Mitochondrial Eve
In human genetics, the Mitochondrial Eve (also mt-Eve, mt-MRCA) is the matrilineal most recent common ancestor (MRCA) of all living humans. In other words, she is defined as the most recent woman from whom all living humans descend in an unbroken line purely through their mothers and through the mothers of those mothers, back until all lines converge on one woman.
Haplogroup L | |
---|---|
Possible time of origin | c. 100–230 kya[note 1][note 2] |
Possible place of origin | East Africa |
Ancestor | n/a |
Descendants | Mitochondrial macro-haplogroups L0, L1, and L5 |
Defining mutations | None |
In terms of mitochondrial haplogroups, the mt-MRCA is situated at the divergence of macro-haplogroup L into L0 and L1–6. As of 2013, estimates on the age of this split ranged at around 155,000 years ago,[note 3] consistent with a date later than the speciation of Homo sapiens but earlier than the recent out-of-Africa dispersal.[4][1][5]
The male analog to the "Mitochondrial Eve" is the "Y-chromosomal Adam" (or Y-MRCA), the individual from whom all living humans are patrilineally descended. As the identity of both matrilineal and patrilineal MRCAs is dependent on genealogical history (pedigree collapse), they need not have lived at the same time. As of 2013, estimates for the age Y-MRCA are subject to substantial uncertainty, with a wide range of times from 180,000 to 580,000 years ago[6][7][8] (with an estimated age of between 120,000 and 156,000 years ago, roughly consistent with the estimate for mt-MRCA.).[2][9]
The name "Mitochondrial Eve" alludes to the biblical Eve, which has led to repeated misrepresentations or misconceptions in journalistic accounts on the topic. Popular science presentations of the topic usually point out such possible misconceptions by emphasizing the fact that the position of mt-MRCA is neither fixed in time (as the position of mt-MRCA moves forward in time as mitochondrial DNA (mtDNA) lineages become extinct), nor does it refer to a "first woman", nor the only living female of her time, nor the first member of a "new species".[note 4]
History
−10 — – −9 — – −8 — – −7 — – −6 — – −5 — – −4 — – −3 — – −2 — – −1 — – 0 — |
| |||||||||||||||||||||
Early research
Early research using molecular clock methods was done during the late 1970s to early 1980s. 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.[11] Related work allowed for an analysis of the evolutionary relationships among gorillas, chimpanzees (common chimpanzee and bonobo) and humans.[12] 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.[13] 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).[14][15][16]
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 !Kung respectively, were available. After more than 40 revisions of the draft, the manuscript was submitted to Nature in late 1985 or early 1986[16] 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.[17]
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 origin model.[18]
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.[19] It also appears in the October 1987 article in Science by Roger Lewin, headlined "The Unmasking of Mitochondrial Eve."[20] The biblical connotation was very clear from the start. The accompanying research news in Nature had the title "Out of the garden of Eden."[21]
Wilson himself preferred the term "Lucky Mother"[22] and thought the use of the name Eve "regrettable."[20][23] 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"),[24] and a cover story in Time on 26 January 1987.[25]
Criticism and later research
Shortly after the 1987 publication, criticism of its methodology and secondary conclusions was published.[26] 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;[27][28][29][30] 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."[31]
Cann, Stoneking & Wilson (1987)'s placement 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)[32] when it was found that the mtDNA sequence for the same region was substantially different from the MRCA relative to any human sequence.[33][34]
In 1997, Parsons et al. (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.[35]
Although the original research did have analytical limitations, the estimate on the age of the mt-MRCA has proven robust.[36][37] More recent age estimates have remained consistent with the 140–200 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.[3] 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.[2]
Female and mitochondrial ancestry
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, the DNA located in mitochondria, different from the one in the cells nucleus) 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.[38][39] Matrilineal descent goes back through 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 per nucleotide.[1][40][note 5] 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 (by producing only non-matrilinear descendants at that time), 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: first and most obviously, the time back to her, but second 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.
Popular reception and misconceptions
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.[41]
The popular name "mitochondrial Eve", of 1980s coinage,[20] has contributed to a number of popular misconceptions. At first, the announcement of a "mitochondrial Eve" was even greeted with endorsement from young earth creationists, who viewed the theory as a validation of the biblical creation story.[42][43][44]
Due to such misunderstandings, authors of popular science publications since the 1990s have been emphatic in pointing out that the name is merely a popular convention, and that the mt-MRCA was not in any way the "first woman".[45] Her position is purely the result of genealogical history of human populations later, and as matrilineal lineages die out, the position of mt-MRCA keeps moving forward to younger individuals over time.
In River Out of Eden (1995), Richard Dawkins discussed human ancestry in the context of a "river of genes", including an explanation of the concept of Mitochondrial Eve.[46] The Seven Daughters of Eve (2002) presented the topic of human mitochondrial genetics to a general audience.[47] The Real Eve: Modern Man's Journey Out of Africa by Stephen Oppenheimer (2003)[41] was adapted into a Discovery Channel documentary.[48]
Not the only woman
One common 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.[45] However, nuclear DNA studies indicate that the effective population size of the ancient human never dropped below tens of thousands.[49] Other women living during Eve's time may have descendants alive today but not in a direct female line.[50]
Not a fixed individual over time
The definition of Mitochondrial Eve is fixed, but the woman in prehistory who fits this definition can change. That is, not only can our knowledge of when and where Mitochondrial Eve lived change due to new discoveries, but the actual Mitochondrial Eve can change. The Mitochondrial Eve can change, when a mother-daughter line comes to an end. 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. In every generation mitochondrial lineages end – when a woman with unique mtDNA dies with no daughters. 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 two or more daughters who together have all living humans as their 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.[51]
Because mtDNA mapping of humans is very incomplete, the discovery of living mtDNA lines which predate our current concept of "Mitochondrial Eve" could result in the title moving to an earlier woman. This happened to her male counterpart, "Y-chromosomal Adam," when an older Y line, haplogroup A-00, was discovered.[52]
Not necessarily a contemporary of "Y-chromosomal Adam"
Sometimes Mitochondrial Eve is assumed to have lived at the same time as Y-chromosomal Adam (from whom all living males are descended patrilineally), and perhaps even met and mated 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. A recent study (March 2013) concluded however that "Eve" lived much later than "Adam" – some 140,000 years later.[7] (Earlier studies considered, conversely, that "Eve" lived earlier than "Adam".)[53] More recent studies indicate that Mitochondrial Eve and Y-chromosomal Adam may indeed have lived around the same time.[54]
Not the most recent ancestor shared by all humans
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. The time period that human MRCA lived is unknown. Rohde et. al put forth a "rough guess" that the MRCA could have existed 5000 years ago; however, the authors state that this estimate is "extremely tentative, and the model contains several obvious sources of error, as it was motivated more by considerations of theoretical insight and tractability than by realism."[55] 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 of all humans alive today. However, such a late date is difficult to reconcile with the geographical spread of our species and the consequent isolation of different groups from each other. For example, it is generally accepted that the indigenous population of Tasmania was isolated from all other humans between the rise in sea level after the last ice age some 8000 years ago and the arrival of Europeans. Estimates of the MRCA of even closely related human populations have been much more than 5000 years ago.[56]
- The identical ancestors point. 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 when Mitochondrial Eve was proposed to have lived.[55]
- 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.
See also
- Archaeogenetics
- Coalescent theory
- Eurasian Adam
- Genealogical DNA test
- Genetic genealogy
- Haplogroup L0 (mtDNA)
- Human evolution
- Human mitochondrial DNA haplogroups
- Last universal ancestor
- Macro-haplogroup L (mtDNA)
- Mitochondrial genome
- Monogenism
- Neutral theory of molecular evolution
- Prehistoric demography
- Single origin hypothesis
- Timeline of evolution
- Timeline of human evolution
- Y-chromosomal Aaron
- Y-chromosomal Adam
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 |
Notes
- "the synonymous mutation rate of Kivisild et al. [...] estimates the coalescence time of the mtDNA tree overall at ~160,000 kya [...] We present a revised chronology using the complete mtDNA genome rate and an ML approach for the mtDNA tree in Figure 6, with full details of the age estimates and associated 95% confidence regions in Table S5."[1] See: Supplemental Data.
- "we estimate the time to the most recent common ancestor (TMRCA) of the Y chromosome to be 120 to 156 thousand years and the mitochondrial genome TMRCA to be 99 to 148 thousand years. Our findings suggest that, contrary to previous claims, male lineages do not coalesce significantly more recently than female lineages."[2]
- Two studies published in 2013 had 95% confidence intervals barely overlapping in the neighbourhood of 15 ka, a third study had a 95% confidence interval intermediate between the two others: "99 to 148 ka" according to Poznik, 2013[2] (ML whole-mtDNA age estimate: 178.8 [155.6; 202.2], ρ whole-mtDNA age estimate: 185.2 [153.8; 216.9], ρ synonymous age estimate: 174.8 [153.8; 216.9]), "134 to 188 ka", according to Fu, 2013,[3] and 150 to 234 ka (95% CI) from Soares, 2009.[1]
- "Caution: This does not make Mitochondrial Eve the first woman, or the first human, or the first member of a new species. Further Caution: This does not mean that other women alive when Eve was do not have descendants today; they simply do not have living descendants who are descended only through female links. Yet Further Caution: If a person were to be discovered whose mtDNA showed a pattern of mutations of greater time depth, then the status of Mitochondrial Eve would be reassigned to the most recent female ancestor shared by both that person and the person we now call Mitochondrial Eve."[10]
- 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).
References
- Soares P, Ermini L, Thomson N, Mormina M, Rito T, Röhl A, et al. (June 2009). "Correcting for purifying selection: an improved human mitochondrial molecular clock". American Journal of Human Genetics. 84 (6): 740–759. doi:10.1016/j.ajhg.2009.05.001. PMC 2694979. PMID 19500773.
- Poznik GD, Henn BM, Yee MC, Sliwerska E, Euskirchen GM, Lin AA, et al. (August 2013). "Sequencing Y chromosomes resolves discrepancy in time to common ancestor of males versus females". Science. 341 (6145): 562–565. Bibcode:2013Sci...341..562P. doi:10.1126/science.1237619. PMC 4032117. PMID 23908239.
- Fu Q, Mittnik A, Johnson PL, Bos K, Lari M, Bollongino R, et al. (April 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. PMC 5036973. PMID 23523248.
- Endicott P, Ho SY, Metspalu M, Stringer C (September 2009). "Evaluating the mitochondrial timescale of human evolution". Trends in Ecology & Evolution. 24 (9): 515–521. doi:10.1016/j.tree.2009.04.006. PMID 19682765.
- "New 'molecular clock' aids dating of human migration history". University of Leeds. 3 June 2009. Retrieved 23 December 2019.
- Mendez FL, Krahn T, Schrack B, Krahn AM, Veeramah KR, Woerner AE, et al. (March 2013). "An African American paternal lineage adds an extremely ancient root to the human Y chromosome phylogenetic tree". American Journal of Human Genetics. 92 (3): 454–459. doi:10.1016/j.ajhg.2013.02.002. PMC 3591855. PMID 23453668. (95% confidence interval 237–581 kya)
- Barras C (6 March 2013). "The father of all men is 340,000 years old". New Scientist. Retrieved 13 March 2013.
- Francalacci P, Morelli L, Angius A, Berutti R, Reinier F, Atzeni R, et al. (August 2013). "Low-pass DNA sequencing of 1200 Sardinians reconstructs European Y-chromosome phylogeny". Science. 341 (6145): 565–569. Bibcode:2013Sci...341..565F. doi:10.1126/science.1237947. PMC 5500864. PMID 23908240.
- Cann RL (August 2013). "Genetics. Y weigh in again on modern humans". Science. 341 (6145): 465–467. Bibcode:2013Sci...341..465C. doi:10.1126/science.1242899. PMID 23908212. S2CID 206550892.
- "Jordan: 'Mitochondrial Eve'". weber.ucsd.edu. 2011. Retrieved 7 January 2012.
- Brown WM, George M, Wilson AC (April 1979). "Rapid evolution of animal mitochondrial DNA". Proceedings of the National Academy of Sciences of the United States of America. 76 (4): 1967–1971. Bibcode:1979PNAS...76.1967B. doi:10.1073/pnas.76.4.1967. PMC 383514. PMID 109836.
- Ferris SD, Wilson AC, Brown WM (April 1981). "Evolutionary tree for apes and humans based on cleavage maps of mitochondrial DNA". Proceedings of the National Academy of Sciences of the United States of America. 78 (4): 2432–2436. Bibcode:1981PNAS...78.2432F. doi:10.1073/pnas.78.4.2432. PMC 319360. PMID 6264476.
- Brown WM (June 1980). "Polymorphism in mitochondrial DNA of humans as revealed by restriction endonuclease analysis". Proceedings of the National Academy of Sciences of the United States of America. 77 (6): 3605–3609. Bibcode:1980PNAS...77.3605B. doi:10.1073/pnas.77.6.3605. PMC 349666. PMID 6251473.
- Cann RL, Brown WM, Wilson AC (1982). "Evolution of human mitochondrial DNA: a preliminary report". Progress in Clinical and Biological Research. 103 Pt A (Pt A): 157–165. PMID 6298804.
- Cann RL, Wilson AC (August 1983). "Length mutations in human mitochondrial DNA". Genetics. 104 (4): 699–711. doi:10.1093/genetics/104.4.699. PMC 1202135. PMID 6311667.
- Cann R (May 2010). "All about mitochondrial eve: an interview with Rebecca Cann. Interview by Jane Gitschier". PLOS Genetics. 6 (5): e1000959. doi:10.1371/journal.pgen.1000959. PMC 2877732. PMID 20523888.
- Cann RL, Stoneking M, Wilson AC (1987). "Mitochondrial DNA and human evolution". Nature. 325 (6099): 31–36. Bibcode:1987Natur.325...31C. doi:10.1038/325031a0. PMID 3025745. S2CID 4285418.
- Vigilant L, Stoneking M, Harpending H, Hawkes K, Wilson AC (September 1991). "African populations and the evolution of human mitochondrial DNA". Science. 253 (5027): 1503–1507. Bibcode:1991Sci...253.1503V. doi:10.1126/science.1840702. PMID 1840702.
- Cann RL (1987). "In Search of Eve". The Sciences. 27 (5): 30–37. doi:10.1002/j.2326-1951.1987.tb02967.x. ISSN 2326-1951.
- Lewin R (October 1987). "The unmasking of mitochondrial Eve". Science. 238 (4823): 24–26. Bibcode:1987Sci...238...24L. doi:10.1126/science.3116666. PMID 3116666.
- Wainscoat J (1987). "Human evolution. Out of the garden of Eden". Nature. 325 (6099): 13. Bibcode:1987Natur.325...13W. doi:10.1038/325013a0. PMID 3796736. S2CID 13187170.
- Wilkins A (27 January 2012). "The scientists behind Mitochondrial Eve tell us about the 'lucky mother' who changed human evolution forever". Gizmodo. Retrieved 23 December 2019.
- Cann RL (1997). "Chapter 4: Mothers, Labels, and Misogyny". In Hager LD (ed.). Women in Human Evolution. London: Routledge. pp. 75–89. ISBN 9780415108331.
- Tierney J (1992). "The Search for Adam and Eve". Newsweek. Carter G. Woodson Institute for Afro-American and African Studies via Internet Archive. Archived from the original on 20 March 2015. Retrieved 21 July 2019.
- Lemonick MD (26 January 1987). "Everyone's Genealogical Mother". Time. Archived from the original on 23 September 2008. Retrieved 13 May 2013.
- Darlu P, Tassy P (1987). "Disputed African origin of human populations". Nature. 329 (6135): 111–112. Bibcode:1987Natur.329..111D. doi:10.1038/329111b0. PMID 3114640. S2CID 4313392.
- Maddison DR (1991). "African Origin of human mitochondrial DNA reexamined". Systematic Zoology. 40 (3): 355–63. doi:10.2307/2992327. JSTOR 2992327.
- Nei M (November 1992). "Age of the common ancestor of human mitochondrial DNA". Molecular Biology and Evolution. 9 (6): 1176–1178. doi:10.1093/oxfordjournals.molbev.a040785. PMID 1435241.
- Ayala FJ (December 1995). "The myth of Eve: molecular biology and human origins". Science. 270 (5244): 1930–1936. Bibcode:1995Sci...270.1930A. doi:10.1126/science.270.5244.1930. PMID 8533083. S2CID 42801341.
- Templeton A (March 2002). "Out of Africa again and again". Nature. 416 (6876): 45–51. Bibcode:2002Natur.416...45T. doi:10.1038/416045a. PMID 11882887. S2CID 4397398.
- Templeton AR (December 1997). "Out of Africa? What do genes tell us?". Current Opinion in Genetics & Development. 7 (6): 841–847. doi:10.1016/S0959-437X(97)80049-4. PMID 9468796.
- Nei M (1992). "Age of the common ancestor of human mitochondrial DNA". Molecular Biology and Evolution. 9 (6): 1176–1178. doi:10.1093/oxfordjournals.molbev.a040785. PMID 1435241.
- Krings M, Salem AE, Bauer K, Geisert H, Malek AK, Chaix L, et al. (April 1999). "mtDNA analysis of Nile River Valley populations: A genetic corridor or a barrier to migration?". American Journal of Human Genetics. 64 (4): 1166–1176. doi:10.1086/302314. PMC 1377841. PMID 10090902.
- Krings M, Stone A, Schmitz RW, Krainitzki H, Stoneking M, Pääbo S (July 1997). "Neandertal DNA sequences and the origin of modern humans". Cell. 90 (1): 19–30. doi:10.1016/S0092-8674(00)80310-4. PMID 9230299. S2CID 13581775.
- Parsons TJ, Muniec DS, Sullivan K, Woodyatt N, Alliston-Greiner R, Wilson MR, et al. (April 1997). "A high observed substitution rate in the human mitochondrial DNA control region". Nature Genetics. 15 (4): 363–368. doi:10.1038/ng0497-363. PMID 9090380. S2CID 32812244.
- Holsinger K (29 September 2012). "(Mathematics of the coalescent) An example: Mitochondrial Eve". Holsinger Lab. Retrieved 16 May 2013.
- Cyran KA, Kimmel M (November 2010). "Alternatives to the Wright-Fisher model: the robustness of mitochondrial Eve dating". Theoretical Population Biology. 78 (3): 165–172. doi:10.1016/j.tpb.2010.06.001. PMID 20600209.
- Giles RE, Blanc H, Cann HM, Wallace DC (November 1980). "Maternal inheritance of human mitochondrial DNA". Proceedings of the National Academy of Sciences of the United States of America. 77 (11): 6715–6719. Bibcode:1980PNAS...77.6715G. doi:10.1073/pnas.77.11.6715. PMC 350359. PMID 6256757.
- Birky CW (August 2008). "Uniparental inheritance of organelle genes". Current Biology. 18 (16): R692–R695. doi:10.1016/j.cub.2008.06.049. PMID 18727899. S2CID 9866662.
- Gibbons A (January 1998). "Calibrating the mitochondrial clock". Science. 279 (5347): 28–29. Bibcode:1998Sci...279...28G. doi:10.1126/science.279.5347.28. PMID 9441404. S2CID 29855766.
- Oppenheimer S (2003). The Real Eve: Modern Man's Journey Out of Africa. New York: Basic Books. ISBN 978-0-7867-1192-5.
- Wieland C (2005). "Mitochondrial Eve and biblical Eve are looking good: criticism of young age is premature". Journal of Creation. 19 (1): 57–59.
- Nelson CW (1 April 2003). "Genetics and Biblical Demographic Events". answersingenesis.org. Answers in Genesis. Retrieved 16 May 2013.
- 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.
- Dawkins R (2004). The ancestor's tale: a pilgrimage to the dawn of evolution. Boston: Houghton Mifflin. ISBN 978-0-618-00583-3.
- Dawkins R (1995). River out of eden: a Darwinian view of life. Basic Books. ISBN 978-0-465-06990-3. Retrieved 5 December 2011.
- Sykes B (2002). The Seven Daughters of Eve: The Science That Reveals Our Genetic Ancestry. W. W. Norton & Company. ISBN 978-0-393-32314-6.
- Oppenheimer, Stephen (2003). The Real Eve: Modern Man's Journey Out of Africa. Carroll & Graf. ISBN 978-0-7867-1192-5.
- Takahata N (January 1993). "Allelic genealogy and human evolution". Molecular Biology and Evolution. 10 (1): 2–22. doi:10.1093/oxfordjournals.molbev.a039995. PMID 8450756.
- "Are we all descended from a common female ancestor?". HowStuffWorks. 14 January 2008. Retrieved 17 May 2022.
- Learn JP. "No, a Mitochondrial "Eve" Is Not the First Female in a Species". Smithsonian Magazine. Retrieved 23 February 2021.
- Mendez FL, Krahn T, Schrack B, Krahn AM, Veeramah KR, Woerner AE, et al. (March 2013). "An African American paternal lineage adds an extremely ancient root to the human Y chromosome phylogenetic tree". American Journal of Human Genetics. 92 (3): 454–459. doi:10.1016/j.ajhg.2013.02.002. PMC 3591855. PMID 23453668.
- Cruciani F, Trombetta B, Massaia A, Destro-Bisol G, Sellitto D, Scozzari R (June 2011). "A revised root for the human Y chromosomal phylogenetic tree: the origin of patrilineal diversity in Africa". American Journal of Human Genetics. 88 (6): 814–818. doi:10.1016/j.ajhg.2011.05.002. PMC 3113241. PMID 21601174.
- Callaway E (6 August 2013). "Genetic Adam and Eve did not live too far apart in time". Nature. doi:10.1038/nature.2013.13478. S2CID 170608686.
- Rohde DL, Olson S, Chang JT (September 2004). "Modelling the recent common ancestry of all living humans". Nature. 431 (7008): 562–566. Bibcode:2004Natur.431..562R. CiteSeerX 10.1.1.78.8467. doi:10.1038/nature02842. PMID 15457259. S2CID 3563900.
- Zhou, Jin; Teo, Yik-Ying (August 2016). "Estimating time to the most recent common ancestor (TMRCA): comparison and application of eight methods". European Journal of Human Genetics. 24 (8): 1195–1201. doi:10.1038/ejhg.2015.258. ISSN 1476-5438. PMC 4970674. PMID 26669663. S2CID 965600.
Further reading
- Atkinson QD, Gray RD, Drummond AJ (January 2009). "Bayesian coalescent inference of major human mitochondrial DNA haplogroup expansions in Africa". Proceedings. Biological Sciences. 276 (1655): 367–373. doi:10.1098/rspb.2008.0785. PMC 2674340. PMID 18826938.
- Ayala FJ (December 1995). "The myth of Eve: molecular biology and human origins". Science. 270 (5244): 1930–1936. Bibcode:1995Sci...270.1930A. doi:10.1126/science.270.5244.1930. PMID 8533083.
- Balloux F, Handley LJ, Jombart T, Liu H, Manica A (October 2009). "Climate shaped the worldwide distribution of human mitochondrial DNA sequence variation". Proceedings. Biological Sciences. 276 (1672): 3447–3455. doi:10.1098/rspb.2009.0752. PMC 2817182. PMID 19586946.
- Behar DM, Villems R, Soodyall H, Blue-Smith J, Pereira L, Metspalu E, et al. (May 2008). "The dawn of human matrilineal diversity". American Journal of Human Genetics. 82 (5): 1130–1140. doi:10.1016/j.ajhg.2008.04.002. PMC 2427203. PMID 18439549.
- Brown WM (June 1980). "Polymorphism in mitochondrial DNA of humans as revealed by restriction endonuclease analysis". Proceedings of the National Academy of Sciences of the United States of America. 77 (6): 3605–3609. Bibcode:1980PNAS...77.3605B. doi:10.1073/pnas.77.6.3605. PMC 349666. PMID 6251473.
- Cox MP (August 2008). "Accuracy of molecular dating with the rho statistic: deviations from coalescent expectations under a range of demographic models". Human Biology. 80 (4): 335–357. doi:10.3378/1534-6617-80.4.335. PMID 19317593. S2CID 207701422.
- Dawkins R (2004). The ancestor's tale: a pilgrimage to the dawn of evolution. Boston: Houghton Mifflin. ISBN 978-0-618-00583-3.
- Endicott P, Ho SY (April 2008). "A Bayesian evaluation of human mitochondrial substitution rates". American Journal of Human Genetics. 82 (4): 895–902. doi:10.1016/j.ajhg.2008.01.019. PMC 2427281. PMID 18371929.
- Endicott P, Ho SY, Metspalu M, Stringer C (September 2009). "Evaluating the mitochondrial timescale of human evolution". Trends in Ecology & Evolution. 24 (9): 515–521. doi:10.1016/j.tree.2009.04.006. PMID 19682765.
- Excoffier L, Yang Z (October 1999). "Substitution rate variation among sites in mitochondrial hypervariable region I of humans and chimpanzees". Molecular Biology and Evolution. 16 (10): 1357–1368. doi:10.1093/oxfordjournals.molbev.a026046. PMID 10563016.
- Felsenstein J (April 1992). "Estimating effective population size from samples of sequences: inefficiency of pairwise and segregating sites as compared to phylogenetic estimates". Genetical Research. 59 (2): 139–147. doi:10.1017/S0016672300030354. PMID 1628818.
- Ferris SD, Brown WM, Davidson WS, Wilson AC (October 1981). "Extensive polymorphism in the mitochondrial DNA of apes". Proceedings of the National Academy of Sciences of the United States of America. 78 (10): 6319–6323. Bibcode:1981PNAS...78.6319F. doi:10.1073/pnas.78.10.6319. PMC 349030. PMID 6273863.
- Gibbons A (January 1998). "Calibrating the mitochondrial clock". Science. 279 (5347): 28–29. Bibcode:1998Sci...279...28G. doi:10.1126/science.279.5347.28. PMID 9441404. S2CID 29855766.
- Gonder MK, Mortensen HM, Reed FA, de Sousa A, Tishkoff SA (March 2007). "Whole-mtDNA genome sequence analysis of ancient African lineages". Molecular Biology and Evolution. 24 (3): 757–768. doi:10.1093/molbev/msl209. PMID 17194802.
- Hazelwood L, Steele J (2004). "Spatial dynamics of human dispersals: Constraints on modelling and archaeological validation". Journal of Archaeological Science. 31 (6): 669–79. doi:10.1016/j.jas.2003.11.009.
- Ho SY, Larson G (February 2006). "Molecular clocks: when times are a-changin'". Trends in Genetics. 22 (2): 79–83. doi:10.1016/j.tig.2005.11.006. PMID 16356585.
- Ingman M, Kaessmann H, Pääbo S, Gyllensten U (December 2000). "Mitochondrial genome variation and the origin of modern humans". Nature. 408 (6813): 708–713. Bibcode:2000Natur.408..708I. doi:10.1038/35047064. PMID 11130070. S2CID 52850476.
- Kaessmann H, Pääbo S (January 2002). "The genetical history of humans and the great apes". Journal of Internal Medicine. 251 (1): 1–18. doi:10.1046/j.1365-2796.2002.00907.x. PMID 11851860. S2CID 7453672.
- Kimura M, Ota T (2001). "Theoretical aspects of population genetics". Monographs in Population Biology. Princeton University Press. 4: 1–219. ISBN 978-0-691-08098-7. PMID 5162676.
- Loewe L, Scherer S (November 1997). "Mitochondrial Eve: the plot thickens". Trends in Ecology & Evolution. 12 (11): 422–423. doi:10.1016/S0169-5347(97)01204-4. PMID 21238138.
- Loogväli EL, Kivisild T, Margus T, Villems R (December 2009). O'Rourke D (ed.). "Explaining the imperfection of the molecular clock of hominid mitochondria". PLOS ONE. 4 (12): e8260. Bibcode:2009PLoSO...4.8260L. doi:10.1371/journal.pone.0008260. PMC 2794369. PMID 20041137.
- Maca-Meyer N, González AM, Larruga JM, Flores C, Cabrera VM (2001). "Major genomic mitochondrial lineages delineate early human expansions". BMC Genetics. 2 (1): 13. doi:10.1186/1471-2156-2-13. PMC 55343. PMID 11553319.
- Mishmar D, Ruiz-Pesini E, Golik P, Macaulay V, Clark AG, Hosseini S, et al. (January 2003). "Natural selection shaped regional mtDNA variation in humans". Proceedings of the National Academy of Sciences of the United States of America. 100 (1): 171–176. Bibcode:2003PNAS..100..171M. doi:10.1073/pnas.0136972100. PMC 140917. PMID 12509511.
- Nei M (November 1992). "Age of the common ancestor of human mitochondrial DNA". Molecular Biology and Evolution. 9 (6): 1176–1178. doi:10.1093/oxfordjournals.molbev.a040785. PMID 1435241.
- Nielsen R, Beaumont MA (March 2009). "Statistical inferences in phylogeography". Molecular Ecology. 18 (6): 1034–1047. doi:10.1111/j.1365-294X.2008.04059.x. PMID 19207258. S2CID 13613087.
- Oppenheimer S (2004). The Real Eve: Modern Man's Journey Out of Africa. New York: Carroll & Graf. ISBN 978-0-7867-1334-9.
- Pritchard JK, Seielstad MT, Perez-Lezaun A, Feldman MW (December 1999). "Population growth of human Y chromosomes: a study of Y chromosome microsatellites". Molecular Biology and Evolution. 16 (12): 1791–1798. doi:10.1093/oxfordjournals.molbev.a026091. PMID 10605120.
- Reed FA, Tishkoff SA (December 2006). "African human diversity, origins and migrations". Current Opinion in Genetics & Development. 16 (6): 597–605. doi:10.1016/j.gde.2006.10.008. PMID 17056248.
- Rohde DL, Olson S, Chang JT (September 2004). "Modelling the recent common ancestry of all living humans". Nature. 431 (7008): 562–566. Bibcode:2004Natur.431..562R. CiteSeerX 10.1.1.78.8467. doi:10.1038/nature02842. PMID 15457259. S2CID 3563900.
- Santos C, Sierra B, Alvarez L, Ramos A, Fernández E, Nogués R, Aluja MP (August 2008). "Frequency and pattern of heteroplasmy in the control region of human mitochondrial DNA". Journal of Molecular Evolution. 67 (2): 191–200. Bibcode:2008JMolE..67..191S. doi:10.1007/s00239-008-9138-9. PMID 18618067. S2CID 1143395.
- Schaffner SF (January 2004). "The X chromosome in population genetics". Nature Reviews. Genetics. 5 (1): 43–51. doi:10.1038/nrg1247. PMID 14708015. S2CID 15772901.
- Soares P, Ermini L, Thomson N, Mormina M, Rito T, Röhl A, et al. (June 2009). "Correcting for purifying selection: an improved human mitochondrial molecular clock". American Journal of Human Genetics. 84 (6): 740–759. doi:10.1016/j.ajhg.2009.05.001. PMC 2694979. PMID 19500773.
- Suissa S, Wang Z, Poole J, Wittkopp S, Feder J, Shutt TE, et al. (May 2009). Desalle R (ed.). "Ancient mtDNA genetic variants modulate mtDNA transcription and replication". PLOS Genetics. 5 (5): e1000474. doi:10.1371/journal.pgen.1000474. PMC 2673036. PMID 19424428.
- Sykes B (2001). The Seven Daughters of Eve. New York: Norton. ISBN 978-0-393-02018-2.
- Sykes BD (2003). Adam's curse: a future without men. London: Bantam. ISBN 978-0-593-05005-7.
- Takahata N (January 1993). "Allelic genealogy and human evolution". Molecular Biology and Evolution. 10 (1): 2–22. doi:10.1093/oxfordjournals.molbev.a039995. PMID 8450756.
- Takahata N, Lee SH, Satta Y (February 2001). "Testing multiregionality of modern human origins". Molecular Biology and Evolution. 18 (2): 172–183. doi:10.1093/oxfordjournals.molbev.a003791. PMID 11158376.
- Tamura K, Nei M (May 1993). "Estimation of the number of nucleotide substitutions in the control region of mitochondrial DNA in humans and chimpanzees". Molecular Biology and Evolution. 10 (3): 512–526. doi:10.1093/oxfordjournals.molbev.a040023. PMID 8336541.
- Tang H, Siegmund DO, Shen P, Oefner PJ, Feldman MW (May 2002). "Frequentist estimation of coalescence times from nucleotide sequence data using a tree-based partition". Genetics. 161 (1): 447–459. doi:10.1093/genetics/161.1.447. PMC 1462078. PMID 12019257.
- Tishkoff SA, Gonder MK, Henn BM, Mortensen H, Knight A, Gignoux C, et al. (October 2007). "History of click-speaking populations of Africa inferred from mtDNA and Y chromosome genetic variation". Molecular Biology and Evolution. 24 (10): 2180–2195. doi:10.1093/molbev/msm155. PMID 17656633.
- Tishkoff SA, Reed FA, Friedlaender FR, Ehret C, Ranciaro A, Froment A, et al. (May 2009). "The genetic structure and history of Africans and African Americans". Science. 324 (5930): 1035–1044. Bibcode:2009Sci...324.1035T. doi:10.1126/science.1172257. PMC 2947357. PMID 19407144.
- Vigilant L, Pennington R, Harpending H, Kocher TD, Wilson AC (December 1989). "Mitochondrial DNA sequences in single hairs from a southern African population". Proceedings of the National Academy of Sciences of the United States of America. 86 (23): 9350–9354. Bibcode:1989PNAS...86.9350V. doi:10.1073/pnas.86.23.9350. PMC 298493. PMID 2594772.
- Vigilant L, Stoneking M, Harpending H, Hawkes K, Wilson AC (September 1991). "African populations and the evolution of human mitochondrial DNA". Science. 253 (5027): 1503–1507. Bibcode:1991Sci...253.1503V. doi:10.1126/science.1840702. PMID 1840702.
- Watson E, Forster P, Richards M, Bandelt HJ (September 1997). "Mitochondrial footprints of human expansions in Africa". American Journal of Human Genetics. 61 (3): 691–704. doi:10.1086/515503. PMC 1715955. PMID 9326335.
- White TD, Asfaw B, Beyene Y, Haile-Selassie Y, Lovejoy CO, Suwa G, WoldeGabriel G (October 2009). "Ardipithecus ramidus and the paleobiology of early hominids". Science. 326 (5949): 75–86. Bibcode:2009Sci...326...75W. doi:10.1126/science.1175802. PMID 19810190. S2CID 20189444.
- Wilder JA, Mobasher Z, Hammer MF (November 2004). "Genetic evidence for unequal effective population sizes of human females and males". Molecular Biology and Evolution. 21 (11): 2047–2057. doi:10.1093/molbev/msh214. PMID 15317874.
- Wilson AC, Cann RL, Carr SM, George M, Gyllensten UB, Helm-Bychowski KM, et al. (1985). "Mitochondrial DNA and two perspectives on evolutionary genetics". Biological Journal of the Linnean Society. 26 (4): 375–400. doi:10.1111/j.1095-8312.1985.tb02048.x.