Книга: Кто мы и как сюда попали. Древняя ДНК и новая наука о человеческом прошлом
Назад: Пояснения к иллюстрациям
Дальше: Примечания

Примечания

Введение

1 L. L. Cavalli-Sforza, P. Menozzi, A. Piazza. The History and Geography of Human Genes (Princeton, NJ: Princeton University Press, 1994).

2 L. L. Cavalli-Sforza, F. Cavalli-Sforza. The Great Human Diasporas: The History of Diversity and Evolution (Reading, MA: Addison-Wesley, 1995).

3 N. A. Rosenberg et al. “Genetic Structure of Human Populations”, Science 298 (2002): 2381–2385.

4 P. Menozzi, A. Piazza, L. L. Cavalli-Sforza. “Synthetic Maps of Human Gene Frequencies in Europeans”, Science 201 (1978): 786–792; L. L. Cavalli-Sforza, P. Menozzi, A. Piazza. “Demic Expansions and Human Evolution”, Science 259 (1993): 639–646.

5 A. J. Ammerman, L. L. Cavalli-Sforza. The Neolithic Transition and the Genetics of Populations in Europe (Princeton, NJ: Princeton University Press, 1984).

6 J. Novembre, M. Stephens. “Interpreting Principal Component Analyses of Spatial Population Genetic Variation”, Nature Genetics 40 (2008): 646–649.

7 O. François et al. “Principal Component Analysis Under Population Genetic Models of Range Expansion and Admixture”, Molecular Biology and Evolution 27 (2010): 1257–68.

8 A. Keller et al. “New Insights into the Tyrolean Iceman’s Origin and Phenotype as Inferred by Whole-Genome Sequencing”, Nature Communications 3 (2012): 698; P. Skoglund et al. “Ori gins and Genetic Legacy of Neolithic Farmers and Hunter-Gatherers in Europe”, Science 336 (2012): 466–469; I. Lazaridis et al. “Ancient Human Genomes Suggest Three Ancestral Populations for Present-Day Europeans”, Nature 513 (2014): 409–13.

9 J. K. Pickrell, D. Reich. “Toward a New History and Geography of Human Genes Informed by Ancient DNA”, Trends in Genetics 30 (2014): 377–89.

10 R. E. Green et al. “A Draft Sequence of the Neandertal Genome”, Science 328 (2010): 710–722.

11 D. Reich et al. “Genetic History of an Archaic Hominin Group from Denisova Cave in Siberia”, Nature 468 (2010): 1053–1060.

12 M. Rasmussen et al. “Ancient Human Genome Sequence of an Extinct Palaeo-Eskimo”, Nature 463 (2010): 757–762.

13 W. Haak et al. “Massive Migration from the Steppe Was a Source for Indo-European Languages in Europe”, Nature 522 (2015): 207–211.

14 M. E. Allentoft et al. “Population Genomics of Bronze Age Eurasia”, Nature 522 (2015): 167–172.

15 I. Mathieson et al. “Genome-Wide Patterns of Selection in 230 Ancient Eurasians”, Nature 528 (2015): 499–503.

16 Q. Fu et al. “DNA Analysis of an Early Modern Human from Tianyuan Cave, China”, Proceedings of the National Academy of Sciences of the U.S.A. 110 (2013): 2223–2227.

17 H. Shang et al. “An Early Modern Human from Tianyuan Cave, Zhoukoudian, China”, Proceedings of the National Academy of Sciences of the U.S.A. 104 (2007): 6573–6578.

18 Haak et al. “Massive Migration”.

19 I. Lazaridis et al. “Genomic Insights into the Origin of Farming in the Ancient Near East”, Nature 536 (2016): 419–424.

20 P. Skoglund et al. “Genomic Insights into the Peopling of the Southwest Pacific”, Nature 538 (2016): 510–513.

21 Lazaridis et al. “Ancient Human Genomes”.

22 Pickrell, Reich. “Toward a New History”.



Глава 1. Как геном объясняет, кто мы такие

1 J. D. Watson and F. H. Crick. “Molecular Structure of Nucleic Acids; a Structure for Deoxyribose Nucleic Acid”, Nature 171 (1953): 737–738.

2 R. L. Cann, M. Stoneking, A. C. Wilson. “Mitochondrial DNA and Human Evolution”, Nature 325 (1987): 31–36.

3 Cann et al. “Mitochondrial DNA and Human Evolution”.

4 Q. Fu et al. “A Revised Timescale for Human Evolution Based on Ancient Mitochondrial Genomes”, Current Biology 23 (2013): 553–559.

5 D. E. Lieberman, B. M. McBratney, G. Krovitz. “The Evolution and Development of Cranial Form in Homo sapiens”, Proceedings of the National Academy of Sciences of the U.S.A. 99 (2002):1134– 39; Richter et al. “The Age of the Hominin Fossils from Jebel Irhoud, Morocco, and the Origins of the Middle Stone Age”, Nature 546 (2017): 293–296.

6 H. S. Groucutt et al. “Rethinking the Dispersal of Homo sapiens Out of Africa”, Evolutionary Anthropology 24 (2015): 149–164.

7 C.-J. Kind et al. “The Smile of the Lion Man: Recent Excavations in Stadel Cave (Baden-Württemberg, South-Western Germany) and the Restoration of the Famous Upper Palaeolithic Figurine”, Quartär 61 (2014): 129–145.

8 T. Higham et al. “The Timing and Spatiotemporal Patterning of Neanderthal Disappearance”, Nature 512 (2014): 306–309.

9 Richard G. Klein, Blake Edgar. The Dawn of Human Culture (New York: Wiley, 2002).

10 J. Doebley. “Mapping the Genes That Made Maize”, Trends in Genetics 8 (1992): 302–307.

11 S. McBrearty, A. S. Brooks. “The Revolution That Wasn’t: A New Interpretation of the Origin of Modern Human Behavior”, Journal of Human Evolution 39 (2000): 453–563.

12 C. S. L. Lai et al. “A Forkhead-Domain Gene Is Mutated in a Severe Speech and Language Disorder”, Nature 413 (2001): 519–523.

13 W. Enard et al. “Molecular Evolution of FOXP2, a Gene Involved in Speech and Language”, Nature 418 (2002): 869–872.

14 W. Enard et al. “A Humanized Version of FOXP2 Affects Cortico-Basal Ganglia Circuits in Mice”, Cell 137 (2009): 961–971.

15 J. Krause et al. “The Derived FOXP2 Variant of Modern Humans Was Shared with Neandertals”, Current Biology 17 (2007): 1908–1912.

16 T. Maricic et al. “A Recent Evolutionary Change Affects a Regulatory Element in the Human FOXP2 Gene”, Molecular Biology and Evolution 30 (2013): 844–852.

17 S. Pääbo. “The Human Condition – a Molecular Approach”, Cell 157 (2014): 216–226.

18 R. E. Green et al. “A Draft Sequence of the Neandertal Genome”, Science 328 (2010): 710–722; K. Prüfer et al. “The Complete Genome Sequence of a Neanderthal from the Altai Mountains”, Nature (2013): doi: 10.1038/nature 1288.

19 R. Lewin. “The Unmasking of Mitochondrial Eve”, Science 238 (1987): 24–26.

20 A. Kong et al. “A High-Resolution Recombination Map of the Human Genome”, Nature Genetics 31 (2002): 241–247.

21 “Descent of Elizabeth II from William I”, Familypedia, http:// familypedia wikia.com/wiki/Descent_of_Elizabeth_II_from_ William_I#Shorter_line_of_descent.

22 S. Mallick et al. “The Simons Genome Diversity Project: 300 Genomes from 142 Diverse Populations”, Nature 538 (2016): 201–206.

23 Green et al. “Draft Sequence”.

24 H. Li, R. Durbin. “Inference of Human Population History from Individual Whole-Genome Sequences”, Nature 475 (2011): 493–496.

25 Там же.

26 S. Schiffels, R. Durbin. “Inferring Human Population Size and Separation History from Multiple Genome Sequences”, Nature Genetics 46 (2014): 919–925.

27 Mallick et al. “Simons Genome Diversity Project”.

28 I. Gronau et al. “Bayesian Inference of Ancient Human Demography from Individual Genome Sequences”, Nature Genetics 43 (2011): 1031–1034.

29 Mallick et al. “Simons Genome Diversity Project”.

30 P. C. Sabeti et al. “Detecting Recent Positive Selection in the Human Genome from Haplotype Structure”, Nature 419 (2002): 832–837; B. F. Voight, S. Kudaravalli, X. Wen, J. K. Pritchard. “A Map of Recent Positive Selection in the Human Genome”, PLoS Biology 4 (2006): e72.

31 K. M. Teshima, G. Coop, M. Przeworski. “How Reliable Are Empirical Genomic Scans for Selective Sweeps?”, Genome Research 16 (2006): 702–712.

32 R. D. Hernandez et al. “Classic Selective Sweeps Were Rare in Recent Human Evolution”, Science 331 (2011): 920–924.

33 S. A. Tishkoff et al. “Convergent Adaptation of Human Lactase Persistence in Africa and Europe”, Nature Genetics 38 (2006): 31–40.

34 M. C. Turchin et al. “Evidence of Widespread Selection on Standing Variation in Europe at Height-Associated SNPs”, Nature Genetics 44 (2012): 1015–1019.

35 I. Mathieson et al. “Genome-Wide Patterns of Selection in 230 Ancient Eurasians”, Nature 528 (2015): 499–503.

36 Y. Field et al. “Detection of Human Adaptation During the Past 2000 Years”, Science 354 (2016): 760–764.

37 D. Welter et al. “The NHGRI GWAS Catalog, a Curated Resource of SNP-Trait Associations”, Nucleic Acids Research 42 (2014): D1001–1006.

38 D. B. Goldstein. “Common Genetic Variation and Human Traits”, New England Journal of Medicine 360 (2009): 1696–1698.

39 A. Okbay et al. “Genome-Wide Association Study Identifies 74 Loci Associated with Educational Attainment”, Nature 533 (2016): 539–542; M. T. Lo et al. “Genome-Wide Analyses for Personality Traits Identify Six Genomic Loci and Show Correlations with Psychiatric Disorders”, Nature Genetics 49 (2017): 152–156; G. Davies et al. “Genome-Wide Association Study of Cognitive Functions and Educational Attainment in UK Biobank (N=112 151)”, Molecular Psychiatry 21 (2016): 758–767.



Глава 2. Встреча с неандертальцами

1 Charles Darwin. The Descent of Man, and Selection in Relation to Sex (London: John Murray, 1871).

2 Erik Trinkaus. The Shanidar Neanderthals (New York: Academic Press, 1983).

3 D. Radovcˇ i, A. O. Sršen, J. Radovcˇ ic´, D. W. Frayer. “Evidence for Neandertal Jewelry: Modified White-Tailed Eagle Claws at Krapina”, PLoS One 10 (2015): e0119802.

4 J. Jaubert et al. “Early Neanderthal Constructions Deep in Bruniquel Cave in Southwestern France”, Nature 534 (2016): 111–114.

5 W. L. Straus, A. J. E. Cave. “Pathology and the Posture of Neanderthal Man”, Quarterly Review of Biology 32 (1957): 348–363.

6 William Golding. The Inheritors (London: Faber and Faber, 1955).

7 Jean M. Auel. The Clan of the Cave Bear (New York: Crown, 1980).

8 T. Higham et al. “The Timing and Spatiotemporal Patterning of Neanderthal Disappearance”, Nature 512 (2014): 306–309.

9 T. Higham et al. “Chronology of the Grotte du Renne (France) and Implications for the Context of Ornaments and Human Remains Within the Châtelperronian”, Proceedings of the National Academy of Sciences of the U.S.A. 107 (2010): 20234–20239; O. Bar-Yosef, J.-G. Bordes. “Who Were the Makers of the Châtelperronian Culture?”, Journal of Human Evolution 59 (2010): 586–593.

10 R. Grün et al. “U-series and ESR Analyses of Bones and Teeth Relating to the Human Burials from Skhul”, Journal of Human Evolution 49 (2005): 316–334.

11 H. Valladas et al. “Thermo-Luminescence Dates for the Neanderthal Burial Site at Kebara in Israel”, Nature 330 (1987): 159–160.

12 E. Trinkaus et al. “An Early Modern Human from the Pe¸stera cu Oase, Romania”, Proceedings of the National Academy of Sciences of the U.S.A. 100 (2003): 11231–11236.

13 M. Krings et al. “Neandertal DNA Sequences and the Origin of Modern Humans”, Cell 90 (1997): 19–30.

14 C. Posth et al. “Deeply Divergent Archaic Mitochondrial Genome Provides Lower Time Boundary for African Gene Flow into Neanderthals”, Nature Communications 8 (2017): 16046.

15 Krings et al. “Neandertal DNA Sequences”.

16 M. Currat, L. Excoffier. “Modern Humans Did Not Admix with Neanderthals During Their Range Expansion into Europe”, PLoS Biology 2 (2004): e421; D. Serre et al. “No Evidence of Neandertal mtDNA Contribution to Early Modern Humans”, PLoS Biology 2 (2004): e57; M. Nordborg.“On the Probability of Neanderthal Ancestry”, American Journal of Human Genetics 63 (1998): 1237–1240.

17 R. E. Green et al. “Analysis of One Million Base Pairs of Neanderthal DNA”, Nature 444 (2006): 330–336.

18 J. D. Wall and S. K. Kim. “Inconsistencies in Neanderthal Genomic DNA Sequences”, PLoS Genetics 3 (2007): 1862–1866.

19 Krings et al. “Neandertal DNA Sequences”.

20 S. Sankararaman et al. “The Date of Interbreeding Between Neandertals and Modern Humans”, PLoS Genetics 8 (2012): e1002947.

21 P. Moorjani et al. “A Genetic Method for Dating Ancient Genomes Provides a Direct Estimate of Human Generation Interval in the Last 45,000 Years”, Proceedings of the National Academy of Sciences of the U.S.A. 113 (2016): 5652–5657.

22 G. Coop. “Thoughts On: The Date of Interbreeding Between Neandertals and Modern Humans”, Haldane’s Sieve, September 18, 2012, https://haldanessieve.org/2012/09/18/thoughts-on-neandertal-article/

23 K. Prüfer et al. “The Complete Genome Sequence of a Neanderthal from the Altai Mountains”, Nature (2013): doi: 10.1038/nature 12886.

24 Там же.

25 Там же; M. Meyer et al. “A High-Coverage Genome Sequence from an Archaic Denisovan Individual”, Science 338 (2012): 222–226; J. D. Wall et al. “Higher Levels of Neanderthal Ancestry in East Asians Than in Europeans”, Genetics 194 (2013): 199–209.

26 Q. Fu et al. “The Genetic History of Ice Age Europe”, Nature 534 (2016): 200–205.

27 I. Lazaridis et al. “Genomic Insights into the Origin of Farming in the Ancient Near East”, Nature 536 (2016): 419–424.

28 Trinkaus et al. “An Early Modern Human”.

29 “An Early Modern Human from Romania with a Recent Neanderthal Ancestor”, Nature 524 (2015): 216–219.

30 N. Teyssandier, F. Bon, and J.-G. Bordes. “Within Projectile Range: Some Thoughts on the Appearance of the Aurignacian in Europe”, Journal of Anthropological Research 66 (2010): 209–229; P. Mellars. “Archeology and the Dispersal of Modern Humans in Europe: Deconstructing the ‘Aurignacian’ ”, Evolutionary Anthropology 15 (2006): 167–182.

31 M. Currat, L. Excoffier. “Strong Reproductive Isolation Between Humans and Neanderthals Inferred from Observed Patterns of Introgression”, Proceedings of the National Academy of Sciences of the U.S.A. 108 (2011): 15129–15134.

32 S. Sankararaman et al. “The Genomic Landscape of Neanderthal Ancestry in Present-Day Humans”, Nature 507 (2014): 354–357; B. Vernot, J. M. Akey. “Resurrecting Surviving Neandertal Lineages from Modern Human Genomes”, Science 343 (2014): 1017–1021.

33 N. Patterson et al. “Genetic Evidence for Complex Speciation of Humans and Chimpanzees”, Nature 441 (2006): 1103–1108.

34 Там же; R. Burgess, Z. Yang. “Estimation of Hominoid Ancestral Population Sizes Under Bayesian Coalescent Models Incorporating Mutation Rate Variation and Sequencing Errors”, Molecular Biology and Evolution 25 (2008): 1975–1994.

35 J. A. Coyne and H. A. Orr. “Two Rules of Speciation”, in Speciation and Its Consequences, ed. Daniel Otte and John A. Endler (Sunderland, MA: Sinauer Associates, 1989), 180–207.

36 P. K. Tucker et al. “Abrupt Cline for Sex-Chromosomes in a Hybrid Zone Between Two Species of Mice”, Evolution 46 (1992): 1146–1163.

37 H. Li, R. Durbin. “Inference of Human Population History from Individual Whole-Genome Sequences”, Nature 475 (2011): 493–496.

38 T. Mailund et al. “A New Isolation with Migration Model Along Complete Genomes Infers Very Different Divergence Processes Among Closely Related Great Ape Species”, PLoS Genetics 8 (2012): e1003125.

39 J. Y. Dutheil et al. “Strong Selective Sweeps on the X Chromosome in the Human-Chimpanzee Ancestor Explain Its Low Divergence”, PLoS Genetics 11 (2015): e1005451.

40 Sankararaman et al. “Genomic Landscape”; B. Jégou et al. “Meiotic Genes Are Enriched in Regions of Reduced Archaic Ancestry”, Molecular Biology and Evolution 34 (2017): 1974–1980.

41 Q. Fu et al. “Ice Age Europe”.

42 I. Juric, S. Aeschbacher, G. Coop. “The Strength of Selection Against Neanderthal Introgression”, PLoS Genetics 12 (2016): e1006340; K. Harris, R. Nielsen. “The Genetic Cost of Neanderthal Introgression”, Genetics 203 (2016): 881–891.

43 G. Bhatia et al. “Genome-Wide Scan of 29, 141 African Americans Finds No Evidence of Directional Selection Since Admixture”, American Journal of Human Genetics 95 (2014): 437–444.

44 Johann G. Fichte. Grundlage der gesamten Wissenschaftslehre (Jena, Germany: Gabler, 1794).



Глава 3. Древняя ДНК открывает шлюзы

1 J. Krause et al. “Neanderthals in Central Asia and Siberia”, Nature 449 (2007): 902–904.

2 J. Krause et al. “The Complete Mitochondrial DNA Genome of an Unknown Hominin from Southern Siberia”, Nature 464 (2010): 894–897.

3 C. Posth et al. “Deeply Divergent Archaic Mitochondrial Genome Provides Lower Time Boundary for African Gene Flow into Neanderthals”, Nature Communications 8 (2017): 16046.

4 Krause et al. “Unknown Hominin”.

5 D. Reich et al. “Genetic History of an Archaic Hominin Group from Denisova Cave in Siberia”, Nature 468 (2010): 1053–1060.

6 K. Prüfer et al. “The Complete Genome Sequence of a Neanderthal from the Altai Mountains”, Nature (2013): doi: 10.1038/nature 12886.

7 Jerry A. Coyne, H. Allen Orr. Speciation (Sunderland, MA: Sinauer Associates, 2004).

8 S. Sankararaman, S. Mallick, N. Patterson, D. Reich. “The Combined Landscape of Denisovan and Neanderthal Ancestry in Present-Day Humans”, Current Biology 26 (2016): 1241–1247.

9 P. Moorjani et al. “A Genetic Method for Dating Ancient Genomes Provides a Direct Estimate of Human Generation Interval in the Last 45,000 Years”, Proceedings of the National Academy of Sciences of the U.S.A. 113 (2016): 5652–5657.

10 Sankararaman et al. “Combined Landscape”.

11 D. Reich et al. “Denisova Admixture and the First Modern Human Dispersals into Southeast Asia and Oceania”, American Journal of Human Genetics 89 (2011): 516–528.

12 Q. Fu et al. “DNA Analysis of an Early Modern Human from Tianyuan Cave, China”, Proceedings of the National Academy of Sciences of the U.S.A. 110 (2013): 2223–2227; M. Yang et al. “40,000-Year-Old Individual from Asia Provides Insight into Early Population Structure in Eurasia”, Current Biology 27 (2017): 3202–3208.

13 Prüfer et al. “Complete Genome”.

14 C. B. Stringer and I. Barnes. “Deciphering the Denisovans”, Proceedings of the National Academy of Sciences of the U.S.A. 112 (2015): 15542–15543.

15 G. A. Wagner et al. “Radiometric Dating of the Type-Site for Homo Heidelber-gensis at Mauer, Germany”, Proceedings of the National Academy of Sciences of the U.S.A. 107 (2010): 19726–19730.

16 C. Stringer. “The Status of Homo heidelbergensis (Schoetensack 1908)”, Evolutionary Anthropology 21 (2012): 101–107.

17 A. Brumm et al. “Age and Context of the Oldest Known Hominin Fossils from Flores”, Nature 534 (2016): 249–253.

18 Reich et al. “Denisova Admixture”.

19 Prüfer et al. “Complete Genome”.

20 Там же; Sankararaman et al. “Combined Landscape”.

21 E. Huerta-Sánchez et al. “Altitude Adaptation in Tibetans Caused by Introgression of Denisovan-like DNA”, Nature 512 (2014): 194–197.

22 F. H. Chen et al. “Agriculture Facilitated Permanent Human Occupation of the Tibetan Plateau After 3600 B.P.”, Science 347 (2015): 248–250.

23 S. Sankararaman et al. “The Genomic Landscape of Neanderthal Ancestry in PresentDay Humans”, Nature 507 (2014): 354–357; B. Vernot and J. M. Akey. “Resurrecting Surviving Neandertal Lineages from Modern Human Genomes”, Science 343 (2014): 1017–1021.

24 Prüfer et al. “Complete Genome”.

25 G. P. Rightmire. “Homo erectus and Middle Pleistocene Hominins: Brain Size, Skull Form, and Species Recognition”, Journal of Human Evolution 65 (2013): 223–252.

26 M. Martinón-Torres et al. “Dental Evidence on the Hominin Dispersals During the Pleistocene”, Proceedings of the National Academy of Sciences of the U.S.A. 104 (2007): 13279–13282; M. Martinón-Torres, R. Dennell, J. M. B. de Castro. “The Denisova Hominin Need Not Be an Out of Africa Story”, Journal of Human Evolution 60 (2011): 251–255; J. M. B. de Castro, M. Martinón-Torres. “A New Model for the Evolution of the Human Pleistocene Populations of Europe”, Quaternary International 295 (2013): 102–112.

27 De Castro, Martinón-Torres. “A New Model”.

28 J. L. Arsuaga et al. “Neandertal Roots: Cranial and Chronological Evidence from Sima de los Huesos”, Science 344 (2014): 1358–1363; M. Meyer et al. “A Mitochondrial Genome Sequence of a Hominin from Sima de los Huesos”, Nature 505 (2014): 403–406.

29 M. Meyer et al. “Nuclear DNA Sequences from the Middle Pleistocene Sima de los Huesos Hominins”, Nature 531 (2016): 504–507.

30 Meyer et al. “A Mitochondrial Genome”; Meyer et al. “Nuclear DNA Sequences”.

31 Krause et al. “Unknown Hominin”; Reich et al. “Genetic History”.

32 Posth et al. “Deeply Divergent Archaic”.

33 Там же.

34 Prüfer et al. “Complete Genome”.

35 S. McBrearty, A. S. Brooks. “The Revolution That Wasn’t: A New Interpretation of the Origin of Modern Human Behavior”, Journal of Human Evolution 39 (2000): 453–563.

36 M. Kuhlwilm et al. “Ancient Gene Flow from Early Modern Humans into Eastern Neanderthals”, Nature 530 (2016): 429–433.



Глава 4. Призраки человечества

1 Charles R. Darwin. On the Origin of Species by Means of Natural Selection, or the Preservation of Favoured Races in the Struggle for Life (London: John Murray, 1859).

2 C. Becquet et al. “Genetic Structure of Chimpanzee Populations”, PLoS Genetics 3 (2007): e66.

3 R. E. Green et al. “A Draft Sequence of the Neandertal Genome”, Science 328 (2010): 710–722.

4 N. J. Patterson et al. “Ancient Admixture in Human History”, Genetics 192 (2012): 1065–1093.

5 Ernst Mayr. Systematics and the Origin of Species from the Viewpoint of a Zoologist (New York: Columbia University Press, 1942).

6 J. K. Pickrell, D. Reich. “Toward a New History and Geography of Human Genes Informed by Ancient DNA”, Trends in Genetics 30 (2014): 377–389.

7 A. R. Templeton. “Biological Races in Humans”, Studies in History and Philosophy of Biological and Biomedical Science 44 (2013): 262–271.

8 M. Raghavan et al. “Upper Palaeolithic Siberian Genome Reveals Dual Ancestry of Native Americans”, Nature 505 (2014): 87–91.

9 I. Lazaridis et al. “Ancient Human Genomes Suggest Three Ancestral Populations for Present-Day Europeans”, Nature 513 (2014): 409–413.

10 I. Lazaridis et al. “Genomic Insights into the Origin of Farming in the Ancient Near East”, Nature 536 (2016): 419–424.

11 Там же.

12 F. Broushaki et al. “Early Neolithic Genomes from the Eastern Fertile Cres cent”, Science 353 (2016): 499–503; E. R. Jones et al. “Upper Palaeolithic Genomes Reveal Deep Roots of Modern Eurasians”, Nature Communications 6 (2015): 8912.

13 B. M. Henn et al. “Genomic Ancestry of North Africans Supports Back-to Africa Migrations”, PLoS Genetics 8 (2012): e1002397.

14 Lazaridis et al. “Genomic Insights”.

15 O. Bar-Yosef. “Pleistocene Connections Between Africa and Southwest Asia: An Archaeological Perspective”, African Archaeological Review 5 (1987): 29–38.

16 Lazaridis et al. “Genomic Insights”.

17 Lazaridis et al. “Ancient Human Genomes”.

18 Q. Fu et al. “The Genetic History of Ice Age Europe”, Nature 534 (2016): 200–205.

19 Q. Fu et al. “Genome Sequence of a 45,000-Year-Old Modern Human from Western Siberia”, Nature 514 (2014): 445–449.

20 Q. Fu et al. “An Early Modern Human from Romania with a Recent Neanderthal Ancestor”, Nature 524 (2015): 216–219.

21 F. G. Fedele, B. Giaccio, I. Hajdas. “Timescales and Cultural Process at 40,000 BP in the Light of the Campanian Ignimbrite Eruption, Western Eurasia”, Journal of Human Evolution 55 (2008): 834–857; A. Costa et al. “Quantifying Volcanic Ash Dispersal and Impact of the Campanian Ignimbrite Super-Eruption”, Geophysical Research Letters 39 (2012): L10310.

22 Fedele et al. “Timescales and Cultural Process”.

23 A. Seguin-Orlando et al. “Genomic Structure in Europeans Dating Back at Least 36,200 Years”, Science 346 (2014): 1113–1118.

24 Fu et al. “Ice Age Europe”.

25 Andreas Maier. The Central European Magdalenian: Regional Diversity and Inter-nal Variability (Dordrecht, The Netherlands: Springer, 2015).

26 Fu et al. “Ice Age Europe”.

27 N. A. Rosenberg et al. “Clines, Clusters, and the Effect of Study Design on the Inference of Human Population Structure”, PLoS Genetics 1 (2005): e70; G. Coop et al. “The Role of Geography in Human Adaptation”, PLoS Genetics 5 (2009): e1000500.

28 Q. Fu et al. “DNA Analysis of an Early Modern Human from Tianyuan Cave, China”, Proceedings of the National Academy of Sciences of the U.S.A. 110 (2013): 2223–2227.

29 Fu et al. “Recent Neanderthal Ancestor”; W. Haak et al. “Massive Migration from the Steppe Was a Source for Indo-European Languages in Europe”, Nature 522 (2015): 207–211.

30 R. Pinhasi et al. “Optimal Ancient DNA Yields from the Inner Ear Part of the Human Petrous Bone”, PLoS One 10 (2015): e0129102.

31 Lazaridis et al. “Genomic Insights”.

32 Там же; Broushaki et al. “Early Neolithic Genomes”.

33 I. Olalde et al. “Derived Immune and Ancestral Pigmentation Alleles in a 7,000YearOld Mesolithic European”, Nature 507 (2014): 225–228.

34 I. Mathieson et al. “Genome-Wide Patterns of Selection in 230 Ancient Eurasians”, Nature 528 (2015): 499–503.

35 I. Mathieson et al. “The Genomic History of Southeastern Europe”, bioRxiv (2017): doi.org/10.1101/135616.

36 Haak et al. “Massive Migration”; M. E. Allentoft et al. “Population Genomics of Bronze Age Eurasia”, Nature 522 (2015): 167–172.

37 Templeton. “Biological Races”.



Глава 5. Как сложилась современная Европа

1 B. Bramanti et al. “Genetic Discontinuity Between Local Hunter-Gatherers and Central Europe’s First Farmers”, Science 326 (2009): 137–40.

2 A. Keller et al. “New Insights into the Tyrolean Iceman’s Origin and Phenotype as Inferred by Whole-Genome Sequencing”, Nature Communications 3 (2012): 698.

3 W. Muller et al. “Origin and Migration of the Alpine Iceman”, Science 302 (2003): 862–866.

4 P. Skoglund et al. “Origins and Genetic Legacy of Neolithic Farmers and Hunter-Gatherers in Europe”, Science 336 (2012): 466–469.

5 Albert J. Ammerman, L. L. Cavalli-Sforza. The Neolithic Transition and the Genetics of Populations in Europe (Princeton, NJ: Princeton University Press, 1984).

6 N. J. Patterson et al. “Ancient Admixture in Human History”, Genetics 192 (2012): 1065–1093.

7 M. Raghavan et al. “Upper Palaeolithic Siberian Genome Reveals Dual Ancestry of Native Americans”, Nature (2013): doi: 10.1038/nature12736.

8 I. Lazaridis et al. “Ancient Human Genomes Suggest Three Ancestral Populations for Present-Day Europeans”, Nature 513 (2014): 409–413.

9 C. Gamba et al. “Genome Flux and Stasis in a Five Millennium Transect of European Prehistory”, Nature Communications 5 (2014): 5257; M. E. Allentoft et al. “Population Genomics of Bronze Age Eurasia”, Nature 522 (2015): 167–172; W. Haak et al. “Massive Migration from the Steppe Was a Source for Indo-European Languages in Europe”, Nature 522 (2015): 207–211; I. Mathieson et al. “Genome-Wide Patterns of Selection in 230 Ancient Eurasians”, Nature 528 (2015): 499–503.

10 L. L. Cavalli-Sforza, Paolo Menozzi, Alberto Piazza. The History and Geography of Human Genes (Princeton, NJ: Princeton University Press, 1994).

11 Haak et al. “Massive Migration”; Mathieson et al. “Genome-Wide Patterns”.

12 Q. Fu et al. “The Genetic History of Ice Age Europe”, Nature 534 (2016): 200–205.

13 I. Mathieson. “The Genomic History of Southeastern Europe”, bioRxiv (2017): doi.org/10.1101/135616.

14 K. Douka et al. “Dating Knossos and the Arrival of the Earliest Neolithic in the Southern Aegean”, Antiquity 91 (2017): 304–321.

15 Haak et al. “Massive Migration”; M. Lipson et al. “Parallel Palaeogenomic Transects Reveal Complex Genetic History of Early European Farmers”, Nature 551 (2017): 368–372.

16 Colin Renfrew. Before Civilization: The Radiocarbon Revolution and Prehistoric Europe (London: Jonathan Cape, 1973).

17 M. Gimbutas. The Prehistory of Eastern Europe, Part I: Mesolithic, Neolithic and Copper Age Cultures in Russia and the Baltic Area (American School of Prehistoric Research, Harvard University, Bulletin № 20) (Cambridge, MA: Peabody Museum, 1956).

18 David W. Anthony. The Horse, the Wheel, and Language: How Bronze-Age Riders from the Eurasian Steppes Shaped the Modern World (Princeton, NJ: Princeton University Press, 2007).

19 Там же.

20 Там же.

21 Haak et al. “Massive Migration”.

22 Там же; I. Lazaridis et al. “Genomic Insights into the Origin of Farming in the Ancient Near East”, Nature 536 (2016): 419–424.

23 M. Ivanova. “Kaukasus Und Orient: Die Entstehung des ‘MaikopPhänomens’ im 4. Jahrtausend v. Chr”, Praehistorische Zeitschrift 87 (2012): 1–28.

24 Haak et al. “Massive Migration”; Allentoft et al. “Bronze Age Eurasia”.

25 Там же.

26 G. Kossinna. “Die Deutsche Ostmark: Ein Heimatboden der Germanen”, Berlin (1919).

27 B. Arnold. “The Past as Propaganda: Totalitarian Archaeology in Nazi Germany”, Antiquity 64 (1990): 464–478.

28 H. Härke. “The Debate on Migration and Identity in Europe”, Antiquity 78 (2004): 453–456.

29 V. Heyd. “Kossinna’s Smile”, Antiquity 91 (2017): 348–359; M. Van der Linden. “Population History in Third-Millennium BC Europe: Assessing the Contribution of Genetics”, World Archaeology 48 (2016): 714–728; N. N. Johannsen, G. Larson, D. J. Meltzer, M. Van der Linden. “A Composite Window into Human History”, Science 356 (2017): 1118–1120.

30 Vere Gordon Childe. The Aryans: A Study of Indo-European Origins (London and New York: K. Paul, Trench, Trubner and Co. and Alfred A. Knopf, 1926).

31 Härke. “Debate on Migration and Identity”.

32 P. Bellwood. First Migrants: Ancient Migration in Global Perspective (Chichester, West Sussex, UK / Malden, MA: Wiley-Blackwell, 2013).

33 C. McEvedy, R. Jones. Atlas of World Population History (Harmonds-worth, Middlesex, UK: Penguin, 1978).

34 K. Kristiansen. “The Bronze Age Expansion of Indo-European Languages: An Archaeological Model” in: Becoming European: The Transformation of Third Millennium Northern and Western Europe, ed. Christopher Prescott and Håkon Glørstad (Oxford: Oxbow Books, 2011), 165–181.

35 S. Rasmussen et al. “Early Divergent Strains of Yersinia pestis in Eurasia 5,000 Years Ago”, Cell 163 (2015): 571–582.

36 A. P. Fitzpatrick. The Amesbury Archer and the Boscombe Bowmen: Bell Beaker Burials at Boscombe Down, Amesbury, Wiltshire (Salisbury, UK: Wessex Archaeology Reports, 2011).

37 I. Olalde et al. “The Beaker Phenomenon and the Genomic Transformation of Northwest Europe”, bioRxiv (2017): doi. org/10.1101/135962.

38 L. M. Cassidy et al. “Neolithic and Bronze Age Migration to Ireland and Establishment of the Insular Atlantic Genome”, Proceedings of the National Academy of Sciences of the U.S.A. 113 (2016): 368–373.

39 C. Renfrew. Archaeology and Language: The Puzzle of Indo-European Origins (Cambridge: Cambridge University Press, 1997).

40 Там же.

41 P. Bellwood. “Human Migrations and the Histories of Major Language Families” in The Global Prehistory of Human Migration (Chichester, UK, and Malden, MA: Wiley-Blackwell, 2013), 87–95.

42 С. Renfrew. Archaeology and Language; P. Bellwood, First Farmers: The Origins of Agricultural Societies (Malden, MA: Blackwell, 2005).

43 Haak et al. “Massive Migration”; Allentoft et al. “Bronze Age Eurasia”.

44 D. W. Anthony, D. Ringe. “The Indo-European Homeland from Linguistic and Archaeological Perspectives”, Annual Review of Linguistics 1 (2015): 199–219.

45 L. Poliakov. The Aryan Myth: A History of Racist and Nationalist Ideas in Europe (New York: Basic Books, 1974).



Глава 6. Столкновение, сотворившее Индию

1 The Rigveda, trans. Stephanie W. Jamison and Joel P. Brereton (Oxford: Oxford University Press, 2014), hymns 1.33, 1.53, 2.12, 3.30, 3.34, 4.16, 4.28.

2 M. Witzel. “Early Indian History: Linguistic and Textual Parameters”, in The Indo-Aryans of Ancient South Asia: Language, Material Culture and Ethnicity, ed. George Erdosy (Berlin: Walter de Gruyter, 1995), 85–125.

3 Rita P. Wright. The Ancient Indus: Urbanism, Economy, and Society (Cambridge: Cambridge University Press, 2010); Gregory L. Possehl. The Indus Civilization: A Contemporary Perspective (Lanham, MD: AltaMira Press, 2002).

4 Там же.

5 A. Parpola. Deciphering the Indus Script (Cambridge: Cambridge University Press, 1994); S. Farmer, R. Sproat, M. Witzel. “The Collapse of the Indus-Script Thesis: The Myth of a Literate Harappan Civilization”, Electronic Journal of Vedic Studies 11 (2004): 19–57.

6 Richard H. Meadow, ed. Harappa Excavations 1986–1990: A Multidisciplinary Approach to Third Millennium Urbanism (Madison, WI: Prehistory Press, 1991); A. Lawler. “Indus Collapse: The End or the Beginning of an Asian Culture?”, Science 320 (2008): 1281–1283.

7 J. Puhvel. Comparative Mythology (Baltimore: Johns Hopkins University Press, 1987).

8 Wright. The Ancient Indus; Possehl. The Indus Civilization.

9 A. Rosenberg. The Myth of the Twentieth Century: An Evaluation of the Spiritual-Intellectual Confrontations of Our Age, trans. Vivian Bird (Torrance, CA: Noontide Press, 1982).

10 L. Poliakov. The Aryan Myth: A History of Racist and Nationalist Ideas in Europe (New York: Basic Books, 1974).

11 B. Arnold. “The Past as Propaganda: Totalitarian Archaeology in Nazi Germany”, Antiquity 64 (1990): 464–478.

12 B. Ward-Perkins. The Fall of Rome and the End of Civilization (Oxford: Oxford University Press, 2005).

13 P. Bellwood. First Farmers: The Origins of Agricultural Societies (Malden, MA: Blackwell, 2005).

14 Там же.

15 M. Witzel. “Substrate Languages in Old Indo-Aryan (Rigvedic, Middle and Late Vedic)”, Electronic Journal of Vedic Studies 5 (1999): 1–67.

16 K. Thangaraj et al. “Reconstructing the Origin of Andaman Islanders”, Science 308 (2005): 996; K. Thangaraj et al. “In situ Origin of Deep Rooting Lineages of Mitochondrial Macrohaplogroup ‘M’ in India”, BMC Genomics 7 (2006): 151.

17 R. S. Wells et al. “The Eurasian Heartland: A Continental Perspective on Y-chromosome Diversity”, Proceedings of the National Academy of Sciences of the U.S.A. 98 (2001): 10244–10249; M. Bamshad et al. “Genetic Evidence on the Origins of Indian Caste Populations”, Genome Research 11 (2001): 994–1004; I. Thanseem et al. “Genetic Affinities Among the Lower Castes and Tribal Groups of India: Inference from Y Chromosome and Mitochondrial DNA”, BMC Genetics 7 (2006): 42.

18 Thangaraj et al. “Andaman Islanders”.

19 D. Reich et al. “Reconstructing Indian Population History”, Nature 461 (2009): 489–494.

20 R. E. Green et al. “A Draft Sequence of the Neandertal Genome”, Science 328 (2010): 710–722.

21 Thangaraj et al. “Deep Rooting Lineages”.

22 Reich et al. “Reconstructing Indian Population History”; P. Moorjani et al. “Genetic Evidence for Recent Population Mixture in India”, American Journal of Human Genetics 93 (2013): 422–438.

23 Там же.

24 I. Karve. Hindu Society – An Interpretation (Pune, India: Deccan College Post Graduate and Research Institute, 1961).

25 P. A. Underhill et al. “The Phylogenetic and Geographic Structure of Y-Chromosome Haplogroup R1a”, European Journal of Human Genetics 23 (2015): 124–131.

26 S. Perur. “The Origins of Indians: What Our Genes Are Telling Us”, Fountain Ink, December 3, 2013, http://fountainink.in/?p=4669&all=1.

27 K. Bryc et al. “The Genetic Ancestry of African Americans, Latinos, and European Americans Across the United States”, American Journal of Human Genetics 96 (2015): 37–53.

28 L. G. Carvajal-Carmona et al. “Strong Amerind/White Sex Bias and a Possible Sephardic Contribution Among the Founders of a Population in Northwest Colombia”, American Journal of Human Genetics 67 (2000): 1287–1295; G. Bedoya et al. “Admixture Dynamics in Hispanics: A Shift in the Nuclear Genetic Ancestry of a South American Population Isolate”, Proceedings of the National Academy of Sciences of the U.S.A. 103 (2006): 7234–7239.

29 Moorjani et al. “Recent Population Mixture”.

30 Там же.

31 R. Thapar. Early India: From the Origins to AD 1300 (Berkeley: University of California Press, 2002); Karve. Hindu Society; Susan Bayly, Caste, Society and Politics in India from the Eighteenth Century to the Modern Age (Cambridge: Cambridge University Press, 1999); M. N. Srinivas. Caste in Modern India and Other Essays (Bombay: Asia Publishing House, 1962); L. Dumont. Homo Hierarchicus: The Caste System and Its Implications (Chicago: University of Chicago Press, 1980).

32 K. Suresh Singh. People of India: An Introduction (People of India National Series) (New Delhi: Oxford University Press, 2002); K. C. Malhotra, T. S. Vasulu. “Structure of Human Populations in India” in Human Population Genetics: A Centennial Tribute to J. B. S. Haldane, ed. P. P. Majumder (New York: Plenum Press, 1993), 207–234.

33 Karve. “Hindu Society”.

34 Там же.

35 N. B. Dirks. Castes of Mind: Colonialism and the Making of Modern India (Princeton, NJ: Princeton University Press, 2001); N. Boivin. “Anthropological, Historical, Archaeological and Genetic Perspectives on the Origins of Caste in South Asia” in The Evolution and History of Human Populations in South Asia, ed. M. D. Petraglia and B. Allchin (Dordrecht, The Netherlands: Springer, 2007), 341–362.

36 Reich et al. “Reconstructing Indian Population History”.

37 M. Arcos-Burgos, M. Muenke. “Genetics of Population Isolates”, Clinical Genetics 61 (2002): 233–247.

38 N. Nakatsuka et al. “The Promise of Discovering PopulationSpecific Disease-Associated Genes in South Asia”, Nature Genetics 49 (2017): 1403–1407.

39 Reich et al. “Reconstructing Indian Population History”.

40 I. Manoharan et al. “Naturally Occurring Mutation Leu307Pro of Human Butyrylcholinesterase in the Vysya Community of India”, Pharmacogenetics and Genomics 16 (2006): 461–468.

41 A. E. Raz. “Can Population-Based Carrier Screening Be Left to the Community?”, Journal of Genetic Counseling 18 (2009): 114–118.

42 I. Lazaridis et al. “Genomic Insights into the Origin of Farming in the Ancient Near East”, Nature 536 (2016): 419–424; F. Broushaki et al. “Early Neolithic Genomes from the Eastern Fertile Crescent”, Science 353 (2016): 499–503.

43 Там же.

44 Lazaridis et al. “Genomic Insights”.

45 Неопубликованные результаты из лаборатории Дэвида Райха.



Глава 7. В поисках предков индейцев

1 B. Mindlin. Unwritten Stories of the Suruí Indians of Rondônia (Austin: Institute of Latin American Studies; distributed by the University of Texas Press, 1995).

2 D. Reich et al. “Reconstructing Native American Population History”, Nature 488 (2012): 370–374.

3 P. Skoglund et al. “Genetic Evidence for Two Founding Populations of the Americas”, Nature 525 (2015): 104–108.

4 P. D. Heintzman et al. “Bison Phylogeography Constrains Dispersal and Viability of the Ice Free Corridor in Western Canada”, Proceedings of the National Academy of Sciences of the U.S.A. 113 (2016): 8057–8063; M. W. Pedersen et al. “Postglacial Viability and Colonization in North America’s Ice-Free Corridor”, Nature 537 (2016): 45–49.

5 J. Acosta. Historia Natural y Moral de las Indias: En que se Tratan las Cosas Notables del Cielo y Elementos, Metales, Plantas y Animales de Ellas y los Ritos, Ceremonias, Leyes y Gobierno y Guerras de los Indios (Seville: Juan de León, 1590).

6 D. J. Meltzer. First Peoples in a New World: Colonizing Ice Age America (Berkeley: University of California Press, 2009).

7 J. H. Greenberg, C. G. Turner II, S. L. Zegura. “The Settlement of the Americas: A Comparison of the Linguistic, Dental, and Genetic Evidence”, Current Anthropology 27 (1986): 477–497.

8 P. Forster, R. Harding, A. Torroni, H.-J. Bandelt. “Ori gin and Evolution of Native American mtDNA Variation: A Reappraisal”, American Journal of Human Genetics 59 (1996): 935–45; E. Tamm et al. “Beringian Standstill and Spread of Native American Founders”, PloS One 2 (2017): e829.

9 T. D. Dillehay et al. “Monte Verde: Seaweed, Food, Medicine, and the Peopling of South America”, Science 320 (2008): 784–786.

10 D. L. Jenkins et al. “Clovis Age Western Stemmed Projectile Points and Human Coprolites at the Paisley Caves”, Science 337 (2012): 223–228.

11 M. Rasmussen et al. “The Genome of a Late Pleistocene Human from a Clovis Burial Site in Western Montana”, Nature 506 (2014): 225–229.

12 Povos Indígenas No Brasil. “Karitiana: Biopiracy and the Unauthor-ized Collection of Biomedical Samples”, https://pib.socioambiental. org/en/povo/ karitiana/389.

13 N. A. Garrison, M. K. Cho. “Awareness and Acceptable Practices: IRB and Researcher Reflections on the Havasupai Lawsuit”, AJOB Primary Research 4 (2013): 55–63; A. Harmon. “Indian Tribe Wins Fight to Limit Research of Its DNA”, New York Times, April 21, 2010.

14 Ronald P. Maldonado. “Key Points for University Researchers When Considering a Research Project with the Navajo Nation”, http://nptao.arizona.edu/sites/nptao/files/navajonationkey- researchrequirements_0.pdf

15 R. Skloot. The Immortal Life of Henrietta Lacks (New York: Crown, 2010).

16 B. L. Shelton. “Consent and Consultation in Genetic Research on American Indians and Alaska Natives”, http://www.ipcb.org/publi- cations/briefing_papers/files/consent.html.

17 R. R. Sharp, M. W. Foster. “Involving Study Populations in the Review of Genetic Research”, Journal of Law, Medicine and Ethics 28 (2000): 41–51; International HapMap Consortium. “The International HapMap Project”, Nature 426 (2003): 789–796.

18 T. Egan. “Tribe Stops Study of Bones That Challenge History”, New York Times, September 30, 1996; D. W. Owsley, R. L. Jantz. Kennewick Man: The Scientific Investigation of an Ancient American Skeleton (College Station: Texas A&M University Press, 2014); D. J. Meltzer. “Kennewick Man: Coming to Closure”, Antiquity 348 (2015): 1485–1493.

19 M. Rasmussen et al. “The Ancestry and Affiliations of Kennewick Man”, Nature 523 (2015): 455–458.

20 Там же.

21 J. Lindo et al. “Ancient Individuals from the North American Northwest Coast Reveal 10,000 Years of Regional Genetic Continuity”, Proceedings of the National Academy of Sciences of the U.S.A. 114 (2017): 4093–4098.

22 S. J. Redman. Bone Rooms: From Scientific Racism to Human Prehistory in Museums (Cambridge, MA, and London: Harvard University Press, 2016).

23 M. Rasmussen et al. “An Aboriginal Australian Genome Reveals Separate Human Dispersals into Asia”, Science 334 (2011): 94–98.

24 Rasmussen et al. “Genome of a Late Pleistocene Human”.

25 Rasmussen et al. “Ancestry and Affiliations of Kennewick Man”.

26 A. S. Malaspinas et al. “A Genomic History of Aboriginal Australia”, Nature 538 (2016): 207–214.

27 E. Callaway. “Ancient Genome Delivers ‘Spirit Cave Mummy’ to US tribe”, Nature 540 (2016): 178–179.

28 Там же.

29 M. Livi-Bacci. “The Depopulation of Hispanic America After the Conquest”, Population and Development Review 32 (2006): 199–232; L. H. Morgan. Ancient Society; Or, Researches in the Lines of Human Progress from Savagery Through Barbarism to Civilization (Chicago: Charles H. Kerr, 1909).

30 Reich et al. “Reconstructing Native American Population History”.

31 Lindo et al. “Ancient Individuals”.

32 L. Campbell, M. Mithun. The Languages of Native America: Historical and Comparative Assessment (Austin: University of Texas Press, 1979).

33 L. Campbell. “Comment on Greenberg, Turner and Zegura”, Current Anthropology 27 (1986): 488.

34 P. Bellwood. First Migrants: Ancient Migration in Global Perspective (Chichester, West Sussex, UK / Malden, MA: Wiley-Blackwell, 2013).

35 Reich et al. “Reconstructing Native American Population History”.

36 W. A. Neves, M. Hubbe. “Cranial Morphology of Early Americans from Lagoa Santa, Brazil: Implications for the Settlement of the New World”, Proceedings of the National Academy of Sciences of the U.S.A. 102 (2005): 18309–18314.

37 Rasmussen et al. “Ancestry and Affiliations of Kennewick Man”.

38 P. Skoglund et al. “Genetic Evidence for Two Founding Populations of the Americas”, Nature 525 (2015): 104–108.

39 Povos Indígenas No Brasil. “Surui Paiter: Introduction”, https://pib.socioambiental.org/en/povo/surui-paiter; R. A. Butler. “Amazon Indians Use Google Earth, GPS to Protect Forest Home”, Mongabay: News and Inspiration from Nature’s Frontline, November 15, 2006, https://news.mongabay.com/2006/11/amazon-indians-use-googleearth-gps-to-protect-forest-home/

40 “Karitiana: Biopiracy and the Unauthorized Collection”.

41 Povos Indígenas No Brasil. “Xavante: Introduction”, https://pib.socioambiental.org/en/povo/xavante

42 M. Raghavan et al. “Genomic Evidence for the Pleistocene and Recent Population History of Native Americans”, Science 349 (2015): aab3884.

43 E. J. Vajda. “A Siberian Link with Na-Dene Languages”, in Anthropological Papers of the University of Alaska: New Series, ed. James M. Kari and Ben Austin Potter, 5 (2010): 33–99.

44 Reich et al. “Reconstructing Native American Population History”.

45 M. Rasmussen et al. “Ancient Human Genome Sequence of an Extinct Palaeo-Eskimo”, Nature 463 (2010): 757–762.

46 M. Raghavan et al. “The Genetic Prehistory of the New World Arctic”, Science 345 (2014): 1255832.

47 P. Flegontov et al. “Paleo-Eskimo Genetic Legacy Across North America”, bioRxiv (2017): doi.org/10.1101.203018.

48 Flegontov et al. “Paleo-Eskimo Genetic Legacy”.

49 T. M. Friesen. “Pan-Arctic Population Movements: The Early Paleo-Inuit and Thule Inuit Migrations” in: The Oxford Handbook of the Prehistoric Arctic, ed. T. Max Friesen and Owen K. Mason (New York: Oxford University Press, 2016), 673–692.

50 Reich et al. “Reconstructing Native American Population History”.

51 J. Diamond, P. Bellwood. “Farmers and Their Languages: The First Expansions”, Science 300 (2003): 597–603; P. Bellwood. First Farmers: The Origins of Agricultural Societies (Malden, MA: Blackwell, 2005).

52 R. R. da Fonseca et al. “The Origin and Evolution of Maize in the Southwestern United States”, Nature Plants 1 (2015): 14003.



Глава 8. Генетические корни восточных азиатов

1 X. H. Wu et al. “Early Pottery at 20,000 Years Ago in Xianrendong Cave, China”, Science 336 (2012): 1696–1700.

2 R. X. Zhu et al. “Early Evidence of the Genus Homo in East Asia”, Journal of Human Evolution 55 (2008): 1075–85.

3 C. C. Swisher III et al. “Age of the Earliest Known Hominids in Java, Indonesia”, Science 263 (1994): 1118–1121; P. Bellwood. First Islanders: Prehistory and Human Migration in Island Southeast Asia (Oxford: Wiley-Blackwell, 2017).

4 D. Richter et al. “The Age of the Hominin Fossils from Jebel Irhoud, Morocco, and the Origins of the Middle Stone Age”, Nature 546 (2017): 293–296; J. G. Fleagle, Z. Assefa, F. H. Brown, J. J. Shea. “Paleoanthropology of the Kibish Formation, Southern Ethiopia: Introduction”, Journal of Human Evolution 55 (2008): 360–365.

5 T. Sutikna et al. “Revised Stratigraphy and Chronology for Homo floresiensis at Liang Bua in Indonesia”, Nature 532 (2016): 366–369.

6 Y. Ke et al. “African Origin of Modern Humans in East Asia: A Tale of 12,000 Y Chromosomes”, Science 292 (2001): 1151–1153.

7 J. Qiu. “The Forgotten Continent: Fossil Finds in China Are Challenging Ideas About the Evolution of Modern Humans and Our Closest Relatives”, Nature 535 (2016): 218–220.

8 R. J. Rabett, P. J. Piper. “The Emergence of Bone Technologies at the End of the Pleistocene in Southeast Asia: Regional and Evolutionary Implications”, Cambridge Archaeological Journal 22 (2012): 37–56; M. C. Langley, C. Clarkson, S. Ulm. “From Small Holes to Grand Narratives: The Impact of Taphonomy and Sample Size on the Modernity Debate in Australia and New Guinea”, Journal of Human Evolution 61 (2011): 197–208; M. Aubert et al. “Pleistocene Cave Art from Sulawesi, Indonesia”, Nature 514 (2014): 223–227.

9 Langley, Clarkson, Ulm. “From Small Holes to Grand Narratives”; J. F. Connell, J. Allen. “The Process, Biotic Impact, and Global Implications of the Human Colonization of Sahul About 47,000 Years Ago”, Journal of Archaeological Science 56 (2015): 73–84.

10 J.-J. Hublin. “The Modern Human Colonization of Western Eurasia: When and Where?”, Quaternary Science Reviews 118 (2015): 194–210.

11 R. Foley, M. M. Lahr. “Mode 3 Technologies and the Evolu tion of Modern Humans”, Cambridge Archaeological Journal 7 (1997): 3–36.

12 M. M. Lahr, R. Foley. “Multiple Dispersals and Modern Human Origins”, Evolutionary Anthropology 3 (1994): 48–60.

13 H. Reyes-Centeno et al. “Testing Modern Human Out-ofAfrica Dispersal Models and Implications for Modern Human Origins”, Journal of Human Evolution 87 (2015): 95–106.

14 H. S. Groucutt et al. “Rethinking the Dispersal of Homo sapiens Out of Africa”, Evolutionary Anthropology 24 (2015): 149–164.

15 R. Grün et al. “U-series and ESR Analyses of Bones and Teeth Relating to the Human Burials from Skhul”, Journal of Human Evolution 49 (2005): 316–334.

16 S. J. Armitage et al. “The Southern Route ‘Out of Africa’: Evidence for an Early Expansion of Modern Humans into Arabia”, Science 331 (2011): 453–456; M. D. Petraglia. “Trailblazers Across Africa”, Nature 470 (2011): 50–51.

17 M. Kuhlwilm et al. “Ancient Gene Flow from Early Modern Humans into Eastern Neanderthals”, Nature 530 (2016): 429–433.

18 M. Rasmussen et al. “An Aboriginal Australian Genome Reveals Separate Human Dispersals into Asia”, Science 334 (2011): 94–98.

19 D. Reich et al. “Genetic History of an Archaic Hominin Group from Denisova Cave in Siberia”, Nature 468 (2010): 1053–1060;

M. Meyer et al. “A High-Coverage Genome Sequence from an Archaic Denisovan Individual”, Science 338 (2012): 222–226.

20 S. Mallick et al. “The Simons Genome Diversity Project: 300 Genomes from 142 Diverse Populations”, Nature 538 (2016): 201–206.

21 Q. Fu et al. “Genome Sequence of a 45,000-Year-Old Modern Human from Western Siberia”, Nature 514 (2014): 445–449; S. Sankararaman, S. Mallick, N. Patterson, D. Reich. “The Combined Landscape of Denisovan and Neanderthal Ancestry in PresentDay Humans”, Current Biology 26 (2016): 1241–1247; P. Moorjani et al. “ A Genetic Method for Dating Ancient Genomes Provides a Direct Estimate of Human Generation Interval in the Last 45,000 Years”, Proceedings of the National Academy of Sciences of the U.S.A. 113 (2016): 5652–5627.

22 Mallick et al. “Simons Genome Diversity Project”; M. Lipson, D. Reich. “A Working Model of the Deep Relationships of Diverse Modern Human Genetic Lineages Outside of Africa”, Molecular Biology and Evolution 34 (2017): 889–902.

23 Mallick et al. “Simons Genome Diversity Project”; A. S. Malaspinas et al. “A Genomic History of Aboriginal Australia”, Nature 538 (2016): 207–214; L. Pagani et al. “Genomic Analyses Inform on Migration Events During the Peopling of Eurasia”, Nature 538 (2016): 238–242.

24 Hublin. “Modern Human Colonization of Western Eurasia”.

25 M. Raghavan et al. “Upper Palaeolithic Siberian Genome Reveals Dual Ancestry of Native Americans”, Nature (2013): doi: 10.1038/nature12736.

26 Hugo Pan-Asian SNP Consortium. “Mapping Human Genetic Diversity in Asia”, Science 326 (2009): 1541–45.

27 S. Ramachandran et al. “Support from the Relationship of Genetic and Geographic Distance in Human Populations for a Serial Founder Effect Originating in Africa”, Proceedings of the National Academy of Sciences of the U.S.A. 102 (2005): 15942–15947; B. M. Henn, L. L. Cavalli-Sforza, M. W. Feldman. “The Great Human Expansion”, Proceedings of the National Academy of Sciences of the U.S.A. 109 (2012): 17758–17764.

28 J. K. Pickrell, D. Reich. “Toward a New History and Geography of Human Genes Informed by Ancient DNA”, Trends in Genetics 30 (2014): 377–389.

29 Неопубликованные данные лаборатории Дэвида Райха.

30 V. Siska et al. “Genome-Wide Data from Two Early Neolithic East Asian Individuals Dating to 7700 Years Ago”, Science Advances 3 (2017): e1601877.

31 P. Bellwood. First Farmers: The Origins of Agricultural Societies (Malden, MA: Blackwell, 2005).

32 J. Diamond, P. Bellwood. “Farmers and Their Languages: The First Expansions”, Science 300 (2003): 597–603.

33 S. Xu et al. “Genomic Dissection of Population Substructure of Han Chinese and Its Implication in Association Studies”, American Journal of Human Genetics 85 (2009): 762–774; J. M. Chen et al. “Genetic Structure of the Han Chinese Population Revealed by Genome-Wide SNP Variation”, American Journal of Human Genetics 85 (2009): 775–785.

34 B. Wen et al. “Genetic Evidence Supports Demic Diffusion of Han Culture”, Nature 431 (2004): 302–305.

35 F. H. Chen et al. “Agriculture Facilitated Permanent Human Occupation of the Tibetan Plateau After 3600 B.P.”, Science 347 (2015): 248–250.

36 Неопубликованные данные лаборатории Дэвида Райха.

37 T. A. Jinam et al. “Unique Characteristics of the Ainu Population in Northern Japan”, Journal of Human Genetics 60 (2015): 565–571.

38 Там же; P. R. Loh et al. “Inferring Admixture Histories of Human Populations Using Linkage Disequilibrium”, Genetics 193 (2013): 1233–1254.

39 Неопубликованные данные лаборатории Дэвида Райха; Bellwood, First Migrants.

40 Diamond, Bellwood. “Farmers and Their Languages”.

41 M. Lipson et al. “Reconstructing Austronesian Population History in Island Southeast Asia”, Nature Communications 5 (2014): 4689.

42 R. Blench. “Was There an Austroasiatic Presence in Island Southeast Asia Prior to the Austronesian Expansion?”, Bulletin of the Indo-Pacific Prehistory Association 30 (2010): 133–144.

43 Bellwood. First Migrants.

44 A. Crowther et al. “Ancient Crops Provide First Archaeological Signature of the Westward Austronesian Expansion”, Proceedings of the National Academy of Sciences of the U.S.A. 113 (2016): 6635–6640.

45 Lipson et al. “Reconstructing Austronesian Population History”.

46 A. Wollstein et al. “Demographic History of Oceania Inferred from Genome-Wide Data”, Current Biology 20 (2010): 1983–1992; M. Kayser. “The Human Genetic History of Oceania: Near and Remote Views of Dispersal”, Current Biology 20 (2010): R194–201; E. Matisoo-Smith. “Ancient DNA and the Human Settlement of the Pacific: A Review”, Journal of Human Evolution 79 (2015): 93– 104.

47 D. Reich et al. “Denisova Admixture and the First Modern Human Dispersals into Southeast Asia and Oceania”, American Journal of Human Genetics 89 (2011): 516–528; P. Skoglund et al. “Genomic Insights into the Peopling of the Southwest Pacific”, Nature 538 (2016): 510–513.

48 R. Pinhasi et al. “Optimal Ancient DNA Yields from the Inner Ear Part of the Human Petrous Bone”, PLoS One 10 (2015): e0129102.

49 Skoglund et al. “Genomic Insights”.

50 Там же.

51 Неопубликованные данные лабораторий Дэвида Райха и Йоханнеса Краузе.

52 Там же.



Глава 9. Африка снова включается в историю человечества

1 J. Lachance et al. “Evolutionary History and Adaptation from High-Coverage Whole-Genome Sequences of Diverse African Hunter-Gatherers”, Cell 150 (2012): 457–469.

2 V. Plagnol, J. D. Wall. “Possible Ancestral Structure in Human Populations”, PLoS Genetics 2 (2006): e105; J. D. Wall,

K. E. Lohmueller, V. Plagnol. “Detecting Ancient Admixture and Estimating Demographic Parameters in Multiple Human Populations”, Molecular Biology and Evolution 26 (2009): 1823–1827.

3 M. F. Hammer et al. “Genetic Evidence for Archaic Admix ture in Africa”, Proceedings of the National Academy of Sciences of the U.S.A. 108 (2011): 15123–15128.

4 K. Harvati et al. “The Later Stone Age Calvaria from Iwo Eleru, Nigeria: Morphology and Chronology”, PLoS One 6 (2011): e24024; I. Crevecoeur, A. Brooks, I. Ribot, E. Cornelissen, and P. Semal. “The Late Stone Age Human Remains from Ishango (Democratic Republic of Congo): New Insights on Late Pleistocene Modern Human Diversity in Africa”, American Journal of Physical Anthropology 96 (2016): 35–57.

5 Неопубликованные данные лаборатории Дэвида Райха.

6 D. Richter et al. “The Age of the Hominin Fossils from Jebel Irhoud, Morocco, and the Origins of the Middle Stone Age”, Nature 546 (2017): 293–296; J. G. Fleagle, Z. Assefa, F. H. Brown, J. J. Shea. “Paleoanthropology of the Kibish Formation, Southern Ethiopia: Introduction”, Journal of Human Evolution 55 (2008): 360–365.

7 H. Li and R. Durbin. “Inference of Human Population Histo ry from Individual Whole-Genome Sequences”, Nature 475 (2011): 493–496.

8 Li, Durbin. “Inference of Human Population History”; K. Prüfer et al. “The Complete Genome Sequence of a Neanderthal from the Altai Mountains”, Nature (2013): doi: 10.1038/nature12886.

9 P. H. Dirks et al. “The Age of Homo Naledi and Associated Sediments in the Rising Star Cave, South Africa”, eLife 6 (2017): e24231.

10 I. Gronau et al. “Bayesian Inference of Ancient Human Demography from Individual Genome Sequences”, Nature Genetics 43 (2011): 1031–1034.

11 P. Skoglund et al. “Reconstructing Prehistoric African Population Structure”, Cell 171 (2017): 5694.

12 S. Mallick et al. “The Simons Genome Diversity Project: 300 Genomes from 142 Diverse Populations”, Nature 538 (2016): 201–206; Gronau et al. “Bayesian Inference”.

13 S. A. Tishkoff et al. “The Genetic Structure and History of Africans and African Americans”, Science 324 (2009): 1035–1044.

14 C. J. Holden. “Bantu Language Trees Reflect the Spread of Farming Across Sub-Saharan Africa: A Maximum-Parsimony Analysis”, Proceedings of the Royal Society B – Biological Sciences 269 (2002): 793–799; P. de Maret. “Archaeologies of the Bantu Expansion”, in The Oxford Handbook of African Archaeology, ed. Peter Mitchell and Paul J. Lane (Oxford: Oxford University Press, 2013), 627–644.

15 K. Bostoen et al. “Middle to Late Holocene Paleoclima tic Change and the Early Bantu Expansion in the Rain Forests of Western Central Africa”, Current Anthropology 56 (2016): 354–384; K. Manning et al. “4,500-Year-Old Domesticated Pearl Millet (Pennisetum glaucum) from the Tilemsi Valley, Mali: New Insights into an Alternative Cereal Domestication Pathway”, Journal of Archaeological Science 38 (2011): 312–322.

16 D. Killick. “Cairo to Cape: The Spread of Metallurgy Through Eastern and Southern Africa”, Journal of World Prehistory 22 (2009): 399–414.

17 De Maret. “Archaeologies of the Bantu Expansion”.

18 Holden. “Bantu Language Trees”.

19 Bostoen et al. “Middle to Late Holocene”; Manning et al. “4,500-Year-Old”.

20 D. J. Lawson, G. Hellenthal, S. Myers, D. Falush. “Inference of Population Structure Using Dense Haplotype Data”, PLoS Genetics 8 (2012): e1002453; G. Hellenthal et al. “A Genetic Atlas of Human Admixture History”, Science 343 (2014): 747–751; C. de Filippo, K. Bostoen, M. Stoneking, B. Pakendorf. “Bringing Together Linguistic and Genetic Evidence to Test the Bantu Expansion”, Proceedings of the Royal Society B – Biological Sciences 279 (2012): 3256–3263; E. Patin et al. “Dispersals and Genetic Adaptation of Bantu-Speaking Populations in Africa and North America”, Science 356 (2017): 543–546; G. B. Busby et al. “Admixture Into and Within Sub-Saharan Africa”, eLife 5(2016): e15266.

21 Tishkoff et al. “Genetic Structure and History”; G. Ayodo et al. “Combining Evidence of Natural Selection with Association Analysis Increases Power to Detect Malaria-Resistance Variants”, American Journal of Human Genetics 81 (2007): 234–242.

22 C. Ehret. “Reconstructing Ancient Kinship in Africa”, in Ear ly Human Kinship: From Sex to Social Reproduction, ed. Nicholas J. Allen, Hilary Callan, Robin Dunbar, Wendy James (Malden, MA: Blackwell, 2008), 200–231; C. Ehret, S. O. Y. Keita, P. Newman. “The Origins of Afroasiatic”, Science 306 (2004): 1680–1681.

23 J. Diamond, P. Bellwood. “Farmers and Their Languages: The First Expansions”, Science 300 (2003): 597–603; P. Bellwood. “Response to Ehret et al. ‘The Origins of Afroasiatic,’ ” Science 306 (2004): 1681.

24 D. Q. Fuller, E. Hildebrand. “Domesticating Plants in Africa” in: The Oxford Handbook of African Archaeology, ed. Peter Mitchell, Paul J. Lane (Oxford: Oxford University Press, 2013), 507–526; M. Madella et al. “Microbotanical Evidence of Domestic Cereals in Africa 7000 Years Ago”, PLoS One 9 (2014): e110177.

25 I. Lazaridis et al. “Genomic Insights into the Origin of Farming in the Ancient Near East”, Nature 536 (2016): 419–424; Skoglund et al. “Reconstructing Prehistoric African Population Structure”.

26 Lazaridis et al. “Genomic Insights”; Skoglund et al. “Reconstructing Prehistoric African Population Structure”; V. J. Schuenemann et al. “Ancient Egyptian Mummy Genomes Suggest an Increase of Sub-Saharan African Ancestry in Post-Roman Periods”, Nature Communications 8 (2017): 15694.

27 T. Güldemann. “A Linguist’s View: Khoe-Kwadi Speakers as the Earliest Food-Producers of Southern Africa”, Southern African Humanities 20 (2008): 93–132.

28 J. K. Pickrell et al. “Ancient West Eurasian Ancestry in Southern and Eastern Africa”, Proceedings of the National Academy of Sciences of the U.S.A. 111 (2014): 2632–2637.

29 Pagani et al. “Ethiopian Genetic Diversity”.

30 Skoglund et al. “Reconstructing Prehistoric African Population Structure”.

31 L. L. Cavalli-Sforza, F. Cavalli-Sforza. The Great Human Diasporas: The History of Diversity and Evolution (Reading, MA: Addison-Wesley, 1995).

32 M. Gallego Llorente et al. “Ancient Ethiopian Genome Reveals Extensive Eurasian Admixture Throughout the African Continent”, Science 350 (2015): 820–822.

33 D. N. Levine. Greater Ethiopia: The Evolution of a Multiethnic Society (Chicago: University of Chicago Press, 2000).

34 L. Van Dorp et al. “Evidence for a Common Origin of Blacksmiths and Cultivators in the Ethiopian Ari Within the Last 4500 Years: Lessons for Clustering-Based Inference”, PLoS Genetics 11 (2015): e1005397.

35 D. Reich et al. “Reconstructing Indian Population History”, Nature 461 (2009): 489–494.

36 Skoglund et al. “Reconstructing Prehistoric African Population Structure”.

37 Там же.

38 Там же.

39 J. K. Pickrell et al. “The Genetic Prehistory of Southern Africa”, Nature Communications 3 (2012): 1143; C. M. Schlebusch et al. “Genomic Variation in Seven Khoe-San Groups Reveals Adaptation and Complex African History”, Science 338 (2012): 374–379; Mallick et al. “Simons Genome Diversity Project”.

40 M. E. Prendergast et al. “Continental Island Formation and the Archaeology of Defaunation on Zanzibar, Eastern Africa”, PLoS One 11 (2016): e0149565.

41 Skoglund et al. “Reconstructing Prehistoric African Population Structure”.

42 P. Ralph, G. Coop. “Parallel Adaptation: One or Many Waves of Advance of an Advantageous Allele?”, Genetics 186 (2010): 647–668.

43 S. A. Tishkoff et al. “Convergent Adaptation of Human Lactase Persistence in Africa and Europe”, Nature Genetics 39 (2007): 31–40.

44 Ralph, Coop. “Parallel Adaptation”.



Глава 10. Геномика неравенства

1 P. Wade. Race and Ethnicity in Latin America (London and New York: Pluto Press, 2010).

2 Trans-Atlantic Slave Trade Database, www.slavevoyages.org/assess- ment/ estimates

3 K. Bryc et al. “The Genetic Ancestry of African Americans, Latinos, and European Americans Across the United States”, American Journal of Human Genetics 96 (2015): 37–53.

4 А. Piers. Race Against Time (New York: Hawthorn Books, 1973).

5 Первая государственная перепись в 1790 г. зарегистрировала в Виргинии 292627 рабов-мужчин, а общее число мужчин в штате составляло 747610; данные с сайта www.nationalgeographic.org/ media/us-census-1790/

6 J. D. Rothman. Notorious in the Neighborhood: Sex and Families Across the Color Line in Virginia, 1787–1861 (Chapel Hill: University of North Carolina Press, 2003).

7 E. A. Foster et al. “Jefferson Fathered Slave’s Last Child”, Nature 396 (1998): 27–28.

8 “Statement on the TJMF Research Committee Report on Thomas Jefferson and Sally Hemings”, January 26, 2000, в свободном доступе на сайте https://www.monticello.org/sites/default/files/ inline-pdfs/jefferson-hemings_report.pdf

9 M. Hemings. “Life Among the Lowly, No. 1”, Pike County (Ohio) Republican, March 13, 1873.

10 E. J. Parra et al. “Ancestral Proportions and Admixture Dynamics in Geographically Defined African Americans Living in South Carolina”, American Journal of Physical Anthropology 114 (2001): 18–29.

11 Там же.

12 Bryc et al. “Genetic Ancestry”.

13 J. N. Fenner. “Cross-Cultural Estimation of the Human Generation Interval for Use in Genetics-Based Population Divergence Studies”, American Journal of Physical Anthropology 128 (2005): 415–423.

14 D. Morgan. The Mongols (Malden, MA, and Oxford: Blackwell, 2007).

15 T. Zerjal et al. “The Genetic Legacy of the Mongols”, American Journal of Human Genetics 72 (2003): 717–721.

16 L. T. Moore et al. “A Y-Chromosome Signature of Hegemony in Gaelic Ireland”, American Journal of Human Genetics 78 (2006): 334–338.

17 S. Lippold et al. “Human Paternal and Maternal Demographic Histories: Insights from High-Resolution Y Chromosome and mtDNA Sequences”, Investigative Genetics 5 (2014): 13; M. Karmin et al. “A Recent Bottleneck of Y Chromosome Diversity Coincides with a Global Change in Culture”, Genome Research 25 (2015): 459–466.

18 Там же.

19 A. Sherratt. “Plough and Pastoralism: Aspects of the Secondary Products Revolution”, in Pattern of the Past: Studies in Honour of David Clarke, ed. Ian Hodder, Glynn Isaac, Norman Hammond (Cambridge: Cambridge University Press, 1981), 261–306.

20 D. W. Anthony. The Horse, the Wheel, and Language: How BronzeAge Riders from the Eurasian Steppes Shaped the Modern World (Princeton, NJ: Princeton University Press, 2007).

21 W. Haak et al. “Massive Migration from the Steppe Was a Source for Indo-European Languages in Europe”, Nature 522 (2015): 207–211; M. E. Allentoft et al. “Population Genomics of Bronze Age Eurasia”, Nature 522 (2015): 167–172.

22 E. Murphy, A. Khokhlov. “A Bioarchaeological Study of Prehistoric Populations from the Volga Region” in: A Bronze Age Landscape in the Russian Steppes: The Samara Valley Project, Monumenta Archaeologica 37, ed. David W. Anthony, Dorcas R. Brown, Aleksandr A. Khokhlov, Pavel V. Kuznetsov, Oleg D. Mochalov (Los Angeles: Cotsen Institute of Archaeology Press, 2016), 149–216.

23 M. Gimbutas. The Prehistory of Eastern Europe, Part I: Mesolithic, Neolithic and Copper Age Cultures in Russia and the Baltic Area (American School of Prehistoric Research, Harvard University, Bulletin No. 20) (Cambridge, MA: Peabody Museum, 1956).

24 Haak et al. “Massive Migration”.

25 R. S. Wells et al. “The Eurasian Heartland: A Continental Perspective on Y-Chromosome Diversity”, Proceedings of the National Academy of Sciences of the U.S.A. 98 (2001): 10244–10249.

26 R. Martiniano et al. “The Population Genomics of Archaeological Transition in West Iberia: Investigation of Ancient Substructure Using Imputation and Haplotype-Based Methods”, PLoS Genetics 13 (2017): e1006852.

27 M. Silva et al. “A Genetic Chronology for the Indian Subcontinent Points to Heavily Sex-Biased Dispersals”, BMC Evolutionary Biology 17 (2017): 88.

28 Martiniano et al. “West Iberia”; неопубликованные результаты лаборатории Дэвида Райха.

29 J. A. Tennessen et al. “Evolution and Functional Impact of Rare Coding Variation from Deep Sequencing of Human Exomes”, Science 337 (2012): 64–69.

30 A. Keinan, J. C. Mullikin, N. Patterson, D. Reich. “Accelerated Genetic Drift on Chromosome X During the Human Dispersal out of Africa”, Nature Genetics 41 (2009): 66–70; A. Keinan, D. Reich. “Can a Sex-Biased Human Demography Account for the Reduced Effective Population Size of Chromosome X in Non-Africans?”, Molecular Biology and Evolution 27 (2010): 2312–2321.

31 P. Verdu et al. “Sociocultural Behavior, Sex-Biased Admixture, and Effective Population Sizes in Central African Pygmies and NonPygmies”, Molecular Biology and Evolution 30 (2013): 918–937.

32 S. Mallick et al. “The Simons Genome Diversity Project: 300 Genomes from 142 Diverse Populations”, Nature 538 (2016): 201–206.

33 L. G. Carvajal-Carmona et al. “Strong Amerind/White Sex Bias and a Possible Sephardic Contribution Among the Founders of a Population in Northwest Colombia”, American Journal of Human Genetics 67 (2000): 1287–1295.

34 Bedoya et al. “Admixture Dynamics in Hispanics: A Shift in the Nuclear Genetic Ancestry of a South American Population Isolate”, Proceedings of the National Academy of Sciences of the U.S.A. 103 (2006): 7234–7239.

35 P. Moorjani et al. “Genetic Evidence for Recent Population Mixture in India”, American Journal of Human Genetics 93 (2013): 422–438.

36 M. Bamshad et al. “Genetic Evidence on the Origins of In dian Caste Populations”, Genome Research 11 (2001): 994–1004; D. Reich et al. “Reconstructing Indian Population History”, Nature 461 (2009): 489–944.

37 Bamshad et al. “Genetic Evidence”; I. Thanseem et al. “Genetic Affinities Among the Lower Castes and Tribal Groups of India: Inference from Y Chromosome and Mitochondrial DNA”, BMC Genetics 7 (2006): 42.

38 M. Kayser. “The Human Genetic History of Oceania: Near and Remote Views of Dispersal”, Current Biology 20 (2010): R194–201; P. Skoglund et al. “Genomic Insights into the Peopling of the Southwest Pacific”, Nature 538 (2016): 510–513.

39 F. M. Jordan, R. D. Gray, S. J. Greenhill, R. Mace. “Matrilocal Residence Is Ancestral in Austronesian Societies”, Proceedings of the Royal Society B – Biological Sciences 276 (2009): 1957–1964.

40 Skoglund et al. “Genomic Insights”.

41 I. Lazaridis, D. Reich. “Failure to Replicate a Genetic Signal for Sex Bias in the Steppe Migration into Central Europe”, Proceedings of the National Academy of Sciences of the U.S.A. 114 (2017): E3873–74.



Глава 11. Геномика рас и национальностей

1 Центр профилактики и контроля заболеваний: “Prostate Cancer Rates by Race and Ethnicity”, https://www.cdc.gov/cancer/prostate/ statistics/race.htm

2 N. Patterson et al. “Methods for High-Density Admixture Mapping of Disease Genes”, American Journal of Human Genetics 74 (2004): 979–1000; M. W. Smith et al. “A High-Density Admixture Map for Disease Gene Discovery in African Americans”, American Journal of Human Genetics 74 (2004): 1001–1013.

3 M. L. Freedman et al. “Admixture Mapping Identifies 8q24 as a Prostate Cancer Risk Locus in African-American Men”, Proceedings of the National Academy of Sciences of the U.S.A. 103 (2006): 14068–14073.

4 C. A. Haiman et al. “Multiple Regions within 8q24 Independently Affect Risk for Prostate Cancer”, Nature Genetics 39 (2007): 638–644.

5 Freedman et al. “Admixture Mapping Identifies 8q24”.

6 M. F. Ashley Montagu. Man’s Most Dangerous Myth: The Fallacy of Race (New York: Columbia University Press, 1942).

7 R. C. Lewontin. “The Apportionment of Human Diversity”, Evolutionary Biology 6 (1972): 381–398.

8 J. M. Stevens. “The Feasibility of Government Oversight for NIH-Funded Population Genetics Research”, in Revisiting Race in a Genomic Age (Studies in Medical Anthropology), ed. Barbara A. Koenig, Sandra Soo-Jin Lee, Sarah S. Richardson (New Brunswick, NJ: Rutgers University Press, 2008), 320–341; J. Stevens. “Racial Meanings and Scientific Methods: Policy Changes for NIH-Sponsored Publications Reporting Human Variation”, Journal of Health Policy, Politics and Law 28 (2003): 1033–1087.

9 N. A. Rosenberg et al. “Genetic Structure of Human Populations”, Science 298 (2002): 2381–2385.

10 D. Serre, S. Pääbo. “Evidence for Gradients of Human Gene tic Diversity Within and Among Continents”, Genome Research 14 (2004): 1679–1685; F. B. Livingstone. “On the Non-Existence of Human Races”, Current Anthropology 3 (1962): 279.

11 J. Dreyfuss. “Getting Closer to Our African Origins”, The Root, October 17, 2011, www.theroot.com/getting-closer-to-our-african- origins-1790866394

12 N. A. Rosenberg et al. “Clines, Clusters, and the Effect of Study Design on the Inference of Human Population Structure”, PLoS Genetics 1 (2005): e70.

13 E. G. Burchard et al. “The Importance of Race and Ethnic Background in Biomedical Research and Clinical Practice”, New England Journal of Medicine 348 (2003): 1170–75.

14 J. F. Wilson et al. “Population Genetic Structure of Variable Drug Response”, Nature Genetics 29 (2001): 265–269.

15 D. Fullwiley. “The Biologistical Construction of Race: ‘Admixture’ Technology and the New Genetic Medicine”, Social Studies of Science 38 (2008): 695–735.

16 Lewontin. “The Apportionment of Human Diversity”; A. R. Templeton. “Biological Races in Humans”, Studies in History and Philosophy of Biological and Biomedical Science 44 (2013): 262–271.

17 Razib Khan, www.razib.com/wordpress

18 Dienekes’ Anthropology Blog, dienekes.blogspot.com.

19 Eurogenes Blog, http://eurogenes.blogspot.com.

20 L. Poliakov. The Aryan Myth: A History of Racist and Nationalist Ideas in Europe (New York: Basic Books, 1974).

21 B. Arnold. “The Past as Propaganda: Totalitarian Archaeology in Nazi Germany”, Antiquity 64 (1990): 464–478.

22 J. K. Pritchard, J. K. Pickrell, G. Coop. “The Genetics of Human Adaptation: Hard Sweeps, Soft Sweeps, and Polygenic Adaptation”, Current Biology 20 (2010): R208–15; R. D. Hernandez et al. “Classic Selective Sweeps Were Rare in Recent Human Evolution”, Science 331 (2011): 920–924.

23 M. C. Turchin et al. “Evidence of Widespread Selection on Standing Variation in Europe at Height-Associated SNPs”, Nature Genetics 44 (2012): 1015–1019.

24 Y. Field et al. “Detection of Human Adaptation During the Past 2000 Years”, Science 354 (2016): 760–764.

25 A. Okbay et al. “Genome-Wide Association Study Identifies 74 Loci Associated with Educational Attainment”, Nature 533 (2016): 539–542.

26 В 2016 году Бенджамин с коллегами предложили модель, связывающую определенные генетические аллели и время, потраченное на образование. Используя эту модель, я подсчитал, сколько времени должны потрать на учебу верхние и нижние 5 % индивидов из модельного распределения (то есть те, кто по генетическим вариантам должен получить минимальное образование или, наоборот, максимальное). Для этого я провел следующие расчеты. (1) Время учебы в проанализированной Бенджамином выборке равно 14,3 ± 3,7 лет. Стандартное отклонение 3,7 года получается из указанной в работе величины отклонения в размерности неделя/аллель “0,014 – 0,048 стандартных отклонений на аллель (2,7–9,0 недель учебы)”. Эти значения дают от 188 (=9,0/0,048) до 193 (=2,7 / 0,014) недель. Так как в году 52 недели, отклонение составит 3,7 года. (2) Бенджамин с коллегами указывают, что генетическая модель объясняет 3,2 % дисперсии данного признака (время учебы). Это означает, что корреляция между ожидаемым и реальным значением равна √0,032 = 0,18. Это следует из математических моделей двумерного нормального распределения. (3) У индивида, попавшего по своим генетическим характеристикам в нижние 5 % (то есть более 1,64 стандартных отклонения левее среднего), вероятность проучиться больше 12 лет рассчитывается как число людей в этих 5 % распределения, соотнесенное с числом тех из них, кто учился больше 12 лет (то есть нужно взять площадь под кривой распределения левее 5 % распределения), а затем разделить на 0,05. Это дает вероятность 60 %. Ту же процедуру проделаем с верхними 5 % и получим вероятность 84 %. (4) Бенджамин предположил, что, имея достаточно большую выборку, можно будет с помощью генетических признаков объяснить 20 % дисперсии вместо 3,2 %. Выполним те же расчеты, но только с этими 20 % дисперсии. И получится, что вероятность получить 12-летнее образование и продолжить его у нижних 5 % составит 37 %, а у тех, кто попал в правый 5-процентный хвост распределения, – 96 %.

27 A. Kong et al. “Selection Against Variants in the Genome Associated with Educational Attainment”, Proceedings of the National Academy of Sciences of the U.S.A. 114 (2017): E727–732.

28 Kong et al. “Selection Against Variants”, estimate that the genetically predicted number of years of education has decreased by an estimated 0.1 standard deviations over the last century under the pressure of natural selection.

29 G. Davies et al. “Genome-Wide Association Study of Cognitive Functions and Educational Attainment in UK Biobank (N=112 151)”, Molecular Psychiatry 21 (2016): 758–767; M. T. Lo et al. “GenomeWide Analyses for Personality Traits Identify Six Genomic Loci and Show Correlations with Psychiatric Disorders”, Nature Genetics 49 (2017): 152–156.

30 S. Sniekers et al. “Genome-Wide Association Meta-Analysis of 78,308 Individuals Identifies New Loci and Genes Influencing Human Intelligence”, Nature Genetics 49 (2017): 1107–1112.

31 I. Mathieson et al. “Genome-wide Patterns of Selection in 230 Ancient Eurasians”, Nature 528 (2015): 499–503; Field et al. “Detection of Human Adaptation”.

32 N. A. Rosenberg et al. “Genetic Structure of Human Populations”, Science 298 (2002): 2381–2385.

33 S. Ramachandran et al. “Support from the Relationship of Genetic and Geographic Distance in Human Populations for a Serial Founder Effect Originating in Africa”, Proceedings of the National Academy of Sciences of the U.S.A. 102 (2005): 15942–15947; B. M. Henn, L. L. Cavalli-Sforza, M. W. Feldman. “The Great Human Expansion”, Proceedings of the National Academy of Sciences of the U.S.A. 109 (2012): 17758–17764.

34 J. K. Pickrell, D. Reich. “Toward a New History and Geography of Human Genes Informed by Ancient DNA”, Trends in Genetics 30 (2014): 377–389.

35 M. Raghavan et al. “Upper Palaeolithic Siberian Genome Reveals Dual Ancestry of Native Americans”, Nature (2013): doi: 10.1038/nature 12736.

36 I. Lazaridis et al. “Genomic Insights into the Origin of Farming in the Ancient Near East”, Nature 536 (2016): 419–424.

37 N. Wade. A Troublesome Inheritance: Genes, Race and Human History (New York: Penguin Press, 2014).

38 G. Coop et al. “A Troublesome Inheritance” (letters to the editor), New York Times, August 8, 2014.

39 G. Cochran, J. Hardy, H. Harpending. “Natural History of Ashkenazi Intelligence”, Journal of Biosocial Science 38 (2006): 659–693.

40 P. F. Palamara, T. Lencz, A. Darvasi, I. Pe’er. “Length Distributions of Identity by Descent Reveal Fine-Scale Demographic History”, American Journal of Human Genetics 91 (2012): 809–822; M. Slatkin. “A Population-Genetic Test of Founder Effects and Implications for Ashkenazi Jewish Diseases”, American Journal of Human Genetics 75 (2004): 282–293.

41 H. Harpending. “The Biology of Families and the Future of Civilization” (minute 38), Preserving Western Civilization, 2009 Conference, audio available at www.preservingwesternciv.com/audio/07 % 20Prof._Henry_Harpending – The_Biology_of_Families_and_the_ Future_of_Civilization.mp3 (2009).

42 G. Clark. “Genetically Capitalist? The Malthusian Era, Institutions and the Formation of Modern Preferences” (2007), www. econ.ucdavis.edu/faculty/gclark/papers/Capitalism%20Genes.pdf; G. Clark. A Farewell to Alms: A Brief Economic History of the World (Princeton, NJ: Princeton University Press, 2007).

43 Wade. A Troublesome Inheritance.

44 C. Hunt-Grubbe. “The Elementary DNA of Dr. Watson”, The Sunday Times, October 14, 2017.

45 Coop et al. letters. New York Times.

46 D. Epstein. The Sports Gene: Inside the Science of Extraordinary Athletic Performance (New York: Current, 2013).

47 Там же.

48 Эти расчеты я проделал следующим образом. (1) 99,9999999-й процентиль распределения признака соотносится с 6 стандартными отклонениями от среднего, тогда как 99.99999-й процентиль ограничивается 5,2 стандартного отклонения. Значит, 0,8 стандартного отклонения в сто раз увеличивает число индивидов с данным признаком. (2) Я предположил, что 1,33-кратное увеличение генетической изменчивости в популяциях Черной Африки ассоциировано не только со случайными мутациями, но и с такими, которые формируют те или иные биологические признаки. Следовательно, увеличение стандартного отклонения будет равно √1,33=1,15 (здесь применяется формула из статьи: J. Berg, G. Coop. “A Population Genetic Signal of Polygenic Adaptation”, PLoS Genetics 10 (2014): e1004412). Таким образом, изменчивость, заключенная в пределах 6 стандартного отклонения у неафриканского населения, будет примерно такая же, как заключенная в рамки 5,2 (5,2=6/1,15) стандартного отклонения у африканцев Черной Африки, что и составит как раз то самое стократное увеличение в крайнем правом 0,00000001-м процентиле.

49 W. Haak et al. “Massive Migration from the Steppe Was a Source for Indo-European Languages in Europe”, Nature 522 (2015): 207–211; M. E. Allentoft et al. “Population Genomics of Bronze Age Eurasia”, Nature 522 (2015): 167–172.

50 D. Reich et al. “Reconstructing Indian Population History”, Nature 461 (2009): 489–494; Lazaridis et al. “Genomic Insights”.

51 M. F. Robinson. The Lost White Tribe: Explorers, Scientists, and the Theory That Changed a Continent (New York: Oxford University Press, 2016).

52 A. Haley. Roots: The Saga of an American Family (New York: Doubleday, 1976).

53 “Episode 4: (2010) Know Thyself ” (minute 17) in Faces of America with Henry Louis Gates Jr., http://www.pbs.org/wnet/facesofameri- ca/video/episode-4-know – thyself/237/

54 African Ancestry. “Frequently Asked Questions”, “About the Results”, question 3 (2016), http://www.africanancestry.com/faq/

55 Dreyfuss. “Getting Closer to Our African Origins”.

56 S. Sailer. “African Ancestry Inc. Traces DNA Roots”, United Press International, April 28, 2003, www.upi.com/inc/view.php?StoryID= 20030428-074922-7714r

57 Неопубликованные данные лаборатории Дэвида Райха.

58 H. Schroeder et al. “Genome-Wide Ancestry of 17th-Century Enslaved Africans from the Caribbean”, Proceedings of the National Academy of Sciences of the U.S.A. 112 (2015): 3669–3673.

59 R. E. Green et al. “A Draft Sequence of the Neanderthal Genome”, Science 328 (2010): 710–722.

60 E. Durand, 23andMe: “White Paper 23–05: Neanderthal Ancestry Estimator” (2011), https://23andme.https.internapcdn.net/res/pdf/ hXitekfSJe1lcIy7 – Q72XA_23-05_Neanderthal_Ancestry.pdf; S. Sankararaman et al. “The Genomic Landscape of Neanderthal Ancestry in Present-Day Humans”, Nature 507 (2014): 354–357.

61 Sankararaman et al. “Genomic Landscape”.

62 https://customercare.23andme.com/hc/en-us/articles/212873707Neanderthal-Report-Basics, #13514.



Глава 12. Будущее древней ДНК

1 J. R. Arnold, W. F. Libby. “Age Determinations by Radiocarbon Content – Checks with Samples of Known Age”, Science 110 (1949): 678–680.

2 C. Renfrew. Before Civilization: The Radiocarbon Revolution and Prehistoric Europe (London: Jonathan Cape, 1973).

3 L. R. Binford. In Pursuit of the Past: Decoding the Archaeological Record (Berkeley: University of California Press, 1983).

4 M. Rasmussen et al. “Ancient Human Genome Sequence of an Extinct Palaeo-Eskimo”, Nature 463 (2010): 757–762; M. Rasmussen et al. “The Genome of a Late Pleistocene Human from a Clovis Burial Site in Western Montana”, Nature 506 (2014): 225–229; M. Raghavan et al. “Upper Palaeolithic Siberian Genome Reveals Dual Ancestry of Native Americans”, Nature (2013): doi: 10.1038/nature 12736.

5 P. Skoglund et al. “Genomic Insights into the Peopling of the Southwest Pacific”, Nature 538 (2016): 510–513.

6 J. Dabney et al. “Complete Mitochondrial Genome Sequence of a Middle Pleistocene Cave Bear Reconstructed from Ultrashort DNA Fragments”, Proceedings of the National Academy of Sciences of the U.S.A. 110 (2013): 15758–15763; M. Meyer et al. “A High-Coverage Genome Sequence from an Archaic Denisovan Individual”, Science 338 (2012): 222–226; Q. Fu et al. “DNA Analysis of an Early Modern Human from Tianyuan Cave, China”, Proceedings of the National Academy of Sciences of the U.S.A. 110 (2013): 2223–2227; R. Pinhasi et al. “Optimal Ancient DNA Yields from the Inner Ear Part of the Human Petrous Bone”, PLoS One 10 (2015): e0129102.

7 I. Lazaridis et al. “Genomic Insights into the Origin of Farming in the Ancient Near East”, Nature 536 (2016): 419–424.

8 I. Olalde et al. “The Beaker Phenomenon and the Genomic Transformation of Northwest Europe”, bioRxiv (2017): doi. org/10.1101/135962.

9 P. F. Palamara, T. Lencz, A. Darvasi, I. Pe’er. “Length Distributions of Identity by Descent Reveal Fine-Scale Demographic History”, American Journal of Human Genetics 91 (2012): 809–822; D. J. Lawson, G. Hellenthal, S. Myers, D. Falush. “Inference of population structure using dense haplotype data”, PLoS Genetics 8 (2012): e1002453.

10 S. Leslie et al. “The Fine-Scale Genetic Structure of the British Population”, Nature 519 (2015): 309–314.

11 S. R. Browning and B. L. Browning. “Accurate Non-parametric Estimation of Recent Effective Population Size from Segments of Identity by Descent”, American Journal of Human Genetics 97 (2015): 404–418.

12 M. Lynch. “Rate, Molecular Spectrum, and Consequences of Human Mutation”, Proceedings of the National Academy of Sciences of the U.S.A. 107 (2010): 961–968; A. Kong et al. “Selection Against Variants in the Genome Associated with Educational Attainment”, Proceedings of the National Academy of Sciences of the U.S.A. 114 (2017): E727–732.

13 J. K. Pritchard, J. K. Pickrell, G. Coop. “The Genetics of Human Adaptation: Hard Sweeps, Soft Sweeps, and Polygenic Adaptation”, Current Biology 20 (2010): R208–215.

14 S. Haensch et al. “Distinct Clones of Yersinia pestis Caused the Black Death”, PLoS Pathogens 6 (2010): e1001134; K. I. Bos et al. “A Draft Genome of Yersinia pestis from Victims of the Black Death”, Nature 478 (2011): 506–510.

15 I. Wiechmann, G. Grupe. “Detection of Yersinia pestis DNA in Two Early Medieval Skeletal Finds from Aschheim (Upper Bavaria, 6th Century AD)”, American Journal of Physical Anthropology 126 (2005): 48–55; D. M. Wagner et al. “Yersinia pestis and the Plague of Justinian 541–543 AD: A Genomic Analysis”, Lancet Infectious Diseases 14 (2014): 319–326.

16 S. Rasmussen et al. “Early Divergent Strains of Yersinia pestis in Eurasia 5,000 Years Ago”, Cell 163 (2015): 571–582.

17 P. Singh et al. “Insight into the Evolution and Origin of Leprosy Bacilli from the Genome Sequence of Mycobacterium lepromatosis”, Proceedings of the National Academy of Sciences of the U.S.A. 112 (2015): 4459–4464.

18 K. I. Bos et al. “Pre-Columbian Mycobacterial Genomes Reveal Seals as a Source of New World Human Tuberculosis”, Nature 514 (2014): 494–497.

19 K. Yoshida et al. “The Rise and Fall of the Phytophthora infestans Lineage That Triggered the Irish Potato Famine”, eLife 2 (2013): e00731.

20 C. Warinner et al. “Pathogens and Host Immunity in the Ancient Human Oral Cavity”, Nature Genetics 46 (2014): 336–344.

21. T. Higham et al. “The Timing and Spatiotemporal Patterning of Neanderthal Disappearance”, Nature 512 (2014): 306–309.

22 E. Callaway. “Ancient Genome Delivers ‘Spirit Cave Mummy’ to US Tribe”, Nature 540 (2016): 178–179.

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