Genetic history of Europe
The genetic history of Europe includes information around the formation, ethnogenesis, and other DNA-specific information about populations indigenous, or living in Europe.
The most significant recent dispersal of modern humans from
As a result of the population movements during the Mesolithic to Bronze Age, modern European populations are distinguished by differences in WHG, EEF and Ancient North Eurasian (ANE) ancestry.[12][13][14]
Admixture rates varied geographically; in the late Neolithic, WHG ancestry in farmers in Hungary was at around 10%, in Germany around 25% and in Iberia as high as 50%.
Ethnogenesis of the modern
expansions.Research into the genetic history of Europe became possible in the second half of the 20th century, but did not yield results with high resolution before the 1990s. In the 1990s, preliminary results became possible, but they remained mostly limited to studies of mitochondrial and Y-chromosomal lineages. Autosomal DNA became more easily accessible in the 2000s, and since the mid-2010s, results of previously unattainable resolution, many of them based on full-genome analysis of ancient DNA, have been published at an accelerated pace.[16][17]
Prehistory
Due to natural selection, the percentage of Neanderthal DNA in ancient Europeans gradually decreased over time. From 45,000 BP to 7,000 BP, the percentage dropped from around 3–6% to 2%.[17] The removal of Neanderthal-derived alleles occurred more frequently around genes than other parts of the genome.[17]
Palaeolithic
There has also been speculation about the inheritance of specific genes from Neanderthals. For example, one
Upper Paleolithic
It is thought that modern humans began to inhabit Europe during the Upper Paleolithic about 40,000 years ago. Some evidence shows the spread of the Aurignacian culture.[27]: 59
From a purely patrilineal,
may be those with the oldest presence in Europe. They have been found in some very old human remains in Europe. However, other haplogroups are far more common among living European males because of later demographic changes.Currently the oldest sample of Haplogroup I (M170), which is now relatively common and widespread within Europe, has been found to be Krems WA3 from Lower Austria dating back to about 30–31,000 ybp.[28] At about this time, an Upper Palaeolithic culture also appeared, known as the Gravettian.[29]
Earlier research into Y-DNA had instead focused on haplogroup R1 (M173): the most populous lineage among living European males; R1 was also believed to have emerged ~ 40,000 BP in Central Asia.[29][30] However, it is now estimated that R1 emerged substantially more recently: a 2008 study dated the most recent common ancestor of haplogroup IJ to 38,500 and haplogroup R1 to 18,000 BP. This suggested that haplogroup IJ colonists formed the first wave and haplogroup R1 arrived much later.[31]
Thus the genetic data suggests that, at least from the perspective of patrilineal ancestry, separate groups of modern humans took two routes into Europe: from the Middle East via the Balkans and another from Central Asia via the Eurasian Steppe, to the north of the Black Sea.
Martin Richards et al. found that 15–40% of extant mtDNA lineages trace back to the Palaeolithic migrations (depending on whether one allows for multiple founder events).[32] MtDNA haplogroup U5, dated to be ~ 40–50 kYa, arrived during the first early upper Palaeolithic colonisation. Individually, it accounts for 5–15% of total mtDNA lineages. Middle U.P. movements are marked by the haplogroups HV, I and U4. HV split into Pre-V (around 26,000 years old) and the larger branch H, both of which spread over Europe, possibly via Gravettian contacts.[29][33]
Haplogroup H accounts for about half the gene lines in Europe, with many subgroups. The above mtDNA lineages or their precursors, are most likely to have arrived into Europe via the Middle East. This contrasts with Y DNA evidence, whereby some 50%-plus of male lineages are characterised by the R1 superfamily, which is of possible central Asian origin.[citation needed] Ornella Semino postulates that these differences "may be due in part to the apparent more recent molecular age of Y chromosomes relative to other loci, suggesting more rapid replacement of previous Y chromosomes. Gender-based differential migratory demographic behaviors will also influence the observed patterns of mtDNA and Y variation"[citation needed].
Last Glacial Maximum
The Last Glacial Maximum ("LGM") started c. 30 ka BCE, at the end of
- Northern Iberia and Southwest France, together making up the "Franco-Cantabrian" refugium
- The Balkans
- Ukraine and more generally the northern coast of the Black Sea[29]
- Italy.[34]
This event decreased the overall genetic diversity in Europe, a "result of drift, consistent with an inferred population bottleneck during the Last Glacial Maximum".[30] As the glaciers receded from about 16,000–13,000 years ago, Europe began to be slowly repopulated by people from refugia, leaving genetic signatures.[29]
Some Y haplogroup I clades appear to have diverged from their parental haplogroups sometime during or shortly after the LGM.[35]
Cinnioglu sees evidence for the existence of an Anatolian refuge, which also harboured Hg R1b1b2.[36] Today, R1b dominates the y chromosome landscape of western Europe, including the British Isles, suggesting that there could have been large population composition changes based on migrations after the LGM.
Semino, Passarino and Pericic place the origins of haplogroup R1a within the Ukrainian
From an mtDNA perspective, Richards et al. found that the majority of mtDNA diversity in Europe is accounted for by post-glacial re-expansions during the late upper Palaeolithic/ Mesolithic. "The regional analyses lend some support to the suggestion that much of western and central Europe was repopulated largely from the southwest when the climate improved. The lineages involved include much of the most common haplogroup, H, as well as much of K, T, W, and X." The study could not determine whether there were new migrations of mtDNA lineages from the near east during this period; a significant input was deemed unlikely.[32]
The alternative model of more refugees was discussed by Bilton et al.[39]
From a study of 51 individuals, researchers were able to identify five separate genetic clusters of ancient Eurasians during the LGM: the
From around 37,000 years ago, all ancient Europeans began to share some ancestry with modern Europeans.[17] This founding population is represented by GoyetQ116-1, a 35,000 year old specimen from Belgium.[17] This lineage disappears from the record and is not found again until 19,000 BP in Spain at El Mirón, which shows strong affinities to GoyetQ116-1.[17] During this interval, the distinct Věstonice Cluster is predominant in Europe, even at Goyet.[17] The re-expansion of the El Mirón Cluster coincided with warming temperatures following the retreat of the glaciers during the Last Glacial Maximum.[17] From 37,000 to 14,000 years ago, the population of Europe consisted of an isolated population descended from a founding population that didn't interbreed significantly with other populations.[40]
Mesolithic
Around 14,000 years ago, the
Neolithic
A big cline in genetic variation that has long been recognised in Europe seems to show important dispersals from the direction of the Middle East. This has often been linked to the spread of farming technology during the Neolithic, which has been argued to be one of the most important periods in determining modern European genetic diversity.
The Neolithic started with the introduction of farming, beginning in SE Europe approximately 10,000–3000 BCE, and extending into NW Europe between 4500 and 1700 BCE. During this era, the
- In a late European Mesolithic prelude to the Neolithic, it appears that Near Eastern peoples from areas that already had farming, and who also had sea-faring technology, had a transient presence in Greece (for example at Franchthi Cave).[44][45]
- There is consensus that agricultural technology and the main breeds of animals and plants which are farmed entered Europe from somewhere in the area of the Fertile Crescent and specifically the Levant region from the Sinai to Southern Anatolia.[27]: 1143, 1150 [46] (Less certainly, this agricultural revolution is sometimes argued to have in turn been partly triggered by movements of people and technology coming across the Sinai from Africa.) For more see Fertile Crescent: Cosmopolitan diffusion.
- A later stage of the Neolithic, the so-called Pottery Neolithic, saw an introduction of pottery into the Levant, Balkans and Southern Italy (it had been present in the area of modern Sudan for some time before it is found in the Eastern Mediterranean, but it is thought to have developed independently), and this may have also been a period of cultural transfer from the Levant into the Balkans.
An important issue regarding the genetic impact of neolithic technologies in Europe is the manner by which they were transferred into Europe. Farming was introduced by a significant migration of farmers from the Near East (Cavalli-Sforza's biological
Martin Richards estimated that only 11% of European mtDNA is due to immigration in this period, suggesting that farming was spread primarily due to being adopted by indigenous Mesolithic populations, rather than due to immigration from Near East. Gene flow from SE to NW Europe seems to have continued in the Neolithic, the percentage significantly declining towards the British Isles. Classical genetics also suggested that the largest admixture to the European Paleolithic/Mesolithic stock was due to the Neolithic revolution of the 7th to 5th millennia BCE.[47] Three main mtDNA gene groups have been identified as contributing Neolithic entrants into Europe: J, T1 and U3 (in that order of importance). With others, they amount up to around 20% of the gene pool.[32]
In 2000, Semino's study on Y DNA revealed the presence of haplotypes belonging to the large clade
Concerning timing the distribution and diversity of V13 however, Battaglia[51] proposed an earlier movement whereby the E-M78* lineage ancestral to all modern E-V13 men moved rapidly out of a Southern Egyptian homeland and arrived in Europe with only Mesolithic technologies. They then suggest that the E-V13 sub-clade of E-M78 only expanded subsequently as native Balkan 'foragers-cum-farmers' adopted Neolithic technologies from the Near East. They propose that the first major dispersal of E-V13 from the Balkans may have been in the direction of the Adriatic Sea with the Neolithic Impressed Ware culture often referred to as Impressa or Cardial,[37] rather propose that the main route of E-V13 spread was along the Vardar-Morava-Danube river 'highway' system.
In contrast to Battaglia, Cruciani[52] tentatively suggested (i) a different point where the V13 mutation happened on its way from Egypt to the Balkans via the Middle East, and (ii) a later dispersal time. The authors proposed that the V13 mutation first appeared in western Asia, where it is found in low but significant frequencies, whence it entered the Balkans sometime after 11 kYa. It later experienced a rapid dispersal which he dated to c. 5300 years ago in Europe, coinciding with the Balkan Bronze Age. Like Peričic et al. they consider that "the dispersion of the E-V13 and J-M12 haplogroups seems to have mainly followed the river waterways connecting the southern Balkans to north-central Europe".
More recently, Lacan
The migration of Neolithic farmers into Europe brought along several new adaptations.[41] The variation for light skin colour was introduced to Europe by the neolithic farmers.[41] After the arrival of the neolithic farmers, a SLC22A4 mutation was selected for, a mutation which probably arose to deal with ergothioneine deficiency but increases the risk of ulcerative colitis, coeliac disease, and irritable bowel syndrome.
Bronze Age
The Bronze Age saw the development of long-distance trading networks, particularly along the Atlantic Coast and in the Danube valley. There was migration from Norway to Orkney and Shetland in this period (and to a lesser extent to mainland Scotland and Ireland). There was also migration from Germany to eastern England. Martin Richards estimated that there was about 4% mtDNA immigration to Europe in the Bronze Age.
Another theory about the origin of the Indo-European language centres around a hypothetical Proto-Indo-European people, who, according to the Kurgan hypothesis, can be traced to north of the Black and Caspian Seas at about 4500 BCE.[55] They domesticated the horse and possibly invented the wooden disk wheel, and are considered to have spread their culture and genes across Europe.[56] The Y haplogroup R1a is a proposed marker of these "Kurgan" genes, as is the Y Haplogroup R1b, although these haplogroups as a whole may be much older than the language family.[57]
In the far north, carriers of the
The relationship between roles of European and Asian colonists in the prehistory of Finland is a point of some contention, and some scholars insist that Finns are "predominantly Eastern European and made up of people who trekked north from the Ukrainian refuge during the Ice Age".
The Yamnaya component contains partial ancestry from an Ancient North Eurasian component, a Paleolithic Siberian lineage but closely related to European hunter-gatherers, first identified in
Up to a half of the Yamnaya component may have come from a
According to Lazaridis et al. (2016), a population related to the people of the Chalcolithic Iran contributed to roughly half of the ancestry of Yamnaya populations of the Pontic–Caspian steppe. These Iranian Chalcolithic people were a mixture of "the Neolithic people of western Iran, the Levant, and Caucasus Hunter Gatherers."[66]
The genetic variations for
Recent history
During the period of the
Given their small numbers and varied origins, Romanization does not appear to have left distinct genetic signatures in Europe. Indeed, Romance-speaking populations in the Balkans, like Romanians, Aromanians, Moldovans, etc. have been found to genetically resemble neighbouring Greek and South Slavic-speaking peoples rather than modern Italians.[70][71] Steven Bird has speculated that E1b1b1a was spread during the Roman era through Thracian and Dacian populations from the Balkans into the rest of Europe.[54]
Concerning the late Roman period of (not only)
Genetics of modern European populations
Patrilineal studies
There are four main Y-chromosome DNA haplogroups that account for most of Europe's patrilineal descent.[29]
- Haplogroup eastern Germany, and northern Italy. It drops outside this area and is around 30% or less in areas such as southern Italy, Poland, the Balkans and Cyprus. R1b remains the most common clade as one moves east to Germany, while farther east, in Poland, R1a is more common (see below).[77] In Southeast Europe, R1b drops behind R1a in the area in and around Hungary and Serbia but is more common both to the south and north of this region.[37] R1b in Western Europe is dominated by at least two sub-clades, R-U106, which is distributed from the east side of the Rhine into northern and central Europe (with a strong presence in England) and R-P312, which is most common west of the Rhine, including the British Isles.[74][75]
- Haplogroup R1a, almost entirely in the R1a1a sub-clade, is prevalent in much of Eastern and Central Europe (also in South and Central Asia). For example, there is a sharp increase in R1a1 and decrease in R1b1b2 as one goes east from Germany to Poland.[77] It also has a substantial presence in Scandinavia (particularly Norway).[78][79] In the Baltic countries R1a frequencies decrease from Lithuania (45%) to Estonia (around 30%).[80]
- Haplogroup I is found in the form of various sub-clades throughout Europe and is found at highest frequencies in the Nordic countries as I1 (Norway, Denmark, Sweden, Finland) and in the Balkan Peninsula as I2a (Bosnia and Herzegovina 65%,[60] Croatia and Serbia). I1 is also frequent in Germany, Great Britain and Netherlands, while I2a is frequent also in Sardinia, Romania/Moldova, Bulgaria and Ukraine. This clade is found at its highest expression by far in Europe and may have been there since before the LGM.[35]
- Haplogroup North Africa subclade E-M81 is also present in Sicily and Andalusia.
Putting aside small enclaves, there are also several haplogroups apart from the above four that are less prominent or most common only in certain areas of Europe.
- Haplogroup G, a common haplogroup among European Neolithic farmers, is common in most parts of Europe at a low frequency, reaching peaks above 70% around Georgia and among the Madjars (although living in Asia they border the eastern perimeter of Europe), up to 10% in Sardinia, 12% in Corsica and Uppsala (Sweden), 11% in the Balkans and Portugal, 10% in Spain and 9% in European Russia. This clade is also found in the Near East.
- Haplogroup N, is common only in the northeast of Europe and in the form of its N1c1 sub-clade reaches frequencies of approximately 60% among Finns and approximately 40% among Estonians, Latvians, and Lithuanians.
- Western Asia and the Eastern Mediterranean.[82]
Matrilineal studies
There have been a number of studies about the mitochondrial DNA haplogroups (mtDNA) in Europe. In contrast to Y DNA haplogroups, mtDNA haplogroups did not show as much geographical patterning, but were more evenly ubiquitous. Apart from the outlying Saami, all Europeans are characterised by the predominance of haplogroups H, U and T. The lack of observable geographic structuring of mtDNA may be due to socio-cultural factors, namely the phenomena of polygyny and patrilocality.[48]
Genetic studies suggest some maternal gene flow to eastern Europe from eastern Asia or southern Siberia 13,000 – 6,600 years BP.[83] Analysis of Neolithic skeletons in the Great Hungarian Plain found a high frequency of eastern Asian mtDNA haplogroups, some of which survive in modern eastern European populations.[83] Maternal gene flow to Europe from sub-Saharan Africa began as early as 11,000 years BP, although the majority of lineages, approximately 65%, are estimated to have arrived more recently, including during the Romanization period, the Arab conquests of southern Europe, and during the Atlantic slave trade.[84]
European population sub-structure
Genetically, Europe is relatively homogeneous, but distinct sub-population patterns of various types of genetic markers have been found,[85] particularly along a southeast–northwest cline.[86] For example, Cavalli-Sforza's principal component analyses revealed five major clinal patterns throughout Europe, and similar patterns have continued to be found in more recent studies.[85]: 291–296
- A cline of genes with highest frequencies in the Middle East, spreading to lowest levels northwest. Cavalli-Sforza originally described this as faithfully reflecting the spread of agriculture in Neolithic times. This has been the general tendency in interpretation of all genes with this pattern.
- A cline of genes with highest frequencies among Samiin the extreme north east, and spreading to lowest frequencies in the south west.
- A cline of genes with highest frequencies in the area of the lower Saami speakers in the extreme north of Scandinavia. Cavalli-Sforza associated this with the spread of Indo-European languages, which he links in turn to a "secondary expansion" after the spread of agriculture, associated with animal grazing.
- A cline of genes with highest frequencies in the Balkans and Southern Italy, spreading to lowest levels in Britain and the Basque country. Cavalli-Sforza associates this with "the Greek expansion, which reached its peak in historical times around 1000 and 500 BCE but which certainly began earlier".
- A cline of genes with highest frequencies in the Basque country, and lower levels beyond the area of Iberia and Southern France. In perhaps the most well-known conclusion from Cavalli-Sforza, this weakest of the five patterns was described as isolated remnants of the pre-Neolithic population of Europe, "who at least partially withstood the expansion of the cultivators". It corresponds roughly to the geographical spread of rhesus negative blood types. In particular, the conclusion that the Basques are a genetic isolate has become widely discussed, but also a controversial conclusion.
He also created a phylogenetic tree to analyse the internal relationships among Europeans. He found four major 'outliers'-
A study conducted in May of 2009[88] researching 19 populations from Europe using 270,000 SNPs highlighted the genetic diversity of European populations corresponding to the northwest to southeast gradient and distinguished "four several distinct regions" within Europe:
- Finland, showing the greatest distance to the rest of Europeans.
- the Baltic region (Estonia, Latvia and Lithuania), western Russia and eastern Poland.
- Central and Western Europe.
- Italy, due to the alps acting as a great genetic barrier.
In this study, barrier analysis revealed "genetic barriers" between Finland, Italy and other countries and that barriers could also be demonstrated within Finland (between Helsinki and Kuusamo) and Italy (between northern and southern part, Fst=0.0050). Fst (Fixation index) was found to correlate considerably with geographic distances ranging from ≤0.0010 for neighbouring populations to 0.0200–0.0230 for Southern Italy and Finland. For comparisons, pair-wise Fst of non-European samples were as follows: Europeans – Africans (Yoruba) 0.1530; Europeans – Chinese 0.1100; Africans (Yoruba) – Chinese 0.1900.[89]
A study by Chao Tian in August 2009 extended the analysis of European population genetic structure to include additional southern European groups and Arab populations (
Autosomal DNA
Seldin (2006) used over 5,000 autosomal SNPs. It showed "a consistent and reproducible distinction between ‘northern’ and ‘southern’
A similar study in 2007 using samples predominantly from Europe found that the most important genetic differentiation in Europe occurs on a line from the north to the south-east (northern Europe to the Balkans), with another east–west axis of differentiation across Europe. Its findings were consistent with earlier results based on mtDNA and Y-chromosomal DNA that support the theory that modern Iberians (Spanish and Portuguese) hold the most ancient European genetic ancestry, as well as separating Basques and Sami from other European populations.[92]
It suggested that the English and Irish cluster with other Northern and Eastern Europeans such as Germans and Poles, while some Basque and Italian individuals also clustered with Northern Europeans. Despite these stratifications, it noted that "there is low apparent diversity in Europe with the entire continent-wide samples only marginally more dispersed than single population samples elsewhere in the world".[92]
In 2008, two international research teams published analyses of large-scale genotyping of large samples of Europeans, using over 300,000 autosomal SNPs. With the exception of usual isolates such as
Two whole-genome studies of the two Eastern European populations in Ukraine (
According to geneticist David Reich, based on ancient human genomes that his laboratory sequenced in 2016, Europeans descend from a mixture of four West-Eurasian ancestral components, namely WHG (Western Hunter-gatherers), EHG (Eastern Hunter-gatherers), Neolithic farmers from the Levant/Anatolia as well as from Neolithic farmers from Iran (often summarized as "EEF"; Early European farmers), in varying degrees.[98][99]
Siberian geneflow is found among several Uralic-speaking European ethnic groups. This Siberian component is itself a composition of Ancient North Eurasian and East Asian-related ancestry from Eastern Siberia, maximized among
Modern Europeans are genetically rather homogeneous and derive their ancestry - predominantly to exclusively - from up to five West-Eurasian lineages, in varying degrees. Modern Europeans show affinity and continuity to ancient European hunter-gatherers (
Autosomal genetic distances (Fst) based on SNPs (2009)
The genetic distance between populations is often measured by
The values range from 0 to 1. A zero value implies that the two populations are panmictic, that they are interbreeding freely. A value of one would imply that the two populations are completely separate. The greater the Fst value, the greater the genetic distance. Essentially, these low Fst values suggest that the majority of genetic variation is at the level of individuals within the same population group (~ 85%); whilst belonging to a different population group within same ‘race’/ continent, and even to different racial/ continental groups added a much smaller degree of variation (3–8%; 6–11%, respectively).
Italian Americans | Palestinians | Swedes | Druzes
|
Spaniards | Germans | Russians | Irish | Greek Americans | Ashkenazi Jews | Circassians | |
---|---|---|---|---|---|---|---|---|---|---|---|
Italian Americans | 0.0064 | 0.0064 | 0.0057 | 0.0010 | 0.0029 | 0.0088 | 0.0048 | 0.0000 | 0.0040 | 0.0067 | |
Palestinians | 0.0064 | 0.0191 | 0.0064 | 0.0101 | 0.0136 | 0.0202 | 0.0170 | 0.0057 | 0.0093 | 0.0108 | |
Swedes | 0.0064 | 0.0191 | 0.0167 | 0.0040 | 0.0007 | 0.0030 | 0.0020 | 0.0084 | 0.0120 | 0.0117 | |
Druzes | 0.0057 | 0.0064 | 0.0167 | 0.0096 | 0.0121 | 0.0194 | 0.0154 | 0.0052 | 0.0088 | 0.0092 | |
Spaniards | 0.0010 | 0.0101 | 0.0040 | 0.0096 | 0.0015 | 0.0070 | 0.0037 | 0.0035 | 0.0056 | 0.0090 | |
Germans | 0.0029 | 0.0136 | 0.0007 | 0.0121 | 0.0015 | 0.0030 | 0.0010 | 0.0039 | 0.0072 | 0.0089 | |
Russians | 0.0088 | 0.0202 | 0.0030 | 0.0194 | 0.0070 | 0.0030 | 0.0038 | 0.0108 | 0.0137 | 0.0120 | |
Irish | 0.0048 | 0.0170 | 0.0020 | 0.0154 | 0.0037 | 0.0010 | 0.0038 | 0.0067 | 0.0109 | 0.0110 | |
Greek Americans | 0.0000 | 0.0057 | 0.0084 | 0.0052 | 0.0035 | 0.0039 | 0.0108 | 0.0067 | 0.0042 | 0.0054 | |
Ashkenazi Jews | 0.0040 | 0.0093 | 0.0120 | 0.0088 | 0.0056 | 0.0072 | 0.0137 | 0.0109 | 0.0042 | 0.0107 | |
Circassians | 0.0067 | 0.0108 | 0.0117 | 0.0092 | 0.0090 | 0.0089 | 0.0120 | 0.0110 | 0.0054 | 0.0107 |
Austria | Bulgaria | Czech Republic | Estonia | Finland (Helsinki) | Finland (Kuusamo) | France | Northern Germany | Southern Germany | Hungary | Northern Italy | Southern Italy | Latvia | Lithuania | Poland | Russia | Spain | Sweden | Switzerland | CEU | ||
---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|
Austria | 1.14 | 1.08 | 1.58 | 2.24 | 3.30 | 1.16 | 1.10 | 1.04 | 1.04 | 1.49 | 1.79 | 1.85 | 1.70 | 1.19 | 1.47 | 1.41 | 1.21 | 1.19 | 1.12 | Austria | |
Bulgaria | 1.14 | 1.21 | 1.70 | 2.19 | 2.91 | 1.22 | 1.32 | 1.19 | 1.10 | 1.32 | 1.38 | 1.86 | 1.73 | 1.29 | 1.53 | 1.30 | 1.47 | 1.13 | 1.29 | Bulgaria | |
Czech Republic | 1.08 | 1.21 | 1.42 | 2.20 | 3.26 | 1.35 | 1.15 | 1.16 | 1.06 | 1.69 | 2.04 | 1.62 | 1.48 | 1.09 | 1.27 | 1.63 | 1.26 | 1.37 | 1.21 | Czech Republic | |
Estonia | 1.58 | 1.70 | 1.42 | 1.71 | 2.80 | 2.08 | 1.53 | 1.70 | 1.41 | 2.42 | 2.93 | 1.24 | 1.28 | 1.17 | 1.21 | 2.54 | 1.49 | 2.16 | 1.59 | Estonia | |
Finland (Helsinki) | 2.24 | 2.19 | 2.20 | 1.71 | 1.86 | 2.69 | 2.17 | 2.35 | 1.87 | 2.82 | 3.37 | 2.31 | 2.33 | 1.75 | 2.10 | 3.14 | 1.89 | 2.77 | 1.99 | Finland (Helsinki) | |
Finland (Kuusamo) | 3.30 | 2.91 | 3.26 | 2.80 | 1.86 | 3.72 | 3.27 | 3.46 | 2.68 | 3.64 | 4.18 | 3.33 | 3.37 | 2.49 | 3.16 | 4.21 | 2.87 | 3.83 | 2.89 | Finland (Kuusamo) | |
France | 1.16 | 1.22 | 1.35 | 2.08 | 2.69 | 3.72 | 1.25 | 1.12 | 1.16 | 1.38 | 1.68 | 2.40 | 2.20 | 1.44 | 1.94 | 1.13 | 1.38 | 1.10 | 1.13 | France | |
Northern Germany | 1.10 | 1.32 | 1.15 | 1.53 | 2.17 | 3.27 | 1.25 | 1.08 | 1.11 | 1.72 | 2.14 | 1.84 | 1.66 | 1.18 | 1.49 | 1.62 | 1.12 | 1.36 | 1.06 | Northern Germany | |
Southern Germany | 1.04 | 1.19 | 1.16 | 1.70 | 2.35 | 3.46 | 1.12 | 1.08 | 1.08 | 1.53 | 1.85 | 1.20 | 1.84 | 1.23 | 1.58 | 1.40 | 1.21 | 1.17 | 1.07 | Southern Germany | |
Hungary | 1.04 | 1.10 | 1.06 | 1.41 | 1.87 | 2.68 | 1.16 | 1.11 | 1.08 | 1.42 | 1.63 | 1.58 | 1.46 | 1.14 | 1.28 | 1.32 | 1.22 | 1.16 | 1.13 | Hungary | |
Northern Italy | 1.49 | 1.32 | 1.69 | 2.42 | 2.82 | 3.64 | 1.38 | 1.72 | 1.53 | 1.42 | 1.54 | 2.64 | 2.48 | 1.75 | 2.24 | 1.42 | 1.86 | 1.36 | 1.56 | Northern Italy | |
Southern Italy | 1.79 | 1.38 | 2.04 | 2.93 | 3.37 | 4.18 | 1.68 | 2.14 | 1.85 | 1.63 | 1.54 | 3.14 | 2.96 | 1.99 | 2.68 | 1.67 | 2.28 | 1.54 | 1.84 | Southern Italy | |
Latvia | 1.85 | 1.86 | 1.62 | 1.24 | 2.31 | 3.33 | 2.40 | 1.84 | 1.20 | 1.58 | 2.64 | 3.14 | 1.20 | 1.26 | 1.84 | 2.82 | 1.89 | 2.52 | 1.87 | Latvia | |
Lithuania | 1.70 | 1.73 | 1.48 | 1.28 | 2.33 | 3.37 | 2.20 | 1.66 | 1.84 | 1.46 | 2.48 | 2.96 | 1.20 | 1.20 | 1.26 | 2.62 | 1.74 | 2.29 | 1.74 | Lithuania | |
Poland | 1.19 | 1.29 | 1.09 | 1.17 | 1.75 | 2.49 | 1.44 | 1.18 | 1.23 | 1.14 | 1.75 | 1.99 | 1.26 | 1.20 | 1.18 | 1.66 | 1.30 | 1.46 | 1.28 | Poland | |
Russia | 1.47 | 1.53 | 1.27 | 1.21 | 2.10 | 3.16 | 1.94 | 1.49 | 1.58 | 1.28 | 2.24 | 2.68 | 1.84 | 1.26 | 1.18 | 2.32 | 1.59 | 1.20 | 1.56 | Russia | |
Spain | 1.41 | 1.30 | 1.63 | 2.54 | 3.14 | 4.21 | 1.13 | 1.62 | 1.40 | 1.32 | 1.42 | 1.67 | 2.82 | 2.62 | 1.66 | 2.32 | 1.73 | 1.16 | 1.34 | Spain | |
Sweden | 1.21 | 1.47 | 1.26 | 1.49 | 1.89 | 2.87 | 1.38 | 1.12 | 1.21 | 1.22 | 1.86 | 2.28 | 1.89 | 1.74 | 1.30 | 1.59 | 1.73 | 1.50 | 1.09 | Sweden | |
Switzerland | 1.19 | 1.13 | 1.37 | 2.16 | 2.77 | 3.83 | 1.10 | 1.36 | 1.17 | 1.16 | 1.36 | 1.54 | 2.52 | 2.29 | 1.46 | 1.20 | 1.16 | 1.50 | 1.21 | Switzerland | |
CEU | 1.12 | 1.29 | 1.21 | 1.59 | 1.99 | 2.89 | 1.13 | 1.06 | 1.07 | 1.13 | 1.56 | 1.84 | 1.87 | 1.74 | 1.28 | 1.56 | 1.34 | 1.09 | 1.21 | CEU | |
Austria | Bulgaria | Czech Republic | Estonia | Finland (Helsinki) | Finland (Kuusamo) | France | Northern Germany | Southern Germany | Hungary | Northern Italy | Southern Italy | Latvia | Lithuania | Poland | Russia | Spain | Sweden | Switzerland | CEU |
CEU – Utah residents with ancestry from Northern and Western Europe.
History of research
Classical genetic markers (by proxy)
One of the first scholars to perform genetic studies was
From this, he constructed phylogenetic trees that showed genetic distances diagrammatically. His team also performed principal component analyses, which is good at analysing multivariate data with minimal loss of information. The information that is lost can be partly restored by generating a second principal component, and so on.[85]: 39 In turn, the information from each individual principal component (PC) can be presented graphically in synthetic maps. These maps show peaks and troughs, which represent populations whose gene frequencies take extreme values compared to others in the studied area.[85]: 51
Peaks and troughs usually connected by smooth gradients are called clines. Genetic clines can be generated by adaptation to environment (natural selection), continuous gene flow between two initially different populations or a demographic expansion into a scarcely populated environment, with little initial admixture with existing populations.[107]: 390 Cavalli-Sforza connected these gradients with postulated pre-historic population movements, based on archaeological and linguistic theories. However, given that the time depths of such patterns are not known, "associating them with particular demographic events is usually speculative".[48]
Direct DNA analysis
Studies using direct DNA analysis are now abundant and may use mitochondrial DNA (mtDNA), the non-recombining portion of the Y chromosome (NRY), or even autosomal DNA. MtDNA and NRY DNA share some similar features, which have made them particularly useful in genetic anthropology. These properties include the direct, unaltered inheritance of mtDNA and NRY DNA from mother to offspring and father to son, respectively, without the 'scrambling' effects of genetic recombination. We also presume that these genetic loci are not affected by natural selection and that the major process responsible for changes in base pairs has been mutation (which can be calculated).[27]: 58
The smaller effective population size of the NRY and mtDNA enhances the consequences of drift and founder effect, relative to the autosomes, making NRY and mtDNA variation a potentially sensitive index of population composition.[48][32][29] These biologically plausible assumptions are not concrete; Rosser suggests that climatic conditions may affect the fertility of certain lineages.[48]
The underlying mutation rate used by the geneticists is more questionable. They often use different mutation rates and studies frequently arrive at vastly different conclusions.[48] NRY and mtDNA may be so susceptible to drift that some ancient patterns may have become obscured. Another assumption is that population genealogies are approximated by allele genealogies. Guido Barbujani points out that this only holds if population groups develop from a genetically monomorphic set of founders. Barbujani argues that there is no reason to believe that Europe was colonised by monomorphic populations. This would result in an overestimation of haplogroup age, thus falsely extending the demographic history of Europe into the Late Paleolithic rather than the Neolithic era.[108] Greater certainty about chronology may be obtained from studies of ancient DNA (see below), but so far these have been comparatively few.
Whereas Y-DNA and mtDNA haplogroups represent but a small component of a person's DNA pool, autosomal DNA has the advantage of containing hundreds of thousands of examinable genetic loci, thus giving a more complete picture of genetic composition. Descent relationships can only be determined on a statistical basis, because autosomal DNA undergoes recombination. A single chromosome can record a history for each gene. Autosomal studies are much more reliable for showing the relationships between existing populations, but do not offer the possibilities for unravelling their histories in the same way as mtDNA and NRY DNA studies promise, despite their many complications.
Genetic studies operate on numerous assumptions and suffer from methodological limitations, such as selection bias and confounding phenomena like genetic drift, foundation and bottleneck effects cause large errors, particularly in haplogroup studies. No matter how accurate the methodology, conclusions derived from such studies are compiled on the basis of how the author envisages their data fits with established archaeological or linguistic theories.[citation needed]
See also
- General
- Genetics by European group
- Genetic history of Italy
- Genetic history of the British Isles
- Genetic history of the Iberian Peninsula
- Genetic studies on Bosniaks
- Genetic studies on Bulgarians
- Genetic studies on Croats
- Genetic studies on Jews
- Genetic studies on Russians
- Genetic studies on Sami
- Genetic studies on Serbs
- Genetic studies on Turkish people
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Further reading
- Rodríguez-Varela, Ricardo et al. "The genetic history of Scandinavia from the Roman Iron Age to the present". In: Cell. Volume 186, Issue 1, 5 January 2023, Pages 32–46.e19.
- Skourtanioti, E., Ringbauer, H., Gnecchi Ruscone, G.A. et al. "Ancient DNA reveals admixture history and endogamy in the prehistoric Aegean". In: Nature Ecology & Evolution (2023). https://doi.org/10.1038/s41559-022-01952-3