Table 6. Lower triangular matrix based on F_ST GD, upper triangular matrix - Nei's GD.

Noteworthy, the Exact test revealed that there are not any statistically significant differences between the Armenian groups, which is in contrast with previous studies [Weale et al., 2001, Yepiskoposyan et al., 2001] performed on Armenians using Y-chromosomal markers. The most plausible explanation of this discrepancy is the social practice of patrilocality in the Armenian communities. Patrilocality, or a higher rate of women's spatial dispersal, is a widely spread cultural tradition for human societies. This practice is being used to explain the following patterns detected in ethnically different populations: relatively low Y chromosome and high mtDNA variability within populations, large between-group genetic distances for the male chromosome and small distances for the mtDNA [Seielstad et al., 1998; Perez-Lezaun et al., 1999; Jorde et al., 2000; Wilson et al., 2001]. It is usually explained by higher rate of female versus male migration which takes place when women move to their husbands' residence after marriage. Traditionally, the Armenian society is patrilocal, and the deviation from this cultural practice is very rare and only due to specific circumstances [Redgate, 2000].

In order to determine the possible gene flow between Armenians and neighboring groups, the populations from Levant, the Caucasus and Europe were also included in Exact test. The results demonstrate that EA group does not differ from Syrians, Georgians and Chechens, that might indicate gene flows between these groups, which took place on Ararat Valley - the active ancient trade route. However, these pattern of genetic homogeneity ends in north-western part of the Caucasus, which might be illustrated by the significant difference between all the Armenian groups and the Adyghs. The absence of differentiation is also found between Armenians from western and central parts of the Armenian highland and Syrians, and between CA and Georgians, which could be explained by geographic proximity of these groups. Europeans, presented in this analysis by population from France, significantly differ from all considered groups.

Table 7. The values of the exact test of population differentiation of Armenians and comparative data sets. In lower triangular matrix “-” indicates the absence of significant difference (p>0.05), “+” - the presence, upper triangular matrix - p-values, with SD in parentheses.

To further elucidate the structure of mtDNA gene pool of Armenians among neighboring groups, we have performed the Analysis of Molecular Variance (AMOVA), where we have grouped populations according to their geographic locations: CA, EA, WA and SK as the groups of the Armenian Highland; Syria, Lebanon and Jordan as populations from Levant, and Chechens, Ingushes and Dargins as the ethnic groups from the Caucasus. The AMOVA results are shown in Table 8.

The greatest part of variation is found within populations (98.6 %), whilst among groups the percentage of variation is negligible (0.91%), which indicates a very high level of homogeneity (based on matrilineal gene pool) of the populations of studied regional groups.

Table 8. The results of AMOVA analysis conducted for populations from three regional groups - Armenian Highland, Levant and the Caucasus.

We have also assessed the genetic relationships between Armenian groups and some other populations using the principal coordinate analysis based on Nei's GD.

Figure 11. Principal coordinates plots (first two axes) based on Nei's GD values calculated from mtDNA haplogroup frequency data. Numbers in brackets show the percentage of total variation explained by each axis.

However, the PC plot did not show any clear clustering of the studied groups. Nevertheless, four Armenian populations and Syrian, representing Levant, plotted relatively close to each other. It should be mentioned, that PC1 axis describes the majority of variation (ca. 53%), in contrast with PC2, that includes only 24 % of the total variation. Thus, the genetic proximity of studied populations should be inferred predominantly by PC1 axis. To visualize the pattern of genetic affinity of Armenians and neighboring populations in more accurate way, the phylogenetic trees based on F_ST and Reinolds's GDs using UPGMA method were built (Fig. 12). Both trees evidently unify Armenian populations in a single cluster, with the population of Levant being the closest neighbor.

Figure 12. The UPGMA tree of studied population based on F_ST (left) and Reinolds's (right) GDs.

This result is in a good acordance with those of other analyses performed in our study, as well as with the results obtained for general Armenian population.

SUMMARY

The investigation of historical and demographic events that characterize modern human populations is crucial for the reconstruction of human evolutionary history. Novel approaches based on the advances in molecular genetics alongside with traditional methods of investigation of genetic structure of human populations might shed light on fundamental issues of the spread and dispersal of Homo sapiens. Rapidly accumulating data on human genetic variation of three widely used types of DNA markers, i.e. of non-recombining region of Y-chromosome, mtDNA and autosomal markers have been extensively used for the reconstruction of genetic history of modern humans.

One of the most interesting regions in terms of population genetics is the Middle East, which served as a key crossroad for human disseminations. Since ancient times Armenians inhabited the territory of the Armenian Highland, which occupies the central positions on the Middle East landscape. Numerous historical records and studies show that this region was an important ancient migratory pathway for different ethnic groups, which undoubtedly had an influence on the genetic structure of the Armenian population. Hence, the study of the Armenian gene pool is an important issue to understand the genetic impact of ancient migrations in this region. Today, there are a limited number of genetic studies of Armenians. Most of them were conducted mainly using Y-chromosomal markers, which reveal only patrilineal portion of genetic structure. However, the matrilineal component of the Armenian genetic history based on mitochondrial DNA (mtDNA) markers is still poorly investigated. Summarizing all the studies of Armenians performed using mtDNA markers so far, it should be mentioned that Armenian matrilineal genetic portion was described just in the context of investigations about other Middle Eastern and Caucasian populations. Moreover, all studies that were carried out with the inclusion of Armenians didn't contain the appropriate representative datasets and the origin of these individuals was not clearly determined. These data do not allow making any precise conclusions and inferences on the Armenian matrilineal genetic structure. Considering above mentioned, the main aim of the work was to reconstruct the matrilineal genetic history of Armenians and identify the location of the Armenian population on the genetic landscape of the Middle East using mitochondrial genetic markers, which directly reflect the maternal component of the population's gene pool.

In order to elucidate the questions posed, we have analyzed the unpublished mtDNA HVRI typing data of relatively large dataset of 400 ethnic Armenian individuals, whose maternal ancestors inhabited different geographic areas of the Armenian Highland. Informed consent and the information about birthplace of parents and grandparents were gained from all the individuals. This dataset was considered as a general Armenian population. To determine further the genetic structure of Armenians, we have divided the general Armenian population into three geographically distinct groups, according to the place of origin of individual's maternal ancestor. After the division we have got following subpopulations: East Armenians (n=61), West Armenians (n=44), and Central Armenians (n=85). Afterwards, to this dataset we have also added corresponding unpublished data of two hundred Armenian subjects from the region Salmast/Khoy. The data of Armenians was compared with those of other population, representing five large geographic areas - Europe, the Caucasus, the Middle East, the East Asia and the North Africa.

The investigation of the Armenian matrilineal gene pool in our study was made in two steps. First, we have analyzed the genetic features of the general Armenian population and compared the results with those of other populations. Second, we have performed an inter-population analysis of Armenians from different geographic groups, and juxtaposed the obtained result with those of some neighboring ethnic groups.

Our results have revealed that the mtDNA haplogroup content of general Armenian population is composed mainly by four haplogroups H, U, J and D (ca. 70% in aggregate), which are widely distributed today in Europeans. On the other hand, the Asian genetic component was presented in quite restricted amount (A, B, C, etc., ca. 4%), while found in significant concentrations in some neighboring groups. Additionally, the frequency of African macro-haplogroup L did not reach 1% in Armenians.

The analysis of genetic diversity has revealed 244 different haplotypes in general Armenian population (n=400) (H=0.988, the second highest value in the region after the Levant groups) and 126 segregating sites with total number of 140 mutations. Highly negative value of Tajima's D parameter might indicate a recent population expansion or the presence of purifying selection of some mtDNA HVRI haplotypes in Armenians.

Further, to determine the location of Armenians on the genetic landscape of the region, we have performed the Principal Coordinates analysis and constructed the UPGMA tree based on different genetic distances. Despite the minor differences in the location of some population in both PCA plot and phylogenetic tree, which are due to the application of several genetic distances, it was clearly demonstrated that Armenians have the closest links with the Levantine ethnic groups. This finding indicates the high genetic proximity between matrilineal genetic structures of Levant populations and Armenians.

In order to elucidate the intra-population diversity of the Armenian matrilineal gene pool, we have analyzed the mtDNA HVRI data of individual, representing four regional groups of the Armenian plateau, i.e. Western, Central, Eastern and Salmast/ Khoy.

The analysis of haplogroup distribution has demonstrated that, as it was observed in general population, the majority (~70%) of mtDNA haplogroups in these sub-populations is represented by four haplogroups H, U, J and T, with haplogroup H being a modal one in all the groups. The Asian haplogroups are absent from the mtDNA gene pool of all the Armenian groups studied. Only in the group of Salmast/ Khoy the haplogroups C and D reach 2% of frequency each. Interestingly, the African haplogroup L is found only in Central Armenians with 1% of frequency.

The summary statistics have revealed that Armenian groups have very high rate of gene diversity H, which reaches 0.99 in average, with the highest value observed in the group of Central Armenians (H=0.995), making them one of the most diverse population in the Middle East. This pattern of genetic diversity corroborates with the results obtained by the Median-Joining network analysis. The MJ networks of four major haplogroups widely distributed in Armenians (i.e. H, U, J and T) were the most diverse in the Central Armenians. Additionally, the “star”-like topology of the H haplogroup network alongside with the strongly negative values of Tajima's D obtained for all the Armenian subpopulations might indicate the recent population expansions or the presence of negative selection.

Further, we aimed to assess the genetic relatedness of four Armenian groups observed. For this purpose, we have carried out the Exact test of population differentiation, where some other neighboring groups were also included. The results clearly demonstrated that there was no any significant differentiation (p>0.05) between the Armenian groups. However, previous studies on the Armenian Y-chromosomal genetic structure have nevertheless revealed a significant geographic stratification of the Armenian patrilineal genetic pool. This incongruence between matri- and patrilineal genetic structures could be mainly interpreted by taking into account the cultural practice of patrilocality, i.e. a higher rate of women's versus men's mobility, among the Armenians, which is a widely used tradition in many human populations. Moreover, the lack of differentiation was also found between all Armenian groups (except Salmast/Khoy) and Syrians, Central and East Armenians and Georgians, and East Armenians and Chechens. This pattern of genetic differentiation might indicate that possible gene flow took place between these populations during long period of time. It is notable, that the pattern of genetic homogeneity ends on the north-west of the Caucasus while considering the population of the Adyghs.

To investigate the amount of genetic variation among regional groups we have performed the AMOVA analysis to partition the total genetic variation among regional groups, and populations within these groups. The results have shown that the greatest part of variation is found within populations (98.6 %), whilst among groups the percentage of variation is negligible (0.91%), which indicates a very high level of homogeneity of matrilineal gene pool of the populations from the observed regional groups.

On the other hand, we have also conducted the PCA analysis and constructed UPGMA tree based on different genetic distance in order to visualize the genetic relationships of the studied populations. While the PCA plots did not show a good clustering, the phylogenetic tree has united all the Armenian sub-populations into a distinct from other populations clade, thus indicating a very high level of relatedness between geographically different Armenian groups.

In general, the results obtained indicate that the modern genetic approaches provide very strong and sensitive tools to disentangle some crucial events related to the origin of Armenians, their ancient migrations and contacts with neighboring populations and invaders.

CONCLUSIONS

Based on the results obtained in our project we have made the following conclusions:

1. The lack of the mtDNA lineages of the Central Asian and African origin witnesses the very weak genetic contacts of the Armenians with other ethnically different populations during last millennia despite active ancient migrations in the region.

2. The Armenian matrilineal genetic pool demonstrates very high level of diversity in the region, which might reflect the existence of numerous source populations contributed to the origin of Armenians.

3. According to mtDNA markers the Armenians have the highest genetic affinity with the Levantine populations which possibly points out that the modern Armenians by the maternal lineages are the descendants of the pre-Neolithic and Neolithic migrants from the Near East.

4. Armenians from different geographic groups do not differ significantly from each other, which might be explained by the cultural practice of patrilocality during long centuries.

5. Close genetic affinity of the Armenian matrilineal genetic structure to the Eastern Mediterranean populations may indicate the indigenous origin of the Armenians in West Asia.

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