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{{evolutionary biology}}
{{evolutionary biology}}


The '''last universal common ancestor''' ('''LUCA'''), also called the '''last universal ancestor''' ('''LUA'''), '''cenancestor''', or (incorrectly<ref group=R>There is a common misconception that definitions of LUCA and progenote are the same; however, progenote is defined as an organism “still in the process of evolving the relationship between genotype and phenotype”, and it is only hypothesed that LUCA is a progenote.<br/>{{cite journal|last1=Gogarten|first1=Johann Peter|last2=Deamer|first2=David|title=Is LUCA a thermophilic progenote?|journal=Nature Microbiology|volume=1|date=25 November 2016|page=16229|doi=10.1038/nmicrobiol.2016.229}}</ref>) '''progenote''', is the most recent population of organisms from which all organisms now [[Life|living on Earth]] have a [[common descent]].<ref name="theobald">{{cite journal |author=Theobald DL |title=A formal test of the theory of universal common ancestry |journal=Nature |volume=465 |issue=7295 |pages=219–22 |date=May 2010 |pmid=20463738 |doi=10.1038/nature09014 |bibcode=2010Natur.465..219T}}</ref> LUCA is the [[most recent common ancestor]] of all current life on Earth. LUCA is not thought to be the [[first living organism on Earth]], but only one of many early organisms, whereas the others became extinct.
The '''last universal common ancestor''' ('''LUCA'''), also called the '''last universal ancestor''' ('''LUA'''), '''cenancestor''', or (incorrectly<ref group=R>There is a common misconception that definitions of LUCA and progenote are the same; however, progenote is defined as an organism “still in the process of evolving the relationship between genotype and phenotype”, and it is only hypothesed that LUCA is a progenote.<br/>{{cite journal|last1=Gogarten|first1=Johann Peter|last2=Deamer|first2=David|title=Is LUCA a thermophilic progenote?|journal=Nature Microbiology|volume=1|issue=12|date=25 November 2016|page=16229|doi=10.1038/nmicrobiol.2016.229|pmid=27886195}}</ref>) '''progenote''', is the most recent population of organisms from which all organisms now [[Life|living on Earth]] have a [[common descent]].<ref name="theobald">{{cite journal |author=Theobald DL |title=A formal test of the theory of universal common ancestry |journal=Nature |volume=465 |issue=7295 |pages=219–22 |date=May 2010 |pmid=20463738 |doi=10.1038/nature09014 |bibcode=2010Natur.465..219T}}</ref> LUCA is the [[most recent common ancestor]] of all current life on Earth. LUCA is not thought to be the [[first living organism on Earth]], but only one of many early organisms, whereas the others became extinct.


While there is no specific fossil evidence of LUCA, it can be studied by [[phylogenetic bracketing|comparing the genomes of its descendants]], all organisms known to be living today. By this means, a 2016 study identified a set of 355 [[gene]]s inferred to have been present in LUCA. This would imply it was already a complex life form with many [[co-adapted]] features, including [[transcription (biology)|transcription]] and [[translation (biology)|translation]] mechanisms to convert information between [[DNA]], [[RNA]], and [[protein]]s.<ref name="nature" /><ref name="NYT-20160725" /> However, some of those genes could have developed later and spread universally by [[horizontal gene transfer]] between [[archaea]] and [[bacteria]].<ref name="GogartenDeamer2016" />
While there is no specific fossil evidence of LUCA, it can be studied by [[phylogenetic bracketing|comparing the genomes of its descendants]], all organisms known to be living today. By this means, a 2016 study identified a set of 355 [[gene]]s inferred to have been present in LUCA. This would imply it was already a complex life form with many [[co-adapted]] features, including [[transcription (biology)|transcription]] and [[translation (biology)|translation]] mechanisms to convert information between [[DNA]], [[RNA]], and [[protein]]s.<ref name="nature" /><ref name="NYT-20160725" /> However, some of those genes could have developed later and spread universally by [[horizontal gene transfer]] between [[archaea]] and [[bacteria]].<ref name="GogartenDeamer2016" />


LUCA is estimated to have lived some 3.5 to 3.8 billion years ago in the [[Paleoarchean]] [[era]],<ref>{{cite journal|author=Doolittle WF|date=February 2000|title=Uprooting the tree of life|journal=Scientific American|volume=282|issue=2|pages=90–95|bibcode=2000SciAm.282b..90D|doi=10.1038/scientificamerican0200-90|pmid=10710791}}</ref><ref>{{cite journal|vauthors=Glansdorff N, Xu Y, Labedan B|year=2008|title=The last universal common ancestor: emergence, constitution and genetic legacy of an elusive forerunner|journal=Biology Direct|volume=3|issue=|page=29|doi=10.1186/1745-6150-3-29|pmc=2478661|pmid=18613974}}</ref> a few hundred million years after the [[Earliest known life forms|earliest evidence of life on Earth]], for which there are several candidates. [[Microbial mat]] [[fossils]] have been found in 3.48 billion-year-old [[sandstone]] from [[Western Australia]],<ref name="AP-20131113">{{cite news|url=http://apnews.excite.com/article/20131113/DAA1VSC01.html|title=Oldest fossil found: Meet your microbial mom|last=Borenstein|first=Seth|date=13 November 2013|accessdate=15 November 2013|agency=Associated Press}}</ref><ref name="AST-20131108">{{cite journal|author=Noffke, N.|author2=Christian, D.|author3=Wacey, D.|author4=Hazen, R.M.|date=December 2013|title=Microbially induced sedimentary structures recording an ancient ecosystem in the ca. 3.48 billion-year-old Dresser Formation, Pilbara, Western Australia|journal=Astrobiology|volume=13|issue=12|pages=1103–24|bibcode=2013AsBio..13.1103N|doi=10.1089/ast.2013.1030|pmc=3870916|pmid=24205812}}</ref> while [[Biogenic substance|biogenic]] [[graphite]] has been found in 3.7 billion-year-old [[Metasediment|metamorphized sedimentary rocks]] from [[Western Greenland]].<ref name="NG-20131208">{{cite journal|last1=Ohtomo|first1=Yoko|last2=Kakegawa|first2=Takeshi|last3=Ishida|first3=Akizumi|last4=Nagase|first4=Toshiro|last5=Rosing|first5=Minik T.|year=2013|title=Evidence for biogenic graphite in early Archaean Isua metasedimentary rocks|journal=Nature Geoscience|volume=7|pages=25–28|bibcode=2014NatGe...7...25O|doi=10.1038/ngeo2025}}</ref><ref>{{Cite journal|last=Hassenkam|first=T.|last2=Andersson|first2=M. P.|last3=Dalby|first3=K. N.|last4=Mackenzie|first4=D. M. A.|last5=Rosing|first5=M. T.|year=2017|title=Elements of Eoarchean life trapped in mineral inclusions|url=http://www.nature.com/doifinder/10.1038/nature23261|journal=Nature|volume=548|issue=7665|pages=78–81|bibcode=2017Natur.548...78H|doi=10.1038/nature23261}}</ref> Recent studies have tentatively proposed evidence of life as early as 4.28 billion years ago.<ref name="AP-20151019">{{cite news |last=Borenstein |first=Seth |title=Hints of life on what was thought to be desolate early Earth |url=https://apnews.com/e6be2537b4cd46ffb9c0585bae2b2e51 |date=19 October 2015 |work=AP News |publisher=[[Associated Press]] |accessdate=9 October 2018}}</ref><ref>{{Cite journal|last=Bell|first=Elizabeth A.|last2=Boehnke|first2=Patrick|last3=Harrison|first3=T. Mark|last4=Mao|first4=Wendy L.|date=24 November 2015|title=Potentially biogenic carbon preserved in a 4.1 billion-year-old zircon|journal=Proceedings of the National Academy of Sciences of the United States of America|volume=112|issue=47|pages=14518–21|bibcode=2015PNAS..11214518B|doi=10.1073/pnas.1517557112|issn=1091-6490|pmc=4664351|pmid=26483481}}</ref><ref>{{cite journal|author=Dodd, Matthew S.|author2=Papineau, Dominic|author3=Grenne, Tor|author4=slack, John F.|author5=Rittner, Martin|author6=Pirajno, Franco|author7=O'Neil, Jonathan|author8=Little, Crispin T. S.|date=2 March 2017|title=Evidence for early life in Earth's oldest hydrothermal vent precipitates|url=https://www.nature.com/articles/nature21377|journal=Nature|volume=543|pages=60–64|bibcode=2017Natur.543...60D|doi=10.1038/nature21377|archiveurl=http://eprints.whiterose.ac.uk/112179/|archivedate=10 February 2017}}</ref>
LUCA is estimated to have lived some 3.5 to 3.8 billion years ago in the [[Paleoarchean]] [[era]],<ref>{{cite journal|author=Doolittle WF|date=February 2000|title=Uprooting the tree of life|journal=Scientific American|volume=282|issue=2|pages=90–95|bibcode=2000SciAm.282b..90D|doi=10.1038/scientificamerican0200-90|pmid=10710791}}</ref><ref>{{cite journal|vauthors=Glansdorff N, Xu Y, Labedan B|year=2008|title=The last universal common ancestor: emergence, constitution and genetic legacy of an elusive forerunner|journal=Biology Direct|volume=3|issue=|page=29|doi=10.1186/1745-6150-3-29|pmc=2478661|pmid=18613974}}</ref> a few hundred million years after the [[Earliest known life forms|earliest evidence of life on Earth]], for which there are several candidates. [[Microbial mat]] [[fossils]] have been found in 3.48 billion-year-old [[sandstone]] from [[Western Australia]],<ref name="AP-20131113">{{cite news|url=http://apnews.excite.com/article/20131113/DAA1VSC01.html|title=Oldest fossil found: Meet your microbial mom|last=Borenstein|first=Seth|date=13 November 2013|accessdate=15 November 2013|agency=Associated Press}}</ref><ref name="AST-20131108">{{cite journal|author=Noffke, N.|author2=Christian, D.|author3=Wacey, D.|author4=Hazen, R.M.|date=December 2013|title=Microbially induced sedimentary structures recording an ancient ecosystem in the ca. 3.48 billion-year-old Dresser Formation, Pilbara, Western Australia|journal=Astrobiology|volume=13|issue=12|pages=1103–24|bibcode=2013AsBio..13.1103N|doi=10.1089/ast.2013.1030|pmc=3870916|pmid=24205812}}</ref> while [[Biogenic substance|biogenic]] [[graphite]] has been found in 3.7 billion-year-old [[Metasediment|metamorphized sedimentary rocks]] from [[Western Greenland]].<ref name="NG-20131208">{{cite journal|last1=Ohtomo|first1=Yoko|last2=Kakegawa|first2=Takeshi|last3=Ishida|first3=Akizumi|last4=Nagase|first4=Toshiro|last5=Rosing|first5=Minik T.|year=2013|title=Evidence for biogenic graphite in early Archaean Isua metasedimentary rocks|journal=Nature Geoscience|volume=7|issue=1|pages=25–28|bibcode=2014NatGe...7...25O|doi=10.1038/ngeo2025}}</ref><ref>{{Cite journal|last=Hassenkam|first=T.|last2=Andersson|first2=M. P.|last3=Dalby|first3=K. N.|last4=Mackenzie|first4=D. M. A.|last5=Rosing|first5=M. T.|year=2017|title=Elements of Eoarchean life trapped in mineral inclusions|journal=Nature|volume=548|issue=7665|pages=78–81|bibcode=2017Natur.548...78H|doi=10.1038/nature23261|pmid=28738409|url=http://orbit.dtu.dk/en/publications/elements-of-eoarchean-life-trapped-in-mineral-inclusions(9f420096-58b8-43a9-8fc1-092b0b1f549c).html}}</ref> Recent studies have tentatively proposed evidence of life as early as 4.28 billion years ago.<ref name="AP-20151019">{{cite news |last=Borenstein |first=Seth |title=Hints of life on what was thought to be desolate early Earth |url=https://apnews.com/e6be2537b4cd46ffb9c0585bae2b2e51 |date=19 October 2015 |work=AP News |publisher=[[Associated Press]] |accessdate=9 October 2018}}</ref><ref>{{Cite journal|last=Bell|first=Elizabeth A.|last2=Boehnke|first2=Patrick|last3=Harrison|first3=T. Mark|last4=Mao|first4=Wendy L.|date=24 November 2015|title=Potentially biogenic carbon preserved in a 4.1 billion-year-old zircon|journal=Proceedings of the National Academy of Sciences of the United States of America|volume=112|issue=47|pages=14518–21|bibcode=2015PNAS..11214518B|doi=10.1073/pnas.1517557112|issn=1091-6490|pmc=4664351|pmid=26483481}}</ref><ref>{{cite journal|author=Dodd, Matthew S.|author2=Papineau, Dominic|author3=Grenne, Tor|author4=slack, John F.|author5=Rittner, Martin|author6=Pirajno, Franco|author7=O'Neil, Jonathan|author8=Little, Crispin T. S.|date=2 March 2017|title=Evidence for early life in Earth's oldest hydrothermal vent precipitates|url=https://www.nature.com/articles/nature21377|journal=Nature|volume=543|issue=7643|pages=60–64|bibcode=2017Natur.543...60D|doi=10.1038/nature21377|pmid=28252057|archiveurl=http://eprints.whiterose.ac.uk/112179/|archivedate=10 February 2017}}</ref>


[[Charles Darwin]] proposed the theory of universal common descent through an [[evolution]]ary process in his book ''[[On the Origin of Species]]'' in 1859, saying, "Therefore I should infer from analogy that probably all the organic beings which have ever lived on this earth have descended from some one primordial form, into which life was first breathed."<ref name="origin">{{Citation |last=Darwin |first=C. |year=1859 |title=[[The Origin of Species by Means of Natural Selection]] |page=490 |publisher=[[John Murray (publisher)|John Murray]]}}</ref> Later biologists have separated the problem of the [[origin of life]] from that of the LUCA.
[[Charles Darwin]] proposed the theory of universal common descent through an [[evolution]]ary process in his book ''[[On the Origin of Species]]'' in 1859, saying, "Therefore I should infer from analogy that probably all the organic beings which have ever lived on this earth have descended from some one primordial form, into which life was first breathed."<ref name="origin">{{Citation |last=Darwin |first=C. |year=1859 |title=The Origin of Species by Means of Natural Selection |page=490 |publisher=[[John Murray (publisher)|John Murray]]|title-link=The Origin of Species by Means of Natural Selection }}</ref> Later biologists have separated the problem of the [[origin of life]] from that of the LUCA.


==Features==
==Features==
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With the later gene pool of the LUCA's descendants, with their common framework of the [[Chargaff's rules|AT/GC]] rule and the standard twenty amino acids, horizontal gene transfer would have been feasible and could have been very common.
With the later gene pool of the LUCA's descendants, with their common framework of the [[Chargaff's rules|AT/GC]] rule and the standard twenty amino acids, horizontal gene transfer would have been feasible and could have been very common.


In an earlier hypothesis, [[Carl Woese]] (1988) had proposed that: (1) no individual organism can be considered a LUCA, and (2) the genetic heritage of all modern organisms derived through [[horizontal gene transfer]] among an ancient community of organisms.<ref>{{cite journal |author=Woese C |title=The universal ancestor |journal=Proceedings of the National Academy of Sciences of the United States of America |volume=95 |issue=12 |pages=6854–59 |date=June 1998 |pmid=9618502 |pmc=22660 |doi=10.1073/pnas.95.12.6854 |bibcode=1998PNAS...95.6854W}}</ref> While the results described by Theobald (2010) and Saey (2010) demonstrate the existence of a single LUCA, Woese's argument can still be applied to Ur-organisms (initial products of abiogenesis) before the LUCA. At the beginnings of life, ancestry was not as linear as it is today because the genetic code had not evolved.<ref name="majortransinevolution">{{cite book | title=The Major Transitions in Evolution | last=Maynard Smith | first=John | authorlink=John Maynard Smith |first2=Eörs |last2=Szathmáry |author2link=Eörs Szathmáry | publisher=Oxford University Press | year=1995 | location=Oxford, England | isbn=0-19-850294-X |url=https://books.google.com/books?id=UGCmIVB5dhMC}}{{page needed|date=June 2014}}</ref> Before high fidelity replication, organisms could not be easily mapped on a phylogenetic tree. However, the LUCA lived after the origin of the genetic code and at least some rudimentary early form of molecular proofreading.
In an earlier hypothesis, [[Carl Woese]] (1988) had proposed that: (1) no individual organism can be considered a LUCA, and (2) the genetic heritage of all modern organisms derived through [[horizontal gene transfer]] among an ancient community of organisms.<ref>{{cite journal |author=Woese C |title=The universal ancestor |journal=Proceedings of the National Academy of Sciences of the United States of America |volume=95 |issue=12 |pages=6854–59 |date=June 1998 |pmid=9618502 |pmc=22660 |doi=10.1073/pnas.95.12.6854 |bibcode=1998PNAS...95.6854W}}</ref> While the results described by Theobald (2010) and Saey (2010) demonstrate the existence of a single LUCA, Woese's argument can still be applied to Ur-organisms (initial products of abiogenesis) before the LUCA. At the beginnings of life, ancestry was not as linear as it is today because the genetic code had not evolved.<ref name="majortransinevolution">{{cite book | title=The Major Transitions in Evolution | last=Maynard Smith | first=John | authorlink=John Maynard Smith |first2=Eörs |last2=Szathmáry |author2link=Eörs Szathmáry | publisher=Oxford University Press | year=1995 | location=Oxford, England | isbn=978-0-19-850294-4 |url=https://books.google.com/books?id=UGCmIVB5dhMC}}{{page needed|date=June 2014}}</ref> Before high fidelity replication, organisms could not be easily mapped on a phylogenetic tree. However, the LUCA lived after the origin of the genetic code and at least some rudimentary early form of molecular proofreading.


==Location of the root==
==Location of the root==
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{{for|branching of Bacteria phyla|Bacterial phyla}}
{{for|branching of Bacteria phyla|Bacterial phyla}}


The most commonly accepted [[Tree of life (biology)|tree of life]], based on several molecular studies, has its root between a [[monophyletic]] [[domain (biology)|domain]] [[Bacteria]] and a [[clade]] formed by [[Archaea]] and [[Eukaryota]].<ref>Brown, J. R., and W. F. Doolittle. 1995. “Root of the Universal Tree of Life Based on Ancient Aminoacyl-tRNA Synthetase Gene Duplications.” Proc Natl Acad Sci USA 92 (7): 2441–45. [https://www.ncbi.nlm.nih.gov/pubmed/?term=7708661 PMID 7708661]</ref><ref>Gogarten, J. P., H. Kibak, P. Dittrich, L. Taiz, E. J. Bowman, B. J. Bowman, M. F. Manolson, et al. 1989. “Evolution of the Vacuolar H+-ATPase: Implications for the Origin of Eukaryotes.” Proc Natl Acad Sci USA 86 (17): 6661–65. [https://www.ncbi.nlm.nih.gov/pubmed/?term=2528146 PMID 2528146]</ref><ref>{{cite journal | last1 = Gogarten | first1 = J.P. | last2 = Taiz | first2 = L. | year = 1992 | title = Evolution of Proton Pumping ATPases: Rooting the Tree of Life | url = https://link.springer.com/article/10.1007%2FBF00039176 | journal = Photosynthesis Research | volume = 33 | issue = | pages = 137–46 | doi = 10.1007/BF00039176 }}</ref><ref>Gribaldo, S, and P Cammarano. 1998. “The Root of the Universal Tree of Life Inferred from Anciently Duplicated Genes Encoding Components of the Protein-Targeting Machinery.” Journal of Molecular Evolution 47 (5): 508–16. PMID: [https://www.ncbi.nlm.nih.gov/pubmed/?term=9797401 9797401]</ref><ref>Iwabe, Naoyuki, Kei-Ichi Kuma, Masami Hasegawa, Syozo Osawa, Takashi Miyata Source, Masami Hasegawat, Syozo Osawat, and Takashi Miyata. 1989. “Evolutionary Relationship of Archaebacteria, Eubacteria, and Eukaryotes Inferred from Phylogenetic Trees of Duplicated Genes.” Proc Natl Acad Sci USA 86 (86): 9355–59. [https://www.ncbi.nlm.nih.gov/pubmed/?term=2531898 PMID 2531898]</ref><ref>{{cite book |editor1-first=David R. |editor1-last=Boone |editor2-first=Richard W. |editor2-last=Castenholz |editor3-first=George M. |editor3-last=Garrity |title=The ''Archaea'' and the Deeply Branching and Phototrophic ''Bacteria'' |series=Bergey's Manual of Systematic Bacteriology |isbn=978-0-387-21609-6 |url=https://www.springer.com/life+sciences/microbiology/book/978-0-387-98771-2 |doi=10.1007/978-0-387-21609-6}}{{page needed|date=June 2014}}</ref> However, a very small minority of studies place the root in the domain Bacteria, either in the phylum [[Firmicutes]]<ref>{{cite journal |author1=Valas, R.E. |author2=Bourne, P.E. |title=The origin of a derived superkingdom: how a gram-positive bacterium crossed the desert to become an archaeon |journal=Biology Direct |volume=6 |issue= |page=16 |year=2011 |pmid=21356104 |pmc=3056875 |doi=10.1186/1745-6150-6-16}}</ref> or state that the phylum [[Chloroflexi (phylum)|Chloroflexi]] is [[Basal (phylogenetics)|basal]] to a clade with Archaea and Eukaryotes and the rest of Bacteria (as proposed by [[Thomas Cavalier-Smith]]).<ref name=CS2>{{cite journal |author=Cavalier-Smith T |title=Rooting the tree of life by transition analyses |journal=Biology Direct |volume=1 |page=19 |year=2006 |pmid=16834776 |pmc=1586193 |doi=10.1186/1745-6150-1-19}}</ref>
The most commonly accepted [[Tree of life (biology)|tree of life]], based on several molecular studies, has its root between a [[monophyletic]] [[domain (biology)|domain]] [[Bacteria]] and a [[clade]] formed by [[Archaea]] and [[Eukaryota]].<ref>Brown, J. R., and W. F. Doolittle. 1995. “Root of the Universal Tree of Life Based on Ancient Aminoacyl-tRNA Synthetase Gene Duplications.” Proc Natl Acad Sci USA 92 (7): 2441–45. [https://www.ncbi.nlm.nih.gov/pubmed/?term=7708661 PMID 7708661]</ref><ref>Gogarten, J. P., H. Kibak, P. Dittrich, L. Taiz, E. J. Bowman, B. J. Bowman, M. F. Manolson, et al. 1989. “Evolution of the Vacuolar H+-ATPase: Implications for the Origin of Eukaryotes.” Proc Natl Acad Sci USA 86 (17): 6661–65. [https://www.ncbi.nlm.nih.gov/pubmed/?term=2528146 PMID 2528146]</ref><ref>{{cite journal | last1 = Gogarten | first1 = J.P. | last2 = Taiz | first2 = L. | year = 1992 | title = Evolution of Proton Pumping ATPases: Rooting the Tree of Life | journal = Photosynthesis Research | volume = 33 | issue = 2| pages = 137–46 | doi = 10.1007/BF00039176 | pmid = 24408574 }}</ref><ref>Gribaldo, S, and P Cammarano. 1998. “The Root of the Universal Tree of Life Inferred from Anciently Duplicated Genes Encoding Components of the Protein-Targeting Machinery.” Journal of Molecular Evolution 47 (5): 508–16. PMID: [https://www.ncbi.nlm.nih.gov/pubmed/?term=9797401 9797401]</ref><ref>Iwabe, Naoyuki, Kei-Ichi Kuma, Masami Hasegawa, Syozo Osawa, Takashi Miyata Source, Masami Hasegawat, Syozo Osawat, and Takashi Miyata. 1989. “Evolutionary Relationship of Archaebacteria, Eubacteria, and Eukaryotes Inferred from Phylogenetic Trees of Duplicated Genes.” Proc Natl Acad Sci USA 86 (86): 9355–59. [https://www.ncbi.nlm.nih.gov/pubmed/?term=2531898 PMID 2531898]</ref><ref>{{cite book |editor1-first=David R. |editor1-last=Boone |editor2-first=Richard W. |editor2-last=Castenholz |editor3-first=George M. |editor3-last=Garrity |title=The ''Archaea'' and the Deeply Branching and Phototrophic ''Bacteria'' |series=Bergey's Manual of Systematic Bacteriology |isbn=978-0-387-21609-6 |url=https://www.springer.com/life+sciences/microbiology/book/978-0-387-98771-2 |doi=10.1007/978-0-387-21609-6|publisher=Springer |year=2001 }}{{page needed|date=June 2014}}</ref> However, a very small minority of studies place the root in the domain Bacteria, either in the phylum [[Firmicutes]]<ref>{{cite journal |author1=Valas, R.E. |author2=Bourne, P.E. |title=The origin of a derived superkingdom: how a gram-positive bacterium crossed the desert to become an archaeon |journal=Biology Direct |volume=6 |issue= |page=16 |year=2011 |pmid=21356104 |pmc=3056875 |doi=10.1186/1745-6150-6-16}}</ref> or state that the phylum [[Chloroflexi (phylum)|Chloroflexi]] is [[Basal (phylogenetics)|basal]] to a clade with Archaea and Eukaryotes and the rest of Bacteria (as proposed by [[Thomas Cavalier-Smith]]).<ref name=CS2>{{cite journal |author=Cavalier-Smith T |title=Rooting the tree of life by transition analyses |journal=Biology Direct |volume=1 |page=19 |year=2006 |pmid=16834776 |pmc=1586193 |doi=10.1186/1745-6150-1-19}}</ref>


Ground-breaking research by [[William F. Martin]] (2016) genetically analyzed 6.1 million protein-coding genes and 286,514 protein clusters from sequenced prokaryotic genomes of various phylogenetic trees, and identified 355 protein clusters that were probably common to the LUCA. The results "depict LUCA as [[anaerobic organism|anaerobic]], CO<sub>2</sub>-fixing, H<sub>2</sub>-dependent with a [[Wood–Ljungdahl pathway]] (the reductive [[Acetyl-CoA|acetyl-coenzyme A]] pathway), N<sub>2</sub>-fixing and thermophilic. LUCA's biochemistry was replete with FeS clusters and radical reaction mechanisms." The [[Cofactor (biochemistry)|cofactors]] also reveal "dependence upon [[transition metal]]s, [[Flavin mononucleotide|flavins]], [[S-adenosyl methionine]], [[coenzyme A]], [[ferredoxin]], [[molybdopterin]], [[corrin]]s and [[selenium]]. Its genetic code required [[nucleoside]] modifications and S-adenosylmethionine-dependent [[methylation]]s."<ref name="nature">"[http://www.nature.com/articles/nmicrobiol2016116 The physiology and habitat of the last universal common ancestor]" by Madeline C. Weiss, FilipaL.Sousa, Natalia Mrnjavac, Sinje Neukirchen, Mayo Roettger, Shijulal Nelson-Sathi and William F. Martin (July 25, 2016) (Nature Microbiology 16116 | {{Doi|10.1038/NMICROBIOL.2016.116}}</ref><ref name="NLane_Funke">Nick Lane: [https://books.google.de/books?id=IfJYBQAAQBAJ&pg=PT77&lpg=PT77&dq=nick+lane+the+vital+question&source=bl&ots=wZ1vmiXh7z&sig=DAn9E_5fw8XGQ8nmsGOm1_R4VlI&hl=de&sa=X&ved=0ahUKEwjKnLWhtuPVAhWLPxQKHXK3B1sQ6AEIIDAA#v=onepage&q=alkaline%20hydrothermal%20vents The Vital Question – Energy, Evolution, and the Origins of Complex Life], WW Norton, 2015, {{ISBN|978-0-393-08881-6}}</ref><ref>Joseph F. Sutherland: [http://prehistoricict.blogspot.de/2014/08/on-origin-of-bacteria-and-archaea.html on The Origin Of Tha Bacteria And The Archaea], auf B.C vom 16. August 2014</ref> The results are "quite specific":<ref name="NYT-20160725">{{cite news |last=Wade |first=Nicholas |title=Meet Luca, the Ancestor of All Living Things |url=https://www.nytimes.com/2016/07/26/science/last-universal-ancestor.html |date=25 July 2016 |work=[[The New York Times]] |accessdate=8 October 2018 }}</ref> they show that [[methanogen]]ic [[clostridium|clostridia]] was a basal clade in the 355 lineages examined, and that the LUCA may therefore have inhabited an anaerobic [[hydrothermal vent]] setting in a geochemically active environment rich in H<sub>2</sub>, CO<sub>2</sub>, and iron.<ref name="nature" />
Ground-breaking research by [[William F. Martin]] (2016) genetically analyzed 6.1 million protein-coding genes and 286,514 protein clusters from sequenced prokaryotic genomes of various phylogenetic trees, and identified 355 protein clusters that were probably common to the LUCA. The results "depict LUCA as [[anaerobic organism|anaerobic]], CO<sub>2</sub>-fixing, H<sub>2</sub>-dependent with a [[Wood–Ljungdahl pathway]] (the reductive [[Acetyl-CoA|acetyl-coenzyme A]] pathway), N<sub>2</sub>-fixing and thermophilic. LUCA's biochemistry was replete with FeS clusters and radical reaction mechanisms." The [[Cofactor (biochemistry)|cofactors]] also reveal "dependence upon [[transition metal]]s, [[Flavin mononucleotide|flavins]], [[S-adenosyl methionine]], [[coenzyme A]], [[ferredoxin]], [[molybdopterin]], [[corrin]]s and [[selenium]]. Its genetic code required [[nucleoside]] modifications and S-adenosylmethionine-dependent [[methylation]]s."<ref name="nature">"[http://www.nature.com/articles/nmicrobiol2016116 The physiology and habitat of the last universal common ancestor]" by Madeline C. Weiss, FilipaL.Sousa, Natalia Mrnjavac, Sinje Neukirchen, Mayo Roettger, Shijulal Nelson-Sathi and William F. Martin (July 25, 2016) (Nature Microbiology 16116 | {{Doi|10.1038/NMICROBIOL.2016.116}}</ref><ref name="NLane_Funke">Nick Lane: [https://books.google.de/books?id=IfJYBQAAQBAJ&pg=PT77&lpg=PT77&dq=nick+lane+the+vital+question&source=bl&ots=wZ1vmiXh7z&sig=DAn9E_5fw8XGQ8nmsGOm1_R4VlI&hl=de&sa=X&ved=0ahUKEwjKnLWhtuPVAhWLPxQKHXK3B1sQ6AEIIDAA#v=onepage&q=alkaline%20hydrothermal%20vents The Vital Question – Energy, Evolution, and the Origins of Complex Life], WW Norton, 2015, {{ISBN|978-0-393-08881-6}}</ref><ref>Joseph F. Sutherland: [http://prehistoricict.blogspot.de/2014/08/on-origin-of-bacteria-and-archaea.html on The Origin Of Tha Bacteria And The Archaea], auf B.C vom 16. August 2014</ref> The results are "quite specific":<ref name="NYT-20160725">{{cite news |last=Wade |first=Nicholas |title=Meet Luca, the Ancestor of All Living Things |url=https://www.nytimes.com/2016/07/26/science/last-universal-ancestor.html |date=25 July 2016 |work=[[The New York Times]] |accessdate=8 October 2018 }}</ref> they show that [[methanogen]]ic [[clostridium|clostridia]] was a basal clade in the 355 lineages examined, and that the LUCA may therefore have inhabited an anaerobic [[hydrothermal vent]] setting in a geochemically active environment rich in H<sub>2</sub>, CO<sub>2</sub>, and iron.<ref name="nature" />


These findings could mean that life on Earth originated in such hydrothermal vents, but it is also possible that life was restricted to such locations at some later time, perhaps by the [[Late Heavy Bombardment]].<ref name="NYT-20160725" /> The identification of these genes as being present in LUCA has also been criticized, as they may simply represent later genes which migrated via [[horizontal gene transfer]]s between archaea and bacteria.<ref name="GogartenDeamer2016">{{cite journal |last1=Gogarten |first1=Johann Peter |last2=Deamer |first2=David |title=Is LUCA a thermophilic progenote? |journal=Nature Microbiology |volume=1 |year=2016 |pages=16229 |doi=10.1038/nmicrobiol.2016.229}}</ref>
These findings could mean that life on Earth originated in such hydrothermal vents, but it is also possible that life was restricted to such locations at some later time, perhaps by the [[Late Heavy Bombardment]].<ref name="NYT-20160725" /> The identification of these genes as being present in LUCA has also been criticized, as they may simply represent later genes which migrated via [[horizontal gene transfer]]s between archaea and bacteria.<ref name="GogartenDeamer2016">{{cite journal |last1=Gogarten |first1=Johann Peter |last2=Deamer |first2=David |title=Is LUCA a thermophilic progenote? |journal=Nature Microbiology |volume=1 |issue=12 |year=2016 |pages=16229 |doi=10.1038/nmicrobiol.2016.229|pmid=27886195 }}</ref>


==See also==
==See also==

Revision as of 11:14, 18 January 2019

For lowest common ancestors in graph theory and computer science, see lowest common ancestor.
"LUCA" redirects here. For other uses, see Luca (disambiguation).
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The last universal common ancestor (LUCA), also called the last universal ancestor (LUA), cenancestor, or (incorrectly[R 1]) progenote, is the most recent population of organisms from which all organisms now living on Earth have a common descent.[1] LUCA is the most recent common ancestor of all current life on Earth. LUCA is not thought to be the first living organism on Earth, but only one of many early organisms, whereas the others became extinct.

While there is no specific fossil evidence of LUCA, it can be studied by comparing the genomes of its descendants, all organisms known to be living today. By this means, a 2016 study identified a set of 355 genes inferred to have been present in LUCA. This would imply it was already a complex life form with many co-adapted features, including transcription and translation mechanisms to convert information between DNA, RNA, and proteins.[2][3] However, some of those genes could have developed later and spread universally by horizontal gene transfer between archaea and bacteria.[4]

LUCA is estimated to have lived some 3.5 to 3.8 billion years ago in the Paleoarchean era,[5][6] a few hundred million years after the earliest evidence of life on Earth, for which there are several candidates. Microbial mat fossils have been found in 3.48 billion-year-old sandstone from Western Australia,[7][8] while biogenic graphite has been found in 3.7 billion-year-old metamorphized sedimentary rocks from Western Greenland.[9][10] Recent studies have tentatively proposed evidence of life as early as 4.28 billion years ago.[11][12][13]

Charles Darwin proposed the theory of universal common descent through an evolutionary process in his book On the Origin of Species in 1859, saying, "Therefore I should infer from analogy that probably all the organic beings which have ever lived on this earth have descended from some one primordial form, into which life was first breathed."[14] Later biologists have separated the problem of the origin of life from that of the LUCA.

Features

Further information: Phylogenetic bracketing

By analysis of the presumed LUCA's offspring groups, the LUCA appears to have been a small, single-celled organism. It likely had a ring-shaped coil of DNA floating freely within the cell, like modern bacteria. Morphologically, it would likely not have stood out within a mixed population of small modern-day bacteria. However, Carl Woese et al., who first proposed the currently-used three domain system based on an analysis of ribosomal RNA (rRNA) sequences of bacteria, archaea, and eukaryotes, stated that in its genetic machinery, the LUCA would have been a "...simpler, more rudimentary entity than the individual ancestors that spawned the three [domains] (and their descendants)".[15]

While the gross anatomy of LUCA can only be reconstructed with much uncertainty, its biochemical mechanisms can be described in some detail, based on the properties currently shared by all independently living organisms on Earth.[16][17][18][19]

The genetic code was most likely based on DNA,[20] though some studies suggest that LUCA may have been defined wholly through RNA.[21] If DNA was present, it was composed exclusively of four nucleotides: deoxyadenosine, deoxycytidine, deoxythymidine, and deoxyguanosine. The DNA was kept double-stranded by a template-dependent enzyme, DNA polymerase. The integrity of the DNA benefited from a group of maintenance and repair enzymes including DNA topoisomerase and DNA ligase. The DNA was also protected by DNA-binding proteins such as histones. The genetic code was composed of 64 three-nucleotide codons which redundantly coded for 20 amino acids.[16][17][18][19] If the code was DNA-based, its genetic code was expressed via single-stranded RNA intermediates. The RNA was produced by a DNA-dependent RNA polymerase using nucleotides similar to those of DNA, with the exception that the DNA nucleotide thymidine was replaced by uridine in RNA.[16][17][18][19]

The genetic code was expressed into proteins. These were assembled from free amino acids by translation of a messenger RNA via a mechanism of ribosomes, transfer RNAs, and a group of related proteins. The ribosomes were composed of two subunits, a big 50S and a small 30S. Each ribosomal subunit was composed of a core of ribosomal RNA surrounded by ribosomal proteins. Both types of RNA molecules (ribosomal and transfer RNAs) played an important role in the catalytic activity of the ribosomes. Only 20 amino acids were used, only in L-isomers, to the exclusion of countless other amino acids. ATP served as an energy intermediate. Several hundred protein enzymes catalyzed chemical reactions to extract energy from fats, sugars, and amino acids, and to synthesize fats, sugars, amino acids, and nucleic acid bases through various chemical pathways.[16][17][18][19]

The cell contained a water-based cytoplasm effectively enclosed by a lipid bilayer membrane. The cell tended to exclude sodium and concentrate potassium by means of specific ion transporters (or ion pumps). The cell multiplied by duplicating all its contents followed by cellular division.[16][17][18][19] The cell used chemiosmosis to produce energy. It also reduced CO2 and oxidized H2 (methanogenesis or acetogenesis) via acetyl-thioesters.[22][23]

The LUCA probably lived in the high-temperature conditions found in deep sea vents caused by ocean water interacting with magma beneath the ocean floor.[24][2]

Hypotheses

A 1990 phylogenetic tree linking all major groups of living organisms to the LUCA (the black trunk at the bottom), based on ribosomal RNA sequence data.[25]

In 1859, Charles Darwin published On the Origin of Species in which he twice stated the hypothesis that there was only one progenitor for all life forms. In the summation he states, "Therefore I should infer from analogy that probably all the organic beings which have ever lived on this earth have descended from some one primordial form, into which life was first breathed."[26] The very last sentence begins with a restatement of the hypothesis: "There is grandeur in this view of life, with its several powers, having been originally breathed into a few forms or into one..."[26]

When the LUCA was hypothesized, cladograms based on genetic distance between living cells indicated that Archaea split early from the rest of life. This was inferred from the fact that the archaeans known at that time were highly resistant to environmental extremes such as high salinity, temperature or acidity, leading some scientists to suggest that the LUCA evolved in areas like the deep ocean vents, where such extremes prevail today. Archaea, however, were later discovered in less hostile environments, and are now believed to be more closely related to the Eukaryota than to the Bacteria, although many details are still unknown.[27][28]

2005 tree of life showing horizontal gene transfers between branches, giving rise to an interconnected network rather than a plain hierarchy

In 2010, based on "the vast array of molecular sequences now available from all domains of life,"[29] a formal test of universal common ancestry was published.[1] The formal test favored the existence of a universal common ancestor over a wide class of alternative hypotheses that included horizontal gene transfer. Basic biochemical principles make it overwhelmingly likely that all organisms do have a single common ancestor. It is extremely unlikely that organisms descended from separate incidents of cell-formation would be able to complete a horizontal gene transfer without garbling each other's genes, converting them into noncoding segments. Further, many more amino acids are chemically possible than the twenty found in modern protein molecules. These lines of chemical evidence, incorporated into the formal statistical test point to a single cell having been the LUCA. While the test overwhelmingly favored the existence of a single LUCA, this does not imply that the LUCA was ever alone: instead, it was one of many early microbes[1] but the only one whose descendents survived beyond the Paleoarchean Era.[30]

With the later gene pool of the LUCA's descendants, with their common framework of the AT/GC rule and the standard twenty amino acids, horizontal gene transfer would have been feasible and could have been very common.

In an earlier hypothesis, Carl Woese (1988) had proposed that: (1) no individual organism can be considered a LUCA, and (2) the genetic heritage of all modern organisms derived through horizontal gene transfer among an ancient community of organisms.[31] While the results described by Theobald (2010) and Saey (2010) demonstrate the existence of a single LUCA, Woese's argument can still be applied to Ur-organisms (initial products of abiogenesis) before the LUCA. At the beginnings of life, ancestry was not as linear as it is today because the genetic code had not evolved.[32] Before high fidelity replication, organisms could not be easily mapped on a phylogenetic tree. However, the LUCA lived after the origin of the genetic code and at least some rudimentary early form of molecular proofreading.

Location of the root

The LUCA used the Wood–Ljungdahl or reductive acetyl–CoA pathway to fix carbon.
For branching of Bacteria phyla, see Bacterial phyla.

The most commonly accepted tree of life, based on several molecular studies, has its root between a monophyletic domain Bacteria and a clade formed by Archaea and Eukaryota.[33][34][35][36][37][38] However, a very small minority of studies place the root in the domain Bacteria, either in the phylum Firmicutes[39] or state that the phylum Chloroflexi is basal to a clade with Archaea and Eukaryotes and the rest of Bacteria (as proposed by Thomas Cavalier-Smith).[40]

Ground-breaking research by William F. Martin (2016) genetically analyzed 6.1 million protein-coding genes and 286,514 protein clusters from sequenced prokaryotic genomes of various phylogenetic trees, and identified 355 protein clusters that were probably common to the LUCA. The results "depict LUCA as anaerobic, CO2-fixing, H2-dependent with a Wood–Ljungdahl pathway (the reductive acetyl-coenzyme A pathway), N2-fixing and thermophilic. LUCA's biochemistry was replete with FeS clusters and radical reaction mechanisms." The cofactors also reveal "dependence upon transition metals, flavins, S-adenosyl methionine, coenzyme A, ferredoxin, molybdopterin, corrins and selenium. Its genetic code required nucleoside modifications and S-adenosylmethionine-dependent methylations."[2][41][42] The results are "quite specific":[3] they show that methanogenic clostridia was a basal clade in the 355 lineages examined, and that the LUCA may therefore have inhabited an anaerobic hydrothermal vent setting in a geochemically active environment rich in H2, CO2, and iron.[2]

These findings could mean that life on Earth originated in such hydrothermal vents, but it is also possible that life was restricted to such locations at some later time, perhaps by the Late Heavy Bombardment.[3] The identification of these genes as being present in LUCA has also been criticized, as they may simply represent later genes which migrated via horizontal gene transfers between archaea and bacteria.[4]

See also

Remark

  1. ^ There is a common misconception that definitions of LUCA and progenote are the same; however, progenote is defined as an organism “still in the process of evolving the relationship between genotype and phenotype”, and it is only hypothesed that LUCA is a progenote.
    Gogarten, Johann Peter; Deamer, David (25 November 2016). "Is LUCA a thermophilic progenote?". Nature Microbiology. 1 (12): 16229. doi:10.1038/nmicrobiol.2016.229. PMID 27886195.

References

  1. ^ a b c Theobald DL (May 2010). "A formal test of the theory of universal common ancestry". Nature. 465 (7295): 219–22. Bibcode:2010Natur.465..219T. doi:10.1038/nature09014. PMID 20463738.
  2. ^ a b c d "The physiology and habitat of the last universal common ancestor" by Madeline C. Weiss, FilipaL.Sousa, Natalia Mrnjavac, Sinje Neukirchen, Mayo Roettger, Shijulal Nelson-Sathi and William F. Martin (July 25, 2016) (Nature Microbiology 16116 | doi:10.1038/NMICROBIOL.2016.116
  3. ^ a b c Wade, Nicholas (25 July 2016). "Meet Luca, the Ancestor of All Living Things". The New York Times. Retrieved 8 October 2018.
  4. ^ a b Gogarten, Johann Peter; Deamer, David (2016). "Is LUCA a thermophilic progenote?". Nature Microbiology. 1 (12): 16229. doi:10.1038/nmicrobiol.2016.229. PMID 27886195.
  5. ^ Doolittle WF (February 2000). "Uprooting the tree of life". Scientific American. 282 (2): 90–95. Bibcode:2000SciAm.282b..90D. doi:10.1038/scientificamerican0200-90. PMID 10710791.
  6. ^ Glansdorff N, Xu Y, Labedan B (2008). "The last universal common ancestor: emergence, constitution and genetic legacy of an elusive forerunner". Biology Direct. 3: 29. doi:10.1186/1745-6150-3-29. PMC 2478661. PMID 18613974.{{cite journal}}: CS1 maint: unflagged free DOI (link)
  7. ^ Borenstein, Seth (13 November 2013). "Oldest fossil found: Meet your microbial mom". Associated Press. Retrieved 15 November 2013.
  8. ^ Noffke, N.; Christian, D.; Wacey, D.; Hazen, R.M. (December 2013). "Microbially induced sedimentary structures recording an ancient ecosystem in the ca. 3.48 billion-year-old Dresser Formation, Pilbara, Western Australia". Astrobiology. 13 (12): 1103–24. Bibcode:2013AsBio..13.1103N. doi:10.1089/ast.2013.1030. PMC 3870916. PMID 24205812.
  9. ^ Ohtomo, Yoko; Kakegawa, Takeshi; Ishida, Akizumi; Nagase, Toshiro; Rosing, Minik T. (2013). "Evidence for biogenic graphite in early Archaean Isua metasedimentary rocks". Nature Geoscience. 7 (1): 25–28. Bibcode:2014NatGe...7...25O. doi:10.1038/ngeo2025.
  10. ^ Hassenkam, T.; Andersson, M. P.; Dalby, K. N.; Mackenzie, D. M. A.; Rosing, M. T. (2017). "Elements of Eoarchean life trapped in mineral inclusions". Nature. 548 (7665): 78–81. Bibcode:2017Natur.548...78H. doi:10.1038/nature23261. PMID 28738409.
  11. ^ Borenstein, Seth (19 October 2015). "Hints of life on what was thought to be desolate early Earth". AP News. Associated Press. Retrieved 9 October 2018.
  12. ^ Bell, Elizabeth A.; Boehnke, Patrick; Harrison, T. Mark; Mao, Wendy L. (24 November 2015). "Potentially biogenic carbon preserved in a 4.1 billion-year-old zircon". Proceedings of the National Academy of Sciences of the United States of America. 112 (47): 14518–21. Bibcode:2015PNAS..11214518B. doi:10.1073/pnas.1517557112. ISSN 1091-6490. PMC 4664351. PMID 26483481.
  13. ^ Dodd, Matthew S.; Papineau, Dominic; Grenne, Tor; slack, John F.; Rittner, Martin; Pirajno, Franco; O'Neil, Jonathan; Little, Crispin T. S. (2 March 2017). "Evidence for early life in Earth's oldest hydrothermal vent precipitates". Nature. 543 (7643): 60–64. Bibcode:2017Natur.543...60D. doi:10.1038/nature21377. PMID 28252057. Archived from the original on 10 February 2017.
  14. ^ Darwin, C. (1859), The Origin of Species by Means of Natural Selection, John Murray, p. 490
  15. ^ Woese, C. R.; Kandler, O.; Wheelis, M. L. (1990). "Towards a natural system of organisms: proposal for the domains Archaea, Bacteria, and Eucarya". Proceedings of the National Academy of Sciences. 87 (12): 4576–79. Bibcode:1990PNAS...87.4576W. doi:10.1073/pnas.87.12.4576. ISSN 0027-8424. PMC 54159. PMID 2112744.
  16. ^ a b c d e Wächtershäuser, Günter (1998). "Towards a Reconstruction of Ancestral Genomes by Gene Cluster Alignment". Systematic and Applied Microbiology. 21 (4): 473–74, IN1, 475–77. doi:10.1016/S0723-2020(98)80058-1.
  17. ^ a b c d e Gregory, Michael. "What is Life?". Clinton College. Archived from the original on 2007-12-13. {{cite web}}: Unknown parameter |deadurl= ignored (|url-status= suggested) (help)
  18. ^ a b c d e Pace NR (January 2001). "The universal nature of biochemistry". Proceedings of the National Academy of Sciences of the United States of America. 98 (3): 805–08. Bibcode:2001PNAS...98..805P. doi:10.1073/pnas.98.3.805. PMC 33372. PMID 11158550.
  19. ^ a b c d e Wächtershäuser, G. (January 2003). "From pre-cells to Eukarya – a tale of two lipids". Molecular Microbiology. 47 (1): 13–22. doi:10.1046/j.1365-2958.2003.03267.x. PMID 12492850.
  20. ^ Garwood, Russell J. (2012). "Patterns In Palaeontology: The first 3 billion years of evolution". Palaeontology Online. 2 (11): 1–14. Retrieved June 25, 2015.
  21. ^ Marshall, Michael. "Life began with a planetary mega-organism". New Scientist.
  22. ^ Martin, W.; Russell, M.J. (October 2007). "On the origin of biochemistry at an alkaline hydrothermal vent". Philosophical Transactions of the Royal Society of London. Series B, Biological Sciences. 362 (1486): 1887–925. doi:10.1098/rstb.2006.1881. PMC 2442388. PMID 17255002.
  23. ^ Lane, N.; Allen, J.F.; Martin, W. (April 2010). "How did LUCA make a living? Chemiosmosis in the origin of life". BioEssays. 32 (4): 271–80. doi:10.1002/bies.200900131. PMID 20108228.
  24. ^ Wade, Nicholas (2016-07-25). "Meet Luca, the Ancestor of All Living Things". The New York Times. ISSN 0362-4331. Retrieved 26 July 2016.
  25. ^ Woese, C.R.; Kandler, O.; Wheelis, M.L. (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): 4576–79. Bibcode:1990PNAS...87.4576W. doi:10.1073/pnas.87.12.4576. PMC 54159. PMID 2112744.
  26. ^ a b Darwin, Charles. On the Origin of Species. London: John Murray, Albermarle Street. 1859. pp. 484, 490.
  27. ^ Xie, Q.; Wang, Y.; Lin, J.; Qin, Y.; Wang, Y.; Bu, W. (2012). "Potential key bases of ribosomal RNA to kingdom-specific spectra of antibiotic susceptibility and the possible archaeal origin of eukaryotes". PLoS ONE. 7 (1): e29468. Bibcode:2012PLoSO...729468X. doi:10.1371/journal.pone.0029468. PMC 3256160. PMID 22247777.{{cite journal}}: CS1 maint: unflagged free DOI (link)
  28. ^ Yutin, N.; Makarova K.S.; Mekhedov S.L.; Wolf, Y.I.; Koonin, E.V. (August 2008). "The deep archaeal roots of eukaryotes". Molecular Biology and Evolution. 25 (8): 1619–30. doi:10.1093/molbev/msn108. PMC 2464739. PMID 18463089.
  29. ^ Steel, M.; Penny, D. (May 2010). "Origins of life: Common ancestry put to the test". Nature. 465 (7295): 168–69. Bibcode:2010Natur.465..168S. doi:10.1038/465168a. PMID 20463725.
  30. ^ Egel, Richard (March 2012). "Primal Eukaryogenesis: On the Communal Nature of Precellular States, Ancestral to Modern Life". Life. 2 (1): 170–212. doi:10.3390/life2010170. PMC 4187143. PMID 25382122.{{cite journal}}: CS1 maint: unflagged free DOI (link)
  31. ^ Woese C (June 1998). "The universal ancestor". Proceedings of the National Academy of Sciences of the United States of America. 95 (12): 6854–59. Bibcode:1998PNAS...95.6854W. doi:10.1073/pnas.95.12.6854. PMC 22660. PMID 9618502.
  32. ^ Maynard Smith, John; Szathmáry, Eörs (1995). The Major Transitions in Evolution. Oxford, England: Oxford University Press. ISBN 978-0-19-850294-4.[page needed]
  33. ^ Brown, J. R., and W. F. Doolittle. 1995. “Root of the Universal Tree of Life Based on Ancient Aminoacyl-tRNA Synthetase Gene Duplications.” Proc Natl Acad Sci USA 92 (7): 2441–45. PMID 7708661
  34. ^ Gogarten, J. P., H. Kibak, P. Dittrich, L. Taiz, E. J. Bowman, B. J. Bowman, M. F. Manolson, et al. 1989. “Evolution of the Vacuolar H+-ATPase: Implications for the Origin of Eukaryotes.” Proc Natl Acad Sci USA 86 (17): 6661–65. PMID 2528146
  35. ^ Gogarten, J.P.; Taiz, L. (1992). "Evolution of Proton Pumping ATPases: Rooting the Tree of Life". Photosynthesis Research. 33 (2): 137–46. doi:10.1007/BF00039176. PMID 24408574.
  36. ^ Gribaldo, S, and P Cammarano. 1998. “The Root of the Universal Tree of Life Inferred from Anciently Duplicated Genes Encoding Components of the Protein-Targeting Machinery.” Journal of Molecular Evolution 47 (5): 508–16. PMID: 9797401
  37. ^ Iwabe, Naoyuki, Kei-Ichi Kuma, Masami Hasegawa, Syozo Osawa, Takashi Miyata Source, Masami Hasegawat, Syozo Osawat, and Takashi Miyata. 1989. “Evolutionary Relationship of Archaebacteria, Eubacteria, and Eukaryotes Inferred from Phylogenetic Trees of Duplicated Genes.” Proc Natl Acad Sci USA 86 (86): 9355–59. PMID 2531898
  38. ^ Boone, David R.; Castenholz, Richard W.; Garrity, George M., eds. (2001). The Archaea and the Deeply Branching and Phototrophic Bacteria. Bergey's Manual of Systematic Bacteriology. Springer. doi:10.1007/978-0-387-21609-6. ISBN 978-0-387-21609-6.[page needed]
  39. ^ Valas, R.E.; Bourne, P.E. (2011). "The origin of a derived superkingdom: how a gram-positive bacterium crossed the desert to become an archaeon". Biology Direct. 6: 16. doi:10.1186/1745-6150-6-16. PMC 3056875. PMID 21356104.{{cite journal}}: CS1 maint: unflagged free DOI (link)
  40. ^ Cavalier-Smith T (2006). "Rooting the tree of life by transition analyses". Biology Direct. 1: 19. doi:10.1186/1745-6150-1-19. PMC 1586193. PMID 16834776.{{cite journal}}: CS1 maint: unflagged free DOI (link)
  41. ^ Nick Lane: The Vital Question – Energy, Evolution, and the Origins of Complex Life, WW Norton, 2015, ISBN 978-0-393-08881-6
  42. ^ Joseph F. Sutherland: on The Origin Of Tha Bacteria And The Archaea, auf B.C vom 16. August 2014

External links

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