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Mtdna and Y-Chromosome Variation in the Talysh of Iran and Azerbaijan

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Mtdna and Y-Chromosome Variation in the Talysh of Iran and Azerbaijan AMERICAN JOURNAL OF PHYSICAL ANTHROPOLOGY 138:82–89 (2009) mtDNA and Y-Chromosome Variation in the Talysh of Iran and Azerbaijan Ivan Nasidze,1* Dominique Quinque,1 Manijeh Rahmani,2 Seyed Ali Alemohamad,3 Pervin Asadova,4 Olga Zhukova,4 and Mark Stoneking1 1Department of Evolutionary Genetics, Max Planck Institute for Evolutionary Anthropology, D-04103 Leipzig, Germany 2Department of Molecular Genetics, Cardiovascular Research Center, Imam Hospital, Tehran University of Medical Sciences, Tehran, Iran 3Department of Human Genetics, School of Public Health, Tehran University of Medical Sciences, Tehran, Iran 4N.I. Vaviliv Institute for General Genetics, Russian Academy of Sciences, Moscow, Russian Federation KEY WORDS Northern Talysh; Southern Talysh; Y-chromosome; mtDNA ABSTRACT The Northern Talysh from Azerbaijan Talysh Y-chromosome variation differs from that of and the Southern Talysh from Iran self-identify as one neighboring groups, probably as a result of genetic drift. ethnic group and speak a Northwestern Iranian language. This genetic drift most likely reflects a founder event in However, the Northern and Southern Talysh dialects are the male gene pool of Northern Talysh: either fewer so different that they may actually be separate lan- males than females migrated to Azerbaijan, or there guages. Does this linguistic differentiation reflect inter- was a higher degree of relatedness among the male nal change due to isolation, or could contact-induced migrants. Since we find no evidence of substantial change have played a role? We analyzed mtDNA HVI genetic contact between either Northern or Southern sequences, 11 Y-chromosome bi-allelic markers, and 9 Y- Talysh and neighboring groups, we conclude that inter- STR loci in Northern and Southern Talysh and compared nal change, rather than contact-induced change, most them with their neighboring groups. The mtDNA data likely explains the linguistic differentiation between show a close relatedness of both groups with each other Northern and Southern Talysh. Am J Phys Anthropol and with neighboring groups, whereas the Northern 138:82–89, 2009. VC 2008 Wiley-Liss, Inc. Determining the extent to which the genetic and lin- differentiation between the Northern and Southern guistic relationships of populations are correlated can Talysh dialects: 1) divergence from a common ancestral provide insights into population history. A number of language, due to isolation; and 2) contact-influenced previous studies of neighboring groups, who speak differ- language change of one or both dialects. Indeed, there is ent languages, has revealed the complex nature of such some linguistic evidence to indicate that Northern relationships (Nasidze and Stoneking, 2001; Nasidze Talysh, and possibly Southern Talysh as well, are et al., 2003; Nasidze et al., 2004; Nasidze et al., 2005; ‘‘mixed’’ languages, exhibiting characteristics of two or Nasidze et al., 2006; Pakendorf et al., 2006; Nasidze possibly even more languages (Donald Stilo, personal et al., 2007; Pakendorf et al., 2007). However, the subjects communication). Genetic studies can help elucidate the of these studies are usually different self-defined groups, historical circumstances that led to the differentiation who speak different languages and often have different between the Northern and Southern Talysh dialects, as histories. Meanwhile, there are cases in which people, language change due to contact between speakers of who identify as one ethnic group with a common history, either (or both) dialects with some other group(s) should and presumably a shared common ancestry, in fact differ also have left a genetic signature of the contact. We have linguistically enough from each other to be classified as therefore examined patterns of mtDNA and non-recom- speaking separate languages. Examples are the North- bining Y-chromosome (NRY) variation in Northern and ern and Southern Talysh (Clifton et al., 2005; Donald Southern Talysh, and compared them with their neigh- Stilo, personal communication). The Talysh are an Iranian people who speak a North- western Iranian language and are found in the northern regions of the Iranian provinces of Gilan and Ardabil Grant sponsor: Max Planck Society, Germany. and in the southern parts of the Republic of Azerbaijan, *Correspondence to: Ivan Nasidze, Department of Evolutionary South Caucasus (Fig. 1). Talysh has two major dialects— Genetics, Max Planck Institute for Evolutionary Anthropology, Northern (in Azerbaijan and Iran), and Southern (in Deutscher Platz 6, 04103 Leipzig, Germany. Iran) (Ethnologue, 2000). These two dialects exhibit sub- E-mail: [email protected] stantial differences in grammatical structure and lexi- con, and low mutual comprehension, and it has been Received 2 April 2008; accepted 31 May 2008 argued that they should be considered different lan- guages (Donald Stilo, personal communication). DOI 10.1002/ajpa.20903 In principle, there are two processes, that are not Published online 18 August 2008 in Wiley InterScience mutually-exclusive, that could have lead to the extreme (www.interscience.wiley.com). VC 2008 WILEY-LISS, INC. mtDNA AND Y-CHROMOSOME VARIATION IN TALYSH 83 Fig. 1. A map of the geographic location of the sampling sites for the Northern and Southern Talysh in this study from Len- koran and Masal, respectively. boring groups in Iran and the Caucasus, to determine if sequenced again in each direction, so that each base was contact-influenced language change may have played a determined twice. role in their history. Published mtDNA HV1 sequences were included from Iran, Central Asia, and the South Caucasus (Comas et al., 1998; Nasidze et al., 2004; Chaix et al., 2007). MATERIALS AND METHODS mtDNA haplogroups that are most informative in Samples and DNA extraction southwest Asia, i.e., haplogroups H, J, N, T, and U (Quintana-Murci et al., 2004), were determined by PCR- A total of 128 whole blood samples from unrelated RFLP assays of the relevant SNP, using previously- individuals, representing two Talysh groups—Southern described methods (Finnila et al., 2000; Quintana-Murci Talysh from Iran (50 samples) and Northern Talysh et al., 2004). from Azerbaijan (78 samples)—were collected (Fig. 1). Genomic DNA from blood samples was extracted using a Qiagen DNA extraction kit (Qiagen GmbH, Germany), Y-Chromosome bi-allelic markers following the protocol recommended by the supplier. All 128 samples were typed for the X- and Y-linked Informed consent and information about birthplace, zinc finger protein genes to confirm the gender of the parents, and grandparents was obtained from all donors. sample (Wilson and Erlandsson, 1998). Genotyping was carried out for 10 NRY SNP markers: M9, M17, M45, mtDNA first hypervariable segment sequences M89, M124, RPS4Y (M130), M170, M173, M172, and M201 (Underhill et al., 2000); the YAP Alu insertion The first hypervariable segment (HV1) of the mtDNA polymorphism (Hammer and Horai, 1995) was also control region was amplified using primers L15996 and typed. The markers M9 and RPS4Y were typed by H16410 (Vigilant et al., 1989), as described previously means of PCR-RFLP as described elsewhere (Kayser (Redd et al., 1995). The nested primers L16001 (Cordaux et al., 2000). The markers M17, M124, M172, M173, et al., 2003) and H16401 (Vigilant et al., 1989) were M45, and M201 were typed using PIRA-PCR (primer used to determine sequences for both strands of the PCR introduced restriction analysis) assays (Yoshimoto et al., products with the DNA Sequencing Kit (Perkin–Elmer), 1993) as described elsewhere (Cordaux et al., 2004). M89 follow the protocol recommended by the supplier, and an was typed as described elsewhere (Ke et al., 2001), while ABI 3700 automated DNA sequencer. Individuals with the YAP Alu insertion was typed as described by Ham- the ‘‘C-stretch’’ between positions 16184 and 16193, mer and Horai (1995). Polymorphism of M170 was which is caused by the 16189C substitution, were detected by direct sequencing of the PCR product on an American Journal of Physical Anthropology 84 I. NASIDZE ET AL. TABLE 1. mtDNA HV1 sequence variability among Southern and Northern Talysh populations and neighboring groups No. of Haplotype Population N haplotypes diversity and SE MPD Tajima’s D Source Southern Talysh (Iran) 49 39 0.987 6 0.008 5.41 21.83* Present study Northern Talysh (Azerbaijan) 78 60 0.981 6 0.009 5.96 22.20** Present study Other groups from Iran Gilaki 87 73 0.995 6 0.003 6.40 22.19** Nasidze et al., 2006; Quintana-Murci et al., 2004 Mazandarani 71 58 0.992 6 0.005 5.98 22.02** Nasidze et al., 2006; Quintana-Murci et al., 2004 Iranians Tehran 79 63 0.984 6 0.008 5.53 22.05** Nasidze et al., 2004 Iranians Isfahan 46 42 0.996 6 0.006 6.17 22.13** Nasidze et al., 2004 Lur 17 15 0.978 6 0.031 5.52 21.85* Quintana-Murci et al., 2004 Iran, Arabs 46 40 0.991 6 0.008 7.56 21.86* Nasidze et al., 2008 Bakhtiari 53 44 0.991 6 0.006 7.14 22.20** Nasidze et al., 2008 Kurds 20 19 0.995 6 0.018 6.13 21.57* Quintana-Murci et al., 2004 Groups from the South Caucasus Georgians 57 40 0.971 6 0.007 5.16 21.99** Nasidze et al., 2004 Armenians 42 35 0.980 6 0.003 5.22 22.18** Nasidze et al., 2004 Azeri 41 37 0.995 6 0.006 5.17 22.13** Nasidze et al., 2004 * P \ 0.05; **P \ 0.01. TABLE 2. Mean pairwise Fst values between Northern and Southern Talysh, and neighboring groups Northern Southern South Caspian The rest West Asia Central Talysh Talysh Caucasus region of Iran of Iran w/o Iran Asia Northern Talysh 0.013 0.016 0 0.007 0.004 0.042 Southern Talysh 0.070 0.019 0.007 0.013 0.011 0.052 South Caucasus 0.080 0.023 0.017 0.020 0.013 0.065 Caspian region of Iran 0.057 0.011 0.013 0.004 0.002 0.047 The rest of Iran 0.129 0.038 0.038 0.050 0.007 0.052 West Asia w/o Iran 0.124 0.063 0.061 0.078 0.045 0.050 Central Asia 0.158 0.111 0.109 0.047 0.106 0.108 Below diagonal: paiwise Fst values based on Y-SNP haplogroups; above diagonal: pairwise Fst values based on mtDNA HVI sequences.
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