Genes Mirror Migrations and Cultures in Prehistoric Europe—A Population
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Available online at www.sciencedirect.com ScienceDirect Genes mirror migrations and cultures in prehistoric Europe — a population genomic perspective Torsten Gu¨ nther and Mattias Jakobsson Genomic information from ancient human remains is beginning Recent years have seen technological and methodological to show its full potential for learning about human prehistory. advances which gave rise to the field of ancient genomics. We review the last few years’ dramatic finds about European By studying the genomes of individuals from the past, we prehistory based on genomic data from humans that lived are able to investigate individuals (and groups) that lived many millennia ago and relate it to modern-day patterns of during those times and during particular events and genomic variation. The early times, the Upper Paleolithic, periods. This new approach has provided unprecedented appears to contain several population turn-overs followed by power and resolution to study human prehistory. In more stable populations after the Last Glacial Maximum and contrast to prior views of population continuity, we during the Mesolithic. Some 11 000 years ago the migrations now know that migrations (from various areas) were driving the Neolithic transition start from around Anatolia and numerous during the past and that they shaped mod- reach the north and the west of Europe millennia later followed ern-day patterns of variation, in combination with isola- by major migrations during the Bronze Age. These findings tion-by-distance processes. European prehistory has show that culture and lifestyle were major determinants of proven to be more complex than what would be conclud- genomic differentiation and similarity in pre-historic Europe ed from the most parsimonious models based on modern- rather than geography as is the case today. day population-genomic data [9]. Here, we review the most recent findings about European prehistory based on Address ancient genomics. Department of Organismal Biology and SciLife Lab, Evolutionary Biology Centre, Uppsala University, Sweden European hunter-gatherers during the Upper Corresponding author: Jakobsson, Mattias ([email protected]) Palaeolithic and the Mesolithic The first anatomically modern humans spread over Eur- ope around 45 000 years ago [10–13] where they met [11] Current Opinion in Genetics & Development 2016, 41:115–123 and mixed with Neandertals [14–16]. The contributions This review comes from a themed issue on Genetics of human origin of these early Europeans to modern populations appear Edited by Joshua Akey and Anna Di Rienzo limited [5 ,16], and mitochondrial data shows that some haplogroups found in some individuals who lived in Europe during the Upper Palaeolithic are not present in modern-day Europeans, which suggests loss of some http://dx.doi.org/10.1016/j.gde.2016.09.004 genetic variation or replacement through later migrations 0959-437X/# 2016 Elsevier Ltd. All rights reserved. [5 ,17]. However, European individuals that lived after 37 000 years ago, in the Upper Palaeolithic, have been suggested to already carry major genetic components found in today’s Europe [4 ,5 ]. The peoples living in Europe during these times were obviously influenced by climatic changes, at least in terms of the geographic areas The genetic makeup of modern-day European groups and that were habitable. For instance, during the Last Glacial the historical processes which generated these patterns of Maximum (LGM) [26 500–19 000 years ago, 18], the population structure and isolation-by-distance have north of Europe became uninhabitable and the remaining attracted great interest from population geneticists, anthro- habitable areas became fragmented [19,20]. The LGM pologists, historians and archaeologists. Early studies of likely caused a severe bottleneck for the groups at the genetic variation among Europeans indicated that the time [21], when people may have moved south into major axes of variation are highly correlated with geogra- specific refugia, which is indicated by low genomic diver- phy [1], a pattern that has been confirmed and refined sity of later hunter-gatherer groups [22 ] and reduced during the age of population-genomics (Figure 1a) mitochondrial diversity [17]. After the LGM, migrants [2,3]. Today, we have learned from studying the genomics from the southeast appear in Europe and mixed with the of past peoples of Europe that the modern-day patterns of local European post ice age groups [5 ] resulting in a genomic variation were shaped by several major demo- population that may have remained largely uniform dur- graphic events in the past. These events include the first ing the late Upper Paleolithic and possibly into the peopling of Europe [4 ,5 ], the Neolithic transition [6], Mesolithic in some areas [5 ,23,24 ,25 ], although this and later migrations during the Bronze Age [7 ,8 ]. interpretation is still uncertain as only a limited number of www.sciencedirect.com Current Opinion in Genetics & Development 2016, 41:115–123 116 Genetics of human origin Figure 1 (a)Today (b) 36,000-7,500 BP Tajik Russian Finnish North_Ossetian LezginChechenBalkar KumykAdygei LithuanianEstonian BelarusianUkrainian Abkhasian Czech NorwegianIcelandicHungarian Georgian OrcadianScottishCroatian Turkish English Bulgarian Armenian PC2 French Albanian PC2 Italian Cypriot Spanish Maltese Basque Druze Eastern Upper Palaeolithic (37kya) Central Upper Palaeolithic (30kya) Iberian Upper Paleolithic (19kya) Southern Late Upper Palaeolithic (14kya) Sardinian Central Late Upper Palaeolithic (14kya) Caucasus Palaeolithic (13kya)/Mesolithic (10kya) –0.10 –0.05 0.00 0.05 0.10 Central Mesolithic (8kya) –0.10Iberian –0.05 Mesolithic 0.00 (8kya) 0.05 0.10 Scandinavian Mesolithic (8kya) Eastern Mesolithic (7.5kya) –0.10 –0.05 0.00 0.05 0.10 –0.10 –0.05 0.00 0.05 0.10 PC1 PC1 (c)8,000-4,500 BP (d) 5,000-3,000 BP PC2 PC2 Northern (farmer) Northern (hunter gatherer) Central –0.10Hungary –0.05 0.00 0.05 0.10 Iberia Pontic steppe –0.10 –0.05 0.00 0.05 0.10 Anatolia Ireland Hungary Central Central Ireland Iberia Southern Northern –0.10 –0.05 0.00 0.05 0.10 –0.10 –0.05 0.00 0.05 0.10 PC1 PC1 Current Opinion in Genetics & Development Principal component analyses for different time periods and focusing on western Eurasia. (a) Modern-day samples, (b) Upper Palaeolithic and Mesolithic samples, including samples identified as representative for prehistoric clusters in [5 ], arrows show transitions during the Upper Palaeolithic, (c) early and middle Neolithic samples, and (d) late Neolithic and Bronze Age samples. Individuals from previous sub-figures are depicted in gray. geographic areas in Europe have been investigated. The from ancient north Eurasian populations [5 ,7 ,27 ] genomic make-up of these hunter-gatherers falls outside (Figure 2). Scandinavian Mesolithic individuals of the genetic variation of modern-day people, including [22 ,24 ] show genetic affinities to both the western western Eurasians (Figure 1B). The genomic patterns of and the eastern Mesolithic groups, suggesting a continu- western [23,26] and central European Mesolithic individ- um of genetic variation, possibly caused by admixture uals [24 ,25 ] appear distinct from eastern European from both geographic areas. Such a scenario would be Mesolithic individuals who showed a stronger influence consistent with patterns seen in many animal and plant Current Opinion in Genetics & Development 2016, 41:115–123 www.sciencedirect.com Population genomics of prehistoric Europe Gu¨ nther and Jakobsson 117 Figure 2 Sample Age [Years before present] 40000 35000 10000 5000 0 GBR IBS TSI FIN Anatolia Pontic Steppe Northern Hungary Ireland Central Northerb Iberia Northern Central + Western Ireland Russia Southern Central Iberia Central Hungary Kostenki14 Upper Paleolithic Early Neolithic Middle Neolithic Late Neolithic + Bronze Age + Mesolithic + Chalcolithic FIN GBR TSI IBS Current Opinion in Genetics & Development Supervised Admixture [75] analysis of ancient samples and modern populations from the 1000 genomes project [76 ] shown together with the approximate geographic origin of the ancient samples and the sample age. Admixture was run in supervised mode using Western hunter- gatherers, Anatolian Neolithic farmers and Yamnaya herders from the Pontic steppe as ‘source’ populations [7 ]. Note that these three ‘source’ groups were themselves mixed groups (see text). Pie charts show admixture proportions for modern European populations from the 1000 genomes project. species which recolonized the Scandinavian peninsula scale networks across Eurasia already during the Upper from the south and north-east after the last glacial period Palaeolithic and Mesolithic periods. [28–30]. Surprisingly, some of these Scandinavian hunter- gatherers were carriers of the derived allele in the EDAR The Neolithic transition gene [27 ]. EDAR affects hair thickness and teeth mor- The Neolithic transition — the transformation from a phology and has been subject to a selective sweep in East hunter-gatherer lifestyle to a sedentary farming life- Asia, but the derived allele is virtually absent from style — has been an exceptionally important change in modern day Europe [27 ]. Together with evidence from human history and forms the basis for emergence of stone tool technology [31] and the potential introduction civilizations. This transition occurred independently in of domesticated dogs from Asia [32–34] this suggests large different parts of the world [35]. For western Eurasia, the www.sciencedirect.com