Primal ancestor

from Wikipedia, the free encyclopedia
Modern representation of the relationships of all recent organisms in the form of a so-called cladogram (to this extent also called “Tree of Life”). The primal ancestor (LUCA) is at the root of this cladogram. The position of the most recent common stem form of plants, animals, fungi and others is also recorded. a. Euglens and Leishmanias (LECA; see below )

The primal ancestor (English Last Universal Ancestor , LUA , or Last Universal Common / Cellular Ancestor , LUCA ) is the - hypothetical - last common ancestral form of all today's ( recent ) cellular organisms . It is postulated by the so-called monophyletic theories of ancestry , which assume that all living beings on earth, plants , animals , fungi and unicellular forms , can be traced back to a single "primitive" archetype, while all other contemporary, likely existing on the early earth, similar primitive forms have not left any descendants in the recent living world or in recent geological and evolutionary history. It is assumed in these hypotheses that this original form lived at least 3.5 billion years ago, i.e. in the late Hadean or early Archean .

background

For the ancestors, according to the thrift principle ( Occam's razor ), it is usually assumed that they had those characteristics of today's living organisms that they have in common: a cell structure , DNA , the (largely identical) genetic code as well as mRNA , tRNA and a ribosomal mediated Translation . Even DNA repair mechanisms show similarities (homologies) across all cellular organisms and some viruses : the family of RecA recombinases with RecA in bacteria, Rad51 and DMC1 in eukaryotes, RadA in archaea and UvsX in bacteriophage T4 , a bacterial virus .

Since the organization of DNA in bacteria differs fundamentally from that in archaea and complex cells ( eukaryotes ), the thesis has recently been advanced that cellular LUCA still belonged to the RNA world . The storage of genetic information in DNA is then seen as a skill that was initially 'invented' by retroviruses and which cellular organisms have acquired several times through the transmission of such viruses . This results in the bacteria on the one hand, and the archaea and eukaryotes on the other. The basic structure of the ribosomes as protein factories and the genetic code, on the other hand, agree so well in all cellular organisms that the LUCA must already have it.

In addition, it is possible that the ancestor was thermophilic . This is, however, increasingly doubted because this hypothesis was based on comparative sequence analyzes of rRNA ( ribosomal RNA ), which give questionable results in thermophilic organisms. It appears that its cell wall components contained murein , a common cell wall component of modern bacteria .

Notes on possible fallacies:

  1. The Last Universal Ancestor is not a specific individual, but a taxon with the rank of a species and thus a population .
  2. This primordial population does not necessarily represent the first existing (cellular) organisms.
  3. Consequently, it was not necessarily the "most primitive" possible forms of life.

The genesis and development of the genetic code and the ribosomes would then have taken place in the RNA world before LUCA. Before, only ribozymes were available as biocatalysts , then real (protein) enzymes (two phases of the RNA world). Modern ribosomes consist of both ribosomal RNA ( rRNA ) and (auxiliary) proteins. If the term Progenot is understood to mean an organism that is still in the process of developing the relationship between genotype and phenotype , then this organism would have to be located in the first phase, i.e. before the LUCA, which is presumably to be located in the second phase. The term protocell or protobiont is sometimes understood similarly: RNA is replicated in a replicating vesicle (see also microsphere ).

A 2010 statistical study found that life is very likely to descend from a single common ancestor. A single common ancestor is 10 2860 times more likely than several. All of his contemporaries have since died out; only the genetic inheritance would have survived to this day.

In contrast, Carl Woese propagated that our genetic pre-LUCA inheritance came from a large number of organisms than from just one species ( biofilm theory , horizontal [lateral] gene transfer ). The ability of primitive living beings for horizontal gene transfer could mean that instead of a single universal ancestor, a community ( gene pool ) of primitive unicellular organisms appears. Since these were in constant gene exchange with one another ('Common ancestral community'), from today's perspective they could appear as a kind of unit (species). This gene exchange could also have been mediated by viruses.

The structure of dendrograms (tree diagrams), based on the genetic distance between all existing cell types, shows a relatively early split between archaea, which are highly resistant to extreme living conditions ( extremophile ), and the other life forms. This has led to some suspicions that the ancestor might have evolved in such extreme ecosystems , such as the deep sea ridge .

Two or three domains

New investigation techniques have changed the picture significantly since around 2015. The previous family trees were based on the comparison of homologous DNA sequences mainly from the smaller subunit of ribosomal RNA . The methods used for the amplification , especially the primers used , now turned out to be unsuitable for a number of unusual prokaryotes. Until then, these organisms had escaped the attention of biologists because they could not be cultivated with the standard microbiological techniques developed over decades, and most of them still cannot be cultivated to this day. Many of them have a very small genome, they lack the synthesis and metabolic pathways essential for life processes, which is why it is assumed that they may only survive as symbionts or parasites , together with other organisms. Since the formal description of prokaryote taxa is tied to cultivation, the newly found lines persist on the status of a Candidatus . But in fact many taxa, due to their divergence with the rank of were tribes must be classified (phyla) found, in which not a member of cultivable included. Their existence is revealed by analytical methods in which genomes in the environment are analyzed directly without detour via cultivation and then assigned to individual lines / species (called metagenomics ) or in which the genome of a single, isolated cell is amplified and sequenced.

One such group, the “ Lokiarchaeota ” belonging to the archaea , unexpectedly turned out to be closely related to the eukaryotes after genetic analysis. Then further sister groups of the "Lokiarchaeota" were found (" Thorarchaeota ", " Odinarchaeota ", " Heimdallarchaeota "); all together are grouped together to form the "Asgard" super group.

A new family tree of life established with the help of the new techniques in 2016, which is based on more than 3000 fully analyzed genomes, found not only a new large group of bacteria, which (with one exception) only contains previously uncultivable bacterial strains. Another result was that the origin of the eukaryotes lies within the archaea and is not, as previously assumed, independent of them. After that, only two (instead of three) domains of life would exist, because the eukaryotes would be reduced to a line within the archaea. However, this had been suggested earlier in the form of the eocyte hypothesis , but was previously considered to be in contradiction to the findings. Now the Asgard supergroup of archaea would come into question as such a starting point for eukaryotes.

A further support for this dichotomy of living beings comes from the knowledge that the DNA replication systems of bacteria on the one hand and archaea and eukaryotes on the other are so different that - in contrast to the ribosomes and the genetic code - for this no common origin ( homology ) can be assumed. It is possible that the cellular LUCA (like many viruses ) still had an RNA genome, i.e. it was a (hypothetical) ribocyte .

The same applies to proteins considered in terms of DNA repair mechanisms. While the proteins from the RecA recombinase family have homologies across all cellular organisms (and some viruses) as mentioned above, there are homologies among eukaryotes and archaea ( Rad54 , Mre11 , and Rad50 ) for another repair protein .

In connection with these discoveries, other previous views on the systematic value of fundamental differences, such as the structure of lipids in the cell membrane or the origin of gram-positive bacteria, were also questioned. However, since the complex basic structure of the ATP synthase is common to all living things (with the exception of a few purely fermenting microbes and primitive multicellular animals such as some myxozoa and corset animals ), the LUCA should already have a basic version of it. Since the ATP synthase is a membrane-bound protein, the LUCA should already have membranes (in some form), so that an at least primitive cellular organism can be assumed in it, for example with a semipermeable membrane envelope (instead of an impermeable one, as is the case with today's cells Membrane envelope and active transport mechanisms). The analysis of the protein families of the different subunits of membrane-bound F- and A- / V-type ATPases allows conclusions to be drawn about the detailed origin and development of these 'motor' enzymes. This further supports the assumption that LUCA was already a cellular organism with at least a primitive form of membranes, and that the eukaryotes evolved from a branch of the archaea.

Another support of the assumption of a common origin of all cellular organisms could be from the comparative stöchiometrichen analysis of fundamental metabolic pathways ( English metabolic pathways ) of model organisms arise. At MIT, Jean-Benoît Lalanne, Gene-Wei Li and colleagues found a remarkable agreement in a publication in March 2018 on a corresponding study on various bacteria and on baker's yeast . The evolutionary forces behind this highly conserved stoichiometry initially remained in the dark.

The results obtained so far are still provisional due to their novelty, they partly contradict the knowledge obtained using traditional methods and therefore still need to be fully examined. However, they suggest the possibility that some prokaryotes with a very small genome may not go back to the reduction of complex, organized ancestors, but that they could be relictary lines that could only survive in the modern world as symbionts.

Urviruses

In contrast to the cellular organisms (biota) there are apparently different primordial viruses (viral LCAs). Although extremely different viruses also have homologous envelope proteins, there are twenty or more fundamentally different such proteins that are not related to one another, which speaks against a common ancestor of all viruses. Viruses seem to have arisen in a period of strong horizontal gene transfer through reduction from primitive cells that had a segmented RNA genome (ribocytes, RNA world), not from a “modern” counterpart. In accordance with the RNA world and ribocyte hypothesis, the original viruses must have been RNA viruses before they discovered the DNA used to store the genetic information.

Urkaryotes, archezoans and LECA

The last common ancestor of the eukaryotes is sometimes referred to as Last Eukaryotic Common Ancestor , LECA for short , in analogy to the Last Universal Common Ancestor .

All eukaryotes examined so far have, in addition to a cell nucleus separated from the cytoplasm by a nuclear membrane , mitochondria or organelles ( hydrogenosome , mitosome, etc.), which evidently descend from mitochondria or have a common ancestor with them. At least genes of mitochondrial origin could be detected everywhere in the cell nucleus DNA as a result of a lateral gene transfer from an earlier mitochondrion to the nucleus. Therefore LECA should already have a nucleus and mitochondrial-like organelles (with their own mtDNA ).

If the acquisition of the cell nucleus preceded that of the mitochondria , there must have previously been amitochondrial organisms with a cell nucleus that acquired the mitochondria through endosymbiosis in the course of their evolution (see endosymbiotic theory ). Carl Woese and George Fox led in 1977 amoeboid for these hypothetical forms of "primitive" predatory protozoans the notion Urkaryoten (Engl. Urkaryotes ) and Thomas Cavalier-Smith led for recent, primitive amitochondriale protozoa, which he regarded as descendants of these Urkaryoten, the group name Archezoa a. However, there was increasing evidence that the Archezoa are secondary amitochondrial real eukaryotes, i.e. descendants of LECA.

As an alternative to the original karyote hypothesis, the acquisition of mitochondria by archaea ( hydrogen hypothesis ) before or simultaneously with the cell nucleus, which may be of viral origin (among the NCLDVs ), is being discussed .

Archaea and Bacteria - LACA and LBCA

By analogy, the last common ancestor of the archaea can be called LACA , that of the bacteria as LCBA . (sometimes also LBCA).

literature

  • Nicolas Glansdorff, Ying Xu, Bernard Labedan: The Last Universal Common Ancestor: emergence, constitution and genetic legacy of an elusive forerunner. Biology Direct 2008, 3:29.

Web links

Individual evidence

  1. Patrick Forterre: Looking for LUCA.
  2. ^ Book presentation by Susanne Liedtke: Matt Ridley, The story of Luga - "Alphabet of Life". In: image of science.
  3. Xiaoxia Liu, Jingxian Zhang, Feng Ni, Xu Dong, Bucong Han, Daxiong Han, Zhiliang Ji, Yufen Zhao: Genome wide exploration of the origin and evolution of amino acids . In: BMC Evolutionary Biology . tape 10 , March 15, 2010, ISSN  1471-2148 , p. 77 , doi : 10.1186 / 1471-2148-10-77 (English).
  4. a b Haseltine CA, Kowalczykowski SC: An archaeal Rad54 protein remodels DNA and stimulates DNA strand exchange by RadA . In: Nucleic Acids Research . 37, No. 8, May 2009, pp. 2757-2770. doi : 10.1093 / nar / gkp068 . PMID 19282450 . PMC 2677860 (free full text).
  5. a b c d e Patrick Forterre: Evolution - The true nature of viruses. In: Spectrum of Science. Issue 8/2017, p. 37 ( online article published on July 19, 2017); Note: The author speaks of "several viral LUCAs", and seems to use the term "LUCA" as a synonym for LCA / MRCA
  6. Dawn J. Brooks, Eric A. Gaucher: A thermophilic last universal ancestor inferred from its estimated amino acid composition. Pp. 200-207 in: David A. Liberles (Ed.): Ancestral Sequence Reconstruction. Oxford University Press, Oxford (UK) 2007, ISBN 978-0-19-929918-8 , Kapitel- doi: 10.1093 / acprof: oso / 9780199299188.001.0001 (alternative full text access: Foundation for Applied Molecular Evolution )
  7. Nicolas Galtier, Nicolas Tourasse, Manolo Gouy: A Nonhyperthermophilic Common Ancestor to Extant Life Forms . In: Science . tape 283 , no. 5399 , January 8, 1999, p. 220–221 , doi : 10.1126 / science.283.5399.220 (English).
  8. Arthur L. Koch: The Exocytoskeleton . In: Journal of Molecular Microbiology and Biotechnology . tape 11 , no. 3–5 , pp. 115-125 , doi : 10.1159 / 000094048 (English).
  9. Arthur L. Koch: The Exoskeleton of Bacterial Cells (the Sacculus): Still a Highly Attractive Target for Antibacterial Agents That Will Last For a Long Time . In: Critical Reviews in Microbiology . tape 26 , no. 1 , January 2000, ISSN  1040-841X , p. 1–35 , doi : 10.1080 / 10408410091154165 (English).
  10. ^ Johann Peter Gogarten, David Deamer: Is LUCA a thermophilic progenote? . In: Nature Microbiology . 1, November 25, 2016, p. 16229. doi : 10.1038 / nmicrobiol.2016.229 .
  11. ^ IA Chen, P. Walde: From self-assembled vesicles to protocells. In: Cold Spring Harbor perspectives in biology. Volume 2, number 7, July 2010, p. A002170, doi: 10.1101 / cshperspect.a002170 , PMID 20519344 , PMC 2890201 (free full text) (review).
  12. ^ Douglas L. Theobald, A formal test of the theory of universal common ancestry . nature.com. Retrieved January 15, 2011. Nature, Volume 465, 2010, pp. 219-222
  13. ^ All Present-day Life Arose From A Single Ancestor . sciencenews.org. Retrieved January 15, 2011.
  14. ^ New Mexico Museum of Natural History and Science: Tree of Life - Introduction , there in particular the interactive information on 'Who were they'
  15. FL Sousa, T. Thiergart, G. Landan, S. Nelson-Sathi, IA Pereira, JF Allen, N. Lane, WF Martin: Early bioenergetic evolution. In: Philosophical transactions of the Royal Society of London. Series B, Biological sciences. Volume 368, number 1622, July 2013, p. 20130088, doi: 10.1098 / rstb.2013.0088 , PMID 23754820 , PMC 3685469 (free full text) (review), Figure 6: The amazing disappearing tree
  16. a b Nick Lane: The Spark of Life - Energy and Evolution , Konrad Theiss Verlag, (C) 2017 by WBG, ISBN 978-3-8062-3484-8 . Original English title: Nick Lane: The Vital Question - Energy, Evolution, and the Origins of Complex Life , Ww Norton, 2015-07-20, ISBN 978-0-393-08881-6 , PDF ( Memento of the original from 10 September 2017 in the Internet Archive ) Info: The archive link was inserted automatically and has not yet been checked. Please check the original and archive link according to the instructions and then remove this notice. . Figure 20 and 16. (the wondrous self-dissolving family tree) @1@ 2Template: Webachiv / IABot / armscoop.com
  17. Christopher T. Brown, Laura A. Hug, Brian C. Thomas, Itai Sharon, Cindy J. Castelle, Andrea Singh, Michael J. Wilkins, Kelly C. Wrighton, Kenneth H. Williams, Jillian F. Banfield (2015): Unusual biology across a group comprising more than 15% of domain Bacteria. Nature 523: 208-211. doi: 10.1038 / nature14486
  18. Lindsey Solden, Karen Lloyd Kelly Wrighton (2016): The bright side of microbial dark matter: lessons learned from the uncultivated majority. Current Opinion in Microbiology 31: 217-226. doi: 10.1016 / j.mib.2016.04.020
  19. Anja Spang, Jimmy H. Saw, Steffen L. Jørgensen, Katarzyna Zaremba-Niedzwiedzka, Joran Martijn, Anders E. Lind, Roel van Eijk, Christa Schleper, Lionel Guy, Thijs JG Ettema (2015): Complex archaea that bridge the gap between prokaryotes and eukaryotes. Nature 521: 173-179. doi: 10.1038 / nature14447
  20. Loki's Castle and the New Tree of Life: Two Domains and the CPR Hidden Folk . University of Hawai'i Reed Lab, Floyd A. Reed, June 24, 2016.
  21. KW Seitz et al .: Genomic reconstruction of a novel, deeply branched sediment archaeal phylum with pathways for acetogenesis and sulfur reduction , Research Gate, January 2016
  22. K et al Zaremba-Niedzwiedzka: Asgard archaea illuminate the origin of eukaryotic cellular complexity . In: Nature . 541, 2017, pp. 353-358. doi : 10.1038 / nature21031 .
  23. Laura A. Hug, Brett J. Baker, Karthik Anantharaman, Christopher T. Brown, Alexander J. Probst, Cindy J. Castelle, Cristina N. Butterfield, Alex W. Hernsdorf, Yuki Amano, Kotaro Ise, Yohey Suzuki, Natasha Dudek , David A. Relman, Kari M. Finstad, Ronald Amundson, Brian C. Thomas, Jillian F. Banfield (2016): A new view of the tree of life. Nature Microbiology 1, Article number: 16048. doi: 10.1038 / nmicrobiol.2016.48
  24. ^ Carl R. Woese, Otto Kandler, Mark L. Wheelis (1990): Towards a natural system of organisms: Proposal for the domains Archaea, Bacteria, and Eucarya. Proceedings of the National Academy of Sciences USA 87: pp. 4576-4579.
  25. Kasie Raymann, Celine Brochier-Armanet, Simonetta Gribaldo (2015): The two-domain tree of life is linked to a new root for the Archaea. Proceedings of the National Academy of Sciences USA 112 (21): pp. 6670-6675. doi: 10.1073 / pnas.1420858112
  26. James A. Lake , E. Henderson, M. Oakes, M. W. Clark: Eocytes: A new ribosome structure indicates a kingdom with a close relationship to eukaryotes . Proceedings of the National Academy of Sciences USA 81, 1984, pp. 3786-3790.
  27. Yarus M: Primordial genetics: phenotype of the ribocyte . In: Annu. Rev. Genet. . 36, 2002, pp. 125-51. doi : 10.1146 / annurev.genet.36.031902.105056 . PMID 12429689 .
  28. Huang Q, Liu L, Liu J, Ni J, She Q, Shen Y: Efficient 5'-3 'DNA end resection by HerA and NurA is essential for cell viability in the crenarchaeon Sulfolobus islandicus . In: BMC Molecular Biology . 16, 2015, p. 2. doi : 10.1186 / s12867-015-0030-z . PMID 25880130 . PMC 4351679 (free full text).
  29. Lin Z, Kong H, Nei M, Ma H: Origins and evolution of the recA / RAD51 gene family: evidence for ancient gene duplication and endosymbiotic gene transfer . In: Proceedings of the National Academy of Sciences of the United States of America . 103, No. 27, July 2006, pp. 10328-10333. bibcode : 2006PNAS..10310328L . doi : 10.1073 / pnas.0604232103 . PMID 16798872 . PMC 1502457 (free full text).
  30. Laura Villanueva, Stefan Schouten, Jaap S. Sinninghe Damsté (2016): Phylogenomic analysis of lipid biosynthetic genes of Archaea shed light on the 'lipid divide'. Environmental Microbiology 19: 54-69. doi: 10.1111 / 1462-2920.13361
  31. ^ Sergio A. Muñoz-Gómez & Andrew J. Roger (2016): Phylogenomics: Leaving negative ancestors behind. eLife 2016; 5: e20061. doi: 10.7554 / eLife.14589
  32. Jan Osterkamp: First animal without breathing and mitochondria , on: Spektrum.de from February 25, 2020
  33. Tel Aviv University researchers discover unique non-oxygen breathing animal , on: EurekAlert! from February 25, 2020
  34. Andy Coghaln: Zoologger: The mud creature did lives without oxygen on: NewScientist from 7 April 2010
  35. Roberto Danovaro et al. : The first metazoa living in permanently anoxic conditions , in: BMC Biology, Volume 8, No. 30, April 6, 2010, doi: 10.1186 / 1741-7007-8-30
  36. Armen Y Mulkidjanian, Michael Y Galperin, Kira S Makarova, Yuri I Wolf and Eugene V Koonin: Evolutionary primacy of sodium bioenergetics . In: Biology Direct . 3, No. 13, 2008. doi : 10.1186 / 1745-6150-3-13 .
  37. arms Y Mulkidjanian, Kira Makarova S, Y Michael Galperin, Eugene V Koonin: Inventing the dynamo machine: the evolution of the F-type and V-type ATPases . In: Nature Reviews Microbiology . 5, No. 11, 2007, pp. 892-899. doi : 10.1038 / nrmicro1767 . This article at Uni Osnabrück: Perspectives (PDF)  ( page no longer available , search in web archivesInfo: The link was automatically marked as defective. Please check the link according to the instructions and then remove this notice.@1@ 2Template: Toter Link / www.macromol.uni-osnabrueck.de  
  38. arms Y Mulkidjanian, Michael Y Galperin, Eugene V Koonin: Co-evolution of primordial membranes and membrane proteins . In: Trends Biochem Sci. . 4, No. 34, 2009, pp. 206-215. doi : 10.1016 / j.tibs.2009.01.005 . PMC 2752816 (free full text).  ( Page no longer available , search in web archivesInfo: The link was automatically marked as defective. Please check the link according to the instructions and then remove this notice.@1@ 2Template: Toter Link / www.macromol.uni-osnabrueck.de  
  39. Justin Chen: New Discovery Suggests All Life May Share This Common Design Principle , on SciTechDaily March 30, 2018
  40. Jean-Benoît Lalanne, James C. Taggart, Monica S. Guo, Lydia Herzel, Ariel Schieler, Gene-Wei Li: Evolutionary Convergence of Pathway-Specific Enzyme Expression Stoichiometry , Cell, Volume 173, No. 3, April 19, 2018 (online March 29, 2018), pp. 749-761.e38, doi: 10.1016 / j.cell.2018.03.007
  41. M. Krupovic et al .: Multiple origins of viral capsid proteins from cellular ancestors , PNAS 114 (12) of February 3, 2017, doi: 10.1073 / pnas.1621061114 , sections Significance and Abstract
  42. A. Nasir et al .: A phylogenomic data-driven exploration of viral origins and evolution , Science Advances AAAS, Vol. 1 (8), September 25, 2015, doi: 10.1126 / sciadv.1500527
  43. Eugene V. Koonin: The origin and early evolution of eukaryotes in the light of phylogenomics , in: BioMed Central: Genome Biology 201011: 209 of May 5, 2010, doi: 10.1186 / gb-2010-11-5-209
  44. Brigitte Boxma, Rob M. de Graaf, Georg WM van der Staay, Theo A. van Alen, Guenola Ricard, Toni Gabaldon, Angela HAM van Hoek, Seung Yeo Moon-van der Staay, Werner JH Koopman, Jaap J. van Hellemond , Aloysius GM Tielens, Thorsten Friedrich, Marten Veenhuis, Martijn A. Huynen, Johannes HP Hackstein: An anaerobic mitochondrion that produces hydrogen . In: Nature . tape 434 , no. 7029 , February 3, 2005, p. 74-79 , doi : 10.1038 / nature03343 .
  45. A. Akhmanove et al .: A hydrogenosomes with a genome . In: Nature . tape 396 , no. 6711 , December 10, 1998, p. 527-528 , doi : 10.1038 / 25023 .
  46. CR Woese, GE Fox: Phylogenetic structure of the prokaryotic domain: the primary kingdoms. In: Proceedings of the National Academy of Sciences . Volume 74, Number 11, November 1977, pp. 5088-5090, PMID 270744 , PMC 432104 (free full text).
  47. Tom Cavalier-Smith: Archaebacteria and Archezoa . In: Nature . 339, No. 6220, May 1989, pp. 100-101. doi : 10.1038 / 339100a0 . PMID 2497352 .
  48. Patrick J. Keeling: A kingdom's progress: Archezoa and the origin of eukaryotes. In: BioEssays. Volume 20, number 1, January 1998, pp. 87–95, doi : 10.1002 / (SICI) 1521-1878 (199801) 20: 1 <87 :: AID-BIES12> 3.0.CO; 2-4 (alternative full text access : SemanticScholar ).
  49. ^ Martin W and Müller M: The hydrogen hypothesis for the first eukaryote . In: Nature . 392, No. 6671, 1998, pp. 37-41. doi : 10.1038 / 32096 . PMID 9510246 .
  50. Marta Kwapisz, Frédéric Beckouët, Pierre Thuriaux: Early evolution of eukaryotic DNA-dependent RNA polymerases. In: Trends in Genetics. 24, 2008, p. 211, doi: 10.1016 / j.tig.2008.02.002 , 3rd illustration.
  51. S. Hou, KS Makarova, et al .: Complete genome sequence of the extremely acidophilic methanotroph isolate V4, Methylacidiphilum infernorum, a representative of the bacterial phylum Verrucomicrobia. In: Biology direct. Volume 3, July 2008, p. 26, doi: 10.1186 / 1745-6150-3-26 , PMID 18593465 , PMC 2474590 (free full text).