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Evolution: Chu et al Proc. Natl. Acad. Sci. USA 95(1998) 11767 Southern origin outhern Chinese Siberia is doubtful, given the fact that the last glacier started an Indian to recede only 15,000 years ago(see Fig. 2, dashed lines) This conclusion can be tested by using simple inductive logic. If the ancestral Altaic-speaking population was of northern origin, the genetic relationship of extant populations should follow the phylogeny presented in the bottom of Fig. 3. The hylogeny generated in the current study apparently supports the upper phylogeny of Fig 3. In this analysis, Altaic popula- Northern Origin tions are represented by Buryat and Yakut. Southern Chinese Indian ulations are those populations from Yunnan and Taiwan that reportedly did not have any admixture with Altaic pop- batons Southern chinese Populations from Middle Asia were not available to this study Now that we have established that populations in East Asia were subjected to genetic contributions from multiple sources: Southeast asia. Altaic from northeast asia, and mid-Asia or Europe. It would be interesting to estimate relative contribu- FIG 3. Phylogenetic relationships of worldwide populations under tions from each source. Unfortunately, the current study two hypotheses: see text fo involved only mostly minority populations. A study involving populations across the country is necessary to reveal such a urrent analysis showed that northern populations belong to two different groups, although statistical support was still weak. One noticeable difference in our study is the employ We thank the le whose dna was ded by L. L Cavalli- ment in the phylogeny reconstruction of the neighbor-joinin Sforza, J. Kidd, M. Hsu, s.O. Mehdi, and J Bertranpetit Informed nethod, which is supposedly more robust in the presence of consent was obtained for the newly collected Chinese samples.This genetic admixture. The use of microsatellites, a different typ project was completed under the organization of Z Chen and B Q of genetic markers from previous studies, and the measures of Qiang and funded by Ational Natural Sciences Foundation of genetic distance introduced further complication. However China. We also thank P. Watkin and P. Morin from Sequana Ther the northern populations in cluster N2 were sampled from the apeutics, Inc, for their generous support. outhwestern part of China, except for Ewenki, where genetic admixture with the southern population was more likely to 1. Grimes, B F(1996)Ethnologue (Summer Institute of Linguistics, ccur. This might explain why this group of northern popula Dallas), 13th Ed 2. Zhao, T. M, Zhang G, Zhu, Y, Zheng, s, Liu, D, Chen, O.& tions clustered with southern populations Another noticeable feature from this analysis is that the 3. Zhao, T M.& Lee, T. D(1989) Hum. Genet. 83, 101-110. linguistic boundaries are often transgressed across the six Weng, Z, Yuan, Y.& du, R.(1989)Acta Anthropol. Sin. 8, language families studied(Sino-Tibetan, Daic, Hmong-Mien, ustro-Asiatic. Altaic, and Austronesian). Such a phenome 5. Zhang, Z. B ( 1988)Acta Anthropol. Sin. 7, 314-323 non is even more pronounced among southern populations, where populations from the same geographic regions tend to A.(1992)Hum.Bio.64,567-58 cluster in the ph A. M., Ruiz-Linares, A. Tomfohrde. J, Minch, E, ny(see 1B). This observation is Kidd, J. R& Cavalli-Sforza, LL(1994)Nature(London)368, consistent with the history of Chinese populations, where population migrations were substantial 9. Deka. R. Jin. L. Shriver. M. D. Yu. L. M. DeCroo The current analysis suggests that the southern populations Hundrieser. J. Bunker C. H. Ferrell. R.e.& in East Asia may be derived from the populations in Southeast (1995)AmJ.Hm. Genet.56,461-474 Asia that originally migrated from Africa, possibly via mid 10. Jorde, L B, Bamshad, M.J., Watkins, w.S., Zenger, R, Fraley, Asia, and the northern populations were under strong genetic A.E., Krakowiak, P. A, Carpenter, K D, Soodyall, H, Jenkins, influences from Altaic populations from the north. But it is T& Rogers, A. R(1995)Am J. Hum. Genet. 57, 523-538 unclear how Altaic populations migrated to Northeast Asia. It 11. Wang, L(1986)Acta Anthropol. Sin. 5, 24.3-258 is possible that ancestral Altaic populations arrived there from 12. Brooks, A.S.& Wood, B (1990) Nature(London)344, 288-289 13. Li, T.& Etler, D. A(1992) Nature(London)357, 404-40 middle Asia, or alternatively they may have originated from 14. Cann, R. L.(1996)in Prehistoric Mongoloid Dispersals, eds East Asia Akazawa, R& Szathmary, E.J. E(Oxford Univ Press, Oxford The analyses of metric and nonmetric cranial traits of nodern and prehistoric Siberian and Chinese populations M.& Takezaki, N. (1996)Mol. Biol. EvoL. 13, 170-176 showed that Siberians are closer to Northern Chinese and 16. Cavalli-Sforza, L L, Menozzi, P. Piazza, A(1994)The History Mongolian than European(26, 27). The same notion holds for and Geograp/ry of Human Genes(Princeton Univ. Press, Prince the facial flatness(26-28). European populations did not ton,NJ),pp.280-287 appear in Siberia, western Mongolia, and China until the Ruiz-Linares, A(1994)in The Origin and Past of Moder s Viewed from DNA, eds. Brenner, s& Hanihara, cranial and dental analyses have linked the Arctic peoples, 18. Saitou, N& Nei, M(198 37)Mol. Biol. Evol. 4, 406-425 Buryat and east Asians with American Indians(31-35), which 19. Cavalli-Sforza, LL. Edwards, A. w.F(1967)Am J. Hum. arrived through Beringia(Bering land bridge) somewhere between 15,000 and 30,000 years ago(36). These observations 20. Shriver. M. D. Jin. L. Boerwinkle E. Deka. R. FerrelL. R.e. are generally consistent with the genetic evidence based on this Chakraborty, R(1995)Mol. Biol. Evol. 12, 914-920 research and mitochondrial dNA data(37-40). Therefore, it Goldstein. D. B Linares. A. Cavalli-Sforza. L. L. Feldman, M. w.(1995)Genetics 139, 463-471 is more likely that ancestors of Altaic-speaking populations 22. Slatkin, M(1995) Genetics 139, 457 originated from an East Asian population that was originally 23. Goldstein.D. B. Ruiz-Linares. A. Cavalli-Sforza. L L derived from Southeast Asia, although the current Altaic speaking populations undeniably admixed with later arrivers 24. Deka, R, Jin, L, Shriver, M D, Yu, L M, Saha, N Barrantes,R from mid-Asia and Europe(see Fig. 2, thin solid lines). The Chakraborty, R. Ferrell,R. E. (1996)Genome Res. 6, 1177-1184current analysis showed that northern populations belong to two different groups, although statistical support was still weak. One noticeable difference in our study is the employ￾ment in the phylogeny reconstruction of the neighbor-joining method, which is supposedly more robust in the presence of genetic admixture. The use of microsatellites, a different type of genetic markers from previous studies, and the measures of genetic distance introduced further complication. However, the northern populations in cluster N2 were sampled from the southwestern part of China, except for Ewenki, where genetic admixture with the southern population was more likely to occur. This might explain why this group of northern popula￾tions clustered with southern populations. Another noticeable feature from this analysis is that the linguistic boundaries are often transgressed across the six language families studied (Sino-Tibetan, Daic, Hmong-Mien, Austro-Asiatic, Altaic, and Austronesian). Such a phenome￾non is even more pronounced among southern populations, where populations from the same geographic regions tend to cluster in the phylogeny (see Fig. 1B). This observation is consistent with the history of Chinese populations, where population migrations were substantial. The current analysis suggests that the southern populations in East Asia may be derived from the populations in Southeast Asia that originally migrated from Africa, possibly via mid￾Asia, and the northern populations were under strong genetic influences from Altaic populations from the north. But it is unclear how Altaic populations migrated to Northeast Asia. It is possible that ancestral Altaic populations arrived there from middle Asia, or alternatively they may have originated from East Asia. The analyses of metric and nonmetric cranial traits of modern and prehistoric Siberian and Chinese populations showed that Siberians are closer to Northern Chinese and Mongolian than European (26, 27). The same notion holds for the facial flatness (26–28). European populations did not appear in Siberia, western Mongolia, and China until the Neolithic and Bronze Age (26, 27, 29, 30). Furthermore, cranial and dental analyses have linked the Arctic peoples, Buryat and east Asians with American Indians (31–35), which arrived through Beringia (Bering land bridge) somewhere between 15,000 and 30,000 years ago (36). These observations are generally consistent with the genetic evidence based on this research and mitochondrial DNA data (37–40). Therefore, it is more likely that ancestors of Altaic-speaking populations originated from an East Asian population that was originally derived from Southeast Asia, although the current Altaic￾speaking populations undeniably admixed with later arrivers from mid-Asia and Europe (see Fig. 2, thin solid lines). The possibility of early northern route migration from mid-Asia to Siberia is doubtful, given the fact that the last glacier started to recede only 15,000 years ago (see Fig. 2, dashed lines). This conclusion can be tested by using simple inductive logic. If the ancestral Altaic-speaking population was of northern origin, the genetic relationship of extant populations should follow the phylogeny presented in the bottom of Fig. 3. The phylogeny generated in the current study apparently supports the upper phylogeny of Fig. 3. In this analysis, Altaic popula￾tions are represented by Buryat and Yakut. Southern Chinese populations are those populations from Yunnan and Taiwan that reportedly did not have any admixture with Altaic pop￾ulations. Populations from Middle Asia were not available to this study. Now that we have established that populations in East Asia were subjected to genetic contributions from multiple sources: Southeast Asia, Altaic from northeast Asia, and mid-Asia or Europe. It would be interesting to estimate relative contribu￾tions from each source. Unfortunately, the current study involved only mostly minority populations. A study involving populations across the country is necessary to reveal such a picture. We thank the people whose DNA was provided by L. L. Cavalli￾Sforza, J. Kidd, M. Hsu, S. Q. Mehdi, and J. Bertranpetit. Informed consent was obtained for the newly collected Chinese samples. This project was completed under the organization of Z. Chen and B. Q. Qiang and funded by the National Natural Sciences Foundation of China. We also thank P. Watkin and P. Morin from Sequana Ther￾apeutics, Inc., for their generous support. 1. Grimes, B. F. (1996) Ethnologue (Summer Institute of Linguistics, Dallas), 13th Ed. 2. Zhao, T. M., Zhang, G., Zhu, Y., Zheng, S., Liu, D., Chen, Q. & Zhang, X. (1986) Acta Anthropol. Sin. 6, 1–8. 3. Zhao, T. M. & Lee, T. D. (1989) Hum. Genet. 83, 101–110. 4. Weng, Z., Yuan, Y. & Du, R. (1989) Acta Anthropol. Sin. 8, 261–268. 5. Zhang, Z. B. (1988) Acta Anthropol. Sin. 7, 314–323. 6. Zhang, H. (1988) Acta Anthropol. Sin. 7, 39–45. 7. Etler, D. A. (1992) Hum. Biol. 64, 567–585. 8. Bowcock, A. M., Ruiz-Linares, A., Tomfohrde, J., Minch, E., Kidd, J. R. & Cavalli-Sforza, L. L. (1994) Nature (London) 368, 455–457. 9. Deka, R., Jin, L., Shriver, M. D., Yu, L. M., DeCroo, S., Hundrieser, J., Bunker, C. H., Ferrell, R. E. & Chakraborty, R. (1995) Am. J. Hum. Genet. 56, 461–474. 10. Jorde, L. B., Bamshad, M. J., Watkins, W. S., Zenger, R., Fraley, A. E., Krakowiak, P. A., Carpenter, K. D., Soodyall, H., Jenkins, T. & Rogers, A. R. (1995) Am. J. Hum. Genet. 57, 523–538. 11. Wang, L. (1986) Acta Anthropol. Sin. 5, 243–258. 12. Brooks, A. S. & Wood, B. (1990) Nature (London) 344, 288–289. 13. Li, T. & Etler, D. A. (1992) Nature (London) 357, 404–407. 14. Cann, R. L. (1996) in Prehistoric Mongoloid Dispersals, eds. Akazawa, R. & Szathmary, E. J. E. (Oxford Univ. Press, Oxford), pp. 41–51. 15. Nei, M. & Takezaki, N. (1996) Mol. Biol. Evol. 13, 170–176. 16. Cavalli-Sforza, L. L., Menozzi, P. & Piazza, A. (1994) The History and Geography of Human Genes (Princeton Univ. Press, Prince￾ton, NJ), pp. 280–287. 17. Ruiz-Linares, A. (1994) in The Origin and Past of Modern Humans as Viewed from DNA, eds. Brenner, S. & Hanihara, K. (World Scientific, Singapore), pp. 123–148. 18. Saitou, N. & Nei, M. (1987) Mol. Biol. Evol. 4, 406–425. 19. Cavalli-Sforza, L. L. & Edwards, A. W. F. (1967) Am. J. Hum. Genet. 19, 233–243. 20. Shriver, M. D., Jin, L., Boerwinkle, E., Deka, R., Ferrell, R. E. & Chakraborty, R. (1995) Mol. Biol. Evol. 12, 914–920. 21. Goldstein, D. B., Ruiz-Linares, A., Cavalli-Sforza, L. L. & Feldman, M. W. (1995) Genetics 139, 463–471. 22. Slatkin, M. (1995) Genetics 139, 457–462. 23. Goldstein, D. B., Ruiz-Linares, A., Cavalli-Sforza, L. L. & Feldman, M. W. (1995) Proc. Natl. Acad. Sci. USA 92, 6723–6727. 24. Deka, R., Jin, L., Shriver, M. D., Yu, L. M., Saha, N., Barrantes, R., Chakraborty, R. & Ferrell, R. E. (1996) Genome Res. 6, 1177–1184. FIG. 3. Phylogenetic relationships of worldwide populations under two hypotheses; see text for discussion. Evolution: Chu et al. Proc. Natl. Acad. Sci. USA 95 (1998) 11767
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