Mitochondrial Eve

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Mitochondrial Eve (mtMRCA) is the name given by researchers to the woman who is defined as the matrilineal most recent common ancestor (MRCA) for all living humans. Passed down from mother to offspring, all mitochondrial DNA (mtDNA) in every living person is directly descended from hers. Mitochondrial Eve is the female counterpart of Y-chromosomal Adam, the patrilineal most recent common ancestor, although they lived thousands of years apart.

There are various estimates given for when Mitochondrial Eve lived, ranging between 234,000 years ago and 82,000 years before present (BP), with the majority of estimates clustered between 200,000 and 160,000 BP.^[1][2] She is believed to have lived within the East African region around Tanzania, within a population with about 3000 to 33000 other females, many of whom will also have been our non-matrilineal ancestors.^[3][4] Mitochondrial Eve lived during a period of time when Homo sapiens sapiens were developing as a species separate from other human species.

Mitochondrial Eve lived much earlier than the out of Africa migration that is thought to have occurred between 95,000 to 45,000 BP.^[5] The dating for 'Eve' was a blow to the multiregional hypothesis, and a boost to the hypothesis that modern humans originated relatively recently in Africa and spread from there, replacing more "archaic" human populations such as Neanderthals. As a result, the latter hypothesis is now the dominant one.

[edit] Female and mitochondrial ancestry

Through random drift or selection the female-lineage may trace back to a single female, such as Mitochondrial Eve

Without a DNA sample, it is not possible to reconstruct the complete genetic makeup (genome) of any individual who died very long ago. By looking at descendant's DNA, however, parts of ancestral genomes are estimated by scientists. Mitochondrial DNA (mtDNA) and Y chromosome 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. 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 a 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 Eve's 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.

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 which survive at least until the next generation approximately once every 3624 years.^[6][7][8] 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 branches dies out, the MRCA will move to a more recent female ancestor, the first branch point in the surviving branch. The amount of mutations which 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.

[edit] Estimating time to MRCA

Examining the mtDNA family tree of modern humans reveals two deep branches that converge. Between the two deepest nodes (the modal haplotypes of L0 and L1'2'5) a collection of genetic markers exists that can be arranged to form possible sequences of the MRCA, the MRCA haplotype. The haplotype cannot be precisely determined, even with information added from other apes' mtDNA or the recent Neanderthal mtDNA sequences. The L0 and L1'2'5 haplogroups evolved from one of these possible haplotypes. Therefore, based on this convergence of lineages and matrilineality one female had at least two female children. This female would be the mitochondrial MRCA, 'Eve'.^[9]

To determine the optimal sequence representation of the MRCA, maximum parsimony analysis is employed to best position the branches of the mtDNA family tree.^[10] Furthermore bootstrap based methods are employed to best find the MRCA.^[3] Once the optimal placements of L0 and L1'2'5 nodes are determined, then the average depth of genetic diversity (measured in mutations along the branches) within each haplogroup can be determined. Adding these two average distances together with the genetic distance between L0 and L1'2'5 nodes gives a distance that is twice the average distance of any living human to the putative MRCA sequence.

Many mutations occur in mitochondrial DNA, and most never make it past a few generations.^[11] Some mutations that are mildly disadvantageous persist for a time as a consequence of variable or relaxed selection. Such mutations are largely excluded over 1000s of years via a process known as purifying selection.^[6] Those markers that persist are often referred to as 'stable' single nucleotide polymorphisms (SNPs).^[6] Some markers can be measured in the shorter term, for example between individuals who migrated into certain world regions (e.g. migrations into East Asia greater than 42,000 BP). These methods can produce mutation rates based on the MRCA of haplogroups (e.g. Haplogroup P and Haplogroup Q). Using such methods an old study and a new study have produced estimates between 290,000 and 82,000 BP.^[12]

Other markers are so stable that they can be measured between chimpanzees and humans, who had a common ancestor between 4 and 13 million years ago. Most studies rely on a such deep anchor point, the time of the chimpanzee-human last common ancestor (T_CHLCA). Chimpanzees and humans share a mitochondrial most recent common ancestor (MRCA). Therefore, a process similar to determining MRCA in humans can be used to estimate the sequence divergence between species. Recent CHLCA-based studies have produced estimates between 194,000 and 162,000 BP.^[3][6] These estimates follow more than 30 years of research and the estimates continue to evolve. The process is described below.

The process that has resulted in multiple determinations of the T_MRCA began during the late 1970s. Allan Wilson and his colleagues found rapidly evolving regions in the mtDNA. Given DNA sequencing technology of the time, this finding was useful because many discrepant mutations could be detected over a short sequences of DNA.^[13][14] In 1980 W.M. Brown, looking at the relative variation between human and other species, recognized that there was a constriction in the human population 180,000 BP.^[15] A year later Brown and Wilson were looking at RFLP fragments and determined the human population expanded more recently than other ape populations and noted that humans had the mtDNA diversity that was comparable to isolated subspecies of other apes.^[16]

[edit] Time estimates based on archaeological methods

Anatomical modern humans (AMH) spread out of Africa and over a large area of Eurasia and left artifacts in Southwest, South, Southeast and East Asia. Evidence of colonization can be used to estimate MtDNA haplogroups that have nodes in Southeast Asia and Oceania to estimate mutation rates. In 1987 Cann, Stoneking & Wilson (1987) used this calibration method with a type of sequence structure analysis known as RFLP to arrive at a median T_MRCA of 215,000 BP.

In 2008, Endicott & Ho (2008) used a similar calibration method with sequence information from the coding region of mitochondrial DNA. They argue that this method is particularly useful in determining when certain haplogroups entered into Europe, Australia, and the Americans. With this technique this group came up with a T_MRCA of 108,000 BP.

[edit] Time estimates based on chimpanzee-human last common ancestor

In the late 1980s, Linda Vigilant began looking at PCR based sequencing of the hypervariable region of mitochondrial DNA as more direct means of measuring sequence diversification. At the time, it was believed that sequencing this region was advantageous because of a higher density of mutations and because the HVR region avoided the implicit non-neutrality of the coding region. In addition they could employ the T_CHLCA anchor and compare the human sequences to chimpanzees sequences. Vigilant et al. (1991) used this technique to produced human mtDNA TMRCA between 166,000 and 249,000 BP (T_CHLCA = 4 to 6 Ma, see table). Their method was challenged by the fact that while more neutral than coding, the high rate of mutation caused genetic saturation which complicated direct rate estimations based on human-chimpanzees comparisons. The indirect method they used decreased confidence in their dates, therefore increasing the confidence range. Nei (1992) and Tamura & Nei (1993) have applied corrections to their methodology and generated considerably wider confidence ranges, with ranges starting at 80 Ka and ending at 760 Ka.

Advances in sequencing made it possible to sequence large numbers of genomic mitochondrial DNA (mitogenome). In 2000, Ingman et al. (2000) analyzing the non-HVR region of mitogenomes estimated a TMRCA of mtEve of 171,500. Because of the recent CHLCA used, the estimate is lower most other studies. Mitogenomic studies more carefully resolve deep branching, and detect evidence of homoplasies and reversions in HVR based lineages.

Because of the sample size this study failed to see evidence of selection or population size growth. Mishmar et al. (2003) was the first to see evidence of selection in the coding region, with an increase in sample size a pattern of regional selection with Eurasian was apparent that was absent in Africa. Gonder et al. (2007) undertook mitogenomic sequencing in areas of Africa were previous studies indicated deep diversity. This new study found new lineages of African mtDNA and more importantly narrowed the region within Africa in which humans ancestors likely arose. This new study indicated that the TMRCA most-likely occurred between 160,000 to 227,000 years ago. Gonder found the most evidence for selection outside of Africa, but also found evidence for selection within Africa, with the lowest evidence for selection in Tanzania's highly diverse mtDNA population.

This study was followed by Soares et al. (2009) which estimated the TMRCA at 192,000 years by singling out sites that were not as subject to purifying selection in the mitogenome. Evolution at these sites was compared with a method correcting for purifying selection at the first and second positions of codons.

[edit] Estimated times of major mtDNA branchpoints

Early studies like Cann, Stoneking & Wilson (1987) used paleoanthropological evidence for human settlement in New Guinea, Australia and the New World allowing them to estimate that an ancestor "c" contained no known African ancestors and they suggest this ancestor lived between 90,000 and 180,000 BP. Ingman et al. (2000) presented with an 'exodus' time from Africa in non-Africans of 52,000 ± 27,500 BP (Assuming T_CHLCA = 5 Ma).^[19] Soares et al. (2009) suggested this ancestor left Africa around 71,000 BP.

The deepest branching lineage within the human mitochondrial population is the L0/L1 branches. Beyond this, the L1 subbranches had largely been described by in the study of HVR regions in the decade previous to that study. The L0 subbranches have undergone intense study in the since 2000. Behar et al. (2008) examined the Khoisan population adding many more sequences. They determined that Khoisan mitogenomes other than the L0d and L0k appear to be the result of recent admixture. Consequently they estimated that Khoisans separated from the core interbreeding population after both the L0d and L0k clades had formed, about 144,000 ± 11,000 BP.

[edit] Inter-comparing rates and studies

Studies such as Cann, Stoneking & Wilson (1987) or Endicott & Ho (2008) draw upon archaeological discoveries that are subject to change.^[20] A growing body of evidence from the Levant (Skhul and Qafzeh), India, China and Australia (Mungo Lake- LM3) that humans had migrated from Africa well before 52 kya.^[21] For example the higher end of the confidence interval used by Endicott and Ho, for certain areas is later than the evidence of human occupation in the greater region. The critique is also a problem for CHLCA based studies. Lower-set T_CHLCA (such as used in Ingman et al.) often place the upper limit of confidence below the age of the earliest non-Neanderthal early modern humans in the Levant.

In addition, CHLCA based techniques have 2 major drawbacks: the obvious uncertainty of the CHLCA remains a problem (see: CHLCA), and molecular clocking of mitochondrial DNA has been criticized because of its inconsistent molecular clock.^[22] The effect of these two sources of errors has largely balanced each other within majority of median mtDNA MRCA estimates that are in close agreement between 180,000 and 210,000 BP. First, the unknown but likely older than 7 Million year CHLCA has a decreasing effect on rates.^[23] Second, the unknown, but likely greater than corrected for presence of slightly deleterious or adaptive mutations^[24] has a tendency to raise mutation rates in the highest (most recent) branches of the mtDNA family tree.

[edit] Mutation rate inconsistencies

For estimating rates intermediate between short and long distance estimates, Endicott et al. (2009) recommend using local or 'soft' calibration of the SNP rate (intra-population based calibration) such as archaeologically defined colonization of different world regions. However, problems with archaeological anchoring become exaggerated when used to calculate the human mtDNA MRCA. Another strategy is to characterize and correct for rate inconsistencies. Soares et al. (2009) used 2196 mtDNA sequences and uncovered 10,683 substitution events within these sequences. Eleven of 16560 sites in the mitogenome produced greater than 11% of all the substitutions with statistically significant rate variation within the 11 sites.^[25] Some rapidly evolving sites were excluded, other sites were corrected in deep branches for saturation, and other sites were corrected for purifying selection. Endicott et al. (2009), having recently reviewed the literature, have cast doubt on the use of global molecular clocks,^[26] but focus their critique on the use of such rates on dating of recent human migrations (e.g. entry of humans into the Americas).

[edit] Early modern human population structure

For anthropology, mitochondrial Eve can be thought of as a symbol of early population structure. The T_MRCAs for mitochondrial Eve do not represent a known special event within the modern human (Homo sapiens sapiens) population. Instead this range of times for mitochondrial Eve has meaning in terms of population structure. By looking at the sequences of human mtDNA scientist can tell the influences that have shaped mtDNA within modern humans.

There is a formula that applies to loci only passed by one sex. The formulation is based on the argument that under neutral selection, the T_MRCA, in generations, for any two alleles within the population is two times the population size therefore; ,^[27] where T_{G, MRCA} is the time in generations of MRCA.^[28] I_G is equal to the length of a generation in years and N_e in this instance is the population size of females that are effectively reproducing (of age and health capable of reproducing) at any given time, this is about 1/3rd the census size of females in the population. Estimates of generation time now lie between 20 and 25 years,^[29] there appears to be a preference for 22 or 23 years. To obtain population size then: .

This formula is an estimate and should be consider to have substantial statistical variance. This formula can be used directly when there is substantial evidence that population size has not grown, or that growth is recent relative to a very ancient T_MRCA. Such examples represent flat population structure. Within a adequate and representative sampling of the human population flat structures are represented by two-directional branching (splits) at branchpoints (or node) in the tree. An example of a nodes in the human population that represent flat population structures are the nodes for Haplogroup L0, Haplogroup L1'5 and the MtDNA MRCA.

Population sizes versus two commonly used generation times in the literature, plot based on the 2N rule, Note: width of varation not shown

A node with multiple independent branches are said to radiate in multiple directions. The nodes of Haplogroup M and Haplogroup N are two examples of radiation in the human population. Both of these nodes represent the oldest branchpoints for mtDNA outside of Africa. And therefore these 'radiation' of these two haplogroups can be represented by expansions Out of Africa as some humans left Africa and most space and resources to live in. Haplogroup M, as an example has 40 or more branches that radiated from the nodal Haplotype, the parent MtDNA type for the haplogroup M family tree.^[30]

[edit] Solitary female Eve as a misconception

Within Africa however the population structure has been more difficult to resolve and different studies have reached different conclusions. Based on Cann et al. 1987 of a T_MRCA of 200 Ka and generation times of 20 years, N_e for females was estimated at 5000 individuals. Based on the early mtDNA studies and other early indirect studies Takahata (1993) estimated that the N_{e, female} was between 3,500 and 4,600 individuals. As sequences accumulated for the mtDNA hypervariable regions with Vigilante et al., and later studies, evidence accumulated for a 'big bang', or explosion in lineages. However, because of the rapid nature of mutations within these hypervariable regions of the mitogenome, resolving when the population grew proved difficult.

One of the misconceptions of the heralding of mitochondrial Eve in the late 1980s was that since all females descended matrilineally from a single female that there was only one female alive at the time.^[31][32] While mtDNA studies cannot absolutely rule out this possibility, studies at other genetic loci suggest that the ancient human population carried longstanding population 10,000s of individuals in size.^[31]

[edit] Proposals of a flat population structure

One early oversight of many early studies is that the fixation of alleles (the object of coalescent theory study) is not a discrete mathematical function, it is a probabilistic function, and it is highly dependent on the ploidy being studied.

T_MRCA of MtEve compared to the 2N rule probability distributions, assuming that the population expanded 75 ka from 11,000 effectively reproducing pop. size

Takahata (1999) pointed out that conclusions drawn from single locus studies suffer from the large randomness of the fixation process. Schaffner (2004) demonstrated the three sets of coalescence ranges, haploid, X-linked and diploid where T_MRCA. Takahata (1993) estimated the effective human population size at 11,000 individuals (CHLCA = 5 Ma), and Schaffner working on an improved set of X-linked markers from low recombination regions of the X-chromosome identified an effective size of approximately 12,000 individuals.^[31][33] Based on the standard application of the 2N rule and it variance, assuming generation times are under 25 years, female to male effective ratios under 2:1, a flat population structure through the majority of African prehistory can produce all cited T_MRCA except Endicott & Ho (2008).

Y chromosomal TMRCA, the time of the Y-chromosomal Adam, lie in the 42 to 168 ky range, which is a little less than half the TMRCA of mtDNA. Importantly, the genetic evidence suggests that the most recent patriarch of all humanity is much more recent than the most recent matriarch, suggesting that 'Adam' and 'Eve' were not alive at the same time. While 'Eve' is believed to have lived more than 140,000 years ago, 'Adam' appears to have lived less than 110,000 years ago.^[32] According to Wilder, Mobasher & Hammer (2004) the lower TMRCA of Y is due to an effective population size of males 1/2 that of females over most of human evolution. This inconsistency maybe explained by some form of Y chromosome selection (cultural, or genetic). Alternatively, Pritchard et al. (1999) found that a Y-chromosomal lineage might have swept the male population. Consequently since Y chromosomal T_MRCA is lower than that for mtDNA a flat population structure couple with a high effective female to male ratio maybe considered compatible.

[edit] Proposals for a population bottleneck

MtEve's mtDNA descendants (gray) exclude older branching alleles (yellow). If population grows many fold, exclusion occurs exclusion of deepest branch occurs within bottleneck

The term bottleneck has been used to describe the population structure that created mtDNA Eve. The appearance of a bottleneck in early studies was a consequence of the appearance of a 'big bang' of HVR branching about the time humans first left Africa. From that point back to the TMRCA was less than 100,000 years and the population size estimate was below 5000 effective females. Looking backwards in time this is what might be called a retrograde bottleneck, however it is an artifact of coalescence process, since the coalescence of mitogenomes on the sequence of the MRCA (the event which initiated with mtDNA Eve and extended to the extant population) conceals the population size from all points earlier than that mutation. Therefore, the human population may not have been going through a constriction at the time of mitochondrial Eve. Instead, it may have existed at its low-stand for a long period of time (100s of thousands of years) until technology allowed its expansion in Africa.

Population structure is heavily influenced by branching in several ways. With the recent expansion of mitogenomic sequences within the last 5 years there is increasing evidence of a population expansion in Africa. Atkinson, Gray & Drummond (2009) argue that there were significant increases in the population size in mulitple lineages (L0, L1, L2 and especially L3) prior to 60 kya. As a consequence of these earlier expansions this group was able to establish a lower limit of effective females at approximately 1000 individuals and an upper limit of population size about 11,000. The median estimate, about 3500 individuals is less than expected number of females based on X-linked studies and the lower range of the confidence interval is below Shaffner's estimates, indicating that a bottleneck might have been present.

[edit] Implications of dating and placement of Eve

The first studies suggesting a recent common ancestor for humans within Africa came a time when a hypothesis for human evolution, known as the Multiregional Evolution hypothesis was popular among some leading Paleoanthropologist (e.g. Milford Wolpoff). The impetus for this hypothesis comes from the belief that humans left Africa first about 2 million years ago and spread globally, these humans were similar to modern humans in many ways. Other biases indicated the temporal (time) difference between Homo erectus and modern Homo sapiens was too short to allow for another new species, and many authors perceived regional evolution from Archaic forms into modern forms. Consequently, the finding of a recent maternal ancestor for all humans in Africa created an extended controversy. Over a decade the MREH theory retracted to theories of admixture (Such as proposed by Eric Trinkhaus) to mostly recent Africa origin hypotheses.

[edit] Geographic and Temporal constraints of early modern humans

The strict Out of Africa and Multiregional Evolution controversy revolved around where to best place the evolution of anatomically modern humans over evolutionary timescales.

Cann, Stoneking & Wilson (1987) 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 using data from many loci has extrapolated origins to the Angola-Namibia border region near the Atlantic Ocean (although this region has poor genetic definition), whereas Behar et al. 2008 places an ancestral population in Ethiopia. These opinions all point toward a sub-Saharan origin. More recent literature on languages and pygmy phenotype indicate that L0 and L1 were carried by click-speaking pygmies from SE Africa to Central and Western Africa, therefore explaining much of the genetic diversity in those regions. Consequently, more recent studies have tended to push the cradle of humanity more toward the South or East of Africa.

To some extent the studies have already revealed that the presence of archaic homo sapiens in Northwest Africa, Jebel Irhoud, were not likely part of the contiguous modern human population. In addition, the older remains at Skhul and Qafzeh are also unlikely part of the constrict human population, evidence currently indicates humans expanded in the region no earlier than 90,000 BP.^{[citation needed]} Tishkoff argues that humans might have migrated to the levant before 90 Ka, but this colony did not persist in SW Asia.^{[citation needed]} Better defined is the genetic separation between Neanderthals, Flores hobbit, Java man, 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 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 larger genetic divide with other hominids.

Since Multiregional evolution hypothesis (MREH) revolved around a belief that regional modern human population evolved in-situ in various regions (Europe - Neandertals to Europeans, Asia - Homo Erectus to East Asians, Australia - Sumatran erectines to Indigeonous Australians), the demonstrated that a pure MREH hypothesis could not explain one important genetic marker.

[edit] Mitochondrial MRCA and the MRCA of all humans

Mitochondrial Eve is the most recent common matrilineal ancestor, not the MRCA. Since the mtDNA are 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. Ironically mtDNA are not human; they are organelles that live within our cells, so it is better to say these are human-mitochondrial Most Recent Common Ancestor. Despite the recent fixation of the mtDNA genome in humans, other genes have evolved that were broadly selective in the human population, genes that have swept through the human population, two such genes having been identified on the X-chromosome.

Rohde, Olson & Chang (2004) indicate that the overwhelming majority of humans have a recent common ancestor within the last 5000 years (albeit between any two individuals, it may not be the same ancestor), however the genetic relationship between well diverged individuals may not reflect the theoretical relationship, as geographic and cultural barriers may slow gene migration. Gene migration is not fluid in humans, as genes are passed in units called chromosomes, which undergo a limited number of recombination on each unit per generation; therefore a common ancestor genealogically may not indicate the passage of DNA from that ancestor to the two divergent individuals. Whereas since mtDNA does not undergo this dilution via recombination, we can argue that the majority of mtDNA sequence (that which has not undergone mutation) from mtDNA ~16000 nts came from a single individual >150,000 years ago. A more recent common ancestor for all males is the much larger Y chromosome (however it codes for very few genes).

[edit] In popular science and culture

Cover of the January 11, 1988 edition of Newsweek

Newsweek Magazine 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.^[34]

Bryan Sykes (2001) presents the theory of human mitochondrial genetics to a general audience.

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.

The Discovery Channel produced a documentary entitled The Real Eve (or Where We Came From in the United Kingdom), based on the book Out of Eden by Stephen Oppenheimer.

In popular culture

• The main villainess of the Parasite Eve series is named Mitochondria Eve.
• In the "reimagined" Battlestar Galactica television series, Hera Agathon (the Cylon-human hybrid) is the Mitochondrial Eve as revealed in the series finale.^[35]

[edit] See also

[edit] Footnotes

[edit] References

[edit] External links

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