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nbsp; * First submitted in 2009, updated in December 2011.
CHAPTER 7
LANGUAGE, GENES, AND CULTURAL INTERACTION
STEPHEN SHENNAN
THE most extensive and long-lasting, sometimes passionate, debate in studies of the European Neolithic has concerned the extent to which farming was introduced through a process of indigenous adoption or the expansion of farming populations. It is in the context of this debate that arguments about languages and genes, and the application of mathematical methods from genetics, were introduced into the study of the European Neolithic and continue to play a major role. In this essay I will review the history of arguments about genes and languages and how they have contributed to this debate, examine the current state of this field, and conclude by relating the arguments to our growing knowledge of demographic patterns.
Following on the work of Clark (1965), Ammerman and Cavalli-Sforza (1971) showed that if the dates of the first arrival of farming in a given location were plotted against distance from the assumed origin of agriculture in south-west Asia (taken to be Jericho) there was a good fit to a straight-line relationship, with a correlation of 0.89, and an overall rate of spread of about 1km per year (using uncalibrated dates). They went on to propose that this rate made sense if farming spread by a process of ‘demic diffusion’ (Ammerman and Cavalli-Sforza 1973), a spatial expansion of population arising from the greater productivity of the new subsistence system compared with hunting and gathering, following a model created to describe the wave of advance of an advantageous gene and subsequently to describe the process of population expansion (see Steele 2009 for a review). In this model the rate of increase in the size of the population at a given place depends on a population growth component and a diffusion component. The rate of growth depends on the size of the initial population, the maximum growth rate, and the nearness of the population size to the local carrying capacity, because as the population nears carrying capacity growth slows down until a density-dependent equilibrium is reached. The value of the diffusion component depends on the average dispersal distance of individuals in the time between birth and reproduction. The speed at which the wave of expanding population travels is dependent on the diffusion value and the maximum population growth value. Ammerman and Cavalli-Sforza found that if they took ethnographically derived values for growth rates and marriage distances in farming populations and plugged these into the relevant equations the resulting rate of spread corresponded well to that actually observed.
Subsequently, a principal components analysis summarizing the variation in a large number of genetic markers in modern European populations (this was before actual DNA information from present-day populations became available) turned out to show that the main pattern of variation corresponded with a strong spatial trend from south-east to north-west Europe (Menozzi et al. 1978; Ammerman and Cavalli-Sforza 1984). The authors postulated that the trend could be accounted for by their demic diffusion model for the spread of farming, with the immigrant farming signal being successively diluted over time by intermarriage with indigenous hunter-gatherers. On this basis, a link was established between a postulated process accounting for the origin and spread of farming into Europe between 9,000 and 6,000 years ago and evidence from the genetics of present-day populations. But it was also possible to link genetic patterns to the distribution of languages, it was argued, since an expanding population would carry its language with it, and its descendant languages as time went on, not just its genes.
As far as Europe is concerned, the key question about languages, going back to the late eighteenth century when its existence was first proposed, has always been how to account for the pattern of relationships between the languages of the Indo-European family, distributed as they are all the way from Ireland to India and believed by historical linguists to descend from a common ancestor, proto-Indo-European, estimated by most historical linguists to have existed about 6,000 years ago (but see below in the Language section). This question had been of interest to archaeologists prior to World War II, and was the subject of one of Gordon Childe’s early books, The Aryans (1926). However, because of its association with Nazi ideas of the existence and superiority of ‘the Aryan race’, questions about the relationship between archaeology and language largely disappeared from the mainstream European prehistory agenda after World War II.
The main exception to this trend, at least to the west of the Russian language sphere, where discussion continued, was the work of Marija Gimbutas (e.g. 1963), who proposed that the distribution could be accounted for by the spread of nomadic peoples east and west from the Ukrainian and Russian steppe north of the Black Sea, her suggested homeland of Proto-Indo-European speakers. These peoples were traceable archaeologically by their use of ‘kurgans’ (Russian for burial mounds), hence the term ‘kurgan hypothesis’ or ‘kurgan invasions’ to describe this spread, which could be dated archaeologically, it was argued, to around 4000–3000 BC, thus fitting the date proposed by many linguists for the initial split of proto-Indo-European. In subsequent work (e.g. 1991) Gimbutas went on to propose that the ‘kurgan invasions’ marked a major break in the prehistory of Europe because they were responsible not just for the introduction of new languages but for the demise of what she regarded as the matri-centred cultures of the early Neolithic—what she called ‘Old Europe’—and their replacement by a new set of male-centred social and cultural institutions.
In 1987 Colin Renfrew, in his book Archaeology and Language, brought the relationship between the two back into the archaeological mainstream in a way that could not be ignored, given his by then established position as one of the leading figures in the development of the New Archaeology and its introduction to Europe. In it he proposed there was no convincing evidence in the European Neolithic and Bronze Age after the initial spread of farming for large-scale migrations of the kind that would be required to account for the spread of Indo-European languages; thus they must have arrived with the expansion of population that introduced farming, by a process of demic diffusion of the kind proposed by Ammerman and Cavalli-Sforza, a suggestion he had first made in outline much earlier (Renfrew 1973). Importantly, their demic diffusion model showed how it was possible for populations to spread without mass migrations, since no individual on this model need move very far at all.
Where are we today? How have the ‘Wave of Advance’ model and genetic and linguistic arguments for it fared, and how do they relate to the archaeology? Both Ammerman and Cavalli-Sforza’s genetic arguments and Renfrew’s linguistic ones have proved extremely controversial, but also extremely productive, in that they have generated an enormous amount of new interdisciplinary research and have also created massive interest in a public well beyond archaeology. For archaeologists they have also proved quite problematic, in that both genetics and linguistics are highly technical subjects whose arguments are difficult to evaluate by those who do not have the appropriate training. In the remainder of this chapter I will review the current state of the various arguments, before concluding by considering the archaeological evidence for their demographic implications.
GENES
It is worth pointing out from the outset that what most geneticists have been concerned with by and large is trying to explain the proportions of gene variants of different putative origins in modern populations. This is not the same as investigating the proportion of migrants involved in the spread of farming, say, 8,000 years ago.
As far as Cavalli-Sforza’s and colleagues’ principal components analysis inferences are concerned, it has subsequently been shown that spatial clines based on such synthetic data can arise from a variety of different processes, including population replacement with successive founder effects (i.e. an immigration process without successive dilution by intermarriage with indigenous populations) (e.g. Barbujani et al. 1995); by gradients in the duration of natural selection, for example, if the farming way of life had an effect on the survival and reproductive success of people with certain g
enes that effect would have lasted more than 2,000 years longer in south-east than north-west Europe because farming arrived much earlier there (Fix 1996); or even as a result of simple ‘isolation by distance’, with gene flow between adjacent stable populations (Novembre and Stephens 2008; Ray and Excoffier 2009). Accordingly, this line of argument can no longer be considered valid.
The genetic studies that followed relied on newly available data on the lineages of mtDNA, relevant to female genealogies, and Y-chromosome DNA, relevant to males (Richards et al. 1996; Semino et al. 2001). On the basis of identifying specific gene lineages (or haplogroups) estimated to have a most recent common ancestor (MRCA) in the Neolithic as opposed to earlier—this is done by counting differences in mt or Y-DNA between present-day individuals, calculating the number of mutations required to get back to a common ancestor for those individuals, and then using the so-called ‘molecular clock’ to give a date in years with an error margin—they proposed a contribution (or admixture) of immigrant Near Eastern genes to the general European population today of 20% for mtDNA and 22% for Y-chromosome DNA, and saw this as strong evidence that the introduction of agriculture largely involved cultural diffusion between indigenous Mesolithic groups. However, it has been shown that there are many problems with this argument. First, the date of a MRCA does not tell us anything about a colonization date. If Mars is colonized in the future by a team consisting of individuals of Chinese, African, and European ancestry, the date of the MRCA will be well back in the Palaeolithic and certainly will not date the colonization of Mars! Second, other studies that attempted to explicitly model demographic events between 8,000 years ago and the present have come up with much higher contributions of ‘Neolithic’ genes to the present-day European gene pool (e.g. Chikhi et al. 2002; Currat and Excoffier 2005). More generally though, if one is trying to evaluate the extent of genetic admixture between two populations and their relative contributions to a descendant population, using one or two genetic loci does not provide enough information to do this with any confidence (Belle et al. 2006). Belle et al. therefore analysed data from 377 different autosomal genes, i.e. those where one copy is inherited from each parent. In order to do such admixture analyses one needs to identify present-day populations that can be regarded as roughly corresponding genetically to Mesolithic European populations of local Palaeolithic ancestry and to incoming Neolithic farmers. Like all previous investigators, they chose the current Basque population to represent the former and present-day Near Eastern populations for the latter. In all cases they found a cline, with the proportion of Near Eastern genes decreasing with distance from that region, at a very high level of correlation; but even at the extreme western end of the distribution the estimated proportion of genes of putatively Near Eastern origin in the present-day European gene pool was at least 50%. It is apparent that the implications of these varied results for the different versions of the demic diffusion and cultural diffusion models of the spread of farming into Europe remain unclear. Arguably, this lack of clarity is unsurprising given the complex chain of links between data from present-day populations and inferences about the origin of populations interacting (or not) 8,000 years ago, not least the questionable modern proxies that have been used for ancestral source populations. Moreover, such studies cannot take into account data from genetic lineages that have since gone extinct.