Neomura (from Ancient Greek neo- "new", and Latin -murus "wall") is a proposed clade of life composed of the two domains Archaea and Eukaryota, coined by Thomas Cavalier-Smith in 2002. [1] Its name reflects the hypothesis that both archaea and eukaryotes evolved out of the domain Bacteria, and one of the major changes was the replacement of the bacterial peptidoglycan cell walls with other glycoproteins.
As of October 2024 [update] , the neomuran hypothesis is not accepted by most scientific workers; many molecular phylogenies suggest that eukaryotes are most closely related to one group of archaeans and evolved from them, rather than forming a clade with all archaeans, and that archaea and bacteria are sister groups both descended from the last universal common ancestor (LUCA). Other scenarios have been proposed based on competing phylogenies, and the relationship between the three domains of life (Archaea, Bacteria, and Eukaryota) was described in 2021 as "one of Biology's greatest mysteries". [2]
Considered as comprising the Archaea and the Eukaryota, the Neomura are a very diverse group, containing all of the multicellular species, as well as all of the most extremophilic species, but they all share certain molecular characteristics. All neomurans have histones to help with chromosome packaging, and most have introns. All use the molecule methionine as the initiator amino acid for protein synthesis (bacteria use formylmethionine). Finally, all neomurans use several kinds of RNA polymerase, whereas bacteria use only one.[ citation needed ]
There are several hypotheses for the phylogenetic relationships between archaeans and eukaryotes.
When Carl Woese first published his three-domain system in 1990, [3] [4] it was believed that the domains Bacteria, Archaea, and Eukaryota were equally old and equally related on the tree of life. However certain evidence began to suggest that Eukaryota and Archaea were more closely related to each other than either was to Bacteria. This evidence included the common use of cholesterols and proteasomes, which are complex molecules not found in most bacteria, leading to the inference that the root of life lay between Bacteria on the one hand, and Archaea and Eukaryota combined on the other, i.e. that there were two primary branches of life subsequent to the LUCA – Bacteria and Neomura (not then called by this name).
| later called Neomura |
The "three primary domains" (3D) scenario was one of the two hypotheses considered plausible in a 2010 review of the origin of eukaryotes. [5]
In a 2002 paper, and subsequent papers, Thomas Cavalier-Smith and coworkers have promulgated a hypothesis that Neomura is a clade deeply nested with Eubacteria with Actinomycetota as its sister group. He wrote, "Eukaryotes and archaebacteria form the clade neomura and are sisters, as shown decisively by genes fragmented only in archaebacteria and by many sequence trees. This sisterhood refutes all theories that eukaryotes originated by merging an archaebacterium and an α-proteobacterium, which also fail to account for numerous features shared specifically by eukaryotes and actinobacteria." [1]
These include the presence of cholesterols and proteasomes in Actinomycetota as well as in Neomura. Features of this complexity are unlikely to evolve more than once in separate branches, so either there was a horizontal transfer of those two pathways, or Neomura evolved from this particular branch of the bacterial tree.
As early as 2010, the major competitor to the three domains scenario for the origin of eukaryotes was the "two domains" (2D) scenario, in which eukaryotes emerged from within the archaea. [5] The discovery of a major group within the Archaea, Lokiarchaeota, to which eukaryotes are more genetically similar than to other archaeans, is not consistent with the Neomura hypothesis. Instead, it supports the hypothesis that eukaryotes emerged from within one group of archaeans: [6]
| Archaea |
A 2016 study using 16 universally-conserved ribosomal proteins supports the two domain view. Its "new view of the tree of life" shows eukaryotes as a small group nested within Archaea, in particular within the TACK superphylum. However, the origin of eukaryotes remains unresolved, and the two domain and three domain scenarios remain viable hypotheses. [7]
An alternative to the placement of Eukaryota within Archaea is that both domains evolved from within Bacteria, which is then the ancestral group. This view is similar to the derived clade view above, but the bacterial group involved is different. The evidence for this phylogeny includes the detection of membrane coat proteins and of processes related to phagocytosis in the bacterial Planctomycetes. Although Archaea and Eukaryota are sisters in this view, their joint sister is a bacterial group called PVC for short (the Planctomycetes-Verrucomicrobia-Chlamydiae superphylum): [2]
| traditional Bacteria |
On this view, the traditional Bacteria taxon is paraphyletic. Eukaryotes were not formed by a symbiotic merger between an archaeon and a bacterium, but by the merger of two bacteria, albeit that one was highly modified. [2] In a 2020 paper, Cavalier-Smith accepted the planctobacterial origins of Archaea and Eukaryota, noting that the evidence was not sufficient to safely distinguish between the two possibilities that eukaryotes are sisters of all archaea (as shown in the cladogram above) or that eukaryotes evolved from filarchaeotes, i.e. within Archaea (the two-domain view above). [8]
Carl Richard Woese was an American microbiologist and biophysicist. Woese is famous for defining the Archaea in 1977 through a pioneering phylogenetic taxonomy of 16S ribosomal RNA, a technique that has revolutionized microbiology. He also originated the RNA world hypothesis in 1967, although not by that name. Woese held the Stanley O. Ikenberry Chair and was professor of microbiology at the University of Illinois Urbana–Champaign.
In biology, a kingdom is the second highest taxonomic rank, just below domain. Kingdoms are divided into smaller groups called phyla.
In biological taxonomy, a domain, also dominion, superkingdom, realm, or empire, is the highest taxonomic rank of all organisms taken together. It was introduced in the three-domain system of taxonomy devised by Carl Woese, Otto Kandler and Mark Wheelis in 1990.
The three-domain system is a taxonomic classification system that groups all cellular life into three domains, namely Archaea, Bacteria and Eukarya, introduced by Carl Woese, Otto Kandler and Mark Wheelis in 1990. The key difference from earlier classifications such as the two-empire system and the five-kingdom classification is the splitting of Archaea from Bacteria as completely different organisms. It has been challenged by the two-domain system that divides organisms into Bacteria and Archaea only, as Eukaryotes are considered as a clade of Archaea.
The Thermoproteota are prokaryotes that have been classified as a phylum of the domain Archaea. Initially, the Thermoproteota were thought to be sulfur-dependent extremophiles but recent studies have identified characteristic Thermoproteota environmental rRNA indicating the organisms may be the most abundant archaea in the marine environment. Originally, they were separated from the other archaea based on rRNA sequences; other physiological features, such as lack of histones, have supported this division, although some crenarchaea were found to have histones. Until 2005 all cultured Thermoproteota had been thermophilic or hyperthermophilic organisms, some of which have the ability to grow at up to 113 °C. These organisms stain Gram negative and are morphologically diverse, having rod, cocci, filamentous and oddly-shaped cells. Recent evidence shows that some members of the Thermoproteota are methanogens.
Euryarchaeota is a kingdom of archaea. Euryarchaeota are highly diverse and include methanogens, which produce methane and are often found in intestines; halobacteria, which survive extreme concentrations of salt; and some extremely thermophilic aerobes and anaerobes, which generally live at temperatures between 41 and 122 °C. They are separated from the other archaeans based mainly on rRNA sequences and their unique DNA polymerase.
In taxonomy, the Methanopyri are a class of the Euryarchaeota.
The two-empire system was the top-level biological classification system in general use from the early 20th century until the establishment of the three-domain system. It classified cellular life into Prokaryota and Eukaryota as either "empires" or "superkingdoms". When the three-domain system was introduced, some biologists preferred the two-superkingdom system, claiming that the three-domain system overemphasized the division between Archaea and Bacteria. However, given the current state of knowledge and the rapid progress in biological scientific advancement, especially due to genetic analyses, that view has all but vanished.
Gracilicutes is a clade in bacterial phylogeny.
The PVC superphylum is a superphylum of bacteria named after its three important members, Planctomycetota, Verrucomicrobiota, and Chlamydiota. Cavalier-Smith postulated that the PVC bacteria probably lost or reduced their peptidoglycan cell wall twice. It has been hypothesised that a member of the PVC clade might have been the host cell in the endosymbiotic event that gave rise to the first proto-eukaryotic cell.
In the taxonomy of microorganisms, the Methanomicrobia are a class of the Euryarchaeota.
Methanococci is a class of methanogenic archaea in the phylum Euryarchaeota. They can be mesophilic, thermophilic or hyperthermophilic.
A prokaryote is a single-cell organism whose cell lacks a nucleus and other membrane-bound organelles. The word prokaryote comes from the Ancient Greek πρό (pró), meaning 'before', and κάρυον (káruon), meaning 'nut' or 'kernel'. In the two-empire system arising from the work of Édouard Chatton, prokaryotes were classified within the empire Prokaryota. However in the three-domain system, based upon molecular analysis, prokaryotes are divided into two domains: Bacteria and Archaea. Organisms with nuclei are placed in a third domain: Eukaryota.
Archaea is a domain of organisms. Traditionally, Archaea only included its prokaryotic members, but this sense has been found to be paraphyletic, as eukaryotes are now known to have evolved from archaea. Even though the domain Archaea includes eukaryotes, the term "archaea" in English still generally refers specifically to prokaryotic members of Archaea. Archaea were initially classified as bacteria, receiving the name archaebacteria, but this term has fallen out of use.
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 1987 paper by Carl Woese.
There are several models of the Branching order of bacterial phyla, one of these was proposed in 2002 and 2004 by Thomas Cavalier-Smith. In this frame of work, the branching order of the major lineage of bacteria are determined based on some morphological characters, such as cell wall structure, and not based on the molecular evidence.
The eocyte hypothesis in evolutionary biology proposes that the eukaryotes originated from a group of prokaryotes called eocytes. After his team at the University of California, Los Angeles discovered eocytes in 1984, James A. Lake formulated the hypothesis as "eocyte tree" that proposed eukaryotes as part of archaea. Lake hypothesised the tree of life as having only two primary branches: prokaryotes, which include Bacteria and Archaea, and karyotes, that comprise Eukaryotes and eocytes. Parts of this early hypothesis were revived in a newer two-domain system of biological classification which named the primary domains as Archaea and Bacteria.
The initial version of a classification system of life by British zoologist Thomas Cavalier-Smith appeared in 1978. This initial system continued to be modified in subsequent versions that were published until he died in 2021. As with classifications of others, such as Carl Linnaeus, Ernst Haeckel, Robert Whittaker, and Carl Woese, Cavalier-Smith's classification attempts to incorporate the latest developments in taxonomy., Cavalier-Smith used his classifications to convey his opinions about the evolutionary relationships among various organisms, principally microbial. His classifications complemented his ideas communicated in scientific publications, talks, and diagrams. Different iterations might have a wider or narrow scope, include different groupings, provide greater or lesser detail, and place groups in different arrangements as his thinking changed. His classifications has been a major influence in the modern taxonomy, particularly of protists.
The two-domain system is a biological classification by which all organisms in the tree of life are classified into two domains, Bacteria and Archaea. It emerged from development of knowledge of archaea diversity and challenges the widely accepted three-domain system that classifies life into Bacteria, Archaea, and Eukarya. It was preceded by the eocyte hypothesis of James A. Lake in the 1980s, which was largely superseded by the three-domain system, due to evidence at the time. Better understanding of archaea, especially of their roles in the origin of eukaryotes through symbiogenesis with bacteria, led to the revival of the eocyte hypothesis in the 2000s. The two-domain system became more widely accepted after the discovery of a large group (superphylum) of archaea called Asgard in 2017, which evidence suggests to be the evolutionary root of eukaryotes, thereby making eukaryotes members of the domain Archaea.