DNA history of Egypt

The genetic history of the demographics of Egypt reflects Egypt's geographical location at the crossroads of several major cultural areas: Northeast Africa, Northwest Africa, the Sahara, Sub-Saharan Africa, the Middle East, and the Mediterranean.

In general, various DNA studies have found that the gene frequencies of modern Egyptian populations are intermediate between those of the Middle East, the Horn of Africa, southern Europe and Sub-Saharan Africa,[1] though NRY frequency distributions of the modern Egyptian population appear to be substantially more similar to those of the Middle East than to any Sub-Saharan African or European population, suggesting a substantially larger Middle Eastern genetic component.[2][3][3][4][5][6][7]

Ancient DNA

Contamination from handling and intrusion from microbes create obstacles to the recovery of ancient DNA.[8] Consequently, most DNA studies have been carried out on modern Egyptian populations with the intent of learning about the influences of historical migrations on the population of Egypt.[9][10][11][12] One successful 1993 study was performed on ancient mummies of the 12th Dynasty, by Dr. Svante Pääbo and Dr. Anna Di Rienzo, which identified multiple lines of descent, some of which originated in Sub-Saharan Africa.[13]

Blood typing and ancient DNA sampling on Egyptian mummies is scant. However, blood typing of dynastic period mummies found their ABO frequencies to be most similar to that of modern Egyptians,[14] and some also to northern Haratin populations.

DNA studies on modern Egyptians

In general, various DNA studies have found that the gene frequencies of modern North African populations are intermediate between those of the Horn of Africa and Eurasia,[15] though possessing a greater genetic affinity with the populations of Eurasia than they do with sub-Saharan Africa.[3][3][4] The present population of the Sahara is more Caucasoid (though still mixed) in the extreme north, with a fairly gradual increase of Negroid component as one goes south.[6][16][17] The results of these genetic studies is consistent with the historical record, which records significant bidirectional contact between Egypt and Nubia, and the Levant/Middle East within the last few thousand years, but with general population continuity from the Early Dynastic Period of Egypt up to the modern day era.[18][19]

Genetic analysis of modern Egyptians reveals that they have paternal lineages common to indigenous North-East African populations primarily (Algeria, Tunisia, Morocco), and to Middle Eastern peoples to a lesser extent—these lineages would have spread during the Neolithic and were maintained by the predynastic period.[6][20]

A study by Krings et al. (1999) on mitochondrial DNA clines along the Nile Valley found that a Eurasian cline runs from Northern Egypt to Southern Sudan and a Sub-Saharan cline from Southern Sudan to Northern Egypt.[21]

Luis et al. (2004) found that the male haplogroups in a sample of 147 Egyptians were E1b1b (36.1%, predominantly E-M78), J (32.0%), G (8.8%), T(8.2%), and R (7.5%). E1b1b and its subclades are characteristic of some Afro-Asiatic speakers and are believed to have originated in either the Middle East, North Africa, or the Horn of Africa. Cruciani et al. (2007) suggests that E-M78, E1b1b predominant subclade in Egypt, originated in "Northeastern Africa", which in the study refers specifically to Egypt and Libya [2][22]

Other studies have shown that modern Egyptians have genetic affinities primarily with populations of Asia, North and Northeast Africa,[23][24][25][26] and to a lesser extent Middle Eastern and European populations.[27]

Some genetic studies done on modern Egyptians suggest a more distant relationship to Sub Saharan Africans.[28] and a closer link to other North Africans.[25] In addition, some studies suggest lesser ties with populations in the Middle East, as well as some groups in southern Europe.[26] A 2004 mtDNA study of upper Egyptians from Gurna found a genetic ancestral heritage to modern Northeast Africans, characterized by a high M1 haplotype frequency and a comparatively low L1 and L2 macrohaplogroup frequency of 20.6%. Another study links Egyptians in general with people from modern Eritrea and Ethiopia.[24][29] Though there has been much debate of the origins of haplogroup M1 a recent 2007 study had concluded that M1 has West Asia origins not a Sub Saharan African origin[30] Origin A 2003 Y chromosome study was performed by Lucotte on modern Egyptians, with haplotypes V, XI, and IV being most common. Haplotype V is common in Berbers and has a low frequency outside North Africa. Haplotypes V, XI, and IV are all predominantly North African/Horn of African haplotypes, and they are far more dominant in Egyptians than in Middle Eastern or European groups.[31]

Y-DNA haplogroups

A study using the Y-chromosome of modern Egyptian males found similar results, namely that North East African haplogroups are predominant in the South but the predominant haplogroups in the North are characteristic of North African and West Eurasian populations.[32]

Population Nb A/B E1b1a E1b1b1 (M35) E1b1b1a (M78) E1b1b1b (M81) E1b1b1c (M123) F K G I J1 J2 R1a R1b Other Study
1 Egyptians 1472.7%2.7%018.4%8.2%9.5%07.5%9.5%019.7%12.2%3.4% 4.1%2.1%Luis et al. (2004)[33]
2 Egyptians from El-Hayez Oasis (Western Desert)3505.70%5.7%28.6%28.6%0000031.4%0000Kujanová et al. (2009)[34]
3 Egyptians from Siwa Oasis (Western Desert)9328.0%6.5%2.2%6.5%1.1%2.2%003.2%07.5% 6.5%028.0%8.3%Dugoujon et al. (2009)[35]
4 Northern Egyptians 442.3%04.5%27.3%11.4%9.1%6.8%2.3%009.1%9.1%2.3%9.9%6.8%Arredi et al. (2004)
5 Southern Egyptians 290.0%0017.2%6.9%6.9%17.2%10.3%03.4%20.7%3.4%013.8%0Arredi et al. (2004)
Distribution of E1b1b1a (E-M78) and its subclades
Population N E-M78 E-M78* E-V12* E-V13 E-V22 E-V32 E-V65 Study
Southern Egyptians 7950.6%44.3%1.3%3.8%1.3% Cruciani et al. (2007)[36]
Egyptians from Bahari 4141.4%14.6%2.4%21.9%2.4% Cruciani et al. (2007)
Northern Egyptians (Delta) 7223.6%5.6%1.4%13.9%2.8% Cruciani et al. (2007)
Egyptians from Gurna Oasis 3417.6%5.9%8.8%2.9% Cruciani et al. (2007)
Egyptian from Siwa Oasis 936.4%2.1%4.3% Cruciani et al. (2007)

Autosomal DNA

On 13 January 2012, an exhaustive genetic study of North Africa's human populations was published.[37] The researchers analyzed around 800,000 genetic markers, distributed throughout the entire genome in 125 North African individuals belonging to seven representative populations in the whole region (Saharawi, South Moroccans, North Moroccans, Algerians, Tunisians Berbers, Libyans and Egyptians) and the information obtained was compared with the information from the neighbouring populations. The results of this study show that there is a native genetic component ("Maghrebi") which defines North Africans. The study identified mainly two distinct, opposite gradients of ancestry: an east-to-west increase of this native North African ancestry and an east-to-west decrease in likely Middle Eastern Arab ancestry.

The study also reveals that the genetic composition of North Africa's human populations is very complex and is the result of five distinct ancestries : a local component (Maghrebi) dating back thirteen thousand years and the varied genetic influence of neighbouring populations on North African groups during successive migrations (European, Middle Eastern, eastern and western Sub-Saharan Africa). According to the authors, the people inhabiting North Africa today are not descendants of either the earliest occupants of this region fifty thousand years ago, or descendants of the most recent Neolithic populations. The data shows that the ancestors of today's North Africans were a group of populations which already lived in the region around thirteen thousand years ago. Furthermore, this local North African genetic component is very different from the one found in the populations in the south of the Sahara, which shows that the ancestors of today's North Africans were members of a subgroup of humanity who left Africa to conquer the rest of the world and who subsequently returned to the north of the continent to settle in the region. As well as this local component, North African populations were also observed to share genetic markers with all the neighbouring regions, as a result of more recent migrations, although these appear in different proportions. There is an influence from the Middle East, which becomes less marked as the distance from the Arabian Peninsula increases, similar proportions of European influence in all North African populations, and, in some populations (South Moroccans, Saharawi...), there are even individuals who present a large proportion of recent influence from the South of the Sahara in their genome.

Admixture analysis

Recent genetic analysis of North African populations have found that, despite the complex admixture genetic background, there is an autochthonous genomic component which is likely derived from "back-to-Africa" gene flow older than 12,000 years ago (ya) (i.e., prior to the Neolithic migrations). This local population substratum seems to represent a genetic discontinuity with the earliest modern human settlers of North Africa (those with the Aterian industry) given the estimated ancestry is younger than 40,000 years ago. North Morocco, Libya and Egypt carry high proportions of European and Middle Eastern ancestral components, whereas Tunisian Berbers and Saharawi are those populations with highest autochthonous North African component.[38]

Copts

According to Y-DNA analysis by Hassan et al. (2008), around 45% of Copts carry the haplogroup J. The remainder mainly belong to the E1b1b clade (21%). Both paternal lineages are common among local Afro-Asiatic-speaking populations,[39] with E1b1b/E3b reaching its highest frequencies among Berbers.[40] The next most common haplogroups borne by Copts are the Western European-linked R1b clade (15%), and the archaic African B lineage (15%).[39]

Maternally, Hassan (2009) found that Copts exclusively carry various descendants of the macrohaplogroup N. This mtDNA clade is likewise closely associated with local Afro-Asiatic-speaking populations. Of the N derivatives borne by Copts, U6 is most frequent (28%), followed by the haplogroup T (17%).[41]

A 2015 study by Dobon et al. identified an ancestral autosomal component of West Eurasian origin that is common to many modern Afroasiatic-speaking populations in Northeast Africa. Known as the Coptic component, it peaks among Egyptian Copts who settled in Sudan over the past two centuries. Copts also formed a separated group in PCA, a close outlier to other Egyptians, Afro-Asiatic-speaking Northeast Africans and Middle East populations. The Coptic component evolved out of a main Northeast African and Middle Eastern ancestral component that is shared by other Egyptians and also found at high frequencies among other Afro-Asiatic-speaking populations in Northeast Africa (~70%). The scientists suggest that this points to a common origin for the general population of Egypt. They also associate the Coptic component with Ancient Egyptian ancestry, without the later Arabian influence that is present among other Egyptians.[42]

See also

References

  1. ↑ Cavalli-Sforza, History and Geography of Human Genes, The intermediacy of North Africa and to a lesser extent Europe is apparent
  2. 1 2 Luis, J. R.; et al. (2004). "The Levant versus the Horn of Africa: Evidence for Bidirectional Corridors of Human Migrations" (PDF). American Journal of Human Genetics. 74 (3): 532–544. doi:10.1086/382286. PMC 1182266Freely accessible. PMID 14973781. Archived from the original (PDF) on February 16, 2012.
  3. 1 2 3 4 Cavalli-Sforza, L.L., P. Menozzi, and A. Piazza. 1994. The History and Geography of Human Genes. Princeton: Princeton University Press.
  4. 1 2 Bosch, E.; Calafell, F; Pérez-Lezaun, A; Comas, D; Mateu, E; Bertranpetit, J (1997). "Population history of north Africa: evidence from classical genetic markers". Human Biology. 69 (3): 295–311. PMID 9164042.
  5. ↑ Arredi B, Poloni E, Paracchini S, Zerjal T, Fathallah D, Makrelouf M, Pascali V, Novelletto A, Tyler-Smith C; Poloni; Paracchini; Zerjal; Fathallah; Makrelouf; Pascali; Novelletto; Tyler-Smith (2004). "A predominantly Nilo Saharan origin for Y-chromosomal DNA variation in North Africa". Am J Hum Genet. 75 (2): 338–45. doi:10.1086/423147. PMC 1216069Freely accessible. PMID 15202071.
  6. 1 2 3 Manni F, Leonardi P, Barakat A, Rouba H, Heyer E, Klintschar M, McElreavey K, Quintana-Murci L; Leonardi; Barakat; Rouba; Heyer; Klintschar; McElreavey; Quintana-Murci (2002). "Y-chromosome analysis in Egypt suggests a genetic regional continuity in Northeastern Africa". Hum Biol. 74 (5): 645–58. doi:10.1353/hub.2002.0054. PMID 12495079.
  7. ↑ Cavalli-Sforza. "Synthetic maps of Africa". The History and Geography of Human Genes. ISBN 978-0-691-08750-4.The present population of the Sahara is Sudan in the extreme north, with an increase of Negroid component as one goes south
  8. ↑ Encyclopedia of the Archaeology of Ancient Egypt By Kathryn A. Bard, Steven Blake Shubert pp 278-279
  9. ↑ S.O.Y. Keita & A. J. Boyce, S. O. Y.; Boyce, A. J. (Anthony J.) (June 2009). "Genetics, Egypt, and History: Interpreting Geographical Patterns of Y Chromosome Variation". History in Africa. 32 (1): 221–246. doi:10.1353/hia.2005.0013.
  10. ↑ Shomarka Keita (2005), S. O. Y. (2005). "Y-Chromosome Variation in Egypt". African Archaeological Review. 22 (2): 61–75. doi:10.1007/s10437-005-4189-4.
  11. ↑ Keita, S.O.Y. (2005). "History in the Interpretation of the Pattern of p49a,f TaqI RFLP Y-Chromosome Variation in Egypt". American Journal of Human Biology. 17 (5): 559–67. doi:10.1002/ajhb.20428. PMID 16136533.
  12. ↑ Shomarka Keita: What genetics can tell us
  13. ↑ Paabo, S., and A. Di Rienzo, A molecular approach to the study of Egyptian history. In Biological Anthropology and the Study of Ancient Egypt. V. Davies and R. Walker, eds. pp. 86-90. London: British Museum Press. 1993
  14. ↑ Borgognini Tarli, S.M.; Paoli, G. (1982). "Survey on paleoserological studies". Homo. 33: 69–89.
  15. ↑ Cavalli-Sforza, History and Geography of Human Genes, The intermediacy of North Africa and to lesser extent East Africa between Africa and Europe is apparent
  16. ↑ Arredi B, Poloni E, Paracchini S, Zerjal T, Fathallah D, Makrelouf M, Pascali V, Novelletto A, Tyler-Smith C; Poloni; Paracchini; Zerjal; Fathallah; Makrelouf; Pascali; Novelletto; Tyler-Smith (2004). "A predominantly neolithic origin for Y-chromosomal DNA variation in North Africa". Am J Hum Genet. 75 (2): 338–45. doi:10.1086/423147. PMC 1216069Freely accessible. PMID 15202071.
  17. ↑ Cavalli-Sforza. "Synthetic maps of Africa". The History and Geography of Human Genes. ISBN 978-0-691-08750-4.The present population of the Sahara is Caucasoid in the extreme north, with a fairly gradual increase of Negroid component as one goes south
  18. ↑ Krings, T; Salem, AE; Bauer, K; Geisert, H; Malek, AK; Chaix, L; Simon, C; Welsby, D; et al. (1992). "mtDNA Analysis of Nile River Valley Populations: Genetic Corridor or a Barrier to Migration?" (PDF). Am J Hum Genet. 64 (5): 1116–76. doi:10.1086/302314. PMC 1377841Freely accessible. PMID 10090902.
  19. ↑ Lucotte, G.; Mercier, G. (2001). "Brief communication: Y-chromosome haplotypes in Egypt". American Journal of Physical Anthropology. 121 (1): 63–6. doi:10.1002/ajpa.10190. PMID 12687584.
  20. ↑ Arredi B, Poloni E, Paracchini S, Zerjal T, Fathallah D, Makrelouf M, Pascali V, Novelletto A, Tyler-Smith C; Poloni; Paracchini; Zerjal; Fathallah; Makrelouf; Pascali; Novelletto; Tyler-Smith (2004). "A Predominantly Neolithic Origin for Y-Chromosomal DNA Variation in North Africa". Am J Hum Genet. 75 (2): 338–45. doi:10.1086/423147. PMC 1216069Freely accessible. PMID 15202071.
  21. ↑ Kings, T; Salem, AE; Bauer, K; Geisert, H; Malek, AK; Chaix, L; Simon, C; Welsby, D; et al. (1992). "mtDNA Analysis of Nile River Valley Populations: Genetic Corridor or a Barrier to Migration?" (PDF). Am J Hum Genet. 64 (5): 1116–76. doi:10.1086/302314. PMC 1377841Freely accessible. PMID 10090902.
  22. ↑ Underhill (2002), Bellwood and Renfrew, ed., Inference of Neolithic Population Histories using Y-chromosome Haplotypes, Cambridge: McDonald Institute for Archaeological Research, ISBN 978-1-902937-20-5.
  23. ↑ Kivisild T, Reidla M, Metspalu E, et al. (2004). "Ethiopian mitochondrial DNA heritage: tracking gene flow across and around the gate of tears". Am. J. Hum. Genet. 75 (5): 752–70. doi:10.1086/425161. PMC 1182106Freely accessible. PMID 15457403.
  24. 1 2 Stevanovitch A, Gilles A, Bouzaid E, et al. (2004). "Mitochondrial DNA sequence diversity in a sedentary population from Egypt". Ann. Hum. Genet. 68 (Pt 1): 23–39. doi:10.1046/j.1529-8817.2003.00057.x. PMID 14748828.
  25. 1 2 Arredi B, Poloni E, Paracchini S, Zerjal T, Fathallah D, Makrelouf M, Pascali V, Novelletto A, Tyler-Smith C; Poloni; Paracchini; Zerjal; Fathallah; Makrelouf; Pascali; Novelletto; Tyler-Smith (2004). "A predominantly neolithic origin for Y-chromosomal DNA variation in North Africa". Am J Hum Genet. 75 (2): 338–45. doi:10.1086/423147. PMC 1216069Freely accessible. PMID 15202071.
  26. 1 2 Manni F, Leonardi P, Barakat A, Rouba H, Heyer E, Klintschar M, McElreavey K, Quintana-Murci L; Leonardi; Barakat; Rouba; Heyer; Klintschar; McElreavey; Quintana-Murci (2002). "Y-chromosome analysis in Egypt suggests a genetic regional continuity in Northeastern Africa". Hum Biol. 74 (5): 645–58. doi:10.1353/hub.2002.0054. PMID 12495079.
  27. ↑ Luca Cavalli-Sforza, Luigi; Paolo Menozzi; Alberto Piazza (1996-08-05). The History and Geography of Human Genes. Princeton University Press. ISBN 978-0-691-02905-4.
  28. ↑ Cavalli-Sforza, L.L., P. Menozzi, and A. Piazza. 1994, The History and Geography of Human Genes. Princeton:Princeton University Press,
  29. ↑ Kivisild T, Reidla M, Metspalu E, Rosa A, Brehm A, Pennarun E, Parik J, Geberhiwot T, Usanga E; et al. (2004). "Ethiopian mitochondrial DNA heritage: tracking gene flow across and around the gate of tears". Am J Hum Genet. 75 (5): 752–70. doi:10.1086/425161. PMC 1182106Freely accessible. PMID 15457403.
  30. ↑ González, A. M.; Larruga, J. M.; Abu-Amero, K. K.; Shi, Y; Pestano, J; Cabrera, V. M. (2007). "Mitochondrial lineage M1 traces an early human backflow to Africa". BMC Genomics. 8: 223. doi:10.1186/1471-2164-8-223. PMC 1945034Freely accessible. PMID 17620140.
  31. ↑ Keita, S.O. (2005). "History in the interpretation of the pattern of p49a, f TaqI RFLP Y-chromosome variation in Egypt: a consideration of multiple lines of evidence". Am J Hum Biol. 17 (5): 559–67. doi:10.1002/ajhb.20428. PMID 16136533.
  32. ↑ Lucotte, G.; Mercier, G. (2001). "Brief communication: Y-chromosome haplotypes in Egypt". American Journal of Physical Anthropology. 121 (1): 63–6. doi:10.1002/ajpa.10190. PMID 12687584.
  33. ↑ Luis, JR; Rowold, DJ; Regueiro, M; Caeiro, B; Cinnioglu, C; Roseman, C; Underhill, PA; Cavalli-Sforza, LL; Herrera, RJ (2004). "The Levant versus the Horn of Africa: evidence for bidirectional corridors of human migrations". Am J Hum Genet. 74 (3): 532–544. doi:10.1086/382286. PMC 1182266Freely accessible. PMID 14973781.
  34. ↑ Martina Kujanová, Luísa Pereira, Verónica Fernandes, Joana B. Pereira, Viktor Černý, Near Eastern Neolithic genetic input in a small oasis of the Egyptian Western Desert, American Journal of Physical Anthropology, Volume 140, Issue 2, pages 336–346, October 2009
  35. ↑ Dugoujon J.M., Coudray C., Torroni A., Cruciani F., Scozzari F., Moral P., Louali N., Kossmann M. The Berber and the Berbers: Genetic and linguistic diversities
  36. ↑ Cruciani, F.; La Fratta, R.; Trombetta, B.; Santolamazza, P.; Sellitto, D.; Colomb, E. B.; Dugoujon, J.-M.; Crivellaro, F.; et al. (2007). "Tracing Past Human Male Movements in Northern/Eastern Africa and Western Eurasia: New Clues from Y-Chromosomal Haplogroups E-M78 and J-M12" (PDF). Molecular Biology and Evolution. 24 (6): 1300–1311. doi:10.1093/molbev/msm049. PMID 17351267.
  37. ↑ Henn, BM; Botigué, LR; Gravel, S; Wang, W; Brisbin, A; et al. (2012). "Genomic Ancestry of North Africans Supports Back-to-Africa Migrations". PLoS Genet. 8 (1): e1002397. doi:10.1371/journal.pgen.1002397. PMC 3257290Freely accessible. PMID 22253600.
  38. ↑ North African Populations Carry the Signature of Admixture with Neandertals. Sánchez-Quinto F, Botigué LR, Civit S, Arenas C, Ávila-Arcos MC, et al. (2012) North African Populations Carry the Signature of Admixture with Neandertals" PLoS ONE 7(10) e47765. doi:10.1371/journal.pone.0047765
  39. 1 2 Hassan, Hisham Y. et al. (2008). "Y‐chromosome variation among Sudanese: Restricted gene flow, concordance with language, geography, and history" (PDF). American Journal of Physical Anthropology. 137 (3): 316–323. Retrieved 14 October 2016.
  40. ↑ Trombetta, Beniamino et al. (2015). "Phylogeographic refinement and large scale genotyping of human Y chromosome haplogroup E provide new insights into the dispersal of early pastoralists in the African continent" (PDF). Genome Biology and Evolution. 7 (7): 1940–1950. doi:10.1093/gbe/evv118. Retrieved 13 October 2016.; Supplementary Table 7
  41. ↑ Mohamed, Hisham Yousif Hassan. "Genetic Patterns of Y-chromosome and Mitochondrial DNA Variation, with Implications to the Peopling of the Sudan" (PDF). University of Khartoum. Retrieved 13 October 2016.
  42. ↑ Begoña Dobon, Hisham Y. Hassan, Hafid Laayouni, Pierre Luisi, Isis Ricaño-Ponce, Alexandra Zhernakova, Cisca Wijmenga, Hanan Tahir, David Comas, Mihai G. Netea, Jaume Bertranpetit (28 May 2015). "The genetics of East African populations: a Nilo-Saharan component in the African genetic landscape" (PDF). Scientific Reports. 5: 9996. doi:10.1038/srep09996. Retrieved 13 June 2015.
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