally of defined size and they are flanked on both sides by conserved regions to ensure their reliability. While indels can be arbitrary inserts or deletions, CSIs are defined as only those protein indels that are present within conserved regions of the protein.
Prokaryotic ubiquitin-like protein (Pup) is a functional analog of ubiquitin found in the prokaryote Mycobacterium tuberculosis. Like ubiquitin, Pup serves to direct proteins to the proteasome for degradation in the Pup-proteasome system (PPS). However, the enzymology of ubiquitylation and pupylation is different, owing to their distinct evolutionary origins. In contrast to the three-step reaction of ubiquitylation, pupylation requires only two steps, and thus only two enzymes are involved in pupylation. The enzymes involved in pupylation are descended from glutamine synthetase.
Hydrobacteria is a taxon containing approximately one-third of prokaryote species, mostly gram-negative bacteria and their relatives. It was found to be the closest relative of an even larger group of Bacteria, Terrabacteria, which are mostly gram-positive bacteria. The name Hydrobacteria refers to the moist environment inferred for the common ancestor of those species. In contrast, species of Terrabacteria possess adaptations for life on land. Since 2024, the only validly published name for this group is Pseudomonadati.
Asgard or Asgardarchaeota is a proposed superphylum belonging to the domain Archaea that contain eukaryotic signature proteins. It appears that the eukaryotes, the domain that contains the animals, plants, and fungi, emerged within the Asgard, in a branch containing the Heimdallarchaeota. This supports the two-domain system of classification over the three-domain system.
The candidate phyla radiation is a large evolutionary radiation of bacterial lineages whose members are mostly uncultivated and only known from metagenomics and single cell sequencing. They have been described as nanobacteria or ultra-small bacteria due to their reduced size (nanometric) compared to other bacteria.