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In human genetics, Haplogroup G (M201) is a Y-chromosome haplogroup. It is a branch of Haplogroup F (M89). Haplogroup G has an overall low frequency in most populations but is widely distributed within many ethnic groups of the Old World in Europe, northern and western Asia, northern Africa, the Middle East, India, Sri Lanka and Malaysia.
[edit] OriginsVarious estimated dates and locations have been proposed for the origin of Haplogroup G. The man who developed by chance the M-201 mutation of haplogroup G would not know that anything was different, and it probably took some time before the descendants of this M-201 man dispersed to different locales. The father of the man who mutated belonged to haplogroup F, and other mutations off of haplogroup F may not have then yet occurred. The earliest authoritative comments about the origin of G suggested the mutation took place in the Himalayan foothills in Pakistan or India. While not addressing locations, the 2007 pre-publication of the Y chromosome consortium (YCC) report assigned the origin of G to 20,000 years ago. The National Geographic Society places haplogroup G origins in the Middle East 10-20,000 years ago and presumes that people carrying the haplogroup took part in the spread of the Neolithic [1] Two scholarly papers have also suggested an origin in the Middle East, while differing on the date. Semino et al. suggested 17,000 years ago.[2] Cinnioglu et al. suggested the mutation took place 9,500 years ago.[3] The latter date, however, is probably too recent because a skeleton found in Alaska belonging to haplogroup Q (another haplogroup branching off haplogroup F) was radiocarbon-dated to 9,200 years ago.[4] The oldest skeletons confirmed by ancient DNA testing as haplogroup G men date only from the 7th century C.E. as found in present-day Bavaria, Germany.[5] [edit] Subgroup Tree Listing
The # indicates newly identified in 2009 The five most commonly occurring subgroups of haplogroup G with their corresponding SNPs in parentheses are: G1 (M285), G2a1 (P16) G2a3a (M406), G2a3b1 (P303) and G2c (M377). [edit] Characteristics of Haplogroup G SubgroupsThe International Society of Genetic Genealogy (ISOGG)[6] maintains the most up-to-date version of haplogroup categories. These classifications are based on shared SNP mutations. The discovery of new SNPs can result in assignment of new names to haplogroup categories. There were only a few G categories until 2008 when major revisions to categories were made. Even more G SNPs were identified in 2009 leading to more changes. Until 2008, new G SNPs were reported from labs at the University of Arizona (P designations), Stanford University (M designations) or the University of Central Florida (U designations). Beginning in 2008, additional G SNPs were identified at Family Tree DNA (L designations) and Ethnoancestry (S designations). These latter labs also made use of raw data results reported by individuals tested for about 2,000 SNPs at 23andMe to provide new L or S-designated SNP tests. In 2009, Family Tree DNA's Walk through the Y Project, sequencing certain Y-chromosome segments, provided a number of new G SNPs with the L designation. Because SNPs provide the most reliable method of categorization, they are allowed to represent official G categories with alternating letters and numbers. But unusual values or unusual value combinations found at short tandem repeat markers (STRs) can also provide the basis of additional subgrouping. [edit] G1 (M285+)Main article: Haplogroup G1 (Y-DNA) Almost all haplogroup G1 persons have the value of 12 at short tandem repeat (STR) marker DYS392 and all will have the M285 SNP mutation which characterizes this group. This value of 12 is uncommon in other haplogroups. Subgroups of G1a and G1b exist with several new SNPs within G1 under investigation. The highest reported concentration of G1 and its subgroups in a single country is in Iran, with next most frequent concentrations in neighboring countries to the west. There are distinctive Ashkenazi Jewish and Kazakh G1 subgroups based on STR marker value combinations. They subgroups do not have SNP mutations to identify them. [edit] G2a1 (P16+)Main article: Haplogroup G2a1 (Y-DNA) Haplogroup G2a1 and its one subgroup G2a1a represent the majority of haplogroup G samples in some parts of the Caucasus Mountains area. G2a1 is found only in tiny numbers elsewhere. Almost all G2a1 persons have a value of 10 at STR marker DYS392. They also typically have higher values for DYS385b, such as 16, 17 or 18, than seen in most G persons. The North Ossetians in the mid northern Caucasus area of Russia belong overwhelmingly to the G2a1 subgroup based on available samples. The presence of the SNP P18 mutation characterizes G2a1's only subgroup, G2a1a. The reliability of both P16 and P18 in identifying everyone in each of these categories has been questioned and individual components of the SNP have to be examined. Ashkenazi Jewish G2a1 men with northeastern European ancestry form a distinct cluster based on STR marker values. Men from the Caucasus and men from eastern Europe also form distinctive STR clusters. [edit] G2a2 (M286+)This group is tiny. Samples indicating British Isles and Turkish ancestry have been identified. The British samples have distinctive double values for STR marker DYS19. [edit] G2a3a (M406+)Main article: Haplogroup G2a3a (Y-DNA) G2a3a and its several subgroups seem most commonly found in Turkey and Greece and the costal areas of the eastern Mediterranean where it can constitute up to 50% of haplogroup G samples. G2a3a is more common in southern Europe than northern Europe. In Europe -- except in Greece and Italy -- G2a3a constitutes less than 20% of G samples. G2a3a so far has seldom surfaced in northern Africa or southern Asia, but represents a small percentage of the G population in the Caucasus Mountains region and in Iran. A relatively high percentage of G2a3a persons have a value of 21 at STR marker DYS390. The DYS391 marker has mostly a value of 10, but sometimes 11, in G2a3a persons, and DYS392 is almost always 11. If a sample meets the criteria indicated for these three markers, it is likely the sample is G2a3a. G2a3a has two known subgroups. G2a3a1 is relatively common, but the frequency of G2a3a2 which was first identified in 2009 is unknown. [edit] G2a3b* (L141+)The SNP that defines this group was identified only in mid-2009 at Family Tree DNA, handicapping the ability to describe the features of this group. Initial testing seems to indicate this is a significant G group. Samples from persons with British Isles and Turkish ancestry have been identified. [edit] G2a3b1 (P303+)Main article: Haplogroup G2a3b1 (Y-DNA) The G2a3b1 definable subgroups are heavily concentrated throughout Europe west of the Black Sea and Russia where G2a3b1 is often in the majority among G persons. Small percentages of G2a3b1 are found primarily in the area encompassed by Turkey, the southern Caucasus countries, Iran and the Middle East where the G2a3b1 SNP may have originated. G2a3b1 is also found in India. The largest G2a3b1 subgroup based on available samples is one in which almost all persons have the value of 13 at STR marker DYS388. There are additional subgroups of DYS388=13 men characterized by the presence of specific SNPs or uncommon STR marker oddities. The next largest G2a3b1 subgroup is characterized by the presence of the L13/S13 SNP. This subgroup is most common in north central Europe. The final major subgroup is characterized in a high percentage of cases by the values of 9 at marker DYS568 and 20,21 at marker YCA. This latter DYS568=9 subgroup contains a further large subgroup consisting overwhelmingly of Ashkenazi Jews. The highest percentage of G2a3b1 persons in a discrete population so far described is in the island of Ibiza off the eastern Spanish coast. [edit] G2a4 (L91+)The SNP that characterizes this group (L91) was identified only in spring 2009 at Family Tree DNA. G2a4 would seem to encompass a significant group of G persons, but the recent identification and the paucity of samples are obstacles to describing characteristics of this group. Included within G2a4 are some men with double values for STR marker DYS19, but there are also G2a men with this finding who are not G2a4. The double 19 value situation is not seen in the G2a1 and G2a3 subgroups. [edit] G2c (M377+)Main article: Haplogroup G2c (Y-DNA) A clade of closely related Ashkenazi Jews represent virtually all G2c persons, with just three other G2c haplotypes having been reported so far: one Turk from Kars in northeast Turkey near Armenia, one Pashtun, and one Burusho in Pakistan. The extreme rarity of G2c in northern Pakistan could indicate that G2c in this area originates outside the region and was brought there in the historic period, perhaps from further west (Pakistan 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 Pakistani G2c haplotypes are quite divergent from the Ashkenazi Jewish clade, and therefore do not at all indicate a recent common origin. The Turkish G2c is somewhat closer, but not identical. It remains to be seen if testing will reveal G2c haplotypes in other populations — this is some indication that G2c occurs at low levels in the Near East. Early reports that Ashkenazi G men were all G1 are now proven incorrect. There are also Jewish genetic clades within G2 and G1 whose members are not closely related to the G2c men. All G2c men tested so far have a rare null value for the DYS425 marker, (a missing "T" allele of the DYS371 palindromic STR), the result of a RecLOH event, a finding not yet seen among most other G haplotypes. Among Jews worldwide, Haplogroup G comprises between 10–20% of the population. Though forming some recognizable clades, Jews today comprise a small percentage of the total number of G men worldwide. [edit] Geographical DistributionKnowing the distribution of haplogroup G in general is not as useful as that of the distribution of its subgroups. The subgroups likely spread to new areas of the world in different time periods and to different locations. All available G samples derive from studies or collections that do not meet criteria for random sampling, and conclusions based on them are only rough approximations of what is seen in populations. In Europe west of the Black Sea Haplogroup G is found at about 5% of the population on average throughout most of the continent.[7] The concentration of G falls below this average in Scandinavia, the westernmost former Soviet republics and Poland, as well as in Iceland and the British Isles. There are seeming pockets of unusual concentrations within Europe. In Wales, a distinctive G2a3b1 type (DYS388=13 and DYS592=12) dominates there and pushes the G percentage of the population higher than in England. In western Austria, the G percentage can reach 8% or more. In the northern and highland areas of the island of Sardinia off western Italy), G percentages reach 11% of the population in one study[8] and reached 21% in the town of Tempio in another study. In the Greek island of Crete (approx. 7%[9] to 11%[10]) is G. In Russia, the Ukraine and central Asia, the G percentage is around 1% or less. The northern slope of the Caucasus Mountains represents a major exception where cencentrations in the Kabardinian and Ossetian populations are noted. In Digora, North Ossetia the highest known concentration of G in a single town in the world is reached where 74% of the tested men were G.[11] The Madyars of Kazakhstan were found to be 87% G1. Haplogroup G is also found in 12% of sampled Pashtun.[12] Haplogroup G is found as far east as the Han population of China in small numbers where G can reach 2%. In Turkey, the southern Caucasus region and Iran, haplogroup G reaches the highest percentage of a regional population worldwide. Among Turkish males 11% of the population is G.[3] In Iran, Haplogroup G reaches 13 to 15% of the population in various parts of the country. While it is found in percentages higher than 10% among the Bakhtiari, Gilaki and Mazandarani, it is closer to 5% among the Iranian Arabs and in some large cities.[13] Among the samples in the YHRD database from the southern Caucasus countries, 29% of the samples from Abazinia, 31% from Georgia, 18% from Azerbaijan and 11% from Armenia appear to be G samples. In southern Asia, haplogroup G is found in concentrations of approximately 20%[14] of Kalash and approximately 16% of Brahui,[14] but in very small percentages overall in Pakistan and India. The many groups in India and Bangla Desh have not been well studied. About 6% of the samples from Sri Lanka and Malaysia were reported as haplogroup G, but none were found in the other coastal lands of the Indian Ocean or Pacific Ocean in Asia.[15] In the Middle East, haplogroup G accounts for about 3% of the population in almost all areas.[16][17] Among the Druze mostly residents of Israel 10% were found to be haplogroup G.[18] In Africa, haplogroup G is rarely found in sub-Saharan Africa or south of the horn of Africa among native populations. In Egypt, studies have provided information that pegs the G percentage there to be between 6%.[19] and 10%.[20] Three percent of North African Berbers were found to be haplogroup G.[21] Two percent of Arab Moroccans and 8% of Berber Moroccans were likewise found to be G.[22] In the Americas, the percentage of haplogroup G corresponds to the numbers of persons from Old World countries who emigrated. It is not found among Native Americans except where intermarriage with non-native persons has occurred. Around 10% of Jewish males have Haplogroup G. [edit] Migrations in the Historical PeriodHaplogroup G seems to have expanded from southwestern Asia or the Middle East to Europe west of the Black Sea and to the western Mediterranean and to central Asia primarily about 1,700 to 3,000 years ago. The time period is based on time calculations derived by comparing the number of STR marker mutations among men in these three areas and the presumed region of origin. This region of origin for the major G groups of today was most likely somewhere in Turkey, the Caucasus Mountains region, Iran or the eastern Mediterranean where G is found both in significant numbers and with the diversity expected with a long presence. The principal spread of G from the mentioned region was to Europe west of the Black Sea. As to the most common G groups there, their (a) presence in 4% or more of the population there plus (b) seeming ancestral splits (based on mutations in 67-marker STR samples) between persons in southern and northern Europe no more recently than the early Middle Ages suggest the G ancestors came as part of identifiable groups (or s group) rather than as isolated movements of individuals. There are Jewish components of the major groups which require explanation when advocating any particular ancestral group for these major G groups now in Europe. The likeliest geographical sources for spread of each of the following principal G subgroups is: G1 -- Iran or eastern Middle East...... G2a1 -- the northern or southern Caucasus region..... G2a3a -- Turkey or the eastern Mediterrean...... G2a3b1 (Caucasus type) -- the northern or southern Caucasus..... G2a3b1 (DYS388=13 type) -- perhaps Iran or southern Caucasus region..... G2a3b1 (DYS568=9 type -- unknown..... G2a3b1a1 -- Iran or southern Caucasus region...... G2c -- northern Middle East [edit] North of the Caucasus MountainsIf G migrations originated in the area north of the Caucasus Mountains, surviving traces of an original G population in today's Ukraine and almost all of Russia are few. The exceptions today are in Russia along the north slope of the Caucasus where G2a3b1 in common among the Kabardinians, G2a1 is common among the Ossetians, and an unknown G type is found in Dagestan Other G types are found in small numbers in these three areas. The variety of G types there could indicate a long G presence in the northern Caucasus if G did not arrive piecemeal from southern locations a long time after the formation of each of these G types. There were two major groups immediately north of the Caucasus area in the period of interest who engaged in migrations: the Sarmatians and the Huns. Both groups have cultural similarities, such as manipulating infant head shapes. One after another, the two moved into the area north of the Caucasus from the east when a westward movement provided better options than resisting groups to the east. The Sarmatians subsumed an existing population of Scythians during their occupation of this area. Like the Scythians, the Sarmatians spoke an Iranian language. The more central portion of the Sarmatians were identified as Alan Sarmatians. Strabo claimed these groups could assemble hundreds of thousands of warriors. [23] Being nomadic with a history of moving en masse, it is conceivable that the Sarmatians when they began the Great Migration westward in the 1st century B.C.E. took with them a high percentage of their population. The Huns were also nomadic and had been as far east as central Asia. Their language is not well defined. Complicating the discovery of the genetic make-up of these groups, scholars have described both Sarmatians and Huns as confederations of groups rather than solitary ethnic groups. Greek historian Diodorus Siculus related that the Scythians had carried Medes to the River Don north of the Caucasus presumably from today's northwestern Iran, this event giving rise to their name Sauromatians.[24] Scythians had occupied Media 653-625 B.C.E. Pliny also suggested Sarmatians descended from Medes. [25]. But historians have difficulty explaining how the Sarmatians instead seemed to come from central Asia off to the east in their occupation of Scythian lands around the 5th century B.C.E. The Huns in their westward migration had reached today's Ukraine by the 100s C.E. replacing the Sarmatians who left, and they moved dramatically into more western lands in the 400s. Under Hun rule, some Alans who had not migrated remained in the area north of the Caucasus. In the Middle Ages, invading Mongols reportedly forced them down to the Caucasus area. But it is also known that 30,000 Alans formed the royal guard of the Yuan court in Dadu (Beijing, China). While it is conceivable that many of the G persons in Europe proper today had ancestors part of the Great Migration that carried large populations out of southwestern Russia, they left few close relatives in the area they vacated if so. G2a1 which predominates among the North Ossetians is rare in Europe proper. The Caucausus G2a3b1 common among the Kabardinians (DYS390=23, DYS388=12, DYS385=13,14), for example, is seen only rarely in Europe proper where the DYS388=13 type (DYS390=22, DYS385=14,15) predominates. In addition, none of the ancient DNA recovered so far from skeletons in the Scythian/Sarmatian area of occupation has been haplogroup G. While these mentioned northern Caucasus types of G are uncommon in lands to the north, they have significant equivalents in the southeastern Caucasus, particularly in Georgia, including South Ossetia which is often included with Georgia. If these persons in the north Caucasus were originally Medes who became Sauromatians (later Sarmatians and Alans) when carried to the north Caucasus by the Scythians, the G groups in the land of the Kingdom of the Medes in northwestern Iran, and also the rest of Iran, are different from what is seen in the northern Caucasus. G1, for example, is common in Iran, but rare in the north Caucasus. The DYS388=13 G2a3b1 and G2a3b1a1 groups are found in Iran but seemingly not in the north Caucasus. [edit] South of the Caucasus MountainsIn contrast to the regional types of G seen north of the Caucasus Mountains, the three countries of Georgia, Armenia and Azerbaijan provide STR samples consistent with all types of haplogroup G seen today in Europe west of the Black Sea with the exception of G1 and G2c. And the regional STR types north of the Caucasus are seen in smaller numbers south of the Caucasus. Because of the rich diversity of these G types, this area of the world is a leading candidate for the starting point for most G migrations especially those to western and central Europe. The STR marker samples in the YHRD database[26] from these three countries obtained from the Nasidze study,[27] as well as samples in the Haplogroup G Project,[28] contain examples of (1) G2a1, (2) G2a3a, (3) G2a3b1 (DYS388=13 type), and (4) G2a3b1a1. These four G types represent over 90% of G types found in western and central Europe. The three countries also have the north Caucasus type of G2a3b1* seen especially among the Kabardinians (DYS390=23, DYS388=12, DYS385=13,14) ). Both the Babylonian Empire and Persian Empire once included the southern Caucasus and stretched to the Mediterranean, but there is no seeming report of groups from the Caucasus being relocated to the western limits of these empires. But the Assyrians deported Jews to Armenia, and 10,000 Jews moved there on their own. The commonly accepted view is that the Mountain Jews of Azerbaijan were early medieval immigrants from Persia and this immigration continued throughout the Middle Ages. One possible source of a G migration would be men recruited into military units during the Roman Empire. Armenia which encompassed nearly all the southern Caucasus region was occupied for a while in the 100s C.E. None of the Roman legions bore the name of locations in the southern Caucasus. Other possible sources of southern Caucasus G migrations were the Jewish communities. There are significant Jewish subgroups within G2a3a, G2a1 and certain types of G2a3b1. The Jewish Radhanites had significant presences in western and central Europe in the early Medieval period with trade routes that included the eastern shore of the Caucasus together with some known Jewish communities in the Caucasus region. One problem with identifying the southern Caucasus as the source of most of the G in western and central Europe is the minority status there of the DYS388=13 type of G2a3b1. This group is dominant in Europe but its rarity elsewhere actually makes no area the most logical location as a source for its migration. [edit] IranIn Iran G1 is found in significant numbers in all tested areas. The Persian Empire once extended to the Mediterranean, and Persian military forces may have carried G persons into the southern Caucasus as well as to the shores of the Mediterranean. In fact, the highest percentages of G1 outside of Iran are in the countries just to the west of southern Iran that were once part of the Persian Empire. If the G movement occurred instead before the Persian Empire, G1 could have originated either in the Middle East or Iran. While G2a3a occurs in Iran, it is apparently too rare there to account for any source group for G2a3a persons. The same situation applies to G2a1 there. G2a3b1 Iranian samples are available, especially G2a3b1a1, and several unclassified types of G2a and G2a3 are noted. Because the DYS388=13 type of G2a3b1 dominates G samples in western central Europe, the existence of a few such samples in Iran make it possible Iran was a source for these, but there is no Iranian concentration of such G types needed to argue for this. Neither G2c nor the Caucasus type of G2a3b1 samples have surfaced among Iranians. The Jews trace their heritage in Iran to the Babylonian Exile of the 6th century B.C.E. The Sassanids temporarily suppressed Judaism in the early centuries of the Current Era and may have perhaps caused some Jewish emigration out of Persia. Because there are significant Jewish components to G1 and G2a3b1 types, the Jewish Radhanites could also have spread these G types in their travels in the early Middle Ages. They maintained a significant presence on the trade routes that connected Iran to the rest of the world. [edit] Turkey and the eastern MediterraneanThis region has unusual percentages of G2a3a. Beginning with the Phoenicians, any of the trading and conquering governments of the Mediterranean could have spread G2a3a to the western Mediterranean where it is more common than in Europe to the north. G2a3a also has a Jewish-Italian subgroup, and any of the Jewish diaspora movements may have contributed to the spread. The percentages of G2a1 and the DYS388=13 type of G2a3b1 are so tiny in this area, that it is unlikely to be the starting point for a migration of these G groups. G1 is common in the northern Middle East, and that area may have been the source of G movements to Iran or vice versa. G2c is found in small numbers in the northern Middle East, and the Middle East could have been the source from which European G2c and Pakistani G2c1 derived. [edit] OtherWithin the Old World, due to the rarity of samples it is currently unclear how G2a and G2c spread into Uzbekistan, Kazakhstan, Pakistan, southern India, Sri Lanka, Malaysia and northern China. Haplogroup G was carried to the New World beginning in the 1500s by individual migrations principally from the British Isles, Germany, Iberia and Italy. [edit] Famous membersJoseph Stalin, from a genetic test on his grandson (his son Vasily's son; Alexander Burdonsky) shows his Y-DNA haplogroup to be G2a1a [1]
[edit] See also[edit] References
[edit] External links
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