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[[Image:Haplogroup G5 (Y-DNA) distribution.jpg|left|thumb|325px|Map of the distribution of Y-DNA haplogroup G5 in Eastern Europe showing the distribution of G5 within the borders of the historic Polish-Lithuanian Commonwealth.]]
[[Image:Haplogroup G5 (Y-DNA) distribution.jpg|left|thumb|325px|Map of the distribution of Y-DNA haplogroup G5 in Eastern Europe showing the distribution of G5 within the borders of the historic Polish-Lithuanian Commonwealth.]]
|[[Image:Rzeczpospolita.png|right|thumb|400px|Map of the Polish-Lithuanian Commonwealth, 1569-1765.
|[[Image:Rzeczpospolita.png|right|thumb|400px|Map of the Polish-Lithuanian Commonwealth, 1569-1765.
The areas eventually occupied by Jews exclude the light pink regions to the north and east which were lost to the [[Cossacks]] and then Russia during [[Khmelnytsky_Uprising#Jewish | Khmelytsky's Pogrom]]. The Jews in these regions were massacred or driven out. The more westerly striped region was [[East Prussia]], where Jews were generally not allowed to reside before the late 17th century, and later only under onerous tax restrictions.<ref> {{cite web|url=http://www.jewishencyclopedia.com/view.jsp?artid=570&letter=P#2131 |title=Jewish Encyclopedia (1906): Prussia - Under the Hohenzollerens |accessdate=2007-09-16 | first=Gotthard | last=Deutsch |date=1906 }}</ref>. The northern striped regions were [[Courland ]] and [[Livonia]] from which the Jews were excluded before 1670, and were expelled several times, first in 1714 finally and permanently in 1760.<ref>{{cite web|url=http://www.jewishencyclopedia.com/view.jsp?artid=831&letter=C&search=Courland#2839|title=Jewish Encyclopedia (1906): Courland|accessdate=2007-09-16 | first=Herman | last=Rosenthal |date=1906 }}</ref>]]
The areas eventually occupied by Jews exclude the light pink regions to the north and east which were lost to the [[Cossacks]] and then Russia during [[Khmelnytsky_Uprising#Jewish | Khmelytsky's Pogrom]]. The Jews in these regions were massacred or driven out. The more westerly striped region was [[East Prussia]], where Jews were generally not allowed to reside before the late 17th century, and later only under onerous tax restrictions.<ref> {{cite web|url=http://www.jewishencyclopedia.com/view.jsp?artid=570&letter=P#2131 |title=Jewish Encyclopedia (1906): Prussia - Under the Hohenzollerens |accessdate=2007-09-16 | first=Gotthard | last=Deutsch |date=1906 }}</ref>. The northern striped regions were [[Courland ]] and [[Livonia]] from which the Jews were excluded before 1670, and were expelled several times, first in 1714 finally and permanently in 1760.<ref>{{cite web|url=http://www.jewishencyclopedia.com/view.jsp?artid=831&letter=C&search=Courland#2839| title=Jewish Encyclopedia (1906): Courland|accessdate=2007-09-16 | first=Herman | last=Rosenthal |date=1906 }}</ref>]]
|}
|}

===== Western Germany =====
G5 is also found among Ashkenazi Jews from Western Germany. Jews were not allowed to reside in most parts of Germany in the 16th and 17th centuries, aside from the [[Frankfurter Judengasse | Frankfurt Jewish Ghetto]]. Jews were expelled in 1670 from Vienna and the Archduchy of Austria. <ref>{{cite web|url=http://www.jewishencyclopedia.com/view.jsp?artid=2152&letter=A&search=Austria#6712 | title=Jewish Encyclopedia (1906): Austria - From the Expulsion of 1420 to that of 1670 | accessdate=2007-09-16 | first=Gotthard | last=Deutsch |date=1906 }}</ref> After [[Khmelnytsky_Uprising#Jewish | Khmelytsky's Pogrom]] in Poland in 1648, there began a migration of Jews from Poland and Lithuania to Western Germany, which accelerated and continued into the 19th century. It isn't clear at this point whether German Jewish G5s represent an independent settlement, or the result of a migration from Eastern Europe (although there is some evidence for the latter).


===== Southern Italy and Sicily =====
===== Southern Italy and Sicily =====

Revision as of 06:37, 17 September 2007

In human genetics, Haplogroup G5 is a Y-chromosome haplogroup and is defined by the presence of the M377 mutation.[1] It is a branch of Haplogroup G (which is in turn defined by the presence of the M201 mutation).

Phylogenetic position

Extensive SNP testing by the Haplogroup G SNP project found that G5 is an independent branch of haplogroup G characterized by only one SNP, M377. A forthcoming study by the Y Chromosome Consortium at the University of Arizona found that within haplogroup G, G5 and G2 share a new SNP, P387, which is not found in the other well-attested branch of G, haplogroup G1. As a result, it will be proposed that Haplogroup G5 be renamed "G2c".

A study of median-joining networks[2] generated for a set of approximately 180 67 Y-STR haplotypes in haplogroup G shows that G5 might cluster most closely with sub-haplogroup G2a.

Y-STR haplotype characteristics

All G5 samples tested so far have a null value for the DYS425 marker, (a missing "T" allele of the DYS371 palindromic STR), the result of a RecLOH event. This change is extremely uncommon in the rest of haplogroup G, but apparently happened early in the history of G5.

G5 Y-STR distinguishing allele values
Y-STR Allele range Modals
DYS393 12-13 13
DYS390 23-24 23
DYS19 15-17 15
17
DYS391 10-11 10
11
DYS385 13,15
13,16
14,16		 13,16
DYS426 11
DYS388 12
DYS439 11-12 11
DYS392 11
DYS389 13,30
13,31
14,31
14,32
14,33
15,33		 14,32
DYS459 8,9
DYS464 13,13,14,15
13,13,15,15
13,14,14,15
13,14,15,15		 13,14,15,15
DYS395S1a 16,16
DYS425 null
DYS413 21,22
DYS436 11
DYS481 19

Distribution

Ashkenazi Jews

A cluster of closely related Ashkenazi Jews represent virtually all confirmed G5 persons worldwide, both from private testing, and from academic studies. G5 makes up about 7% of all Ashkenazi Jewish Y chromosome haplotypes, as was found in Behar et al. (2004) (n=442, GxG2=33).[3] A much smaller group of Ashkenazi Jews however are in haplogroup G1, so not all GxG2 Ashkenazi Jews in the above study would be G5. The ratio of G5:G1 among Ashkenazim is approximately 10:1.

Eastern Europe

The distribution of G5 among Ashkenazi Jews in Eastern Europe very closely reflects the 16th and 17th century settlement patterns of Jews in the Polish-Lithuanian Commonwealth:

File:Haplogroup G5 (Y-DNA) distribution.jpg
Map of the distribution of Y-DNA haplogroup G5 in Eastern Europe showing the distribution of G5 within the borders of the historic Polish-Lithuanian Commonwealth.
Map of the Polish-Lithuanian Commonwealth, 1569-1765. The areas eventually occupied by Jews exclude the light pink regions to the north and east which were lost to the Cossacks and then Russia during Khmelytsky's Pogrom. The Jews in these regions were massacred or driven out. The more westerly striped region was East Prussia, where Jews were generally not allowed to reside before the late 17th century, and later only under onerous tax restrictions.[4]. The northern striped regions were Courland and Livonia from which the Jews were excluded before 1670, and were expelled several times, first in 1714 finally and permanently in 1760.[5]
Western Germany

G5 is also found among Ashkenazi Jews from Western Germany. Jews were not allowed to reside in most parts of Germany in the 16th and 17th centuries, aside from the Frankfurt Jewish Ghetto. Jews were expelled in 1670 from Vienna and the Archduchy of Austria. [6] After Khmelytsky's Pogrom in Poland in 1648, there began a migration of Jews from Poland and Lithuania to Western Germany, which accelerated and continued into the 19th century. It isn't clear at this point whether German Jewish G5s represent an independent settlement, or the result of a migration from Eastern Europe (although there is some evidence for the latter).

Southern Italy and Sicily

Among Europeans, there are a few significant exceptions to this almost exclusive Ashkenazi Jewish distribution - there are a small handful of samples from Southern Italians, or people with traditions of patrilineal descent from 15th century Sicily.

The Hindu Kush and Kashmir (Pakistan)

There are just two other confirmed G5 samples that have been reported in the academic literature so far, one Pashtun in the Northwest Frontier Provinces of Pakistan (the Hindu Kush Range), and one Burusho in the Hunza Valley in Kashmir. These two G5s are Y-STR haplotypes 731 and 794 from Table 3 in the study by Sengupta et al. (2006) of Indian (n=728), Pakistani (n=176), and East Asian (n=175) Y chromosome lineages.[1]

Other possible G5 Jewish haplotypes

Two possible G5 Y-STR haplotype samples in the literature are from the study of Jewish and non-Jewish Near Eastern Y chromosomes by Nebel et al. (2001) (in the Appendix Table A1), haplotype 51 which was found in 1 Ashkenazi Jew (n=79) and 3 Kurdish Jews (n=99), and haplotype 47 which was found in 1 Iraqi Jew (n=23). These belong to what was termed at the time "Haplogroup 2", which consisted of Y haplogroups F*, G, and I,[7] and within this set of haplogroups these display a Y-STR allele pattern unique to haplogroup G5.[8]

Possible Eastern Anatolian G5

The one other possible G5 Y-STR haplotype found in the literature is haplotype 54 from a study of Anatolian Y chromosomes (n=523) by Cinnioglu et al. (2004) which was found in Eastern Turkey.[9]

Major regions where G5 is not found

There is a good deal of negative evidence that shows where G5 is not found: A large study of Iranian (n=150) Y chromosomes by Regueiro et. al (2006) failed to find any G5s,[10] a study of hundreds of Iberian (n=860) and North African Berber (n=75) Y chromosomes by Alonso et al. (2005) found not a single G5,[11], and likewise a study of samples from Amman (n=101) and the Dead Sea area (n=45) in Jordan by Flores et al. (2005) also failed to find any G5.[12]

Upcoming studies

Other Y chromosome samples taken from an upcoming study of Sephardi and Near Eastern (Mizrahi) Jews have found only a few GxG2 (in Y chromosome haplogroup G but not in G2) samples. However, these include among others a couple of Turkish Jews, and single [[Moroccan Jews | Moroccan], Kurdish, Iraqi, and Yemenite Jewish samples. These are being tested for M377/G5.

The Kurdish and Iraqi Jewish samples from Nebel et al. (2002) are also being tested for M377/G5 by a different group for another study. Y chromosome haplogroup G1 is also found among Jewish populations, but it is likely that some of these will turn out to be in haplogroup G5.

Time to the Most Recent Common Ancestor (tMRCA)

Possible history

The preliminary conclusions from this evidence is that haplogroup G5 is not native to Europe, [TODO}

The rarity of G5 in northeast Pakistan could indicate that G5 in this area originates outside the region and was brought there in the historic period, perhaps from further west (this area was part of both the Achaemenid Persian Empire, conquered by Alexander the Great, and then formed a part of the Greco-Bactrian Kingdom). These two reported G5 haplotypes are quite divergent from the Ashkenazi Jewish clade, and therefore do not at all indicate a recent common origin.

References

  1. ^ a b Sengupta S, Zhivotovsky LA, King R; et al. (2006). "For the primer sequence of M377 see: 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 (Table 2)". Am. J. Hum. Genet. 78 (2): 202–21. doi:10.1086/499411. PMID 16400607. {{cite journal}}: Explicit use of et al. in: |author= (help)CS1 maint: multiple names: authors list (link) Cite error: The named reference "pmid16400607" was defined multiple times with different content (see the help page).
  2. ^ Bandelt HJ, Forster P, Röhl A (1999). "Median-joining networks for inferring intraspecific phylogenies". Mol. Biol. Evol. 16 (1): 37–48. PMID 10331250.{{cite journal}}: CS1 maint: multiple names: authors list (link)
  3. ^ Behar DM, Garrigan D, Kaplan ME; et al. (2004). "Contrasting patterns of Y chromosome variation in Ashkenazi Jewish and host non-Jewish European populations (Table 2)". Hum. Genet. 114 (4): 354–65. doi:10.1007/s00439-003-1073-7. PMID 14740294. {{cite journal}}: Explicit use of et al. in: |author= (help)CS1 maint: multiple names: authors list (link)
  4. ^ Deutsch, Gotthard (1906). "Jewish Encyclopedia (1906): Prussia - Under the Hohenzollerens". Retrieved 2007-09-16.
  5. ^ Rosenthal, Herman (1906). "Jewish Encyclopedia (1906): Courland". Retrieved 2007-09-16.
  6. ^ Deutsch, Gotthard (1906). "Jewish Encyclopedia (1906): Austria - From the Expulsion of 1420 to that of 1670". Retrieved 2007-09-16.
  7. ^ Y Chromosome Consortium (2002). "YCC NRY Tree 2002 v2002.01.18". Retrieved 2007-09-16.
  8. ^ Nebel A, Filon D, Brinkmann B, Majumder PP, Faerman M, Oppenheim A (2001). "The Y chromosome pool of Jews as part of the genetic landscape of the Middle East (Appendix Table A1)". Am. J. Hum. Genet. 69 (5): 1095–112. PMID 11573163.{{cite journal}}: CS1 maint: multiple names: authors list (link)
  9. ^ Cinnioğlu C, King R, Kivisild T; et al. (2004). "Excavating Y-chromosome haplotype strata in Anatolia. Haplotype 54 was found in Region 4 on the map in Figure 1". Hum. Genet. 114 (2): 127–48. doi:10.1007/s00439-003-1031-4. PMID 14586639. {{cite journal}}: Explicit use of et al. in: |author= (help)CS1 maint: multiple names: authors list (link)
  10. ^ Regueiro M, Cadenas AM, Gayden T, Underhill PA, Herrera RJ (2006). "Iran: tricontinental nexus for Y-chromosome driven migration". Hum. Hered. 61 (3): 132–43. doi:10.1159/000093774. PMID 16770078.{{cite journal}}: CS1 maint: multiple names: authors list (link)
  11. ^ Alonso S, Flores C, Cabrera V; et al. (2005). "The place of the Basques in the European Y-chromosome diversity landscape". Eur. J. Hum. Genet. 13 (12): 1293–302. doi:10.1038/sj.ejhg.5201482. PMID 16094307. {{cite journal}}: Explicit use of et al. in: |author= (help)CS1 maint: multiple names: authors list (link)
  12. ^ Flores C, Maca-Meyer N, Larruga JM, Cabrera VM, Karadsheh N, Gonzalez AM (2005). "Isolates in a corridor of migrations: a high-resolution analysis of Y-chromosome variation in Jordan". J. Hum. Genet. 50 (9): 435–41. doi:10.1007/s10038-005-0274-4. PMID 16142507.{{cite journal}}: CS1 maint: multiple names: authors list (link)

See also

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