The genetic history of the British Isles is the subject of research within the larger field of human population genetics. It has developed in parallel with DNA testing technologies capable of identifying genetic similarities and differences between both modern and ancient populations. The conclusions of population genetics regarding the British Isles in turn draw upon and contribute to the larger field of understanding the history of the human occupation of the area, complementing work in linguistics, archaeology, history and genealogy. Research concerning the most important routes of migration into the British Isles is the subject of debate. Apart from the most obvious route across the narrowest point of the English Channel into Kent, other routes may have been important over the millennia, including a land bridge in the Mesolithic period, as well as maritime connections along the Atlantic coasts. Genetic studies have revealed multiple migration waves into Britain and Ireland from the Palaeolithic onwards, with detectable regional differences among present-day populations. After the Last Glacial Maximum, hunter-gatherer groups carrying two distinct ancestries (GoyetQ2-related and Villabruna-related) repopulated Britain and Ireland, with the latter eventually becoming dominant. This hunter-gatherer ancestry was substantially replaced during the Neolithic revolution, c. 4000 BC, by groups carrying Early European Farmer (EEF) ancestry from the European mainland, who admixed to a certain extent with the existing hunter-gatherer population in some regions. At the start of the Bronze Age, another major population replacement occurred when migrating Bell Beaker groups, carrying a high proportion of Steppe-related ancestry, replaced around 90% of the Neolithic gene pool. Throughout the Bronze and Iron Ages, further migration from mainland Europe raised the proportion of Early European Farmer ancestry in southern Britain. This has been proposed as one possible mechanism for the introduction of Celtic languages. Other potentially important historical periods of migration that have been subject to consideration in this field include the Roman era, the period of early Germanic influx, the Viking era, the Norman invasion of 1066, and the era of the European wars of religion.
History of research Early studies by Luigi Cavalli-Sforza used polymorphisms from proteins found within human blood (such as the ABO blood groups, Rhesus blood antigens, HLA loci, immunoglobulins, G6PD isoenzymes, amongst others). One of the lasting proposals of this study with regards to Europe is that within most of the continent the majority of genetic diversity may best be explained by immigration coming from the southeast towards the northwest or in other words from the Middle East towards Britain and Ireland. Cavalli-Sforza proposed at the time that the invention of agriculture might be the best explanation for this. With the advent of DNA analysis modern populations were sampled for mitochondrial DNA to study the female line of descent and Y chromosome DNA to study male descent. As opposed to large scale sampling within the autosomal DNA, Y DNA and mitochondrial DNA represent specific types of genetic descent and can therefore reflect only particular aspects of past human movement. Later projects began to use autosomal DNA to gather a more complete picture of an individual's genome. For Britain, major research projects aimed at collecting data include the Oxford Genetic Atlas Project (OGAP) and more recently the People of the British Isles, also associated with Oxford. Owing to the difficulty of modelling the contributions of historical migration events to modern populations based purely on modern genetic data, such studies often varied significantly in their conclusions. One early Y DNA study estimated a complete genetic replacement by the Anglo-Saxons, whilst another argued that it was impossible to distinguish between the contributions of the Anglo-Saxons and Vikings and that the contribution of the latter may even have been higher.
Analyses of nuclear and ancient DNA
Paleolithic The earliest modern humans to inhabit Britain during the Initial Upper Paleolithic (c. 45-40,000 years ago) produced stone tools of the Lincombian-Ranisian-Jerzmanowician (LRJ) technocomplex. While no Initial Upper Paleolithic remains from Britain have been genetically sampled, sampling of LRJ-associated individuals from Germany suggests that they belonged to an early diverging branch of the Out of Africa population, less closely related to modern non-Africans than all modern non-Africans are to each other, and unrelated to later Upper Palaeolithic European hunter-gatherers. These people carried the introgressed Neanderthal ancestry found in all modern non-Africans, from an interbreeding episode thought to have occurred around 45,000–49,000 years ago. European Upper Paleolithic hunter-gatherers from at least 37,000 years ago onwards are part of the "West Eurasian" branch, more closely related to modern Europeans than East Asians. After the Last Glacial Maximum (LGM) resulted in Britain being deserted, there is evidence of repopulation of Britain and Ireland during the late Upper Paleolithic from c. 13,500 BC. The earliest inhabitants of Britain after recolonisation (as evidenced by sites such as Gough's Cave) were members of the Magdalenian culture, which evidence suggests had expanded from southern France and Spain after the end of the LGM, and had genetic continuity from members of the earlier Gravettian culture. From around 14,000 years ago, Magdalenian peoples were replaced across northern Europe, including in Britain, by people belonging to the genetically distinct Villabruna cluster, associated with the Epigravettian and related archaeological cultures. In Britain, this transition is evidenced by an individual from Kendrick's Cave, dated to approximately 14,100–13,460 years Before Present.
Mesolithic population British Mesolithic hunter-gatherers, such as the famous Cheddar Man, were closely related to other Mesolithic people throughout Western Europe (the so-called Western Hunter Gatherer cluster, which are direct descendants of people from the Villabruna cluster) This population probably had blue or green eyes, lactose intolerance, dark hair and dark to very dark skin. British Mesolithic people probably contribute negligible ancestry to modern British people.
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