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==History== {{Human timeline}} ===Early research=== Early research using [[molecular clock]] methods was done during the late 1970s to early 1980s. [[Allan Wilson (biologist)|Allan Wilson]], [[Mark Stoneking]], [[Rebecca L. Cann]] and Wesley Brown found that [[mutation]] in human mtDNA was unexpectedly fast, at 0.02 substitution per base (1%) in a million years, which is 5–10 times faster than in [[nuclear DNA]].<ref>{{cite journal |vauthors=Brown WM, George M, Wilson AC |title=Rapid evolution of animal mitochondrial DNA |journal=Proceedings of the National Academy of Sciences of the United States of America |volume=76 |issue=4 |pages=1967–1971 |date=April 1979 |pmid=109836 |pmc=383514 |doi=10.1073/pnas.76.4.1967 |doi-access=free |bibcode=1979PNAS...76.1967B }}</ref> Related work allowed for an analysis of the evolutionary relationships among gorillas, [[Pan (genus)|chimpanzee]]s ([[common chimpanzee]] and [[bonobo]]) and humans.<ref>{{cite journal |vauthors=Ferris SD, Wilson AC, Brown WM |title=Evolutionary tree for apes and humans based on cleavage maps of mitochondrial DNA |journal=Proceedings of the National Academy of Sciences of the United States of America |volume=78 |issue=4 |pages=2432–2436 |date=April 1981 |pmid=6264476 |pmc=319360 |doi=10.1073/pnas.78.4.2432 |doi-access=free |bibcode=1981PNAS...78.2432F }}</ref> With data from 21 human individuals, Brown published the first estimate on the age of the mt-MRCA at 180,000 years ago in 1980.<ref>{{cite journal |vauthors=Brown WM |title=Polymorphism in mitochondrial DNA of humans as revealed by restriction endonuclease analysis |journal=Proceedings of the National Academy of Sciences of the United States of America |volume=77 |issue=6 |pages=3605–3609 |date=June 1980 |pmid=6251473 |pmc=349666 |doi=10.1073/pnas.77.6.3605 |doi-access=free |bibcode=1980PNAS...77.3605B }}</ref> A statistical analysis published in 1982 was taken as evidence for [[recent African origin]] (a hypothesis which at the time was competing with Asian origin of ''H. sapiens'').<ref name=Cann82>{{cite journal |vauthors=Cann RL, Brown WM, Wilson AC |title=Evolution of human mitochondrial DNA: a preliminary report |journal=Progress in Clinical and Biological Research |volume=103 Pt A |issue=Pt A |pages=157–165 |year=1982 |pmid=6298804 }}</ref><ref>{{cite journal |vauthors=Cann RL, Wilson AC |title=Length mutations in human mitochondrial DNA |journal=Genetics |volume=104 |issue=4 |pages=699–711 |date=August 1983 |pmid=6311667 |pmc=1202135 |doi=10.1093/genetics/104.4.699 }}</ref><ref name="git" /> ===1987 publication=== By 1985, data from the mtDNA of 145 women of different populations, and of two [[cell lines]], [[HeLa]] and GM 3043, derived from an African American and a [[ǃKung people|ǃKung]] respectively, were available. After more than 40 revisions of the draft, the manuscript was submitted to [[Nature (journal)|''Nature'']] in late 1985 or early 1986<ref name="git">{{cite journal |vauthors=Cann R |title=All about mitochondrial eve: an interview with Rebecca Cann. Interview by Jane Gitschier |journal=PLOS Genetics |volume=6 |issue=5 |pages=e1000959 |date=May 2010 |pmid=20523888 |pmc=2877732 |doi=10.1371/journal.pgen.1000959 |doi-access=free }}</ref> and published on 1 January 1987. The published conclusion was that all current human mtDNA originated from a single population from Africa, at the time dated to between 140,000 and 200,000 years ago.<ref name="pmid3025745">{{cite journal |vauthors=Cann RL, Stoneking M, Wilson AC |title=Mitochondrial DNA and human evolution |journal=Nature |volume=325 |issue=6099 |pages=31–36 |year=1987 |pmid=3025745 |doi=10.1038/325031a0 |s2cid=4285418 |bibcode=1987Natur.325...31C }}</ref> The dating for "Eve" was a blow to the [[multiregional hypothesis]], which was debated at the time, and a boost to the theory of the [[recent African origin of modern humans|recent origin model]].<ref name="pmid1840702">{{cite journal |vauthors=Vigilant L, Stoneking M, Harpending H, Hawkes K, Wilson AC |title=African populations and the evolution of human mitochondrial DNA |journal=Science |volume=253 |issue=5027 |pages=1503–1507 |date=September 1991 |pmid=1840702 |doi=10.1126/science.1840702 |bibcode=1991Sci...253.1503V }}</ref> Cann, Stoneking and Wilson did not use the term "Mitochondrial Eve" or even the name "Eve" in their original paper. It is however used by Cann in an article entitled "In Search of Eve" in the September–October 1987 issue of ''The Sciences''.<ref>{{cite journal|vauthors=Cann RL |year=1987 |title=In Search of Eve |url=https://onlinelibrary.wiley.com/doi/abs/10.1002/j.2326-1951.1987.tb02967.x |journal=The Sciences |volume=27 |issue=5 |pages=30–37 |doi=10.1002/j.2326-1951.1987.tb02967.x}}</ref> It appears in the October 1987 article in [[science (journal)|''Science'']] by [[Roger Lewin]], headlined "The Unmasking of Mitochondrial Eve".<ref name=lew>{{cite journal |vauthors=Lewin R |title=The unmasking of mitochondrial Eve |journal=Science |volume=238 |issue=4823 |pages=24–26 |date=October 1987 |pmid=3116666 |doi=10.1126/science.3116666 |bibcode=1987Sci...238...24L }}</ref> The biblical connotation was very clear from the start. The accompanying research news in ''Nature'' had the title "Out of the garden of Eden".<ref>{{cite journal |vauthors=Wainscoat J |title=Human evolution. Out of the garden of Eden |journal=Nature |volume=325 |issue=6099 |pages=13 |year=1987 |pmid=3796736 |doi=10.1038/325013a0 |s2cid=13187170 |bibcode=1987Natur.325...13W |doi-access=free }}</ref> Wilson himself preferred the term "Lucky Mother"<ref>{{cite web|url=https://io9.gizmodo.com/the-scientists-behind-mitochondrial-eve-tell-us-about-t-5879991 |vauthors=Wilkins A |date=27 January 2012 |title=The scientists behind Mitochondrial Eve tell us about the 'lucky mother' who changed human evolution forever |work=Gizmodo |access-date=23 December 2019}}</ref> and thought the use of the name Eve "regrettable".<ref name=lew/><ref>{{cite book|vauthors=Cann RL |year=1997 |chapter=Chapter 4: Mothers, Labels, and Misogyny |title=Women in Human Evolution |editor=Hager LD |publisher=Routledge |location=London |pages=75–89 |isbn=9780415108331}}</ref> But the concept of Eve caught on with the public and was repeated in a ''[[Newsweek]]'' cover story (11 January 1988 issue featured a depiction of Adam and Eve on the cover, with the title "The Search for Adam and Eve"),<ref>{{cite web|vauthors=Tierney J |year=1992 |title=The Search for Adam and Eve|url=http://www.virginia.edu/woodson/courses/aas102%20(spring%2001)/articles/tierney.html |work=Newsweek |publisher=Carter G. Woodson Institute for Afro-American and African Studies |via=Internet Archive |archive-url=https://web.archive.org/web/20150320110036/http://www.virginia.edu/woodson/courses/aas102%20(spring%2001)/articles/tierney.html |access-date=21 July 2019|archive-date=20 March 2015 }}</ref> and a cover story in ''[[Time (magazine)|Time]]'' on 26 January 1987.<ref>{{cite magazine|vauthors=Lemonick MD |date=26 January 1987|title=Everyone's Genealogical Mother|url=http://www.time.com/time/magazine/article/0,9171,963320,00.html |archive-url=https://web.archive.org/web/20080923010301/http://www.time.com/time/magazine/article/0,9171,963320,00.html |url-status=dead |archive-date=23 September 2008 |magazine=Time |access-date=13 May 2013}}</ref> ===Reception and later research=== Shortly after the 1987 publication, its methodology and secondary conclusions were criticised.<ref>{{cite journal |vauthors=Darlu P, Tassy P |title=Disputed African origin of human populations |journal=Nature |volume=329 |issue=6135 |pages=111–112 |year=1987 |pmid=3114640 |doi=10.1038/329111b0 |s2cid=4313392 |bibcode=1987Natur.329..111D |doi-access=free }}</ref> Both the dating of mt-Eve and the relevance of the age of the purely matrilineal descent for population replacement were subjects of controversy during the 1990s;<ref>{{cite journal |vauthors=Maddison DR |title=African Origin of human mitochondrial DNA reexamined |journal=Systematic Zoology |volume=40 |issue=3 |pages=355–63 |year=1991 |doi=10.2307/2992327 |jstor=2992327 }}</ref><ref>{{cite journal |vauthors=Nei M |title=Age of the common ancestor of human mitochondrial DNA |journal=Molecular Biology and Evolution |volume=9 |issue=6 |pages=1176–1178 |date=November 1992 |pmid=1435241 |doi=10.1093/oxfordjournals.molbev.a040785 |doi-access=free }}</ref><ref>{{cite journal |vauthors=Ayala FJ |title=The myth of Eve: molecular biology and human origins |journal=Science |volume=270 |issue=5244 |pages=1930–1936 |date=December 1995 |pmid=8533083 |doi=10.1126/science.270.5244.1930 |s2cid=42801341 |doi-access=free |bibcode=1995Sci...270.1930A }}</ref><ref>{{cite journal |vauthors=Templeton A |title=Out of Africa again and again |journal=Nature |volume=416 |issue=6876 |pages=45–51 |date=March 2002 |pmid=11882887 |doi=10.1038/416045a |s2cid=4397398 |bibcode=2002Natur.416...45T }}</ref> [[Alan Templeton]] (1997) asserted that the study did "not support the hypothesis of a recent African origin for all of humanity following a split between Africans and non-Africans 100,000 years ago" and also did "not support the hypothesis of a recent global replacement of humans coming out of Africa."<ref>{{cite journal |vauthors=Templeton AR |title=Out of Africa? What do genes tell us? |journal=Current Opinion in Genetics & Development |volume=7 |issue=6 |pages=841–847 |date=December 1997 |pmid=9468796 |doi=10.1016/S0959-437X(97)80049-4 }}</ref> The placement by {{Harvtxt|Cann|Stoneking|Wilson|1987}} of a relatively small population of humans in sub-Saharan Africa was consistent with the hypothesis of Cann (1982) and lent considerable support for the "recent out-of-Africa" scenario. In 1999, Krings et al. eliminated problems in molecular clocking postulated by Nei (1992)<ref>{{cite journal |vauthors=Nei M |year=1992 |title=Age of the common ancestor of human mitochondrial DNA |url=https://www.researchgate.net/publication/21720959 |journal=Molecular Biology and Evolution |volume=9 |issue=6 |pages=1176–1178|doi=10.1093/oxfordjournals.molbev.a040785 |pmid=1435241 |doi-access=free }}</ref> when it was found that the mtDNA sequence for the same region was substantially different from the MRCA relative to any human sequence.<ref>{{cite journal |vauthors=Krings M, Salem AE, Bauer K, Geisert H, Malek AK, Chaix L, Simon C, Welsby D, Di Rienzo A, Utermann G, Sajantila A, Pääbo S, Stoneking M |display-authors=6 |title=mtDNA analysis of Nile River Valley populations: A genetic corridor or a barrier to migration? |language=English |journal=American Journal of Human Genetics |volume=64 |issue=4 |pages=1166–1176 |date=April 1999 |pmid=10090902 |pmc=1377841 |doi=10.1086/302314 }}</ref><ref>{{cite journal |vauthors=Krings M, Stone A, Schmitz RW, Krainitzki H, Stoneking M, Pääbo S |title=Neandertal DNA sequences and the origin of modern humans |language=English |journal=Cell |volume=90 |issue=1 |pages=19–30 |date=July 1997 |pmid=9230299 |doi=10.1016/S0092-8674(00)80310-4 |s2cid=13581775 |doi-access=free |hdl=11858/00-001M-0000-0025-0960-8 |hdl-access=free }}</ref> In 1997, {{Harvtxt|Parsons|Muniec|Sullivan|Woodyatt|1997}} published a study of mtDNA mutation rates in a single, well-documented family (the [[Romanov]] family of Russian royalty). In this study, they calculated a mutation rate upwards of twenty times higher than previous results.<ref>{{cite journal |last1=Parsons | first1 = Thomas J. |last2 = Muniec | first2 = David S. | last3 = Sullivan | first3= Kevin | last4 = Woodyatt | first4 = Nicola | last5= Alliston-Greiner | first5=Rosemary |last6 = Wilson | first6 = Mark R. |last7= Berry | first7= Dianna L. | last8 = Holland | first8 = Koren A. |last9 = Weedn | first9 = Viktor W. | last10 = Gill | first10 = Peter| last11= Holland | first11 = Mitchell M. |title=A high observed substitution rate in the human mitochondrial DNA control region |journal=Nature Genetics |volume=15 |issue=4 |pages=363–368 |date=April 1997 |pmid=9090380 |doi=10.1038/ng0497-363 |s2cid=32812244 }}</ref> Although the original research did have analytical limitations, the estimate on the age of the mt-MRCA has proven robust.<ref>{{cite web|vauthors=Holsinger K|url=http://darwin.eeb.uconn.edu/eeb348/lecturenotes/coalescent/node5.html |title=(Mathematics of the coalescent) An example: Mitochondrial Eve |work=Holsinger Lab |date=29 September 2012 |access-date=16 May 2013}}</ref><ref>{{cite journal |vauthors=Cyran KA, Kimmel M |title=Alternatives to the Wright-Fisher model: the robustness of mitochondrial Eve dating |journal=Theoretical Population Biology |volume=78 |issue=3 |pages=165–172 |date=November 2010 |pmid=20600209 |doi=10.1016/j.tpb.2010.06.001 }}</ref> More recent age estimates have remained consistent with the 140–200 [[Year#Abbreviations yr and ya|kya]] estimate published in 1987: A 2013 estimate dated Mitochondrial Eve to about 160 kya (within the reserved estimate of the original research) and [[Out of Africa II]] to about 95 kya.<ref name=Fu2013>{{cite journal |vauthors=Fu Q, Mittnik A, Johnson PL, Bos K, Lari M, Bollongino R, Sun C, Giemsch L, Schmitz R, Burger J, Ronchitelli AM, Martini F, Cremonesi RG, Svoboda J, Bauer P, Caramelli D, Castellano S, Reich D, Pääbo S, Krause J |display-authors=6 |title=A revised timescale for human evolution based on ancient mitochondrial genomes |journal=Current Biology |volume=23 |issue=7 |pages=553–559 |date=April 2013 |pmid=23523248 |pmc=5036973 |doi=10.1016/j.cub.2013.02.044 }}</ref> Another 2013 study (based on [[genome sequencing]] of 69 people from 9 different populations) reported the age of Mitochondrial Eve between 99 and 148 kya and that of the [[Y-MRCA]] between 120 and 156 kya.<ref name=poz/>
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