merhaba tahran universitesinin epidemology bolumunde olan ermeni

arastirmaci ve bazi fars dostlari ve birde ermenistan biology institut

arastirmacilarin uluslar asrasi bir bilimsel dergiye makale gondermisler

ve bu makalede 35 milyon guney azerbaycan ve 20 milyon kuzey azerbaycan

halklarinin genetik yapisinin turkiye halkindan farkli oldunu one

surmusler lutfen bu konuda butun bilimsel arastirmacilari haberdar edin ve

genetik konusunda uzmanlara bu konuyu iletin ,lutfen bu konu cok onemli

,cevabinizi bekliyorum



http://journals.tums.ac.ir/abs.aspx?org_id=59&culture_var=en&journal_id=5&issue_id=2114&manuscript_id=17783&segment=en



makale linki



saygilarla



guney azerbaycan univrsite ogrencisi



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Iranian Journal of Public Health

2011;40(1) : 119-123



Original Article

Iranian Azeri's Y-Chromosomal Diversity in the Context of Turkish-Speaking Populations of the Middle East



*L Andonian1, S Rezaie2, A Margaryan3, DD Farhud4, K Mohammad1, K Holakouie Naieni1, 5, MR Khorramizadeh6, MH Sanati7, M Jamali8, P Bayatian1, L Yepiskoposyan3



1Dept. of Epidemiology and Biostatistics, School of Public Health, Tehran University of Medical Sciences, Tehran Iran

2Division of Molecular Biology, Dept. of Medical Parasitology and Mycology, School of Public Health, Tehran University of Medical Sciences, Tehran, Iran

3Human Genetics Group, Institute of Molecular Biology, National Academy of Sciences, Yerevan, Armenia,

4School of Public Health, Tehran University of Medical Sciences, Tehran Iran

5Iranian Epidemiological Association

6Dept. of Medical Biotechnology, School of Advanced Medical Technologies, Tehran University of Medical Sciences, Tehran Iran

7National Institute for Genetic Engineering and Biotechnology, Pajoohesh Blvd., 17 Km Karaj HWY, Tehran, Iran,

8Academic Member of Ministry of Health and Education, Tehran Iran

Corresponding Author:





L Andonian



E-mail: andonian@tums.ac.ir

Received: September 5,2010

Accepted: February 16,2011

Abstract:



Background: The main goal of this study was to conduct a comparative population genetic study of Turkish speaking Iranian Azeris as being the biggest ethno-linguistic community, based on the polymorph markers on Y chromosome.

Methods: One hundred Turkish-speaking Azeri males from north-west Iran (Tabriz, 2008-2009) were selected based on living 3 generations paternally in the same region and not having any relationship with each other. Samples were collected by mouth swabs, DNA extracted and multiplex PCR done, then 12 Single Nucleotide Polymorphisms (SNPs) and 6 Microsatellites (MS) were sequenced. Obtained data were statistically analyzed by Arlequin software.

Results: SNPs and Microsatellites typing were compared with neighboring Turkish-speaking populations (from Turkey and Azerbaijan) and Turkmens representing a possible source group who imposed the Turkish language during 11-15th centuries AD. Azeris demonstrated high level of gene diversity compatible with patterns registered in the neighboring Turkish-speaking populations, whereas the Turkmens displayed significantly lower level of genetic variation. This rate of genetic affiliation depends primarily on the geographic proximity.

Conclusion: The imposition of Turkish language to this region was realized predominantly by the process of elite dominance, i.e. by the limited number of invaders who left only weak patrilineal genetic trace in modern populations of the region.





Keywords:

Iranian Azeris ، Y chromosome diversity ، Microsatellites ، SNPs ، Iran

Permanent Link: http://journals.tums.ac.ir/abs/17783



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Iranian J Publ Health, Vol. 40, No.1, 2011, pp.119-127 Original Article

119

Iranian Azeri's Y-Chromosomal Diversity in the Context of

Turkish-Speaking Populations of the Middle East

*L Andonian1, S Rezaie2, A Margaryan3, DD Farhud4, K Mohammad1, K Holakouie Naieni1, 5,

MR Khorramizadeh6, M H Sanati7, M Jamali8, P Bayatian1, L Yepiskoposyan3

1Dept. of Epidemiology and Biostatistics, School of Public Health, Tehran University of Medical Sciences,

Tehran Iran

2Division of Molecular Biology, Dept. of Medical Parasitology and Mycology, School of Public Health, Tehran

University of Medical Sciences, Tehran, Iran

3Human Genetics Group, Institute of Molecular Biology, National Academy of Sciences, Yerevan, Armenia,

4School of Public Health, Tehran University of Medical Sciences, Tehran Iran

5Iranian Epidemiological Association

6Dept. of Medical Biotechnology, School of Advanced Medical Technologies, Tehran University of Medical

Sciences, Tehran Iran

7National Institute for Genetic Engineering and Biotechnology, Pajoohesh Blvd., 17 Km Karaj HWY, Tehran, Iran,

8Academic Member of Ministry of Health and Education, Tehran Iran

(Received 5 Sep 2010; accepted 16 Feb 2011)

Introduction

Due to its geo-strategic location in the Middle

East, the Iranian plateau has served as a key

crossroad for human disseminations and played

a critical role in the migratory waves between

the populations of the Middle East and beyond

(1-5). The most important long-term factor in

this process was human adaptation to the Iranian

plateau and its geographical, topographical, and

climatic conditions with the subsequent development

of agriculture, pastoralism, and pastoral nomadism.

The spread of these technological innovations,

along with a series of major demographic

and historical events, has resulted in a large diversity

and dispersal of ethnic groups and languages

(1, 6, 7).

Abstract

Background: The main goal of this study was to conduct a comparative population genetic study of Turkish speaking

Iranian Azeris as being the biggest ethno-linguistic community, based on the polymorph markers on Y chromosome.

Methods: One hundred Turkish-speaking Azeri males from north-west Iran (Tabriz, 2008-2009) were selected based

on living 3 generations paternally in the same region and not having any relationship with each other. Samples were

collected by mouth swabs, DNA extracted and multiplex PCR done, then 12 Single Nucleotide Polymorphisms (SNPs)

and 6 Microsatellites (MS) were sequenced. Obtained data were statistically analyzed by Arlequin software.

Results: SNPs and Microsatellites typing were compared with neighboring Turkish-speaking populations (from Turkey

and Azerbaijan) and Turkmens representing a possible source group who imposed the Turkish language during 11-15th

centuries AD. Azeris demonstrated high level of gene diversity compatible with patterns registered in the neighboring

Turkish-speaking populations, whereas the Turkmens displayed significantly lower level of genetic variation. This rate

of genetic affiliation depends primarily on the geographic proximity.

Conclusion: The imposition of Turkish language to this region was realized predominantly by the process of elite

dominance, i.e. by the limited number of invaders who left only weak patrilineal genetic trace in modern populations of

the region.

Keywords: Iranian Azeris, Y chromosome diversity, Microsatellites, SNPs, Iran

*Corresponding author: E-mail: andonian@tums.ac.ir

L Andonian et al: Iranian Azeri's Y-Chromosomal Diversity …

120

Between the third and second millennia BC, the

Iranian plateau became exposed to incursions of

pastoral nomads from the Central Asian steppes

(1), which were a difficult environment for agriculture

but ideally suited to animal husbandry and

pastoral nomadism.

Presumably, via an elite-dominance process, existing

Dravidian language across the regeon was

substituted by Indo-Iranian language, which is a

branch of Indo-European language (8-10). Also

their genetic impacts were as significant as the

imposition of their language, which is clearly observed

in Iran (11), Pakistan (12) and northern

India (13).

In the period of the eleventh to thirteenth centuries

AD the Arab-Muslim, Seljuk and subsequent

Turkic-Mongol invasions signaled the arrival of a

new people with flocks and culture. Specifically,

in a series of rapid Arab-Muslim conquests in

the seventh century, the Arab armies swept through

most of the Middle East, completely engulfing the

Persian lands (7).

The dominance of the Arabs came to a sudden

end in the mid-eleventh century with the arrival

of Seljuk Turks, a clan of the Oguz Turks (8).

The expanding waves of these Altaic-speaking

nomads from Central Asia involved regions farther

to the west, such as Iran, Iraq, Anatolia, and

the Caucasus, where they imposed Altaic (Turkish)

languages. In these western regions, however, the

genetic contribution is low or undetectable (14),

even though the power of these invaders was

sometimes strong enough to impose a language

replacement, as in Turkey and Azerbaijan (1).

Later, the Mongol armies also moved westward

and, by the early thirteenth century, established

their rule over a vast region, including Iran and

advancing as far west as the Caucasus and Turkey

(1, 7). These waves of various invasions and

subsequent migrations resulted in major demographic

expansions in the region, which added

new languages and culture to the mix of peoples

that had pre-existed in Iran.

In general, a considerable genetic diversity is observed

in Iranian populations, which resembles

to that of Middle East patterns as a whole and

strengthens the idea of Persia, being the main

crossroad for human dissemination (3, 11, 15-18).

This area is remarkable for its high level of ethnic

and linguistic diversity, comprising the major

language families (Indo-European, Altaic, and

Afro-Asiatic) currently spoken by more than seventy

ethnically different populations (http://www.

ethnologue.org/). This demonstrates the role of

Iran, which played in population dispersal across

the latitudinal belt spanning from western Anatolia

to the Indus Valley. However, there have been

gaps in high-resolution genetic analyses for this

region to uncover population history at a fine scale,

for example, for particular ethnic and linguistic

groups.

In this project, we intended to get relatively

comprehensive information about the Y chromosome

diversity in Azeris living in Iran. Subsequently,

the principal aim of this paper was to identify the

place of Azeris in the frame of Turkic-speaking

populations of the Middle East and to test the

extent of gene flow from Central Asia. We used

both SNP and STR genetic markers on the nonrecombining

portion of Y chromosome, which provide

high level of genetic resolution and are consistent

with other sets of markers applied to the

studies on patrilineal genetic history of various populations

(19, 20).

Material and Methods

Buccal swab specimens were collected from 100

ethnical Azeri men currently living in Tabriz, Iran

in 2008. One sample was later discarded, as the

Y-chromosome typing was unsuccessful.

All donors were selected only if their paternal

grandfathers were from the same region and they

were unrelated to other donors at the grandfather

level. The samples were stored in a DNA preservative

solution consisting of 0.5% sodium dodecyl

sulphate and 0.05M ethylenediaminetetraacetic acid

for transport purposes. Samples were collected

anonymously and informed consent was obtained

from all individuals before samples were taken.

The comparative data sets have been taken from

previously published papers (19, 21, 22); the reIranian

J Publ Health, Vol. 40, No.1, 2011, pp.119-127

121

sults for the Turkmen population are available only

for microsatellite loci, therefore the comparison with

this data set was conducted only at STR markers.

Standard phenol-chloroform DNA extractions were

performed. The strategy adopted for typing samples

was designed to ensure informative comparison

with existing published data. NRY were characterized

by 12 binary Y chromosome polymorphisms:

92R7, M9, M13, M20, sY81, SRY+465,

SRY4064, SRY10831, Tat, M17, Alu insert-YAP,

and p12f2, (19, 23) and screened for six microsatellite

(MS) markers: DYS19, DYS388, DYS390,

DYS391, DYS392, and DYS393, as described

by Thomas et al. (24). Haplogroups (hg) were

defined by single nucleotide polymorphism (SNP)

markers according to the Y Chromosome Consortium

nomenclature (25). Microsatellite repeat numbers

were assigned according to Kayser et al. (26).

Either the microsatellite PCR products and UEP

digestion products were run on an ABI-310 capillary-

based genetic analyzer or a gel based system

such as an ABI-377 automated sequencer. For the

ABI-310 genetic analyzer, 1.2 ul aliquots of the

microsatellite PCR products or UEP digestion

products were mixed with 0.5 ul size standard

labeled with the fluorescent dye TAMRA (PEApplied

Biosystems) and 12 ul of de-ionized formamide.

Samples were denaturated at 96° C for

3 min and chilled on ice for 5 min before being

run, using POP-4 polymer and a 36 cm POP-4

capillary. For the ABI-377 automated sequencer,

1.0 ul aliquots of the microsatellite PCR products

or UEP digestion products were mixed with

2.0 ul of a loading buffer (formamide: dextran blue:

TAMRA-labeled size standard, in the ratio of

23: 4: 2).

Unbiased genetic diversity index, h, and its standard

error were calculated using the formulae of

Nei (27). Nei’s Genetic Identity, I, was calculated

in accordance with Nei (27). Pairwise genetic

distances (FST) were estimated from analysis of

molecular variance (AMOVA) JST values with the

aid of Arlequin software (28). Tests for significant

population differentiation were carried out using

the exact test for population differentiation (27, 29).

Testing for differences in h between populations

was performed by bootstrapping method (30).

Principal Coordinates Analysis was conducted on

similarity matrices calculated as one minus Genetic

distance (FST, RST) or based on Nei’s Genetic Identity

values. Figures along the main diagonal, representing

the similarity of each population sample to

itself, were calculated from the estimated genetic

distance between two copies of the same sample.

Signature haplotype analysis (high frequency modal

haplotypes and modal clusters) (19, 31-33)

was performed by hand.

Results

The number of haplogroups detected in Azeris is

the highest, nine, whereas in Eastern Turks, it is

seven, and in Azerbaijanis, it is only five. The latter

might be explained by the limited number of specimens

in this data set. Nevertheless, we tried to

make some inferences about the genetic structure

of these groups based on haplogroup frequencies.

The most common haplogroup in all data sets is

haplogroup J that is present at almost equal rate

in the three groups: 39.39% in Azeris, 40.00% in

Azerbaijanis and 39.02% in Eastern Turks. The

next frequently encountered haplogroup is hg BR*

(xDE,JR) which is registered at 23.23%, 40.00%,

and 9.76%, respectively. The haplogroups P*(xR1a),

E*(xE3a) and R1a1 are also rather frequent in

Eastern Turks (19.51%, 14.63% and 10.98%, respectively).

It is necessary to add that haplogourp

N3 defined by Tat mutation which presumably

originated in Central Asia (34) is detected

only in Azeris and Azerbaijanis.

The overall comparison of population structures

between the groups shows that Azerbaijanis differ

significantly from Eastern Turks according to

exact test of population differentiation (P< 0.0001);

in the same time, there is no significant disparity

between the Iranian Azeris and the two other

(Azerbaijanis and Eastern Turks) comparative

data sets (P> 0.05). This pattern of genetic relatedness

is supported also by Fst values, which

indicate closer genetic affinity between the Azeris

and Eastern Turks, as well as Azeris and AzerbaiL

Andonian et al: Iranian Azeri's Y-Chromosomal Diversity …

122

janis; the biggest genetic distance was detected

between Azerbaijanis and Eastern Turks.

At STR level, we had possibility to add Turkmens

in comparative data sets. Microsatellite markers

identified 137 haplotypes in total while considering

the four groups. The most diverse pattern

of haplotypes is observed in Azerbaijanis

and Azeris (32 haplotypes in 40 samples, and 76

in 99, respectively). In contrast, Eastern Turks

and Turkmens display much lower level of variability

(49 in 82 and 18 in 51, respectively). It is

worth mentioning that Azeris bear 52 unique

haplotypes, i.e. they are not encountered in other

three data sets; only two haplotypes were shared

by all groups considered.

The above-mentioned pattern of genetic variability

is reflected in the actual values of gene diversity,

h (Fig. 1). The Azeris show the highest level of genetic

diversity (h= 0.9934, bootstrapped value -

0.9834). The dramatically lower rate of this parameter

was registered for Turkmens (actual value

-0.8267, bootstrapped value -0.8068). Azerbaijanis

and Eastern Turks also have rather high

level of gene diversity, although they are still lower

than in Azeris. While comparing differences in h

values we found two not significant differencesbetween

Azerbaijanis and Azeris, as well as between

Azerbaijanis and Eastern Turks- using bootstrap

method. In case of Bayesian approach, all

possible comparisons show significant level of

differences (P< 0.0001, Table 1).

The AMOVA analysis revealed that the bulk of

observed genetic diversity is explained by withinpopulation

differentiation (94.16%), and only about

5.84% reflects inter-population variability. Once

again, this result supports the general rule that

within-group variability is the main source of

human genetic diversity.

The Azeri modal haplotypes, ht 14-15-23-10-11-

12, is detected at 5.05% rate and is modal in

Azerbaijani data set (7.50%). The comparable

level of this haplotype is found in Eastern Turks

(6.10%), while in Turkmens it is at 1.96%. The

modal haplotypes in Eastern Turks data set, each

at 8.54%, ht 13-12-24-10-11-13 and ht 14-12-

24-11-13-12, are detected also in Azerbaijanis

(2.50% each) and in Azeris (3.03% and 2.02%,

respectively), being absent in Turkmens. Two

modal haplotypes of Turkmens, ht 15-12-24-10-

11-13 (35.29%) and 16-12-24-10-11-13 (21.57%)

display very low frequency only in Azeris (2.02%

and 1.01%, respectively).

Actually, these two haplotypes are one-step neighbors

and therefore might be considered as one

modal cluster. The modal cluster in Turkmens, the

most pronounced one, accounting for 60.78%; was

found at low frequency only in Azeris (3.03%)

and is totally absent in Azerbaijanis and Eastern

Turks (Table 2). The modal cluster of Azeris and

Azerbaijanis, being the same, accounts, respectively,

for 20.00% and 14.14%; it is at comparable

rate in Eastern Turkey (9.76%) and at much

lower level in Turkmens (1.96%). Eastern Turks’

modal clusters, accounting for 10.98% and 9.76%,

are present only in neighboring populations (Azerbaijanis

and Azeris) and are absent in Turkmens.

It displays some important patterns of relationships

between the groups. First, it is rather evident

that Turkmens are almost equally distant from

the rest of Turkic-speaking populations studied.

In the same time, Azeris, Azerbaijanis, and Eastern

Turks form some sort of dense cluster, possibly

reflecting the close genetic contacts between these

groups compared to Central Asian Turkic-speaking

peoples. While constructing PCO plots based

on SNP+MS haplotypes, the relationship between

Middle Eastern populations becomes more refined

and it is in a full accordance with the results of

genetic distance comparison.

Once again, the plot proves that Azeris occupy

intermediate position between their close neighbors

that might witness to some extent a common origin

and/or intense genetic contacts since ancient times.

The mentioned relationships between the populations

are fully supported by the exact test of

population differentiation (Table 3 based on MS

data only).

Genetic distance data (Rst based on MS only)

were used to visualize the spatial relationships

of the groups (Fig. 2).

Iranian J Publ Health, Vol. 40, No.1, 2011, pp.119-127

123

Fig. 1: Genetic diversity, h, with bootstrap 95% confidence intervals

Table 1: Pairwise differences in h values based on SNP+MS haplotype (lower left table, based on bootstrap approach; upper

right table, based on Bayesian method)

Azeri Azerb ET Turkmen

(n=99) (n=40) (n=82) (n=51)

h

Azeri - 0.0000 0.0000 0.0000 0.9934

Azerb 0.5721 - 0.0000 0.0000 0.9885

ET 0.0122 0.9004 - 0.0000 0.9744

Turkmen 0.0000 0.0001 0.0003 - 0.8267

Significant values in underlined

Table 2: Frequently Encountered Clusters

Azerb Azeri ET Turkmen

MS

(n=40) (n=99) (n=82) (n=51)

14 15 23 10 11 12 0.2000 0.1414 0.0976 0.0196

13 12 24 10 11 13 0.0250 0.0606 0.1098 0.0000

14 12 24 11 13 12 0.1000 0.0303 0.0976 0.0000

15 12 24 10 11 13 0.0000 0.0303 0.0000 0.6078

Significant values in underlined

L Andonian et al: Iranian Azeri's Y-Chromosomal Diversity …

124

Fig. 2: Principal coordinates plot based on RST values

Table 3: P values for the exact test of population differentiation

based on MS data

Azerb

(n=40)

Azeri

(n=99)

ET

(n=82)

Turkmen

(n=51)

Azerb -

Azeri 0.363 -

ET 0.009 0.060 -

Turkmen 0.000 0.000 0.000 -

Discussion

The results obtained show that the rate of genetic

relatedness between the populations considered

depends in the first instance on the spatial

proximity than on the belonging to the same linguistic

group. In this context, these outcomes were

highly expected taking into consideration the actual

geographic location of the three populations. Azeris,

being situated in between Azerbaijanis and Eastern

Turks, had more possibilities of genetic contacts

with both groups as the closest neighbors,

while gene flow between Azerbaijan and Eastern

Turkey could have been rather limited.

In general, we can make rather strong inferences

about the genetic relatedness between the populations

under consideration. The principal one is that

Iranian Azeris have much weaker genetic affinity

with Turkmens than with their immediate neighbors.

The same statement could be attributed to

the Azerbaijanis and Eastern Turks. It seems

that Turkmens had no marked input in the gene

pool of Azeris, Azerbaijanis, and Eastern Turks,

despite very close linguistic affinity of these groups

belonging to Turkic-speaking populations. We have

all grounds to suggest that language replacement

took place through elite dominance phenomenon

rather than demic diffusion model (35).

A weak genetic affinity between Middle Eastern

Turkic-speaking populations and Turkmens is

possibly explained by the fact that Central Asian

populations had not any essential gene flow to

the origin of Turkic speaking peoples of South

Caucasus and Asia Minor, which is supported

also by the results of CinnioOlu et al. (21).

Therefore, the imposition of Turkic language to

this region was realized predominantly by limited

number of invaders who left only weak genetic

signal in modern populations of the region.

The same pattern of geographic vs. genetic relatedness

was revealed while comparing Indo-

European speaking Bakhtiari, and Semitic-speaking

Arabs (36). Both mtDNA and the Y chromosome,

showed a close relationship of these groups with

each other and with neighboring geographic groups,

irrespective of the language spoken. Moreover,

Semitic-speaking North African groups are more

distant genetically from Semitic-speaking groups

from the Near East and Iran. Similar results were

recently obtained in the region of north-west Iran:

the Uromian people (Iranian Muslim group) display

a particularly close genetic relationship to the

Armenians living in the same area (4). Thus, geographical

proximity better explains genetic relatedness

between populations than linguistic relatedness

in this part of the world.

As it was shown in our recently published results

(37), based on multivariate classification Iranian

Azeris and their close neighbors, Persians and

Armenians, form a rather distinct cluster of Middle

East origin. This pattern was obtained both while

using Principal Coordinate Analysis and Neighbor-

Joining method for phylogenetic inferences.

As a whole, the results obtained indicate that the

used set of genetic markers is an appropriate tool

Iranian J Publ Health, Vol. 40, No.1, 2011, pp.119-127

125

for population genetics study of such an ethnically

and linguistically complex area as the Middle

East. The methods applied allowed distinguishing

fine specific features of each population and

making inferences about their origin and possible

ancient genetic contacts.

We also realize that the Y chromosome represents

only one locus in the human genome and

describes only one, patrilineal, facet of the genetic

history of human populations. The more comprehensive

results on the origin, ancient relationships,

and migrations of the populations of the Middle

East can be achieved while using other complementary

genetic systems, i.e. mtDNA and autosomal

markers. Nevertheless, the Y chromosome

markers provide rather strong information on the

genetic history of the populations studied in the

frame of this project. In addition, the main outcome

of the project is that all the three populations

could be considered as indigenous representatives

of the area of their inhabitance with very limited

genetic influence from East.

Ethical Considerations

Ethical issues including plagiarism, informed

consent, misconduct, data fabrication and/or

falsification, double publication and/or submission,

redundancy, etc. have been completely observed

by the authors.

Acknowledgements

The authors would want to thank all people who

had contribution to this study (Armenian and Persian

volunteers in Isfahan as well as Azeri volunteers

in Tabriz). This research was planned according

to a MOU between Tehran University of

Medical Sciences (TUMS) from one side and

The Centre for Genetic Anthropology (TCGA) in

University College London and The Institute of

Man in Yerevan from the other sides. Therefore,

the authors wish to thank all the colleagues in all

the three sides, which have been involved in this

research. This research has been supported by

Tehran University of Medical Sciences under grant

number 5213. The authors declare that they have

no conflicts of interest.

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