Multiregional origin of modern humans: Wikis

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A graph detailing the evolution to modern humans using the Multiregional theory of human evolution. The horizontal lines represent 'multiregional evolution' gene flow between regional lineages.

The multiregional hypothesis is a model to account for the pattern of human evolution proposed by Milford H. Wolpoff[1] in 1988.[2] Multiregional origin holds that the evolution of humanity from the beginning of the Pleistocene 2.5 million years BP to the present day has been within a single, continuous human species, evolving worldwide to modern Homo sapiens.

A competing theory, the recent African origin of modern humans (also known as "Out of Africa"), has emerged as the near consensus view since the 1990s,[3][4] proposing that modern humans arose in Africa around 100-200,000 years ago, moving out of Africa around 50-60,000 years ago to replace existing human species such as Homo erectus and the Neanderthals.[5]

Proponents of multiregional origin point to fossil and genomic data[6] as support for their hypothesis. The gene flow and sexual reproduction between modern and ancestral human population has not been ruled out.[7][8] There is no evidence for a contribution of Neanderthal mitochondrial DNA (37 genes) [9] or Y-DNA (78 genes) to modern humans; the studies claiming to find evidence of older diversity in ancestry are on the autosomal DNA (20-25k genes) making up the vast majority of the human genome.

Contents

Regional continuity

The term "multiregional hypothesis" was first coined in the early 1980s by Milford H. Wolpoff and colleagues as an explanation for the apparent similarities seen in Homo erectus and Homo sapiens fossils from the same region, what they called regional continuity.[2]

Wolpoff rejected the earlier proposal by Coon of parallel evolution,[2] and proposed a theory based on clinal variation that would allow for the necessary balance between local selection and a global species. He proposed that Homo erectus, Neanderthals, Homo sapiens and other humans were a single species. This species arose in Africa two million years ago as H. erectus and then spread out over the world, developing adaptations to regional conditions. It was proposed that for periods of time some populations became isolated, developing in a different direction, but through continuous interbreeding, replacement, genetic drift and selection, adaptations that were an advantage anywhere on earth would spread, keeping the development of the species in the same overall direction, while maintaining adaptations to regional factors. Eventually, the more unusual local varieties of the species would have disappeared in favor of modern humans, retaining some regional adaptations, but with many common features.[2]

Fossil evidence

Some proponents of the multiregional hypothesis, including Wolpoff, argue that fossil evidence is more reliable than estimates based on genetic evidence and molecular clocks, which they contend are subject to genetic drift, bottlenecks and other complicating factors.

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Neanderthals

Multiregionalists claimed that the discovery of a possible hybrid Homo sapiens X neanderthalensis fossil child at the Abrigo do Lagar Velho rock-shelter site in Portugal in 1999 further supports the multiregional hypothesis, by reflecting the admixture of diverse human populations.[10] Two other archaeologists dispute this: "the analysis by Duarte et al. of the Lagar Velho child's skeleton is a brave and imaginative interpretation, of which it is unlikely that a majority of paleoanthropologists will consider proven."[11]

In an article appearing in the Proceedings of the National Academy of Sciences[12] in 2007, Erik Trinkaus has brought together the available data, which shows that early modern humans did exhibit evidence of Neanderthal traits, saying, "When you look at all of the well dated and diagnostic early modern European fossils, there is a persistent presence of anatomical features that were present among the Neandertals but absent from the earlier African modern humans...Early modern Europeans reflect both their predominant African early modern human ancestry and a substantial degree of admixture between those early modern humans and the indigenous Neandertals."[13]

Peking man

Shang et al. see continuity in skeletal remains of archaic people from east Asia.[14]

Early modern humans

Wolpoff and colleagues published an analysis in 2001 of character traits of the skulls of early modern human fossils, which failed to reject a theory of dual ancestry from Javan Homo erectus for Australian early modern humans and Neanderthals for Central European modern humans, and which they said ruled out a replacement model.[15] A subsequent analysis comparing differences of Neanderthal skulls to those of modern humans using 3D morphometric techniques showed a large difference between the two populations, such that Harvati & al concluded that "we interpret the evidence presented here as supporting the view that Neanderthals represent an extinct human species and therefore refute the regional continuity model for Europe."[16] It has been argued that these differences are consistent with an evolving lineage, as ancestors are never identical to their descendants.[17]

New early modern human remains were unearthed in 2003 in Tianyuan Cave, Zhoukoudian. 14C dated 42-39 ky Tianyuan 1 holotype is the oldest, directly dated EMH in eastern Eurasia. Tianyuan 1 exhibits a series of typical modern, derived modern human features and few archaic traits. Some late archaic human traits include a large hamulus length, anterior to posterior dental proportions and a broad and rounded distal phalangeal tuberosityhis.[18]

The oldest European EMH remains were discovered in 2002 in cave named Peştera cu Oase near the Iron Gates in the Danubian corridor. Oase 1 holotype revealed specific traits combining a variety of archaic Homo traits, derived early modern humans, and possibly Neanderthal features. Modern human attributes place it close to European early modern humans among Late Pleistocene samples. The fossil belongs to the few findings in Europe which could be directly dated and is considered the oldest known early modern human fossil from Europe. Two laboratories independently yielded microfiltrated collagen with 14C averaging to 34,950 B.P.[19] In Europe around 40-30 ka evolved latitudinal cline between intermixed Neanderthal traits on west and EEMH on east. The human population phenotypic continuity exist in subsequent generations.[10][19]

Genetic evidence

By analysing haplotype data, Alan Templeton found support for three waves of human migration out of Africa, the first being 1.9 million years ago, and concluded that it was impossible that existing Eurasian populations had not interbred with African migrants.[20]

Studies on past population bottlenecks that can be inferred from molecular data have led multiregionalists to conclude that the recent single-origin hypothesis is untenable because there are no population size bottlenecks affecting all genes that are more recent than 2 million years ago.

  • Microcephalin D allele introgressed into the modern human gene pool points to the Neanderthal lineage as possible source and compelling evidence of admixture among the human loci.[21][22][23]
  • RRM2P4[24] (ribonucleotide reductase M2 subunit pseudogene 4) [6]. Present day human population clades A and B have 2.3 Ma time to most recent common ancestor (TMRCA).[25] The gene tree branches of RRM2P4 point to eastern Asian ancestry.[26]
  • PDHA1 (pyruvate dehydrogenase) locus on X chromosome has estimated coalescent-time depth of 1.86 Ma.[26][27][28]
  • MAPT locus 17q21.3 split into deep genetic lineages H1 and H2 . H2 lineage in European population suggest inheritance from Neanderthals [29][30][31][32][33].
  • ASAH1. Related to mental activity N-Acylsphingosine Amidohydrolase gene two V and M deep genetic lineages[34] have TMRCA 2.4±.4 Ma.[35] Linkage disequilibrium 62% and small nucleotide diversity 0.05% indicate a signature of positive Darwinian selection for the V lineage. The M lineage is attributed to ancient population structure of humans in Africa.[36]
  • X-chromosome genes DMD44, APXL, AMELX, TNFSF5 show S-N heterogeneous patterns of variation and may play role in diversity-reducing selection in non-Africans[37].
  • CMAH CMP-N-acetylneuraminic acid hydroxylase pseudogene show 2.9 Ma genetic history [38].
  • NAT2 [39] SNPs cluster predictably with linages originating in sub-Saharan Africa, Europe, and East Asia.[40] NAT1*11A 0.29 Ma suggest ancient structured population with gene tree rooted in Eurasia.[41][42].
  • ALMS1 suggest ancient and complex evolutionary history.[43]
  • Genome polymorphism: Inversion polymorphism: known 5-million-base pair (Mbp) 8p23.1, 1-Mbp on 17q21.3 and novel 1.2-Mbp on 15q24, 2.1-Mbp 15q13, 1.7-Mbp 17q12 [10] . In the sample of 8 gnomes from worldwide sample including Yuruba Kidd&al group found 4 million SNPs and 796,273 small indels (1−100 bp in size); 15 large regions of excess nucleotide variation 500 kbp to 3 Mbp. Two of variable sites are described detailed above.[44]

Proponents of the multiregional hypothesis show genetic sequences of several loci in the human genome with million year old genealogy.[45][46][47][48][49][50] Those data of deep genetic lineages are explained in the multiregional theory framework as a result of heredity from structured ancestral population.[51] The data are not interpreted in light of the RAO hypothesis postulating recent replacement where separated million years ago genetic lineages are at best unpredicted.[52][53]

Criticism of the multiregional hypothesis

A competing theory, the recent African origin of modern humans (also known as "Out of Africa"), has emerged as the near consensus view since the 1990s,[3][4] proposing that modern humans arose in Africa around 100-200,000 years ago, moving out of Africa around 50-60,000 years ago to replace existing human species such as Homo erectus and the Neanderthals.[5]

Ancient mitochondrial DNA sequence, extracted from 37,000 years old Neanderthal specimen, and mitochondrial DNA from present day humans have different sequence. However, the largest example of sequenced Neanderthal nuclear DNA comprised 1 million base pairs compared to a human nuclear genome size of roughly 3 billion base pairs. This amounts to a comparison of only 0.033% of the genomes.

See also

References

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  11. ^ Chunky Gravettian child; Ian Tattersall and Jeffrey H. Schwartz
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  19. ^ a b Trinkaus, E; Moldovan, O; Milota, S; Bîlgăr, A; Sarcina, L; Athreya, S; Bailey, Se; Rodrigo, R; Mircea, G; Higham, T; Ramsey, Cb; Van, Der, Plicht, J (Sep 2003). "An early modern human from the Peştera cu Oase, Romania" (Free full text). Proceedings of the National Academy of Sciences of the United States of America 100 (20): 11231–6. doi:10.1073/pnas.2035108100. ISSN 0027-8424. PMID 14504393.& PMC 208740. http://www.pnas.org/cgi/pmidlookup?view=long&pmid=14504393. ""When multiple measurements are undertaken, the mean result can be determined through averaging the activity ratios. For Oase 1, this provides a weighted average activity ratio of 〈14a〉 = 1.29 ± 0.15%, resulting in a combined OxA-GrA 14C age of 34,950, +990, and –890 B.P."". 
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  22. ^ Trinkaus, E (May 2007). "European early modern humans and the fate of the Neandertals" (Free full text). Proceedings of the National Academy of Sciences of the United States of America 104 (18): 7367–72. doi:10.1073/pnas.0702214104. ISSN 0027-8424. PMID 17452632.& PMC 1863481. http://www.pnas.org/cgi/pmidlookup?view=long&pmid=17452632. 
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  24. ^ sequence and gene tree for RRM2P4 haplotypes oxfordjournals.org
  25. ^ Time-scaled gene tree of the RRM2P4. TMRCA between A, B clades 2.3 Mya jpeg
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  27. ^ Rosalind M. Harding (March 16, 1999). "More on the X files". Proceedings of the National Academy of Sciences 96 (6): 2582–2584. doi:10.1073/pnas.96.6.2582. http://www.pnas.org/cgi/content/full/96/6/2582. ""the pattern of diversity at the PDHA1 locus unexpected is that this extreme structure is observed in a polymorphism with an estimated total coalescent-time depth of 1.86 million years"". 
  28. ^ Harris, E. E. ;Jody Hey (1999). [pdf "X chromosome evidence for ancient human histories"]. Proceedings of the National Academy of Sciences 96: 3320. doi:10.1073/pnas.96.6.3320. pdf. 
  29. ^ J. Hardy, A. Pittman, A. Myers, K. Gwinn-Hardy, H.C. Fung, R. de Silva, M. Hutton and J. Duckworth (2005). "Evidence suggesting that Homo neanderthalensis contributed the H2 MAPT haplotype to Homo sapiens". Biochemical Society Transactions 33, part 4;. http://www.google.com/url?sa=t&source=web&ct=res&cd=1&url=http%3A%2F%2Fwww.biochemsoctrans.org%2Fbst%2F033%2F0582%2F0330582.pdf&ei=HqA7SpzqFoPoNJXMtbQO&rct=j&q=Evidence+suggesting+that+Homo+neanderthalensis+contributed+the+H2+MAPT+haplotype+to+Homo+sapiens&usg=AFQjCNFvcjikMrJcuAZsfxmKc_bZ6f0vMA. ""We suggest that the H2 haplotype is derived from Homo neanderthalensis and entered H. sapiens populations during the coexistence of these species in Europe from approx. 45 000 to 18 000 years ago and that the H2 haplotype has been under selection pressure since that time, possibly because of the role of this H1 haplotype in neurodegenerative disease."..."The tau (MAPT ) locus is very unusual. Over a region of approx. 1.8 Mb, there are two haplotype clades in European populations, H1 and H2 [6,7]. In other populations, only the H1 occurs and shows a normal pattern of recombination"". 
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  34. ^ ASAH1 SL and ML region SNP DNA seguences jpeg
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  40. ^ Multidimensional scaling of genetic distances indicating a very good fit of the projection to the original samples data from sub-Saharan Africa, Europe, and East Asia map
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  47. ^ X chromosome evidence for ancient human histories; Eugene E. Harris and Jody Hey; PNAS March 16, 1999 vol. 96 no. 6 3320-3324 [4]
  48. ^ A common inversion under selection in Europeans; Stefansson H, Helgason A, Thorleifsson G, Steinthorsdottir V, Masson G, Barnard J, Baker A, Jonasdottir A, Ingason A, Gudnadottir VG, et al. Nature Genetics 37, 129 - 137 (2005) Published online: 16 January 2005; doi:10.1038/ng1508
  49. ^ Evidence that the adaptive allele of the brain size gene microcephalin introgressed into Homo sapiens from an archaic Homo lineage; Patrick D. Evans, Nitzan Mekel-Bobrov, Eric J. Vallender, Richard R. Hudson and Bruce T. Lahn; PNAS November 28, 2006 vol. 103 no. 48 18178-18183 [5]
  50. ^ Early modern human diversity suggests subdivided population structure and a complex out-of-Africa scenario Philipp Gunza, Fred L. Booksteina, Philipp Mitteroeckera, Andrea Stadlmayra, Horst Seidlera and Gerhard W. Webera; 10.1073/pnas.0808160106 [6]
  51. ^ [7]
  52. ^ Ancient lineages in the genome: A response to Fagundes et al; Daniel Garrigan and Michael F. Hammer; doi:10.1534/genetics.105.041095 [8]
  53. ^ Reply to Garrigan and Hammer: Ancient lineages and assimilation; Nelson J. R. Fagundes, Nicolas Ray, Mark Beaumont, Samuel Neuenschwande, Francisco M. Salzano†, Sandro L. Bonatto and Laurent Excoffier ;10.1073/pnas.0711261105 [9] quote:We must repeat that our results do not exclude the occurrence of some admixture events between modern and archaic humans,

Reviews

External links

  • [11] - 'Genomics refutes an exclusively African origin of humans' (pdf) Vinayak Eswaran, Henry Harpending, Alan R. Rogers, Journal of Human Evolution (2005)
  • [12] - 'Templeton tree'
  • [13] - 'The Hybrid Child from Portugal'
  • Biochem. Soc. Trans (2005) 33, 582-585 - J. Hardy and others - Molecular Mechanisms of Neurodegeneration (Evidence suggesting that Homo neanderthalensis contributed the H2 MAPT haplotype to Homo sapiens)
  • Kent Holsinger's web site - 'Drift and migration' (only 1 migrant per generation between populations of reasonable big sizes can prevent divergence in allelic frequencies)
  • Genetics - 'Deep Haplotype Divergence and Long-Range Linkage Disequilibrium at Xp21.1 Provide Evidence That Humans Descend From a Structured Ancestral Population' (first genetic evidence that statistically rejects the null hypothesis that our species descends from a single, historically panmictic population), Daniel Garrigan, Zahra Mobasher, Sarah B. Kingan, Jason A. Wilder, and Michael F. Hammer, University of Arizona, Tucson, Genetics, Vol. 170, 1849–1856, August 2005
  • Linfield.edu - 'The Origin of Modern Humans: Multiregional and Replacement Theories', Michael Roberts, Linfield College
  • [14] - 'Evidence for Archaic Asian Ancestry on the Human X Chromosome' (suggests ancient RRM2P4 lineage is remnant of introgressive hybrid of anatomically modern humans from Africa and archaic populations in Eurasia), Daniel Garrigan, Zahra Mobasher, Tesa Severson, Jason A. Wilder, Michael F. Hammer, University of Arizona, Tucson, Molecular Biology and Evolution, vol 22, no 2, p 189–192 (2005)
  • PNAS.org - 'Mitochondrial DNA sequences in ancient Australians: Implications for modern human origins', Gregory J. Adcock, Elizabeth S. Dennis, Simon Easteal, Gavin A. Huttley, Lars S. Jermiin, W. James Peacock, Alan Thorne, Australian National University, Proceedings of the National Academy of Sciences, vol 98, no 2, p 537-542 (January 16, 2001)
  • StephenJayGould.org - 'Out of Africa vs. Multiregionalism', Tod Billings (December 7, 1999)
  • TalkOrigins.org - 'The evolution of modern humans: where are we now?' Christopher B. Stringer, General Anthropology, vol 7, no 2, p 1–5 (2001)
  • Selection, nuclear genetic variation, and mtDNA

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