Early Back-to-Africa Migration into the Horn of Africa
The Horn of Africa (HOA) occupies a central place in our understanding of modern human origins. This region is the location of the earliest known modern human fossils, a possible source for the out-of-Africa migration, and one of the most genetically and linguistically diverse regions of the world. Numerous genetic studies over the last decades have identified substantial non-African ancestry in populations in this region. Because there is archaeological, historical, and linguistic evidence for contact with non-African populations beginning about 3,000 years ago, it has often been assumed that the non-African ancestry in HOA populations dates to this time. In this work, we find that the genetic composition of non-African ancestry in the HOA is distinct from the genetic composition of current populations in North Africa and the Middle East. With these data, we demonstrate that most non-African ancestry in the HOA cannot be the result of admixture within the last few thousand years, and that the majority of admixture probably occurred prior to the advent of agriculture. These results contribute to a growing body of work showing that prehistoric hunter-gatherer populations were much more dynamic than usually assumed.
Vyšlo v časopise:
Early Back-to-Africa Migration into the Horn of Africa. PLoS Genet 10(6): e32767. doi:10.1371/journal.pgen.1004393
Kategorie:
Research Article
prolekare.web.journal.doi_sk:
https://doi.org/10.1371/journal.pgen.1004393
Souhrn
The Horn of Africa (HOA) occupies a central place in our understanding of modern human origins. This region is the location of the earliest known modern human fossils, a possible source for the out-of-Africa migration, and one of the most genetically and linguistically diverse regions of the world. Numerous genetic studies over the last decades have identified substantial non-African ancestry in populations in this region. Because there is archaeological, historical, and linguistic evidence for contact with non-African populations beginning about 3,000 years ago, it has often been assumed that the non-African ancestry in HOA populations dates to this time. In this work, we find that the genetic composition of non-African ancestry in the HOA is distinct from the genetic composition of current populations in North Africa and the Middle East. With these data, we demonstrate that most non-African ancestry in the HOA cannot be the result of admixture within the last few thousand years, and that the majority of admixture probably occurred prior to the advent of agriculture. These results contribute to a growing body of work showing that prehistoric hunter-gatherer populations were much more dynamic than usually assumed.
Zdroje
1. HowellsWW (1976) Explaining modern man: Evolutionists Versus Migrationists. J Hum Evol 5: 477–495 doi:10.1016/0047-2484(76)90088-9
2. StringerCB, AndrewsP (1988) Genetic and fossil evidence for the origin of modern humans. Science 239: 1263–1268 doi:10.1126/science.3125610
3. RosenbergNA, PritchardJK, WeberJL, CannHM, KiddKK, et al. (2002) Genetic structure of human populations. Science 298: 2381–2385 doi:10.1126/science.1078311
4. Mitchell P (2005) African Connections: Archaeological Perspectives on Africa and the Wider World. Walnut Creek, CA: AltaMira Press.
5. TempletonAR (2005) Haplotype trees and modern human origins. Yrbk Phys Anthropol 48: 33–59 doi:10.1002/ajpa.20351
6. Templeton AR (2007) Population biology and population genetics of Pleistocene Hominins. In: Henke W, Tattersall I, editors. Handbook of Paleoanthropology. Berlin: Springer-Verlag, Vol. 3. pp. 1825–1859.
7. TempletonAR (2013) Biological races in humans. Stud Hist Phil Biol Biomed Sci 44: 262–271 doi:10.1016/j.shpsc.2013.04.010
8. DiamondJ, BellwoodP (2003) Farmers and their languages: the first expansions. Science 300: 597–603 doi:10.1126/science.1078208
9. Bellwood P, Oxenham M (2008) The Expansions of Farming Societies and the Role of the Neolithic Demographic Transition. In: Bocquet-Appel J-P, Bar-Yosef O, editors. The Neolithic Demographic Transition and its Consequences. New York: Springer. pp. 13–34.
10. PriceTD, Bar-YosefO (2011) The Origins of Agriculture: New Data, New Ideas. Curr Anthropol 52: S163–S174 doi:10.1086/659964
11. Levine DM (2000) Greater Ethiopia: The Evolution of a Multiethnic Society. Second Edition. Chicago: University of Chicago Press.
12. Cavalli-Sforza LLL, Menozzi P, Piazza A (1994) History And Geography Of Human Genes. Princeton: Princeton University Press.
13. PassarinoG, SeminoO, Quintana-MurciL, ExcoffierL, HammerMF, et al. (1998) Different genetic components in the Ethiopian population, identified by mtDNA and Y-chromosome polymorphisms. Am J Hum Genet 62: 420–434 doi:10.1086/301702
14. Ehret C (2002) The Civilizations of Africa: A History to 1800. Charlottesville: University of Virginia Press.
15. NonAL, Al-MeeriA, RaaumRL, SanchezLF, MulliganCJ (2011) Mitochondrial DNA reveals distinct evolutionary histories for Jewish populations in Yemen and Ethiopia. Am J Phys Anthropol 144: 1–10 doi:10.1002/ajpa.21360
16. PaganiL, KivisildT, TarekegnA, EkongR, PlasterC, et al. (2012) Ethiopian Genetic Diversity Reveals Linguistic Stratification and Complex Influences on the Ethiopian Gene Pool. Am J Hum Genet 91: 83–96 doi:10.1016/j.ajhg.2012.05.015
17. PickrellJK, PattersonN, LohP-R, LipsonM, BergerB, et al. (2014) Ancient west Eurasian ancestry in southern and eastern Africa. Proc Natl Acad Sci USA 111: 2632–2637 doi:10.1073/pnas.1313787111
18. KitchenA, EhretC, AssefaS, MulliganCJ (2009) Bayesian phylogenetic analysis of Semitic languages identifies an Early Bronze Age origin of Semitic in the Near East. Proc Roy Soc B Biol Sci 276: 2703–2710 doi:10.1098/rspb.2009.0408
19. Cerulli E (1960) Punti di vista sulla storia dell'Etiopia. Atti del Convegno Internazionale di Studi Etiopici, Roma 1959. Rome: Accademia Nazionale dei Lincei. pp. 5–27.
20. Bent T (1893) The Sacred City of the Ethiopians. London: Longmans, Green.
21. Conti Rossini C (1928) Storia D'Etiopia. Bergamo: Istituto Italiano d'Arti Grafiche.
22. Ullendorff E (1973) The Ethiopians. London: Oxford University Press.
23. RobinC, de MaigretA (1998) Le grand temple de Yèha (Tigray, Éthiopie) après la première campagne de fouilles de la mission française (1998). CR Acad Inscr Belle 142: 737–798.
24. PhillipsonDW (2009) The First Millennium BC in the Highlands of Northern Ethiopia and South–Central Eritrea: A Reassessment of Cultural and Political Development. Afr Archaeol Rev 26: 257–274 doi:10.1007/s10437-009-9064-2
25. FattovichR (2010) The Development of Ancient States in the Northern Horn of Africa, c. 3000 BC–AD 1000: An Archaeological Outline. J World Prehist 23: 145–175 doi:10.1007/s10963-010-9035-1
26. ManzoA (2009) Capra nubiana in Berbere Sauce? Afr Archaeol Rev 26: 291–303 doi:10.1007/s10437-009-9066-0
27. FattovichR (2009) Reconsidering Yeha, c. 800–400 BC. Afr Archaeol Rev 26: 275–290 doi:10.1007/s10437-009-9063-3
28. CurtisMC (2009) Relating the Ancient Ona Culture to the Wider Northern Horn: Discerning Patterns and Problems in the Archaeology of the First Millennium BC. Afr Archaeol Rev 26: 327–350 doi:10.1007/s10437-009-9062-4
29. SchmidtPR (2009) Variability in Eritrea and the Archaeology of the Northern Horn During the First Millennium BC: Subsistence, Ritual, and Gold Production. Afr Archaeol Rev 26: 305–325 doi:10.1007/s10437-009-9061-5
30. Boivin N, Blench R, Fuller DQ (2010) Archaeological, Linguistic and Historical Sources on Ancient Seafaring: A Multidisciplinary Approach to the Study of Early Maritime Contact and Exchange in the Arabian Peninsula. In: Petraglia MD, Rose JI, editors. The Evolution of Human Populations in Arabia. New York: Springer. pp. 251–278.
31. Khalidi L (2010) Holocene Obsidian Exchange in the Red Sea Region. In: Petraglia MD, Rose JI, editors. The Evolution of Human Populations in Arabia. New York: Springer. pp. 279–291.
32. Quintana-MurciL, SeminoO, BandeltH-J, PassarinoG, McElreaveyK, et al. (1999) Genetic evidence of an early exit of Homo sapiens sapiens from Africa through eastern Africa. Nat Genet 23: 437–441 doi:10.1038/70550
33. KivisildT, ReidlaM, MetspaluE, RosaA, BrehmA, et al. (2004) Ethiopian mitochondrial DNA heritage: tracking gene flow across and around the gate of tears. Am J Hum Genet 75: 752–770 doi:10.1086/425161
34. OlivieriA, AchilliA, PalaM, BattagliaV, FornarinoS, et al. (2006) The mtDNA legacy of the Levantine early Upper Palaeolithic in Africa. Science 314: 1767–1770 doi:10.1126/science.1135566
35. GonzálezAM, LarrugaJM, Abu-AmeroKK, ShiY, PestanoJ, et al. (2007) Mitochondrial lineage M1 traces an early human backflow to Africa. BMC Genomics 8: 223 doi:10.1186/1471-2164-8-223
36. PennarunE, KivisildT, MetspaluE, MetspaluM, ReisbergT, et al. (2012) Divorcing the Late Upper Palaeolithic demographic histories of mtDNA haplogroups M1 and U6 in Africa. BMC Evol Biol 12: 234 doi:10.1186/1471-2148-12-234
37. FernandesV, AlshamaliF, AlvesM, CostaMD, PereiraJB, et al. (2012) The Arabian Cradle: Mitochondrial Relicts of the First Steps along the Southern Route out of Africa. Am J Hum Genet 90: 347–355 doi:10.1016/j.ajhg.2011.12.010
38. ČernýV, MulliganCJ, FernandesV, SilvaNM, AlshamaliF, et al. (2011) Internal Diversification of Mitochondrial Haplogroup R0a Reveals Post-Last Glacial Maximum Demographic Expansions in South Arabia. Mol Biol Evol 28: 71–78 doi:10.1093/molbev/msq178
39. MusilováE, FernandesV, SilvaNM, SoaresP, AlshamaliF, et al. (2011) Population history of the Red Sea—genetic exchanges between the Arabian Peninsula and East Africa signaled in the mitochondrial DNA HV1 haplogroup. Am J Phys Anthropol 145: 592–598 doi:10.1002/ajpa.21522
40. CrucianiF, FrattaRL, SantolamazzaP, SellittoD, PasconeR, et al. (2004) Phylogeographic analysis of haplogroup E3b (E-M215) Y chromosomes reveals multiple migratory events within and out of Africa. Am J Hum Genet 74: 1014–1022 doi:10.1086/386294
41. CrucianiF, FrattaRL, TrombettaB, SantolamazzaP, SellittoD, 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. Mol Biol Evol 24: 1300–1311 doi:10.1093/molbev/msm049
42. ArrediB, PoloniES, ParacchiniS, ZerjalT, FathallahDM, et al. (2004) A predominantly neolithic origin for Y-chromosomal DNA variation in North Africa. Am J Hum Genet 75: 338–345 doi:10.1086/423147
43. HennBM, BotiguéLR, GravelS, WangW, BrisbinA, et al. (2012) Genomic Ancestry of North Africans Supports Back-to-Africa Migrations. PLoS Genet 8: e1002397 doi:10.1371/journal.pgen.1002397
44. SanchezJJ, HallenbergC, BørstingC, HernandezA, MorlingN (2005) High frequencies of Y chromosome lineages characterized by E3b1, DYS19-11, DYS392-12 in Somali males. Eur J Hum Genet 13: 856–866 doi:10.1038/sj.ejhg.5201390
45. MendezFL, KarafetTM, KrahnT, OstrerH, SoodyallH, et al. (2011) Increased Resolution of Y Chromosome Haplogroup T Defines Relationships among Populations of the Near East, Europe, and Africa. Hum Biol 83: 39–53 doi:10.3378/027.083.0103
46. Plaster CA (2011) Variation in Y chromosome, mitochondrial DNA and labels of identity on Ethiopia [Doctoral]. UCL (University College London). Available: http://discovery.ucl.ac.uk/1331901/. Accessed 4 September 2013.
47. LuisJR, RowoldDJ, RegueiroM, CaeiroB, CinnioğluC, et al. (2004) The Levant versus the Horn of Africa: evidence for bidirectional corridors of human migrations. Am J Hum Genet 74: 532–544 doi:10.1086/382286
48. BeharDM, YunusbayevB, MetspaluM, MetspaluE, RossetS, et al. (2010) The genome-wide structure of the Jewish people. Nature 466: 238–242 doi:10.1038/nature09103
49. BrycK, AutonA, NelsonMR, OksenbergJR, HauserSL, et al. (2010) Genome-wide patterns of population structure and admixture in West Africans and African Americans. Proc Natl Acad Sci USA 107: 786–791 doi:10.1073/pnas.0909559107
50. Hunter-ZinckH, MusharoffS, SalitJ, Al-AliKA, ChouchaneL, et al. (2010) Population Genetic Structure of the People of Qatar. Am J Hum Genet 87: 17–25 doi:10.1016/j.ajhg.2010.05.018
51. SchlebuschCM, SkoglundP, SjödinP, GattepailleLM, HernandezD, et al. (2012) Genomic Variation in Seven Khoe-San Groups Reveals Adaptation and Complex African History. Science 338: 374–379 doi:10.1126/science.1227721
52. AltshulerDM, GibbsRA, PeltonenL, AltshulerDM, GibbsRA, et al. (2010) Integrating common and rare genetic variation in diverse human populations. Nature 467: 52–58 doi:10.1038/nature09298
53. LiJ, AbsherD, TangH, SouthwickA, CastoA, et al. (2008) Worldwide Human Relationships Inferred from Genome-Wide Patterns of Variation. Science 319: 1100–1104 doi:10.1126/science.1153717
54. RosenbergNA, MahajanS, RamachandranS, ZhaoC, PritchardJK, et al. (2005) Clines, clusters, and the effect of study design on the inference of human population structure. PLoS Genet 1: e70 doi:10.1371/journal.pgen.0010070
55. PattersonNJ, PriceAL, ReichD (2006) Population structure and eigenanalysis. PLoS Genet 2: e190 doi:10.1371/journal.pgen.0020190
56. MaJ, AmosCI (2012) Principal Components Analysis of Population Admixture. PLoS ONE 7: e40115 doi:10.1371/journal.pone.0040115
57. AlexanderDH, NovembreJ, LangeK (2009) Fast model-based estimation of ancestry in unrelated individuals. Genome Res 19: 1655–1664 doi:10.1101/gr.094052.109
58. AlexanderD, LangeK (2011) Enhancements to the ADMIXTURE Algorithm for Individual Ancestry Estimation. BMC Bioinformatics 12: 246 doi:10.1186/1471-2105-12-246
59. TishkoffSA, ReedFA, FriedlaenderFR, EhretC, RanciaroA, et al. (2009) The Genetic Structure and History of Africans and African Americans. Science 324: 1035–1044 doi:10.1126/science.1172257
60. HaberM, GauguierD, YouhannaS, PattersonN, MoorjaniP, et al. (2013) Genome-Wide Diversity in the Levant Reveals Recent Structuring by Culture. PLoS Genet 9: e1003316 doi:10.1371/journal.pgen.1003316
61. Sánchez-QuintoF, BotiguéLR, CivitS, ArenasC, Ávila-ArcosMC, et al. (2012) North African Populations Carry the Signature of Admixture with Neandertals. PLoS ONE 7: e47765 doi:10.1371/journal.pone.0047765
62. PattersonNJ, MoorjaniP, LuoY, MallickS, RohlandN, et al. (2012) Ancient Admixture in Human History. Genetics 192: 1065–1093 doi:10.1534/genetics.112.145037
63. LohP-R, LipsonM, PattersonN, MoorjaniP, PickrellJK, et al. (2013) Inferring Admixture Histories of Human Populations Using Linkage Disequilibrium. Genetics 193: 1233–1254 doi:10.1534/genetics.112.147330
64. LawsonDJ, HellenthalG, MyersS, FalushD (2012) Inference of Population Structure using Dense Haplotype Data. PLoS Genet 8: e1002453 doi:10.1371/journal.pgen.1002453
65. ExcoffierL, SmousePE, QuattroJM (1992) Analysis of molecular variance inferred from metric distances among DNA haplotypes: application to human mitochondrial DNA restriction data. Genetics 131: 479–491.
66. UrbanekM, GoldmanD, LongJC (1996) The apportionment of dinucleotide repeat diversity in Native Americans and Europeans: a new approach to measuring gene identity reveals asymmetric patterns of divergence. Mol Biol Evol 13: 943–953.
67. LongJC, KittlesRA (2003) Human genetic diversity and the nonexistence of biological races. Hum Biol 75: 449–471 doi:10.3378/027.081.0621
68. Lewis MP, Simons GF, Fennig CD (2013) Ethnologue. Ethnologue: Languages of the World, 17th Edition. Available: http://www.ethnologue.com/. Accessed 17 December 2013.
69. Dimmendaal GJ (2011) Historical Linguistics and the Comparative Study of African Languages. Philadelphia, PA: John Benjamins Publishing.
70. MoorjaniP, PattersonN, HirschhornJN, KeinanA, HaoL, et al. (2011) The History of African Gene Flow into Southern Europeans, Levantines, and Jews. PLoS Genet 7: e1001373 doi:10.1371/journal.pgen.1001373
71. PetersenDC, LibigerO, TindallEA, HardieR-A, HannickLI, et al. (2013) Complex Patterns of Genomic Admixture within Southern Africa. PLoS Genet 9: e1003309 doi:10.1371/journal.pgen.1003309
72. VerduP, RosenbergNA (2011) A General Mechanistic Model for Admixture Histories of Hybrid Populations. Genetics 189: 1413–1426 doi:10.1534/genetics.111.132787
73. HolsingerKE, WeirBS (2009) Genetics in geographically structured populations: defining, estimating and interpreting FST. Nat Rev Genet 10: 639–650 doi:10.1038/nrg2611
74. GutenkunstRN, HernandezRD, WilliamsonSH, BustamanteCD (2009) Inferring the Joint Demographic History of Multiple Populations from Multidimensional SNP Frequency Data. PLoS Genet 5: e1000695 doi:10.1371/journal.pgen.1000695
75. TishkoffSA, ReedFA, RanciaroA, VoightBF, BabbittCC, et al. (2007) Convergent adaptation of human lactase persistence in Africa and Europe. Nat Genet 39: 31–40 doi:10.1038/ng1946
76. EnattahNS, SahiT, SavilahtiE, TerwilligerJD, PeltonenL, et al. (2002) Identification of a variant associated with adult-type hypolactasia. Nat Genet 30: 233–237 doi:10.1038/ng826
77. SwallowDM (2003) Genetics of Lactase Persistence and Lactose Intolerance. Ann Rev Genet 37: 197–219 doi:10.1146/annurev.genet.37.110801.143820
78. ImtiazF, SavilahtiE, SarnestoA, TrabzuniD, Al-KahtaniK, et al. (2007) The T/G 13915 variant upstream of the lactase gene (LCT) is the founder allele of lactase persistence in an urban Saudi population. J Med Genet 44: e89 doi:10.1136/jmg.2007.051631
79. EnattahNS, JensenT, NielsenM, LewinskiR, KuokkanenM, et al. (2008) Independent Introduction of Two Lactase-Persistence Alleles into Human Populations Reflects Different History of Adaptation to Milk Culture. Am J Hum Genet 82: 57–72 doi:10.1016/j.ajhg.2007.09.012
80. EhretC (1979) On the Antiquity of Agriculture in Ethiopia. J Afr Hist 20: 161–177 doi:10.1017/S002185370001700X
81. Lovejoy PE (2000) Transformations in Slavery: A History of Slavery in Africa. 2nd ed. Cambridge: Cambridge University Press.
82. Hoelzmann P, Gasse F, Dupont LM, Salzmann U, Staubwasser M, et al.. (2004) Palaeoenvironmental changes in the arid and sub arid belt (Sahara-Sahel-Arabian Peninsula) from 150 kyr to present. In: Battarbee RW, Gasse F, Stickley CE, editors. Past Climate Variability through Europe and Africa. Developments in Paleoenvironmental Research. Springer Netherlands. pp. 219–256.
83. HetheringtonR, WiebeE, WeaverAJ, CartoSL, EbyM, et al. (2008) Climate, African and Beringian subaerial continental shelves, and migration of early peoples. Quatern Int 183: 83–101 doi:10.1016/j.quaint.2007.06.033
84. WilliamsMAJ (2009) Late Pleistocene and Holocene environments in the Nile basin. Global Planet Change 69: 1–15 doi:10.1016/j.gloplacha.2009.07.005
85. Parker AG (2010) Pleistocene Climate Change in Arabia: Developing a Framework for Hominin Dispersal over the Last 350 ka. In: Petraglia MD, Rose JI, editors. The Evolution of Human Populations in Arabia. New York: Springer. pp. 39–49.
86. WilliamsMAJ, WilliamsFM, DullerGAT, MunroRN, El TomOAM, et al. (2010) Late Quaternary floods and droughts in the Nile valley, Sudan: new evidence from optically stimulated luminescence and AMS radiocarbon dating. Quatern Sci Rev 29: 1116–1137 doi:10.1016/j.quascirev.2010.02.018
87. BrandtSA (1986) The Upper Pleistocene and early Holocene prehistory of the Horn of Africa. Afr Archaeol Rev 4: 41–82 doi:10.1007/BF01117035
88. Brandt SA (1997) Horn of Africa: History of Archaeology. In: Vogel JO, editor. Encyclopedia of Precolonial Africa. Walnut Creek, CA: AltaMira Press. pp. 69–75.
89. BrandtSA (1988) Early Holocene Mortuary Practices and Hunter-Gatherer Adaptations in Southern Somalia. World Archaeol 20: 40–56 doi:10.2307/124524
90. MayerDEB-Y, BeyinA (2009) Late Stone Age Shell Middens on the Red Sea Coast of Eritrea. J Island Coastal Archaeol 4: 108–124 doi:10.1080/15564890802662171
91. BeyinA (2010) Use-wear analysis of obsidian artifacts from Later Stone Age shell midden sites on the Red Sea Coast of Eritrea, with experimental results. J Archaeol Sci 37: 1543–1556 doi:10.1016/j.jas.2010.01.015
92. Beyin (2011) Early to Middle Holocene human adaptations on the Buri Peninsula and Gulf of Zula, coastal lowlands of Eritrea. Azania 46: 123–140 doi:10.1080/0067270X.2011.580139
93. BraceCL, SeguchiN, QuintynCB, FoxSC, NelsonAR, et al. (2006) The questionable contribution of the Neolithic and the Bronze Age to European craniofacial form. Proc Natl Acad Sci USA 103: 242–247 doi:10.1073/pnas.0509801102
94. IrishJD, KonigsbergL (2007) The Ancient Inhabitants of Jebel Moya Redux: Measures of Population Affinity Based on Dental Morphology. Int J Osteoarchaeol 17: 138–156 doi:10.1002/oa.868
95. RicautFX, WaelkensM (2008) Cranial Discrete Traits in a Byzantine Population and Eastern Mediterranean Population Movements. Hum Biol 80: 535–564 doi:10.3378/1534-6617-80.5.535
96. Sellers TA (2008) The Influence of Subsistence Shift on Dental Reductions: A Comparison of Prehistoric and Modern Nubian and Somalian Dental Samples [M.A. Thesis]. Cincinatti, OH: University of Cincinnati.
97. FerembachD (1985) On the origin of the iberomaurusians (Upper palaeolithic: North Africa). A new hypothesis. J Hum Evol 14: 393–397 doi:10.1016/S0047-2484(85)80047-6
98. IrishJD (2000) The Iberomaurusian enigma: North African progenitor or dead end? J Hum Evol 39: 393–410 doi:10.1006/jhev.2000.0430
99. RandoJC, PintoF, GonzálezAM, HernándezM, LarrugaJM, et al. (1998) Mitochondrial DNA analysis of northwest African populations reveals genetic exchanges with European, near-eastern, and sub-Saharan populations. Ann Hum Genet 62: 531–550 doi:10.1046/j.1469-1809.1998.6260531.x
100. Maca-MeyerN, GonzálezAM, LarrugaJM, FloresC, CabreraVM (2001) Major genomic mitochondrial lineages delineate early human expansions. BMC Genet 2: 13 doi:10.1186/1471-2156-2-13
101. Maca-MeyerN, GonzálezAM, PestanoJ, FloresC, LarrugaJM, et al. (2003) Mitochondrial DNA transit between West Asia and North Africa inferred from U6 phylogeography. BMC Genet 4: 15 doi:10.1186/1471-2156-4-15
102. PlazaS, CalafellF, HelalA, BouzernaN, LefrancG, et al. (2003) Joining the Pillars of Hercules: mtDNA Sequences Show Multidirectional Gene Flow in the Western Mediterranean. Ann Hum Genet 67: 312–328 doi:10.1046/j.1469-1809.2003.00039.x
103. CherniL, FernandesV, PereiraJB, CostaMD, GoiosA, et al. (2009) Post-last glacial maximum expansion from Iberia to North Africa revealed by fine characterization of mtDNA H haplogroup in Tunisia. Am J Phys Anthropol 139: 253–260 doi:10.1002/ajpa.20979
104. EnnafaaH, CabreraVM, Abu-AmeroKK, GonzálezAM, AmorMB, et al. (2009) Mitochondrial DNA haplogroup H structure in North Africa. BMC Genet 10: 8 doi:10.1186/1471-2156-10-8
105. Fadhlaoui-ZidK, Rodríguez-BotiguéL, NaouiN, Benammar-ElgaaiedA, CalafellF, et al. (2011) Mitochondrial DNA structure in North Africa reveals a genetic discontinuity in the Nile Valley. Am J Phys Anthropol 145: 107–117 doi:10.1002/ajpa.21472
106. PritchardJK, StephensM, DonnellyP (2000) Inference of population structure using multilocus genotype data. Genetics 155: 945–959.
107. Greenberg JH (1971) African languages. In: Dil A, editor. Language, Culture, and Communication. Stanford: Stanford University Press. pp. 126–136.
108. EhretC, KeitaSOY, NewmanP (2004) The Origins of Afroasiatic. Science 306: 1680–1680 doi:10.1126/science.306.5702.1680c
109. Militarev A (2000) Towards the chronology of Afrasian (Afroasiatic) and its daughter families. In: Renfrew C, McMahon A, Trask L, editors. Time Depth in Historical Linguistics. Cambridge: The McDonald Institute for Archaeological Research, Vol. 1. pp. 267–307.
110. Militarev A (2002) The Prehistory of a Dispersal: the Proto-Afrasian (Afroasiatic) Farming Lexicon. In: Bellwood P, Renfrew C, editors. Examining the Farming/Language Dispersal Hypothesis. Cambridge: The McDonald Institute for Archaeological Research. pp. 135–150.
111. McCallDF (1998) The Afroasiatic Language Phylum: African in Origin, or Asian? Curr Anthropol 39: 139–144 doi:10.1086/204702
112. HodgsonJA, MulliganCJ, Al-MeeriA, RaaumRL (2014) Data from: Early Back-to-African Migration into the Horn of Africa. Dryad Digital Repository doi:10.5061/dryad.d9s74
113. PurcellS, NealeB, Todd-BrownK, ThomasL, FerreiraMAR, et al. (2007) PLINK: A Tool Set for Whole-Genome Association and Population-Based Linkage Analyses. Am J Hum Genet 81: 559–575 doi:10.1086/519795
114. RosenbergNA (2006) Standardized Subsets of the HGDP-CEPH Human Genome Diversity Cell Line Panel, Accounting for Atypical and Duplicated Samples and Pairs of Close Relatives. Ann Hum Genet 70: 841–847 doi:10.1111/j.1469-1809.2006.00285.x
115. PembertonTJ, WangC, LiJZ, RosenbergNA (2010) Inference of Unexpected Genetic Relatedness among Individuals in HapMap Phase III. Am J Hum Genet 87: 457–464 doi:10.1016/j.ajhg.2010.08.014
116. ManichaikulA, MychaleckyjJC, RichSS, DalyK, SaleM, et al. (2010) Robust relationship inference in genome-wide association studies. Bioinformatics 26: 2867–2873 doi:10.1093/bioinformatics/btq559
117. MatiseTC, ChenF, ChenW, VegaFMDL, HansenM, et al. (2007) A second-generation combined linkage–physical map of the human genome. Genome Res 17: 1783–1786 doi:10.1101/gr.7156307
118. François O (n.d.) How to display admixture coefficients (Q matrix) spatially? Available: http://membres-timc.imag.fr/Olivier.Francois/admix_display.html. Accessed 29 January 2013.
119. Venables WN, Ripley BD (2002) Modern applied statistics with S. Fourth Edition. New York: Springer.
120. Becker RA, Wilks AR, Brownrigg R, Minka TP (2012) maps: Draw Geographical Maps. Available: http://CRAN.R-project.org/package=maps.
121. Lewin-Koh NJ, Bivand R, Pebesma EJ, Archer E, Baddeley A, et al.. (2012) maptools: Tools for reading and handling spatial objects. Available: http://CRAN.R-project.org/package=maptools.
122. R Development Core Team (2013) R: A Language and Environment for Statistical Computing. Vienna, Austria. Available: http://www.R-project.org.
123. DelaneauO, ZaguryJ-F, MarchiniJ (2013) Improved whole-chromosome phasing for disease and population genetic studies. Nat Meth 10: 5–6 doi:10.1038/nmeth.2307
124. DrayS, DufourA-B (2007) The ade4 Package: Implementing the Duality Diagram for Ecologists. J Stat Softw 22: 1–20.
125. TremblayM, VézinaH (2000) New Estimates of Intergenerational Time Intervals for the Calculation of Age and Origins of Mutations. Am J Hum Genet 66: 651–658 doi:10.1086/302770
126. HelgasonA, HrafnkelssonB, GulcherJR, WardR, StefánssonK (2003) A Populationwide Coalescent Analysis of Icelandic Matrilineal and Patrilineal Genealogies: Evidence for a Faster Evolutionary Rate of mtDNA Lineages than Y Chromosomes. Am J Hum Genet 72: 1370–1388 doi:10.1086/375453
127. MatsumuraS, ForsterP (2008) Generation time and effective population size in Polar Eskimos. Proc Roy Soc B Biol Sci 275: 1501–1508 doi:10.1098/rspb.2007.1724
128. PriceAL, TandonA, PattersonN, BarnesKC, RafaelsN, et al. (2009) Sensitive Detection of Chromosomal Segments of Distinct Ancestry in Admixed Populations. PLoS Genet 5: e1000519 doi:10.1371/journal.pgen.1000519
129. ScheetP, StephensM (2006) A Fast and Flexible Statistical Model for Large-Scale Population Genotype Data: Applications to Inferring Missing Genotypes and Haplotypic Phase. Am J Hum Genet 78: 629–644 doi:10.1086/502802
Štítky
Genetika Reprodukčná medicínaČlánok vyšiel v časopise
PLOS Genetics
2014 Číslo 6
- Je „freeze-all“ pro všechny? Odborníci na fertilitu diskutovali na virtuálním summitu
- Gynekologové a odborníci na reprodukční medicínu se sejdou na prvním virtuálním summitu
Najčítanejšie v tomto čísle
- Early Back-to-Africa Migration into the Horn of Africa
- PINK1-Mediated Phosphorylation of Parkin Boosts Parkin Activity in
- OsHUS1 Facilitates Accurate Meiotic Recombination in Rice
- An Operon of Three Transcriptional Regulators Controls Horizontal Gene Transfer of the Integrative and Conjugative Element ICE in B13