Abstract
Mitochondrial deoxyribonucleic acid (DNA) from 201 unrelated Mongolian individuals in the three different regions was analyzed. The Mongolians took the dominant East Asian-specific haplogroups, and some European-prevalent haplogroups were detected. The East Asians-specific haplogroups distributed from east to west in decreasing frequencies, and the European-specific haplogroups distributed conversely. These genetic data suggest that the Mongolian empire played an important role in the maternal genetic admixture across Mongolians and even Central Asian populations, whereas the Silk Road might have contributed little in the admixture between the East Asians and the Europeans.
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Introduction
The phylogenetic analysis of mitochondrial deoxyribonucleic acid (DNA) (mtDNA) have revealed a regional clustering of continent habitation (Herrnstadt et al. 2002) and revealed that humans had a recent African origin and migrated through the Eurasian continent (Cann et al. 1987). The mtDNA distribution of the populations who lived in the intermediate region is crucial for elucidating migrations and genetic admixture. Central Asia is located between East Asia and Europe, and there is debate regarding the origin of East Asians (Shi et al. 2005). The genetic distribution of the East Asian and European populations has specific features, such as the distribution of the mtDNA: A, B, D, G, C, F, and M7 are the main mtDNA haplogroups of East Asian populations (Ballinger et al. 1992; Yao et al. 2002), and HV, TJ, IK, U, W, and X are the European-specific haplogroups (Torroni et al. 1996). The mtDNA data from East Asians (Kivisild et al. 2002; Kong et al. 2003b) and Europeans (Ingman et al. 2000) could be a effective tool to reconstruct the origin and migration in the populations of the two continents. There was extensive genetic admixture of eastern Asians and western Eurasians in Central Asia where the Silk Road crossed between the two areas (David et al. 1998; Comas et al. 1998). Xinjiang Province in northwestern China is an important part of the Silk Road, where extensive genetic admixture was observed in local populations (Yao et al. 2004). European-specific mtDNA haplogroups were identified in all five major Xinjiang populations, and 14% of these haplogroups were found in the Xinjiang Mongolians but only 2% in the Mongolians who lived in the east part of plateau of Mongolia (Kong et al. 2003b). European-specific mtDNA haplogroups were also identified in the populations of northeast Mongolia far away the Silk Road (Kolman et al. 1996). The genetic characters of the Mongolians were important data to elucidate the origin and the late admixture history of the populations in the plateau of Mongolia.
The Mongolian empire had a great impact on the history of Europe and Central Asia and could represent an important role in the genetic admixture. An increasing frequency of European-specific haplogroups in the upper three regional Mongolian populations from northeastern China to Central Asia has been observed. However, no or fewer European-specific haplogroups have been found in the Han and Tibetan populations, who also live along the Silk Road (Yao et al. 2002; Qian et al. 2001). There are now nearly 8.2 million Mongolians living primarily in Mongolia, Inner Mongolia of China, and the Siberia region of Russia. Most Mongolians live outside the region of the Silk Road.
There is a Mongolian population living in Tonghai County of Yunnan Province in southwestern China. Ancestors of Tonghai Mongolians migrating to extant inhabitation at the beginning of the Mongolian empire would have carried the original genetic components of Mongolians (Du et al. 1993). These Mongolians then formed a relatively independent Mongolian group (You 1997). In this study, mtDNA haplotypes from 201 Mongolian samples were analyzed. Phylogenetic relationships among the Mongolian populations from the central plateau of Mongolia (Inner Mongolians) and Tonghai County were revealed. The genetic impact of the Mongolian empire was detected by comparing the original Mongolians with populations in the regions where the empire had explored.
Materials and methods
Sampling
A total 201 unrelated Mongolians individuals were analyzed: 155 from central Inner Mongolia of China (107 from Hohhot City and 48 from Zhenglanqi County), and 46 from Tonghai County in Yunnan Province of China. All individuals were able to confirm their maternal origin and gave informed consent.
Previously published Mongolian mtDNA data was used for comparison. These data include 103 mtDNA from Mongolia with HVS-I and eight additional coding-region information (Kolman et al. 1996); 50 mtDNA from Xingjing Province with HVS-I and 9-bp-deletion data (Yao et al. 2004); 48 mtDNA from Xin Barage Zuoqi County in the eastern part of Inner Mongolia of China with HVS-I and HVS-II, and an additional one or two coding-regions (Kong et al. 2003b) (Fig. 1).
Genotyping of mtDNA polymorphisms
Genomic DNA was extracted by standard phenol/chloroform methods. The control region 15926-619 and four parts of the coding regions containing 2796-5441, 5126-7701, 8215-10735, and 14054-16048 were amplified using five pairs of primers, L15926/H619, L2796/H5441, L5126/H7701, L8215/H10735, and L14054/H16048, respectively. Polymerase chain reaction (PCR) was performed in a 15-μl reaction mixture that contained 20 ng DNA, reaction buffer [10 mM Tris hydrochloride (Tris-HCl), pH 8.3, 1.5 mM magnesium chloride (MgCl2), 50 mM potassium chloride (KCl), 0.001% gelatin, and 0.5 μl 1 mM of each primer, 200 μM deoxynucleotide triphosphate (dNTPs), and 0.2 U Taq polymerase (Promega). The PCR amplification was performed in either a PE-9600 or PE-9700 (ABI Applied Biosystems, Fostor City, CA, USA) with 95°C for 2 min followed by 30 cycles of 95°C for 30 s, 58°C for 30 s, 72°C for 30 s, then 72°C 2 min. Each PCR product was diluted by adding 60 μl of water. Using upper PCR product as a template, the control region and parts of the coding regions containing 3010-3430, 4640-5204, 10170-10660, and 14478-15086 were amplified and sequenced, respectively, by six pairs of nested primers: L15996/H107, L29/H590, L3010/H3430, L4640/H5204, L10170/H10660, and L14478/H15086. Another six pairs of confirming primers, L3953/H4508, L5126/H5441, L8215/H8297, L9766/H1063, L14989/H15400, and L15391/H16048, were used when needed to unambiguously define a sample. All the primer sequences were cited from Yao et al. (2002) or designed by the primer 3 software (http://www-genome.wi.mit.edu/genome_software/other/primer3.html) (Table 1).
The nested or second PCR products were examined on a 2% agarose gel then purified with the Wizard Purification Kit (Promega, Madison, WI, USA). Each fragment was sequenced for both strands with the BigDye and Dye Terminator Cycle Sequence Kit (ABI Applied Biosystems), run on an ABI 377 DNA sequencer (ABI Applied Biosystems), and scored using Sequence Analysis and Sequence Navigator software (Perkin–Elmer).
The sequences were analyzed by the DNA-STAR software (DNASTAR, Inc.) and compared with the Revised Cambridge Reference Sequence (rCRS) (Andrews et al. 1999). We also rechecked all polymorphisms in the Chromas 1.45 software (Griffith University, Australia). We adopted the established phylogeny of Europeans and East Asians proposed by Finnilä et al. (2001), Kivisild et al. (2002), and Kong et al. (2003a). To show genetic relationships among Mongolians in different regions, principal component analysis (PC) was conducted by haplogroup frequency using SPSS12.0 software (SPSS Inc.). Arlequin software (http://anthro.unigo.ch/arlequin) was used to calculate neutrality tests of Tonghai Mongolians (Tajima 1996) and genetic distance (F ST values) of regional Mongolians and other East Asian populations (Schneider et al. 2000). Neighbor-joining (NJ) trees based on F ST values were constructed in MEGA software (http://www.megasoftware.net) (Nei 1987; Kumar et al. 2001).
Result
Among all 201 northern and southern Mongolian samples, 111 samples had the 489T → C, 10400 C → T, 14783 T → C, and 15043 G → A mutations attributed to macrohaplogroup M. These samples were further subcladed to haplogroups D, G, M*, and M7–M11 according to the information in the D-loop region and encoding region. The other 90 samples did not have the above-mentioned four mutations and belonged haplogroups A, N9, F, R9b, R11, HV, K, J, or I, which belong to macrohaplogroup N.
Four parallel mutation sites were found. Deletions were detected at 16227 in four G2a haplogroup samples, 16224 in all five Z haplogroup samples, 16316 in one M9a haplogroup sample, and no 73 G → A were observed in two H haplogroup samples. Table 2 shows the distribution of haplogroup frequencies in norther Inner Mongolians and southern Tonghai Mongolians. Based on M and N macrohaplogroups, 23 haplogroups and 52 subhaplotypes were defined, including nearly all East-Asian-specific haplogroups, HV, J, K, U, and I, European-specific haplogroups (Finnilä et al. 2001), and the classification tree of the mtDNA haplogroups in Mongolians shown in Fig. 2. For Inner Mongolians, there were 39, 18, and 13 individuals belonging to haplotypes D, A, and C, respectively. The D haplotype had the highest frequency (25%), followed by A (12%) and C (9%). The combined frequency of these three haplogroups was more than 46%. F, B, and M7 haplogroups were also present, with relatively high frequencies of F (13.5%), B (7.7%), and M (76.5%), respectively. The European-specific haplogroups K, J, and I were first found in Mongolians. The combined frequencies of European-specific haplogroups K, J, I, H, and U) was 8.4%. In contrast, no European-specific haplogroup was found in Tonghai Mongolians; all 46 individuals were assigned into the East-Asian-specific haplogroups. We found 22%, 17%, and 9% of individuals belonged to haplogroups A, D, and C, respectively. The combined frequency of these three haplogroups was nearly 48%. Haplogroup R9b had the third highest frequency (13%) in Tonghai Mongolians (Table 2). There was no significant difference in the mtDNA haplogroup frequency distribution between Mongolians from Tonghai and those from central Inner Mongolia of China. However, Fisher’s exact test showed that the mtDNA haplogroup frequency distribution between Mongolians from Tonghai was significantly different from that of Mongolians in Mongolia (Kolman et al. 1996), in Xin Barage Zuoqi in the eastern part of Inner Mongolia of China (Kong et al. 2003b), and in Xingjiang Province in the western part of China (Yao et al. 2004) (Table 2).
The basal mtDNA haplogroup profiles of the five Mongolian populations were treated as input vectors for PC analyses. Figure 3a displays the PC map for the first two principal components, of which the first principal component accounts for 79% of the total variation. Southern Tonghai Mongolians and the other four northern Mongolians are separated by PC2 (accounting for 10% of the total variation). Figure 3b presents PC results conducted in regional Mongolians and other East- and Central-Asian populations. Northern East Asians, southern East Asians, and Central Asians are clearly separated by PC2 (accounting for 21.4% of the total variation). Tonghain, Inner Mongolian, and Mongolian samples fall entirely into the northern East Asian group; Xinjing and eastern Inner Mongolia samples cluster together.
The P value of neutrality tests in Tonghai Mongolians was 0.109. The haplotype diversity was 0.9428 in Tonghai Mongolians, 0.9426 in Mongolians from central Inner Mongolia of China, 0.9087 in Mongolians from Mongolia (Kolman et al. 1996), 0.8580 in Mongolians from Xin Barage Zuoqi, eastern part of Inner Mongolia of China (Kong et al. 2003b), and 0.9132 in Mongolians from Xingjiang Province, western part of China (Yao et al. 2004) (Table 2). An unrooted NJ tree was constructed among regional Mongolians and East Asian populations using F ST distances. Since Kolman et al. (1996) only named A, B, C, and D haplogroups in Mongolian samples, we thus excluded them from the phylogenetic analysis. Fig. 4, is largely divided into two different clusters, and all four regional Mongolians grouped with the northern East Asian (NEA) cluster (Fig. 4).
Discussion
The Mongolians have four dominant haplogroups (A, D, G, and M8), including those from the regions of the Mongolia plateau or adjacent region (Yao et al. 2004), central Inner Mongolian and Monglia (Kolman et al. 1996) and eastern Xin Barage Zuoqi (Kong et al. 2003b). Those haplogroups are prevalent in the NEAs, and the prevalent southern East Asians (SEAs) haplogroups are B, M7, R9, and N9a. The European-prevalent haplogroups are HV, U, K, I, and J (Kivisild et al. 2002; Yao et al. 2002; Kong et al. 2003b). The main maternal components of these Mongolians belong to NEA-specific haplogroups A, D, G, and M8, which occupy more than 54% in eastern Xin Barage Zuoqi County samples, nearly 45% in central Inner Mongolian and Mongolian samples, and 40% in western Xingjang Mongolian samples, showing decreasing frequencies from east to west. SEA-prevalent haplogroup is nearly 28% in Inner Mongolians, 22% in eastern Xin Barage Zuoqi County, and Mongolian samples and only 16% in Xingjing Mongolians. The European-prevalent haplogroups (HV, U, K, I, J) are 14% in western Xingjang Mongolian, 10% in Monglia, 8.4% in central Inner Mongolian samples, and only 2% in eastern Xin Barage Zuoqi County samples, showing decreasing frequencies from west to east. The NEA-specific haplogroups distributed in reverse to the European-prevalent haplogroups. The European-specific haplogroups H and U are prevalent in Mongolians, and K, J, and I haplogroups are first detected in Inner Mongolians. Mongolian genetics had a northern East-Asian orientation and admixed with the Southern East Asians. The genetic studies indicate the East Asians had single south origin, and there the early human migration through central East after modern humans came out of Africa did not occur (Shi et al. 2005). European-specific haplogroups K, J, and I indicate a late Mongolian admixture with Europeans. The Mongolian plateau lies in the heart of east Central Asia, where modern humans inhabited 20,000–25,000 years ago (Fiedel et al. 1992). Mongolians living in the Mongolia plateau were separated from Europe by Central Asia, so there must have been certain factors promoting the genetic admixture.
Mitochondrial DNA data indicated that northern East Asians were the ancestors of the Native Americans who migrated about 7,000–9,000 years ago through the Bering strait (http://www.mitomap.org/). Mitochondrial DNA analysis of the 18 Asian and 16 American groups indicate that the Mongolians were the ancestor of Native Americans, and whose mtDNA haplogroups are A, B, C, and D (Kolman et al. 1996). All studies suggested that the Mongolians did not contain the European lineage 7,000–9,000 years ago.
Mongolians migrated south to TongHai in Yunnan Province about 800 years ago (Du et al. 1993). Historically, the Mongolian army was accompanied by their wives at the beginning of the war during the Mongolian empire, and later some of them married local people (You 1997; Hu 2002). Comparison of the 14 different populations in several regions of East Asia (Yao and Zhang 2002; Yao et al. 2002, 2004; Kong et al. 2003b; Qian et al. 2001) revealed that TongHai Mongolians are close to northern Mongolians genetically and belong to the NEAs (Figs. 3b, 4). Our recent study indicated that all Mongolians clustered together in Y-STR data, including the TongHai Mongolians (our unpublished data). The extant TongHai Mongolians take the north-prevalent haplogroups (51.5%), and no European-specific haplotypes were observed. The neutrality tests showed no selective effect (P ≥ 0.05), and there was no population expansion and bottleneck. The study also indicated that Mongolians who migrated south did not contain European-specific mtDNA haplotypes and in Native Americans—the descendants of original Mongolians—the same pattern appeared (Kolman et al. 1996).
The European haplogroups were detected in Mongolians who inhabited Central Asia, such as Xingjing Autonomous region, and central Inner Mongolian. The Silk Road (second century bc to the thirteenth century ad) would have brought the European-specific genetics into those regions. However, few European-specific haplogroups have been found in Han and Tibetan populations who lived along the Silk Road (Yao et al. 2002; Qian et al. 2001), which supports the hypothesis that the Silk Road contributed little genetic admixture between the Europeans and East Asians. In history, the Parthian empire prevented any mass migration of strange people, usually only allowing caravans of traders to pass, so population migration freely between Europe and East Asia could not occur (Dubs 1957). Thus, the genetic admixture of Europeans with northern East Asians in Central Asian can only be explained by the later conquer of the Mongolian empire (1271–1368 ad). The Mongolian populations played an important role in promoting the historical development of Eurasia. Genghis Khan (1162–1227) and his descendants waged three wars in the west and established the united multiethnic Mongolian empire crossing Eurasia, which had 3.5 million km2 and a 100 million people. The Mongolians continued to dominate the region north of the Great Wall of China and west to the Aral Sea for two more centuries after the Mongolian empire ended in 1368. Then, some Mongolian descendants moved back to their hometowns from thousands of kilometers away. Those who migrated back carried some exotic genetic components with them. During their wars, the Mongolian conquerers incorporated some ethnic populations into their groups, and those admixtures expanded the Mongolian gene pool. The European-specific mtDNA haplogroups could be detected in different northern Mongolian populations.
Y-chromosome studies support the fact that Mongolians have kept East-Asian characteristics (Nasidze et al. 2005; Wells et al. 2001; Xue et al. 2006; Yang et al. 2005). The prevalence of the Mongolian Y chromosome in Central Asians support the inequity admixture due to Genghis Khan’s military success (Zerjal et al. 2002, 2003), that is, the Mongolia conqueror contributed the dominant Y chromosome to the descendant admixture populations. This happened in the Indo-Europeans admixture in the caste establishment in India (Bamshad et al. 2001; Thanseem et al. 2006), and similar situations were found among Spanish Conquistadors and British colonialists in North America and Australia (Carvajal-Carmonn et al. 2000). The Y-chromosome data support Mongolian impact in the genetics of Central and northern East Asians, however, Mongolian impact from maternal lineage was not significant (Bamshad et al. 2001; Thanseem et al. 2006).
In summary, the dominant Mongolian mtDNA gene pool belongs to the eastern Asian lineage, which agreed with its origin. The unbalanced genetic contribution of the paternal and maternal in the admixture populations during conquest by the Mongolian empire indicate the developmental history of Mongolian populations.
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Acknowledgments
We thank Junping Xin and the reviewers for their critical comments on the manuscript. We thank all of the DNA donors for making this work possible. This work was financially supported by the National Nature Science Foundation of China (No. 30660076) and the 973 Prophase Project of the Science and Technology Ministry of China and the Science and Technology Bureau of Yunnan Province (No. 2006CB708502, N0. 2006GP10).
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Cheng, B., Tang, W., He, L. et al. Genetic imprint of the Mongol: signal from phylogeographic analysis of mitochondrial DNA. J Hum Genet 53, 905–913 (2008). https://doi.org/10.1007/s10038-008-0325-8
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DOI: https://doi.org/10.1007/s10038-008-0325-8