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Mitochondrial Eve

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Haplogroup Modern humans
Possible time of origin152,000 – 234,000 BP[1]
Possible place of originEast Africa
Ancestorn/a
DescendantsMitochondrial macro-haplogroups L0, L1, and L5
Defining mutationsNone

In the field of human genetics, Mitochondrial Eve, who is estimated to have lived approximately 140,000–200,000 years ago, refers to the matrilineal most recent common ancestor (MRCA) of all currently living anatomically modern humans. In other words, she was the most recent woman from whom all living humans today descend, on their mother’s side, and through the mothers of those mothers, and so on, back until all lines converge on one person. Because all mitochondrial DNA (mtDNA) generally (but see paternal mtDNA transmission) is passed from mother to offspring without recombination, all mtDNA in every living person is directly descended from hers by definition, differing only by the mutations that over generations have occurred in the germ cell mtDNA since the conception of the original "Mitochondrial Eve".

Mitochondrial Eve is named after mitochondria and the biblical Eve.[2] Unlike her biblical namesake, she was not the only living human female of her time. However, her female contemporaries, except her mother, failed to produce a direct unbroken female line to any living woman in the present day.

Mitochondrial Eve is estimated to have lived between 140,000 and 200,000 years ago,[3][4] most likely in East Africa,[5] when Homo sapiens sapiens (anatomically modern humans) were developing as a population distinct from other human sub-species.

Mitochondrial Eve lived later than Homo heidelbergensis and the emergence of Homo neanderthalensis, but earlier than the out of Africa migration.[6] The dating for "Eve" was a blow to the multiregional hypothesis, and a boost to the hypothesis of the origin of modern humans in Africa and spread from there, replacing more "archaic" human populations such as Neanderthals. As a result, the latter hypothesis became dominant.

Analogous to Mitochondrial Eve, Y-chromosomal Adam is the man from whom all living humans are descended patrilineally. The inherited DNA in the male case is his nuclear Y chromosome, rather than the mtDNA. There is no reason why Mitochondrial Eve and Y-chromosomal Adam should necessarily have lived at the same time.[7] Y-chromosomal Adam had been estimated to have lived during a wide range of times from 180,000 to 581,000 years ago.[8][9][10] and another giving between 120,000 and 156,000 years.[11][12]

Discovery

Origin of the science

Mitochondrial Eve was theoretically and experimentally discovered by Allan Wilson, of the University of California, Berkeley, with his doctoral students Mark Stoneking and Rebecca L. Cann through a wave of advances in genetics. Wilson had earned a worldwide recognition with his molecular clock, the technique by which species divergence time can be estimated. Inspired by the discovery of mtDNA and its unique properties, he had been motivated to employ the novel field in the study of human origin. It came to reality when Wesley M. Brown joined him as a post-doc in 1978. Brown had developed a firm grasp of restriction enzyme digestion of mtDNA from University of California, San Francisco and California Institute of Technology. The merger of two different techniques with a similar goal proved to be fruitful. In 1979 they found that mutation in mtDNA was exceptionally fast (0.02 substitution per base (1%) in a million years, which means 5-10 times faster than in nuclear DNA), especially in humans when compared to those of other primates including Guinea baboon, rhesus macaque, and guenon. This was the first crucial step in understanding the usefulness of mtDNA in estimating evolutionary divergence and relationships,[13] and they soon published a paper describing the evolutionary relationships among gorillas, chimpanzees (common chimpanzee and bonobo) and humans.[14] Strictly on the evolution of modern humans, Brown studied mitochondrial fractions from 21 individuals (placentas from labour and delivery) of different ethnicity and geographic origin. In 1980, he discovered incredible variations among humans, and by estimating the rate of mutation, he concluded that modern human species originated as recently as 180,000 years ago.[15] But the shortcomings were that medical records were disassociated and he didn't know about the actual donors' ethnic background.

Enter into the scene a fresh doctoral student, Cann, who had started collecting mtDNA in 1979 based on ethnicity. She collected samples from US hospitals from women (their placenta, to be precise) of different ethnic backgrounds, such as from Asia, South Pacific, Europe and Americans of African descent (she could not get samples from aboriginal Africans). Their initial observation of the mitochondrial genome of 100 individuals led them to resolve that it was possible to trace back individual female lineages back hundreds of generations thereby reaching the point of a common ancestor.[16] However, in 1983, Cann and Wilson failed to show common ancestry from 112 individuals of different ethnic groups, and they had no indication of African samples as the oldest or the lineage of the common ancestor using an analysis called Fitch-Margoliash trees.[17] Even in Cann's thesis in 1982, in which more elaborate estimates (including PAUP and Neighbor joining in addition to Fitch-Margoliash) were used, African origin of all humankind was not yet convincing, especially to Wilson, though the hunch was evident there.[18] Prudence in Wilson was largely in part due to a publication of an eminent geneticist, Luigi Luca Cavalli-Sforza and his team in 1981 in which they concluded that mtDNA pointed most likely to human origin from Asia.[19] In 1981, a fresh graduate student, Stoneking joined Wilson's lab for his PhD and immediately pursued Cann's work in collecting samples from aboriginal Australians and New Guineans from hospitals in Australia and New Guinea. By 1985, armed with mtDNA of 145 women of different population, and of two cell lines, HeLa and GM 3043, derived from a Black American and a !Kung respectively, the reluctant Wilson became convinced. After more than 40 revision of the draft, the manuscript was submitted to Nature in March 1986 (but according to Cann it was in late 1985; see Gitschier, 2010,[18] but the published paper indicated otherwise). The bombshell was eventually dropped on 1 January 1987 in Nature in which their findings clearly showed that all mtDNA originated from a single population, that is from Africa. To paraphrase their conclusion, all living humans were descended from one specific woman, who lived between 140,000 and 200,000 years ago in Africa.[3] The concept is a serious challenge to the prevailing multiregional theory, and directly implies recent African origin of modern humans, hence, the underpinning of the so-called the Out of Africa theory.[20]

Origin of the name

Cann, Stoneking and Wilson did not use the term “Mitochondrial Eve” or even the name “Eve” in their original paper; in fact it appears to be a popularised media interpretation, obviously because of its catchy name.[21] The name stemmed first from research news of the scientific breakthrough in Science in 1987, where Roger Lewin reported with a rather captivating title "The Unmasking of Mitochondrial Eve".[22] The biblical connotation was very clear from the start. The accompanying research news in Nature had the obvious title "Out of the garden of Eden".[23] Wilson was aware of it and wanted to use a more euphemistic "Lucky Mother" as the name.[24] When the use of Eve spread like wildfire, Wilson remarked it as "regrettable".[22] Scientists including the original researchers attempted to avoid the biblical confusion by trying to give alternative name like "African Eve", most ardently by Wilson himself, obviously as a pun.[25] But the subsequent popularisation enamoured the public's eye such as with the Newsweek cover story (11 January 1998 issue had Adam and Eve on the cover with the title "The Search for Adam and Eve"),[26] and a cover story in Time on 26 January 1987,[27] they were the baptism on the christened name. It is interesting that even though Mitochondrial Eve only vaguely denotes the biblical Eve, on immediate aftermath of its publication, Cann received not only scientific criticisms, but also public and religious hostility with hate mails, crank mail, late night calls, and even a visit from FBI.[18]

Female and mitochondrial ancestry

Further information: Genetic genealogy (matrilineal), Mitochondrial DNA, and Human mitochondrial molecular clock
Through random drift or selection the female-lineage will trace back to a single female, such as Mitochondrial Eve. In this example over five generations colors represent extinct matrilineal lines and black the matrilineal line descended from mtDNA MRCA.
Simplified Human mitochondrial phylogeny

Without a DNA sample, it is not possible to reconstruct the complete genetic makeup (genome) of any individual who died very long ago. By analysing descendants' DNA, however, parts of ancestral genomes are estimated by scientists. Mitochondrial DNA (mtDNA) and Y-chromosome DNA are commonly used to trace ancestry in this manner. mtDNA is generally passed un-mixed from mothers to children of both sexes, along the maternal line, or matrilineally.[28][29] Matrilineal descent goes back to our mothers, to their mothers, until all female lineages converge.

Branches are identified by one or more unique markers which give a mitochondrial "DNA signature" or "haplotype" (e.g. the CRS is a haplotype). Each marker is a DNA base-pair that has resulted from an SNP mutation. Scientists sort mitochondrial DNA results into more or less related groups, with more or less recent common ancestors. This leads to the construction of a DNA family tree where the branches are in biological terms clades, and the common ancestors such as Mitochondrial Eve sit at branching points in this tree. Major branches are said to define a haplogroup (e.g. CRS belongs to haplogroup H), and large branches containing several haplogroups are called "macro-haplogroups".

The mitochondrial clade which Mitochondrial Eve defines is the species Homo sapiens sapiens itself, or at least the current population or "chronospecies" as it exists today. In principle, earlier Eves can also be defined going beyond the species, for example one who is ancestral to both modern humanity and Neanderthals, or, further back, an "Eve" ancestral to all members of genus Homo and chimpanzees in genus Pan. According to current nomenclature, Mitochondrial Eve's haplogroup was within mitochondrial haplogroup L because this macro-haplogroup contains all surviving human mitochondrial lineages today, and she must predate the emergence of L0.

The variation of mitochondrial DNA between different people can be used to estimate the time back to a common ancestor, such as Mitochondrial Eve. This works because, along any particular line of descent, mitochondrial DNA accumulates mutations at the rate of approximately one every 3,500 years.[30][31][32] A certain number of these new variants will survive into modern times and be identifiable as distinct lineages. At the same time some branches, including even very old ones, come to an end, when the last family in a distinct branch has no daughters.

Mitochondrial Eve is the most recent common matrilineal ancestor for all modern humans. Whenever one of the two most ancient branch lines dies out, the MRCA will move to a more recent female ancestor, always the most recent mother to have more than one daughter with living maternal line descendants alive today. The number of mutations that can be found distinguishing modern people is determined by two criteria: firstly and most obviously, the time back to her, but secondly and less obviously by the varying rates at which new branches have come into existence and old branches have become extinct. By looking at the number of mutations which have been accumulated in different branches of this family tree, and looking at which geographical regions have the widest range of least related branches, the region where Eve lived can be proposed.

Common misconceptions

Not the only woman

One misconception surrounding mitochondrial Eve is that since all women alive today descended in a direct unbroken female line from her, she must have been the only woman alive at the time.[7][33] However, nuclear DNA studies indicate that the size of the ancient human population never dropped below tens of thousands. Other women living during Eve's time have descendants alive today, but at some point in the past each of their lines of descent included at least one male, thereby breaking the mitochondrial DNA lines of descent. By contrast, Eve's lines of descent to each person alive today includes precisely one purely matrilineal line.[33]

Not a fixed individual over time

The definition of mitochondrial Eve is fixed, but the person in prehistory who will fit this definition can change, not only because of new discoveries, but also because of unbroken mother-daughter lines coming to an end by chance. It follows from the definition of Mitochondrial Eve that she had at least two daughters who both have unbroken female lineages that have survived to the present day. Every generation, mitochondrial lineages end, such as when an only daughter has only sons. When the mitochondrial lineages of daughters of mitochondrial Eve die out, then the title of "Mitochondrial Eve" shifts forward from the remaining daughter through her matrilineal descendants, until the first descendant is reached who had at least two daughters who both have living, matrilineal descendants. Once a lineage has died out it is irretrievably lost and this mechanism can thus only shift the title of "Mitochondrial Eve" forward in time.

Not necessarily a contemporary of "Y-chromosomal Adam"

Sometimes mitochondrial Eve is assumed to have lived at the same time as Y-chromosomal Adam, who all living people are descended from patrilineally, perhaps even meeting and mating with him. Even if this were true, which is currently regarded as highly unlikely, this would only be a coincidence. Like mitochondrial "Eve", Y-chromosomal "Adam" probably lived in Africa; however, according to the latest proposals this "Eve" lived much later than "Adam" – some 140,000 years later.[9] (Earlier studies considered, conversely, that "Eve" lived earlier than "Adam".)[34] But it is also possible that they lived at the same time (then again not necessarily at the same place), as indicated by an August 2013 report.[11][35][36]

Not the most recent ancestor shared by all humans

Main article: Most recent common ancestor

Mitochondrial Eve is the most recent common matrilineal ancestor, not the most recent common ancestor. Since the mtDNA is inherited maternally and recombination is either rare or absent, it is relatively easy to track the ancestry of the lineages back to a MRCA; however, this MRCA is valid only when discussing mitochondrial DNA. An approximate sequence from newest to oldest can list various important points in the ancestry of modern human populations:

  • The Human MRCA. All humans alive today share a surprisingly recent common ancestor, perhaps even within the last 5,000 years, even for people born on different continents.[37]
  • The Identical ancestors point. Just a few thousand years before the most recent single ancestor shared by all living humans was the time at which all humans who were then alive either left no descendants alive today or were common ancestors to all humans alive today. In other words, "each present-day human has exactly the same set of genealogical ancestors" alive at the "Identical ancestors point" in time. This is far more recent than Mitochondrial Eve.[37]
  • Mitochondrial Eve, the most recent female-line common ancestor of all living people.
  • "Y-chromosomal Adam", the most recent male-line common ancestor of all living people, is currently thought to have lived long before Mitochondrial Eve.

Not the biblical Eve

Owing to its figurative reference to the first woman in the Biblical Book of Genesis, the Mitochondrial Eve theory initially met with enthusiastic endorsement from some young earth creationists, who viewed the theory as a validation of the biblical creation story. Some even went so far as to claim that the Mitochondrial Eve theory disproved evolution.[38][39][40] However, the theory does not suggest any relation between biblical Eve and Mitochondrial Eve, or vice versa. Mitochondrial Eve is not a fixed individual, had a mother, was not the only woman of her time, and Y-chromosomal Adam is unlikely to have been her sexual partner, or indeed to have been contemporaneous to her.

Criticism and new evidence

Soon after the watershed publication, some population geneticists criticised the analytical method as flawed, and also criticised some of the secondary conclusions[41] while some claimed the interpretation as dubious.[42][43][44][45] Alan Templeton 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."[46] This criticism was popularised into a general conception that human mtDNA analysis and the hypothesis of recent African origin are false.[47][48]

Although the original research did have analytical limitations, the results were not that seriously flawed.[49][50] However they should be interpreted, estimates of the age of the last common mitochondrial ancestor continue to be refined. A recent estimate in 2013 from the Max Planck Institute for Evolutionary Anthropology shows that Mitochondrial Eve lived about 160,000 years ago (within the reserved estimate of the original research) and also that the non-African humans were separated from Africans about 95,000 years ago.[51] In August 2013, a study led by Stanford University School of Medicine geneticists reported the age of Mitochondrial Eve to be about 99,000 and 148,000 years, and the Y-MRCA to have lived between 120,000 and 156,000 years ago, based on genome sequencing of 69 people from 9 different populations.[11] This also implies that the discrepancies between the age of "Adam" and "Eve" are reconcilable.[35][36]

Implications of dating and placement of Eve

Further information: Multiregional origin of modern humans and Recent African origin of modern humans

Initially there was a lot of resistance against the Mitochondrial Eve hypothesis. This resistance was due, in part, to the popularity of the multiregional evolution hypothesis amongst some leading paleoanthropologists such as Milford Wolpoff. This prevailing theory held that the evolution of humanity from the beginning of the Pleistocene (2.5 million years ago) to the present day has been within a single, continuous human species, evolving worldwide to modern Homo sapiens. More resistance came from those who argued that there was too little time between Homo erectus and modern Homo sapiens to allow for another new species, and others who argued that for regional evolution from archaic hominin forms into modern ones. Consequently, the finding of a recent maternal ancestor for all humans in Africa was very controversial.

Cann, Stoneking & Wilson (1987)'s placement of a relatively small population of humans in sub-saharan Africa, lent appreciable support for the recent Out of Africa hypothesis. The current concept places between 1,500 and 16,000 effectively interbreeding individuals (census 4,500 to 48,000 individuals) within Tanzania and proximal regions. Later, Tishkoff et al. (2009) using data from many loci (not just mitochondrial DNA) extrapolated that the Angola-Namibia border region near the Atlantic Ocean is likely to be near the geographical point of origin of modern human genetic diversity. In its relatively southern origin proposals, this autosomal study was considered by the authors to be broadly consistent with previous mitochondrial DNA studies, including one by some of the same authors who associated the origins of mitochondrial haplogroups L0 and L1 with "click languages" in southern and eastern Africa.[52]

To some extent the studies have already revealed that the archaic Homo sapiens present in Northwest Africa (Jebel Irhoud) were not likely part of the contiguous modern human population. Better defined is the genetic separation among Neanderthals, Flores hobbit, Java man, and Peking man. In 1999 Krings et al., eliminated problems in molecular clocking postulated by Nei, 1992 when it was found the mtDNA sequence for the same region was substantially different from the MRCA relative to any human sequence. Currently there are 6 fully sequenced Neanderthal mitogenomes, each falling within a genetic cluster less diverse than that for humans, and mitogenome analysis in humans has statistically markedly reduced the TMRCA range so that it no longer overlaps with Neandertal/human split times. Of all the non-African hominids, European archaics most closely resembled humans, indicating a wider genetic divide with other hominids.

Since the multiregional evolution hypothesis revolved around a belief that regional modern human populations evolved in situ in various regions (Europe: Neandertals to Europeans, Asia: Homo erectus to East Asians, Australia: Sumatran erectines to indigenous Australians), these results demonstrated that a pure MREH hypothesis could not explain one important genetic marker.

  • Newsweek reported on Mitochondrial Eve based on the Cann et al. study in January 1988, under a heading of "Scientists Explore a Controversial Theory About Man's Origins". The edition sold a record number of copies.[53]
  • The Seven Daughters of Eve presents the theory of human mitochondrial genetics to a general audience.[54]
  • In River Out of Eden, Richard Dawkins discusses human ancestry in the context of a river of genes and shows that Mitochondrial Eve is one of the many common ancestors we can trace back to via different gene pathways.[55]
  • The Discovery Channel produced a documentary entitled The Real Eve: 6 Billion People ... from ONE Woman (or Where We Came From in the United Kingdom), based on the book The Real Eve: Modern Man's Journey Out of Africa by Stephen Oppenheimer.[56]

See also

Phylogenetic tree of human mitochondrial DNA (mtDNA) haplogroups

  Mitochondrial Eve (L)    
L0 L1–6  
L1 L2   L3     L4 L5 L6
M N  
CZ D E G Q   O A S R   I W X Y
C Z B F R0   pre-JT   P   U
HV JT K
H V J T

References

  1. ^ Pedro Soares et al 2009, Correcting for Purifying Selection: An Improved Human Mitochondrial Molecular Clock. and its Supplemental Data. The American Journal of Human Genetics, Volume 84, Issue 6, 740–759, 4 June 2009
  2. ^ "Jordan: 'Mitochondrial Eve'". weber.ucsd.edu. 2011 [last update]. Retrieved 7 January 2012. {{cite web}}: Check date values in: |year= (help)CS1 maint: year (link)
  3. ^ a b Cann RL, Stoneking M, Wilson AC (1987), "Mitochondrial DNA and human evolution", Nature, 325 (6099): 31–36, doi:10.1038/325031a0, PMID 3025745 {{citation}}: Invalid |ref=harv (help)CS1 maint: multiple names: authors list (link)
  4. ^ Soares P, Ermini L, Thomson N; et al. (2009), "Correcting for purifying selection: an improved human mitochondrial molecular clock", Am. J. Hum. Genet., 84 (6): 740–59, doi:10.1016/j.ajhg.2009.05.001, PMC 2694979, PMID 19500773 {{citation}}: Explicit use of et al. in: |author= (help); Unknown parameter |month= ignored (help)CS1 maint: multiple names: authors list (link). University of Leeds – New 'molecular clock' aids dating of human migration history
  5. ^ 'Your Genetic Journey' – The Genographic Project
  6. ^ Endicott, P; Ho, SY; Metspalu, M; Stringer, C (2009), "Evaluating the mitochondrial timescale of human evolution", Trends Ecol. Evol. (Amst.), 24 (9): 515–21, doi:10.1016/j.tree.2009.04.006, PMID 19682765 {{citation}}: Unknown parameter |month= ignored (help)
  7. ^ a b Dawkins, Richard (2004), The ancestor's tale: a pilgrimage to the dawn of evolution, Boston: Houghton Mifflin, ISBN 0-618-00583-8
  8. ^ Mendez, Fernando; Krahn, Thomas; Schrack, Bonnie; Krahn, Astrid-Maria; Veeramah, Krishna; Woerner, August; Fomine, Forka Leypey Mathew; Bradman, Neil; Thomas, Mark (7 March 2013), "An African American paternal lineage adds an extremely ancient root to the human Y chromosome phylogenetic tree" (PDF), American Journal of Human Genetics, 92 (3): 454, doi:10.1016/j.ajhg.2013.02.002 (primary source)
  9. ^ a b Barrass, Colin (6 March 2013). "The father of all men is 340,000 years old". New Scientist. Retrieved 13 March 2013.
  10. ^ Francalacci P, Morelli L, Angius A, Berutti R, Reinier F, Atzeni R, Pilu R, Busonero F, Maschio A, Zara I, Sanna D, Useli A, Urru MF, Marcelli M, Cusano R, Oppo M, Zoledziewska M, Pitzalis M, Deidda F, Porcu E, Poddie F, Kang HM, Lyons R, Tarrier B, Gresham JB, Li B, Tofanelli S, Alonso S, Dei M, Lai S, Mulas A, Whalen MB, Uzzau S, Jones C, Schlessinger D, Abecasis GR, Sanna S, Sidore C, Cucca F (2013). "Low-pass DNA sequencing of 1200 Sardinians reconstructs European Y-chromosome phylogeny". Science. 341 (6145): 565–569. doi:10.1126/science.1237947. PMID 23908240.{{cite journal}}: CS1 maint: multiple names: authors list (link)
  11. ^ a b c Poznik GD, Henn BM, Yee MC, Sliwerska E, Euskirchen GM, Lin AA, Snyder M, Quintana-Murci L, Kidd JM, Underhill PA, Bustamante CD (2013). "Sequencing Y chromosomes resolves discrepancy in time to common ancestor of males versus females". Science. 341 (6145): 562–565. doi:10.1126/science.1237619. PMID 23908239.{{cite journal}}: CS1 maint: multiple names: authors list (link)
  12. ^ Cann RL (2013). "Genetics. Y weigh in again on modern humans". Science. 341 (6145): 465–467. doi:10.1126/science.1242899. PMID 23908212.
  13. ^ Brown WM, George M Jr, Wilson AC; George; Wilson (1979), "Rapid evolution of animal mitochondrial DNA", Proc Natl Acad Sci U S A, 76 (4): 1967–1971, Bibcode:1979PNAS...76.1967B, doi:10.1073/pnas.76.4.1967, PMC 383514, PMID 109836{{citation}}: CS1 maint: PMC format (link) CS1 maint: multiple names: authors list (link)
  14. ^ Ferris SD, Brown WM, Wilson AC; Wilson; Brown (1981), "Evolutionary tree for apes and humans based on cleavage maps of mitochondrial DNA", Proc Natl Acad Sci U S A, 78 (4): 2432–2436, Bibcode:1981PNAS...78.2432F, doi:10.1073/pnas.78.4.2432, PMC 319360, PMID 6264476{{citation}}: CS1 maint: PMC format (link) CS1 maint: multiple names: authors list (link)
  15. ^ Ferris SD, Brown WM, Wilson AC (1980), "Polymorphism in mitochondrial DNA of humans as revealed by restriction endonuclease analysis", Proc Natl Acad Sci U S A, 77 (6): 2432–2436, PMC 319360, PMID 6264476{{citation}}: CS1 maint: PMC format (link) CS1 maint: multiple names: authors list (link)
  16. ^ Cann RL, Brown WM, Wilson AC (1982), "Evolution of human mitochondrial DNA: a preliminary report", Prog Clin Biol Res, 103 (Pt A): 157–165, PMID 6298804{{citation}}: CS1 maint: multiple names: authors list (link)
  17. ^ Cann RL, Wilson AC (1983), "Length mutations in human mitochondrial DNA", Genetics, 104 (4): 699–711, PMC 1202135, PMID 6311667{{citation}}: CS1 maint: PMC format (link)
  18. ^ a b c Gitschier J (2010), "All about Mitochondrial Eve: An interview with Rebecca Cann", PLoS Genetics, 6 (3): e1000959, doi:10.1371/journal.pgen.1000959, PMC 2877732, PMID 20523888{{citation}}: CS1 maint: PMC format (link) CS1 maint: unflagged free DOI (link)
  19. ^ Denaro M, Blanc H, Johnson MJ, Chen KH, Wilmsen E, Cavalli-Sforza LL, Wallace DC (1981), "Ethnic variation in Hpa 1 endonuclease cleavage patterns of human mitochondrial DNA", Proc Natl Acad Sci U S A, 78 (9): 5768–5772, doi:10.1073/pnas.78.9.5768, PMC 348856, PMID 6272318{{citation}}: CS1 maint: PMC format (link) CS1 maint: multiple names: authors list (link)
  20. ^ Vigilant L, Stoneking M, Harpending H, Hawkes K, Wilson AC (1991), "African populations and the evolution of human mitochondrial DNA", Science, 253 (5027): 1503–1507, doi:10.1126/science.1840702, PMID 1840702 {{citation}}: Invalid |ref=harv (help); Unknown parameter |month= ignored (help)CS1 maint: multiple names: authors list (link)
  21. ^ Johnson NA (2007), Darwinian Detectives: Revealing the Natural History of Genes and Genomes, New York (US): Oxford University Press, Inc, p. 92, ISBN 9780195306750
  22. ^ a b Lewin R (1987), "The unmasking of mitochondrial Eve", Science, 238 (4823): 24–26, doi:10.1126/science.3116666, PMID 3116666
  23. ^ Wainscoat J (1987), "Out of the garden of Eden", Nature, 325 (6099): 13, doi:10.1038/325013a0, PMID 3796736
  24. ^ Wilkins A (27 January 2012). "The scientists behind Mitochondrial Eve tell us about the "lucky mother" who changed human evolution forever". On the Human. Gawker Media 2013. Retrieved 3 May 2013.
  25. ^ Cann RL (1997), "Chapter 4: Mothers, Labels, and Misogyny", in Hager LD (ed.), Women in Human Evolution, London, UK: Routledge, pp. 75–89, ISBN 9780415108331
  26. ^ Tierney J (1992). "The Search for Adam and Eve". Newsweek. Carter G. Woodson Institute for Afro-American and African Studies. Retrieved 13 May 2013.
  27. ^ Lemonick MD (26 January 1987). "Everyone's Genealogical Mother". Time. Time Inc. Retrieved 13 May 2013.
  28. ^ Giles, Richard E (1980), "Maternal inheritance of human mitochondrial DNA", PNAS, 77 (11): 6715–6719, doi:10.1073/pnas.77.11.6715, PMC 350359, PMID 6256757 {{citation}}: Unknown parameter |coauthors= ignored (|author= suggested) (help)
  29. ^ Birky, C. William (2008), "Uniparental inheritance of organelle genes", Current Biology, 18 (16): R692–R695, doi:10.1016/j.cub.2008.06.049, PMID 18727899
  30. ^ Soares, P; Ermini, L; Thomson, N; Mormina, M; Rito, T; Röhl, A; Salas, A; Oppenheimer, S; Macaulay, V (2009), "Correcting for purifying selection: an improved human mitochondrial molecular clock", American Journal of Human Genetics, 84 (6): 740–59, doi:10.1016/j.ajhg.2009.05.001, PMC 2694979, PMID 19500773 {{citation}}: Unknown parameter |month= ignored (help)
  31. ^ Gibbons, A (1998), "Calibrating the Mitochondrial Clock", Science, 279 (5347): 28–29, doi:10.1126/science.279.5347.28, PMID 9441404 {{citation}}: Unknown parameter |month= ignored (help)
  32. ^ There are sites in mtDNA (such as: 16129, 16223, 16311, 16362) that evolve more rapidly, have been noted to change within intragenerational timeframes – Excoffier & Yang (1999).
  33. ^ a b Takahata, N (1993), "Allelic genealogy and human evolution", Mol. Biol. Evol., 10 (1): 2–22, PMID 8450756 {{citation}}: Unknown parameter |month= ignored (help)""
  34. ^ Cruciani, F; Trombetta, B; Massaia, A; Destro-Bisol, G (2011), "A Revised Root for the Human Y Chromosomal Phylogenetic Tree: The Origin of Patrilineal Diversity in Africa", The American Journal of Human Genetics, 88 (6): 814–818, doi:10.1016/j.ajhg.2011.05.002, PMC 3113241, PMID 21601174 {{citation}}: Unknown parameter |coauthors= ignored (|author= suggested) (help); Unknown parameter |month= ignored (help)
  35. ^ a b University of Michigan Health System (1 August 2013). "The when and where of the Y: Research on Y chromosomes uncovers new clues about human ancestry". ScienceDaily. Retrieved 10 August 2013.
  36. ^ a b Stanford University Medical Center (1 August 2013). "Common genetic ancestors lived during roughly same time period, Stanford scientists find". EurekaAlert. AAAS, the science society. Retrieved 10 August 2013.
  37. ^ a b Rohde, DL; Olson, S; Chang, JT (2004), "Modelling the recent common ancestry of all living humans", Nature, 431 (7008): 562–6, doi:10.1038/nature02842, PMID 15457259 {{citation}}: Unknown parameter |month= ignored (help)
  38. ^ Wieland C (2005), "Mitochondrial Eve and biblical Eve are looking good: criticism of young age is premature", Journal of Creation, 19 (1): 57–59
  39. ^ Nelson CW (1 April 2003). "Genetics and Biblical Demographic Events". answersingenesis.org. Answers in Genesis. Retrieved 16 May 2013.
  40. ^ Oakes J. "Can the human arguments about mitochondrial Eve and y-chromosome Adam be extended to the animal world to test the reality of the flood of Noah?". Evidence for Christianity. Answers in Genesis. Retrieved 16 May 2013.
  41. ^ Darlu P, Tassy P (1987). "Disputed African origin of human populations". Nature. 329 (6135): 111–112. doi:10.1038/329111b0. PMID 3114640.
  42. ^ Maddison DR (1991). "African Origin of human mitochondrial DNA reexamined". Systematic Zoology. 40 (3): 355–363. JSTOR 2992327.
  43. ^ Nei M (1992). "Age of the common ancestor of human mitochondrial DNA". Mol Biol Evol. 9 (6): 1176–1178. PMID 1435241.
  44. ^ Ayala FJ (1995). "The myth of Eve: molecular biology and human origins". Science. 270 (5244): 1930–1936. doi:10.1126/science.270.5244.1930. PMID 8533083.
  45. ^ Templeton A (2002). "Out of Africa again and again". Nature. 416 (6876): 45–51. doi:10.1038/416045a. PMID 11882887.
  46. ^ Templeton A (1997). "Out of Africa? What do genes tell us?". Curr Opin Genet Dev. 7 (6): 841–847. doi:10.1016/S0959-437X(97)80049-4. PMID 9468796.
  47. ^ Harrub B, Thompson B (2003). "The Demise of Mitochondrial Eve". Apologetics Press. Apologetics Press, Inc. Retrieved 16 May 2013.
  48. ^ Thompson B, Harrub B (2003). "How Many Times does "Mitochondrial Eve" have to Die?". Apologetics Press. Apologetics Press, Inc. Retrieved 16 May 2013.
  49. ^ Holsinger K (29 September 2012). "(Mathematics of the coalescent) An example: Mitochondrial Eve". Holsinger Lab. Retrieved 16 May 2013.
  50. ^ Cyran KA, Kimmel M (2010). "Alternatives to the Wright-Fisher model: the robustness of mitochondrial Eve dating". Theor Popul Biol. 78 (3): 165–172. doi:10.1016/j.tpb.2010.06.001. PMID 20600209.
  51. ^ Fu Q, Mittnik A, Johnson PLF, 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 (2013). "A revised timescale for human evolution based on ancient mitochondrial genomes". Current Biology. 23 (7): 553–559. doi:10.1016/j.cub.2013.02.044.{{cite journal}}: CS1 maint: multiple names: authors list (link)
  52. ^ Tishkoff et al. (2007); also see Behar et al. (2008) for example.
  53. ^ Oppenheimer, Stephen (2004), The Real Eve: Modern Man's Journey Out of Africa, New York: Carroll & Graf, ISBN 0-7867-1334-8
  54. ^ Bryan Sykes (2002), The Seven Daughters of Eve: The Science That Reveals Our Genetic Ancestry, W. W. Norton & Company, ISBN 978-0-393-32314-6
  55. ^ Richard Dawkins (1995), River out of eden: a Darwinian view of life, Basic Books, ISBN 978-0-465-06990-3, retrieved 5 December 2011
  56. ^ Stephen Oppenheimer (2003), The Real Eve: Modern Man's Journey Out of Africa, New York: Basic Books, ISBN 0-7867-1192-2, retrieved 5 December 2011

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