One of the great innovators of science, New Zealander Allan Wilson revolutionised the study of human evolution. He was shortlisted for the Nobel Prize and is the only New Zealander to win the prestigious US MacArthur “Genius” Award. Allan Wilson’s scientific achievements are nothing short of profoundly significant.
His investigations into the origins of humanity through biochemistry were revolutionary, yet at the time of his death in 1991, he was still a controversial figure. His theories on the evolution and age of modern humans still flew in the face of anthropological thinking of the time, not to mention inciting anger from American creationists.
After Wilson’s death Charles Laird published some thoughts on his lost colleague and friend. He examined his work and his personality and theorised about how the two combined:
“I have wondered about the parts of his personality that were so unusual even among first-rate scientists – his courage, his openness, his ability to focus on a problem and not let go, his special vision to see the final experiment and not to get distracted by intermediate ones and the details in between. Where did these traits come from? Do we learn them? If so can we learn them as young adults, or teach them to our children and students? Or do we have to learn them from our parents when we are young?”
Into Biochemistry
Allan Wilson was born in Ngaruawahia, New Zealand, and raised on a farm at Helvetia, Pukekohe. He attended King’s College in Auckland and excelled in maths and chemistry. After school he gained a BSc from Otago University. It was here, as a Masters student, that Wilson met Professor C.P. ‘Mac’ McMeekan, a New Zealand pioneer in soil and water science. He suggested that Wilson further his study in biochemistry instead of genetics.
In 1955, Wilson was invited to do his Ph.D at the University of California, Berkeley. His father was reluctant to let his son go to America, for fear that he would never return, but Wilson’s mother saw her son’s potential and persuaded her husband to let him go. At the time the family thought Wilson would only be gone two years; instead he stayed at Berkeley for 35 years, setting up one of the world’s most creative biochemistry labs and turning ideas of evolution on their ear.
The Molecular Clock
Allan Wilson first came to world attention when he published a paper titled ‘Immunological Time-Scale For Human Evolution’ in Science magazine in December 1967. Together with doctoral student Vince Sarich, Wilson argued that the origins of the human species could be seen through, what he termed, a ‘molecular clock’. This was a way of dating, not from fossils, but from the genetic mutations that had accumulated since they parted from a common ancestor. The molecular clock estimated the length of time from divergence, given a certain rate of change.
When Wilson and Sarich analysed and compared genetic material from humans with chimpanzees they found the material to be 99 percent identical. From this, using the ‘molecular clock’ reasoning (bigger differences equate to greater time since their last common ancestor) they deduced that the earliest proto-hominids evolved only five million years old. This was fifteen million years younger than stated by conventional anthropology.
An obituary written by Joseph Felsenstein and published in Nature magazine describes how Wilson turned around, not only the study of molecular evolution, but also, the way it was studied:
“Systematists, microbiologists and biochemists came to Berkeley to learn the techniques involved. But Wilson’s influence went beyond this. While others concentrated on what evolution could tell them about molecules, Wilson always looked for ways that molecules could say something about evolution.”
Berkeley in the 1960s was a hotbed of protest against the Vietnam War and the American military industrial establishment, and rife with academic liberalism. By all accounts Wilson was highly active in the tide of action sweeping the campus.
Crop of Hybrids
Yet it was in the lab where his influence was felt the strongest. The Wilson Lab at Berkeley pioneered new techniques, such as the Polymerase Chain Reaction and the Relative Rate Test. These pushed the limits of DNA analysis to include extinct species. DNA samples from extinct species such as Moa and Tasmanian wolves were tested, as was fossil bacteria and tissue from a 7000 year old human brain.
In a feature on the relationship between molecular and evolutionary biology published in Science magazine, science writer Ann Gibbons recognises Wilson’s influence on modern scientific thought and practices through his teaching and his foresight to combine disciplines:
“The first recognised prototype of the molecular systematics lab was set up at Berkeley, where Wilson’s lab in the department of genetics collaborated with a wide array of faculty and other researchers on campus. And out of that lab – and others like it at Berkeley – came a crop of hybrids: graduate students trained in both systematics and molecular techniques who have seeded the combination around the world.
“At a celebration for the 25th anniversary of Wilson’s lab, former students jokingly showed a slide of a map of the world, showing the paths the Berkeley researchers have taken had taken to other molecular evolution labs around the world. You could trace Berkeley students and collaborators to brand new labs at Pennsylvania State University, Louisiana State University, the University of Hawaii, the Museo Nacional de Ciecias Naturales in Madrid, and the Smithsonian’s Museum of National History among others.”
Despite Wilson’s molecular evidence being as strong as the conventional fossil evidence, he remained on the fringes of the anthropological community for the next twenty years. While this was definitely a case of academic politics and a demonstration of the difficulty of changing accepted norms, Wilson, according to accounts given by his team, appeared to relish his role as an outsider, an edge-dweller. He gathered the brightest students around him to test his theories on a multitude of plant and animal species.
Slowly through the 1970s his ideas gained credibility, and through the course of his career he was a visiting professor at Harvard, St Louis, Kansas, Carmel and MIT, and at universities in Israel and Kenya. He edited scientific magazines and journals including: Biochemical Genetics, Chemical Abstracts, Journal of Molecular Evolution, Journal of Human Evolution, Systematic Zoology and Geonomics.
Evolutionary Bombshell
In the early 1980s, as his findings for the age of the proto-humans were starting to be more widely accepted, Wilson again dropped a bombshell on traditional anthropological thinking.
In studying gene types he started to focus on mitochondria DNA (mDNA) – genes that sit in the cell, but not in the nucleus, and are passed from mother to child. This DNA is material important because it mutates quickly, thus making it easy to plot changes over relatively short time spans. By comparing differences in the (m)DNA Wilson believed it was possible to estimate the time, and the place, modern human first evolved. With his discovery that human mDNA is genetically much less diverse than chimpanzee mDNA, he concluded that modern human races had diverged recently from a single population while older human races such as Neanderthal, Java Erectus and Pekin Erectus had become extinct.
He and his team compared mDNA in people of different racial backgrounds and concluded that all modern humans evolved from one ‘lucky mother’ in Africa about 200,000 years ago.
This finding was as, if not more, controversial than his 1967 findings. Accepted thinking had various human groups evolving from different ancestors, over a million years in separate geographic regions, but at basically the same rate around the world. In Europe with Homo Sapien Neanderthals, in Indonesia with Java Man, in China with Peking Man. Again, like in the 1960s, many palaeontologists rejected Wilson’s conclusions; fossil scientists were unfamiliar with biochemistry and trusted their own data more than molecular data. It took 20 years to convince palaeontologists of the value of Wilson’s theory, but when they did, it married their science with that of genetics. It was Wilson’s legacy to turn genetics into a study of inherited traits to a biochemical science.
The media concerning this discovery initially offended the religious lobby. Time magazine ran the story as the ‘Black Eve Theory’; other magazines followed with headlines proclaiming ‘African Eve’ and other derivations.
The ‘Out of Africa’ theory is now the accepted account of modern human origins. Further computer analysis of mDNA data, studies of the male ‘Y’ chromosome (indicating a single male ancestor living in Africa around 270,000 years ago) and re-analysis of the original data show that European Neanderthal and Java Erectus are not ancestral to modern humans.
Allan Wilson died in 1991 in Seattle, aged 55, at the height of his career. He left behind a wife, son and daughter and, back in New Zealand, a mother, brother and sister. He had kept his New Zealand ties (his brother Gary donated bone marrow as part of Wilson’s leukaemia treatment). He also remained close to Otago University, addressing the graduates of 1988, returning the next year to teach, being awarded an honorary doctorate that year and being the recipient of the William Evans Fellow for science.
Marvellous Legacy
Many of his Berkeley students are at the forefront of biochemistry today. In a obituary written by British evolutionary theorist, John Maynard Smith, and published in The Independent, Smith points out that Wilson’s teaching ability should be remembered as well as his theories:
“I find it impossible to write about Allan Wilson without, in fact, writing about his science, because his science was always such fun. He succeeded in passing on his enthusiasm, his open-mindedness, and his breadth of interest to a succession of young scientists who worked with him at Berkeley. They are a wonderful advertisement for Allan as a teacher: there is not a narrow-minded one among them. Evolutionary biology can ill-afford to do without him, but he has left us a marvellous legacy of ideas and a splendid group of students.”
Further Reading
Thank you to Rowan Taylor for his comments and contributions. Rowan has contributed a comprehensive selection of articles, website references and list of papers by Allan Wilson and colleagues.
Sites which have linked to this page:
Allan Wilson Centre for Molecular Ecology and Evolution, Massey University, NZ. The centre was set up as part of the NZ Government’s Centres of Research Excellence initiative and continues research into the stuff of our origins under the inspiration of Allan’s example.
(A) Articles About Allan Wilson
Rebecca Andrews (1991) FASEB honours Berkeley biochemist. The Scientist, Vol 5 (No. 12) 19 June. Can be downloaded from:
http://www.the-scentist.library.upenn.edu/yr1991/june/peop_910610.html
Bob Brockie (1992) The Origin of Our Species. Evening Post (22 July)
Rebecca Cann, (1993) Obituary: Allan C. Wilson, 1935-1991. Human Biology Vol.65 (No.3), p. 343-58.
“Six Who Dared”, Cornucopia (Washington State University), Fall 1990
Eve, genetics. The Concise Columbia Electronic Encylopedia, 3rd Edition
http://www.encyclopedia.com/articles/04296.html
J.Felsenstein (1991) Allan Charles Wilson (1934-1991). Nature Vol.353 (5 September) p.19
Ann Gibbons (1991) Systematics goes molecular. Science Vol.251, pp.872-874.
Thomas Jukes and Emile Zuckerkandl (1991) Remembering Allan Wilson. Journal of Molecular Evolution Vol.33 (18 July) p.ii
John Maynard Smith (1991) Obituary: Allan C. Wilson. The Telegraph (10 August)
Rowan Taylor (1998) Common Bonds. New Zealand Listener (31 January)
H.B.White (1995) Dating Eve. Case Study Problem in Molecular Evolution No. 3
(Course notes for CHEM-647 BIOCHEMICAL EVOLUTION)
http://www.udel.edu/pbl/curric/chem647prob.html
Gilbert Wong (21 July 1991) The Evolution of Eve. NZ Herald
Gilbert Wong (12 August 1989) Answers Hinge on DNA Theory. NZ Herald
(B) Selected Papers by Allan Wilson
Abstracts of many of the following papers can be seen at the following website:
http://www.ncbi.nlm.nih.gov/htbin-post/Entrez/query?
A.C.Wilson (1975) Evolutionary importance of gene regulation. Stadler Genetics Symposium Vol.7, pp.117-134.
A.C.Wilson (1985) The molecular basis of evolution. Scientific American Vol.253 (No.4, October), pp.164-173.
A.C.Wilson (1986) Time scale for bird evolution. Proceedings of the International Ornithological Congress, Ottawa.
A.C.Wilson (1990) Will sequencing the human genome revolutionize biology? New Biology, Vol.2 (No.7), pp.585-586.
A.C.Wilson and N.O.Kaplan (1963) Enzymes and nucleic acids in systematics. Proceedings of the XVI International Congress of Zoology Vol.4, pp.125-127.
A.C.Wilson and N.O.Kaplan (1964) Enzyme structure and its relation to taxonomy. In: C.A.Leone (editor) Taxonomic Biochemistry and Serology Ronald Press, New York.
A.C.Wilson and R.L.Cann (1992) The recent African genesis of humans. Scientific American Vol.266 (No 4, April): pp.68-73
A.C.Wilson and V.M.Sarich (1969) A molecular time scale for human evolution. Proceedings of the National Academy of Sciences USA Vol.63 (No.4), pp.1088-1093
A.C.Wilson, G.L.Bush, S.M.Case and M.C.King (1975) Social structuring of mammalian populations and rate of chromosomal evolution. Proceedings of the National Academy of Sciences USA Vol.72,5061-5065
A.C.Wilson, L.R.Maxson and V.M.Sarich (1974) Two types of molecular evolution: Evidence from studies of interspecific hybridization. Proceedings of the National Academy of Sciences USA Vol.71, pp.2843-2847
A.C.Wilson, R.L.Cann, S.M.Carr, M.George, U.B.Gyllensten, K.M.Bychowski, R.G.Higuchi, S.R.Palumbi, E.M.Prager, R.D.Sage and M.Stoneking (1985) Mitochondrial DNA and two perspectives on evolutionary genetics. Biological Journal of the Linnaean Society Vol.26, pp.375-400
A.C.Wilson, S.S.Carlson and T.J.White (1977) Biochemical evolution. Annual Review of Biochemistry Vol.46, pp.573-639.
A.C.Wilson, V.M.Sarich and L.R.Maxson (1974) The importance of gene rearrangement in evolution: evidence from studies on rates of chromosomal, protein, and anatomical evolution. Proceedings of the National Academy of Sciences USA Vol.71, pp.3028-3030
A.Cooper, C.Mourer-Chauviré, G.K.Chambers, A.von Haeseler, A.C.Wilson and S.Pääbo (1992) Independent origins of New Zealand moas and kiwis. Proceedings of the National Academy of Sciences USA Vol.89, pp.8741-8744
A.DiRienzo and A.C.Wilson (1991) Branching pattern in the evolutionary tree for human mitochondrial DNA Proceedings of the National Academy of Sciences USA Vol.88, pp.1597-1601.
A.Larson, E.M.Prager and A.C.Wilson (1984) Chromosomal evolution, speciation and morphological change in vertebrates: The role of social behaviour. In M.D. Bennett and A.Gropp (editors) Chromosomes Today (Vol.8) Allen and Unwin, pp. 215-228.
A.Meyer and A.C.Wilson (1990) Origin of tetrapods inferred from their mitochondrial DNA affiliation to lungfish. Journal of Molecular Evolution Vol.31 (No.5), pp 359-364
A.Sidow and A.C.Wilson (1990) Compositional statistics: an improvement of evolutionary parsimony and its application to deep branches in the tree of life. Journal of Molecular Evolution Vol.31 (No.1), pp.51-68
A.Sidow, A.C.Wilson and S.Pääbo (1991) Bacterial DNA from Clarkia fossils. Philosophical Transactions of the Royal Society B. Vol.333, pp.429-433
C.B.Stewart, J.W.Shilling and A.C.Wilson (1987) Adaptive evolution in the stomach lysozymes of foregut fermenters. Nature Vol.330, pp.401-404 (see also commentary p.315)
C.Y.K.Ho, E.M.Prager, A.C.Wilson, D.T.Osuga and R.E.Feeney (1976) Penguin evolution: Protein comparisons demonstrate phylogenetic relationship to flying aquatic birds. Journal of Molecular Evolution Vol.8, pp.271-282.
D.G.Wallace, L.R.Maxson and A.C.Wilson (1971) Albumin evolution in frogs: A test of the evolutionary clock hypothesis. Proceedings of the National Academy of Sciences USA Vol.68, pp.3127-3129.
D.M.Irwin, T.D.Kocher and A.C.Wilson (1991) Evolution of the cytochrome b gene of mammals. Journal of Molecular Evolution Vol.32 (No.2), pp.128-144
E.M.Prager, A.C.Wilson, D.T.Osuga and R.E.Feeney (1976) Evolution of flightless land birds on southern continents: transferrin comparison shows monophyletic origin of ratites. Journal of Molecular Evolution Vol.8(No.3), pp.283-294
E.M.Prager, D.P.Fowler and A.C.Wilson (1976) Rates of evolution in conifers (Pinaceae). Evolution Vol.30, pp.637-649.
E.M.Prager, A.C.Wilson, J.M.Lowenstein and V.M.Sarich (1980) Mammoth albumin. Science Vol.209, pp.287-289
G.B.Kitto and A.C.Wilson (1966) Evolution of malate dehydrogenase in birds. Science Vol.153, pp.1408-1410.
L.L.Cavalli-Sforza, A.C.Wilson, C.R.Cantor, R.M.Cook-Deegan and M.C.King (1991) Call for a worldwide survey of human genetic diversity: a vanishing opportunity for the Human Genome Project. Genomics Vol.11 (No.2) pp.490-491.
L.M.Cherry, S.M.Case and A.C.Wilson (1978) Frog perspective on the morphological difference between humans and chimpanzees. Science Vol.200, pp.209-211.
L.M.Cherry, S.M.Case and A.C.Wilson (1979) Comparison of frogs, humans and chimpanzees. Science Vol.204, p.435.
L.R.Maxson, V.M.Sarich and A.C.Wilson (1975) Continental drift and the use of albumin as an evolutionary clock. Nature Vol.255, pp.397-400.
L.R.Maxson and A.C.Wilson (1979) Rates of molecular and chromosomal evolution in salamanders. Evolution Vol.33, pp.734-740.
L.Vigilant, M.Stoneking, H.Harpending, K.Hawkes and A.C.Wilson (1991) African populations in the evolution of human mitochondrial DNA. Science Vol.253, pp.1503-1507.
M.C.King and A.C.Wilson (1975) Evolution at two levels in humans and chimpanzees. Science Vol.188, pp.107-116 (see also commentary Science Vol.189, pp.446-447).
M.C.King and A.C.Wilson (1975) Our close cousin, the chimpanzee. New Scientist Vol.67, pp.16-18
M.Hasegawa, A.DiRienzo, T.D.Kocher and A.C.Wilson (1993) Toward a more accurate time scale for the human mitochondrial tree. Journal of Molecular Evolution Vol.37, pp.347-354.
P. Goloubinoff, S.Pääbo and A.C.Wilson (1993) The evolution of maize according to nuclear DNA sequences from archaeological specimens. Proceedings of the National Academy of Sciences USA Vol.90, pp.1997-2001
R.G.Higuchi, B.Bowman B, M.Freiberger, O.A.Ryder and A.C.Wilson (1984) DNA sequences from the quagga, an extinct member of the horse family. Nature Vol.312, pp.282-284.
R.G.Higuchi, L.A.Wrischnik, E.Oakes, M.George, B.Tong and A.C.Wilson (1987) Mitochondrial DNA of the extinct quagga: relatedness and extent of postmortem change. Journal of Molecular Evolution Vol.25 (No.4), pp.283-287.
R.H.Thomas, W.Schaffner, A.C.Wilson and S.Pääbo (1989) DNA phylogeny of the extinct marsupial wolf. Nature Vol.340, pp.465-467. (See also R.H.Thomas, A.C.Wilson and S.Pääbo (1990) Reply to Faith. Nature Vol.345, p.390)
R.L.Cann and A.C.Wilson (1982) Models of human evolution. Science Vol.217, pp.303-304.
R.L.Cann and A.C.Wilson (1983) Length mutations in human mitochondrial DNA. Genetics Vol.104, pp.699-711.
R.L.Cann, W.M.Brown and A.C.Wilson (1984) Polymorphic sites and the mechanism of evolution in human mitochondrial DNA. Genetics Vol.106, pp.479-499.
*THE ‘EVE’ PAPER
R.L.Cann, M.Stoneking and A.C.Wilson (1987) Mitochondrial DNA and human evolution. Nature Vol.325, pp. 31-36. Can be downloaded from http://artsci.wustl.edu/~landc/html/cann/
S.D.Ferris, A.C.Wilson and W.M.Brown (1981) Evolutionary tree for apes and humans based on cleavage maps of mitochondrial DNA. Proceedings of the National Academy of Sciences USA Vol.78, pp.2432-2436.
S.D.Ferris, W.M.Brown, W.S.Davidson and A.C.Wilson (1981) Extensive polymorphism in the mitochondrial DNA of apes. Proceedings of the National Academy of Sciences USA Vol.78, pp.6319-6323.
S.D.Ferris, R.D.Sage, E.M.Prager, U.Ritte and A.C.Wilson (1983) Mitochondrial DNA evolution in mice. Genetics Vol.105, pp.681-721.
S.M.Beverley and A.C.Wilson (1984) Molecular evolution in Drosophila and the higher diptera II: A time scale for fly evolution. Journal of Molecular Evolution Vol.21, pp.1-13.
S.Pääbo and A.C.Wilson (1991) Miocene DNA sequences – A dream come true? Current Biology Vol.1, pp.45-46.
S.Paabo, J.A.Gifford and A.C.Wilson (1988) Mitochondrial DNA sequences from a 7,000-year-old brain. Nucleic Acids Research. Vol.16, pp.9775-9787.
S.Pääbo, R.G.Higuchi and A.C.Wilson (1989) Ancient DNA and the polymerase chain reaction: The emerging field of molecular archaeology (Minireview). Journal of Biological Chemistry. Vol.264, pp.9709-9712
S.Pääbo, W.K.Kelley, K.M.Whitfield, Y.Kamazawa and A.C.Wilson (1991) Rearrangements of mitochondrial tRNA genes in marsupials. Journal of Molecular Evolution Vol.33, pp.426-430.
S.S.Carlson, A.C.Wilson and R.D.Maxson (1978) Do albumin clocks run on time? Science Vol.200, pp.1183-1185.
T.J.White and A.C.Wilson (1978) Molecular anthropology. Evolution Vol.32, pp.693-694.
V.M.Sarich and A.C.Wilson (1967) Immunological time scale for hominid evolution. Science Vol.158, pp.1200-1203.
V.M.Sarich and A.C.Wilson (1967) Rates of albumin evolution in primates. Proceedings of the National Academy of Sciences USA Vol.58 (No.1), pp.142-148.
V.M.Sarich and A.C.Wilson (1973) Generation time and genomic evolution in primates. Science Vol 179, pp.1144-1147.
W.K.Thomas, S.Pääbo, F.X.Villablanca and A.C.Wilson (1990) Spatial and temporal continuity of kangaroo rat populations shown by sequencing mitochondrial DNA from museum specimens. Journal of Molecular Evolution Vol.31, pp.101-112.
W.M.Brown, E.M. Prager, A.Wang and A.C.Wilson (1982) Mitochondrial DNA sequences of primates: Tempo and mode of evolution. Journal of Molecular Evolution Vol.18, pp.225-239.
W.M.Brown, M.George Jr. and A.C.Wilson (1979) Rapid evolution of animal mitochondrial DNA. Proceedings of the National Academy of Sciences USA Vol.76, pp.1967-71
(C) Research Supporting the Eve/Out of Africa Hypothesis
J.A.L.Armour, T.Anttinen, C.A.May, E.E.Vega, A.Sajantila, J.R.Kidd,
K.K.Kidd, J.Bertranpetit, S.Pääbo and A.Jeffreys (1996) Minisatellite diversity supports a recent African origin for modern humans. Nature Genetics Vol.13, pp.154-160.
F.J.Ayala (1995) The myth of Eve: molecular biology and human origins. Science Vol.270, pp.1930-1936.
A.M.Bowcock, A.Ruiz-Linares, J.Tomfohrde, E.Minch, J.R.Kidd and L.L.Cavalli-Sforza (1994) High resolution of human evolutionary trees with polymorphic microsatellites. Nature Vol.368, pp.455-457.
R.L.Cann (1992) The search for Eve [letter]. Science, Vol.256, p.79.
L.L.Cavalli-Sforza (1991) Genes, people and languages. Scientific American (November) pp. 70-78.
L.L.Cavalli-Sforza, P. Manozzi and A.Piazza (1993) Demic expansions and human evolution. Science Vol.259, pp.639-646. (Has gene distribution maps showing the imprint of ancient migration waves out of Africa). [See also published erratum in Science Vol.261 (17 September 1993) p.1508].
L.L.Cavalli-Sforza and F.Cavalli-Sforza (1995) The Great Human Diasporas. Addison-Wesley, New York.
J.Y.Chu, W.Huang, S.Q.Kuang, J.M.Wang, J.J.Xu, Z.T.Chu, Z.Q.Yang, K.Q.Lin, P.Li, M.Wu, Z.C.Geng, C.C.Tan, R.F.Du, L.Jin (1998) Genetic relationship of populations in China. Proceedings of the National Academy of Sciences USA Vol.95, pp.11763-11768
R.L.Dorit, H.Akashi and W.Gilbert (1995) Absence of polymorphism at the ZFY’ locus of the human Y chromosome. Science Vol.268, pp.1183-1185. (Finds that the Y chromosome, passed on only by males, tells a similar evolutionary story to the mDNA passed on by females.)
Ann Gibbons (1994) African origins theory goes nuclear. Science Vol.264, pp.350-351.
Ann Gibbons (1995) Out of Africa – at last? Science Vol.267, pp.1272-1273.
Ann Gibbons (1997) Y chromosome shows that Adam was African [news]. Science Vol.278, pp.804-805.
D.B.Goldstein, A.Ruiz Linares, L.L.Cavalli-Sforza and M.W.Feldman (1995) Genetic absolute dating based on microsatellites and the origin of modern humans. Proceedings of the National Academy of Sciences USA Vol.92, pp.6723-6727.
M.F.Hammer (1995) A recent common ancestry for human Y chromosomes. Nature Vol.378, pp.376-378.
M.F.Hammer et al. (1997) The geographic distribution of human Y chromosome variation. Genetics Vol.145 (No.3), pp.787-805.
S.Horai, K.Hayasaka, R.Kondo, K.Tsugane, N.Takahata (1995) Recent African origin of modern humans revealed by complete sequences of hominoid mitochondrial DNAs. Proceedings of the National Academy of Sciences USA Vol.92, pp.532-536.
L.B.Jorde, M.Bamshad and A.R.Rogers (1998) Using mitochondrial and nuclear DNA markers to reconstruct human evolution. Bioessays Vol.20(No.2 February), pp.126-136
M. Krings, A.Stone, R.W.Schmitz, H.Krainitzki, M.Stoneking and S.Pääbo (1997) Neandertal DNA sequences and the origin of modern humans. Cell Vol.90, pp.19-30 (Finds that Neandertals were cousins not ancestors of modern humans.)
M.Krings, H.Geisert, R.W.Schmitz, H.Krainitzki and S.Pääbo (1999) DNA sequence of the mitochondrial hypervariable region II from the Neandertal type specimen Proceedings of the National Academy of Sciences USA Vol.96, pp.5581-5585. (Confirms earlier finding that Neandertals were cousins not ancestors of modern humans.)
M. Nei (1995) Genetic support for the out-of-Africa theory of human evolution. Proceedings of the National Academy of Sciences USA Vol.92, pp.6720-6722.
S.Pääbo (1995) The Y chromosome and the origin of all of us (men). Science Vol.268, pp.1141-1142
D.Penny, E.Watson and M.Steel (1993) Trees from languages and genes are very similar. Systematic Biology Vol.42, pp.382-384.
D.Penny, M.A.Steel, P.J.Waddell and M.D.Hendy (1995) Improved analyses of human mtDNA sequences support a recent African origin for Homo Sapiens. Molecular Biology and Evolution. Vol.12, pp.863-882.
D.E.Reich and D.B.Goldstein (1998) Genetic evidence for a Paleolithic human population expansion in Africa. Proceedings of the National Academy of Sciences USA Vol.95 (No.14), pp.8119-8123. [See also the published erratum Proceedings of the National Academy of Sciences USA Vol.95 (No.18), p.11026.]
M.Ruvolo (1996) A new approach to studying modern human origins: hypothesis testing with coalescent time distributions. Molecular Phylogenetics and Evolution Vol.5 (No.1), pp.202-219.
M.Seielstad, E.Bekele, M.Ibrahim, A.Toure and M.Traore (1999) A view of modern human origins from Y chromosome microsatellite variation. Genome Research Vol.9 (No 6, June), pp.558-567.
M.D.Shriver, L.Jin, R.E.Ferrell and R. Deka (1997) Microsatellite data support an early population expansion in Africa. Genome Research Vol.7 (No.6), pp.586-591.
C..B.Stringer and P.Andrews (1988) Genetic and fossil evidence for the origin of modern humans. Science Vol.239, pp.1263-1268.
Chris B.Stringer (1990) The emergence of modern humans. Scientific American (December), pp.98-104.
Chris Stringer and Robin McKie (1996) African Exodus: The Origins of Modern Humanity. Jonathan Cape, London. C.C.Swisher III, W.J.Rink, S.C.Antón, H.P.Schwarcz, G.H.Curtis, A.Suprijo and Widiasmoro (1996) Latest Homo erectus of Java: potential contemporaneity with Homo sapiens in Southeast Asia. Science Vol.274, pp.1870-1874. (New dating method shows that Homo erectus was in Java after the arrival of Homo sapiens less than 30,000 years ago – making them cousins rather than ancestors.)
S.A.Tishkoff, E. Dietzsch, W.Speed, A.J. Pakstis, J.R. Kidd, K. Cheung, B. Bonne-Tamir, A.S.Santachiara-Benerecetti, P.Moral, M.Krings, S.Pääbo, E.Watson, N.Risch, T.Jenkins and K.K.Kidd (1996) Global patterns of linkage disequilibrium at the CD4 locus and modern human origins. Science Vol.271, pp.1380-1387.
D.M.Waddle, (1994) Matrix correlation tests support a single origin for modern humans. Nature Vol.368, pp.452-454. (This first quantitative test of the fossil evidence supports the African origin hypothesis.)
L.Whitfield, L.Simon, J.E.Sulston and P.N.Goodfellow (1995) Sequence variation of the human Y chromosome. Nature Vol.378, pp.379-380.
(D) Crictics of the Eve/Out of Africa Hypothesis
M.Barinaga (1992) ‘African Eve’ backers beat a retreat Science Vol.255, pp.686-687.
P.Darlu and P.Tassy (1987) Disputed African origin of human populations. Nature Vol.329, p.111 (with response from Cann et al., pp.111-112).
H.Gee (1992) Statistical cloud over African Eden Nature Vol.355, p.583.
R.M.Harding, S.M.Fullerton, R.C.Griffiths, J.Bond, M.J.Cox, J.A.Schneider,
D.S.Moulin and J.B.Clegg (1997) Archaic African and Asian lineages in the genetic ancestry of modern humans. American Journal of Human Genetics. Vol.60 (No. 4), pp.772-789.
A.R.Templeton (1992) Human origins and analysis of mitochondrial DNA sequences. Science Vol.255, p.737.
A.R.Templeton (1993) The ‘Eve’ Hypothesis: A genetic critique and reanalysis. American Anthropologist Vol.95, pp.51-72.
A.Thorne and M.Wolpoff (1992) The multiregional evolution of Humans. Scientific American (April) pp.28-33
A.G.Thorne, M.H.Wolpoff and R.B.Eckhardt (1993) Genetic variation in Africa. Science Vol.261 (No.5128, 17 September), pp.1507-8
M.Wolpoff and A.Thorne (1991) The case against Eve. New Scientist pp.37-41.
