Текст книги "Кто мы и как сюда попали. Древняя ДНК и новая наука о человеческом прошлом"

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.