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Haplogroup R1a (Y-DNA)



  R1a (Y-DNA) is a specific sequence of nucleotides in the male Y chromosome and a Y-chromosome haplogroup identified with the genetic marker M17. R1a originated as a single mutation of one male, the R1a originator considered to be the ancestor of all individuals carrying R1a, and descends of Haplogroup R1.

R1a is present at high frequency (40 per cent plus) from the Czech Republic across to the Altai Mountains in Siberia and south throughout Central Asia. The first carriers of the R1a haplotype are believed to have been peoples living about 15,000 years ago[1] confined by an area within the Ukrainian LGM refuge. The gene spread by a nomadic lifestyle and proliferated on Eurasian steppes.

Contents

Subclades

  • R1a (SRY1532)
    • R1a1 (M17, M198)
      • R1a1a (M56)
      • R1a1b (M157)
      • R1a1c (M64.2, M87, M204)
      • R1a1*
    • R1a*

There is some suggestion that R1a and R1a1 split within Southern and Western Asia. Several possibilities for the spread of R1a and R1a1 exist, which include recent Slavic migration from the 5th century AD, movement of Kurgan people (that is, the spread of the Indo-European languages) associated with the domestication of the horse and/or the re-colonization of Europe following the end of the last ice-age.

Origins

  Absolute dating methods suggest that this marker is 10–15,000 years old, and the microsatellite diversity is greatest in southern Russia and Ukraine, suggesting that it arose there. Investigations suggest the gene expanded from the Dniepr-Don Valley, between 13 000 and 7600 years ago, and was linked to the reindeer hunters of the Ahrensburg culture that started from the Dniepr valley in Ukraine and reached Scandinavia 12 000 years ago.[2] Ornella Semino et al. (see [3]) propose a postglacial spread of the R1a1 gene from the Ukrainian LGM refuge, subsequently magnified by the expansion of the Kurgan culture into Europe and eastward. R1a1 is most prevalent in Poland, Russia, Ukraine, Hungary and is also observed in Pakistan, India and central Asia. Wells suggests the origin, distribution and age of R1a1 points to an ancient migration, possibly corresponding to the spread by the Kurgan people in their expansion across the Eurasian steppe around 3000 BC.[3]

Current theories point to the gene being tied to speakers of the Proto-Indo-European language in the Kurgan scenario, spreading the gene further to Asia and most of Europe. The low occurrence of R1a1 in Western European Indo-European speaking populations, most notably the region west of the Vistula[4] — including the enigmatic Nordwestblock — shows that this correlation with PIE cannot be extended to the "kurganized" western Corded ware and subsequent Beaker culture.[5][6] This corresponds to the now widely accepted view that kurganisation never occurred.[7]

Highest haplotype incidence suggests that haplogroup R1a1 originated among the ancestors of the Balto-Slavic speakers of Eastern and Central Europe.

Marijana Perii &all in 2005 hypothesize that:

At this level of resolution, it is not clear what temporal and effective population size differences contributed to this deep Paleolithic signal as high R1a variance in SEE might be explained by either ancient demography or more recent bottlenecks and founder effects in different Slavic tribes. At least three major episodes of gene flow might have enhanced R1a variance in the region: early post-LGM recolonizations expanding from the refugium in Ukraine, migrations from northern Pontic steppe between 3000 and 1000 B.C., as well as possibly massive Slavic migration from A.D. 5th to 7th centuries.

Distribution Overview

  In Europe, the highest frequencies are found in Central and Eastern Europe. Today it is found at its highest levels in Tajiks (64%), Kyrgyz (63%), Poland and Hungary (56%–60%), Ukraine (44-54%[8], depending on the source), and Russia, where one out of two men has this haplogroup. In Hungary contradicting frequencies are reported 60% or 20%. Relatively high frequencies are also found among the ethnic Sorbs (63%) in Eastern Germany and in Scandinavia[2] (the largest being 23% in Iceland). The modern population of Ukraine has the highest level of diversity of the gene making it the likeliest location of its origin.[2] High haplotype diversity was detected in northern Poland where for 508 males Pawlowski et al[9] found 328 different and 264 unique haplotypes, he wrote "Model for a Polish population haplotype …is almost 15 times more frequent in our population than in a cumulative European one" (for better picture compare this diversity to this map of R1a1 frequency) or more accurate map C on this map[10].

Even in South Eastern Europe (not a major concentration of R1a1) microsatellite networks of major Y chromosomal lineages show high diveristy of R1a1 graph C[10][11]. The variance cluster in South Eastern Europe (SEE) is located in the Republic of Macedonia.

Europe

R1a1 is spread across the whole of Europe, with the highest concentrations found in Poland. The two main directional components of the spread are consistent with an East to West migration as well as a radial spread from the Balkans. The latter is claimed to be a trace of the re-population of Europe after the Last Glacial Maximum from the Ukrainian refuge area.[12]

"At least three major episodes of gene flow might have enhanced R1a variance in the region: early post-LGM recolonizations expanding from the refugium in Ukraine, migrations from northern Pontic steppe between 3000 and 1000 B.C., as well as possibly massive Slavic migration from A.D. 5th to 7th centuries." ref The last possibility is less probable, the distribution of Paleolithic pattern depth is unexplained by massive people flow. Genetic data support autochtonic school of Slovian historiography.

India

Further information: Genetics and Archaeogenetics of South Asia: R1a1 and R2

In India initial studies with limited samples observed a correlation between the Brahmin caste and the R1a haplogroup which was consistent with an Indo-Aryan migration from Central Asia (Bamshad et al. 2001),[13] in line with earlier suggestions (Cavalli-Sforza 1994).[14] The frequency gradients of the haplogroup, falling off eastward across Siberia to the Altai mountains and southward into India, were held to perfectly reflect the inferred migrations of the (pre-)Proto-Indo-Iranians and Indo-Iranians during the period 3000 to 1000 BC (Wells et al 2001).[1] The northern migration theory is also supported by the dating of the haplogroup (Wells et al 2003).

However, Studies of India scholars showed the R1a lineage forms around 35–45% among all the castes in North Indian population (Namita Mukherjee et al. 2001) and the Badagas of the Nilgiris making the association with the Brahmin caste more vague. A further study (Saha et al 2005)[15] examined R1a1 in South Indian tribals and Dravidian population groups more closely, and questioned the concept of its Indo-Iranian origin. Most recently Sengupta et al. (2006)[16] have confirmed R1a's diverse presence including even Indian tribal and lower castes (the so-called untouchables) and populations not part of the caste system. From the diversity and distinctiveness of microsatellite Y-STR variation they conclude that there must have been an independent R1a1 population in India dating back to a much earlier expansion than the Indo-Aryan migration.

The pattern of clustering does not support the model that the primary source of the R1a1-M17 chromosomes in India was Central Asia or the Indus Valley via Indo-European speakers.[17]

According to Sengupta et al. (table 5),[16] R1* is virtually absent in Southeast and East Asia.

Asia

In Asia, high R1a1 frequencies are detected in populations of Ishkashimi (68%), Tajiks (64%), and Kyrgyz (63%).[1][18] "The exceptionally high frequencies of this marker in the Kyrgyz, TajikyKhojant, and Ishkashim populations are likely to be due to drift, as these populations are less diverse, and are characterized by relatively small numbers of individuals living in isolated mountain valleys". If the size of a population decreases, for example, in a particular fraternal family all male members will have 100% of R1a1 or 0% of this marker.

The gene has proven to be a diagnostic Indo-Iranian marker[1] and is believed to have been inherited from people who left a clear pattern of archaeological remains known as the Kurgan culture, generally identified as early Indo-Europeans, and later by the Vikings,[2] which accounts for the existence of it in, among other places, the British Isles.[19][20] Lower frequencies of R1a1 are found among populations of West Asia. Iran appears to have had little genetic influence from the R1a1-carrying Indo-Iranians,[1] attributed to language replacement through the "elite-dominance" model.

Frequency distribution

R1a frequency is expressed as percentage of population samples.

Europe

                             N   *R1     R1a1    source
Sorbs                       112     -    63.39   2
Hungarian                    45   13.3   60.0    1     ?-14
Poles                        55   16.4   56.4    1,14
Ukrainian                    50    2.0   54.0    1,14     
Belarusian                  306          50.98   2     ?-14
Russian                     122    7.0   47.0   14
Belarusian                    -          46      4
Belarusian                   41   10.0   39.0   14
Ukrainian                     -          44      3     ?
Ukrainians, Rashkovo         53          41.5   10     ?
Russian, North               49      0   43      5
Latvian                      34   15.0   41.0   14
Udmurt                       43   11.6   37.2    1
Pomor                        28      0   36      5
Macedonian, R.Macedonia      79          15.2   15
Macedonian, North.Greece     20   10.0   35.0    1
Moldavians, Karahasan        72          34.7   10
Croatian                                 34     15e
Lithuanian                   38     6    34     14
Croatian                     58   10.3   29.3    1
UK Orkney                    26     65   27      5
Gagauzes, Etulia             41          26.8   10
Czech + Slovakian            45   35.6   26.7    1,14
Norwegian                    83          26.5   13 
Bosnians                                 25     15e
Icelander                   181   41.4   23.8   14
Norwegian                    87          21.69   2
Moldavians, Sofia            54          20.4   10
Romanians                    54          20.4   10 (Buhusi, Piatra-Neamt) 
Hungarian                    45   13.3   20.4   14
Orcandin                     71   66.0   19.7   14
Swedish (Northern)           48   23.0   19.0   14
Swedish                     110   20.0   17.3   14
Danish                       12   41.7   16.7   14
Herzegovinian                           15     15e    
Mari                         46      0   13.0    1
German                       88          12.50   2
German                       48   47.9   8.1    14
Greek                        76   27.6   11.8    1
Albanian                     51   17.6    9.8    1
Albanian                                 10     15e
Saami                        24    8.3    8.3    1
UK Isle of Man               62   15      8     11
Greek                                     8     15e 
UK Orkney                   121   23      7     11     ?? 7% <> 23% *5
UK                          309          ~7     13     see references
Georgian                     63 ` 14.3    7.9    1
Turks                                     7     15e
Turkish                      30    6.6    6.6    1
UK Shetland                  63   17      6     11
UK Chippenham                51   16      6     11
UK Cornwall                  52   25      6     11
Dutch                        27   70.4    3.7    1   
German                       16   50.0    6.2    1
Italian central/north        50   62.0    4.0    1
Italians                                  3     15e
British                  ~1000          ~4     11  
Irish                       222   81.5    0.5   14
Calabrian                    37   32.4      0    1
Sardinian                    77   22.1           1
British                     25     72      0    5
    
Poles                       913                  9
Germans                    1215                  9
Dniester-Carpathian           -          50.06  10  
Gagauzes, Kongaz             48          12.5   10

empty or - = no data in sample.
?          = datasets differences, [?-x]:= ^x=# source
  • 1 http://hpgl.stanford.edu/publications/Science_2000_v290_p1155.pdf
  • 2 http://www.familytreedna.com/pdf/Levite%20paper.pdf
  • 3 http://www.springerlink.com/content/r60m403330h204l0/
  • 4 http://www.springerlink.com/content/n2883j06628r5515/
  • 9 http://www.springerlink.com/content/w75j6048545350g5/
  • 10 http://edoc.ub.uni-muenchen.de/archive/00005868/01/Varzari_Alexander.pdf
  • 11 http://www.ucl.ac.uk/tcga/tcgapdf/capelli-CB-03.pdf, table 1, more data % < 6
  • 13 http://mbe.oxfordjournals.org/cgi/reprint/19/7/1008.pdf
  • 14 http://mbe.oxfordjournals.org/cgi/content/full/22/10/1964/TBL1 + (15'th primary sources?)
  • 15e http://mbe.oxfordjournals.org/cgi/content/full/22/10/1964#FIG5


  N *R1 R1a1 source
Sorbs 112 - 63.39 Behar et al (2003)[21]
Hungarian 45 13.3 60.0 Semino et al (2000)[8]
113 20.4 Pericic et al (2005)[12]
Poles 55 16.4 56.4 Semino et al (2000),[8] Pericic et al (2005)[12]
Ukrainian 50 2.0 54.0 Semino et al (2000),[8] Pericic et al (2005)[12]
Belarusian 306 50.98 Behar et al (2003)[21]  ?- Pericic et al (2005)[12]
Russian 122 7.0 47.0 Pericic et al (2005)[12]
Belarusian - 46 4
Belarusian 41 10.0 39.0 Pericic et al (2005)[12]
Ukrainian - 44 3  ?
Ukrainians, Rashkovo 53 41.5 10  ?
Russian, North 49 0 43 5
Latvian 34 15.0 41.0 Pericic et al (2005)[12]
Udmurt 43 11.6 37.2 Semino et al (2000)[8]
Pomor 28 0 36 5
Macedonian 20 10.0 35.0 Semino et al (2000)[8]
Moldavians, Karahasan 72 34.7 10
Lithuanian 38 6 34 Pericic et al (2005)[12]
Croatian 58 10.3 29.3 Semino et al (2000)[8]
UK Orkney 26 65 27 5
Gagauzes, Etulia 41 26.8 10
Czech + Slovakian 45 35.6 26.7 Semino et al (2000)[8] ,14
Norwegian 83 26.5 13
Icelander 181 41.4 23.8 Pericic et al (2005)[12]
Norwegian 87 21.69 Behar et al (2003)[21]
Moldavians, Sofia 54 20.4 10
Romanians 54 20.4 10 (Buhusi, Piatra-Neamt)
Hungarian 45 13.3 20.4 Pericic et al (2005)[12]
Orcandin 71 66.0 19.7 Pericic et al (2005)[12]
Swedish (Northern) 48 23.0 19.0 Pericic et al (2005)[12]
Swedish 110 20.0 17.3 Pericic et al (2005)[12]
Danish 12 41.7 16.7 Pericic et al (2005)[12]
Mari 46 0 13.0 Semino et al (2000)[8]
German 88 12.50 Behar et al (2003)[21]
German 48 47.9 8.1 Pericic et al (2005)[12]
Greek 76 27.6 11.8 Semino et al (2000)[8]
Albanian 51 17.6 9.8 Semino et al (2000)[8]
Saami 24 8.3 8.3 Semino et al (2000)[8]
UK Isle of Man 62 15 8 Capelli et al (2003)[19]
UK Orkney 121 23 7 Capelli et al (2003)[19]  ?? 7% <> 23% *5
UK 309 ~7 13 see references
Georgian 63 14.3 7.9 Semino et al (2000)[8]
Turkish 30 6.6 6.6 1
UK Shetland 63 17 6 Capelli et al (2003)[19]
UK Chippenham 51 16 6 Capelli et al (2003)[19]
UK Cornwall 52 25 6 Capelli et al (2003)[19]
Dutch 27 70.4 3.7 Semino et al (2000)[8]
German 16 50.0 6.2 Semino et al (2000)[8]
Italian central/north 50 62.0 4.0 Semino et al (2000)[8]
Brithish ~1000 ~4 Capelli et al (2003)[19]
Irish 222 81.5 0.5 Pericic et al (2005)[12]
Calabrian 37 32.4 0 Semino et al (2000)[8]
Sardinian 77 22.1 Semino et al (2000)[8]
Brithish 25 72 0 5
Poles 913 9
Germans 1215 9
Dniester-Carpathian - 50.06 10
Gagauzes, Kongaz 48 12.5 10 
empty or - = no data in sample.
?          = datasets differences, [?-x]:= ^x=# source
  • 1 http://hpgl.stanford.edu/publications/Science_2000_v290_p1155.pdf
  • 2 http://www.familytreedna.com/pdf/Levite%20paper.pdf
  • 3 http://www.springerlink.com/content/r60m403330h204l0/
  • 4 http://www.springerlink.com/content/n2883j06628r5515/
  • 9 http://www.springerlink.com/content/w75j6048545350g5/
  • 9 http://www.springerlink.com/content/w75j6048545350g5/
  • 10 http://edoc.ub.uni-muenchen.de/archive/00005868/01/Varzari_Alexander.pdf
  • 11 http://www.ucl.ac.uk/tcga/tcgapdf/capelli-CB-03.pdf, table 1, more data % < 6
  • 13 http://mbe.oxfordjournals.org/cgi/reprint/19/7/1008.pdf
  • 14 http://mbe.oxfordjournals.org/cgi/content/full/22/10/1964/TBL1 + (15'th primary sources?)

Asia

                             N      R1*    R1a1(%)  Sr. Published
                            
Ishkashimi                   25      4     68        5 Spencer Wells,2001
Tajiks                       -             64        6
Tajiks/Khojant               22            64        5 Spencer Wells,2001   
Tajiks/Dushanbe              16            19        5 Spencer Wells,2001   
Tajiks/Samarkand             40            25        5 Spencer Wells,2001   
Kyrgyz                       52      2     63        5 Spencer Wells,2001

Tashkent IE                  69      7     47        ?
India Upper Caste            86      -     45.35     8
Sourasthran                  46      0     39        5 Spencer Wells,2001
Abkhazians                   12      8     33        7 Nasidze,2004
Chenchus (India-Drav.)        -      -     26       12  
Kazan Tatar                  38      3     24        5 Spencer Wells,2001
Saami                        23      9     22        5 Spencer Wells,2001
Dongxiang                    49     <10    28          Wei Wang et al.,2003
Bonan                        47      0     26          Wei Wang et al.,2003
Salar                        52     <10    17          Wei Wang et al.,2003
Iran (Tehran)                24      4      4        5 Spencer Wells,2001
Iran (Tehran)                80      8     20        7 Nasidze,2004 

Iran (Isfahan)               50      0     18        7 Nasidze,2004
Pakistan  ??                 85      1.10  16.47     8 ?
Pakistan                    175      0.57  24.43     8 ?
Pakistan south               91      0     31.87     8 ?
India                       728      0     15.8      8 ?
India                       325      0.3   27       12 ? 
Tuvian                       42      2     14        5 Spencer Wells,2001(*5)
Abazinians                   14      0     14        7 Nasidze,2004(*7)
Turks                        39     31     13        7 Nasidze,2004(*7)
Georgians                    77     10     10        7 Nasidze,2004(*7)
Kurd                         17     29     12        5 Spencer Wells,2001(*5)
Nenets                       54      4     11        5 Spencer Wells,2001(*5)
Syrian                       20     15     10        1

Lebanese                     31      6.4    9.7      1
Turkmen                      37     36      9          ?
Turkmen                      30     37      7        5 Spencer Wells,2001(*5)
Lezgi(S.Caucasus)            12     17      8        7 Nasidze,2004(*7)
Svans                        25      0      8        7 Nasidze,2004(*7)
Azerbaijanians               72     11      7        7 Nasidze,2004(*7)
Armenians                   100     19      6        7 Nasidze,2004(*7)
Armenians                    47     36      9        5 Spencer Wells,2001(*5)
S.Ossetians                  17     12      6        5 Spencer Wells,2001(*5)
Kazaks                       54      6      4        5 Spencer Wells,2001(*5)
Chechenians                  19      0      5        7 Nasidze,2004(*7)
Kallar Dravidian             84      0      4        5 Spencer Wells,2001(*5)
Mongolian                    24      0      4        5 Spencer Wells,2001(*5)
Ossetians (Ardon)            28      0      4        7 Nasidze,2004(*7)
Kazbegi                      25      8      4        7 Nasidze,2004(*7)
India Dravidian (Tribal)    180      -      2.78     8 
Kabardinians                 59      2      2        7 Nasidze,2004(*7)
Lezgi(Dagestan)              25      4      0        7 Nasidze,2004(*7)
Ossetians (Digora)           31      0      0        7 Nasidze,2004(*7)
Rutulians                    24      0      0        7 Nasidze,2004(*7)
Darginians                   26      4      0        7 Nasidze,2004(*7)
Ingushians                   22      0      0        7 Nasidze,2004(*7)
Cambodia                      6      0      0        8 ?
China                       127      0      0        8    
Japan                        23      0      0        8
Siberia                      18      0      0        8 ?

Publications:

  • (*5) http://www.pnas.org/cgi/reprint/98/18/10244.pdf[1]
  • (*6) http://www.journals.uchicago.edu/AJHG/journal/issues/v71n3/023927/023927.web.pdf[18]
  • (*7) http://www.eva.mpg.de/genetics/pdf/Caucasus_big_paper.pdf[22]
  • (*8) http://www.journals.uchicago.edu/AJHG/journal/issues/v78n2/42812/42812.html table 5, 6 & 7[16]
  • (*12) T. Kivisild & all , http://evolutsioon.ut.ee/publications/Kivisild2003b.pdf Fig3 more detailed data for regions, but no caste[23]

Popular culture

Bryan Sykes in his book Blood of the Isles gives (from his fantasy) the populations associated with R1a in Europe the name of Sigurd for a clan patriarch, much as he did for mitochondrial haplogroups in his work The Seven Daughters of Eve.

See also

Relationship to other haplogroups

Human Y-chromosome DNA (Y-DNA) haplogroups

Y-most recent common ancestor
|
A BR
B CR
DE CF
D E C F
G H IJ K
I J L M NO P
N O Q R
Haplogroup R
Haplogroup R1
Haplogroup R1a

Haplogroup R1a1




Haplogroup R1b




Haplogroup R2




References

  1. ^ a b c d e f Wells, RS; Yuldasheva N, Ruzibakiev R, Underhill PA, Evseeva I, Blue-Smith J, Jin L, Su B, Pitchappan R, Shanmugalakshmi S, Balakrishnan K, Read M, Pearson NM, Zerjal T, Webster MT, Zholoshvili I, Jamarjashvili E, Gambarov S, Nikbin B, Dostiev A, Aknazarov O, Zalloua P, Tsoy I, Kitaev M, Mirrakhimov M, Chariev A, Bodmer WF (2001). "The Eurasian Heartland: A continental perspective on Y-chromosome diversity". Proc. Natl. Acad. Sci. U. S. A. 98 (18): 10244–9. PMID 11526236.
  2. ^ a b c d Passarino, G; Cavalleri GL, Lin AA, Cavalli-Sforza LL, Borresen-Dale AL, Underhill PA (2002). "Different genetic components in the Norwegian population revealed by the analysis of mtDNA and Y chromosome polymorphisms". Eur. J. Hum. Genet. 10 (9): 521–9. PMID 12173029.
  3. ^ Wells, Spencer (2002), , Princeton University Press.
  4. ^ Barrier analysis (Alexander Varzari, 5.2.4) show a clear gene barrier along the Vistula: "This finding suggests that across the history the geographic boundary, dividing Southeast Europe from Eastern Europe was more transparent for the reciprocal flows than the boundary between Eastern and Western Europe."
  5. ^ correlated with the "secondary Urheimat" or early Centum dialects; Mallory says (1987, p257): "Perhaps our only recourse is to return to our strict definition of the Proto-Indo-European homeland as where the Indo-European languages were spoken in the period 4500–2500 BC."
  6. ^ European R1a1 measurements (referred to as M17 or Eu19) in Semino et al 2000 read 6.2% to Germans (a 4X drop to Czechs and Slovakians reading 26,7%) and 3.7% to Dutch
  7. ^ The Concise Oxford Dictionary of Archaeology. Copyright © 2002, 2003 by Oxford University Press[1]
  8. ^ a b c d e f g h i j k l m n o p q r Semino, A; Passarino G, Oefner PJ, Lin AA, Arbuzova S, Beckman LE, De Benedictis G, Francalacci P, Kouvatsi A, Limborska S, Marcikiae M, Mika A, Mika B, Primorac D, Santachiara-Benerecetti AS, Cavalli-Sforza LL, Underhill PA (2000). "The Genetic Legacy of Paleolithic Homo sapiens sapiens in Extant Europeans: A Y Chromosome Perspective". Science 290: 1155–59. PMID 11073453.
  9. ^ Pawlowski, R; Dettlaff-Kakol A, Maciejewska A, Paszkowska R, Reichert M, Jezierski G (2002). "Population genetics of 9 Y-chromosome STR loci in Northern Poland". Arch Med Sadowej Kryminol. 52 (4): 261–77. PMID 14669672. (in Polish; English abstract)
  10. ^ a b High-Resolution Phylogenetic Analysis of Southeastern Europe Traces Major Episodes of Paternal Gene Flow Among Slavic Populations; Marijana Perii & ally.
  11. ^ MBE Advance Access originally published online on June 8, 2005 Molecular Biology and Evolution 2005 22(10):1964–75; doi:10.1093/molbev/msi185.
  12. ^ a b c d e f g h i j k l m n o p q Pericic, M; Lauc LB, Klaric IM, Rootsi S, Janicijevic B, Rudan I, Terzic R, Colak I, Kvesic A, Popovic D, Sijacki A, Behluli I, Dordevic D, Efremovska L, Bajec DD, Stefanovic BD, Villems R, Rudan P (2005). "High-resolution phylogenetic analysis of southeastern Europe traces major episodes of paternal gene flow among Slavic populations". Mol. Biol. Evol. 22 (10): 1964–75. PMID 15944443. Haplogroup frequency data in table 1
  13. ^ Bamshad, M; Kivisild T, Watkins WS, Dixon ME, Ricker CE, Rao BB, Naidu JM, Prasad BV, Reddy PG, Rasanayagam A, Papiha SS, Villems R, Redd AJ, Hammer MF, Nguyen SV, Carroll ML, Batzer MA, Jorde LB (2001). "Genetic evidence on the origins of Indian caste populations". Genome Research 11 (6): 994–1004. PMID 11381027.
  14. ^ Cavalli-Sforza, Luigi Luca (1994). The History and Geography of Human Genes. Princeton University Press. ISBN 0-691-08750-4. 
  15. ^ Saha, A; Sharma S, Bhat A, Pandit A, Bamezai R (2005). "Genetic affinity among five different population groups in India reflecting a Y-chromosome gene flow". J. Hum. Genet. 50 (1): 49–51. PMID 15611834.
  16. ^ a b c Sengupta, S; Zhivotovsky LA, King R, Mehdi SQ, Edmonds CA, Chow CE, Lin AA, Mitra M, Sil SK, Ramesh A, Usha Rani MV, Thakur CM, Cavalli-Sforza LL, Majumder PP, Underhill PA (2006). "Polarity and Temporality of High-Resolution Y-Chromosome Distributions in India Identify Both Indigenous and Exogenous Expansions and Reveal Minor Genetic Influence of Central Asian Pastoralists". Am. J. Hum. Genet. 78 (2): 202–21. PMID 16400607.
  17. ^ Polarity and Temporality of High-Resolution Y-Chromosome Distributions in India Identify Both Indigenous and Exogenous Expansions and Reveal Minor Genetic Influence of Central Asian Pastoralists - Sanghamitra Sengupta et al [2] 2005, by The American Society of Human Genetics
  18. ^ a b Zerjal, T; Wells RS, Yuldasheva N, Ruzibakiev R, Tyler-Smith C. (2002). "A Genetic Landscape Reshaped by Recent Events: Y-Chromosomal Insights into Central Asia". Am. J. Hum. Genet. 71 (2): 466–482. 12145751.
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This article is licensed under the GNU Free Documentation License. It uses material from the Wikipedia article "Haplogroup_R1a_(Y-DNA)". A list of authors is available in Wikipedia.
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