Neomura

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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.

Contents

As of October 2024, 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]

Morphology

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 ]

Phylogeny

There are several hypotheses for the phylogenetic relationships between archaeans and eukaryotes.

Three-domain view

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).

Bacteria

Eukaryota 

Archaea

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]

Derived clade view

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.

Two domains view

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]

Bacteria

archaeans

archaeans 

Eukaryota

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]

One domain view

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]

various groups of bacteria 

PVC bacteria

Archaea

Eukaryota

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]

See also

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<span class="mw-page-title-main">Carl Woese</span> American microbiologist (1928–2012)

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.

<span class="mw-page-title-main">Kingdom (biology)</span> Taxonomic rank

In biology, a kingdom is the second highest taxonomic rank, just below domain. Kingdoms are divided into smaller groups called phyla.

<span class="mw-page-title-main">Domain (biology)</span> Taxonomic rank

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.

<span class="mw-page-title-main">Three-domain system</span> Hypothesis for classification of life

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.

<span class="mw-page-title-main">Thermoproteota</span> Phylum of archaea

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<span class="mw-page-title-main">Euryarchaeota</span> Phylum of archaea

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<span class="mw-page-title-main">Two-empire system</span> Biological classification system

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<span class="mw-page-title-main">PVC superphylum</span> Superphylum of bacteria

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<span class="mw-page-title-main">Methanococci</span> Class of archaea

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<span class="mw-page-title-main">Archaea</span> Domain of organisms

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<span class="mw-page-title-main">Eocyte hypothesis</span> Hypothesis in evolutionary biology

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<span class="mw-page-title-main">Two-domain system</span> Biological classification system

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References

  1. 1 2 Cavalier-Smith T (March 2002). "The phagotrophic origin of eukaryotes and phylogenetic classification of Protozoa". Int. J. Syst. Evol. Microbiol. 52 (Pt 2): 297–354. doi:10.1099/00207713-52-2-297. PMID   11931142.
  2. 1 2 3 Devos, Damien P (2021), "Reconciling Asgardarchaeota Phylogenetic Proximity to Eukaryotes and Planctomycetes Cellular Features in the Evolution of Life", Molecular Biology and Evolution, 38 (9): 3531–3542, doi:10.1093/molbev/msab186, PMC   8382908 , PMID   34229349
  3. Woese CR, Fox GE (November 1977). "Phylogenetic structure of the prokaryotic domain: the primary kingdoms". Proceedings of the National Academy of Sciences of the United States of America. 74 (11): 5088–90. Bibcode:1977PNAS...74.5088W. doi: 10.1073/pnas.74.11.5088 . PMC   432104 . PMID   270744.
  4. Woese CR, Kandler O, Wheelis ML (June 1990). "Towards a natural system of organisms: proposal for the domains Archaea, Bacteria, and Eucarya". Proceedings of the National Academy of Sciences of the United States of America. 87 (12): 4576–9. Bibcode:1990PNAS...87.4576W. doi: 10.1073/pnas.87.12.4576 . PMC   54159 . PMID   2112744.
  5. 1 2 Gribaldo, Simonetta; Poole, Anthony M.; Daubin, Vincent; Forterre, Patrick & Brochier-Armanet, Céline (2010-10-01). "The origin of eukaryotes and their relationship with the Archaea: are we at a phylogenomic impasse?". Nature Reviews Microbiology. 8 (10): 743–752. doi:10.1038/nrmicro2426. PMID   20844558. S2CID   12111029.
  6. López-García, Purificación & Moreira, David (2015). "Open Questions on the Origin of Eukaryotes". Trends in Ecology and Evolution. 30 (11): 697–708. Bibcode:2015TEcoE..30..697L. doi:10.1016/j.tree.2015.09.005. PMC   4640172 . PMID   26455774.
  7. Hug, Laura A.; Baker, Brett J.; Anantharaman, Karthik; Brown, Christopher T.; Probst, Alexander J.; Castelle, Cindy J.; Butterfield, Cristina N.; Hernsdorf, Alex W.; Amano, Yuki; Ise, Kotaro; Suzuki, Yohey; Dudek, Natasha; Relman, David A.; Finstad, Kari M.; Amundson, Ronald; Thomas, Brian C. & Banfield, Jillian F. (2016-04-11). "A new view of the tree of life". Nature Microbiology. 1 (5): 16048. doi: 10.1038/nmicrobiol.2016.48 . PMID   27572647.
  8. Cavalier-Smith, Thomas & Chao, Ema E-Yung (2020). "Multidomain ribosomal protein trees and the planctobacterial origin of neomura (eukaryotes, archaebacteria)". Protoplasma. 257 (3): 621–753. doi:10.1007/s00709-019-01442-7. PMC   7203096 . PMID   31900730.

Further reading