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A new press release is circulating on the paper which I blogged a few months ago, Ancient Admixture in Human History. Unlike the paper, the title of the press release is misleading, and unfortunately I notice that people are circulating it, and probably misunderstanding what is going on. Here’s the title and first paragraph:

Native Americans and Northern Europeans More Closely Related Than Previously Thought

Released: 11/30/2012 2:00 PM EST
Source: Genetics Society of America

Newswise — BETHESDA, MD – November 30, 2012 — Using genetic analyses, scientists have discovered that Northern European populations—including British, Scandinavians, French, and some Eastern Europeans—descend from a mixture of two very different ancestral populations, and one of these populations is related to Native Americans. This discovery helps fill gaps in scientific understanding of both Native American and Northern European ancestry, while providing an explanation for some genetic similarities among what would otherwise seem to be very divergent groups. This research was published in the November 2012 issue of the Genetics Society of America’s journal GENETICS

 

The reality is ta Native Americans and Northern Europeans are not more “closely related” genetically than they were before this paper. There has been no great change to standard genetic distance measures or phylogeographic understanding of human genetic variation. A measure of relatedness is to a great extent a summary of historical and genealogical processes, and as such it collapses a great deal of disparate elements together into one description. What the paper in Genetics outlined was the excavation of specific historically contingent processes which result in the summaries of relatedness which we are presented with, whether they be principal component analysis, Fst, or model-based clustering.

What I’m getting at can be easily illustrated by a concrete example. To the left is a 23andMe chromosome 1 “ancestry painting” of two individuals. On the left is me, and the right is a friend. The orange represents “Asian ancestry,” and the blue represents “European” ancestry. We are both ~50% of both ancestral components. This is a correct summary of our ancestry, as far as it goes. But you need some more information. My friend has a Chinese father and a European mother. In contrast, I am South Asian, and the end product of an ancient admixture event. You can’t tell that from a simple recitation of ancestral quanta. But it is clear when you look at the distribution of ancestry on the chromosomes. My components have been mixed and matched by recombination, because there have been many generations between the original admixture and myself. In contrast, my friend has not had any recombination events between his ancestral components, because he is the first generation of that combination.

So what the paper publicized in the press release does is present methods to reconstruct exactly how patterns of relatedness came to be, rather than reiterating well understood patterns of relatedness. With the rise of whole-genome sequencing and more powerful computational resources to reconstruct genealogies we’ll be seeing much more of this to come in the future, so it is important that people are not misled as to the details of the implications.

(Republished from Discover/GNXP by permission of author or representative)
 
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The Pith: You’re Asian. Yes, you!

A conclusion to an important paper, Nick Patterson, Priya Moorjani, Yontao Luo, Swapan Mallick, Nadin Rohland, Yiping Zhan, Teri Genschoreck, Teresa Webster, and David Reich:

In particular, we have presented evidence suggesting that the genetic history of Europe from around 5000 B.C. includes:

1. The arrival of Neolithic farmers probably from the Middle East.

2. Nearly complete replacement of the indigenous Mesolithic southern European populations by Neolithic migrants, and admixture between the Neolithic farmers and the indigenous Europeans in the north.

3. Substantial population movement into Spain occurring around the same time as the archaeologically attested Bell-Beaker phenomenon (HARRISON, 1980).

4. Subsequent mating between peoples of neighboring regions, resulting in isolation-by-distance (LAO et al., 2008; NOVEMBRE et al., 2008). This tended to smooth out population structure that existed 4,000 years ago.

Further, the populations of Sardinia and the Basque country today have been substantially less influenced by these events.

 

It’s in Genetics, Ancient Admixture in Human History. Reading through it I can see why it wasn’t published in Nature or Science: methods are of the essence. The authors review five population genetic statistics of phylogenetic and evolutionary genetic import, before moving onto the novel results. These statistics, which measure the possibility of admixture, the extent of admixture, and the date of admixture, are often presented, but nested into supplements, in previous papers by the same group. On the one hand this removes from view the engines which are driving the science. On the other hand I have always appreciated that a benefit of this injustice to the methods which make insight possible is that those without academic access can actually bite into the meat of the researcher’s mode of thought.

I did read through the methods. Twice. I’ve encountered all the statistics before, and I’ve read how they were generated, but I’ll be honest and admit that I haven’t internalized them. That has to end now, because the authors have finally released a software package which implements the statistics, ADMIXTOOLS. I plan to use it in the near future, and it is generally best if you understand the underlying mechanisms of a software package if you are at the bleeding end of analytics. I will review the technical points in more detail in future posts, more for my own edification than yours. But for the moment I’ll be a bit more cursory. Four of the tests use comparisons of allele frequencies along explicit phylogenetic trees. That’s so general as to be uninformative as a description, but I think it’s accurate to the best of my knowledge. In the basics the tests are seeing if a model fits the data (as opposed to TreeMix, which finds the best model out of a range to fit the data). The last method, rolloff, infers the timing of an admixture event based upon the decay of linkage disequilibrium. In short, admixture between two very distinct populations has the concrete result of producing striking genomic correlations. Over time these correlations dissipate due to recombination. The magnitude of dissipation can allow one to gauge the time in the past when the original admixture occurred.

Let’s look at some results. To the left is a section of a table which illustrates the most significant 3-population test scores in the HGDP. The authors checked all the various combinations, and these came out at the top as likely admixtures (i.e., the two sources produce particular patterns in the target). Please remember that these triads should not be taken literally. The Uygur are not descended from Japanese and Italians. Rather, they are descended from populations with genetic affinities to these two sources. Precisely, the Uygurs are descended from Northeast Asian Turks, who assimilated an Indo-European speaking substratum. Most of the results are rather obvious and explicable. Several Middle Eastern populations are known to have Sub-Saharan African admixture, and this is shows up in the results. Others may be more confusing because of the obscurity of the populations, but the Burusho clearly have ancient East Asian ancestry on clustering algorithms, so their presence is not surprising to me. Similarly, the Russians in the HGDP data set have an ‘eastern’ affinity (or at least some do), either due to Finno-Ugric or Turkic ancestry (Tatars regularly assimilated into a Russian ethnic identity as the Tsars expanded their domains).

Some of the other results are more confusing, but one can still find a historical explanation. I have seen evidence that some of the Cambodian samples may have old Indian admixture, though it is not entirely clear to me. But that could explain why there is a signature of West Eurasian admixture into this population (though one wonders why the donor was not Baloch or Pathan.). The Xibo and Tu are Northeast Asian groups, on the border between China proper and the great Eurasian interior. West Eurasian admixture into these groups is not unexpected. West Eurasians are historically attested among the mercenaries and soldiers who arrived on the North China plain after the collapse of the Han dynasty, down to the Alans who served under Kublai Khan. Some of Mongolian and Turkic peoples have individuals who are attested as having characteristics more typical of Europeans (e.g., red hair), so it is likely that this admixture was relatively old and widespread, well before the era of the Pax Mongolica.

There is a minor dissonant note in these results above. The authors used rolloff and inferred an admixture of ~800 years before the present. This is far lower than earlier estimates, which were >2,000 years before the present. First, I have to say that I was mildly skeptical of the higher value reported earlier. From what little I know the roiling of Turco-Mongol peoples which reordered the Inner Asian landscape did not really establish itself beyond the Chinese fringe at this time. Recall that Central Asia was the domain of the Iranians from prehistory down to the Islamic age (the full transition of Central Asia from Persianate to Turkic has not completed itself to this date, though it has progressed over the centuries since 1000 A.D.). Is it creditable that the Turkic hordes were shut on the other side of the Pamirs for ~1,000 years? Perhaps. But it should warrant skepticism, and openness to the lower values proffered here. The technical reason that the authors consider is that STRUCTURE based inferences may overestimate admixture when reference populations are not appropriate. And yet the authors still concede that 800 years is simply difficult to credit when one consults the historical literature. Strangely though it does align with the date of the Mongol ascendancy, during which time the Uygurs served as civil servants in the barbarian empire (Mongol script derives from the old Uygur script). I managed to dig up a cave painting of Uygurs from this period. There is surely artistic license, but they look rather East Asian to me, as opposed to the hybrid Eurasian appearance modal among modern Uygurs. I won’t touch upon the rather fraught and complex ethnology and ethnogenesis of modern Uygurs, and their relationship to Russian and Chinese ethnographers, but suffice it to say that one needs to be careful about excessive reliance on the literality of historical documents in this area, because of semantic confusions.

So let’s move to the main course: what’s going on in Europe? Before putting the spotlight on the macro picture, let’s highlight one secondary aspect: the authors detect evidence of massive gene flow into Spain from Northern Europe ~4,000 years before the present. I’ll let them speak here:

We hypothesize that we are seeing here a genetic signal of the ‘Bell-Beaker culture’ (HARRISON, 1980). Initial cultural flow of the Bell-Beakers appears to have been from South to North, but the full story may be complex. Indeed one hypothesis is that after an initial expansion from Iberia there was a reverse flow back to Iberia (CZEBRESZUK, 2003); this ‘reflux’ model is broadly concordant with our genetic results, and if this is the correct explanation it suggests that this reverse flow may have been accompanied by substantial population movement.

Two things to hammer home here. First, pots move with people. That’s the inference being drawn from the results. It’s not pots-not-people, it’s people-and-pots. Second, the idea of reversals in the direction of gene flow are intriguing, and, I think need to be taken more seriously. It seems the most plausible candidate here are the people who later became the Celtiberians. Celts have been associated with the Bell Beakers before.

But the bigger shock is that Europeans, and especially Northern Europeans, seem to have a substantial Northeast Asian component. From the nature of the prose I feel that the authors were definitely taken aback. They basically say so in so many words. In the process of resolving their confusion they skinned the cat every which way. And it does look to me that Northern Europeans are truly descended in part from a population which has affinities to the “First Americans.” I say this specifically because the Siberian samples they tested actually gave a weaker result than the South American Amerindians on the 3-population test.

So what’s the proportion of ancestry? Using the Siberian population they came up with an interval of 5-18 percent in Northern Europeans. The authors used the Sardinians as their “pure” European reference, and admit that it is likely that their admixture estimate is lower than real value due to this fact. Inference is inference, do you trust this result? As it happens the authors also checked Ötzi the Iceman, and found that like the modern Sardinians he had very little Northeast Asian ancestry. Ötzi is dated to ~5,000 years before he present. Using rolloff the authors estimate an admixture date of ~4,000 years before the present, with an error of nearly 1,000. Additionally, using a different data set they came with an admixture date of ~2,000 years before the present. The latter is obviously wrong (they explain why this could happen in the text). But Ötzi seems to put a boundary on how early it could have been, at least in Southern Europe.

As of publication the authors did not have time to include a reference to this interesting nugget from the abstracts of ASHG 2012:

The complete genome of the 5,300 year old mummy of the Tyrolean Iceman, found in 1991 on a glacier near the border of Italy and Austria, has recently been published and yielded new insights into his origin and relationship to modern European populations. A key finding of this study has been an apparent recent common ancestry with individuals from Southern Europe, in particular Sardinians…We used unpublished data from whole genome sequencing of 452 Sardinian individuals, together with publicly available data from Complete Genomics and the 1000 Genomes project, to confirm that the Iceman is most closely related to contemporary Sardinians. An analysis of these data together with ancient DNA data from a recently published study on Neolithic farmers and hunter-gatherers from Sweden shows the Iceman most closely related to the farmer individual, but not the hunter-gatherers, with the Sardinians again being the contemporary Europeans with the highest affinity. Strikingly, an analysis including novel ancient DNA data from an early Iron Age individual from Bulgaria also shows the strongest affinity of this individual with modern-day Sardinians. Our results show that the Tyrolean Iceman was not a recent migrant from Sardinia, but rather that among contemporary Europeans, Sardinians represent the population most closely related to populations present in the Southern Alpine region around 5000 years ago. The genetic affinity of ancient DNA samples from distant parts of Europe with Sardinians also suggests that this genetic signature was much more widespread across Europe during the Bronze Age.

I’m betting that this Bulgarian sample won’t exhibit Northeast Asian ancestry, though who knows?

There is a definite geographic pattern within Europe to the strength of the signature of admixture. Northern European populations have the greatest, Southern European populations less, and islanders like Cypriots hardly any. Recall that Sardinians seem to be the best reference, so the ~0 floor may just be a statistical artifact of the measuring stick we have. All that being said, what went on <5,000 years before the present to reorder the European landscape?

The answer may sound crazy, but I think the most probable explanation (even if it is unlikely) is something to do with the Indo-Europeans. We know that Indo-European languages were spoken in Greece by ~1500 BC at the latest. One thing that is clear from less advanced clustering algorithms is that Basques and Finns are somewhat distinctive in relation to their neighbors. Though they are not genetically that different, they still lack some “interesting”elements. The results to the left are from Dienekes, though I’ve replicated it. You can see a similar difference between French, and French Basques. The Basques seem to lack something which has affinities with West Asia. These results, and hints elsewhere, imply that the Basque may not be descended from hunter-gatherers, but the first European farmers. So who came after them?

Though it strikes me as a bizarre conjecture, but I can’t help but imagine the rapid expansion of Indo-European populations into Europe, pushing into the peninsulas of the south. These people may have been a newly formed cosmopolitan mix of West Asians, Northern European Mesolithics, and Northeast Asians. I am at a loss to hazard a guess as to who the First American-like Northeast Asians were, though perhaps they were a western offshoot of the Kets? These people were then absorbed into a melange of tribes who themselves emerged from a synthesis between immigrant West Asian farmers and Northern Europeans. In shorthand: perhaps the Indo-Europeans were mongrels! This is not an entirely crazy proposition if you look at the historical record. Conquest populations often synthesized and absorbed those who they conquered. Sometimes they even became the conquered in deep cultural ways (e.g., the Bulgars).


To ward off accusations of glib and facile speculations, I well understand that much of what I suggest above is likely wrong. But bizarre results are going to elicit unhinged hypotheses. And I shouldn’t overplay how strange these results are, I think they are going to stand the test of time. The authors are top notch, and Dr. Joseph Pickrell found the same pattern (a connection between Europeans and Native Americans) with TreeMix! If we sit back and reflect on phenotype it shouldn’t be entirely surprising. Some Scandinavians have always struck me as having a generalized Eurasian cast to their features. Obviously this tendency is stronger among the Sami and Finns, but you can see it in Swedes and others. This is far less evident to me among Southern European peoples. I doubt one would ever confuse a Sardinian for a Eurasian, and I never had that feeling when I spent some time in Italy a few years back (in contrast, some Finns did look Asiatic to me).

Finally, this paper highlights the reality that population genetics has little to do with Plato. A population within a species is simply not clear and distinct in a sense which would satisfy an Idealist. The authors of the above paper nod to this, illustrating how their tests for admixture are confounded and confused by constant gene flow via isolation-by-distance dynamics. These results indicate that Northern Europeans are on the order of 10% Northeast Asian. Does this mean that Northern Europeans are 10% non-white? Well, it turns out that white people were always 10% non-white! We just didn’t know. Is my daughter (who is 50% Northern European) now majority non-white? Oh wait, I’m South Asian. That means I’m ~50% white! Is my friend who is 25% Japanese now more than 25% Northeast Asian? Words and concepts fail us on the boundary of unfamiliarity, in time and space. Populations and genealogies don’t brook our categorizations. On a deep level we are all admixtures, and partitioning of ancestry along phylogenetic trees are useful and comprehensible fictions. These techniques put flesh upon the bones of archaeology and smoke out the outlines of history. But we always need to be aware that that history is not made by humans, rather, we excavating it, and then giving it appropriate glosses in our museums. And yet it is.

Related: Dienekes has much to say (obviously).

Image credit: Wikipedia, Wikipedia, and Wikipedia.

Cite: 10.1534/genetics.112.145037

(Republished from Discover/GNXP by permission of author or representative)
 
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I cropped the image above from the paper Inference of Population Structure using Dense Haplotype Data. The main reason was emphasize the distinctiveness of the Sardinian cluster, on the bottom right. As you can see this population exhibits a lot of coancestry across individuals. This isn’t too surprising, Sardinia is an island, and islands are often genetically distinctive. Random genetic drift prevents populations from diverging through gene flow, but water is a major impediment to gradual isolation by distance dynamics. The original Sardinians are naturally going to diverge from mainlanders over time, and begin to share the same set of common ancestors in the recent past, because their space of reasonable mating possibilities is constrained. The other population which is similar in the heat map above are the residents of the Orkneys, off the north coast of Scotland (the Orkneys has a much smaller population than Sardinia, but, it is also much closer to the mainland).

This is on my mind because Dienekes has a long post where he explores the D-statistic results of various European populations, using Sardinians as one of the references. You don’t need to know the details, just that Northern European populations seem to exhibit an affinity to East Asians. Our favorite human genomicsts at Broad highlighted this tendency, and Dienekes went looking (if David Reich et al. were inclined toward mischief they should just posit some crazy scenario, and see if Dienekes assembles the requisite data set!). Which is all fine, and we’ll see more of this in the near future. It isn’t as if others aren’t using Sardinians.

If you play around with model-based clustering packages (e.g., ADMIXTURE, STRUCTURE, etc.) or PCA the distinctiveness of the Sardinians does jump out at you. In short they are the least Asian of the European populations. If there is a cline, they generally define one of the antipodes of the cline (often, along with the Basques). They’re a natural reference. But we need to be cautious. On occasion a reader has casually asserted that understanding the relationships between populations is easy enough, just find appropriate reference groups. Easier said that done.

A few years ago there was a great deal of debate whether the Sardinians were Neolithic “newcomers” or the descendants of Paleolithic populations. The latter is actually somewhat conjectural in regards to archaeology. There is some evidence of Paleolithic habitation, but it is fragmentary, implying perhaps very low population densities, and likely local extinctions over time. The reason the argument roiled for so long is that if Sardinians are the descendants of Paleolithic Europeans you have the perfect reference. But what if they’re not? I think that question is up in the air, especially in light of the fact that Ötzi the Iceman has affinities to the Sardinians. Ötzi was a mainlander, and an agriculturalist. That does not preclude descent from a common Paleolithic Western European population which adopted agriculture, but I think it tilts us more toward some sort of demographic expansion (likely from the Eastern Mediterranean, but perhaps a secondary expansion from somewhere in the Western Mediterranean).

Ten years ago we were looking at a two-factor model. Europeans the Ice Age hunters, or Europeans the Middle Eastern farmers. Today the past is much more crisp, and yet that crispness has unveiled a befuddling complexity. If I had to put money down I would bet that in another ten years we’ll have to add into our understanding both demographic pulses during the Ice Age, and multiple ones which post-date the Neolithic. It may be that the signals of a relationship between Northern Europeans and the First Americans is very old, and dates back to the Gravettian culture, which spanned Western Europe eastward toward the fringes of the Black Sea. With ancient DNA I believe that we’ll get a sense of who the “First Europeans” were, and it may be that their genetic contribution to modern Europeans is in the same range as the Neandertals whom they assimilated and exterminated!

All something to keep in mind in the coming years, when European paleogenetics may be measured by the Sardinian….

Image credit: Wikimedia

(Republished from Discover/GNXP by permission of author or representative)
 
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genmap1A few years ago you started seeing the crest of studies which basically took several hundred individuals (or thousands) from a range of locations, and then extracted out the two largest components of genetic variation from the hundreds of thousands of variants. The clusters which fell out of the genetic data, with each point being an individual’s position, were transposed onto a geographical map. The figure to the left (from this paper) has been widely circulated. You don’t have to be a deep thinker to understand why things shake out this way; people are more closely related to those near than those far because gene flow ties populations together, and its power decreases as a function of distance.

Of course the world isn’t flat, and history perturbs regularities. Jews for example often don’t shake out where they “should” geographically, because of their historical mobility contingent upon random and often capricious geopolitical or social pressures. The Hazara of Afghanistan have their ethnogenesis in the melange of peoples who were thrown together after the Mongol conquest of Central Asia and Iran in the 13th century, and the subsequent collapse of the Ilkhan dynasty. Though the Hazara have mixed with their Persian, Tajik and Pashtun neighbors, they still retain a strong stamp of Mongolian ancestry which means that they are at some remove on the “genetic map” from their geographical neighbors.


So when interpreting these sorts of results you have two extreme dynamics operative. On the one hand you have an equilibrium state where gene flow is mediated through continuous but small flows of migration; women moving between villages, younger sons venturing out of the village in search of better opportunities. Then you have the random (or perhaps modeled as a poisson distribution) “shocks” which are attributed to world-historical (or region-historical) events which leave an outsized and often perplexing stamp and distort the genetic map from the geographic one. Sometimes the two are not in balance. In much of the New World and Australasia the native populations were genetically replaced by settlers from the outside. Thousands of years of genetic variation accumulated and shaped by localized gene flow events were wiped clean off the map by the demographic tsunami.

Obviously that’s an extreme scenario. The macroscale does not always render the microscale irrelevant in such a fashion. A new short paper in The European Journal of Human Genetics gives us an example. Genes predict village of origin in rural Europe:

The genetic structure of human populations is important in population genetics, forensics and medicine. Using genome-wide scans and individuals with all four grandparents born in the same settlement, we here demonstrate remarkable geographical structure across 8–30 km in three different parts of rural Europe. After excluding close kin and inbreeding, village of origin could still be predicted correctly on the basis of genetic data for 89–100% of individuals.

Here’s the ubiquitous PC chart, except on the scale of villages:

village1

As noted above they excluded close relatives, out to second cousins. They judge the genetic time depth is about ~120 years into the past back to the common ancestry. Remember that if their grandparents are from this village they obviously are going to be somewhat inbred, from the perspective of an American whose ancestors are from different nations. But for most of history the European case was the typical one, not the American one where people from different continents mingled.

Here’s part of the discussion which I think needs highlighting:

To explore how many markers are required to recover these fine scale patterns of structure, we ranked SNPs by FST among villages and repeated the PCA for the most differentiated subsets of 30 000, 10 000, 3000 and 300 SNPs in each population. In all three populations, 10 000 or more high FST SNPs recovered an essentially identical picture to that using the full data set, and even 3000 SNPs preserved considerable separation between the villages (not shown). Using only the most discriminating 300 SNPs, little structure could be observed between the two Croatian villages; however, in Scotland and Italy one of the three settlements included in each location remained completely differentiated from the other two (not shown). We note that these results are only indicative of the minimum number of SNPs required to separate these populations, as by necessity SNPs have been selected intrinsically on the basis of FST within the same data set, rather than extrinsically from other data.

The slightly lower differentiation of the Croatian villages is not surprising given the fact that they are physically the closest of those considered here, being 8 km apart, with only low hills separating them. In contrast, the settlements in the Scottish Isles and Italy are separated by 15–30 km of sea in the former case, and of 3000 m mountains in the latter, although there are deep connecting valleys.

First, we get a sense of the range of informative markers necessary to discern population structure well in much of the Old World. For continental races (e.g., Europeans vs. East Asians) you need on the order of 10-100 markers to distinguish them with a high degree of confidence (closer to the low bound than the high). It looks like in the case of village vs. village differences, it will be on the order of 100-1000 markers. I suspect in Iraq or the Caucasus you’ll need less than 300 markers, because genetic differentiation is higher over a shorter distance due to inbreeding, ethnic diversity, and geography (more the former in Iraq, more the latter in the Caucasus). In contrast, in regions where geography is conducive to transport and local norms enforce exogamy I wouldn’t be surprised if you need more like a thousand markers.

Second, observe the importance of topographical detail. I have observed before than Sardinia is a genetic outlier in Europe. That’s not because Sardinians interbred with native elves of that island. Rather, a water barrier serves as a major check on continuous gene flow mediated by banal contacts (e.g., going to the market and meeting a person from the neighboring village). Islands become worlds unto themselves. Though they are effected by the exogenous shocks, they are less subject to the continuous gene flow at the equilibrium because the water serves as a barrier. Similarly mountains can produce genetic barriers as well, because they make travel rather difficult. In Consanguinity, Inbreeding, and Genetic Drift in Italy L. L. Cavalli-Sforza documents in detail through Roman Catholic Church records what a big impact modern roads had on inbreeding coefficients, which plunged in the 19th century. Distortions of the genetic map tells about variations in elevation in the third dimension on the geographic map!

The utility of this sort of data collection and analysis in the modern world is an empirical question. On the one hand many Europeans are relatively less inclined to move in comparison to Americans. And yet the breaking down of borders with the European Union and the likely need for a more productive economic sector on that continent because of changing demographics point to greater mobility, migration and mixing, which would make these sorts of studies of only near-term use. Of more interest to me are going to be fine-grained analyses of social groups. For example the Indian caste system. Last fall in the Reich et al. paper the authors seemed to be indicating the likelihood of a lot of between population variance groups these groups. It doesn’t matter if a particular Bania sub-caste from Gujarat is scattered across the world, from Kenya to England to the United States. They may all still marry amongst a set of individuals who hale from the same original few villages.

Good times.

Citation: O’Dushlaine, C., McQuillan, R., Weale, M., Crouch, D., Johansson, Aulchenko, Y., Franklin, C., Polašek, O., Fuchsberger, C., Corvin, A., Hicks, A., Vitart, V., Hayward, C., Wild, S., Meitinger, T., van Duijn, C., Gyllensten, U., Wright, A., Campbell, H., Pramstaller, P., Rudan, I., & Wilson, J. (2010). Genes predict village of origin in rural Europe European Journal of Human Genetics DOI: 10.1038/ejhg.2010.92

(Republished from Discover/GNXP by permission of author or representative)
 
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Razib Khan
About Razib Khan

"I have degrees in biology and biochemistry, a passion for genetics, history, and philosophy, and shrimp is my favorite food. If you want to know more, see the links at http://www.razib.com"